COMITÉ DE REDACCIÓN

ÉDITOR

Ángel Guerra Sierra

EDITORES ADJUNTOS

Eugenia M* Martínez Cueto-Felgueroso

Francisco Javier Rocha Valdés

Gonzalo Rodríguez Casero

Comité EDITORIAL

Kepa Altonaga Sustacha

Eduardo Angulo Pinedo

Rafael Araujo Armero

Thierry Backeljau

Rúdiger Bieler

Sigurd v. Boletzky

Jose Castillejo Murillo

Karl Edlinger

Antonio M. de Frias Martins

José Carlos García Gómez

Edmund Gittenberger

Serge Gofas

Gerhard Hoszprunar

Yuri 1. Kantor

Ángel Antonio Luque del Villar

María Yolanda Manga González

Jordi Martinell Callico

Ron K. 0'Dor

Tokashi Okutani

Marco Oliverio

Pablo E. Penchaszadeh

Winston E. Ponder

Carlos Enrique Prieto Sierra

Me de los Ángeles Ramos Sánchez

Paul G. Rodhouse

Joandoménec Ros ¡ Aragones

María Carmen Salas Casanovas

Gerhard Steiner

José Templado González

Victoriano Urgorri Carrasco

Anders Warén

PORTADA DE lberus

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 Poís Vasco, Bilbao, España

Universidad del País Vasco, Bilbao, España

Museo Nacional de Ciencias Naturales, Madrid, España

Institut Royal des Sciences Naturelles de Belgique, Bruselas, Bélgica

The Field Museum, Chicago, Estados Unidos

Laboratoire Arago, Banyuls-sur-Mer, Francia

Universidad de Santiago de Compostela, Santiago de Compostela, España

Noturhistorisches Museum Wien, Viena, Austria

Universidade dos Acores, Acores, Portugal

Universidad de Sevilla, Sevilla, España

National Natuurhistorisch Museum, Leiden, Holanda

Universidad de Málaga, España

Zoologische Staatssammlung Múnchen, Múnchen, Alemania

AN. Severtzov Institute of Ecology and Evolution, Moscú, Rusia

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Nihon University, Fujisawa City, Japón

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Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina

Australian Museum, Sydney, Australia

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Museo Nacional de Ciencias Naturales, CSIC, Madrid, España

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Universidad de Barcelona, Barcelona, España

Universidad de Málaga, Málaga, España

Institut fir Zoologie der Universitút Wien, Viena, Austria

Museo Nacional de Ciencias Naturales, CSIC, Madrid, España

Universidad de Santiago de Compostela, Santiago de Compostela, España

Swedish Museum of Natural History, Estocolmo, Suecia

lberus gualterianus (Linnaeus, 1758), una especie emblemática de la península Ibérica, que da

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

Iberus

REVISTA DE LA

SOCIEDAD ESPAÑOLA

DE MALACOLOGÍA

Vol. 13 (1) Oviedo, junio 2000

Iberus

Revista de la

SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

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

admiten también notas breves. /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

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

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trónico: backhuysCeuronet.nl

Los resumenes de los artículos editados en esta revista se publican en Aquatic Science

and Fisheries Abstracts (ASFA) y en el Zoological Records, BIOSIS.

Contents list published in Aquatic Science and Fisheries Abstracts and Zoological Records,

BIOSIS.

Dep. Leg. B-43072-81

ISSN 0212-3010

Diseño y maquetación: Gonzalo Rodríguez

Impresión: LOREDO, $. L. - Gijón

O Sociedad Española de Malacología ——_____———— lIberus, 1% (1): 1-14, 2000

Contribución al conocimiento de los Moluscos Opistobran-

quios de la costa de Granada (sureste de la Península Ibérica)

Contribution to the knowledge of the Opisthobranch Molluscs from

the coast of Granada (southeastern Spain)

Luis SÁNCHEZ-TOCINO*, Amelia OCAÑA?, Francisco J. GARCÍA**

Recibido el 26-IV-1999. Aceptado el 11-VII1-1999

RESUMEN

En un estudio realizado en el litoral granadino durante los años 1994-1999 se identificaron

84 especies de opistobranquios, siendo la distribución de especies por órdenes la siguiente:

Cephalaspidea, 4; Thecosomata, 1; Anaspidea, 3; Sacoglossa, 5; Notaspidea, 6 y Nudi-

branchia, 65. En el presente artículo se aportan datos sobre abundancia de las especies,

rangos de variación en la profundidad, distribución temporal y características del substrato

donde fueron observadas cada una de ellas. Del total de especies encontradas, 55 son nue-

vas citas para la costa granadina. Polycera faeroensis y Pruvotfolia pselliotes son nuevas ci-

tas para el Mar Mediterráneo, Geitodoris planata para el Mediterráneo ibérico y Hermaea

paucicirra lo es para el litoral Mediterráneo andaluz. La especie Tambja marbellensis es ci-

tada por primera vez desde su descripción.

ABSTRACT

In a study conducted on the coast of Granada (S. Spain) from 1994 to 1999, 84 opistho-

branchs species were identified. The distribution of the species by orders was: Cephalaspi-

dea, 4; Thecosomata, 1; Anaspidea, 3; Sacoglossa, 5; Notaspidea, 6 and

Nudibranchia, 65. Data are presented for species abundance, depth-variation ranges,

temporal distribution, and characteristics of the substrate were each species was found. Of

the species found, 55 are new citations for the Granada coast. Polycera faeroensis and

Pruvotfolia pselliotes are new citations for the Mediterranean Sea, Geitodoris planata for

the Iberian Mediterranean and Hermaea paucicirra for Andalusian Mediterranean. The

species Tambja marbellensis is collected for the first time since its description.

PALABRAS CLAVE: Mollusca, Opisthobranchia, Mediterráneo, Costa de Granada.

KEY WORDS: Mollusca, Opisthobranchia, Mediterranean, Coast of Granada

INTRODUCCIÓN

Pocos son los estudios malacológicos Granada. LUQUE (1983, 1986) estudió los

realizados hasta la fecha en la costa de gasterópodos de las costas de Málaga y

* Departamento de Biología Animal y Ecología, Facultad de Ciencias, Universidad de Granada, 18071 Granada

(Spain); e-mail: IstocinoCretemail.es; ameliaCgoliat.ugr.es

** Departamento de Fisiología y Biología Animal, Facultad de Biología, Universidad de Sevilla; Avda. Reina

Mercedes, 6; Apdo. 1095; 41080 Sevilla (Spain); e-mail: fjgarciaCcica. es

Iberus, 18 (1), 2000

Tabla 1. Coordenadas geográficas (GTM), de las distintas estaciones de muestreo.

Table I. Geographical coordinates (GTM) for the different sampling stations.

Estaciones de muestreo

Coordenadas GTM

1. La Rábita

2.Peñón de San Patricio

3. Castell de Ferro

4. La Rijana

5. Calahonda

6. Peñón de Salobreña

7. Barranco del Cambrón

8. Bahía de Velilla

9. Peñones de Almuñecar

10. Punta de San José

11. Punta de la Mona

12. Cerro Gordo

36” 45,4” N; 03” 10,5" O

SON AOS MIA O

36” 44,2" N; 03” 20,5" O

JOR AZS NOIA ZO

NAS NOOO)

SO AN AOS AS OM O

36” 44,5" N; 03” 37,4 O

36" 44,8" N; 032 39,1 O - 36 44,5' N; 03 39,9 O

36” 43,6" N; 03 41.50

35” 43,9 N; 03” 42,6" O

367 43,5" N; 03 43,7" O - 36” 43,6" N; 03” 44,4 O

367 44,3" N; 03% 45,7" O- 36” 44,7" N; 03 46, 6 O

Granada, centrándose principalmente

en la primera de las dos provincias, ya

que sólo una de las 15 estaciones de

muestreo establecidas, la de La Herra-

dura, pertenece al litoral granadino. Del

mismo modo, TEMPLADO, LUQUE Y Mo-

RENO (1988) eligieron La Herradura co-

mo única estación de muestreo en la

costa de Granada. Posteriormente, como

consecuencia de la la campaña oceano-

gráfica “Fauna I”, TEMPLADO, GUERRA,

BEDOYA, MORENO, REMÓN, MALDONADO

Y RAMOS (1993) encontraron varias espe-

cies de opistobranquios en el circalitoral

frente a las costas granadinas. MORENO

Y TEMPLADO (1998) completan los estu-

dios anteriores en las provincias de Al-

mería y Granada, seleccionando, en esta

última, sólo 4 puntos, de los 23 muestre-

ados.

Con el presente estudio se pretende

ampliar el inventario de la fauna de

opistobranquios del litoral de la provin-

cia de Granada. Además, se aportan

datos sobre el número de ejemplares,

periodo de avistamiento a lo largo de

los años de muestreo, profundidad y

tipo de substrato donde se encontraron

cada una de las especies, así como las

citas de otros autores que han encon-

trado las especies relacionadas en el pre-

sente trabajo, en las distintas provincias

del litoral andaluz.

MATERIAL Y MÉTODOS

Se han seleccionado en total trece

estaciones de muestreo, doce en el

litoral granadino y una treceava que

corresponde a las muestras recolectadas

en barcos de arrastre. La selección de

dichas estaciones queda justificada por

la variabilidad de substratos y por tanto

de hábitats diferentes que se encuentran

en ellas, ya que es frecuente la combina-

ción, en varias de las mismas, de fondos

de cascajo, arena, fango, praderas de

fanerógamas o rocosos que en algunas

ocasiones, como es el caso de la Punta

de la Mona, alcanzan grandes profundi-

dades.

El área de estudio comprende una

linea litoral de unos 60 Km, desde el

límite con la provincia de Málaga por el

Oeste, hasta La Rábita por el Este. El

nombre de cada una de las doce estacio-

nes de muestreo aparecen en la Tabla 1

acompañadas de las correspondientes

coordenadas geográficas, G. T. M. Las

estaciones 1, 2, 3, 4, 5, 6, 7, 9 y 10 son

puntuales, mientras que las 8, 11 y 12

quedan definidas por la línea de costa

comprendida entre las dos coordenadas

geográficas que se citan en la Tabla 1. De

la estacion 13 se recogen muestras

mediante barcos de arrastre que faenan

frente a la costa de Motril a profundida-

SÁNCHEZ-TOCINO ET AL.: Opistobranquios de la costa de Granada

GRANADA

ESPAÑA

14]

Oo:

Figura 1. Mapa ilustrando las situación geográfica de las diferentes zonas de muestreo.

Figure 1. Illustrated map showing the geographic situation of the different sampling localities.

des comprendidas entre 50 y 150 m. En

la Figura 1 se muestra la ubicación de

las distintas estaciones de muestreo.

Se realizaron una serie de muestreos

preliminares desde Enero de 1994 hasta

Diciembre de 1995, que han servido

para seleccionar aquellas estaciones con

una mayor diversidad de opistobrán-

quios, que correspondieron con la 8, 11

y 12, y en las que se centraron los mues-

treos durante los años 1996, 1997, 1998 y

hasta Marzo de 1999.

El material examinado, a excepción

del recogido mediante arrastres, fué

obtenido en inmersiones con escafandra

autónoma, entre la zona mediolitoral y

40 m. de profundidad.

Para la recogida de datos se realiza-

ron en cada estación de muestreo tran-

sectos de longitud variable. En la esta-

ción 8, se utilizó también un segundo

método que consistió en colgar de una

boya, a 3 m de profundidad, una tela

mosquitera enrollada de 1,5 m de longi-

tud. El crecimiento de algas e hidroideos

fue bastante rápido, por lo que al cabo

de unos dos meses se pudieron recolec-

tar opistobranquios.

Los opistobranquios que ofrecían

alguna duda para su identificación

fueron colectados, fotografiados y gra-

bados en vídeo. Posteriormente los

ejemplares capturados se anestesiaron

mediante congelación en agua de mar o

añadiendo lentamente alcohol. Se fijaron

con formol al 4% y se conservaron en

alcohol al 70%.

RESULTADOS

En la Tabla II se aportan, para cada

una de las especies identificadas, los

siguientes datos: número que define la

zona de muestreo, mes/es de observa-

ción, año/s, abundancia, longitud (en

milímetros), tipo de substrato y profun-

didad (en metros). La abundancia es

referida con el siguiente criterio: E,

especie escasa de la que sólo se han

observado, en todo el periodo de

estudio, entre 1 y 10 ejemplares; PA,

especie poco abundante de la que se han

observado entre 10 y 50 individuos; A,

especie abundante de la que se han

observado entre 50 y 100 ejemplares;

Iberus, 18 (1), 2000

Tabla II. Especies observadas en la provincia de Granada acompañadas del número que define la

estación de muestreo donde aparecen cada una de ellas, mes/es de avistamiento, año/s, abundancia

relativa (E: entre 1 y 10 ejemplares; PA: entre 10 y 50 ejemplares; A: entre 50 y 100 individuos;

MA: más de 100 individuos observados en toda la campaña de muestreo), longitud (mm), tipo de

sustrato (A: algas; B: fondos blandos; BP: bajo piedra; BR: briozoos; C: cuevas, CA: cascajo; E:

esponjas; G: gorgonáceos; H: hidroideos; R: rocoso; S: flotando en superficie) y profundidad (m).

Table II. Species observed in the province of Granada, accompanied by the number that identifes the

sampling station where each specimen was observed, together with month/s, yearls, relative abundance

(E: between 1 and 10 specimens; PA: between 10 and 50 specimens; A: between 50 and 100 specimens;

MA: more than 100 specimens observed over the entire sampling campaign), length (mm), type of'subs-

trate (A: algas; soft sediments; BP: low rock; BR: briozoa; C: caves; CA: gravel; E: sponges; G: gorgona-

cea; H: hydroids; R: rock; S: floating on surface) and depth (m).

Estación Mes Año Abun. Long. Subs. Prof.

Orden CEPHALASPIDEA P. Fischer, 1883

Familia Bullidae Lamorck, 1801.

Bulla striata Bruguiére, 1792 8 10 95 E 15 B 5

Familia Aglojidae Pilsbry, 1895

Aglaja tricolorata Renier, 1804 6 1,8 97 E 10:12 B 5-8

Philinopsis depicta (Renier, 1807) 8 8 95 E 25 B 5

Familia Cylichnidae Adams H.£.A., 1854

Scaphander lignarius (Linnaeus, 1758) 13 6 97 E 40 B 50-150

Orden THECOSOMATA Bloinville, 1824

Familia Cymbuliidae Gray, 1840

Cymbulia peronii Lamarck, 1819 8 4,9 94 E 30 S 0

Orden ANASPIDEA P. Fischer, 1883

Aplysia fasciata Poiret, 1789 8, 11,12 7,10, 12 94,9 MA 200-300 BR 512

Aplysia parvula Guilding in Mórch, 1863 8,11 4,8 95, 97 ESO ISA BRASS

Aplysia punctata (Cuvier, 1803) 8,11 9-10 94,96 EA ES

Orden NOTASPIDEA P. Fischer, 1883

Fomilia Umbraculidae Dall, 1889

Umbraculum umbraculum (Lighfoot, 1786) 12 1 97,98 E 120-150 CR 59

Fomilia Plevrobranchidae Férussac, 1822

Pleurobranchus testudinarivs Cantraine, 1835 11 4, 10,11 97 E 130-150 CA 520

Berthella ocellata (Delle-Chiaje, 1828) 12 2,17 % E 30-35 c 10-13

Berthella plumula (Montagu, 1803) M6 12 SSA E la AZ

Berthella stellata (Risso, 1826) 3,5, 12 1,11 96, 99 E 15 BP 37

Pleurobranchaea meckelii Bloinville, 1825 8,11, 12 2-5, 8, 10 95,96 MA 10-100 B,BP 39

Orden SACOGLOSSA Von Ihering, 1876

Familia Placobranchidae Rang, 1829

Thuridilla hopei (Vérany, 1853) AZ 1,3, 511 9599 PA 1020. AR 515

Elysia viridis (Montagu, 1804) 12 11,12 98 PAS A 30)

Familia Hermaeidae H. Adams y A. Adams, 1854

Hermaea paucicirra Pruvot-Fol, 1953 1 1 99 E 1-2 A 5

Familia Polybranchiidae ODonoghue, 1929

Coliphylla mediterranea A. Costa, 1867 5 1 99 E 5-7 A 3

Familia Limapontiidae Gray, 1847

Placida verticillata Ortea, 1981 11,12 ZETA! 98,99 A 13 A 510

Orden NUDIBRANCHIA Blainville, 1814

Suborden DORIDACEA Odhner, 1934

Familia Aegiretidae P. Fischer, 1883

Aegires leuckarti Vérany, 1853 11 | 99 E 3 R 10

Familia Goniodorididoe Adams H.y A., 1854

Trapania hispalensis Cervera y García-Gómez, 1989 2,4, 11,12 1-7, 9-11 95-99 PA 8-15 als

Trapania lineata Haefelfinger, 1960 1,12 1, 6-8, 10, 11 97,98 E 8-15 BPE 68

Trapania maculata Haefelfinger, 1960 11,12 1,3, 57, 9-12 95-98 PA 8-20 ER 615

SÁNCHEZ-TOCINO ET AL.: Opistobranquios de la costa de Granada

Tabla IL Continuación.

Table IT. Continuation

Familia Onchidorididae Alder y Hancock, 1845

Diaphorodoris luteocincta (Sars M., 1870)

Diaphorodoris papillata Portmann y Sandmeier, 1960

Familia Triophidae Odhner in Franc, 1968

Crimora papillata Alder y Hancock, 1862

Familia Aldisidae Odhner, 1933

Aldisa banyulensis Pruvot-Fol, 1951

Familia Chromodorididae Bergh,1891

Chromodoris britoi Ortea y Perez, 1983

Chromodoris krohni (Vérany, 1846)

Chromodoris luteorosea (Rapp, 1827)

Chromodoris purpurea (Risso in Guérin, 1831)

Cadlina pellucida (Risso, 1826)

Hypselodoris bilineata (Pruvotfol, 1953)

Hypselodoris cantabrica Bouchet y Ortea, 1980

Hypselodoris fontandraui (Pruvo+Fol, 1951)

Hypselodoris picta (Schultz in Philippi, 1836)

Hypselodoris midatlantica Gosliner, 1990

Hypselodoris villafranca (Risso, 1818)

Familia Discodorididae Bergh, 1891

Discodoris atromaculata (Bergh, 1880)

Discodoris fragilis (Alder y Hancock, 1864)

Discodoris rosi Ortea, 1979

Geitodoris portmanni (Schmekel, 1972)

Geitodoris planata (Alder y Hancock, 1846b)

Familia Kentrodorididue Bergh, 1892

Estación

11, 12

a UU) ee

2, WU, 2

10,11, 12

11, 12

11,12

111, 12

11,12

11, 12

1,4,5,7,8,9, 11, 12

4,11, 12

4,5,7,11, 12

4,11, 12

11, 12

9,11

Jorunna onubensis Cervera, Gorcio-cómez y Gorcía, 1986 12

Familia Plotydorididae Bergh, 1891

Platydoris argo (Linneo, 1767)

Familia Rostangidae PruvotFol, 1951

Rostanga rubra (Risso, 1818)

Familia Polyceridae Alder y Hancock, 1845

11,12

Polycera aurantiomarginata García-Gómez y Bobo, 1984 12

Polycera faeroensis Lemche, 1929

Polycera quadrilineata (0.F. Múller, 1776)

Limacia clavigera (0.F. Miller, 1776)

Roboastra europea García-Gómez, 1985

Tambja ceutae Gorcia-cómez y Ortea, 1988

Tambja marbellensis Schick y Cervera, 1998

Familia Dendrodorididae O'Donoghue, 1924

Dendrodoris limbata (Cuvier, 1804)

Doriopsilla areolata Bergh, 1880

Suborden DENDRONOTACEA Odhner, 1934

Familia Dotoidae Gray, 1853

Doto koenneckeri Lemche, 1976

Familia Hancockiidae Pruvot-Fol, 1954

Hancockia uncinata (Hesse, 1872)

Familia Tethyidae Alder y Hancock, 1855

Tethys fimbria Linné, 1767

Fomilia Tritoniidae Lamarck, 1809

Tritonia manicata Deshayes, 1853

Tritonia nilsodhneri Marcus Ev., 1983

Tritonia striata Haefelfinger, 1963

8, 11, 12

10,11, 12

11, 12

2 1H We

2,11, 12

Mes Año

1,2,4,8 96, 97-99

1-5, 8, 9-11 96-99

1,3,4 97-99

8,10 96, 97,98

1,2,7,8,11,12 96,98, 99

1-12 95-99

2,4,911,12 96-98

1-3, 5-7, 9-12 95-99

6,11, 12 95-98

15,7, 8, 10, 12 95-99

3 98

1,3:6,11,12 96-99

1-12 95-99

1-8,10, 11 95-99

1-5, 8, 10-12 95-99

1,3-8,11 94, 96-99

8 97

3,10 96, 97

8,9 96, 97

11 97

12 %

1-8,10,11 95-99

11 97

1-3, 11, 12 96-99

5,8 95, 97

1,3,6,7,12 96-98

1-4, 10-12 96, 99

1,7 95, 97

10,11 %

1-4 98

3-5, 7-11 95-97

2-6, 8, 10,12 94,95, 97-99

1,2, 10

8, 10

96, 91

96, 97

97,98

UE)

96-99

Abun.

Long.

10-20

510

10-15

15-20

515

3-30

10-25

8-10

130, 150

10-15

8-10

Subs. Prof.

LOS

el erS

All

BP,R 8-10

BP,R 5-20

R 720

BP,R 5-20

PESO

BP,E,R 6-15

R 59

RO 25

RE 6:28

BP,E,R 520

BP,E,R 5-12

BP, C, E, R 7-28

ELO

BPE 7-10

Hp 5%)

DEA al,

R 6

BP,E,R 5-20

E /

R 2530

RH 1-10

BP, ER 4-10

R 536

BP, BR,R 7-8

BP, BR 57

BP,CA,R 6-15

BP, ER 3-25

B 50-150

DIAAESO

Iberus, 18 (1), 2000

Tabla II. Continuación.

Table II. Continuation

Estación Mes Año Abun. long. Subs. Prof.

Mariona blainvillea (Risso, 1818) 10,11, 12 1-3, 5, 10,11 95-99 PA 1040 BP.G,R 5-10

Suborden ARMINACEA Odhner, 1934

Familia Arminidae Pruvo+Fol, 1927

Armina maculata Rofinesque, 1814 13 5,6 96, 97 PA 7090 B 50-150

Familia Zephyrinidae Iredale y O'Donoghue, 1923

Janolus cristatus (Delle Chiaje, 1841) 1,3411 12 1,47. 12 NI EOS R 6-7

Suborden AEOLIDACEA Odhner, 1934

Familia Aeolidiidae d'Orbigny, 1834

Aeolidiella soemmeringi (Leuckart, 1828) 4,11 2,4, 9-12 94-96,99 E 1520 BP 4-12

Berghia coerulescens Laurillard, 1830 11 2-8, 10,11 95,97-99 PA 20-30 BP 515

Berghia verrucicomis (A. Costa, 1867) 8 9,10 95,9 E 2025 H 1-3

Spurilla neapolitana (Delle-Chiaje, 1841) 3,8, 11 1,2, 611 97-98 PA 3-25 HBP 15

Familia Eubranchidae Odhner, 1934

Eubranchus farrani (Alder y Hancock, 1844) 6,8,11,12 1,2,5,9,10,12 96-99 PA 8-10 AHBP 1-3

Familia Focelinidae Bergh in Corus, 1889

Focelina annulicornis(Chamisso y Heysenhardt,1821) 8, 11 1-3, 6, 9,10, 12 96-98 A 1020 HBP 16

Facelina coronata (Forbes y Goodsir, 1839) 8,11 1-4, 6-8, 10, 12 96-98 A 10-25 CA 19

Facelina rubrovitatta (A. Costa, 1866) 11 49,10 97,98 PA 10-15 H 7-10

Caloria elegans (Alder y Hancock, 1845) 4, 5,11, 12 1-12 95-99 A 5-15 BP. HR 515

Cratena peregrina (Gmelin, 1791) 4,11, 12 1-11 95-99 MA 10-30 H 7-20

Pruvotfolia pselliotes (Labbé, 1923) 2 2 99 E 15 BP 4

Dicata odhneri Schmekel, 1967 11 4 98 E 3 (AH 9

Dondice banyulensis Portmann y Sandmeier,1960 4,11, 12 25,7,9-11 95-99 PA 1535 RH 1520

Familia Flobellinidae Bergh, 1889

Flabellina affinis (Gmelin, 1791) 4,5,7, 8, 9,10,11, 12 1-12 95-99 MA 5-30 HR 225

Flabellina ischitana Hirano y Thompson, 1990 11,12 1,2,5,8, 12 94-99 A 1020 HR 525

Flabellina babai Schmekel, 1972 11 3,4,8, 10 94, 97,98 PA 1060 CAR 9-16

Flabellina pedata (Montagu, 1815) 2,4,11,12 1,2,411 95-99 MA 525 H 615

Calmella cavolini (Vérany, 1846) 11,12 1,2,911 98, 99 PA 8-15 H 710

Fomilia Piseinotecidae Edmunds, 1970

Piseinotecus gabinieri (Vicente, 1975) 11, 12 2-10 95-99 A 1530 HR 915

Familia Tergipedidae Bergh, 1889

Cuthona genovae (0Donoghue, 1929) 8 5 98 E 6 H |

MA, especie muy abundante de la que

se han observado más de 100 individuos

durante todo el estudio. Los tipos de

substratos vienen indicados con las

abreviaturas: A (algas), B (fondos

blandos), BP (bajo piedra), BR (brio-

zO0OS), C (cuevas), CA (cascajo), E

(esponjas), G (gorgonáceos), H (hidroi-

deos), R (rocoso, cubierto fundamen-

talmente de algas fotófilas e hidroideos)

y S (especie encontrada flotando en

superficie).

En el litoral granadino se han identi-

ficado 84 especies de opistobranquios

distribuidas, en órdenes y subórdenes,

de la siguiente manera: Cephalaspidea, 4;

Thecosomata, 1; Anaspidea, 3; Notaspidea,

6; Sacoglossa, 5 y Nudibranchia, 65, repar-

tidos en Doridacea, 36; Dendronotacea, 7;

Arminacea, 2 y Aeolidacea, 20.

Como se desprende de la Tabla Il,

existen una serie de especies que han

aparecido en tres o más estaciones de

muestreo, prácticamente a lo largo de

todo el año y a lo largo de toda la

campaña de muestreo, aunque en oca-

siones de forma escasa O poco frecuente.

Éstas son: Berthella plumula, Chromodoris

britoi, C. krohni, Hypselodoris picta, H.

villafranca, Doriopsilla areolata, Mariona

SÁNCHEZ-TOCINO ET AL.: Opistobranquios de la costa de Granada

blainvillea, Eubranchus farrani, Caloria

elegans, Dondice banyulensis, Elabellina

affinis y F. pedata. Dichas especies las

podemos considerar como las más cons-

tantes del litoral granadino.

Por el contrario otras especies desta-

can por haber aparecido en una sola

estación de muestreo y únicamente en

una o dos ocasiones, siendo éste el caso

de: Umbraculum umbraculum, Berthella

ocellata, Elysia viridis, Hermaea pucicirra,

Caliphylla mediterranea, Aegires leuckarti,

Hypselodoris cantabrica, Discodoris fragilis,

Geitodoris planata, Jorunna onubenis, Ros-

tanga rubra, Polycera faeroensis, Roboastra

europea, Tambja ceutae, Hancockia unci-

nata, Tritonia manicata, Berghia verrucicor-

nis, Pruvotfolia pselliotes, Dicata odhnert y

Cuthona genovae. Dichas especies

pueden considerarse raras o de presen-

cia accidental en nuestras costas. No

hemos tenido en cuenta, en este caso, los

Cefalaspideos pues de las cuatro esta-

ciones en las que se intensificaron los

muestreos una sóla correspondía a

fondos blandos, por lo que el número de

inmersiones ha sido menor en este tipo

de fondos, que es donde preferente-

mente se encuentran las especies de

dicho grupo. Tampoco hemos tenido en

cuenta las especies procedentes de

arrastreros.

Es igualmente destacable que las tres

especies de Trapania: T. hispalensis, T.

lineata y T. maculata se han encontrado

durante los mismos meses del año, en

las mismas estaciones de muestreo y

sobre el mismo tipo de substrato. Este

hecho justificaría un estudio taxonómico

y ecológico de las especies del género en

el Mediterráneo.

La mayor abundancia de opistobran-

quios se ha observado en la zona com-

prendida entre 5 y 15 m de profundi-

dad, durante los meses de octubre,

noviembre, enero, febrero y marzo, que

corresponden con otoño e invierno.

En la estación de muestreo 11 es

donde aparecen un mayor número de

especies, dado que en ella se alcanza una

mayor profundidad, por lo que la varie-

dad de substratos y por tanto de habitats

diferentes es mayor. Sin embargo las

especies Tambja ceutae, T. marbellensis y

Polycera aurantiomarginati, se recolecta-

ron exclusivamente como juveniles en la

cercana estación 12 y debajo de piedras,

sobre el briozoo Sessibugula barrosoi, del

que hemos comprobado se alimentan, al

menos durante la etapa juvenil. Sólo el

ejemplar de T. ceutae de mayor tamaño,

unos 20 mm de longitud, fué encontrado

directamente sobre una pared a plomo,

en un substrato diferente, suponiendo

que en una fase juvenil más avanzada

dicha especie cambia de habitos alimen-

ticios. En general la mayor parte de las

especies fueron observadas sobre epi-

biontes de substratos rocosos y como

infralapidícolas.

Respecto a la distribución de las

especies que se han encontrado en la

costa granadina y en base a la Tabla III,

de las especies citadas, 53 son nuevas

citas para las costas de Granada. Poly-

cera faeroensis y Pruvotfolia pselliotes son

primeras citas para el Mediterráneo.

Geitodoris planata es nueva cita para el

Mediterráneo ibérico y Hermaea paucici-

rra lo es para el Mediterráneo andaluz.

Polycera faeroensis ha sido citada

antes de ahora en el litoral gallego por

ORTEA Y URGORRI (1981) y URGORRI Y

BESTEIRO (1983) y en el Estrecho de

Gibraltar por GARCÍA- GÓMEZ (1983;

1984).

Por su parte Pruvotfolia pselliotes ha

sido citada con anterioridad en el litoral

atlántico de la Península Ibérica desde

Galicia hasta el Estrecho de Gibraltar y

en las Islas Canarias (CERVERA, TEM-

PLADO, GARCÍA-GÓMEZ, BALLESTEROS,

ORTEA, GARCÍA, ROS Y LUQUE, 1988).

Debido a la confusión que ha habido

entre las especies Geitodoris planata y Dis-

codoris stellifera (CERVERA, (GARCÍA-

GÓMEZ Y GARCÍA., 1985; CERVERA ET AL.,

1988) resulta dificil establecer las áreas de

distribución de ambas, siendo necesario,

como señalan estos últimos la revisión de

las diferentes citas de las dos especies.

Aunque G. planata no ha sido citada en

las costas mediterráneas ibéricas, desde

hace una década, es relativamente fre-

cuente en diferentes localidades de la

costa catalana (Ballesteros, com. pers.).

Los ejemplares observados en las: costas

catalanas junto con los observados en la

Iberus, 18 (1), 2000

Tabla IT. Artículos donde se citan, en cada una de las provincias andaluzas, la presencia de las diferen-

tes especies observadas en la costa de Granada. Simbolos empleados: * especie citada por primera vez en

la provincia de Granada; - especie citada por primera vez en el litoral mediterráneo andaluz; = especie citada

por primera vez en el litoral mediterráneo ibérico, + especie citada por primera vez en el Mediterráneo.

Abreviaturas y números: C: Cervera; HU: Huelva; CA: Cádiz; MA: Málaga; GR: Granada; AL: Almería;

1: AARTSEN, MENKHORST Y GITTENBERGER (1984); 2: BALLESTEROS (1980); 3: BALLESTEROS,

BARRAJÓN, LUQUE, MORENO, TALAVERA Y TEMPLADO (1986); 4: BALLESTEROS Y TEMPLADO (1987);

5: BOBO (1998); 6: CERVERA (1988); 7: CERVERA Y GARCÍA-GÓMEZ (1986); 8: CERVERA Y GARCÍA-

GÓMEZ (1988); 9: CERVERA, GARCÍA-GÓMEZ Y GARCÍA (1985); 10: GARCÍA-GÓMEZ Y CERVERA

(1985); 11: CERVERA, GARCÍA-GÓMEZ Y GARCÍA (1986); 12: CERVERA, GARCÍA-GÓMEZ Y ORTEA

(1988); 13: CERVERA, LÓPEZ GONZÁLEZ Y GARCÍA-GÓMEZ (1998); 14: CERVERA, MEDINA Y GARCÍA

(1986); 15: CERVERA ET AL. (1988); 16: GARCÍA, GARCÍA-GÓMEZ Y CERVERA (1986); 17: GARCÍA-

GÓMEZ (1982); 18: GARCÍA-GÓMEZ (1983); 19: GARCÍA-GÓMEZ (1984); 20: GARCÍA-GÓMEZ (1985);

21: GARCÍA-GÓMEZ (1986); 22: GARCÍA-GÓMEZ (1987); 23: GARCÍA-GÓMEZ Y BOBO (1984); 24:

GARCÍA-GÓMEZ Y ORTEA (1988); 25: GARCÍA RASO, LUQUE, TEMPLADO, SALAS, HERGUETA, MORENO

Y CALVO (1992); 26: HERGUETA (1985); 27: HERGUETA Y SALAS (1987); 28: HIDALGO (1916); 29:

HIDALGO (1917); 30: LUQUE (1983); 31: LUQUE (1986); 32: GARCÍA-GÓMEZ Y GARCÍA (1984);

33: MARTÍNEZ ET AL. (1993); 34: MORENO Y TEMPLADO (1998); 35: NOBRE (1932); 40: RAMPAL

(1968); 41: Ros (1975); 42: SALAS Y HERGUETA (1986); 43: SALAS Y LUQUE (1986); 44: SCHICK Y

CERVERA (1998); 45: SIERRA, GARCÍA Y LLORIS (1978); 46: TEMPLADO, LUQUE Y MORENO (1988);

47: TEMPLADO ET AL. (1993); 49: VAYSSIERE (1913); 50: VIVES, SANTAMARÍA Y TREPART (1975).

Table III. Articles citing, for each Andalusian province, the presence of the different species observed on the

coast of the province of Granada. Symbols used: * species cited for the first time in-the province of Granada;

- species cited for the first time on the Mediterranean coast of Andalusia; = species cited for the first time on

the Mediterranean coast of the Iberian Peninsula; + species cited for the first time in the Mediterranean sea.

Abbreviations and numbers: C: Cervera; HU: Huelva; CA: Cádiz; MA: Málaga; GR: Granada; AL:

Almería; 1: AARTSEN, MENKHORST AND GITTENBERGER (1984); 2: BALLESTEROS (1980); 3:

BALLESTEROS, BARRAJÓN, LUQUE, MORENO, TALAVERA Y TEMPLADO (1986); 4: BALLESTEROS AND

TEMPLADO (1987); 5: BOBO (1998); 6: CERVERA (1988); 7: CERVERA AND GARCÍA-GÓMEZ (1986); 8:

CERVERA AND GARCÍA-GÓMEZ (1988); 9: CERVERA, GARCÍA-GÓMEZ AND GARCÍA (1985); 10: GARCÍA-

GÓMEZ AND CERVERA (1985); 11: CERVERA, GARCÍA-GÓMEZ AND GARCÍA (1986); 12: CERVERA,

GARCÍA-GÓMEZ AND ORTEA (1988); 13: CERVERA, LÓPEZ GONZÁLEZ AND GARCÍA-GÓMEZ (1998);

14: CERVERA, MEDINA AND GARCÍA (1986); 15: CERVERA ET AL. (1988); 16: GARCÍA, GARCÍA-GÓMEZ

AND CERVERA (1986); 17: GARCÍA-GÓMEZ (1982); 18: GARCÍA-GÓMEZ (1983); 19: GARCÍA- GÓMEZ

(1984); 20: GARCÍA-GÓMEZ (1985); 21: GARCÍA-GÓMEZ (1986); 22: GARCÍA-GÓMEZ (1987); 23:

GARCÍA-GÓMEZ AND BOBO (1984); 24: GARCÍA- GÓMEZ AND ORTEA ( 1988); 25: GARCÍA RASO,

LUQUE, TEMPLADO, SALAS, HERGUETA, MORENO Y CALVO (1992); 26: HERGUETA (1985); 27:

HERGUETA AND SALAS (1987); 28: HIDALGO (1916); 29: HIDALGO (1917); 30: LUQUE (1983); 31:

LUQUE (1986); 32: GARCÍA-GÓMEZ AND GARCÍA (1984); 33: MARTÍNEZ ET AL. (1993); 34: MORENO

AND TEMPLADO (1998); 35: NOBRE (1932); 40: RAMPAL (1968); 41: ROS (1975); 42: SALAS AND

HERGUETA (1986); 43: SALAS AND LUQUE (1986); 44: SCHICK AND CERVERA (1998); 45: SIERRA,

GARCÍA AND LLORIS (1978); 46: TEMPLADO, LUQUE AND MORENO (1988); 47: TEMPLADO ET AL.

(1993); 49: VAYSSIERE (1913); 50: VIVES, SANTAMARÍA AND TREPART (1975).

ESPECIE HU CA MA GR AL 1* Cito

Bulla striata 29,17 29,17,1 29,30,31 3,45 ES

Aglaja tricolorata 33 33 34 34

Philinopsis depicta 4] 34 E

Scaphander lignarius 29 29,1,47 29 y

Cymbulia peronii 40,50 25 S

Aplysia fasciata C(obs. pers.) 17 30,31 3,25 z

Aplysia parvula 47 4 25 5

Aplysia punctata 7 17,47 30,31,3,43 3,25 z

Umbraculum umbraculum 29 29 29,30,31 29 E

SÁNCHEZ-TOCINO E7 AL.: Opistobranquios de la costa de Granada

Tabla MI. Continuación.

Table IT. Continuation

ESPECIE

Pleurobranchus testudinarius

Berthella ocellata

Berthella plumula

Berthella stellata

Pleurobranchaea meckelii

Thuridilla hopei

Elysia viridis

Hermaea paucicirra

Coliphylla mediterranea

Placida verticillata

Aegires leuckarti

Trapania hispalensis

Trapania lineata

Trapania maculata

Diaphorodoris luteocincta

Diaphorodoris popillata

Crimora papillata

Aldisa banyulensis

Chromodoris britoi

Chromodoris krohni

Chromodoris luteorosea

Chromodoris purpurea

Cadlina pellucida

Hypselodoris bilineata

Hypselodoris cantabrica

Hypselodoris fontandravi

Hypselodoris picta

Hypselodoris midatlantica

Hypselodoris vilafranca

Discodoris atromaculata

Discodoris fragilis

Discodoris rosi

Geitodoris portmanni

Geitodoris planata

Jorunna onubensis

Platydoris argo

Rostanga rubra

Polycera aurantiomarginata

Polycera faeroensis

Polycera quadrilineata

Limacia clavigera

Roboastra europea

Tambja ceutae

Tambja marbellensis

Dendrodoris limbata

Doriopsilla areolata

Doto koenneckeri

Hancockia uncinata

Tethys fimbria

Tritonia manicata

Tritonia nilsodhneri

Tritonia striata

Mariona blainvillea

HU CA MA

22,47

14 7

14 Neza

8,5 4] 30,31,47

17,19,7 30,31

6 17,19 3

1,12

30,31

19,22

19 19,8

18,19 30,31

5 18,19

17,19,16

5 22

17,19,7 —— 26,31,42

5 18,19,7

5 18,19,7 30,31

5 18,19,7

5 18,19 30,31

18,19 30,31

5 17,18,19 — 41,30,31

18,19

5 19,7 30,31

18,19

18,19 31

11,5 7

41,18,19,47 30,31

25,19,5 /

5 18,19

17,19,5 19 30,31

18,19

19,20

17,19 17,19,7 30,31

19,5 17,19,7,47 30,31,47,42

47 30,31,47

18,19,7 30,31

18,19

49,28,35 17,19 47

47,34

30,31

Iberus, 18 (1), 2000

Tabla HI. Continuación.

Table HT. Continuation

ESPECIE HU CA MA GR AL 12 Cita

Armina moculata 17 30,31 :

Janolus cristatus 19 18,19,7 30,31 46,25 E

Aeolidiella soemmeringi 19 17,197 E

Berghia coerulescens 18,19 26,42,27 34

Berghia verrucicornis 19 18,19,7 3,25 E

Spurilla neapolitana 19,10,5 18,19,10,7 30,31 3,25 Ñ

Eubranchus farrani 7 46 46,25

Facelina annulicornis 18 46

Facelina coronata 18,7 46 25

Facelina rubrovitatta 7 30,31 3 S

Coloria elegans 18,19 3,25 ES

Cratena peregrina 19 11977 3,25 hi

Pruvotfolia pselliotes 19,7,22 +

Dicata odhneri 19 34 ES

Dondice banyulensis 17,32,47 30,31 46,25

Flabellina affinis 19 17,18,19,21,7 30,31 30,31 3,25

Flabellina ischitana 5 13 25 %

Flabellina babai 19,21 31 25 a

Flabellina pedata 19 17,19,7 30,31 46 3

Calmella cavolini 25 Ey

Piseinotecus gabinieri 46

Cuthona genovae 18,19,7 3,25 E

costa granadina constituyen la primera

cita para el Mediterráneo.

Hermaea paucicirra ha sido encon-

trada con anterioridad en el litoral astu-

riano (ORTEA, 1977a, b), en el litoral

gallego (URGORRI Y BESTEIRO, 1983), en

las costas de Alicante y Murcia (MARÍN

Y Ros, 1988), en el litoral catalan

(BALLESTEROS, 1980) así como, en las

costas de Cádiz (CERVERA Y GARCÍA-

GÓMEz, 1986; CERVERA, GARCÍA-GÓMEZ

Y ORTEA, 1988). Por tanto el hallazgo de

esta especie en la costa de Granada

supone su primera cita para el litoral

mediterráneo andaluz.

Resulta de interes el haber obser-

vado en una franja costera tan reducida

los dos tipos de coloraciones que se dan

en el Mediterráneo español, para Hypse-

lodoris villafranca, según ORTEA, VALDÉS

Y GARCÍA-GÓMEZ (1996): la que se da en

las costas catalanas y la propia del sur

de la Península, siendo más frecuente

ésta última.

Por otro lado, el género Tambja es el

que ha proporcionado los resultados

taxonómicos más interesantes, al encon-

trarse en la costa granadina todas las

especies del género citadas por ahora en

las costas europeas. Tambja marbellensis se

(Página derecha) Figura 2. A: Geitodoris planata, ejemplar de 25 mm; B: Pruvotfolia pselliotes, ejem-

plar de 15 mm; C: Polycera faeroensis, ejemplar de 25 mm; D: Hermaea paucicirra, ejemplar de 2

mm; E: Caliphylla mediterranea, ejemplar de 5 mm; E: Tambja marbellensis, ejemplar de 12 mm;

G: Polycera aurantiomarginata, ejemplar de 3 mm: H: Placida verticillata, ejemplar de 2 mm.

(Right page) Figure 2. A: Geitodoris planata, specimen of 25 mm; B: Pruvotfolia pselliotes, specime of

15 mm; C: Polycera faeroensis, specimen of 25 mm; D: Hermaea paucicirra, specimen of 2 mm: E:

Caliphylla mediterranea, specimen of 5 mm: F: Tambja marbellensis, specimen of 12 mm; G: Poly-

cera aurantiomarginata, specimen of 3 mm; H: Placida verticillata, specimen of 2 mm.

SÁNCHEZ-TOCINO ET 4L.: Opistobranquios de la costa de Granada

Iberus, 18 (1), 2000

cita por primera vez tras su descripción

original por SCHICK Y CERVERA (1998)

Respecto a las veces que han sido

citadas las especies contempladas en el

presente trabajo, en las distintas provin-

cias del litoral andaluz (ver Tabla III)

resulta resaltable, en primer lugar, que

éstas no han sido homogéneas, lo que

está motivado por la peculiaridad de la

orografía del litoral de cada una de ellas

y por la profusión de los muestreos, más

intensa en unas que en otras. En cual-

quier caso es destacable que en las pro-

vincias andaluzas más estudiadas

(Cádiz y Almería) determinadas espe-

cies son repetidamente citadas por más

de un autor. Así, en la costa gaditana,

Bulla striata, Scaphander lignarius, Berthe-

lla stellata, Thuridilla hopei, Aldisa banyu-

lensis, Chromodoris krohni, C. luteorosea, C.

purpurea, Hypselodoris bilineata, H. picta,

Platidoris argo, Rostanga rubra, Dendrodo-

ris limbata, Doriopsilla areolata, Tritonia

manicata, Janolus cristatus, Aeolidiella

soemmeringi, Bergia verrucicornis, Spurilla

neapolitana, Pruvotfolia pselliotes, Dondice

banyulensis, Flabellina affinis, Flabellina

pedata, y Cuthona genovae, son las espe-

cies más frecuentes. De la misma forma,

en la costa almeriense, Aplysia fasciata,

Aplysia punctata, Berthella plumula, B. ste-

llata, Thuridilla hopei, Trapania lineata, T.

maculata, Diaphorodoris luteocincta, D.

papillata, Chromodoris britoi, C. krohni, C.

BIBLIOGRAFÍA

AARTSEN, J. J., MENKHORST, H. P. M. G. Y GIT-

TENBERGER, E., 1984. The marine mollusca of

the Bay of Algeciras, Spain, with general no-

tes on Mitrella, Marginellidae and Turridae.

Basteria, Suppl. 2: 1-135.

BALLESTEROS, M., 1980. La presencia en las cos-

tas catalanas de Hermaea paucicirra y Hermaea

cremoniana (Opisthobranchia: Sacoglossa).

Publicaciones del Departamento de Zoología de

Barcelona, 5: 19-23.

BALLESTEROS, M., BARRAJÓN, A., LUQUE, A. A.,

MORENO, D., TALAVERA, P. Y TEMPLADO, J.,

1986. Contribución al conocimiento de los gas-

terópodos marinos de Almería. Iberus, 6: 39-55.

BALLESTEROS, M. Y TEMPLADO, J., 1987. Aplysia

parvula Guilding in Morch, 1863 en las costas

de la Península Ibérica. Publicaciones del

Departamento de Zoología de Barcelona, 13: 55-62.

purpurea, Platydoris argo, Rostanga rubra,

Polycera quadrilineata, Dendrodoris

limbata, Doto koenneckeri, Janolus crista-

tus, Berghia verrucicornis, Spurilla neapoli-

tana, Eubranchus farrani, Caloria elegans,

Cratena peregrina, Dondice banyulensis,

Flabellina affinis, Calmella cavolini y

Cuthona genovae, presentan una mayor

frecuencia. Teniendo en cuenta los datos

de todas las provincias, las especies:

Bulla striata, Aglaja tricolorata, Aplysia fas-

ciata, A. punctata, Umbraculum umbracu-

lum, Pleurobranchaea meckelii, Elysia

viridis, Chromodoris purpurea, Hypselodo-

ris picta, H. villafranca, Polycera quadrili-

neata, Dendrodoris limbata, Doriopsilla are-

olata, Mariona blainvillea, Janolus cristatus,

Spurilla neapolitana, Flabellina affinis y E.

pedata han sido citadas en todo el litoral

andaluz.

AGRADECIMIENTOS

Nuestro más sincero agradecimiento

al Dr. José Templado por su ayuda en la

identificación de algunos ejemplares, así

como a Angel Gaytan y Antonio de la

Linde por acompañarnos en muchas de

las inmersiones. Este trabajo queda englo-

bado dentro del Proyecto “Fauna Ibérica

IV” (DGICYT PB95-0235), subproyecto de

Moluscos Gasterópodos Opistobranquios,

y en el proyecto CICYT PB98-1121.

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O Sociedad Española de Malacología

Iberus, 18 *1): 15-19, 2000

Onoba oliverioi n. sp. (Prosobranchia, Rissoidae), a new

gastropod from the Mediterranean

Onoba oliverioi n. sp. (Prosobranchia, Rissoidae), un nuevo gasteró-

podo para el Mediterráneo

Carlo SMRIGLIO* and Paolo MARIO TTINI**

Recibido el 1-IX-1999. Aceptado el 1-X-1999

ABSTRACT

Onoba oliverioi n. sp. (Prosobranchia, Rissoidae) is here described from material collected

on muddy bathyal bottoms surrounding a deep-sea coral bank located off-shore Latium (Cen-

tral Tyrrhenian Sea). The new species is known from shells only and it has been placed in the

genus Onoba H. and A. Adams, 1852. Onoba oliverioi n. sp. is compared to Onoba gianni-

nii [Nordsieck, 1974) (deep-shelf), Onoba dimassai Amati and Nofroni, 1991 (shallow

water) and Alvania wareni [lemplado and Rolan, 1986) (deep-shelf), which it resembles. A

list of mollusc species found associated with the new taxon ¡is also given.

RESUMEN

En el presente trabajo se describe, en base exclusivamente a caracteres de la concha, una

nueva especie de Onoba, a partir de material recogido en un fondo batial de arena y

fango, en las proximidades de una comunidad de corales blancos, localizada a lo largo de

las costas del Lazio. El nuevo taxon, denominado Onoba oliverioi n. sp., se compara con

las especies similares del género Onoba, O. gianninii y O. dimassai Amati y Nofroni,

1991, y Alvania wareni (Templado y Rolan, 1986). Se incluye la lista de las especies que

se han encontrado en el mismo muestreo.

KEY WORDS: Onoba, new species, Mediterranean Sea.

PALABRAS CLAVE: Onoba, nueva especie, Mediterráneo.

INTRODUCTION

The genus Onoba H. and A. Adams,

1852 includes small size cylindrical

species which share certain shell featu-

res with members of Alvania Risso, 1826

s. l. and Crisilla Monterosato, 1917 s. 1.,

regarded as a genus, according to

BOUCHET AND WARÉN (1993). Anatomi-

cal differences have been found between

* Via di Valle Aurelia 134, 00167 Roma, Italy.

some representative taxa of the two

genera Onoba and Rissoa, leading to the

idea that is an interesting case of shell

morphological convergence (PONDER,

1985; OLIVERIO, 1988; BOUCHET AND

WARÉN, 1993). The Mediterranean

species of Onoba, well illustrated by

GIANNUZZI- SAVELLI, PUSATERI, PALMERI

** Dipartimento di Biologia, Universita degli Studi Roma Tre, Viale Marconi 446, 00146 Roma, Italy.

Corresponding author: Paolo Mariottini, Dipartimento di Biologia, Universita degli Studi Roma Tre, Viale

Marconi 446, 00146 Roma, Italy. E-mail: mariotpaCbio. uniroma3. it

Iberus, 18 (1), 2000

AND EBREO (1996), consist of mainly

shallow-water shells. In fact, the only

deep-water taxon is Onoba gianninii

(Nordsieck F., 1974), figured by SEM

photographs by AMATI AND NOFRONI

(1991), who also designate the lectotype,

and BOUCHET AND WARÉN (1993). In this

paper we describe, on the base of shell

morphology, a new species of Onoba

from material dredged on some muddy

bathyal bottoms surrounding a deep-sea

coral bank located off Latium coast and

currently under investigation (SMRIGLIO

AND MARIOTTINI, 1999). The new species

RESULTS

is compared to the similar O. gianninii

and Onoba dimassai Amati and Nofroni,

1991. A list of mollusc species found in

the same material is also given.

Abbreviations:

MZB: Museo di Zoologia dell Universita

di Bologna, Italy.

MZR: Reparto Malacologico del Museo

Civico di Zoologia di Roma, Italy.

CS: C. Smriglio private collection.

MO: M. Oliverio private collection.

PM: P. Mariottini private collection.

Family RISSOIDAE Gray J. E., 1847

Genus Onoba H. and A. Adams, 1852

Onoba oliverioi n. sp. (Figs. 1-6)

Type material: Holotype (MZB 14000), 1 sh., dredged, june 1987. Paratype D (MZR), 1 sh., type

locality, june 1987. Paratypes A, B, C, E, F, G, H, Iand L (CS), 9 sh., type locality, june 1987. Paratype

M (MO) 1 sh., type locality, may 1990. Paratype N (PM), 1 sh., type locality, may 1990.

Type locality: Central Tyrrhenian Sea (41? 51 N, 11? 28' E), off coast of Latium, on muddy bottom

in a deep-sea coral biocoenosis, [biocoenosis VB and CB sensu PÉRES AND PICARD (1964) ], at a depth

of 350-600 m.

Material examined: O. oliverioi n. sp., type locality, 32 sh. (CS).

Derivatio nominis: This species is named in honour of Dr Marco Oliveiro, expert malacologist and

good friend of the authors, who has contributed a great deal to the knowledge of the Mediterra-

nean malacofauna.

Description: Shell small (from 1.61 to

2.32 mm in height), conical-ovate, with a

large aperture, blunt apex. Protoconch

dome-shaped consisting of about 1.5

whorls, with a diameter of 400-440 um,

sculptured with 6-8 fine and irregular spi-

ral cordlets. Among them, several other

interrupted fine furrows create a sort of mi-

cro-tuberculated sculpture. Teleoconch of

about 3.0 rounded convex whorls, the last

one is about ?/3 of the entire length, ave-

rage ratio H/W=1.55, average ratio H/ Ha

= 1.99. Suture pronounced and shallowly

channelled, axial growing lines evident,

spiral sculpture consisting of about 27

evenly spaced ribs, with about 2-3 much

smaller furrows in the inter-spaces. Aper-

ture ovoid, umbilical crevice slightly visi-

ble. Colour milky-white or yellowish trans-

lucent. Operculum and animal unknown.

Remarks: O. oliverioi (Figs. 1-6) is con-

chologically very similar to O. gianninil

(Figs. 7, 8), resulting in the new taxon ha-

ving been already recorded in the past by

BOUCHET AND WARÉN (1993: fig. 1520, p.

663), but misidentified as O. gianninti. M.

Oliverio already mentioned the possibi-

lity to face a complex of species when de-

aling with O. gianninii: “Studying the spe-

cimens usually classified as gianninit it re-

sulted that more than one species could be

involved” OLIVERIO (1988). Indeed, O. oli-

verioi can be separated from the last spe-

cies on the ground of some protoconch

and teleoconch features. In particular, the

protoconchs differ in the two species, being

the one of O. oliverioi much flatter, more

sculptured and slightly smaller. The tele-

oconch outline of O. oliverioi differs from

the one of O. giannini being less slender

SMRIGLIO AND MARIOTTINI: Onoba oliverioí n. sp., a new Mediterranean Rissoidae

Figures 1-6. Onoba oliverioz. 1: holotype. 2.0 x 1.3 mm, (MZB 14000); 2: paratype D, 1.8 x 1.2

mm, (MZR); 3: paratype C, 1.8 x 1.2 mm, (CS); 4: paratype E, 2.2 x 1.3 mm, (CS); 5: paratype

A; 6: protoconch, paratype A, 2.2 x 1.4 mm, (CS). Figures 7, 8. O. gianninii. 7: lectotype; 8: pro-

toconch,lectotype, 2.2 x 1.2 mm, (MZR). Strait of Bonifacio, off Capo Comino (200-220 m).

Lectotype figure after AMATI AND NOFRONI (1991). Scale bars, shells: 1 mm; protoconchs: 200

mm.

Figuras 1-6. Onoba oliverioi. 1: holotipo. 2,0 x 1,3 mm, (MZB 14000); 2: paratipo D, 1,8 x 1,2

mm, (MZR); 3: paratipo C, 1,8 x 1,2 mm, (CS); 4: paratipo E, 2,2 x 1,3 mm, (CS); 5: paratipo A; 6:

protoconcha, paratipo A, 2,2 x 1,4 mm, (CS). Figuras 7, 8. O. gianninii. 7: lectotipo; 8: protoconcha

del lectotipo, 2,2 x 1,2 mm, (MZR), Estrecho de Bonifacio, Capo Comino (200-220 m). Lectotipo

figura original de AMATI Y NOFRONI (1991). Escalas, conchas: 1 mm; protoconchas: 200 mm.

Iberus, 18 (1), 2000

(H/W = 1.55 versus H/W = 1.75, respec-

tively), having a smaller average number

of whorls (about 2.5-3 versus 3.5-4, res-

pectively) and a bigger aperture (H/ Ha =

1.99 versus H / Ha = 2.40, respectively). On

the contrary, the sculpture of the teleo-

conch in the two species results very si-

milar, consisting in rounded spiral ribs of

equal breadth and number (26-31). O. oli-

verioi also differs from Alvania wareni (Tem-

plado and Rolan, 1986), a species closely

related to O. gianninil, as stressed by BOU-

CHET AND WARÉN (1993: 663): “They may

prove to be conspecific, when material beco-

mes known from the intermediate area”. The

differences outlined between the new spe-

cies and O. gianninii, hold when compa-

ring the new species to A. wareni as well.

Furthermore, the shell of A. wareni has an

additional and diagnostic sculpture fea-

ture at the protoconch-teleoconch demar-

DISCUSSION

The shell features observed in the

specimens of O. oliverioi from the Central

Tyrrhenian Sea, summarised in Table L,

are clearly visible in the individual figu-

red by BOUCHET AND WARÉN (1993),

which is from the Strait of Bonifacio (off

Capo Comino, 200-300 m). So, it seems

that the morphological characters shown

by O. oliverioi are very constant regard-

less the collecting spot. Since shells of

both O. gianninii and O. oliverioi have

been collected together in that area (Bou-

CHET AND WARÉN, 1993), the two species

could be sympatric in the Strait of Bonifa-

cio, corroborating the idea that are in-

deed two different taxa and not just two

extreme forms of the same species. The

shells of O. oliverioi from the Central Tyrr-

henian Sea were dredged on a bathyal

bottom, at a depth of 350-600 m, resulting

in a deeper record than the ones of the

specimen from Capo Comino and of O.

gianninti (AMATI AND NOFRONI, 1991).

The new taxon probably belongs to the

biocoenosis VB sensu PÉRES AND PICARD

(1964), the other mollusc species found in

the same dredged material seem to sup-

port this assumption. The identified spe-

cies occurring with O. oliverioi are: Propi-

cation (BOUCHET AND WARÉN, 1993: 646;

fig. 1462); in fact, the last part of the pro-

toconch whorl shows about 20 fine cord-

lets irregularly interrupted, which are not

present in O. olíverioi nor in O. gianninil.

The new taxon is distinguishable from O.

dimassai too, a shallow water species which

has a similar shell shape. O. dimassai has

a smooth protoconch with no evident mi-

crosculpture, of about 1.20-1.25 whorls

and a diameter of about 300-380 um. Its

teleoconch sculpture shows 24-30 major

spiral ribs which present 4-5 fine furrows,

together with the 3-4 furrows at the inter-

spaces, the shell surface appears to be more

tightly covered by these spiral ribs; furt-

hermore, the last whorl does not show any

umbilical crevice (AMATI AND NOFRONI,

1991). These shell features pointed out

allow an easy separation of O. dimassai

from O. oliveriol.

lidium exiguum Thompson, 1843, Lepetella

cf. laterocompressa (De Rayneval and

Ponzi, 1854), Emarginula tenera Locard,

1892, Clelandella miliaris (Brocchi, 1814),

Danilia otaviana (Cantraine, 1835), Putzey-

sia wiseri (Calcara, 1842), Alvanta cimicoi-

des (Forbes, 1844), Alvania subsoluta (Ara-

das, 1847), Orbitestella dariae (Liuzzi and

Stolfa Zucchi, 1979), Trophon muricatus

var. barvicensis (Johnston, 1825), Nassarius

lima (Dillwin, 1817), Amphissa acuteo-

costata (Philippi, 1844), Granulina gofasi

Smriglio and Mariottini, 1996, Gymnobela

abyssorum (Locard, 1897), Microdrillia lo-

prestiana (Calcara, 1841), Pleurotomella de-

mosia (Dautzenberg and Fischer P., 1896),

Pleurotomella gibbera Bouchet and Warén,

1980 ex Jeffreys ms., Teretia teres (Reeve,

1844), Conopleura aliena Smriglio, Mariot-

tini and Calascibetta, 1999, Heliacus alleryi

(Seguenza G., 1876), Mathilda cochlaefor-

mis Brugnone, 1873, Japonacteon pusillus

(McGillivray, 1843), Asperarca nodulosa

(Múller, 1776), Chlamys bruei (Payrau-

deau, 1826) and Cadulus subfusiformis

(Sars M., 1865). At the present time, the

distribution of O. oliverioi is limited to the

Tyrrhenian Sea: off Latium and Sardinia

(Strait of Bonifacio) coasts.

SMRIGLIO AND MARIOTTINI: Onoba oliverioi n. sp., a new Mediterranean Rissoidae

Table 1. Shell morphological measurements of Onoba oliverioi type material. A- »reviations. H:

height in mm; W: width in mm; Ha: height aperture in mm; Pd: protoconch diame:er in um; Sr:

spiral ribs of the last whorl; ND: not determined.

Tabla 1. Medidas morfológicas de la concha del material tipo de Onoba oliverioi. Abreviaciones. H:

altura en mm; W. ancho en mm; Ha: altura de la boca en mm; Pd: diámetro de la protoconcha en um;

Sr: estrias espirales en la ultima vuelta; ND: no determinado.

Specimen H wW

Holotype 1.85 1.20

Paratype Á 27 1.38

Paratype B 2.26 1.47

Paratype € 2.05 1.26

Paratype D 1.70 17

Paratype E DD 1635

Paratype F 1591 129

Paratype G 179 1.20

Paratype H 179 1.20

Paratype | ZW 1.41

Paratype L 178 1.20

Paratype M 1.61 1.08

Paratype N DSZ 1.47

ACKNOWLEDGEMENTS

We would like to express our grati-

tude to Dr Marco Oliverio (Dipartimento

di Biologia, Universita di Roma “La Sa-

pienza”, Italy) for providing SEM photo-

graphs and for critical comments and ad-

vice on the present paper. Sincere thanks

are due to Dr Antonio Bonfitto (Museo di

Zoologia dell'Universita di Bologna,

Italy) and to Dr Vincenzo Vomero (Mu-

BIBLIOOGRAPHY

AMATI, B. AND NOFRONL I., 1991. Designazione

del lectotipo di “Setia” gianninii F. Nordiesck,

1974 e descrizione di Onoba dimassai nuova

specie (Prosobranchia: Rissoidae). Notiziario

C.I.S.M.A., 13: 30-37.

BOUCHET, P. AND WARÉN, A., 1993. Revision of

the Northeast Atlantic Bathyal and Abyssal

Mesogastropoda. Bollettino Malacologico,

suppl. 3: 579-849.

GIANNUZZI-SAVELLI, R., PUSATERI, F., PALMERI,

A. AND EBREO, C., 1996. Atlante delle Conchi-

glie Marine del Mediterraneo. Vol. 2. “La Con-

chiglia”, Roma, 258 pp.

OLIVERIO, M., 1988. On the systematics of “Se-

tia” gianninii (Gastropoda: Prosobranchia). Bo-

llettino Malacologico, 24 (5-8): 112-114.

0.97

1.08

0.94

0.88

1.05

1.00

0.94

1.11

0.94

0.85

H/W H/Ha Pd Sr

1.54 1.91 440 26

1.57 2.00 400 31

1.53 2.09 420 27

1.62 2.18 440 28

1.58 2.03 400 28

1.65 2.12 440 26

1.48 1.91 440 26

1.49 1.90 440 26

1.49 1.90 A40 28

1.54 1.95 400 ND

1.44 1.84 420 26

1.49 1.89 400 23

1.57 2.09 400 29

seo Civico di Zoologia di Roma, Italy) for

the examination of material kept in MZB

and MZR collections related to O. olíve-

rioi. We are really grateful to the revie-

wers Dr Anders Warén (Department of

Invertebrate Zoology, Swedish Museum

of Natural History, Stockholm, Sweden)

and Dr Italo Nofroni (University of Rome

“La Sapienza”, Rome, Italy) for their cri-

tical comments and suggestions in im-

proving the manuscript.

PÉRES, J. M. AND PICARD, J., 1964. Nouveau Ma-

nuel de Bionomie Benthique de la Mer Mé-

diterranée. Recuil des Travaux de la Station

Marine d'Endoume, 31 (47): 1-137.

PONDER, W. F., 1985. A review of the genera of

the Rissoidae (Mollusca: Mesogastropoda:

Rissoacea). Records of the Australian Museum,

suppl. 4: 1-221.

SMRIGLIO, C. AND MARIOTTINI, P., 1999. Mo-

lluschi del Mar Tirreno Centrale. Contributo

XII. Segnalazione di due rari Epitoniidae ba-

tiali per le coste laziali (Gastropoda, Pteno-

glossa). Bollettino Malacologico, 39 (9-12): 137-

140.

O Sociedad Española de Malacología — —__———— Iberus, 18 '1): 21-94, 2000

The subfamily Rissoininae (Mollusca: Gastropoda: Rissoi-

dae) in the Cape Verde Archipelago (West Africa)

La subfamilia Rissoininae (Mollusca: Gastropoda: Rissoidae) en el

archipiélago de Cabo Verde (África Occidental)

Emilio ROLÁN* and Ángel A. LUQUE**

Recibido el 29-1X-1999. Aceptado el 26-X-1999

ABSTRACT

The subfamily Rissoininae (Gastropoda, Rissoidae) is studied in the Cape Verde Archipel-

ago. Twenty-nine species belonging to three genera have been found: Rissoina [Rissoina)

punctostriata (Talavera, 1975), a species with planktotrophic development widely distrib-

uted along West Africa, and currently confused with the also planktotrophic Caribbean

species Rissoina (Rissoina) decussata (Montagu, 1803); one new species of Rissoina

(Ailinzebina); 26 new species of Schwartziella [(Schwartziella), and one new species of

Zebina (Zebina). The paucispiral protoconch of all the new species indicates a non-plank-

totrophic development and strongly suggest all them are endemic of the Cape Verde Archi-

pelago. The new species are compared among them and with other related West African

species. The high level of endemism, the geographical distribution of all the species in the

Archipelago and the possible relationships of the Cape Verde Rissoininae are also com-

mented.

RESUMEN

Se estudia la subfamilia Rissoininae en el Archipiélago de Cabo Verde. Se han encon-

trado 29 especies pertenecientes a tres géneros: Rissoina (Rissoina) punctostriata [Tala-

vera, 1975), una especie con desarrollo larvario planctotrófico ampliamente distribuida a

lo largo de la costa occidental africana y hasta ahora confundida con la especie caribeña

Rissoina (Rissoina) decussata [Montagu, 1803), también de desarrollo planctotrófico; una

nueva especie de Rissoina (Ailinzebina); 26 especies nuevas de Schwartziella [Schwart-

ziella), y una especie nueva de Zebina (Zebina). La protoconcha paucispiral de todas las

especies nuevas indica un desarrollo larvario no planctotrófico y sugiere que todas ellas

son endémicas del archipiélago. Se comparan las nuevas especies descritas con otras del

África occidental con las que están relacionadas, y entre sí. Se comentan el alto grado de

endemismo, la distribución geográfica de todas las especies en el archipiélago y las posi-

bles relaciones de los Rissoininae de Cabo Verde.

KEY WORDS: Rissoininae, Rissoina, Zebina, Schwartziella, new species, Cape Verde Islands, West Africa.

PALABRAS CLAVE: Rissoininae, Rissoina, Zebina, Schwartziella, nuevas especies, Islas de Cabo Verde, África

Occidental.

* Cánovas del Castillo, 22, 36202 Vigo, Spain.

** Laboratorio de Biología Marina, Departamento de Biología, Universidad Autónoma de Madrid, 28049

Madrid, Spain.

Iberus, 18 (1), 2000

INTRODUCTION

Though many papers dealing with

the marine gastropods of the Cape

Verde Archipelago have been

published during the last years (see

BURNAY AND COSEL, 1987; FERNANDES

AND ROLÁN, 1991, and ROLÁN AND

RUBIO, 1999 for a list), several groups

are still awaiting study. One of them is

the subfamily Rissoininae (Rissoidae),

revised at the generic level by PONDER

(1985) and studied in other parts of

the world (LEAL AND MOORE, 1989;

SUAURS, 10, 1, ISE) LL SO

FABER, 1990; SLEURS AND PREECE, 1994;

ROLÁN, 1998). However, there are only

a few papers dealing with Rissoininae

from West Africa and neighbouring

islands. WATSON (1873) described

Eulima paivensis from the Selvagens

Islands, which turned out to be the

common species recorded from the

Canary Islands as Zebina browniana

(d'Orbigny, 1842) or Z. vitrea (C. B.

Adams, 1850) by ODHNER (1932),

NORDSIECK (1972) and GARCÍA-TALA-

VERA (1983) (see GOFAS, 1999). SMITH

(1890) described five new species from

St. Helena and erroneously recorded

Rissoina bryerea (Montagu, 1803), a

Caribbean species. [DAUTZENBERG

(1913) and TALAVERA (1975), described

two new species: Rissoina africana from

Senegal, and Zebina punctostriata from

Mauritania, respectively. GOFAs (1999)

studied Rissoina punctostriata, Schwart-

ziella africana and described a new

species of Zebina. ROLÁN AND RYALL

(1999) recorded Rissoina punctostriata

(Talavera, 1975) from Angola. A few

probably erroneous records are sparse

in different papers: Rissoina elegantula

(Angas, 1880) from Sáo Tomé (TOMLIN

AND SHACKLEFORD, 1914), Rissoina calia

Bartsch, 1915, from Senegal (NICKLES,

1947), Zebina vitrea (A. Adams, 1854)

from Sahara, Mauritania (ALTIMIRA,

1978) and the Canary Islands (ALri-

MIRA, 1978; NORDSIECK, 1982).

FERNANDES AND ROLÁN (1994) rec-

orded six amphiatlantic species of Ris-

soininae which actually have not been

accepted.

The first record of a Rissoininae

from the Cape Verde Islands is that of

DAUTZENBERG AND FISCHER (1906) (Ris-

soina decussata (Montagu, 1803), see

below under remarks of R. punc-

tostriata). MARCHE-MARCHAD (1958)

and SAUNDERS (1977) recorded Rissoina

africana (Dautzenberg, 1913). GARCÍA-

TALAVERA AND PBACALLADO (1978)

recorded Rissoina bryerea (Montagu,

1803), but this is a misidentification of a

Caribbean species. COSEL (1982a, b, c)

recorded R. decussata (Montagu, 1803),

R. africana Dautzenberg, 1913, Zebina cf.

punctostriata (Talavera, 1975) and other

2-3 probably undescribed species of Ris-

soina.

A preliminary revision of the subfa-

mily Rissoininae from the Cape Verde

Archipelago was presented by MORÁN,

ROLÁN AND LUQUE (1989) to the 5th

Symposium Fauna and Flora of the

Cape Verde Islands (Leiden), and an

updated checklist of the marine gastro-

pods by ROLÁN, FERNANDES, LUQUE,

ORTEA AND TEMPLADO (1993) to the First

Symposium Fauna and Flora of the

Atlantic Islands (Madeira). In both ab-

stracts was referred the existence of

seven or more undescribed species of

Rissoininae. During recent years, a tho-

rough revision of the material from dif-

ferent expeditions to the Cape Verde

Islands has shown that the number of

Rissoininae species in this archipelago

has been greatly underestimated. A total

of 29 species of three genera, 28 of

which are new for the science, are de-

scribed in the present paper.

MATERIAL AND METHODS

About 2300 shells and specimens

have been studied from almost all the

Cape Verde Archipelago (Fig. 157). A

part of this material was collected by the

“T Expedición Científica Ibérica al Archi-

piélago de Cabo Verde” (1985), as well

as in several trips of Spanish and Portu-

guese malacologists between 1978 and

1988, most of them with the partic-

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

ipation of the first author. A small part

of the material was collected by dredg-

ing down to 100 m of depth, or by

SCUBA diving down to 30 m, but most

of the material was collected by skin

diving down to 15 m. Additional mate-

rial collected by the CANCAP Expedi-

tions of the National Museum of

Natural History of Leiden has been

included in the present study, so as

some new material collected in 1997 by

the first author during the expedition

“Macaronesia 2” of Las Palmas Univer-

sity. Some types housed in The Natural

History Museum of London and the

Muséum National d'Histoire Naturelle

of Paris have been also studied.

Specimens are illustrated using SEM

micrographs; the views of microsculp-

ture were made at the middle part of the

body whorl.

Abbreviations:

AMNH: American Museum of Natural

History, New York.

BMNH: The Natural History Museum,

London.

DBUA: Departamento de Biología, Uni-

versidad Autónoma, Madrid.

MNCN: Museo Nacional de Ciencias

Naturales, Madrid.

MNHN: Muséum National d'Histoire

Naturelle, Paris.

NNM: Nationaal Natuurhistorisch Mus-

eum, Leiden.

CER: Collection of E. Rolán, Vigo.

The material with no indication of

collection is from CER.

sp: live collected specimen.

s: empty shell.

j: juvenile shell.

f: fragment of shell.

RESULTS

Family RISSOIDAE J. E. Gray, 1847

Subfamily RISSOININAE Stimpson, 1865

Genus Rissoina d'Orbigny, 1840

Subgenus Rissoina s. s.

Type species: Rissoina inca d'Orbigny, 1840, by original designation.

Diagnosis: PONDER (1985, p. 78) and SLEURS (1993, p. 74).

Rissoina (Rissoina) punctostriata (Talavera, 1975) (Figs. 1-3, 6-8)

Zebina punctostriata Talavera, 1975. Bol. Inst. Esp. Oceanog., 192: 3, pl. 1, fig. 1, pl. 4, fig. 7. [Type

locality: SAHMAS-1, st. EO-8, Mauritania].

Material studied: Cape Verde Archipelago: Sal: 2 s, Palmeira; 2 sp, 8 s, Regona,1-3 m;, 1 sp, 5 s,

Rabo de Junco, 3 m; 1 s, 3 f, Mordeira, 5 m; 3 f, off Palmeira, CANCAP Sta. 7.109, 16* 45' N, 22*

59 W, 31 m (31-VIIT-86) (NN M); 2 j, 1 f, Santa Maria Bay, CANCAP Sta. 7.093, 16? 34” N, 22? 54'

W, 42 m (29-VIII-1986) (NNM); 3 j, 1 f, Santa Maria Bay, CANCAP Sta. 7.094, 16” 34” N, 22? 54”

W, 24 m (NNM). Brava: 8 f, 8 s, Furna, 8 m. Boa Vista: 15 s, 3 f, Sal Rei, 3-7 m,; 2 s, Baia Teodora,

5 m; 4 s, Rife de Chaves, 12 m; 10j, CANCAP Sta. 6.056, 15? 59 N, 22? 47” W, 25 m (12-VI-1982)

(NNM) 6 s, 6 f, CANCAP Sta. 6.064, 15? 58' N, 22” 47” W, 29-32 m (12-VI-82) (NN M); 12 s, 1 j,

Ilhéu Calheta do Velho, CANCAP Sta. 7.064, 16? 11" N, 22? 58” W, 25 m (NNM); 2 j, Ilhéu

Calheta do Velho, CANCAP Sta. 7.068, 16? 11” N, 22? 59 W, (27-VII1-1986) (NNM) 15, 5j 2 f,

CANCAP Sta. 7.075, 16? 08' N, 22? 58” W, 33 m (NNM); 1 j, 5 £, CANCAP Sta. 7.079, Ilhéu de Sal

Rei, 16” 10' N, 23* 00” W, 60 m (28-VIII-1986) (NN M). Maio: 1 j, CANCAP Sta. 7.042, Ponta

Inglez /Ponta Preta, 15* 07' N, 23* 14” W, 76 m (25-VIII-1986) (NN M). Santiago: 1 s, Praia Baixa,

5 m; 1 s, Cidade Velha, 4 m; 1 s, CANCAP Sta. 6.005, 14? 54' N, 23 30” W, 75-68 m (5-VI-1982)

(NNM); 1 s, 2j CANCAP Sta. 6.007, 14” 54” N, 23? 30” W, 70-88 m (5-VI-1982) (NNM), 2 j,

CANCAP Sta. 6.015, 14” 53' N, 23? 30” W, 150 m (5-VI-1982) (NN M); 1 j, CANCAP Sta. 6.024,

15% 00' N, 23 44” W, 540 m (7-VI-1982) (NNM); 1 s, CANCAP Sta. 6.054, 14? 54' N, 23? 30” W,

Iberus, 18 (1), 2000

29-33 m (11-V1-1982) (NNM); 1 s, 1 j, CANCAP Sta. 7.120, 16? 36" N, 24? 37" W, 208 m (1-IX-

1986) (NNM). Sáo Vicente: 2 sp, Calhau, 3 m, 1 sp, 2 s, Matiota, 5-7 m; 1 s, Porto Mindelo, 12 m;

1 j, CANCAP Sta. 6.162, 16? 54” N, 25* 01” W, 38-45 m (21-VI-1982) (NN M). Santa Luzia: 1 s,

Praia Francisca, 2 m. Ilhéu Razo: 1 s, 1 j, CANCAP Sta. 7.116, 16? 36' N, 24” 36" W, 75 m (1-IX-

1986) (NN M). Sáo Nicolau: 2 j, CANCAP Sta. 6086, 16” 34” N, 24” 22” W, 35 m (15-VI-1982)

(NNM). Santo Antáo: 1 s, W of Tarrafal, CANCAP Sta. 6.108, 16” 58' N, 25* 20” W, 10 m (6-VI-

1982) (NNM). Mauritania: 3 s, Baie de lEtoile, Nouadhibou, 3 m; 1 s, off Bank d'Arguin, 20* 01'

N, 17? 32 W, 53 m (14-VI-1988) (NNM). Ghana: 7 s, 6 f, 3j, Mianmia, 25-35 m; 2 s, Busua, 6 m;

3 s, 4 f, Takoradi, 1-4 m. Senegal: 4 s, Almadies, 30 m; 2 s, N'Gor, Dakar, 5 m; 2 s, Madeleines,

Dakar. Sáo Tomé and Principe: 1 s, Esprainha, 3 m, 1 s, Lagoa Azul, 4 m, 7 s, Praia Mutamba, 5

m; 2 s, 3 f, Sáo Tomé city, 4 m. Angola: 4 sp, 18 s, Corimba, Luanda, 20 m; 2 s, Cacuaco, 7 m; 5

s, 10 f, off Luanda, 50 m; 5 s, 4, 4 f, Palmeirinhas, 30 m; 1 sp, 2 s, 3j, Buraco, near Palmeirinhas,

prov. Bengo; 3 s, Santa Maria, 15 m.

Description: See TALAVERA (1975).

Shell (Figs. 1, 2) length 5-10 mm, width

2.5-3.5 mm, not solid, elongate-conical.

Protoconch (Fig. 3) of three spiral

smooth whorls and about 400 ym of di-

ameter of last whorl, of planktotrophic

type. A spiral cord in the middle of the

last quarter of the last whorl reaches the

lower margin of the deep sinusigera

notch.

Teleoconch of 6-7 whorls; adapical 3-

4 spire whorls angulated; subsequent

whorls gradually convex; last whorls

convex. Suture shallow, with a subsutu-

ral depression gradually better marked

which gives a slightly undulated profile

to the last whorls. Colour white.

Axial sculpture of adapical whorls

consisting of somewhat prominent,

rounded, narrow, closely spaced,

slightly curved, opisthocline ribs,

becoming gradually less prominent and

more numerous (up to 30 weakly promi-

nent ribs on the last whorl) (Figs. 1, 2).

Spiral sculpture of about 8-10 prominent

spiral cords on adapical whorls, up to

more than 60 on the last whorl, those of

the base more prominent; interspaces a

little wider or of similar width. Micro-

sculpture (Fig. 8) of fine spiral threads,

with interspaces of the same width or

wider. Aperture D-shaped, large; inner

lip thin, slightly concave; anterior

channel short, shallow; outer lip with

thin external varix, slightly opisthocline

in profile.

Operculum (Fig. 7) yellowish, thick,

pyriform, with a prominent and long

inner peg.

Radula (Fig. 6): central tooth with

two pairs of basal denticles, and a pro-

minent central cusp with 3-4 small cusps

at each edge; lateral teeth with 6-7 cusps

on the inner and outer edge; inner mar-

ginal teeth with cusps on about distal

one third of the outer edge; outer margi-

nal teeth with cusps on about distal one

third of the inner and outer edge.

Habitat: Living specimens were

collected at the base of rocks on sandy

bottom.

Distribution: R. punctostriata is

known from Mauritania (TALAVERA,

1975), Senegal, Ivory Coast, Sáo Tomé,

Gabon, Cameroun (GorFas, 1999),

Angola (Goras, 1999; ROLÁN AND

RYALL, 1999), and Ghana and the Cape

Verde Archipelago (pers. obs.).

Remarks: GARCÍA-TALAVERA (1983)

considered this species a junior synonym

(Right page) Figures 1-3. Rissoina (Rissoina) punctostriata (Talavera, 1975). 1: shell from Corimba,

Luanda, Angola; 2: shell from Sal Rei, Boa Vista, Cape Verde Archipelago; 3: protoconch of a shell

from Regona, Sal. Figures 4-5: Rissoina decussata (Montagu, 1803), 4: shell from Los Canarreos

Archipelago, Cuba; 5: protoconch of the same shell.

(Página derecha) Figuras 1-3. Rissoina punctostriata (Talavera, 1975). 1: concha de Corimba,

Luanda, Angola; 2: concha de Sal Rei, Boa Vista, Archipiélago de Cabo Verde; 3: protoconcha de una

concha de Regona, Sal. Figuras 4-5: Rissoina decussata (Montagu, 1803), 4: concha del Archipiélago

de Los Canarreos, Cuba; 5: protoconcha de la misma concha.

in the Cape Verde Archipelago

ininae

The subfamily Risso

tur Z

ROLÁN AND LUQUE

Iberus, 18 (1), 2000

of Rissoina decussata (Montagu, 1803)

from the Caribbean, probably due to the

shell similarity and the protoconch of

planktotrophic type. But the compara-

tive study of the shells and protoconchs

(Figs. 1-5 and 8-9) of R. decussata from

Cuba and R. punctostriata has showed

enough differences to consider both

them different species (Table I, see page

86). So, R. decussata is not an amphiatlan-

tic species, and all the previous West

African records of this species (Senegal,

Gabon, Ivory Coast, Sáo Tomé, Came-

roun and Angola) should be attributed

to R. punctostriata. The record of Rissoina

elegantula (Angas, 1880), a similar species

described from S Australia, from Sáo

Tomé (TOMLIN AND SHACKLEFORD, 1914),

must be also referred to R. punctostriata.

CoseL (1982a, c) recorded this

species (as Zebina cf. punctostriata) from

the Cape Verde Islands. The previous

records of Rissoina decussata from Santa

Luzia and Boa Vista (DAUTZENBERG AND

FIsCHER, 1906), and the more recent of

CosEL (1982b), who cited the previous

authors, should be attributed to R. punc-

tostriata.

The study of the best preserved of

two syntypes of each R. decussata

(BMNH, no. 4239) and R. striatocostata

(d'Orbigny, 1842) (BMNH 1854.10.4.209)

has proved that the last name is a junior

synonym of R. decussata.

Subgenus Ailinzebina Ladd, 1966

Type species: Zebina (Ailinzebina) abrardi Ladd, 1966, by original designation.

Diagnosis: SLEURS (1993, p. 112).

Rissoina (Ailinzebina) onobiformis n. sp. (Figs. 10-14, 158)

Type material: Holotype (Fig. 11) 1 s of 3.1 x 1.2 mm and 2 paratypes, 2 s, from Rabo de Junco, Sal

Island, Cape Verde Archipelago, 6 m (MNCN 15.05/31713). Other paratypes: 4 s (Fig. 10, broken

during the study), 3 f, from the type locality, 2 m (CER); 1 s, Derrubado, Boa Vista, 3 m (CER); 1 s,

Pedrinha, Brava, 6 m (CER); 2 s, 2 f, 1 j, Furna, Brava, 20-30 m (CER); 1 s, Furna, Brava, 30 m (AMNH);

1 s, Sal Rei, Boa Vista, 6 m (NNM 58020); 1 s, South of Santiago, CANCAP Sta. 6.015, 14? 53" N, 23?

30' W, 50 m (5-V1-1982) (NNM 59417); 1 s, 5 m, Derrubado, Boa Vista (MNHN); 1 s, Cidade Velha,

Santiago, 5 m (DBUA).

Other material studied: Boa Vista: 2 s, Morro de Areia, 1 m; 1 f Baijos de Joáo Valente, 20 m; 3 j,

Baia Teodora, 4 m. Santiago: 1 f, Cidade Velha, 4 m; 1 f, South of Santiago, CANCAP Sta. 6.015, 14?

53 N, 23" 30” W, 50 m (5-VI-1982) (NNM). Brava: 2 s, Furna, 30 my 6 s, Porto do Anciáo, 6 m.

Etymology: The specific name alludes to the resemblance of the shell with that of the genus Onoba.

Description: Shell (Figs. 10, 11) length

up to 3.5 mm, maximum width 1.4 mm,

not solid, elongate, subcylindrical, with

pupoid apex.

Protoconch (Figs. 12, 13) of a little

more than one whorl and 360 um of

maximum diameter, of non-planktotro-

phic type; transition to teleoconch

abrupt; surface smooth with some rather

elevated marks like an Arabic writing.

Teleoconch of 4 */2 whorls in holo-

type, but usually 4, weakly convex, not

(Right page) Figures 6-8. Rissoina punctostriata (Talavera, 1975). 6: radula of a specimen of Mor-

deira, Sal, Cape Verde Archipelago; 7: operculum of the same specimen; 8: teleoconch microsculp-

ture of the shell of Figure 2. Figure 9. Rissoina decussata (Montagu, 1803), teleoconch microsculp-

ture of the shell of Figure 4.

(Página derecha) Figuras 6-8. Rissoina punctostriata (Talavera, 1975). 6: rádula de un ejemplar de

Mordeira, Sal, Archipiélago de Cabo Verde; 7: opérculo del mismo ejemplar; 8: microescultura de la

teloconcha del ejemplar de la Figura 2. Figura 9. Rissoina decussata (Montagu, 1803), microescultura

de la teloconcha del ejemplar de la Figura 4.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Iberus, 18 (1), 2000

angulated below suture, but with a

small angulation near the base; last

whorl weakly convex; suture shallow.

Colour cream-whitish.

Axial sculpture consisting of very

weak, narrow, slightly opisthocline, dis-

tantly spaced axial ribs, gradually more

closely spaced, 30-37 in the last whorl, a

little prominent on the suture. Spiral

sculpture very fine, appreciable at low

magnification. Microsculpture (Fig. 14)

formed by bands of 6-10 very fine

threads between two irregular fine

spiral cords.

Aperture D-shaped, relatively large,

with an acute angulation on its upper

part; inner lip thick; columellar side

weakly concave; anterior channel

almost absent; outer lip with a weak

external varix; peristome wide and flat

when well developed, with an internal

and an external sharp rims; inner part

with two small depressions, one on the

anterior channel and other on the part

corresponding, to the anal sinus.

Habitat: Shells from sand sediments

between 2 and 50 m.

Distribution: Sal, Boa Vista, Santiago,

Brava (Fig. 158). This species probably

will be found in all the islands of the

Cape Verde Archipelago.

Remarks: PONDER (1985) considered

Ailinzebina a synonym of Rissoína s. s., but

SLEURS (1993) considered the radular and

head-foot characters of Rissoina (Ailinze-

bina) elegantissima enough different from

those of Rissoina s. s. to warrant a subge-

neric status for Ailinzebina. SLEURS (1993)

included in this subgenus four Pacific

species and Rissoina (Ailinzebina) elegan-

tissima d'Orbigny, 1842, from the Carib-

bean. Considering the distinctive shell

features (and also the known anatomical

ones, see SLEURS, 1993) of the species of

this subgenus and its wide distribution,

we think that Ailinzebina may be elevated

to the generic level, but we prefer to do

not any taxonomical change waiting for

further anatomical information.

Rissoina (Ailinzebina) elegantissima is

the only other known Atlantic species of

this subgenus and it differs from R. (4.)

onobiformis n. sp. by the planktotrophic

type of protoconch (see LEAL AND

MOORE, 1989, fig. 9, and SLEURS, 1993,

fig. 41), the rather solid shell with

strongly convex whorls, the more pro-

minent and opisthocline axial ribs and

the more densely spaced spiral cords.

Moreover, the sculpture of the proto-

conch of R. onobiformis is different from

any other known species of Rissoina.

Genus Schwartziella Nevill, 1881

Subgenus Schwartziella s. s.

Type species: Rissoina orientalis Nevill, 1881 (= Rissoina triticea Pease, 1861), by original designation.

Diagnosis: PONDER (1985, p. 98-99).

Schwartziella (Schwartziella) robusta n. sp. (Figs. 15-19, 144, 148, 159)

Type material: Holotype (Fig. 15) 1 s of 3.7 x 1.9 mm, and 1 paratype, 1 s, Fiura, Sal Island, Cape

Verde Archipelago (MNCN 15.05/31718). Other paratypes: 3 s, Regona, 10 m, and 3 s, Punta Preta

(Right page) Figures 10-14: Rissoina (Ailinzebina) onobiformis n. sp. 10: paratype (broken during

study), Rabo de Junco, Sal (CER); 11: holotype, Rabo de Junco, Sal (MNCN 15.05/31713); 12-

13: protoconchs of paratypes, Rabo de Junco, Sal (CER); 14: teleoconch microsculpture of a

paratype, Furna, Brava (CER).

(Página derecha) Figuras 10-14: Rissoina (Ailinzebina) onobiformis spec. nov. 10: paratipo (roto

durante su estudio), Rabo de Junco, Sal (CER); 11: holotipo, Rabo de Junco, Sal (MNCN

15.05/31713); 12-13: protoconchas de paratipos, Rabo de Junco, Sal (CER); 14: microescultura de la

teloconcha de un paratipo, Furna, Brava (CER).

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in the Cape Verde Archipelago

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ROLÁN AND LUQUE

Iberus, 18 (1), 2000

(DBUA); 1 s, Rabo do Junco, 4 m (AMNH); 1 s, Rabo do Junco, 3 m (NNM 58028); 4 s, Regona, 2-3

m, and 6s, 1 f, Mordeira Bay, 5 m (CER); 1 s, Regona, 2 m (MNHN). All the type material from Sal.

Other material studied: Sal: 2 sp, Mordeira Bay, 4 m (broken for radular study); 2 s, Regona. Boa

Vista: 3 s, Sal Rei, 8 m; 8 s, Porto da Cruz, 4 m.

Etymology: The specific name alludes to the very solid shell.

Description: Shell (Figs. 15, 16) length

up to 4.3 mm, maximum width 2.0 mm,

very solid, elongate-conic.

Protoconch (Fig. 17) of 1 whorl and

400 ym of maximum diameter, of non-

planktotrophic type, with a spiral cord

and a slight angulation below, transition

to teleoconch abrupt. Microsculpture

(Fig. 18) formed by numerous small pits.

Teleoconch of 5 whorls; two adapical

spire whorls convex below sutures, sub-

sequent spire whorls becoming grad-

ually angulate only in subsutural part;

suture evident; last whorl strongly

convex. Colour whitish.

Axial sculpture consisting of promi-

nent, strongly convex, rounded, almost

orthocline, widely spaced axial ribs,

slightly shouldered in their subsutural

part, between 10-12 in the last whorl.

Spiral sculpture inconspicuous. Micro-

sculpture (Fig. 19) formed by very

numerous fine spiral threads, with a

wider one between each 5-13 fine

threads, sometimes more evident in the

subsutural region.

Aperture D-shaped, relatively small;

inner lip thick; columellar side weakly

concave; anterior channel shallow; outer

lip opisthocline, with strong varix with

several concentric lines towards the

inner part of the aperture.

Operculum (Fig. 148) translucent,

thin, paucispiral, with a very eccentric

nucleus, and without any peg on inner

side; the insertion area is elongate and

close to the edge.

Radula (Fig. 144): central tooth with 1

pair of basal denticles, and a prominent

central cusp with 3-4 small cusps at each

side; inner marginal teeth finely denticu-

late; outer marginal teeth without any

denticles on the external margin.

Habitat: Sandy sediments in shallow

water.

Distribution: Only known from Sal

and Boa Vista Islands (Fig. 159).

Remarks: The holotype of Schwartzie-

lla africana (Dautzenberg, 1913) (MNHN)

from “Pointe de Bel-Air (baie de Hann)””,

Dakar (Senegal) lacks protoconch, but it

is smaller (2.9x1.2 mm) than S. robusta n.

sp., the whorls are not angulate, the axial

ribs are 4-5 times narrower than inter-

spaces and there are fine growth lines

more evident at the subsutural region

and very fine spiral threads more

evident at the middle of the whorls. The

sole specimen of Rissoina africana var.

crassior (Dautzenberg, 1913), from the

same locality (MNHN), is very different

of the holotype in having an undulate

suture between the penultimate and the

last whorls (almost straight in the

holotype), axial ribs only a little nar-

rower than interspaces, and apparently

no microsculpture. The protoconch is

paucispiral (one whorl) and apparently

smooth. It is also smaller (2.9x1.4 mm)

than S. robusta n. sp.

The specimens described and illus-

trated by GOFAS (1999) under the name

of Schwartziella africana from Senegal are

also different from S. robusta n. sp.: they

(Right page) Figures 15-19: Schwartziella (Schwartziella) robusta n. sp. 15: holotype, Fiura, Sal

(MNCN 15.05/31718); 16: paratype, Fiura, Sal (UNCN 15.05/31718); 17: protoconch of a

paratype, Regona, Sal (CER); 18: protoconch microsculpture of the same paratype; 19: teleoconch

microsculpture of the same paratype.

(Página derecha) Figuras 15-19: Schwartziella (Schwartziella) robusta spec. nov. 15: holotipo, Fiura,

Sal (MNCN 15.05/31718); 16: paratipo, Fiura, Sal (MNCN 15.05/31718); 17: protoconcha de un

paratipo, Regona, Sal (CER); 18: microescultura de la protoconcha del mismo paratipo; 19: microescul-

tura de la teloconcha del mismo paratipo.

Archipelago

The subfamily Rissoininae in the Cape Verde

ROLÁN AND LUQUE

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Iberus, 18 (1), 2000

are larger (up to 5.25x2.15 mm), with the

whorls not angulate but convex, and the

protoconch has no spiral cords but

rounded spots. According to (GOFAS

(1999), S. africana is only known with

certainty from a small stretch of coast-

line around Dakar, Senegal. The only

shell recorded from Maio island (Cape

Verde Archipelago) must be regarded as

an erroneous record, since no specimens

of this species were found in the large

material examined in this paper.

The differences of S. robusta with the

other new species of Schwartziella descri-

bed below are given in the remarks of

each species.

Schwartziella (Schwartziella) obesa n. sp. (Figs. 20-24, 160)

Type material: Holotype (Fig. 20) 1 s of 5.5 x 2.4 mm from Furna Bay, Brava Island, Cape Verde

Archipelago, 15-20 m (MNCN 15.05/31712). Paratypes: 3 s (DBUA); 2 s (CER); 1 s (AMNH); 1 s

(NNM 58019); 1 s (MNAHN,), all from the type locality.

Other material studied: Brava: 2 f, Furna, 30 m; 2 s, Pedrinha, 4 m; 1 s, Porto do Anciáo, 30 m. San-

tiago: 1 s, Praia, 5 my 1 sp, 5 s (Fig. 16), 4 f Tarrafal, 3 m; 2 s, Cidade Velha, 5 m, 2 £ CANCAP Sta.

6.015, S of Santiago, 14” 53' N, 23” 30' W, 150 m (5-VI-1982) (NN M); 2 j, 1 £ CANCAP Sta. 6.024, 15?

00' N, 23? 44” W, 540 m (7-VI-1982) (NN M); 2j, CANCAP Sta. 6.054, 14? 54” N, 23? 30" W, 29-33 m

(11-VI-1982) (NNM) 13, CANCAP Sta. 7.008, 14” 54” N, 23” 38" W, 320 m (20-VII-1986) (NNM).

Fogo: 1 j, CANCAP Sta. 6.052, 14” 53 N, 24? 31” W, 85 m (10-VI-1982) (NNM). Sáo Vicente: 1 s,

Calhau, 4 m. Maio: 2 f, CANCAP Sta. 7.050, 15? 06' N, 23? 14' W, 380 m (25-VII1-1986) (NNM).

Etymology: The specific name alludes to the wide shell.

Description: Shell (Figs. 20, 21) length

up to 7.0 mm, maximum width 2.8 mm,

solid, elongate-conic, strongly scalari-

form.

Protoconch (Fig. 22) of 1 whorl and

about 300 um of maximum diameter, of

non-planktotrophic type, with a spiral

cord in its upper part; transition to

teleoconch abrupt. Microsculpture

formed by very small pits.

Teleoconch of 5-6 whorls; spire

strongly scalariform, whorls with a

prominent subsutural shoulder and

rapidly enlarging; suture shallow but

clearly visible. Colour whitish.

Axial sculpture consisting of promi-

nent, sharp, narrow and widely spaced

axial ribs, which are aligned across

several whorls, slightly opisthocline on

the first whorls and almost orthocline in

the body whorl, on where there are

about 14-15 ribs, clearly arched to the

base. Spiral sculpture formed by very

fine cords. Microsculpture (Figs. 23, 24):

all the surface of the shell is covered by

very fine threads, both on the spiral

cords and the interspaces.

Aperture D-shaped, small; inner lip

thick; columellar side strong, weakly

concave; anterior channel shallow; outer

lip opisthocline, with thick external

varix, with about eight parallel lines

towards the inner part of the aperture.

Habitat: Sandy sediments from 3 to

600 m.

Distribution: Known from the group

of Brava-Santiago-Fogo Islands, but

some fragments from Maio and one

shell from Sáo Vicente seem to be this

species (Fig. 160).

(Right page) Figures 20-24: Schwartziella (Schwartziella) obesa n. sp. 20: holotype, Furna, Brava

(MNCN 15.05/31712); 21: shell from Tarrafal, Santiago (CER); 22: protoconch of a shell from

Tarrafal, Santiago (CER); 23: teleoconch microsculpture of the holotype; 24: teleoconch micro-

sculpture of a shell from Tarrafal (CER).

(Página derecha) Figuras 20-24: Schwartziella (Schwartziella) obesa spec. nov. 20: holotipo, Furna,

Brava (MNCN 15.05/31712); 21: concha de Tarrafal, Santiago (CER); 22: protoconcha de una

concha de Tarrafal, Santiago (CER); 23: microescultura de la teloconcha del holotipo; 24: microescul-

tura de la teloconcha de una concha de Tarrafal (CER).

in the Cape Verde Archipelago

ininae

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The subfamily Risso

ROLÁN AND LUQUE

Iberus, 18 (1), 2000

Remarks: The shells of Schwartziella

obesa n. sp. from Brava seem to be a little

different from those of Santiago and Sáo

Vicente: the latter are a little smaller, the

axial ribs are a little closer, the subsutu-

ral angle is slightly more elevate and the

spiral threads are more depressed (see

Figs. 18, 19). All these differences seem

to be not relevant in order to consider

both populations to be not conspecific,

and probably they are the expression of

a difficult genetic flow between popula-

tions of different islands, but only more

detailed studies on living specimens

will clarify this matter.

S. robusta n. sp. also has a thick shell,

but it lacks the subsutural angulation

and the scalariform profile of S. obesa, the

axial ribs are less numerous, sharper and

more arched in its subsutural part and to

the base. The spiral threads of the subsu-

tural part are more evident in S. robusta,

whereas in S. obesa are more attenuated.

S. africana (Dautzenberg, 1913) and

S. africana var. crassior (Dautzenberg,

1913) are smaller, their whorls are not

angulate, and the axial ribs and the

spiral sculpture and microsculpture are

different. The protoconch of S. africana

var. crassior is apparently smooth,

without any spiral cord (see under

remarks of S. robusta). The specimens of

Schwartziella africana described and illus-

trated by GOFAs (1999) from Senegal are

also different: they are smaller, the

whorls are not angulate, have a lower

number of ribs on the body whorl (ca.

12), the protoconch has no spiral cords

and its microsculpture consists of

rounded spots.

Schwartziella (Schwartziella) corrugata n. sp. (Figs. 25-29, 161)

Type material: Holotype (Fig. 25) 1 s of 5.6 x 2.1 mm from Furna, Brava Island, Cape Verde Archi-

pelago, 30 m (MNCN 15.05/31703). Paratypes: 1 s (MNHN) and 3 s (CER) (Fig. 26), all from the

type locality.

Other material studied: Brava: 2 s, 1 f, 2] (1 broken), Pedrinha, 10 m;, 1 s, 2 f, Porto do Anciáo, 3 m;

1 s, llhéus do Rombo, 3 m.

Etymology: The specific name alludes to the sutural undulation formed by the axial ribs of the shell.

Description: Shell (Fig. 25) length up

to 5.6 mm, maximum width 2.1 mm,

very solid, elongate-conic.

Protoconch (Fig. 26) of 1 whorl and

360 um of maximum diameter, of non-

planktotrophic type, with one spiral

cord in its upper part and an angulation

below; transition to teleoconch abrupt.

Microsculpture (Fig. 29) formed by very

small pits.

Teleoconch of 5 whorls, weakly

convex, slightly angulated below su-

tures; last whorl weakly convex; suture

(Figs. 26, 27) well marked and undulous

due to the axial ribs. Colour whitish.

Axial sculpture consisting of promi-

nent, rounded, spaced axial ribs, which

are almost orthocline in the body whorl

and opisthocline in previous whorls,

curved subsuturally, and about 12 in last

whorl. Near the base, the interspace

between ribs is deep. Spiral sculpture

formed by fine cords, visible at low

magnification, and more evident in the

(Right page) Figures 25-29: Schwartziella (Schwartziella) corrugata n. sp. 25: holotype, Furna,

Brava (MNCN 15.05/31703); 26: protoconch of a paratype, Furna, Brava (CER); 27: detail of the

suture of a paratype, Furna, Brava (CER); 28: teleoconch microsculpture of a paratype, Furna,

Brava (CER); 29: protoconch microsculpture of a paratype, Furna, Brava (CER).

(Página derecha) Figuras 25-29: Schwartziella (Schwartziella) corrugata spec. nov. 25: holotipo,

Furna, Brava (MNCN 15.05/31703); 26: protoconcha de un paratipo, Furna, Brava (CER); 27:

detalle de la sutura de un paratipo, Furna, Brava (CER); 28: microescultura de la teloconcha de un

paratipo, Furna, Brava (CER); 29: microescultura de la protoconcha de un paratipo, Furna, Brava

(CER).

Archipelago

in the Cape Verd

ininae

The subfamily Risso

ROLÁN AND LUQUE

url ]

Iberus, 18 (1), 2000

subsutural zone. Microsculpture (Fig.

28) formed by many irregular threads

between and on the spiral cords.

Aperture D-shaped, relatively small;

inner lip thick; columellar side weakly

concave; anterior channel shallow; outer

lip thick with external varix with about

seven parallel lines towards the inner

part of the aperture.

Habitat: The material studied was

obtained from sand sediments between

a few meters and 30 m deep.

Distribution: S. corrugata n. sp. is only

known from Brava Island and Ilhéus do

Rombo (Fig. 161).

Remarks: S. corrugata n. sp. has a shell

as solid as the precedent ones (S. robusta

and S. obesa), but it can be differentiated

by its very undulous sutural line due to

the stronger and elevated axial ribs and it

lacks of the subsutural depression. S.

corrugata and S. robusta have similar pro-

toconch and teleoconch microsculpture,

and the allopatric distribution of both

species point out to be different morphs

(or subspecies) of an unique species dis-

tributed in different islands. Nevertheless,

no intermediate forms between the

marked undulous suture of S. corrugata

and the linear suture of S. robusta were

found in other islands, so we consider

them as different species. S. corrugata is

sympatric with S. obesa in Brava.

Schwartziella africana (Dautzenberg,

1913) is smaller, the whorls are not

angulate, and the suture, axial and

spiral sculpture and microsculpture are

also different (see under remarks of S.

robusta).

S. africana var. crassior (Dautzenberg,

1913) has an undulate suture, but it is

also smaller, its axial ribs are wider, and

apparently has no microsculpture both

in protoconch and teleoconch.

Schwartziella (Schwartziella) sanmartini n. sp. (Figs. 30-34, 146, 149-151, 156, 162)

Type material: Holotype (Fig. 30) 1 s of 4.8 x 2.0 mm, and paratypes, 2 s, from Mordeira Bay, Sal

Island, Cape Verde Archipelago, 4 m (MNCN 15.05/31719). Paratypes: Paratypes: 3 s, Mordeira

Bay, Sal, 5 m (CER); 4 sp, 6 s, Rabo de Junco, Sal, 4 m (CER); 2 sp, 1 s, Mordeira, Sal (DBUA); 2 s,

Regona, Sal, 4 m (MNHN); 1 s, Mordeira, Sal, 4 m (AMNH); 1 s, Mordeira, Sal, 4 m (NNM 58029).

Other material studied: Sal: 1 s, 6j, Palmeira, 6 m; 6 s, 1 f, Mordeira, 5 m; 1 s, Rabo de Junco, 2 m;

1 s, Algodoeiro, 4 m. Boa Vista: 5 s, Sal Rei, 4 m; 2 sp, 2 s, Ilhéu de Sal Rei, 5 m; 1 s, Porto Ferreira,

5 m; 3 s, Porto da Cruz, 4 m; 2 s, 4 j, Baia Teodora, 5 m; 3 s, Derrubado, 5 m. Maio: 1 s, Navio Que-

brado, 3 m.

Etymology: The specific name is dedicated to the zoologist Guillermo San Martín, companion of

some research trips.

Description: Shell (Fig. 30) length up

to 5.0 mm, maximum width 2.1 mm,

except by a very slight angulation in its

upper part. Microsculpture (Fig. 34)

relatively solid, elongate-conic.

Protoconch (Fig. 31) of 1 whorl and

360 ym of maximum diameter, of non-

planktotrophic type, no spiral sculpture

formed by very small pits.

Teleoconch of about 5 whorls weakly

convex, slightly angulated below sutures;

last whorl large, weakly convex, repre-

(Right page) Figures 30-34: Schwartziella (Schwartziella) sanmartini n. sp. 30: holotype, Mordeira,

Sal (UNCN 15.05/31719); 31: protoconch of the holotype; 32: detail of the suture of a paratype,

Mordeira (CER); 33: teleoconch microsculpture of the same paratype; 34: protoconch microsculp-

ture of the same paratype.

(Página derecha) Figuras 30-34: Schwartziella (Schwartziella) sanmartini spec. nov. 25: holotipo,

Mordeira, Sal (MNCN 15.05/31719); 26: protoconcha del holotipo; 27: detalle de la sutura de un

paratipo, Mordeira (CER); 28: microescultura de la teloconcha del mismo paratipo; 34: microescultura

de la protoconcha del mismo paratipo..

un 001

in the Cape Verde Archipelago

ininae

The subfamily Risso

A AS

ROLÁN AND LUQUE

Iberus, 18 (1), 2000

senting more than a half of the shell;

suture shallow, slightly undulate due to

the axial sculpture. Colour whitish.

Axial sculpture consisting of depres-

sed, rounded, narrow, almost orthocline

in last whorl and opisthocline in pre-

vious whorls, distantly spaced axial ribs,

about 14 in last whorl. Spiral sculpture

almost not appreciable at low magnifica-

tion. Microsculpture (Figs. 32, 33)

formed by very fine threads axially inter-

rupted and modified by growth lines.

Aperture D-shaped, small; inner lip

thick; columellar side weakly concave;

anterior channel almost absent; outer lip

with ample external varix with several

parallel lines towards the inner of the

aperture.

Operculum (Figs. 149-151) translu-

cent, thin, paucispiral with the nucleus

very eccentric and without any promi-

nent peg on inner side.

Radula (Fig. 146): central tooth with

one pair of basal denticles, a slightly

prominent central cusp, and 3-4 smaller

cusps at each side, the more basal ones

very small; lateral teeth with 6-8 cusps

on the inner edge and 4-5 on the outer

edge; inner marginal teeth with many

small denticles on both edges (serrated);

outer marginal teeth without any denti-

cle on the outer edge.

The animal (Fig. 156) examined in

alcohol is apparently white. The penis

has a cylindrical base; in the middle part

it is curved ahead, and flattened and

enlarged at its distal end; the margin of

both sides of this terminal widening are

denticulate and also two denticulate

fringes appear close to the tip, with a

short and sharp appendix towards the

right part.

Habitat: The material studied was ob-

tained in sand sediments between 3-6 m.

Distribution: Only known from Sal,

Boa Vista and Maio (Fig. 162).

Remarks: Schwartziella sanmartini n.

sp. has a thinner shell than S. robusta, S.

obesa and S. corrugata, and its axial ribs

are more depressed and rounded and

less elevate. Also it differs from S. ro-

busta n. sp. by having more axial ribs,

from S. obesa n. sp. by lacking of any

subsutural angulation, and from $. co-

rrugata by the slightly undulate suture.

S. africana y S. africana var. crassior are

smaller; the first species has narrower

ribs, fine growth lines more evident at

the subsutural region and very fine

spiral threads more evident at the

middle of the whorls. S. africana var. cras-

sior has an undulate suture, axial ribs

only a little narrower than interspaces,

and apparently no microsculpture.

Schwartziella (Schwartziella) similiter n. sp. (Figs. 35-40, 147, 152, 163)

Type material: Holotype (Fig. 35) 1 s of 5.6 x2.1 mm and 5 paratypes, 5 s, Furna, Brava Island, Cape

Verde Archipelago, 8-20 m (MNCN 15.05/31721). Other paratypes: 2 s from the type locality in

each of MNHN, AMNH, DBUA, NNM (58031), and 42 s in CER.

Other material studied: Brava: 3 sp, 115 s, 41 f, 2 j, Furna, 8-20 my 5 s, 8 j, Pedrinha, 4 m; 3 sp, 23 s,

14 j, 3 £, Porto do Anciáo, 3 m; 6 s, 4 j, Ilhéus do Rombo, 3-5 m. Santiago: 31 s, 5 j, 6 f£, Praia, llhéu

de Santa Maria, 6 m; 8 s, 5j, 1 f, Prainha, 5 m; 4 s, Cidade Velha, 4 m; 29 s, 2j, Tarrafal, 4 m; 1 s, 15,

1f CANCAP Sta. 6024, 15% 00' N, 23? 44” W, 540 m (7-VI-1982) (NNM). Sáo Vicente: 13 s, Calhau,

3 m; 1 s, Porto Mindelo, 15 m, 4 s, Salamanca. Santa Luzia: 2 sp, 1 s, Praia Francisca, 3 m.

(Right page) Figures 35-40: Schwartziella (Schwartziella) similiter n. sp. 35: holotype, Furna, Brava

(MNCN 15.05/31721); 36: shell from Tarrafal, Santiago (CER); 37: protoconch of a paratype,

Furna (CER); 38: protoconch microsculpture of the same paratype; 39-40: teleoconch microsculp-

ture of a shell from Furna.

(Página derecha) Figuras 35-40: Schwartziella (Schwartziella) similiter spec. nov. 35: holotipo, Furna,

Brava (MNCN 15.05/31721); 36: concha de Tarrafal, Santiago (CER); 37: protoconcha de un para-

tipo, Furna (CER); 38: microescultura de la protoconcha del mismo paratipo; 39-40: microescultura de

la teloconcha de un ejemplar de Furna.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

a a

Iberus, 18 (1), 2000

Etymology: The specific name alludes to the similarity of the shell of this species with other species

of Cape Verde Schwartziella.

Description: Shell (Figs. 35, 36) length

up to 6.0 mm, maximum width 2.1 mm,

relatively solid, elongate conic.

Protoconch (Fig. 37) of 1 whorl and

about 300 ym of maximum diameter, of

non-planktotrophic type, with a spiral

cord running along a keel on its upper

part; transition to teleoconch abrupt.

Microsculpture (Fig. 38) formed by very

small pits.

Teleoconch of 5 whorls, regularly

convex, not angulated below sutures, last

whorl weakly convex; suture shallow,

slightly undulate. Colour whitish.

Axial sculpture consisting of slightly

prominent, rounded, narrow, almost or-

thocline in the last whorl and opisthocline

in previous, distantly spaced axial ribs not

regularly continued from whorl to whorl.

Spiral sculpture appreciable with diffi-

culty at small magnification. Microsculp-

ture (Figs. 39, 40) consisting in very close

relatively thick spiral threads interrupted

by axial growth lines, with 2-5 thinner

threads between each two of them which

are interrupted in some places.

Aperture D-shaped, small; inner lip

thick; columellar side weakly concave;

anterior channel almost absent; outer lip

with a thick varix; with about 5 parallel

lines.

Operculum (Fig. 152) translucent,

thin, paucispiral, with a very eccentric

nucleus and without any prominent peg

on the inner side.

Radula (Fig. 147): central tooth with

one pair of basal denticles, a few promi-

nent central cusp and 3-4 small cusps at

each side, those of the extremes very

small; lateral teeth with 7-8 cusps on the

inner edge and 4-5 on the outer edge

inner marginal teeth with many small

denticles; outer marginal teeth without

denticles on the external edge.

Habitat: The material studied was

obtained in sand sediments from 4 to 20

m. One sample was collected at 540 m.

Distribution: Santiago, Brava, Sao

Vicente and Santa Luzia (Fig. 163).

Remarks: The shells from the north-

ern (Santa Luzia and Sáo Vicente) and

the southern (Brava and Santiago)

islands are very similar, with only small

differences in size, and we consider

them conspecific though both groups of

islands are quite far for this species with

non-planktotrophic development.

S. similiter m. sp. is similar to S. san-

martini n. sp., and the differences with

other species mentioned in its remarks are

not repeated here. S. sanmartini n. sp. is

wider than S. similiter n. sp., has more axial

ribs and these are more depressed, also, it

has different microsculpture, with less

marked and interrupted spiral threads,

and a slightly smaller protoconch.

Schwartziella (Schwartziella) typica n. sp. (Figs. 41-45, 142, 143, 145, 164)

Type material: Holotype (Fig. 41) 1 s of 3.8 x 1.6 mm and one paratype, 1 s, Palmeira, Sal Island,

Cape Verde Archipelago, 6 m (MNCN 15.05/31705). Paratypes: 1 s from Monte Leste, Sal, in each of

MNHN, AMNH, DBUA, NNM (58006), and 1 s, from Guincho do Ninho, 4 m; 2 s, from Palmeira, 6

m, 1 s, from Punta Preta, 3 m; 16 s, from Rabo de Junco, 6 m, and 4 s, from Regona, 10 m, all in CER.

Other material studied: Sal: 5 s, Palhona, 1 m; 1 sp (broken for radular study), 3 s, 6 £, Monte Leste,

1 m, 4 s, Algodoeiro, 4 m; 4 s, Palmeira, 8 m; 4 s, Regona, 3 m; 3 s, 1 j, Mordeira, 5m;7s, 3j, 3 f,

(Right page) Figures 41-45: Schwartziella (Schwartziella) typica n. sp. 41: holotype, Palmeira, Sal

(MNCN 15.05/31705); 42: protoconch of a paratype, Palmeira (CER); 43: protoconch micro-

sculpture of the same paratype; 44-45: teleoconch microsculpture of the same paratype.

(Página derecha) Figuras 41-45: Schwartziella (Schwartziella) typica spec. nov. 41: holotipo, Pal-

meira, Sal (MNCN 15.05/31705); 42: protoconcha de un paratipo, Palmeira (CER); 43: microescul-

tura de la protoconcha del mismo paratipo; 44-45: microescultura de la teloconcha del mismo paratipo.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

“S

Iberus, 18 (1), 2000

CANCAP Sta. 7.100, off Palmeira, 16 45' N, 23? 01” W, 354 m (30-VIII-1986) (NNM), 2 j, off Pal-

meira, CANCAP Sta. 7.109, 16? 45' N, 22* 59 W, 31 m (31-VIII-1986) (NN M). Boa Vista: 6 s, 2 j, Sal

Rei. Sáo Nicolau: 17 s, 6 j, 6 f Sáo Jorge Bay, CANCAP Sta. 7.129, 16” 33' N, 24? 16" W, 405 m (2-IX-

1986) (NNM); 13 s, 8 £, CANCAP Sta. 7.128, 16? 33/ N, 24? 17" W, 400 m (2-1X-1986) (NNM).

Etymology: The specific name alludes to the typical aspect of the shell for the genus.

Description: Shell (Fig. 41) length up

to 4.5 mm, maximum width 1.8 mm,

solid, elongate-conic.

Protoconch (Fig. 42) of 1 whorl and

about 290 ym of maximum diameter, of

non-planktotrophic type, with a depres-

sed spiral cord running along a keel in

its upper part; transition to teleoconch

abrupt. Microsculpture (Fig. 43) formed

by flat prominences with irregular

edges on an undulated surface.

Teleoconch of about 5 whorls, strongly

convex, not angulated below sutures; last

whorl convex; suture shallow, but evident.

Colour white.

Axial sculpture consisting of promi-

nent, relatively rounded, narrow, almost

orthocline in last whorl and opisthocline

in previous ones, distantly spaced axial

ribs; the ribs are slightly curved in most

of spire, but less in the last whorl. Spiral

sculpture not appreciable at low magnifi-

cation. Microsculpture (Figs. 44, 45)

formed by bands of 4-6 closely packed,

very fine spiral threads with minute pits

between each two of them and 1-3 fine

threads between each two of these bands.

Aperture D-shaped, small; inner lip

thick; columellar side weakly concave;

anterior channel almost absent; outer lip

with thick and ample varix, almost or-

thocline; with 5-6 lines towards the

inner part of the aperture.

Operculum translucent, with very

eccentric nucleus and without any

prominent peg on the inner side.

Radula (Fig. 145): central tooth with

one pair of basal denticles, a slightly

prominent central cusp, and 3-4 cusps at

each side, those of extremes very small;

lateral teeth with 6-8 cusps on the inner

edge and 4-6 on the outer edge; inner

marginal teeth with many small denti-

cles on both edges, outer marginal teeth

without denticles on external edge.

Habitat: The material studied was

obtained in sand sediments from 1 to

400 m.

Distribution: Sal, Boa Vista and Sáo

Nicolau Islands (Fig. 164).

Remarks: S. typica n. sp. is similar to

S. sanmartini n. sp. and S. similiter n. sp.,

but it is smaller than S. sanmartini and S.

similiter, has more elevated axial ribs,

less evident spiral sculpture and differ-

ent teleoconch and protoconch micros-

culpture. S. africana has a similar proto-

conch microsculpture (GOFAS, 1999), but

differs by its oblique axial ribs.

One shell (Figs. 142, 143) found in

Calhau, Sáo Vicente, is similar to this

species and has the same microsculp-

ture, but has a more evident spiral

sculpture; we provisionally consider it

conspecific awaiting for further mate-

rial.

Schwartziella (Schwartziella) angularis n. sp. (Figs. 46-50, 159)

Type material: Holotype (Fig. 46) 1 s of 2.9 x 1.3 mm from Rabo de Junco, Sal Island, Cape Verde

Archipelago, 4 m (MNCN 15.05/31701). Paratypes: 1 s from the type locality in each of MNHN,

(Right page) Figures 46-50: Schwartziella (Schwartziella) angularis n. sp. 46: holotype, Rabo de

Junco, Sal (MNCN 15.05/31701); 47: paratype, Rabo de Junco, Sal (CER); 48: protoconch of a

paratype, Rabo de Junco (CER); 49-50: teleoconch microsculpture of the same paratype.

(Página derecha) Figuras 46-50: Schwartziella (Schwartziella) angularis spec. nov. 46: holotipo, Rabo

de Junco, Sal (MNCN 15.05/31701); 47: paratipo, Rabo de Junco, Sal (CER); 48: protoconcha de un

paratipo, Rabo de Junco (CER); 49-50: microescultura de la teloconcha del mismo paratipo.

Archipelago

in the Cape Verde

ininae

The subfamily Risso

ROLÁN AND LUQUE

var

005

Iberus, 18 (1), 2000

AMNH, DBUA, NNM (58000); further paratypes in CER: 3 s, Rabo de Junco, 4 m; 2 s, Pesqueiro do

Aire, 1 m, and 2 s, Regona, 2 m.

Other material studied: Sal: 1 s, Rabo de Junco, 4 m; 2 f, CANCAP Sta. 7.110, 16” 46' N, 23 02” W,

85 m (31-VIIT-1986) (NN M). Boa Vista: 1 f, Sal Rei, 3 m.

Etymology: The specific name alludes the angulated subsutural shoulder of the shell.

Description: Shell (Figs. 46, 47) length

up to 3.0 mm, maximum width 1.5 mm,

relatively solid, elongate-conic, strongly

scalariform.

Protoconch (Fig. 48) of 1 whorl and

about 400 ym of maximum diameter, of

non-planktotrophic type, with only a

faint spiral angulation on its upper part;

transition to teleoconch abrupt.

Teleoconch of about 5 whorls, which

are strongly scalaroid, with a prominent

subsutural shoulder and rapid develop-

ment. Suture shallow, but evident.

Colour whitish.

Axial sculpture consisting of promi-

nent, straight, sharp, narrow and spaced

axial ribs, regularly continued from

whorl to whorl, orthocline or slightly

opisthocline in first whorls, and about

12 orthocline ribs in the last whorl.

Spiral sculpture formed by very fine

cords. Microsculpture (Figs. 49, 50)

formed by thick spiral threads with

sparse very small pits, with 5-10 very

thin threads between each two thick

ones. The microsculpture almost disap-

pears on the ribs.

Aperture D-shaped, small; inner lip

thick; columellar side weakly concave;

anterior channel very shallow; outer lip

thick, strongly opisthocline, with wide

external varix, with 5-6 parallel lines on

the inner part.

Habitat: The material studied was

obtained in sand sediments from 1 to 85

m of depth.

Distribution: Only known from Sal

and Boa Vista Islands (Fig. 159).

Remarks: S. angularis n. sp. has a sca-

lariform profile like S. obesa n. sp. and S.

gradata n. sp. (see below), but S. obesa is

larger, wider and more solid, and S.

gradata is more slender and has a differ-

ent protoconch sculpture.

Schwartziella (Schwartziella) luisi n. sp. (Figs. 51-55, 153-155, 159)

Type material: Holotype (Fig. 51) 1 s of 2.0 x 1.0 mm and 2 paratypes, 2 s, from Derrubado, Boa

Vista Island, Cape Verde Archipelago, 2-4 m (MNCN 15.05/31722). Paratypes: 1 sin each of MNHN,

AMNH, DBUA, NNM (58032), and 29 s in CER, all from the type locality.

Other material studied: Sal: 1 s, Praia do Cascalho, 1 m; 1 s, 3j, Palmeira, 8 m; 3 s, Mordeira, 4

m; 1j, CANCAP Sta. 7.100, off Palmeira, 16? 45” N, 23” 01 W, 354 m (30-VII1-1986) (NNM); 1 sp,

12 s, 2 f, Rabo do Junco, 2-5 m; 1 sp, 34 s, Regona, 2-10 m, 1 s, Palhona, 1 m,; 1 sp, 8 s, 3 j, Mor-

deira, 4 m. Boa Vista: 18 s, 35 j, 16 f£, Derrubado, 4 m,; 2 sp, 4 s, Baia Teodora, 6 m; 1 sp, 7 s, llhéu

de Sal Rei, 5 m; 52 s, 18 j, 8 f, Porto da Cruz, 6 m, 35 s, 7 f, 1 j, Sal Rei, 8 m, 9 s, Porto Ferreira, 6

m; 3 s, 2 j, Rife de Chaves, 6 m; 34 s, 2 j, 2 f, Morro de Areia, 4 m; 5 s, 1 j, Baijos de Joáo Valente,

23 m.

Etymology: The specific name is dedicated to Luis Murillo, malacologist and Secretary of the Socie-

dad Española de Malacología, by his contribution to the development of Malacology in Spain.

(Right page) Figures 51-55: Schwartziella (Schwartziella) luisi n. sp. 51: holotype, Derrubado,

Boa Vista (MNCN 15.05/31722); 52: protoconch of a paratype, Derrubado (CER); 53: proto-

conch microsculpture of the same paratype; 54-55: teleoconch microsculpture of the same

paratype.

(Página derecha) Figuras 51-55: Schwartziella (Schwartziella) luisi spec. nov. 51: holotipo, Derru-

bado, Boa Vista (MNCN 15.05/31722); 52: protoconcha de un paratipo, Derrubado (CER); 53:

microescultura de la protoconcha del mismo paratipo; 54-55: microescultura de la teloconcha del mismo

paratipo.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Iberus, 18 (1), 2000

Description: Shell (Fig. 51) length up

to 3.0 mm, maximum width 1.3 mm, not

very solid, elongate-conic.

Protoconch (Fig. 52) of a little more

than 1 whorl and about 280 um of

maximum diameter, of non-planktotro-

phic type, with two spiral angulations,

one on the upper part and other a little

below; transition to teleoconch abrupt.

Microsculpture (Fig. 53) formed by very

small and irregular pits.

Teleoconch of 4 strongly convex

whorls, not angulated below sutures;

last whorl strongly convex; suture

shallow but evident. Colour whitish.

Axial sculpture consisting of promi-

nent, sharp, narrow, slightly curved,

weakly opisthocline spaced axial ribs,

which are continued from whorl to

whorl. Spiral sculpture formed by

prominent, widely spaced, very fine

cords, visible under low magnification.

Microsculpture (Figs. 54, 55) formed by

small pits on the spiral cords, and 15-20

very fine threads between two cords.

Aperture D-shaped, medium sized;

inner lip thick; columellar side weakly

concave; anterior channel almost

absent; outer lip opisthocline, with

thick external varix; with several par-

allel lines towards the inner part of the

aperture.

Operculum (Figs. 153-155) translu-

cent, thin, paucispiral with the nucleus

very eccentric and without any promi-

nent peg on the inner side.

Habitat: The material studied was

obtained in sand sediments between 2-

10 m and at 354 m depth.

Distribution: Only known from Sal

and Boa Vista Islands (Fig. 159).

Remarks: S. luisi n. sp. is smaller

than any of the previously described

species. Among the smallest of those, S.

angularis n. sp. differs by having a

subsutural angulation, and S. typica n.

sp. has a different protoconch and tele-

oconch microsculpture. The differences

with the other small species are discus-

sed below.

Schwartziella (Schwartziella) minima n. sp. (Figs. 56-60, 139, 159)

Type material: Holotype (Fig. 56) 1 s of 2.2 x 1.0 mm and 1 paratype, 1 s, from Regona, Sal Island,

Cape Verde Archipelago, 2-4 m (MNCN 15.05/31711). Paratypes: 1 s in each of MNHN, AMNH,

DBUA, NNM (58018), and 30 s in CER, all them from the type locality.

Other material studied: Sal: 5 s, 1 f, Palhona, 1 m, 1 s, Punta do Cascalho, 2 m; 2 sp, 2 s, Rabo de

Junco, 5 m; 8 s, Palmeira, 5 m, 4 s, Pesqueiro do Aire, 1 m; 3 s, 2 f, Mordeira, 4 m; 3 s, CANCAP Sta.

7.100, off Palmeira, 16? 45' N, 23? 01” W, 354 m (30-VIII1-1986) (NN M); 1 s, CANCAP Sta. 7.109, off

Palmeira, 16? 46' N, 22? 59% W, 31 m (31-VII1-1986) (NNM). Boa Vista: 1 s, Ilhéu de Sal Rei; 2 f, Derru-

bado, 4 m.

Etymology: The specific name alludes to the small size of the shell.

Description: Shell (Fig. 56) length up

to 2.8 mm, maximum width 1.3 mm,

relatively solid, elongate-conic.

Protoconch (Fig. 57) of 1 whorl and

320 um of maximum diameter, of non-

planktotrophic type, without spiral

sculpture. Microsculpture (Fig. 58)

formed by irregular flat prominences

with some circular pits on an undulate

surface.

Teleoconch of about 4 strongly con-

vex whorls, not angulated below sutures,

(Right page) Figures 56-60: Schwartziella (Schwartziella) minima n. sp. 56: holotype, Regona, Sal

(MNCN 15.05/31711); 57: protoconch of a shell from Regona, Sal; 58: protoconch microsculp-

ture of a paratype, Regona (CER); 59-60: teleoconch microsculpture of the same paratype.

(Página derecha) Figuras 56-60: Schwartziella (Schwartziella) minima spec. nov. 56: holotipo,

Regona, Sal (MNCN 15.05/31711); 57: protoconcha de una concha de Regona, Sal; 58: microescul-

tura de la protoconcha de un paratipo, Regona (CER); 59-60: microescultura de la teloconcha del

mismo paratipo.

he Cape Verde Archipelago

ininae In t

The subfamily Risso

wn 00S

ROLÁN AND LUQUE

Iberus, 18 (1), 2000

last whorl convex; suture shallow, but

evident. Colour whitish.

Axial sculpture consisting of promi-

nent, rounded, narrow, distantly spaced

axial ribs, almost orthocline in last

whorl and opisthocline in previous

ones. Spiral sculpture formed by very

fine cords. Microsculpture (Figs. 59, 60)

formed by very fine and irregular spiral

threads.

Aperture D-shaped, medium sized,

inner lip thick; columellar side weakly

concave; anterior channel almost

absent; outer lip strongly opisthocline

with thick varix; with about five para-

llel lines towards the inner part of the

aperture.

Habitat: The material studied was

obtained in sand sediments between 1

and 354 m.

Distribution: Only known from Sal

and Boa Vista Islands (Fig. 159).

Remarks: S. minima n. sp. is smaller

than any of the previously described

species. Among the small species, S.

typica n. sp. has a larger shell, different

spiral microsculpture, and the irregular

flat prominences which form the

microsculpture of the protoconch of

smaller size and without pits; S. angula-

ris n. sp. has a subsutural angulation; S.

luisi m. sp. has a different teleoconch and

protoconch microsculpture.

One shell (Fig. 139) found in Calhau,

Sáo Vicente, is similar in size to this

species, but it has more curved axial ribs,

a slightly different teleoconch micros-

culpture and spiral sculpture in the pro-

toconch. We prefer not to describe it as

new awaiting further material.

Schwartziella (Schwartziella) fulgida n. sp. (Figs. 61-65, 165)

Type material: Holotype (Fig. 61) 1 s 0f2.4 x 1.1 mm, from Furna, Brava Island, Cape Verde Archi-

pelago, 30m (MNCN 15.05/31706). Paratypes: 1 sin each of MNHN, AMNH, DBUA, NNM (58007),

and 6 sin CER, all from the type locality.

Other material studied: Brava: 4 f, Furna, 30 m; 2 s, 1j, 1 f, Pedrinha, 6 m; 5 s, Ilhéus do Rombo, 6

m; 18 c, 1 f, Porto do Anciáo, 3 m. Santiago; 1 s, 6 j, 5 f Prainha, 5 m; 7 s, 1 f, Tarrafal, 4 m; 2 f,

CANCAP Sta. 6.004, 14? 54” N, 23? 30” W, 58-63 m (5-VI-1982) (NN M); 4 s, CANCAP Sta. 6.010, 14?

52 N, 23? 30” W, 310 m (5-VI-1982) (NNM); 4 s, 1 f, CANCAP Sta. 6.015, S of the island, 14? 53' N,

23? 30' W, 150 m (5-VI-1982) (NN M); 1 s, CANCAP Sta. 6.024, 15? 00' N, 23? 44” W, 540 m (7-VI-

1982) (NNM); 2 s, 3 f, CANCAP Sta. 7.008, Ponta Grande da Cidade, 14” 54' N, 23? 38" W, 700 m

(20-VIIT-1986) (NN M) 1 s, 2j, 3 f, Cidade Velha, 6 m. Fogo: 2 s, CANCAP Sta. 6.040, 14? 55' N, 24?

31' W, 38-55 m (9-VI-1982) (NNM); 1 s, CANCAP Sta. 6.041. W of the island, 14? 55 N, 24? 31" W,

60 m (9-V1-82) (NNM).

Etymology: The specific name alludes to the apparently smooth shell, which gives it a shining appe-

arance.

Description: Shell (Fig. 61) length up

to 2.8 mm, maximum width 1.3 mm,

solid, elongate-conic, shining.

Protoconch (Fig. 62) of 1 whorl and

300 um of maximum diameter, of non-

planktotrophic type, with only a very

slight angulation at the upper part; tran-

sition to teleoconch abrupt. Microsculp-

ture (Fig. 63) shows irregular flat promi-

nences with some circular pits on an

undulate surface, but the studied proto-

conchs were poorly preserved.

Teleoconch of about 4 whorls, regu-

larly convex, not angulated below

(Right page) Figures 61-65: Schwartziella (Schwartziella) fulgida n. sp. 61: holotype, Furna, Brava

(MNCN 15.05/31706); 62: protoconch of the holotype; 63: protoconch microsculpture of the

holotype; 64-65: teleoconch microsculpture of the holotype.

(Página derecha) Figuras 61-65: Schwartziella (Schwartziella) fulgida spec. nov. 61: holotipo, Furna,

Brava (MNCN 15.05/31706); 62: protoconcha del holotipo; 63: microescultura de la protoconcha del

holotipo; 64-65: microescultura de la teloconcha del holotipo.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Iberus, 18 (1), 2000

sutures; sutures shallow but evident.

Colour whitish.

Axial sculpture consisting of promi-

nent, rounded, wide, spaced axial ribs,

slightly opisthocline in last whorl and a

little more on previous ones; about 14 in

last whorl. At low magnification, appar-

ently there is no spiral sculpture, but there

is a fine microsculpture (Figs. 64, 65) of

very fine threads on the interspaces be-

tween ribs, which disappears on the ribs.

Aperture D-shaped, medium sized;

inner lip thick; columellar side weakly

concave; anterior channel almost absent;

outer lip opisthocline with a thick exter-

nal varix. Peristome prominent in large

specimens, with several parallel lines

towards the inner part of the aperture.

Habitat: Sand sediments between 4

and 700 m.

Distribution: Only known from Fogo,

Brava and Santiago (Fig. 165).

Remarks: S. fulgida n. sp. differs from

the other small species of Schwartziella

previously described: S. minima n. sp. is

smaller, with smaller and less numerous

axial ribs and it has more evident spiral

sculpture; S. luisi n. sp. also has a more

evident spiral sculpture and the proto-

conch microsculpture formed by very

small pits; S. angularis n. sp. has a

subsutural angulation; S. typica n. sp.

has a larger shell, and a fairly evident

and different spiral microsculpture,

uniform in S. fulgida and with two diffe-

rent alternate zones in S. typica.

Schwartziella (Schwartziella) depressa n. sp. (Figs. 66-70, 164)

Type material: Holotype (Fig. 66) 1 s of 3.6 x 1.5 mm and 2 paratypes, 2 s, from Mordeira Bay, Sal

Island, Cape Verde Archipelago, 4 m (MNCN 15.05/31704). Paratypes: 1 s in each of MNHN,

AMNH, DBUA, NNM (58003), and 15 s, 2 fin CER, all from the type locality.

Other material studied: Sal: 1 s, Mordeira, 4 m; 24 s, 4 f, Palmeira, 8 m; 12 s, Regona, 1-3 m; 1 s,

Algodoeiro, 4 m; 10 s, Rabo do Junco, 2 m; Boa Vista: 1 sp, 14 s, 3 f Sal Rei, 5 m; 2 s, 1 f, Ilhéu de

Sal Rei, 3 m; 10 s, 7 f, Porto da Cruz, 6 m; 9 s, 3 j, 9 f, Derrubado, 4 m; 5 s, Rife de Chaves, 8 m; 1 s,

Baijos de Joáo Valente, 23 m. Sáo Nicolau: 13 s, 7 £, CANCAP Sta. 7.128, 16” 33" N, 24? 17 W, 400

m (2-IX-1986) (NNM); 21 s, 11 f, Sáo Jorge Bay, CANCAP Sta. 7.129, 16” 33' N, 24” 16' W, 405 m (2-

1X-1986) (NNM).

Etymology: The specific name alludes to the unusual subsutural depression of the shell.

Description: Shell (Fig. 66) length up

to 4.0 mm, maximum width 1.6 mm,

relatively solid, elongate-conic.

Protoconch (Fig. 67) of 1 whorl and

about 300 ym of maximum diameter, of

non-planktotrophic type, with a promi-

nent spiral cord on its upper part; tran-

sition to teleoconch abrupt. Microsculp-

ture (Fig. 68) formed by irregular flat

Teleoconch of 5 whorls, strongly

convex, last whorl weakly convex;

suture shallow, with a slight subsutural

depression. Colour whitish.

Axial sculpture consisting of promi-

nent, rounded, wide, slightly opistho-

cline axial ribs, with interspaces of

similar size, a little reduced in the

subsutural depression specially in the

last two whorls; about ten axial ribs in

last whorl. At low magnification, appa-

prominences with small pits on an

undulous surface.

(Right page) Figures 66-70: Schwartziella (Schwartziella) depressa n. sp. 66: holotype, Mordeira, Sal

(MNCN 15.05/31704); 67: protoconch of the holotype; 68: protoconch microsculpture of the

holotype; 69: teleoconch microsculpture of the holotype; 70: teleoconch microsculpture of a shell

from Mordeira, Sal.

(Página derecha) Figuras 66-67: Schwartziella (Schwartziella) depressa spec. nov. 66: holotipo, Mor-

deira, Sal (MNCN 15.05/31704); 67: protoconcha del holotipo; 68: microescultura de la protoconcha

del holotipo; 69: microescultura de la teloconcha del holotipo; 70: microescultura de la teloconcha de

una concha de Mordeira, Sal.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Iberus, 18 (1), 2000

rently there is no spiral sculpture, but

there is a fine microsculpture of bands

with small pits and 1-3 fine and irregu-

lar threads between each two bands on

the interspaces between ribs, which

disappear on the ribs (Figs. 69, 70).

Aperture D-shaped, relatively small;

inner lip thick; columellar side weakly

concave; anterior channel shallow; outer

lip with thick external varix; with about

6 lines towards the inner part of the

aperture.

Habitat: The material studied was

obtained from sand sediments between

1-8 and to 400-405 m.

Distribution: Only known from Sal,

Boa Vista and Sáo Nicolau Islands (Fig.

164).

Remarks: The shells from deep water

off Sáo Nicolau are similar in size and

sculpture to those from shallow waters

of Sal and Boa Vista, but the subsutural

depression is less evident.

S. depressa n. sp. differs from $. simi-

liter n. sp. by its smaller shell, less mar-

ked spiral sculpture, narrower interspa-

ces between axial ribs, and different mi-

crosculpture of the protoconch and

teleoconch. S. typica n. sp. has a wider

shell, lacking subsutural depression. S.

angularis and S. luisi have more evident

spiral sculpture. S. minima and S. fulgida

n. sp. have smaller shells without subsu-

tural depression; S. minima also has a

more evident spiral sculpture and lacks

spiral sculpture on the protoconch.

Schwartziella (Schwartziella) gradata n. sp. (Figs. 71-75, 161)

Type material: Holotype (Fig. 71) 1 s of 3.1x 1.3 mm and 1 paratype, 1 s, Furna, Brava Island, Cape

Verde Archipelago, 30 m (MNCN 15.05/31708); 1 paratype in NNM (58010) from the type locality.

Other material studied: Brava: 2 s, 3 j, Pedrinha, 5 m.

Etymology: The specific name alludes to the scalariform profile of the shell.

Description: Shell (Figs. 71, 72) length

up to 3.1 mm, maximum width 1.3 mm,

slightly solid, elongate-conic and rela-

tively narrow.

Protoconch (Fig. 73, 74) of 1 whorl

and 270 ym of maximum diameter, of

non-planktotrophic type, with an apical

spiral cord and two narrower cords at

both sides, and small and irregular axial

threads on the interspaces between

cords; transition to teleoconch abrupt.

The rest of the surface of the protoconch

have irregular flat prominences with

small pits on a smooth surface.

Teleoconch of about 5 whorls weakly

convex, with a strong subsutural angu-

lation. Colour whitish.

Axial sculpture consisting of promi-

nent, sharp, narrow, slightly opisthocline,

distantly spaced axial ribs, about 12 in last

whorl. Spiral sculpture formed by very

fine cords. Microsculpture (Fig. 75) for-

med by very numerous spiral threads,

which disappear on the cords and contin-

ue on the ribs.

Aperture D-shaped, small; inner lip

thick; columellar side weakly concave;

anterior channel absent, with a depression

in the zone; outer lip opisthocline with

thick external varix, with 5-6 parallel lines

towards the inner part of the aperture.

Habitat: The material studied was

obtained from sand sediments between

5 and 30 m.

(Right page) Figures 71-75: Schwartziella (Schwartziella) gradata n. sp. 71: holotype, Furna, Brava

(MNCN 15.05/31708); 72: paratype, Furna, Brava (NNM 58010); 73: protoconch of a paratype,

Furna (CER); 74: detail of the protoconch of the same paratype: 75: teleoconch microsculpture of

the same paratype.

(Página derecha) Figuras 71-75: Schwartziella (Schwartziella) gradata spec. nov. 71: holotipo, Furna,

Brava (MNCN 15.05/31708); 72: paratipo, Furna, Brava (NNM 58010); 73: protoconcha de un

paratipo, Furna (CER); 74: detalle de la protoconcha del mismo paratipo: 75: microescultura de la telo-

concha del mismo paratipo.

wr 001

in the Cape Verd:

ininae

wur ]

The subfamily Risso

ROLÁN AND LUQUE

Iberus, 18 (1), 2000

Distribution: Only known from Brava

Island (Fig. 161).

Remarks: S. gradata n. sp. differs from

any other species of Schwartziella of the

Cape Verde Islands by the protoconch

sculpture. From other species with

subsutural angulation, the sympatric $.

obesa n. sp. is larger, and S. angularis is

smaller and relatively wider

(length /width ratio 2.12-2.23, and 2,32-

2,44 in S. gradata), with only a spiral an-

gulation in the upper part of the proto-

conch, instead of three spiral cords and

irregular axial threads in S. gradata.

Schwartziella (Schwartziella) pavita n. sp. (Figs. 76-79, 162)

Type material: Holotype (Fig. 76) 1 s of 3.3 x 1.3 mm and 1 paratype, 1 s, from Sal Rei, Boa Vista

Island, Cape Verde Archipelago, 5 m (MNCN 15.05/31714). Other paratypes: 1 s (NNM 58022) 1

s (MNHN) and 1 s (CER), all from the type locality; 11 s from Porto da Cruz, north of Sal Rei (CER).

Other material studied: (in poor condition) Boa Vista: 2 s, 1 f, Tlhéu de Sal Rei, 6 m; 1 s, Sal Rei, 6

m; 1 s, Baia Teodora, 4 m; 1 s, Derrubado, 4 m. Sal: 1 s, Mordeira, 4 m. Maio: 1 s, Pau Seco, 5 m.

Etymology: The specific name derives from the Latin verb pavio which means “to level” by the

depressed axial ribs of the shell.

Description: Shell (Fig. 76) length up

to 3.3 mm, maximum width 1.3 mm,

relatively solid, elongate-conic.

Protoconch (Fig. 77) of 1 whorl,

about 300 ym of maximum diameter, in

poor condition in all the material

studied, of non-planktotrophic type,

apparently without spiral sculpture.

Teleoconch of about 5 whorls, almost

flat-sided; last whorl weakly convex;

suture shallow. Colour whitish.

Axial sculpture consisting of

rounded, not prominent, wide, slightly

opisthocline, distantly spaced axial ribs,

about 12 in last whorl. Spiral sculpture

of fine spiral cords densely spaced.

Microsculpture (Figs. 78, 79) formed by

one or two very fine irregular spiral

threads between each cord.

Aperture D-shaped, small, inner lip

thick; columellar side weakly concave;

anterior channel almost absent; outer lip

opisthocline with thick external varix,

with about 3-4 parallel lines.

Habitat: The material studied was

obtained from sand sediments between

4 and 6 m.

Distribution: Only known from

Sal, Boa Vista and Maio Islands (Fig.

162).

Remarks: S. pavita n. sp. differs from

other Cape Verde species of Schwartzie-

lla by the very depressed axial ribs and

the microsculpture with densely

spaced spiral cords and only one-two

threads between cords. The sympatric

S. sanmartini n. sp. also has non-promi-

nent axial ribs, but it has a larger and

wider shell, with a teleoconch micro-

sculpture formed only by very fine

threads. S. similiter n. sp. and S. typica

n. sp. have the axial ribs more promi-

nent and a different teleoconch micro-

sculpture. S. minima n. sp. and S.

fulgida n. sp. are smaller and both have

more prominent axial ribs and micros-

culpture formed almost exclusively by

very fine threads.

(Right page) Figures 76-79: Schwartziella (Schwartziella) pavita n. sp. 76: holotype, Sal Rei,

Boa Vista (MNCN 15.05/31714); 77: protoconch of a shell from Sal Rei, Boavista; 78: teleo-

conch microsculpture of a shell from Sal Rei, Boavista; 79: teleoconch microsculpture of the

holotype.

(Página derecha) Figuras 76-79: Schwartziella (Schwartziella) pavita spec. nov. 76: holotipo, Sal. Rer,

Boa Vista (MNCN 15.05/31714); 77: protoconcha de una concha de Sal Rei, Boavista; 78, microes-

cultura de la teloconcha de una concha de Sal Rei, Boavista; 79: microescultura de la teloconcha del

holotipo.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

S ¿ -

di at, 2 gut “yz rg

An a AA

Iberus, 18 (1), 2000

Schwartziella (Schwartziella) cancapae n. sp. (Figs. 80-84, 165)

Type material: Holotype (Fig. 80) 1 s of 3.8 x 1.5 mm (NNM 58001) and 6 paratypes (5 f), CANCAP

Sta. 6.009, S of Santiago, Cape Verde Archipelago, 14? 54' N, 23? 30” W, 175 m (5-VI-1982) (NNM

58002). Other paratypes: 1 s (MNCN 15.05/31702) and 1 s (CER), Praia, Santiago, 10 m; 9 s, CANCAP

Sta. 6.006, S of Santiago, 14” 54” N, 23” 30” W, 150 m (5-VI-1982) (NN M 59409).

Other material studied: Santiago: 1 s, 1 f£, CANCAP Sta. 6.005, S of the island, 14? 54' N, 23? 30" W,

175 m (5-VI-1982) (NNM); 3 f, CANCAP Sta. 6.010, 14? 52” N, 23? 30” W, 310 m (5-VI-1982) (NN M);

2 s, 18 f, CANCAP Sta. 6.015, S of the island, 14? 53' N, 23? 30” W, 150 m (5-VI-1982) (NNM), 1 s, 3

j, CANCAP Sta. 6.024, 15 00' N, 23? 44” W, 540 m (7-VI-1982) (NN M); 1 s, 4 £, CANCAP Sta. 7.004,

142 54' N, 23? 38" W, 320 m (21-VIII-1986) (NN M); 1 s, CANCAP Sta. 7.005, 14? 54” N, 23? 38' W,

235 m (21-VIII-1986) (NN M); 3 s, 5 f, CANCAP Sta. 7.007, 14? 54' N, 23? 38" W, 420 m (20-VI-1986)

(NNM) 1 s, 7 j CANCAP Sta. 7.008, 14” 54” N, 23” 38" W, 700 m (20-VIII-1986) (NN M); 1 s, 7 j,

CANCAP Sta. 7.014, Ponta Grande da Cidade, 14? 54' N, 23? 38' W, 450-600 m (21-VIII-1986) (NN M).

Fogo: 1 j, Sáo Felipe, 20 m. Brava: 2 s, 1 f, Furna, 30 m.

Etymology: The specific name alludes to the CANCAP expeditions in which has been collected

most of the material of this species and part of the material studied in this paper.

Description: Shell (Fig. 80) length up

to 4.0 mm, maximum width 1.6 mm, not

solid, elongate-conic.

Protoconch (Fig. 81) of 1 whorl and

400 ym of maximum diameter, of non-

planktotrophic type, without spiral

sculpture; transition to teleoconch

abrupt. Microsculpture (Fig. 82) formed

by a rough surface with many small

pits.

Teleoconch of about 5 whorls,

weakly convex; last whorl weakly

convex; suture shallow but evident.

Colour whitish.

Axial sculpture consisting of promi-

nent, rounded, narrow, slightly opistho-

cline, distantly spaced axial ribs, 13 in

last whorl; the ribs are continued from

whorl to whorl, except in first whorls.

Spiral sculpture appreciable with diffi-

culty at low magnification. Microsculp-

ture (Figs. 83, 84) of threads with a few

parallel lines of minute pits, and more

separated, sometimes paired irregularly

interrupted striae.

Aperture D-shaped, medium-small,

inner lip thick; columellar side weakly

concave; anterior channel almost absent;

outer lip opisthocline with thick exter-

nal varix, peristome narrow with a few

parallel lines.

Habitat: Some shells of this species

were collected in relatively shallow

water (20-30 m), but most of material

comes from 60-200 m in depth in sand

and shell gravel bottom.

Distribution: Only known from San-

tiago, Fogo and Brava (Fig. 165).

Remarks: S. cancapae n. sp. differs

from most of the species described

before by its protoconch lacking spiral

sculpture and larger than most of Cape

Verde Schwartziella species. The more

similar species are: S. depressa n. sp.,

with a more evident subsutural depres-

sion, and an evident spiral cord and dif-

ferent microsculpture in the protoconch;

S. fulgida n. sp. is smaller, almost

smooth, with different microsculpture in

the teleoconch and protoconch, S. typica

n. sp. is smaller and has different proto-

conch microsculpture, and S. similiter n.

sp. is larger, wider, and with more

evident spiral sculpture. S. cancapae is

quite similar to S. africana, but the latter

species has strongly opisthocline axial

(Right page) Figures 80-84: Schwartziella (Schwartziella) cancapae n. sp. 80: holotype, Santiago

(NNM 58001); 81: protoconch of the holotype; 82: protoconch microsculpture of the holotype;

83-84: teleoconch microsculpture of the holotype.

(Página derecha) Figuras 80-84: Schwartziella (Schwartziella) cancapae spec. nov. 80: holotipo, San-

tiago (NNM 58001); 81: protoconcha del holotipo; 82: microescultura de la protoconcha del holotipo;

83-84: microescultura de la teloconcha del holotipo.

SÓ

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Iberus, 18 (1), 2000

ribs, different protoconch and teleo-

conch microsculpture (see GOFAS, 1999).

Previous records or S. africana from the

Cape Verde Islands (MARCHE-MAR-

CHAD, 1958; SAUNDERS, 1977) could be

referred to this species.

Schwartziella (Schwartziella) puncticulata n. sp. (Figs. 85-87, 140-141, 166)

Type material: Holotype (Fig. 85, NNM 58023) 1 s of 4.0 x 1.6 mm and 2 paratypes (NNM 58024),

2 f, CANCAP Sta. 7.119, S of Ilhéu Razo, 16" 36' N, 24? 36" W, Cape Verde Archipelago, 140-160 m

(1-D1986). Paratypes: 1 s (MNCN 15.05/31715) and 1 s (CER), both from the type locality; 7 s, 11

f, CANCAP Sta. 7.028, Ilhéu de Cima, 14? 57' N, 24? 39 W, 225 m (23-VIII-1986) (NN M 59414).

Other material studied: Sal: 6 £, CANCAP Sta. 7.100, off Palmeira, 16* 45' N, 23* 01” W, 354 m (30-

VITI-1986) (NN M). Maio: 6 f, CANCAP Sta. 7.050, 15” 06' N, 23” 14” W, 380 m (25-VIII-1986) (NN M).

Santiago: 1 s, 12j, Cidade Velha, 6 m; 10 j, Prainha, 5 m; 6 s, 3 f, CANCAP Sta. 6.095, 16” 35' N, 24?

37' W, 930 m (15-VI-1982) (NNM). Fogo: 9 s, 6 f, CANCAP Sta. 6.041, W of the island, 14? 55' N, 24?

31" W, 60 m (9-VI-82) (NNM). Brava: 12 j, Pedrinha, 6 m; 7 j, Furna, 30 m. Ilhéu de Cima: 2 s, 5 f,

CANCAP Sta. 7.037, 14” 57' N, 24” 38" W, 350-385 m (24-VIII-1986) (NNM); 1 s, 2 CANCAP Sta.

7.038, 14? 57" N, 24? 38" W, 410-460 m (24-VII1-1986) (NN M); 1 s, 6j, CANCAP Sta. 7.031, 14? 57' N,

24? 38" W, 75 m (23-VIII-1986) (NN M); 2 s, CANCAP Sta. 7.037, 14? 57' N, 24? 38" W, 385-350 m (24-

VIII-1986) (NNM). Sáo Vicente: 1 s, CANCAP Sta. 6.147, 16? 48” N, 25” 06" W, 99 m (20-VI-1982)

(NNM) 4 s, 15, 4£ CANCAP Sta. 6.149, 16 47' N, 25 06" W, 293 m (20-VI-1982) (NN M). Ilhéu Razo:

1 s, 5j, CANCAP Sta. 6.093, 16? 36" N, 24” 37” W, 400-430 m (15-VI-1982) (NNM); 3 s, 10 j, 9 f,

CANCAP Sta. 7.119, S of Ilhéu Razo, 16” 36' N, 24? 36" W; 140-160 m (1-1X-1986); 7.s, 21 f CANCAP

Sta. 7.121, 16” 36' N, 24? 37" W, 200-230 m (1-DX-1986) (NN M). Santa Luzia: 9 f, CANCAP Sta. 6.103,

16? 43' N, 24? 46' W, 102 m (16-VI-1982) (NNM); 1 s, 6 f, CANCAP Sta. 6.105, SSW of the island, 16

43' N, 24? 47” W, 204 m (16-VI-1982) (NNM). Sáo Nicolau: 1 s, 3j, CANCAP Sta. 6.085, 16? 34” N,

24 22 W, 100 m (14-VI-1982) (NN M).

Etymology: The specific name alludes to the microsculpture of this species, formed by undulated

rows of small pits.

Description: Shell (Fig. 85) length up

to 4.5 mm, maximum width 1.7 mm,

relatively solid, elongate-conic.

Protoconch (Fig. 86) of 1 whorl and 360

ym of maximum diameter, of non- plank-

totrophic type; no spiral sculpture; tran-

sition to teleoconch abrupt. Microsculp-

ture formed by irregular flat prominences

on a smooth surface, with some pits.

Teleoconch of 4-5 whorls, strongly

convex, and more convex in the subsu-

tural part of the ribs; last whorl regu-

larly convex; suture a little deep. Colour

whitish.

Axial sculpture consisting of promi-

nent, rounded, narrow, a little opistho-

cline distantly spaced axial ribs, which

are continued from whorl to whorl.

Spiral sculpture not visible at low mag-

nification. Microsculpture (Fig. 87)

formed by very fine, densely packed

undulated threads with intermediate

rows of small pits.

Aperture D-shaped, small; inner lip

thick; columellar side weakly concave;

anterior channel shallow; outer lip

strongly opisthocline with a very thick

external varix, peristome with a promi-

nent edge and some parallel lines

towards the inner part.

Habitat: Sand sediments, between 6

and 930 m.

Distribution: Sal, Maio, Santiago,

Fogo, Brava, Ilhéu de Cima, Sáo Vicente,

Santa Luzia, llhéu Razo and Sáo

Nicolau (Fig. 166). Probably this species

is present in all the archipelago, but it is

frequent in some islands and uncom-

mon in others.

Remarks: S. puncticulata n. sp. differs

from the following similar species: S.

similiter n. sp. has no subsutural shoul-

der and a different protoconch and

teleoconch microsculpture; S. typica n.

sp. is smaller, less elongate, uniformly

convex and with different teleoconch

microsculpture, S. fulgida n. sp. is also

smaller, with the whorls less convex,

and the teleoconch microsculpture more

lineal; S. depressa n. sp. is smaller, with a

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Figures 85-87: Schwartziella (Schwartziella) puncticulata n. sp. 85: holotype, llhéu Razo (NNM

58023); 86: protoconch of a paratype, Ilhéu Razo (NNM 58024); 87: teleoconch microsculpture

of the holotype.

Figuras 85-87: Schwartziella (Schwartziella) puncticulata spec. nov. 85: holotipo, llhéu Razo (NNM

58023); 86: protoconcha de un paratipo, llhéu Razo (NNM 58024); 87: microescultura de la telocon-

cha del holotipo.

Iberus, 18 (1), 2000

spiral cord on the protoconch and a

subsutural depression on the teleo-

conch; S. cancapae n. sp. has a more

regular curvature of the whorls, a larger

protoconch, and a different protoconch

and teleoconch microsculpture.

One shell (Figs. 140, 141) found in

Sao Vicente, CANCAP Sta. 6.145, 167 48'

N, 25? 06” W) is smaller, stouter and the

axial ribs are less convex, but we provi-

sionally consider it conspecific since it

has identical microsculpture.

Schwartziella (Schwartziella) hoenselaari n. sp. (Figs. 88-92, 167)

Type material: Holotype (Fig. 88) 1 s 0f2.3 x 1.1 mm (NNM 58011) and 26 paratypes (NNM 58012),

26 s, S of Santiago, CANCAP Sta. 6.001, 14? 54' N, 23? 30 W, 15-20 m (4-VI-1982). Other paratypes:

13 s, CANCAP Sta. 6.014, 14? 54 N, 2329 W, 18 m (5-V1-1982) (NNM 59415); 1 s (MNCN 15.05 /31709)

and 1 s (CER), both from the type locality; 1 s, Prainha, 5 m (CER).

Other material studied: Santiago: 16 s, 2 j, Tarrafal, 4 m; 4 s, CANCAP Sta. 6.003, 14” 54' N, 23? 30"

W, 15-21 m (5-VI-1982) (NN M); 3 s, CANCAP Sta. 6.004, 14? 54” N, 23” 30” W, 58-63 m (5-VI-1982)

(NN M); 2 f, CANCAP Sta. 6.006, S of the island, 14? 54” N, 23? 30' W, 150 m (5-V1-1982) (NN M); 3

s, 2 CANCAP Sta. 6.007, 14? 54' N, 23? 30” W, 70-88 m (5-V1-1982) (NNM); 1 s, CANCAP Sta. 6.008,

14? 54” N, 23? 30” W, 120 m (5-VI-1982) (NNM); 6 s, 1 f, CANCAP Sta. 6.010, 14” 52” N, 23? 30' W,

310 m (5-VI-1982) (NN M); 6 s, 3) CANCAP Sta. 6.015, S of the island, 14? 53 N, 23 30' W, 150 m

(5-VI-1982) (NN M) 4 s, 1 f, CANCAP Sta. 6.024, 15 00' N, 23? 44' W, 540 m (7-V1-1982) (NN M),; 1

s, 1j, CANCAP Sta. 6.025, 15? 00” N, 23? 45” W, 728 m (7-VI-1982) (NNM); 2 s, CANCAP Sta. 7.005,

14? 54' N, 23? 38" W, 235 m (21-VIII-1986) (NNM); 2 f, CANCAP Sta. 7.007, 14? 54” N, 23? 38' W, 420

m (20-VI-1986) (NN M); 3 s, 2 f, Ponta Grande da Cidade, CANCAP Sta. 7.015, 14” 54” N, 23? 38" W,

450-600 m (21-VIII-1986) (NN My; 2 £, CANCAP Sta. 7.020, 14” 45” N, 23” 29 W, 0-2200 m (21-22-VIIT-

1986). Fogo: 2 s, 1 f, in front of Sáo Felipe, 20 m; 6 s, 3j, CANCAP Sta. 6.040, 14” 55' N, 24? 31" W,

38-55 m (9-VI-1982) (NNM). Ilhéu de Cima: 2 s, CANCAP Sta. 7.028, 14? 57" N, 24? 39 W, 225 m

(23-VIII-1986) (NN M); 1 s, CANCAP Sta. 7.037, 14? 57' N, 24? 38' W, 350-385 m (24-VII1-1986) (NNM).

Santa Luzia: 1 s, 1 f CANCAP Sta. 6.103, 16? 43” N, 24? 46” W, 102 m (16-VI-1982) (NN M); 2 s,

CANCAP Sta. 6.107, 16? 44' N, 24? 46' W, 50 m (16-VI-1982) (NNM). Ilhéu Razo: 1 s, CANCAP Sta.

7.120, 16” 36' N, 24? 37" W, 208 m (1-IX-1986); 1 s, CANCAP Sta. 7.121, 16? 36' N, 24” 37" W, 200-230

(1-IX-1986) (NNM). Sáo Nicolau: 2 s, 6 f, CANCAP Sta. 7.128, Sáo Jorge Bay, 16” 33" N, 24? 17' W,

400 m (2-1X-1986) (NNM); 6 s, 3 f, Sáo Jorge Bay, CANCAP Sta. 7.129, 16” 33" N, 24” 16' W, 405 m

(2-1X-1986) (NNM). Sal: 2 s, Palhona, 2 m.

Etymology: The specific name is dedicated to Hink J. Hoenselaar, Dutch malacologist who begun

the study of the Rissoininae material from the CANCAP Expeditions.

Description: Shell (Figs. 88, 89) length

up to 3.0 mm, maximum width 1.4 mm,

relatively solid, elongate-conic.

Protoconch (Fig. 90) of 1 whorl and

regular flat prominences with some pits

on an undulate surface.

Teleoconch of 4-5 whorls, clearly

convex, not angulated below suture; last

whorl convex; suture shallow, but

evident. Colour whitish.

Axial sculpture consisting of promi-

nent, rounded, narrow, almost orthocline

about 300 ym of maximum diameter, of

non-planktotrophic type, with no spiral

sculpture; transition to teleoconch abrupt.

Microsculpture (Fig. 92) formed by ir-

(Right page) Figures 88-92: Schwartziella (Schwartziella) hoenselaari n. sp. 88: holotype, Santiago

(NNM 58011); 89: paratype, Santiago (NNM 58012); 90: protoconch of a paratype, Santiago

(NNM 58012); 91: teleoconch microsculpture of a paratype, Santiago (NNM 58012); 92: proto-

conch microsculpture of a paratype, Santiago (NNM 58012).

(Página derecha) Figuras 88-92: Schwartziella (Schwartziella) hoenselaari spec. nov. 88: holotipo, San-

tiago (NNM 58011); 89: paratipo, Santiago (NNM 58012); 90: protoconcha del paratipo, Santiago

(NNM 58012); 91: microescultura de la teloconcha del paratipo, Santiago (NNM 58012); 92: micro-

escultura de la protoconcha del paratipo, Santiago (NNM 58012).

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Iberus, 18 (1), 2000

(slightly opisthocline on first whorls),

very distantly spaced axial ribs; being 9-

11 in the last whorl. Spiral sculpture

almost unappreciable at low magnifica-

tion. Microsculpture (Fig. 91) with zones

with 2-3 fine threads irregularly distrib-

uted, alternating with wider threads

with spiral rows of small pits.

Aperture D-shaped, small-medium

sized; inner lip thick; columellar side

weakly concave; anterior channel

absent; outer lip strongly opisthocline

with a very thick external varix; peris-

tome narrow with about 5 parallel lines

towards the inner part of the aperture.

Habitat: The material studied was

collected from 2 to 728 m of depth.

Distribution: Sal, Santiago, Fogo,

llhéu de Cima, Santa Luzia, Ilhéu Razo,

Sao Nicolau (Fig. 167).

Remarks: Schwartziella hoenselaari n.

sp. differs from most cf the Cape Verde

species of the genus by having less axial

ribs. It differs from the following species

with similar shape and protoconch

lacking spiral sculpture: S. puncticulata

n. sp. has a larger protoconch, a subsu-

tural curvature of the whorls, and the

teleoconch microsculpture lacks con-

tinuous threads; S. cancapae n. sp. has a

smaller protoconch, a more pointed

shell, with different protoconch and

teleoconch microsculpture; S. depressa n.

sp. and S. typica n. sp. have a similar

shape, but the protoconch of both

species has spiral sculpture and the

microsculpture of the teleoconch is dif-

ferent. Differences with the similar S.

paucicostata n. sp. are discussed under

remarks of this species.

Schwartziella (Schwartziella) paucicostata n. sp. (Figs. 93-96, 168)

Type material: Holotype (Fig. 93) 1s of 3.1 x 1.3 mm (NNM 58004), and 2 paratypes, 2 s (NNM

58005), CANCAP Sta. 7.105, off Palmeira, 16” 45” N, 23? 01” W, 123-142 m. Other paratypes: 1 f,

CANCAP Sta. 7.093, Ilhéu Razo, 16? 34” N, 33" 54” W, 42 m (29-VIII-1986) (NNM 59410); 1 s, 1 j,

CANCAP Sta. 7.095, Ilhéu Razo, 16? 34' N, 22? 53" W, 30-50 m (29-VIII-1986) (NNM 59411); 1s, 1]

(MNCN 15.05/34277), 3 s, 1] (CER) Pau Seco, Maio, 30 m.

Other material studied: Sal: 1 s, CANCAP Sta. 7.088, Ponta do Leme Velho, 16? 34' N, 22? 54 W,

59 m (29-VII1-1986) (NNM). Maio: 2 s, 1 f, CANCAP Sta. 7.050, 15* 06' N, 23” 14” W, 380 m (25-VIII-

1986) (NN M). Santiago: 1 s, CANCAP Sta. 6.017, 14? 53' N, 23? 30” W, 380 m (5-VI-1982).

Etymology: The specific name alludes to the low number of axial ribs of the shell.

Description: Shell (Fig. 93) length up

to 3.3 mm, maximum width 1.4 mm,

relatively solid, elongate-conic.

Protoconch (Fig. 94) of 1 whorl and

270 um of maximum diameter, of non-

planktotrophic type, without spiral

sculpture, smooth.

Teleoconch of about 5 whorls,

strongly convex; last whorl a little less

convex; suture shallow. Colour whitish.

Axial sculpture consisting of promi-

nent, sharp, very distantly spaced (4

times narrower than the interspaces)

axial ribs, almost orthocline in last

whorl and opisthocline in previous

whorls; only 6-7 axial ribs in last whorl.

Spiral sculpture almost unappreciable at

low magnification. Microsculpture (Figs.

95, 96) formed by bands of interrupted

and irregularly disposed fine spiral

threads and wider threads without

sculpture.

Aperture D-shaped, small; inner lip

thick; columellar side weakly concave;

anterior channel almost absent; outer lip

with thick external varix; peristome

narrow with several parallel lines

towards the inner part of the aperture.

Habitat: The material studied was

collected in sand bottom between 30

and 380 m.

Distribution: Sal, Maio, Santiago and

IIhéu Razo (Fig. 168).

Remarks: S. paucicostata n. sp. differs

from any other of the previously described

species of Cape Verde Schwartziella by its

smooth protoconch and the few and dis-

tant axial ribs. The similar S. hoenselaari n.

sp. has a wider protoconch with micro-

sculpture, and more numerous axial ribs.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

[e]

Figures 93-96: Schwartziella (Schwartziella) paucicostata n. sp. 93: holotype, Sal (NNM 58004);

94: protoconch of the holotype; 95-96: teleoconch microsculpture of the holotype.

Figuras 93-96: Schwartziella (Schwartziella) paucicostata spec. nov. 93: holotipo, Sal (NNM 58004);

94: protoconcha del holotipo; 95-96: microescultura de la teloconcha del holoipo.

Iberus, 18 (1), 2000

Schwartziella (Schwartziella) sulcostriata n. sp. (Figs. 97-101, 169)

Type material: Holotype (Fig. 91) 1 s of 3.1 x 1.2 mm (NNM 58033), and 3 paratypes, 3 s (NNM

58034), CANCAP Sta. 7.100, off Palmeira, 16” 45' N, 23? 01” W, 354 m (30-VII1-1986). Other paraty-

pes: 1 s, CANCAP Sta. 7.101, off Palmeira, Sal, 16” 45' N, 23” 01" W, 262-280 m (30-VI11-1986) (NNM

59421); 2 s, 2f, CANCAP Sta. 7.120, 16? 36' N, 24? 37 W, Ilhéu Razo, 208 m (1-1X-1986) (NNM 59422).

Other material studied: Sal: 2 f, CANCAP Sta. 7.110, 16? 46' N, 23? 02” W, 85 m (31-VIII-1986)

(NNM). Sáo Nicolau: 2 s, Sáo Jorge Bay, CANCAP Sta. 7.129, 16” 33' N, 24” 16" W, 405 m (2-1X-1986)

(NNM); 1 s, CANCAP Sta. 7.128, Sáo Jorge Bay, 16” 33" N, 2417" W, 400 m (2-IX-1986) (NNM).

Etymology: The specific name alludes to the microsculpture of the teleoconch.

Description: Shell (Fig. 97) length up

to 3.1 mm, maximum width 1.2 mm, not

very solid, strongly elongate-conic.

Protoconch (Fig. 98) of 1 whorl and

340 ym of maximum diameter, of non-

planktotrophic type, no spiral sculpture;

transition to teleoconch not very abrupt.

Microsculpture (Fig. 101) formed by

irregular flat prominences with some

pits on a smooth surface.

Teleoconch of 5 whorls, weakly

convex, not angulated below suture but

a little on suture; suture shallow, slightly

undulate. Colour whitish.

Axial sculpture consisting of not

prominent, rounded, narrow, slightly

opisthocline, distantly spaced axial ribs,

not always coincident from whorl to

whorl. Spiral sculpture appreciable with

difficulty at low magnification. Micro-

sculpture (Figs. 99, 100) only observable

among the spiral ribs and formed by

smooth spiral bands separated by

furrows with 2-3 spiral striae.

Aperture D-shaped, small; inner lip

thick; columellar side weakly concave;

anterior channel almost absent; outer lip

with thick external varix, peristoma

narrow and with a few parallel lines

towards the inner part of the aperture.

Habitat: Collected in sand sediments

between 85 and 405 m.

Distribution: Known from Sal, Ilhéu

Razo and Sáo Nicolau (Fig. 169). Prob-

ably it is present in all the islands of the

northern group.

Remarks: S. sulcostriata n. sp. is more

elongated than most of the previously

described species and it also differs

from any other by the spiral sculpture of

the teleoconch, formed by alternate

smooth bands and striated furrows. S.

similiter n. sp. has a spiral cord on the

top of the protoconch, rectilinear profile

and a different microsculpture of the

protoconch. S. pavita n. sp. has more

depressed axial ribs. S. depressa n. sp.

has a spiral cord on the top of the proto-

conch and a subsutural depression, S.

cancapae n. sp. has stronger axial ribs

and a wider protoconch with different

microsculpture.

Schwartziella (Schwartziella) gibbera n. sp. (Figs. 102-106, 170)

Type material: Holotype (Fig. 102) 1 s of 3.7 x 1.3 mm (NNM 58008), and 3 paratypes, 3 s (NNM

58009), CANCAP Sta. 7.079, W of Ilhéu de Sal Rei, Boa Vista, Cape Verde Archipelago, 16? 10' N,

23" 00' W, 60 m (28-VIII-1986). Other paratypes: 5 j, Ilhéu de Sal Rei, CANCAP Sta. 7.080, 16” 10'

N, 23? 01” W, 74 m (28-VIII-1986) (NNM 59412); 2 s (Fig. 103), 4 j, off Palmeira, Sal, CANCAP Sta.

(Right page) Figures 97-101: Schwartziella (Schwartziella) sulcostriata n. sp. 97: holotype, Pal-

meira, Sal (NNM 58033); 98: protoconch of the holotype; 99: detail of the spire of the holotype;

100: teleoconch microsculpture of the holotype; 101: protoconch microsculpture of the holotype.

(Página derecha) Figuras 97-101: Schwartziella (Schwartziella) sulcostriata spec. nov. 97: holotipo,

Palmeira, Sal (NNM 58033); 98: protoconcha del holotipo; 99: detalle de la espira del holotipo; 100:

microescultura de la teloconcha del holotipo; 101: microescultura de la protoconcha del holotipo.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Iberus, 18 (1), 2000

7.101, 16 45' N, 23? 01W, 262-280 m (30-VII1-1986) (NNM 59413); 1j, (MNCN 15.05/31707) and 1

j, both from Boa Vista (CER).

Other material studied: Boa Vista: 43 j, 25 f, CANCAP Sta. 6.066, 15* 53" N, 23” 00” W, 53 m (13-VI-

1982) (NNM). Sal: 2 j, off Palmeira, 16"46' N, 23? 01” W, 165 m (30-VIII-1986) (NNM). Santiago: 1 f,

4 m, Tarrafal; 1 s, 5 £ CANCAP Sta. 6.008, 14? 54' N, 23? 30” W, 120 m (5-V1-1982) (NN M); 11 s, 6 f,

CANCAP Sta. 6.010, 14? 52 N, 23? 30' W, 310 m (5-VI-1982) (NN M).

Etymology: The specific name alludes to the subsutural hump on the axial ribs of the shell.

Description: Shell (Figs. 102, 103)

length up to 3.7 mm, maximum width

1.3 mm, relatively solid, strongly elon-

gate-conic.

Protoconch (Fig. 104) of 1 whorl and

about 325 um of maximum diameter, of

non-planktotrophic type, without spiral

sculpture. Microsculpture formed by

irregular flat prominences with some

pits on a smooth surface.

Teleoconch of about 6 strongly

convex whorls; there is a short depres-

sed space below suture, followed by an

evident shoulder on the axial ribs; last

whorl strongly convex; suture shallow,

but evident. Colour whitish.

Axial sculpture consisting of promi-

nent, sharp, narrow, scarcely opistho-

cline, distantly spaced axial ribs, about

12 in last whorl. Spiral sculpture almost

unappreciable at low magnification.

Microsculpture (Figs. 105, 106) shows

groups of 6-9 threads, interrupted by the

growth lines, alternating with bands

with only pits.

Aperture D-shaped, relatively small;

inner lip thick; columellar side weakly

concave; anterior channel almost absent;

outer lip opisthocline with thick exter-

nal varix; peristome with 4-5 parallel

lines towards the inner part of the aper-

ture.

Habitat: In sandy and muddy

bottom, between 53 and 310 m.

Distribution: Known from Sal, Boa

Vista and Santiago Islands (Fig. 170).

Remarks: Schwartziella gibbera n. sp.

differs from the previously described

species by its elongate shell and subsu-

tural hump. S. puncticulata n. sp. also

has a subsutural shoulder, but it has a

smaller and not so elongate shell and

different teleoconch microsculpture. S.

sulcostriata n. sp. is also elongate, but it

lacks subsutural hump and has a differ-

ent teleoconch microsculpture.

Schwartziella (Schwartziella) irregularis n. sp. (Figs. 107-109, 171)

Type material: Holotype (Fig. 107) 1 s of 3.8 (1.4 mm, CANCAP Sta. 7.042, Ponta Inglez/Ponta

Preta, SW of Maio, 15% 07' N, 23* 14” W, 76 m (25-VIII-1986) (NNM 58014). Paratypes: 1 s, CANCAP

Sta. 7.050, SW of Maio, 15 06' N, 23? 14” W, 380 m (25-VIII1-1986) (NNM 58015); 1 s, CANCAP Sta.

7.101, off Palmeira, Sal, 16” 45' N, 23? 01” W, 262-280 m (30-VIII-1986) (NNM 59416).

Etymology: The specific name alludes to the irregular distribution of the axial ribs on the shell.

Description: Shell (Fig. 107) length up

to 3.8 mm, maximum width 1.4 mm, not

solid, narrowly elongate-conic.

Protoconch (Fig. 108) of 1 whorl and

about 175 ym of maximum diameter, of

non-planktotrophic type, smooth.

(Right page) Figures 102-106: Schwartziella (Schwartziella) gibbera n. sp. 102: holotype, llhéu de -

Sal Rei, Boa Vista (NNM 58008); 103: paratype, off Palmeira, Sal (NNM 59413); 104: proto-

conch of a paratype, Sal Rei, Boavista (NNM 58009); 105-106: teleoconch microsculpture of the

holotype.

(Página derecha) Figuras 102-106: Schwartziella (Schwartziella) gibbera spec. nov. 102: holotipo,

Ilhéu de Sal Rei, Boa Vista (NNM 58008); 103: paratipo, Palmeira, Sal (NNM 59413); 104: proto-

concha de un paratipo, Sal Rei, Boavista (NNM 58009); 105-106: microescultura de la teloconcha del

holotipo.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Iberus, 18 (1), 2000

Figures 107-109: Schwartziella (Schwartziella) irregularis n. sp. 107: holotype, Ponta Inglez/Ponta Preta,

Maio (NNM 58014); 108: protoconch of the holotype; 109: teleoconch microsculpture of the holotype.

Figuras 107-109: Schwartziella (Schwartziella) irregularis spec. nov. 107: holotipo, Ponta Inglez/Ponta Preta,

Maio (NNM 58014); 108: protoconcha del holotipo; 109: microescultura de la teloconcha del holotipo.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

DIAS NRE

ar PA torn

Val

Figures 110-113: Schwartziella (Schwartziella) abundata n. sp. 110: holotype, off Palmeira, Sal

(NNM 57998); 111: protoconch of the holotype; 112-113: teleoconch microsculpture of the

holotype.

Figuras 110-113: Schwartziella (Schwartziella) abundata spec. nov. 110: holotipo, Palmeira, Sal

(NNM 57998); 111: protoconcha del holotipo; 112-113: microescultura de la teloconcha del holotipo.

Iberus, 18 (1), 2000

Figures 114-115: Schwartziella (Schwartziella) rectilinea n. sp. 114: holotype, llhéu Razo (NNM

58026); 115: protoconch of a paratype from type locality (NNM 58027).

Figuras 114-115: Schwartziella (Schwartziella) rectilinea spec. nov. 114: holotipo, llhéu Razo (NNM

58026); 115: protoconcha de un paratipo de la localidad tipo (NNM 58027).

Teleoconch of 5 whorls, weakly

convex, not angulated below sutures,

last whorl weakly convex; suture

shallow. Colour whitish.

Axial sculpture consisting of few prom-

inent, sharp, narrow, slightly opisthocline,

distantly spaced axial ribs, 10 in last whorl,

which have not a regular correspondence

from whorl to whorl. Spiral sculpture not

visible at low magnification. Microsculp-

ture (Fig. 109) only formed by irregular

spiral rows of small pits.

Aperture D-shaped, small sized;

inner lip thin; columellar side weakly

concave; anterior channel almost absent;

outer lip opisthocline with thick exter-

nal varix; peristoma simple with a

second elevation in the inner part.

(Right page) Figures 116-120: Schwartziella (Schwartziella) rarilineata mn. sp. 116: holotype, Rabo

de Junco, Sal (MNCN 15.05/31716); 117: protoconch of a paratype; 118: protoconch of the

holotype; 119-120: teleoconch microsculpture of a paratype, Rabo de Junco (CER).

(Página derecha) Figuras 116-120: Schwartziella (Schwartziella) rarilineata spec. nov. 116: holotipo,

Rabo de Junco, Sal (MNCN 15.05/31716); 117: protoconcha del paratipo; 118: protoconcha del holo-

tipo; 119-120: microescultura de la teloconcha de un paratipo, Rabo de Junco (CER).

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Iberus, 18 (1), 2000

Habitat: Collected in sandy sedi-

ments between 76 and 380 m.

Distribution: Only known from Sal

and Maio (Fig. 171).

Remarks: S. irregularis n. sp. differs

from the previously described species of

Schwartziella by the narrow and elon-

gated shell, small protoconch, axial ribs

irregularly disposed and the very faint

microsculpture with only small perfora-

tions. By its elongate shape it is similar

to S. gibbera n. sp., but this species has a

larger protoconch, a subsutural hump,

axial ribs continued over adjacent

whorls and a more marked microsculp-

ture. S. sulcostriata n. sp. also has a

larger protoconch and different proto-

conch and teleoconch microsculpture.

Schwartziella (Schwartziella) abundata n. sp. (Figs. 110-113, 171)

Type material: Holotype (Fig. 110) 1 s of 2.9 x 1.4 mm (NNM 57998) and 2 paratypes, 2 s (NNM

57999), CANCAP Sta. 7.100, off Palmeira, Sal, 16” 45' N, 23” 01” W, 262-280 m (30-VII1-1986).

Other material studied: Maio: 4j, 1 f, CANCAP Sta. 7.050, 15” 06' N, 23” 14” W, 380 m (25-VII!-

1986) (NNM).

Etymology: The specific name is derived of the Latin abundo (to surpass), alluding to the nume-

rous axial ribs of this species, more than in any other Cape Verde Schwartziella.

Description: Shell (Fig. 110) length up

to 2.9 mm, maximum width 1.4 mm, not

solid, elongate-conical.

Protoconch (Fig. 111) of 1 whorl and

420 um of maximum diameter, of non-

planktotrophic type, without spiral

sculpture; transition to teleoconch

abrupt. Microsculpture formed by irreg-

ular flat prominences with some perfo-

rations on a smooth surface.

Teleoconch of 4 strongly convex

whorls, not angulated below suture; last

whorl strongly convex; suture shallow.

Colour whitish.

Axial sculpture consisting of scarcely

prominent, narrow, opisthocline, distantly

spaced axial ribs, absent just below the

suture; about very curved 15-16 ribs on

the last whorl. Spiral sculpture not visible

at low magnification. Microsculpture

(Figs. 112, 113) formed by fine threads

separated by rows of small pits.

Aperture D-shaped, relatively large;

inner lip thick; columellar side weakly

concave; anterior channel shallow; outer

lip opisthocline with thick external

varix; peristome sharp.

Habitat: Muddy bottom between 260

and 380 m.

Distribution: Only known from Sal

and Maio (Fig. 171).

Remarks: S. abundata n. sp. has a larger

protoconch than any other of the Cape

Verde species of Schwartziella, and also

more numerous and curved axial ribs, and

a subsutural area without axial sculpture.

Schwartziella (Schwartziella) rectilinea n. sp. (Figs. 114, 115, 172)

Type material: Holotype (Fig. 114) 1 s of 3.8 x 1.4 mm (NNM 58026), and 16 paratypes, 16] (NNM

58027), CANCAP Sta. 7.121, Ilhéu Razo, 16” 36' N, 24? 37" W, 200-230 m (1-IX-1986). Other paraty-

(Right page) Figures 121-125: Schwartziella (Schwartziella) inscripta n. sp. 121: holotype, Pal-

meira, Sal (MNCN 15.05/31710); 122: shell from Maio (broken during the study); 123, proto-

conch of a paratype, Rabo de Junco, Sal (CER); 124: protoconch of a shell from Palmeira, Sal;

125: detail of the suture in last whorl of the same shell.

(Página derecha) Figuras 121-125: Schwartziella (Schwartziella) inscripta spec. nov. 121: holotipo,

Palmeira, Sal (MNCN 15.05/31710); 122: concha de Maio (rota durante su estudio); 123, protoconch

of a paratype, Rabo de Junco, Sal; 124: protoconcha de una concha de Palmeira, Sal; 125: detalle de la

sutura en la última vuelta de la misma concha.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Iberus, 18 (1), 2000

pes: 1j (MNCN 15.05/31717) and 1 s (CER), both from the type locality; 19 j, CANCAP Sta. 7.119,

S of Ilhéu Razo, 167 36' N, 24? 36' W, 140-160 m (1-1X-1986) (NNM 59418); 1 s, CANCAP Sta. 6.095,

S of Ilhéu Razo, 16” 35 N, 24? 37" W, 930 m (15-VI-1982) (NNM 59419); 1 s, CANCAP Sta. 7.120, S

of Ilhéu Razo, 16 36" N, 24? 36' W, 208 m (1-I1Xx-1986) (NNM 59420).

Other material studied: Sáo Nicolau: 25 j, 1 f, Sáo Jorge Bay, CANCAP Sta. 7.129, 16” 33" N, 24” 16'

W, 405 m (2-1X-1986) (NNM).

Etymology: The specific name alludes to the right profile of the whorls of the shell.

Description: Shell (Fig. 114) length up

to 3.8 mm, maximum width 1.4 mm,

relatively solid, elongate-conical.

Protoconch (Fig. 115) of 1 whorl and

380 um of maximum diameter, of non-

planktotrophic type, without spiral

sculpture, transition to teleoconch at-

tenuated.

Teleoconch of 7 whorls, almost flat-

sided, specially the first ones; last whorl

weakly convex; suture shallow. Colour

cream-whitish.

Axial sculpture consisting of promi-

nent, narrow, scarcely opisthocline, dis-

tantly spaced axial ribs; the ribs are con-

tinued from whorl to whorl. Spiral

sculpture unappreciable at low magnifi-

cation. Microsculpture formed by spiral

rows of very small pits on the first

whorls. Due to poor condition of the

adult shells no microsculpture could be

observed on the last whorl except for

growth lines.

Aperture D-shaped, relatively small;

inner lip thick; columellar side weakly

concave; anterior channel absent; outer

lip opisthocline with thick external varix.

Habitat: Found in muddy and calca-

reous sand between 140 and 930 m.

Distribution: llhéu Razo and Sáo

Nicolau (Fig. 172).

Remarks: S. rectilinea n. sp. is very

elongate, and it differs from the fol-

lowing similar species with elongate

shell: S. irregularis n. sp. has more

convex spire whorls and smaller proto-

conch; S. gibbera n. sp. has very promi-

nent axial ribs and a subsutural hump;

S. sulcostriata n. sp. has a smaller proto-

conch, a more curved profile of the

spiral whorls, and a more marked teleo-

conch microsculpture.

Schwartziella (Schwartziella) rarilineata n. sp. (Figs. 116-120, 173)

Type material: Holotype (Fig. 116) 1 s of 2.3 (1.0 mm, Rabo de Junco, Sal, Cape Verde Archipelago,

6 m (MNCN 15.05 /31716). Paratypes: 1 s, Palmeira, Sal, 8 m (MNHN); 1 s, Regona, Sal, 2 m (NNM

58025); 1 s, Sáo Vicente, CANCAP Sta. 7.161, 16? 54” N, 24? 54” W, 95 m (NNM 59428); 2 s, 2 f, Rabo

de Junco, Sal, 6 m (CER); 1 j, Baia Teodora, Boa Vista, 4 m (CER); 1 s, Sal Rei, Boa Vista, 4 m (AMNH).

Other material studied: Sal: 2 s, 1 j, 1 f, Palmeira, 8 m; 4 s, Regona, 10 m; 2 s (eroded), 1 j, Rabo de

Junco, 4 m. Sáo Nicolau: 1 s (eroded), Sáo Jorge Bay, CANCAP Sta. 7.129, 16? 33" N, 24” 16' W, 405

m (2-1X-1986) (NNM).

Etymology: The specific name alludes to the teleoconch microsculpture formed by a few lines.

Description: Shell (Fig. 116) length

up to 2.5 mm, maximum width 1.1 mm,

shining, relatively solid, elongate-

conic.

Protoconch (Figs. 117, 118) of 1 whorl

and about 320 ym of maximum diame-

ter, of non-planktotrophic type, smooth,

transition to teleoconch not abrupt.

(Right page) Figures 126-129: Schwartziella (Schwartziella) sculpturata n. sp. 126: holotype, Pal-

meira, Sal (MNCN 15.05/31720); 127-128: protoconchs of two paratypes, Rabo de Junco, Sal

(CER); 129: teleoconch microsculpture of the holotype.

(Página derecha) Figuras 126-129: Schwartziella (Schwartziella) sculpturata spec. nov. 126: holotipo,

Palmeira, Sal (MNCN 15.05/31720); 127-128: protoconchas de dos paratipos, Rabo de Junco, Sal

(CER); 129: microescultura de la teloconcha del holotipo.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

[e]

Iberus, 18 (1), 2000

Teleoconch of 4 whorls, almost flat-

sided, last whorl weakly convex; suture

shallow. Colour translucent white.

No axial sculpture. Spiral sculpture

(Fig. 116, 119) formed only by few spiral

bands, composed by groups of 2-3 very

fine threads with intermediate striae on

subsutural areas (Fig. 120).

Aperture D-shaped, relatively large;

inner lip thin; columellar side weakly

concave; anterior channel almost absent;

outer lip slightly opisthocline with thin

external varix; peristome narrow,

rounded.

Habitat: Sand sediments, in shallow

water (2-6 m), except for two shells

dredged from 95 and 405 m, respecti-

vely.

Distribution: Sal, Boa Vista, Sáo

Nicolau and Sáo Vicente (Fig. 173).

Remarks: Schwartziella rarilineata n.

sp. differs from any of the previously

known Eastern Atlantic species of the

genus by its teleoconch without axial

ribs, almost smooth and shining, like a

Zebina. Nevertheless, we include it in

the genus Schwartziella because it lacks

of tubercles inside of the outer lip,

unlike other Atlantic species of Zebina

(Z. paivensis, Z. browniana, Z. robustior,

see GOFAS, 1999, and below under

Zebina villenai).

Schwartziella (Schwartziella) inscripta n. sp. (Figs. 121-125, 174)

Type material: Holotype (Fig. 121) 1 s of 3.4 x 1.4 mm, Palmeira Bay, Sal Island, Cape Verde Archi-

pelago, 6 m (MNCN 15.05 /31710). Paratypes: 1 s, Matiota, Sáo Vicente, 4 m (NNM 58013); 1 s, Pau

Seco, Maio (AMNH); 1 s, Rabo de Junco, Sal (MNHN y; 1 s, Rabo de Junco, Sal (CER); 1 s, 2 f, llhéu

de Sal Rei, Boa Vista (CER); 1 s, 1 f, Porto da Cruz, Boa Vista, 4 m (CER).

Other material studied: Sal: 2 s (1 destroyed during study), 2 j, 1 £, Palmeira, 8 m; Boa Vista: 1 f,

Sal Rei, 5 m; Maio: 1 s (Fig. 121, destroyed during study), Pau Seco; Sáo Vicente: 1 f, Porto

Mindelo, 15 m.

Etymology: The specific name alludes the presence of striae on the first whorl of the shell.

Description: Shell (Figs. 121, 122)

length up to 3.5 mm, maximum width

1.5 mm, relatively solid, shining, elon-

gate-conic.

Protoconch (Figs. 123, 124) of 1

whorl and about 310 ym of maximum

diameter, of non-planktotrophic type,

smooth; transition to teleoconch not

conspicuous.

Teleoconch of 5 whorls, weakly

convex; suture very shallow. Colour

translucent whitish. Axial sculpture

absent. Spiral sculpture only present on

the first whorl of the teleoconch (Fig.

123), formed by 4-5 spiral striae, which

disappear immediately (Fig. 125).

Aperture D-shaped, relatively large;

inner lip thin; columellar side weakly

concave; anterior channel almost absent;

outer lip opisthocline, with thick exter-

nal varix; peristome simple, rounded.

Habitat: Sand sediments from

shallow water (4-15 m).

Distribution: Sal, Boa Vista, Maio, Sao

Vicente (Fig. 174).

Remarks: S. inscripta n. sp. has a larger

shell than S. rarilineata n. sp., and the spiral

striae only appear on the first whorl.

Schwartziella (Schwartziella) sculpturata n. sp. (Figs. 126-129, 175)

Type material: Holotype (Fig. 126) 1 s of 2.1 x 1.0 mm, Palmeiras, Sal Island, Cape Verde Archi-

pelago, 10 m (MNCN 15.05/31720). Paratypes: 1 s (NNM 58030), 1 s (UNHN) 3 s (CER), all from

the type locality; 2 s, 2 f, Sal Rei, Boa Vista (CER); 3 s, Rabo de Junco, Sal, 4 m (CER).

Other material studied: Sal: 2 s (1 broken during the study), 2 f, Rabo de Junco, 4 m; 2 s, Palmeira,

10 m; 6 s, 1 f, Regona, 10 m. Santiago: 1 s, Cidade Velha, 4 m; 1 s, Praia, 5 m. Brava: 1 s, Furna, 30

m, 1 s, Porto do Anciáo, 3 m. Sáo Vicente: 1 s, Porto Mindelo, 15 m.

Etymology: The specific name alludes to the presence of axial and spiral sculpture.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Figures 130-132: Schwartziella (Schwartziella) paradoxa n. sp. 130: holotype, Sáo Vicente (NNM

58021); 131: protoconch of the holotype; 132: teleoconch microsculpture of the holotype.

Figuras 130-132: Schwartziella (Schwartziella) paradoxa spec. nov. 130: holotipo, Sáo Vicente (NNM

58021); 131: protoconcha del holotipo; 132: microescultura de la teloconcha del holotipo.

Iberus, 18 (1), 2000

Description: Shell (Fig. 126) length up

to 2.1 mm, maximum width 1.0 mm, not

solid, elongate-conic.

Protoconch (Figs. 127, 128) of 1

whorl and 270 um of maximum diame-

ter, of non-planktotrophic type, without

spiral sculpture; transition to teleoconch

abrupt. Microsculpture (Fig. 128) shows

a rough surface formed by small depres-

sions with many irregular threads.

Teleoconch of 4 whorls, weakly

convex; suture shallow. Colour cream-

whitish.

Axial sculpture consisting of weakly

prominent, narrow, almost orthocline, dis-

tantly spaced axial ribs, only on the first

2-3 whorls of teleoconch, disappearing on

the last whorl. Spiral sculpture very

regular, formed by fine spiral cords, about

15 on penultimate and 35 on last whorl.

Microsculpture (Fig. 129) formed by spiral

bands of 4-5 threads alternating with

furrows.

Aperture D-shaped, medium sized;

inner lip thin; columellar side weakly

concave; anterior channel absent; outer

lip opisthocline with a thin external

varix; peristome simple with a pair of

parallel lines towards the inner part of

the aperture.

Habitat: Sandy sediments from

shallow water (3-30 m).

Distribution: Known from Sal, Boa

Vista, Santiago, Brava, Sao Vicente (fig.

175). Probably it can be found in the

entire archipelago.

Remarks: S. sculpturata n. sp. differs

from S. rarilineata n. sp. and S. inscripta

n. sp. by its evident teleoconch and pro-

toconch sculpture.

Schwartziella (Schwartziella) paradoxa n. sp. (Figs. 130-132, 137, 138, 176)

Type material: Holotype (Fig. 130) 1 s of 2.7 x 1.1 mm, Baia das Gatas, Sao Vicente, CANCAP Sta.

7.161, 16% 54” N, 24? 54” W, 95 m (NNM 58021).

Etymology: The specific name alludes to the shell characters intermediate between Schwartziella

and Zebina.

Description: Shell (Fig. 130) length 2.7

mm, width 1.1 mm, not solid, elongate-

conic.

Protoconch (Fig. 131) of 1 whorl and

320 ym of maximum diameter, of non-

planktotrophic type, without spiral

sculpture and smooth; transition to

teleoconch not abrupt.

Teleoconch of 4 whorls, weakly

convex; suture shallow. Colour whitish.

Axial sculpture of adapical spire

whorls consisting of few prominent,

sharp, narrow, opisthocline, slightly

undulate and distantly spaced axial ribs,

which begin on the suture and disappear

a little below the middle of the whorl;

axial ribs absent on the last whorl. Spiral

sculpture visible at low magnification in

all the whorls, except on the inferior part

of the last whorl. Microsculpture (Fig.

132) formed by groups of few spiral

threads separated by wider rough spaces.

Aperture D-shaped, relatively large;

inner lip thin; columellar side weakly

concave; anterior channel absent; outer

lip opisthocline with a not too thick

external varix; peristome simple with

some parallel lines towards the inner

part of the aperture.

Habitat: Muddy bottom, at 95 m.

Distribution: Only known from the

type locality (Fig. 176).

(Right page) Figures 133-136: Zebina (Zebina) villenai n. sp. 133: holotype, S of Ilhéu Razo

(NNM 58016); 134: protoconch of the holotype; 135: shell of smaller size, S of Ilhéu Razo

(NNM 58017); 136: protoconch of the same shell.

(Página derecha) Figuras 133-136: Zebina (Zebina) villenai spec. nov. 133: holotipo, Sur de Ilhéu

Razo (NNM 58016); 134: protoconcha del holotipo; 135: concha de pequeño tamaño, Sur de Ilhéu

Razo (NNM 58017); 136: protoconcha de la misma concha.

he Cape Verde Archipelago

ininae 1n t

The subfamily Risso

ROLÁN AND LUQUE

Iberus, 18 (1), 2000

Figures 137, 138: Schwartziella (Schwartziella) cf. paradoxa. 137: shell from Rabo de Junco, Sal

(CER); 138: teleoconch microsculpture.

Figures 137, 138: Schwartziella (Schwartziella) cf. paradoxa. 1. 137: concha de Rabo de Junco, Sal

(CER); 138: microescultura de la teloconcha.

Remarks: Schwartziella paradoxa n. sp.

differs from any other species of Cape

Verde Schwartziella by having prominent

axial ribs on first whorls of teleoconch,

which are absent on last whorl. S. sculptu-

rata n. sp., which also has weak axial ribs

on the first 2-3 whorls, is less elongate,

has more evident spiral sculpture and a

different protoconch microsculpture.

One shell (Figs. 137, 138) of 3.0 x 1.0

mm, found in Rabo de Junco, Sal, is

similar to that of Schwartziella paradoxa

n. sp. but it lacks totally of axial ribs

and has more spiral sculpture in the last

whorl. We think that it is probably a

different species, but we prefer do not

describe it awaiting for further mate-

rial.

Genus Zebina H. and A. Adams, 1854

Subgenus Zebina s. s.

Type species: Rissoina semiglabrata A. Adams, 1854, by subsequent designation (REHDER, 1980).

Diagnosis: PONDER (1985, p. 85).

Zebina (Zebina) villenai n. sp. (Figs. 133-136, 172)

Type material: Holotype (Fig. 133) 1 s of 4.2 (1.8 mm (NNM 58016) and 1 paratype, 1 s (NNM

58017), CANCAP Sta. 7.119, S of Ilhéu Razo, 16” 36' N, 24” 36" W, 140-160 m (1-IX-1986). Other

paratypes: 1 s, CANCAP Sta. 7.122, S of Ilhéu Razo, 16? 36' N, 24? 35 W, 100 m (1-1X-1986) (NNM

59423); 1 s, CANCAP Sta. 6.093, SW of Ilhéu Razo, 16? 36' N, 24” 37" W, 400-430 m (15-V1-1982)

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

100 um

Figures 139-143: Shells of dubious species of Schwartziella (Schwartziella) from the Cape Verde

Islands. 139: Schwartziella (Schwartziella) cf. minima, Calhau, Sáo Vicente. 140, 141: S. (S.) cf. punc-

ticulata. 140: shell, CANCAP Sta. 6145, Sáo Vicente; 141: microsculpture of the same shell. 142,

143: S. (S.) cf. typica; 142: shell from Calhau, Sáo Vicente; 143: microsculpture of the same shell.

Figuras 139-143: Conchas de especies dudosas de Schwartziella (Schwartziella) de Cabo Verde. 139:

Schwartziella (Schwartziella) cf minima, Calhau, Sáo Vicente. 140, 141: S. (S.) cf. puncticulata.

140: concha, CANCAP Sta. 6145, Sáo Vicente; 141: microescultura de la misma concha; 142, 143: S.

(S.) ef typica; 142: concha de Calhau, Sáo Vicente; 143: microescultura de la misma concha.

Iberus, 18 (1), 2000

(NNM 59424); 1 s, 1j, CANCAP Sta. 7.128, Sáo Jorge Bay, Sáo Nicolau, 16” 33' N, 24* 17” W, 400 m

(2-1-1986) (NNM 59425).

Other material studied: Ilhéu Razo: 2 s (Fig. 134), CANCAP Sta. 7.116, S of Ilhéu Razo, 16" 36' N,

24” 36' W, 75 m (1-1X-1986) (NN M).

Etymology: The specific name is dedicated to Miguel Villena, who manages the type collection of

molluscs at the MNCN, for his kind cooperation.

Description: Shell (Figs. 133, 135)

length up to 4.2 mm, maximum width

1.8 mm, very solid, elongate-conic.

Protoconch (Figs. 134, 136) of 1

whorl and about 430 ym of maximum

diameter, of non-planktotrophic type,

without spiral sculpture; transition to

teleoconch appreciable with difficulty.

Teleoconch of about 5 flat-sided

whorls; last whorl strongly convex at

the lower part; suture shallow. Colour

whitish. Axial and spiral sculpture

lacking. Surface smooth and shining.

Aperture D-shaped, relatively small;

inner lip thin; columellar side weakly

concave; anterior channel almost absent;

peristome simple, with some parallel

lines towards the inner part of the aper-

ture; two tubercles not always evident

on an interior elevation of the aperture.

Habitat: Coarse calcareous sand,

gravel and stones, between 75 and 430 m.

CONCLUSIONS

Twenty-nine species of the subfam-

ily Rissoininae belonging to three

genera and four subgenera have been

found in the Cape Verde Archipelago.

Only one of them, the sole species with

a protoconch of planktotrophic type

(Rissoina (Rissoina) punctostriata) has a

wide distribution along the West

African coast, whereas the other 28

species, 1 belonging to the genus Ris-

Distribution: lhéu Razo and Sáo Ni-

colau (Fig. 172).

Remarks: Zebina villenai n. sp. is

similar to Z. paivensis (Watson, 1873)

from the Canary and Selvagens Islands,

a species confused with the Caribbean

species Z. browniana (d'Orbigny, 1842) or

Z. vitrea (C. B. Adams, 1850) by authors

(ODHNER, 1932; NORDSIECK, 1972;

GARCÍA-TALAVERA, 1983, see GOFAS,

1999), but Z. paivensis has a smaller pro-

toconch (340 um) and more convex

whorls. Z. browniana and Z. vitrea have

protoconchs of planktotrophic type. Z.

robustior Gofas, 1999, from Southern

Morocco to Senegal is larger than Z. ville-

nai (up to 5.3 mm) and its protoconch is

smaller (about 350 ym, measurements

from figure 80 of GOFAs, 1999).

A few shells from Ilhéu Razo are of

small size (Fig. 135), but the protoconch

(Fig. 136) shows no differences.

soina (Ailinzebina), 26 to Schwartziella

(Schwartziella), and 1 to Zebina (Zebina)

should be considered endemic of the

archipelago on the basis of their non-

planktotrophic type of protoconch and

the absence of records from the neigh-

bouring continental coasts (Senegal).

The high percentage of endemism of

the Cape Verde Rissoininae (96,5%) is

comparable to that of other Cape Verde

(Right page) Figures 144-147: Radulae of Schwartziella (Schwartziella) from the Cape Verde

Islands. 144: radula of Schwartziella (Schwartziella) robusta n. sp., Sal; 145: radula of S. (S.) typica

n. sp., Sal; 146: radula of S. (S.) sanmartini n. sp., Boa Vista; 147: radula of S. (S.) similiter n. sp.,

Brava.

(Página derecha) Figures 144-147: Rádulas de Schwartziella (Schwartziella) de las islas de Cabo

Verde. 144: rádula de Schwartziella (Schwartziella) robusta spec. nov., Sal; 145: rádula de S. (S.)

typica spec. nov., Sal; 146: rádula de S. (S.) sanmartini spec. nov., Boa Vista; 147: rádula de S. (S.)

similiter spec. nov., Brava.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Iberus, 18 (1), 2000

rissoideans, like Alvania (16 endemic

species from a total of 19, viz. 84,2%;

MOOLENBEEK AND ROLÁN, 1988, HOEN-

SELAAR AND GOUD, 1998), Manzonia (7 of

7, 100%; ROLÁN, 1987a), Crisilla (6 of 6,

100%; TEMPLADO AND ROLÁN, 1994),

Barleeia (3, 100%; GOFas, 1995). Similar

high percentages of endemic rissoids

were found in other Macaronesian lÍs-

lands: Alvania (Canary Islands, 14 of 20,

70%; Madeira, 10 of 14, 71,4%; HOENSE-

LAAR AND GOUD, 1998), and Manzonia

(Canary Islands, 9 of 10, 90%; Madeira, 6

of 7, 85,7%; MOOLENBEEK AND FABER,

1987a, b, c). Other gastropod genera also

show high percentages of endemic spe-

cies in the Cape Verde Islands, like Eato-

nina (2, 100%, MOOLENBEEK, 1985-86;

ROLÁN AND TEMPLADO, 1993), Ammoni-

cera (7 species, 100%, RoLÁN, 1992),

Euthria (7, 100%; CosEL, 1982d; COSEL

AND BURNAY, 1983; ROLÁN, 1985, 1987b),

Conus (45 of 48, 93,7%) (ROLÁN, 1990,

1991) and Volvarina (9 of 9, 100%) (Mo-

RENO AND BURNAY, 1999). From our own

preliminary data, which will be discus-

sed in a forthcoming paper, the number

of endemic species of Cape Verde ma-

rine gastropods reachs 193, from a total

of 588 identified species, i. e., a general

percentage of 32,8% of endemism. This

high percentage of endemic species is

comparable with other even more isola-

ted oceanic islands (Galapagos, Easter

Is., Hawaii), and requires special rese-

arch and conservation efforts. Conside-

ring only data referring to Rissoininae,

Pitcairn Islands, which are 390 km from

the nearest islands, have 12 species,

with a maximum of three endemic ones,

all them with non-planktotrophic proto-

conch (SLEURS AND PREECE, 1994). Only

four species of Rissoininae are reported

from the Galapagos Islands, which are

1000 km off the nearest continental

mainland, two of which are probably

endemic; the Galapagos Rissoininae

fauna is very impoverished compared to

the rather rich tropical eastern Pacific

(SLEURS, 1989). In contrast, there are four

species, three of them endemic in Gua-

dalupe Island which is only 260 km off

the coast of northern Baja California

(SLEURS, 1989).

The occurrence of the studied species

in each one of the islands of the archipel-

ago is showed in Figures 158-176 and

Table II. The currently available informa-

tion is insufficient to know the actual

geographical patterns of distribution of

all the species in the archipelago. We

have studied only a few samples from

deep water from Santo Antáo and Ilhéus

do Rombo, and from shallow water from

Fogo, llhéu Branco, Ilhéu Razo, Santa

Luzia and Sáo Nicolau. Moreover, it is

also possible that the original distribution

of the species could have been altered by

the accidental introduction of species in

some islands due to the human activities.

Anyway, we can be sure that some

species are not uniformly distributed

along the archipelago but they are only

present in some islands, because we com-

monly found them in one or two islands

and not at all in the rest. Nothing can be

said in species in which little material has

been studied, like Schwartziella paradoxa.

In these cases, we are not sure of its pre-

sence in other islands. Sal and Boa Vista

are the islands with a higher number of

species (20 and 14, respectively), and this

Figures 148-155: Opercula of Schwartziella (Schwartziella) from the Cape Verde Islands. 148: S.

(S.) robusta, Sal, inner part; 149: S. (S.) sanmartini, Sal, inner part; 150: S. (S.) sanmartini, Boa

Vista, outer part; 151: S. (S.) sanmartini, Sal, inner part; 152: S. (S.) similiter, Brava, inner part;

153: S. (S.) luisi, Palhona, Sal, inner part; 154: S. (S.) luisi, Palhona, Sal, outer part; 155: S. (S.) cf.

Iuisi, Calhau, Sáo Vicente.

Figures 148-155: Opérculos de Schwartziella (Schwartziella) de las islas de Cabo Verde. 148: S. (S.)

robusta, Sal, parte interna; 149: S. (S.) sanmartini, Sal, parte interna; 150: S. (S.) sanmartini, Boa

Vista, parte externa; 151: S. (S.) sanmartini, Sal, parte interna; 152: S. (S.) similiter, Brava, parte

interna; 153: S. (S.) luisi, Palhona, Sal, parte interna; 154: S. (S.) luisi, Palhona, Sal, parte externa;

155: S. (S.) cf. luisi, Calhau, Sáo Vicente.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Iberus, 18 (1), 2000

Table 1. Differences between Rissoina (Rissoina) punctostriata and R. (R.) decussata.

Tabla I. Diferencias entre Rissoina (Rissoina) punctostriata y R. (R.) decussata.

Rissoina punctostriata

size up to 10 mm

profile

adapical whorls of teleoconch

axial ribs

subsutural part of last whorls

protoconch

sinusigera notch deep

could be related with its older origin as

ROLÁN (1991) pointed for the species of

the genus Conus, but also with the fact

that these islands were the most sampled,

as well as Santiago (11 species), Sáo

Vicente (8) and Brava (9). Some species

have been found in several close islands

of a group and not in the rest. Probably

these species are restricted to this group

of islands, which is compatible with their

non-planktotrophic development and the

isolation of islands, as was mentioned by

ROLÁN (1991) in the Cape Verde species

slightly undulated in the lost

whorls by the subsutural depression

angulated

opisthocline and slightly curved;

weakly prominent on the last whorl

depressed; axial ribs almost disappear

3 whorls, diameter increases rapidly

Rissoina decussata

usually up to 7 mm

almost rectilineal

not angulated

almost orthocline; rectilineal;

prominent on the last whorl

not depressed; axial ribs well marked

2 */2 whorls, diameter increases slowly

absent

of Conus with non-planktotrophic deve-

lopment. More than a half of Schwartziella

species (15) apparently have a wide

bathymetric distribution, since they were

found between shallow water and

bathyal depths. Nevertheless, only shells

were collected of most of species or were

found at deep water, and they are pro-

bably transported from shallow to deeper

bottoms along the abrupt shelf of the

islands. Only S. cancapae are mainly

represented by shells collected below 60

m, and S. irregularis, S. abundata, S. rectili-

Figure 156: Drawing of the anterior part of a male of Schwartziella (Schwartziella) sanmartini,

showing the penis.

Figura 156: Esquema de la parte anterior de un macho de Schwartziella (Schwartziella) sanmartini,

mostrando el pene.

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Table IT. Distribution of the species in the archipelago. Abbreviations: S, Sal; BV, Boavista; M,

Maio; ST, Santiago; E, Fogo; B, Brava; IR, Ilhéus do Rombo ou Secos; C, Ilhéu de Cima; SA,

Santo Antáo; SV, Sáo Vicente; SL, Santa Luzia; R, llhéu Razo; SN, Sáo Nicolau.

Tabla II. Distribución de las especies en el archipiélago. Abreviaturas: S, Sal; BV, Boavista; M, Maio;

ST, Santiago; E Fogo; B, Brava; IR, llhéus do Rombo ou Secos; C, llhéu de Cima; SA, Santo Antáo; SV;

Sáo Vicente; SL, Santa Luzia; R, Ilhéu Razo; SN, Sáo Nicolau.

Ss BM

R. punctostriata + + Se +

R. onobiformis + so +

S. robusta a da

S. obesa e

S. corrugata

S. sanmartini + + de

S. similiter +

S. typica

S. angularis

S. luisi

S. minima

S. fulgida se

S. depressa + e

S. gradata

S. pavita di + +

++ ++

+++

. cancapae

. puncticulata

.-hoenselaari

. paucicostata

. sulcostriata

. gibbera

. Irregularis

. abundata

. rectilinea

. rarilineata + +

. Inscripta +

. sculpturata + + +

. paradoxa

villenai

++++

+++ +++

nea, S. paradoxa and Zebina villenai are

exclusively represented by shells collec-

ted below 75 m of depth. Since only a

small part of the studied material was

collected alive and the sampling effort

was quite different on different islands

and depths, a lot of work remains to be

done for knowing the basic ecological

information of almost all the species

(actual habitat, relative abundance, etc.)

The sole species with planktotrophic

type of protoconch (Rissoina punctostriata)

was found in all the islands. Eight species

B IR C SA SV SL R SN

+ + + + + + + o +

+ + +

+ + + + + +

+ + PE

+ +

of Schwartziella were only known from

the North-east group of islands: two of

them from Sal, Boa Vista and Maio (S.

sanmartini and S. pavita, Fig. 162), four

only from Sal and Boavista (S. robusta, S.

angularis, S. luisi and S. minima, Fig. 159),

and two other only from Sal and Maio (S.

irregularis and S. abundata, Fig. 169). Th-

ree species were known only from the

North-west group: two from Sáo Nico-

lau, lIhéu Razo and Ilhéu de Cima (S. rec-

tilinea and Zebina villenai, Fig. 172), and

one only from Sáo Vicente (S. paradoxa,

Iberus, 18 (1), 2000

Santo ÁAntáo

Sao Vicente

Santa Luzia

llhéu Branco ha z $

Théu Raso” ; S

20 Nicolau

Boavista de 162

Ilhéus do Rombo bh Maio ;

llhéu de Cima =. 4 DAN

Santiago

23 W

Figure 157. Map of the Cape Verde Archipelago. Figures 158-165: Distribution areas. 158: Rís-

soina (Ailinzebina) onobiformis, 159: Schwartziella (Schwartziella) robusta, S. (S.) angularis, S. (S.)

luisi and S. (S.) minima; 160: S. (S.) obesa; 161: S. (S.) corrugata and S. (S.) gradata (only Brava);

162: S. (S.) sanmartini and S. (S.) pavita; 163: S. (S.) similiter; 164: S. (S.) typica and S. (S.)

depressa; 165: S. (S.) fulgida and S. (S.) cancapae.

Figura 157. Mapa del Archipiélago de Cabo Verde. Figuras 158-165. Áreas de distribución. 158: Ris-

soina (Ailinzebina) onobiformis; 159: Schwartziella (Schwartziella) robusta, S. (S.) angularis, S.

(S.) luisi y S. (S.) minima; 160: S. (S.) obesa; 161: S. (S.) corrugata y S. (S.) gradata (sólo Brava);

162: S. (S.) sanmartini y S. (S.) pavita; 163: S. (S.) similiter; 164: S. (S.) typica y S. (S.) depressa;

165: S. (S.) fulgida y S. (S.) cancapae.

Fig. 176). Four species were known only from Santiago, Fogo and Brava, and two

from the South-west group of islands: (S. corrugata and S. gradata, Fig. 161) only

two (S. fulgida and S. cancapae, Fig. 165) from Brava. Finally, five species (Schwart-

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

Figures 166-176. Distribution areas. 166: S. (S.) puncticulata; 167: S. (S.) hoenselaari; 168: S. (S.)

paucicostata; 169: Schwartziella (Schwartziella) sulcostriata; 170: S. (S.) gibbera; 171: S. (S.) irregu-

laris and S. (S.) abundata; 172: S. (S.) rectilinea and Zebina (Zebina) villenai;, 173: S. (S.) rariline-

ata; 174: S. (S.) inscripta; 175: S. (S.) sculpturata; 176: S. (S.) paradoxa.

Figuras 166-176. Áreas de distribución. 166: S. (S.) puncticulata; 167: S. (S.) hoenselaari; 168: S.

(S.) paucicostata; 169: Schwartziella (Schwartziella) sulcostriata; 170: S. (S.) gibbera; 171: S. (S.)

irregularis y S. (S.) abundata; 172: S. (S.) rectilinea y Zebina (Zebina) villenai; 173: S. (S.) rariline-

ata; 174: S. (S.) inscripta; 175: S. (S.) sculpturata; 176: S. (S.) paradoxa.

ziella obesa, S. puncticulata, S. hoenselaari, S.

paucicostata and S. sculpturata) were

found in the three groups of islands, ot-

her five in the NE and NW groups ($. ty-

pica, S. depressa, S. sulcostriata, S. rariline-

ata and S. inscripta), and three other in the

NW and SW groups (S. similiter, S. gibbera

and Rissoina (Ailinzebina) onobiformis).

Since Rissoininae species do not

differ markedly in anatomical features,

Iberus, 18 (1), 2000

except for the morphology of the penis,

species identification is usually based on

shell characters only (PONDER, 1985;

SLEURS, 1993, 1994). However, shell cha-

racters seem to be strongly subject to

parallelism, and this makes identifi-

cation of the apomorphic conditions

extremely difficult (SLEURS, 1994). Only a

thorough revision of the genera of Ris-

soininae, including teleoconch micros-

culpture, protoconch morphology,

anatomy (and especially the penial cha-

racters) and the fossil record, may give

us an accurate idea of the phylogenetic

relationships of the species. Neverthe-

less, in the absence of sufficient systema-

tic and distributional data, we point out

in the following lines some considera-

tions about the Cape Verde species of

Rissoininae that will need further rese-

arch.

Rissoina punctostriata seems to form

with R. decussata from the Caribbean and

R. elegantula (Angas, 1880) from South

Australia a group of closely related

species. The holotype of R. elegantula

(BMNH 1881.4.29.4, Aldinga Bay) has

similar shell size (6.6 x 2.8 mm), shape

and sculpture to the two Atlantic species,

and it also has a protoconch of plankto-

trophic type with sinusigerous disconti-

nuity. As we will say below, this is not the

only group of species with an apparently

disjunct Atlantic-Pacific distribution.

The subgenus Rissoina (Ailinzebina)

comprises at least five Recent species, of

which four are apparently distributed

only in the Western Pacific, and one (R.

(A.) elegantissima) has a tropical western

Atlantic distribution (SLEURS, 1993). The

second Atlantic species described here,

Rissoina (Atlinzebina) onobiformis n. sp., se-

ems to be more closely related by its size,

subcylindrical and thin shell, axial and

spiral sculpture and non-planktotrophic

type or protoconch to the western Pacific

species R. (A.) abrardi (Ladd, 1966). The

shell of the western Atlantic R. (4.) ele-

gantissima is also similar in size, shape

and sculpture, but it is rather solid and

the protoconch is of planktotrophic larval

type. The remaining three Pacific species

of this subgenus clearly differ from the

precedent “group” of species in shell

shape or size and sculpture (see SLEURS,

1993). It should be considered a vicariant

origin of the two non-planktotrophic spe-

cies of this “group” from a widely distri-

buted Tethyan planktotrophic ancestor

(may be R. (4.) elegantissima?), a hypothe-

sis proposed by LEAL AND MOORE (1989)

for two other related species of Rissoina

(Rissoina indiscreta Leal and Moore, 1989,

from Brazil, and R. turricula Pease, 1861,

from the Indo-Pacific). Nevertheless, spe-

cies of the subgenus Ailinzebina are rather

uncommon, and therefore more biogeo-

graphical and anatomical information is

needed to establish reliable relationships.

The oldest known fossils of this subgenus

date from the Early Tertiary of France

and the Lower Miocene of Bikini (SLEURS,

1993), and this supports such hypothesis.

These matter deserves further detailed

research, since similar groups of closely

related species distributed in one (or

both) sides of the Atlantic and the Indo-

Pacific were recorded in several families,

viz. the genus Luria (Cypraeidae), with a

pair of related species (Luria lurida from

the Eastern Atlantic and L. pulchra from

the Red Sea, Gulf of Oman and Persian

Gulf) (ALVARADO AND ÁLVAREZ, 1964),

the Conus venulatus group from the Cape

Verde Islands and Conus suturatus from

Central Indo-West Pacific and Australia

(ROLÁN, 1991), and the genus Volvarina

(Marginellidae), with Western and East-

ern Atlantic and Red Sea related species

(MORENO AND BURNAY, 1999).

The high number of species of Cape

Verde Schwartziella is surprising, since

only one species of this genus (S. afri-

cana) has been recorded from the neigh-

bouring West African coast, and as far

as we know, there are no endemic

species of this genus in the islands of

Sáo Tomé and Principe and the Canary

Islands. The five species of Schwartziella

from St. Helena Island described by

SMITH (1890) also have paucispiral pro-

toconch and seem to be endemic. The

loss of a planktotrophic larval stage and

the isolation of Cape Verde islands seem

to be the main factors of speciation, but

there is yet much work to do on the iso-

lation mechanisms within the archipel-

ago that has led to such a high specia-

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

tion. The present knowledge does not

allow to stablish if such a radiation is

dued to one or more colonizations of the

archipelago, followed by isolation and

speciation. The scarce information on

Atlantic Schwartziella, especially regard-

ing protoconch characters, teleoconch

microsculpture and penial anatomy,

makes very difficult to establish rela-

tionships based solely on the available

1. - Shell with axial sculpture on all whorls

data. It is possible that the non-plankto-

trophic Atlantic Schwartziella derived

from a widespread Atlantic species with

planktotrophic development, but only a

revision of the Atlantic recent and fossil

species could give solid cues about the

speciation of the genus.

On the basis of the shell characters used

in the descriptions we present a key for

the Cape Verde species of Schwartziella.

- Shell without axial sculpture or with weak axial sculpture not present on all

Os A DO EAS EDO

2. - Shell with a strong subsutural angulation

- Shell lacking strong subsutural angulation

y e

- Shell length up to 3 mm

PAE SMA YT IOSS USE IIA 23

4. - Shell relatively wide (1/w ratio 2.12-2.20); protoconch with only a spiral angula-

HONRA up penpar as:

ds RO ALO 26319312 S. angularis

- Shell relatively narrow (1/w ratio 2.32-2.44); protoconch with three spiral cords

and irregular axial threads

5. - Suture markedly undulous due to the axial ribs

nc astcrs Mitpae ¿he Ade S. gradata

ASstturerechlinearonstishtyundulous A A E A 6

do nellivery sold and:strong? soto 0960 ade rola lio IU DAA S. robusta

- HE E A A o A 7

7. - Shell showing aperture with 8 axial ribs visible in last whorl ............. 8

- Shell showing aperture with less than 8 axial ribs visible in last whorl ...... 9

8. - Adult shells longer than 3 mm; protoconch diameter about 360 um . S. sanmartini

- Adult shells length up to 3 mmy protoconch diameter about 420 um

- S. abundata

9. - Shell showing aperture with less than 6 axial ribs visible in last whorl ...... 10

- Shell showing aperture with 6 or more axial ribs visible in last whorl

10. - Shell showing aperture with 5 axial ribs visible in last whorl; protoconch with

sculpture, of about 300 ym of diameter

E A eS: S. hoenselaari

- Shell showing aperture with 4 axial ribs visible in last whorl; protoconch smooth,

of about 270 um of diameter

O S. paucicostata

MIA simo convexinithe subsutural par 12

Aa lbs notmore conve ubsutural pa 13

12. - Teleoconch microsculpture formed by undulated rows of punctae . S. puncticulata

- Teleoconch microsculpture formed by groups of 6-9 spiral threads interrupted by the

growth lines, alternating with bands with only pits ........0..000. S. gibbera

13. - Spiral sculpture of the teleoconch visible at low magnification .......... 14

- Spiral sculpture of the teleoconch not visible at low magnification ....... 19

A nelllens tu. tos ma a e oe Nes eyes. TEL 15

OS o E Á e 16

Iberus, 18 (1), 2000

15. - Shell with evident and separate spiral threads... 00000. S. luisi

- Shell with finer and closer spiral threads

16. - Shell with not prominent axial ribs

- Shell with prominent axial ribs ....

17. - Protoconch with a spiral cord

- Protoconch lacking spiral sculpture

A ad 17

18. - Axial ribs not very prominent; protoconch diameter about 400 ym. . S. sulcostriata

- Axial ribs prominent; protoconch diameter about 340 ym ........ S. cancapae

19. - Shell with rectilinear profile, specially in first whorls

- Shell with convex whorls ........

20. - Axial ribs not regularly disposed; protoconch diameter < 200 ym

po ca S. rectilinea

Me ra ec cal. 20

. . S. trregularis

- Shell with regular ribs; protoconch diameter >200 4M ......00 0000... ZA

AE Shell e E A li de OSO N: S. fulgida

cl O 22

22. - Shell with a slight subsutural depressi0N .....000oooooooo o S. depressa

¡Shelllackinesubsutural depression O S. typica

23 ohelliwitaispiralisculptureonl eo ala O AO 24

¡SheHwitWspiraltandiaxalsculpture a a A O e 25

24. - Shell with spiral striae on first whorl only ...........0...0... S. inscripta

- Shell with spiral sculpture on all whorls

dos dana S. rarilineata

25. - Shell with uniform spiral sculpture in all the shell ........... S. sculpturata

- Shell without spiral sculpture at the base

ACKNOWLEDGEMENTS

This work would not have been pos-

sible without the help of the following

people and institutions: “Dirección

General de Cooperación Técnica y Cien-

tífica del Ministerio de Asuntos Exterio-

res” (Spain), which provided finantial

support for the “Primera Expedición

Científica Ibérica al Archipiélago de

Cabo Verde”; “Secretaria de Estado das

Pescas” and the Government of the

Republic of Cape Verde, which gave the

facilities for undertaking this expedi-

tion; we also acknowledge to our com-

panions in this expedition the help with

field sampling; the late Francisco Fer-

nandes also helped us collecting sedi-

ments; “Consejería Territorial y de

Medio Ambiente del Gobierno de Cana-

rias”, which provided finantial support

within the cooperation program “Cana-

rias-Cabo Verde” to the project “Evalua-

eo S. paradoxa

ción de los recursos naturales litorales

de la República de Cabo Verde”, in

which was included the “Macaronesia

2” expedition; Margarita Mosquera,

sorted a lot of sediments; Jesús Méndez

(CACTI, Centro de Apoyo Científico y

Tecnológico a la Investigación, Universi-

dad de Vigo) made most of the SEM

photographs, and José Bedoya (Museo

Nacional de Ciencias Naturales, Ma-

drid) made some ones at the beginning

of this study; Edmund Gittenberger

(NNM) loaned the material from

CANCAP expeditions; Virginie Heros

(MNHN) loaned the type material of

Rissoina africana; Kathie M. Way

(BMNH) loaned the type material of R.

decussata, R. striatocostata, Rissoina ele-

gantula and the five species described by

SMITH (1890). We specially acknowledge

the kind cooperation of Rosario Morán,

ROLÁN AND LUQUE: The subfamily Rissoininae in the Cape Verde Archipelago

who begun this study with the material

from the “Primera Expedición Científica

Ibérica” and Hink L. Hoenselaar, who

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O Sociedad Española de Malacología —__——T— lberus, 18 (1): 95-123, 2000

A molluscan community from coastal bioclastic bottoms in

the Strait of Gibraltar area

La comunidad de moluscos de un fondo bioclástico costero del

Estrecho de Gibraltar

José RUEDA*!, Carmen SALAS* and Serge GOFAS*

Recibido el 10-1X-1999. Aceptado el 17-X1-1999

ABSTRACT

The molluscan community of a soft bottom of bioclastic gravels and sand has been studied

monthly over two years in four stations of the bay of Barbate, SW Spain at the Atlantic

entrance of the Strait of Gibraltar, between 18 and 29 m depth. Species richness, domi-

nance and frequency over the two years are recorded. Micromolluscs, less than 1 mm in

size, were also recorded from the sample of April 1994.

Over the two years, 203 species of macromolluscs were found, of which 174 species

(115.104 individuals) were collected alive and quantified. A total of 25 species of micro-

molluscs were determined, some of which usually considered as rare species. Most species

richness values range between 20 to 40 species per sample, which is comparable with, or

higher than, the values reported in other studies from soft bottom communities of molluscs.

The qualitative analysis based on Jaccard's and Baroni-Urbani and Buser's indices show

that three stations are significantly similar (p < 0.01), whereas the lowest similarity was

found between the station close to the harbour of Barbate and the other ones. The quanti-

tative analysis based on the Bray and Curtis index is biased by the high number of indivi-

duals of Chamelea striatula but consistent with the qualitative indices when this species is

not taken into account.

The main community is similar to “Biocénose des fonds meubles instables (MI)” and “Bio-

cénose des fonds detritique du large (DL)” of PÉrEs AND PicARD (1964), but occurs shallo-

wer than in the Mediterranean. The concurrence of species from the Lusitanian, Mediterra-

nean and Mauritanian regions results in an increment of the total number of species.

RESUMEN

Se ha estudiado la comunidad de Moluscos de fondos de grava bioclástica y arena

durante dos años en la bahía de Barbate, SW de España cerca del Estrecho de Gibraltar.

Se han elegido cuatro estaciones entre 18 y 29 metros de profundidad. Se ha determi-

nado la riqueza especifica, la dominancia y la frecuencia a lo largo de los dos años. Los

micromoluscos, de menos de 1 mm de talla, se han estudiado sólo para la muestra de

Abril de 1994.

Se han colectado un total de 203 especies de macromoluscos a lo largo de los dos años,

de las cuales 174 especies (115.104 individuos) se cogieron vivos y se cuantificaron. Se

han determinado también 25 especies de micromoluscos, algunas de las cuales conside-

* Depto. Biología Animal, Facultad de Ciencias, Universidad de Málaga, Campus de Teatinos s/n, E-29071-

Málaga

' Present address: Rijksinstituut voor visserijonderzoek (RIVO-DLO), (Centrum voor Schelpdieronderzoek),

Korringaweg, 5, Postbus 77, 4400 AB Yerseke, The Netherlands.

Iberus, 18 (1), 2000

radas habitualmente como especies raras. La mayoría de los valores de la riqueza especi-

fica oscilan entre 20 y 40 especies por muestra, lo que es comparable o incluso mayor

que los valores reseñados en otros estudios para comunidades de moluscos de fondos

blandos.

Los análisis cualitativos de afinidad basados en los índices de Jaccard y de Baroni-Urbani

y Buser muestran que tres estaciones son significativamente similares (p < 0,01), mientras

que la menor afinidad se encontró entre la estación próxima al puerto de Barbate y las

otras. El análisis cuantitativo de afinidad basado en el índice de Bray y Curtis esta ses-

gado por el gran número de individuos de Chamelea striatula, pero si no consideramos

dicha especie los resultados son similares a los obtenidos por índices cualitativos.

La comunidad es similar a la “Biocénose des fonds meubles instables (MI)” y “Biocénose

des fonds detritique du large (DL)” de PÉrEs Y PicARD (1964), pero aparece a menor pro-

fundidad que en el Mediterráneo. La comunidad de la estación próxima al puerto pre-

senta caracteres intermedios con la “Biocénose des sables fins bien calibrés (SFBC)” de

PÉRES Y PICARD (1964). La coexistencia de especies procedentes de las tres regiones Lusitá-

nica, Mediterránea y Mavritánica supone un incremento de la riqueza específica.

KEY WORDS: Molluscan communities, Strait of Gibraltar, Species richness,

PALABRAS CLAVE: Comunidades de moluscos, Estrecho de Gibraltar, riqueza específica.

INTRODUCTION

The Strait of Gibraltar is an interes-

ting area for the study of the marine

fauna, including molluscs, due to the

confluence of Atlantic and Mediterra-

nean waters. According to EKMAN

(1953), it is the meeting point of three

biogeographic regions: Lusitanian, Mau-

ritanian and Mediterranean. As a result

of this, the species richness in this area

is possibly the highest of the European

coasts. In addition, there is a clear pre-

sence of North African fauna in this

zone, and there also occur some

endemic mollusc species (GOFAS, 1999).

Some XIX century expeditions, such

as “Lightning” and “Porcupine”, “Cha-

llenger” “Travailleur” and “Talisman”

reported on the marine molluscs from

the Ibero-Moroccan Gulf (see review in

SALAS, 1996) but were mostly concerned

with the bathyal fauna. Recently a

French expedition “Balgim” (1984) and

a Spanish survey “Fauna 1” (1989)

added more information about the

malacofauna, particularly bivalves

(SALAS, 1996).

There are relatively few studies

regarding the fauna from the Strait of

Gibraltar. Some of them reported only

faunistic lists (FISCHER-PIETTE, 1959;

THORSON, 1965; GARCÍA-GOMEZ, 1983a;

VAN AARTSEN, MENKHORST AND GITTEN-

BERGER, 1984; CERVERA, TEMPLADO,

GARCÍA-GÓMEZ, BALLESTEROS, ORTEA,

GARCÍA, ROS AND LUQUE, 1988) and

others added information about mollus-

can communities (GARCÍA-GÓMEZ,

1983b; TEMPLADO, GUERRA, BEDOYA,

MORENO, REMON, MALDONADO AND

RAMOS, 1993, GOFaAs, 1999) or other

invertebrate groups (CARBALLO,

NARANJO AND GARCÍA-GÓMEZ, 1997;

LÓPEZ DE LA CUADRA AND GARCÍA-

GOMEZ, 1993; MEDEL-SOTERAS, GARCÍA

AND GARCÍA-GÓMEZ, 1991). Some

general studies have been carried out in

the inner Bay of Cádiz (ARlas, 1976;

LÓPEZ DE LA ROsa, 1997; DRAKE, ARIAS

AND CONRADI, 1997), but these were not

directed to molluscan communities. A

review of the decapods from expedi-

tions carried out off the southern Iberian

peninsula and the northern coast of

Morocco is given by GARCÍA RASO

(1996), and new records were added by

LÓPEZ DE LA ROSA, GARCÍA RASO AND

RODRIGUEZ MARTÍN (1998). The decapod

crustacean community from Barbate

RUEDA ET A£.: Molluscan community from bioclastic bottoms in the Strait of Gibraltar

Table I. Physical characteristics of the sampling stations in bay of Barbate.

Tabla I. Características físicas de las estaciones de muestreo en la bahía de Barbate.

Barra 10

Retín 12 Retín 16 Barra 16

Depth 21-22 m 28-29 m 28-29 m 18m

O ent coarse sand coarse sand medium sand fine sand

yP with bioclasts with bioclasts with bioclasts with bioclasts

Organic matter content 0.35- 0.40% 0.40 - 0.60% 0.70- 0.80% 0.80-1.10%

was studied by MANJÓN-CABEZA AND

GARCÍA RASO (1998a), and the popula-

tion structure and growth of the hermit

crab Diogenes pugilator from the same

area by MANJÓN-CABEZA AND GARCÍA

RASO (1998b). None of these previous

studies contained data on molluscan

communities from infralittoral bottoms

and their changes through the year.

The present paper was aimed to a

better knowledge of molluscan commu-

nities from bioclastic infralittoral

bottoms in the Strait of Gibraltar, and of

the influence they receive from the diffe-

rent water masses. The results have

been obtained from monthly samples

over two years of survey. These mollus-

can communities are poorly documen-

ted in the Spanish coasts, although some

information from other European sta-

tions is available in the literature

(CABIOCH, 1968; PÉRES AND PICARD,

1964; GLÉMAREC, 1969; TEMPLADO ET AL.,

1993).

AREA OF STUDY

Samples were collected in the bay of

Barbate (36% 8” N- 5% 56” W), on the

Atlantic side of the Strait of Gibraltar

(Fig. 1). Four sample stations were selec-

ted and sampled monthly throughout

two years. Sample stations were placed

on two transects, R (Retín) and B

(Barra), at different distances from the

estuary and harbour of Barbate. The

main characteristics of sampling stations

are listed in Table 1.

The station B10 is the nearest to the

harbour and the estuary of Barbate

river, with high level of sedimentation

(fine particles) and a higher value of

percentage of organic matter in the sedi-

ment than in the other stations. Hard

bottoms are present in the bay as flags-

tones between the station R12 and the

beach (Fig. 1).

The Bay of Barbate is mainly

influenced by water masses from the

Atlantic Ocean. Most of the flow comes

from South Portugal, but due to the pro-

ximity of North Africa there are also

currents of Southern origin (REY, 1983).

Atlantic water currents flow in this bay

in direction to the Mediterranean Sea,

while Mediterranean water currents

flow below 200-250 meters deep in the

opposite direction (VIVES, SANTAMARIA

AND TREPAT, 1975) and do not affect the

bay. Tidal currents can change the direc-

tion of the dominant currents, produ-

cing local gyres. Water temperature

varies from 21” C (Summer months) to

14” C (Winter months). Water salinity

values (around 34 %o) can change due to

the influence of the river Barbate and

some other fresh water reservoirs (ESTA-

BLIER AND MARGALEF, 1964; SEOANE-

CAMBA, 1965).

The mixed sediment (shell frag-

ments and sand) is a habitat for some

benthic algae, which were reported on

by FLORES-MOYA, SOTO, SÁNCHEZ, ALTA-

MIRANO AND CONDE (1995a, b) and by

CONDE, FLORES-MOYA, SOTO, ALTAMI-

RANO AND SÁNCHEZ (1996).

MATERIAL AND METHODS

Samples were collected from

October 1993 to September 1995 with a

small fishing boat, by towing a rectan-

Iberus, 18 (1), 2000

Figure 1. Location of the sampling points in Bay of Barbate. The dashed line represents rocky

bottoms.

Figura 1. Localización de los puntos de muestreo dentro de los fondos de la bahía. La linea discontinua

indica la presencia de lajas de roca.

gular dredge (42 cm width x 22 cm

height), with a 0.5 mm mesh inner

holding bag, for 15 minutes at a speed

of 1 knot. The dredged area for each

sample was approximately 150 m?.

Biological samples were sieved on

different mesh sizes (10, 5, 3, and 1 mm)

in order to split into different size frac-

tions and sort them. The smaller frac-

tions were sorted under a stereomicros-

cope. Molluscs were separated from the

rest of macrobenthos and fixed in for-

maldehyde 10 % and subsequently pre-

served in neutralized alcohol 70%.

The fraction above 1 mm shell size,

representing the macrofauna, was sor-

ted quantitatively for every sample over

the two years. Both living and dead spe-

cimens were identified, although only li-

ving specimens were quantified. The ta-

xonomical ordination is according to Sa-

BELLI, GIANNUZZI-SAVELLI AND BEDULLI

(1990) and current CLEMAM catalogue

(www.mnhn.fr / base / malaco.html).

Species richness and dominance

(percentage of individuals to the total,

for a particular species) were calculated

in order to describe the community. We

also calculated the frequency or percen-

tage of the samples in which the species

is present over the two years. Three

indices of affinity were calculated to

classify the molluscan composition from

the four stations, in order to check the

different communities present in our

study area. Two of them are qualitative

(JACCARD, 1908; BARONI-URBANI AND

BUSER, 1976), the other one quantitative

(BRAY AND CURTIS, 1957). The quanti-

tative dissimilarity index (I) of BRAY

AND CURTIS (1957) was transformed as

(1-I) so as to use it as a quantitative

similarity index. The qualitative indices

were chosen in view of the possibility to

test the significance, following REAL

AND VARGAS (1996) for the Jaccard index

and Baroni-Urbani and Buser tables

(BARONI-URBANI AND BUSER, 1976) for

their own index. The affinity between

stations was represented for each index

by a dendrogram, using the UPGMA

algorithm.

Micromolluscs, mostly species

which have a shell size smaller than the

bulk of the sediment grain, need special

processing to be recovered efficiently

and this was done mainly on the sample

of April 1994. The fraction not retained

RUEDA ET 4£.: Molluscan community from bioclastic bottoms in the Strait of Gibraltar

by the 1 mm sieve was sieved on the 0.5

and 0.3 mm sieves. Then, each of these

fractions was winnowed in sea water, so

that the water movement will carry

away the lighter fraction including

molluscs. The water outflow was collec-

ted on the 0.3 mm sieve, examined for

living micromolluscs, and dried. The

micromolluscs were sorted in this dry

residue under the stereomicroscope,

using a Strattman micropaleontological

tray and a fine wet brush to manipulate

them. Micromollusc data were studied

apart due to an incomplete cuantifi-

cation, only in the sample of April 1994.

Sediment samples were collected

and analysed for granulometry and per-

centage of organic matter (% OM). The

grain size distribution of the sediment

was determined by sieving. The mud

fraction was separated by wet sieving in

a 80um sieve, and finally the dried sand

fraction was sieved over a stacked set of

grade sieves. Granulometric parameters

were calculated according to the method

of BUCHANAN (1984). Sediment for

determination of % OM was stored in

formaldehyde 10% just after collection.

A fraction was dried at 100% C for 24

hours and weighted for obtaining dry

weight. Later on it was burnt to ashes

for 2 h at 525” C and finally weighted

again. Difference of dry weight and dry

weight after ignition determined %OM

in the sediment.

RESULTS

Analysis of the taxocoenosis of

molluscs

A: Macromolluscs composition and

species richness: Over the two years, a

total of 203 species of macromolluscs

were found. Of these, 174 species (115.

104 individuals) were collected alive

(Tables IL, III). Some individuals were

only identified to genus level, either

because diagnostic characters are seen

only on living animals and could not be

observed (e. g. Triphoridae, many Opist-

hobranchs), or because they belong to

groups where taxonomic problems are

not solved (see remarks below).

Number and percentage of species

by class collected were similar in R12,

R16 and B16. In B10, the number of

species by class was lower, although the

percentages of abundance were similar

to those in the other stations (Table HT).

The families Trochidae (12 sp., Fig. 2),

Nassariidae (9 sp., Fig. 3) and Muricidae

(7 sp.) were the best represented among

the gastropods; the Veneridae (13 sp.)

and Cardiidae (10 sp.) among the bival-

ves.

Mean values for species richness per

month were lower in B10 (between 12 to

36) than in the other sampling points:

R12 (17-45 species); R16 (20-49 species);

B16 (21-51 species).

The number of species of gastropods

and their abundance were higher in R16

and B16, where the sediments were cha-

racterised by a mixture of bioclasts and

of coarse and medium sand, respecti-

vely. Total abundances of bivalves were

high in R16 and B10 due to a strong set-

tlement of the bivalve Chamelea striatula,

but richness for bivalves was higher in

the stations with mixed sediments

(gravels and fine sand) than in the most

homogeneous one (B10). In global

terms, bivalves were more abundant

than gastropods mainly due to their gre-

garious occurrence in soft bottoms. The

dredge was not adequate for the collec-

tion of cephalopods so that a low

number of them were registered and

always small individuals (2-3 cm).

The base of the ascidian Phallusia

mammillata (Cuvier, 1815) clusters shells

and small stones which provide a micro-

habitat for some species of molluscs

such as Chauvetia procerula and Ocine-

brina edwardsii which were abundant

inside the holes and crevices from these

structures. We also noted the presence

of small individuals of the bivalves

Chlamys varia and Striarca lactea both

attached by byssal thread.

Some associations of molluscs with

others organisms were found. The bi-

valve Digitaria digitaria usually suppor-

ted colonies of the hydrozoan Monobra-

chium parasitum Meresckowsky, 1877 lo-

cated on the posterior part of the umbo.

Colonies of another non determined hy-

Iberus, 18 (1), 2000

Table II. Species of macromolluscs collected in the survey. Ab: Abundance, Er: Frequency, ff: Dead

specimen, +: Non quantitative data. “Total” is (Retín 12 + Retín 16 + Barra 16), excluding Barra 10.

Tabla II Especies de macromoluscos recolectadas en los muestreos. Ab: Abundancia, Fr: Frecuencia, ff: Indi-

viduo muerto, 4: Datos no cuantitativos. “Total” es (Retín 12 + Retín 16 + Barra 16), excluyendo Barra 10.

Retín 12

Ab Fr

CLASSIS POLYPLACOPHORA Gray). E., 1821

Family LEPTOCHITONIDAE Dall, 1889

Leptochiton cimicoides (Monterosato, 1879) 36 20.83

Family ISCHNOCHITONIDAE Dall, 1889

Lepidochitona cinerea (Linné, 1767) 148

Lepidochitona corrugata (Reeve, 1848) 2 417

Family ACANTHOCHITONIDAE Pilsbry, 1893

Acanthochitona fascicularis (Linné, 1767) 8 16.66

Acanthochitona sp.

CLASSIS GASTROPODA Cuvier, 1797

Family ACMAEIDAE Carpenter, 1857

Acmaea virginea (Miller O. F., 1776) 14417

Family FISSURELLIDAE Fleming, 1822

Diodora gibberula (Lamarck, 1822)

Diodora graeca (Linné, 1758)

Family SCISSURELLIDAE Gray J. E., 1847

Scissurella costata D'Orbigny, 1824

Family TROCHIDAE Rofinesque, 1815

Calliostoma zizyphinum (Linné, 1758)

Calliostoma sp. 1 28 45.83

Calliostoma sp. 2 14:17,

Gibbula magus (Linné, 1758) 39 41.67

Gibbula fanulum (Gmelin, 1791) 3 1250

Gibbula guttadauri (Philippi, 1836) 167 70.83

Gibbula cineraria — (Linné, 1758) 1417

Jujubinus dispar — Curini-Golletti, 1982

Jujubinus exasperatus (Pennant, 1777) 2 833

Jujubinus montagui (Wood W., 1828) 12 20.83

Jujubinus striatus (Linné, 1758) 1 417

Clanculus cruciatus (Linné, 1758) +1 f —

Family TRICOLIDAE Robertson, 1985

Tricolia pullus — (Linné, 1758) 1 417

Family TURBINIDAE Rafinesque, 1815

Bolma rugosa (Linné, 1767)

Family CERITHINDAE Férussac, 1819

Bittium reticulatum (Da Costa, 1778)

Bittium submamillatum (De Rayneval a Ponzi, 1854) 619 62.50

Bittium simplex (Jeffreys, 1867) ++ l —

Cerithium vulgatum Bruguiére, 1792 +$f

Family TURRITELLIDAE Lovén, 1847

Mesalia varia (Kiener, 1887) 149 79.17

Turritella communis Risso, 1826 246 50.00

Turritella turbona Monterosato, 1877 250 50.00

Family RISSOIDAE Gray J. E, 1847

Rissoa guerinii Récluz, 1843

Rissoa inconspicua (Alder, 1844) (MAN

100

Retín 16

20.83

Barra 16

Ab Fr

18 54.17

65 58.33

1004117

1 417

2 8.33

8 12.50

120 95.83

3 1250

264 79.17

4 833

11 20.83

40 58.33

68 58.33

13 37.50

495 100

525 75.00

61 29.17

Barra 10 Total

Ab Fr Ab

7 21.74 19

16 30.43 582

13 2174 192

3 8/70 40

826

66.67

4.17

75.00

20.83

4.17

8.33

12.50

95.83

16.67

87.50

25.00

75.00

8.33

58.33

8.33

19.17

8.33

4.17

12.50

4.17

75.00

113 86.96 845 100

45 26.09 1764

7114

95.83

75.00

12.50

8.33

RUEDA E7 A£.: Molluscan community from bioclastic bottoms in the Strait of Gibraltar

Table IH. Continuation.

Tabla II Continuación.

Alvania cimex (Linné, 1758) +t

Alvania discors (Allan, 1818) +f

Family APORRHAIDAE Gray J. E., 1850

Aporrhais pespelecani (Linné, 1758)

Family CALYPTRAEIDAE Lomarck, 1809

Calyptraea chinensis (Linné, 1758)

Family LAMELLARIIDAE D'Orbigny, 1841

Lamellaria. sp.

Family TRIVIIDAE Troschel, 1863

Trivia arctica (Pulteney, 1789)

Family NATICIDAE Forbes, 1838

Natica hebraea (Martyn, 1784)

Polinices alderí (Forbes, 1838)

Polinices guillemini (Payraudeau, 1826)

Polinices macilenta (Philippi, 1844)

Family CASSIDAE Latreille, 1825

Phalium saburon (Bruguiere, 1792) +f

Phalium undulatum (Gmelin, 1791) $t

Family RANELLIDAE Gray J. E., 1854

Cymotium corrugatum (Lamarck, 1816)

Ranella olearia (Linné, 1758) +t

Charonia lampas (Linné, 1758) +4

Family TRIPHORIDAE Gray J. E., 1847

Not determined

Family CERITHIOPSIDAE Adams H. 8. A., 1853

Cerithiopsis scalaris Locard, 1892

Cerithiopsis tubercularis (Montagu, 1803)

Family EPITONIIDAE Berry S. S., 1910

Fpitonium commune (Lamarck, 1822)

Epitonium pulchellum (Bivona Ant., 1832)

Epitonium jolyi (Monterosato, 1878)

Cirsotrema cochlea (Sowerby 6. B. 11, 1844)

Family EULIMIDAE Adams H. 8. A., 1853

Eulima glabra (Da Costa, 1778)

Melanella alba (Da Costa, 1778)

Melanella sp.

Family MURICIDAE Rafinesque, 1815

Bolinus brandaris (Linné, 1758)

Hexaplex trunculus (Linné, 1758)

Muricopsis cristata (Brocchi, 1814)

Ocenebra erinaceus (Linné, 1758)

Ocinebrina aciculata (Lamarck, 1822)

Ocinebrina edwardsii (Payraudeau, 1826)

Trophon muricatus (Montagu, 1803)

Buccinulum corneum (Linné, 1758)

Chauvetia procerula Monterosato, 1889

Chauvetia brunnea (Montagu, 1803) $f

Chauvetia sp. +4

Fusinus pulchellus (Philippi, 1844)

Fusinus rostratus (Olivi, 1792)

Retín 12 Retín 16

Ab Fr Ab Fr

104 79.17 113 58.33

450 70.83 1297 91.67

l 417 l 417

2 8.33

8 20.83 9 16.67

6 20.83

1M 2917 5 12.50

1 417

1 417 3 1250

2 8.33

l 417

3 1250 9 20.83

2 8.33

1 417

61 75.00 88 79.17

19 37.50

1 417

2 833 23 58.33

2 8.33

48 58.33 32 58.33

17 3333 44 62.50

1 417

21 37.50

EEN

9 1250 14 29.17

2 8.33

Barra 16 Barra 10

50.00

87.50

4.17

37.50

25.00

20.83

4.17

70.83

12.50

87.50

4.17

19.17

54.17

45.83

75.00

4.17

Ab Fr

104 56.52

22 52.17

3 13.04

19 56.52

1 435

38 65.22

4 17.39

Total

83.33

95.83

4.17

8.33

62.50

33.33

54.17

12.50

4.17

33.33

8.33

4.17

95.83

37.50

4.17

95.83

12.50

91.67

87.50

4.17

58.33

19.17

12.50

101

Iberus, 18 (1), 2000

Table IL. Continuation.

Tabla II Continuación.

Nassarivs elatus (Gould, 1845)

Nassarius heynemanni (Von Maltzan, 1884)

Nassarius vaucheri (Pallary, 1906)

Nassarius incrassatus (Stróm, 1768)

Nassarivs pygmaeus (Lamarck, 1822)

Nassarius reticulatus (Linné, 1758)

Nassarius mutabilis (Linné, 1758)

Nassarivs granum (Lomarck, 1822) +f

Cyclope donovania Risso, 1826 $f

Family COLUMBELLIDAE Swoinson, 1840

Mitrella minor — (Scacchi, 1836)

Retín 12

co — 0) OQ) O

Mitrella bruggeni van Aartsen, Menkh. 8 Gittenb., 1984 +f 4

Columbella rustica (Linné, 1758) +f

Family COSTELLARINDAE Mc Donald, 1860

Vexillum tricolor (Gmelin, 1791) +t

Family CYSTISCIDAE Stimpson, 1865

Gibberula epigrus (Reeve, 1865)

Gibberula miliaria (Linné, 1758)

Family VOLUTIDAE Rafinesque, 1815

Cymbium olla (Linné, 1758) ++

Family CANCELLARIDAE Gray J. E, 1853

Concellaria cancellata (Linné, 1758) +4

Family CONIDAE — Rafinesque, 1815

Conus mediterraneus Hwass in Bruguiére, 1792 +f

Family TURRIDAE —Swoinson, 1840

Bela laevigata- (Philippi, 1836)

Bela striolata (Risso, 1826)

Bela sp. 1

Bela sp. 2

Mangelia attenuata (Montagu, 1803)

Haedropleura septangularis (Montagu, 1803)

Crassopleura maravignae (Bivona, 1838)

Raphitoma aequalis (Jeffreys, 1867)

Comarmondia gracilis (Montagu, 1803)

Family PYRAMIDELLIDAE Gray J. E., 1840

Chrysallida terebellum (Phillippi, 1844)

Eulimella acicula (Phillippi, 1836)

Eulimella scillae (Scacchi, 1835)

Odostomia acuta Jeffreys, 1848

Odostomia conoidea (Brocchi, 1814)

Brachystomia eulimoides (Hanley, 1844)

Turbonilla fulvocincta (Thompson, 1840)

Turbonilla rufa (Phillippi, 1836)

Turbonilla rufescens (Forbes, )

Turbonilla striatula (Linné, 1758) +4

Family ACTEONIDAE D'Orbigny, 1835

Acteon tornatilis- (Linné, 1758)

Family RETUSIDAE Thiele, 1931

Retusa truncatula (Bruguiére, 1792)

Retusa mamillata (Philippi, 1836)

102

12.50

4.17

12.50

50.00

16.67

45.83

Retín 16

143

19.17

12.50

Barra 16

25.00

16.67

62.50

33.33

8.33

29.17

16.67

70.83

33.33

41.67

4.17

29.17

8.33

4.17

37.50

Barra 10

Ab Fr

202 100

17 39.13

1 435

59 65.22

276 100

243 100

E AUR

44 65.22

1 435

118 65.22

9 21.74

2 8.70

13 34.78

1 435

9 26.09

7 13.04

l 434

=— mn ay —

QDD) un

— D O == —= — —. NN) hn)

33.33

58.33

95.83

50.00

12.50

19.17

4.17

12.50

8.33

58.33

4.17

8.33

4.17

70.83

62.50

4.17

8.33

4.17

16.67

RUEDA E7 4£.: Molluscan community from bioclastic bottoms in the Strait of Gibraltar

Table IH. Continuation.

Tabla 1 Continuación.

Fomily RINGICULIDAE Philippi, 1853

Ringicula auriculata (Menard, 1811)

Family BULLIDAE Lamarck, 1801

Bulla striata Bruguiére, 1792 ++

Family HAMINOEIDAE Pilsbry, 1895

Haminoea sp.

Family PHILINIDAE — Gray. E., 1850

Philine aperta (Linné, 1767)

Philine. sp.

Family CYLICHNIDAE Adams H. 8. A., 1854

Scaphander lignarius (Linné, 1758)

Family PLEUROBRANCHIDAE Férussac, 1822

Berthella. sp.

Family APLYSIIDAE Lomarck, 1809

Aplysia fasciata Poiret, 1789

Family ARCHIDORIDIDAE Bergh, 1892

Archidoris tuberculata (Cuvier, 1804)

Archidoris sp.

Family DENDRODORIDIDAE 0'Donoghue, 1924

Doriopsilla sp.

CLASSIS SCAPHOPODA Bronn, 1862

Family DENTALIIDAE Linné, 1758

Dentalium inaequicostatum Dautzenberg, 1891

Dentalium vulgare Da Costa, 1778

CLASSIS BIVALVIA Line, 1758

Family NUCULIDAE Gray J. E., 1824

Nucula hanleyi (Winckworth, 1931)

Family NUCULANIDAE Adams H. 8 A. 1858

Nuculana pella (Linné, 1767)

Family ARCIDAE Lomarck, 1818

Arca noae Linné, 1758

Anadara poli (Mayer, 1868)

Family NOETIDAE Stewart, 1930

Striarca lactea (Linné, 1758)

Family GLYCYMERIDIDAE Newton, 1922

Glycymeris glycymeris (Linné, 1758)

Family MYTILIDAE Rafinesque, 1815

Modiolarca subpicta (Cantraine, 1835)

Modiolus barbatus (Linné, 1758)

Modiolus adriaticus (Lamarck, 1819)

Amygdalum agglutinans (Cantraine, 1835)

Family PINNIDAE Leach, 1819

Atrina fragilis (Pennant, 1777)

Family PTERIIDAE Gray J. E, 1847

Pteria hirundo (Linné, 1758)

Family PECTINIDAE Rafinesque, 1815

Pecten maximus (Linné, 1758)

Retín 12

—

LL NI

147

326

20.83

8.33

4.17

12.50

8.33

70.83

91.67

8.33

29.17

8.33

62.50

12.50

Retín 16

289

41.67

4.17

8.33

4.17

33.33

95.83

4.17

29.17

8.33

16.67

75.00

37.50

4.17

25.00

Barra 16

19 37.50

8.33

29.17

16.67

20.83

95.83 387100

25.00

8.33

4.17

8.33

66.67 107 47.83

33.33

4.17

25.00

Total

56 70.83

4.17

8.33

4.17

8.33

4.17

8.33

87.50

4.17

54.17

12.50

45.83

25.00

8.33

91.67

66.67

4.17

41.67

103

Table IL. Continuation.

Tabla 11 Continuación.

Aequipecten opercularis (Linné, 1758)

Aequipecten commutatus (Monterosato, 1815)

Chlamys varia (Linné, 1758)

Flexopecten flexuosus (Poli, 1795)

Family ANOMIIDAE Rofinesque, 1815

Anomia ephippium Linné, 1758

Family LIMIDAE Rofinesque, 1815

Limatula subauriculata (Montagu, 1808)

Lima lima (Linné, 1758) +t

Family LUCINIDAE Fleming, 1828

Ctena decussata (Costa 0. G., 1829) $t

Family UNGULINIDAE Adoms H. 8. A., 1857

Diplodonta rotundata (Montagu, 1803)

Family KELLIIDAE Forbes 8. Hanley, 1848

Kellia suborbicularis (Montagu, 1803)

Family ASTARTIDAE D'Orbigny, 1844

Digitaria digitaria (Linné, 1758)

Goodallia triangularis- (Montagu, 1803)

Family CARDIIDAE Lomarck, 1819

Acanthocardia aculeata (Linné, 1758)

Acanthocardia echinata (Linné,1758)

Acanthocardia mucronata (Poli, 1795)

Acanthocardia tuberculata (Linné, 1758)

Acanthocardia spinosa (Solander, 1786) ++

Parvicardium scabrum (Philippi, 1844)

Plagiocardium papillosum (Poli, 1795)

Loevicardium crassum (Gmelin, 1791)

Loevicardium oblongum (Gmelin, 1791)

Cerastoderma glaucum (Poiret, 1789) $t

Family MACTRIDAE Lamarck, 1809

Mactra stultorum (Linné, 1758)

Spisula subtruncata (Da Costa, 1778)

Spisula elliptica (Locard, 1890)

Lutraria angustior Philippi, 1844 +4

Family PHARIDAE Adams H. 8. A., 1858

Ensis ensis- (Linné, 1758)

Ensis sp.

Phaxas pellucidus (Pennant, 1777)

Family TELLINIDAE Bloinville, 1814

Arcopagia balaustina Linné, 1758

Tellina incarnata- (Linné, 1758)

Tellina distorta (Poli, 1791)

Tellina pygmaea Lovén, 1846

Tellina donacina Linné, 1758 ++

Family DONACIDAE Fleming, 1828

Donax venustus Poli, 1795

Capsella variegata (Gray J. E., 1851)

Family PSAMMOBIIDAE Fleming, 1828

Gari fervensis (Gmelin, 1791)

104

Iberus, 18 (1), 2000

Retín 12

597

29.17

58.33

37.50

12.50

4.17

91.67

12.50

8.33

41.67

58.33

12.50

100

29.17

19.17

8.33

16.67

4.17

Retín 16

Ab Fr

5 16.67

7 12.50

63 66.67

133 62.50

162 66.67

1188 100

Barra 16 Barra 10

Ab Fr Ab Fr

5 16.67

34 54.17

51 58.33 19 21.74

140 62.50 15 13.04

pra

1 417

1067 95.83 1272 91.30

2 8.70

2 833 1 435

2 8.33

4 12.50 280100

His

20 25.00 3 13.04

8 25.00

115 79.17 255 78.26

417

5 417 3 13.04

182 91.67 1013100

H —

10 17.39

1 435

l 435

2 8.70

1 417

124 73.91

2 417

2 8.33

Total

117

259

341

29.17

12.50

87.50

12.50

RUEDA ET 4L.: Molluscan community from bioclastic bottoms in the Strait of Gibraltar

Table IL. Continuation.

Tabla II Continuación.

Retín 12 Retín 16 Barra 16 Barra 10 Total

Ab Fr Ab Fr Ab Fr Ab Fr Ab Fr

Gari depressa (Pennant, 1777) 1417 4 870 1 417

Gari costulata Turton, 1822 A Y ¡A

Gari pseudoweinkauffi Cosel, 1990 51 6250 12 1667 5 833 6 435 68 66.67

Family SCROBICULARIIDAE Adoms H. 2. A., 1856

Scrobicularia plana (Da Costa, 1778) +f H—

Family SEMELIDAE Stoliczka, 1870

Ervilia castanea (Montagu, 1803) 10 20.83 10 20.83

Family SOLECURTIDAE D'Orbigny, 1846

Solecurtus scopula (Turton, 1822) ++ * — $ —

Azorinus chamasolen (Da Costa, 1778) +t e =

Family VENERIDAE Rofinesque, 1815

Venus verrucosa Linné, 1758 2 417 3 8.33 5 16.67

Venus casina Linné, 1758 1 417 1 417

Globivenus effossa (Philippi ex Bivona ms., 1836) +f —

Chamelea gallina (Linné, 1758) 9 17.39

Chamelea striatula (Da Costa, 1778) 12582 75.0026169 79.17 7028 91.6726052 73.91 45779 100

Clausinella fasciata (Da Costa, 1778) 152 91.67 55 70.83 85 66.67 15 26.09 292 100

Timoclea ovata (Pennant, 1777) Aaa 1 8:33 12 37.50

Gouldia minima (Montagu, 1803) 1074 87.50 1941 100 1142 83.33 299 56.52 4157 100

Dosinia lupinus (Linné, 1758) 4 1250 1 417 5 16.67

Pitar rudis- (Poli, 1795) 12 20.83 55 66.67 42 75.00 109 87.50

Callista chione- (Linné, 1758) 72 87.50 19 41.67 55 62.50 64 69.57 146 100

Tapes rhomboides (Pennant, 1777) 81 50.00 704 75.00 195 75.00 31 21.74 980 91.67

Tapes decussatus (Linné, 1758) +t e ==

Family CORBULIDAE Lomarck, 1818

Corbula gibba- (Olivi, 1792) 4715 87.50 3886 100 4076 100 1115 78.26 12677 100

Family GASTROCHAENOIDEA Gray J. E., 1840

Gastrochaena dubia (Pennant, 1777) JAS IA, 4 8.33

Family HIATELLIDAE Gray J. E, 1824

Hiatella arctica (Linné, 1767) 1 417 19 25.00 8 25.00 28 50.00

Panopea glycymeris (Von Born, 1778) +t H — —

Family THRACIIDAE Stoliczka, 1870

Thracia sp. +4 4 —

Family PANDORIDAE Rofinesque, 1815

Pandora inaequivalvis (Linné, 1758) 140 45.83 132 62.50 107 58.33 322 69.57 379 91.67

Pandora pinna (Montagu, 1803) 17 29.17 116 6250 83 50.00 8 17.39 216 79.17

Family LYONSIIDAE Fischer P., 1887

Lyonsia norwegica (Gmelin, 1791) 5 20.83 11 3333 10 3333 7 26.09 26 62.50

CLASSIS CEPHALOPODA Cuvier, 1798

Family SEPIIDAE Leach, 1817

Sepia officinalis Linneo, 1758 100x497 AD,

Family SEPIOLIDAE Leach, 1817

Sepietta oweniana- (D'Orbigny, 1840) 1 417

Family OCTOPODIDAE D'Orbigny, 1840

Octopus vulgaris Cuvier, 1798 1 417 1 417 3 12.50 5 20.83

105

Iberus, 18 (1), 2000

Table III. Species richness and abundance (in brackets) per sampling station, and percentages of

each molluscan class.

Tabla III. Riqueza específica y abundancia (entre paréntesis) para cada estación de muestreo y porcenta-

ges por clase de moluscos.

CLASS RETÍN 12 RETÍN 16

POLYPLACOPHORA 4 (47) 2 (91)

GASTROPODA 58 (2.580) 60 (4.165)

BIVALVIA 44 (21.205) 44 (35.806)

SCAPHOPODA 2 (16) 1 (16)

CEPHALOPODA 2 (2) 1(

TOTAL 110 (23.850) 108 (40.079)

drozoan were found in some indivi-

duals of Corbula gibba and were mainly

located on the posterior part of the shell.

The high abundance and frequency

of the shells of Panopea glycymeris, Sole-

curtus scopula and Lutraria angustior

indicates their possible presence in these

communities. These species live deeply

buried in the sediment (25-40 cm),

making difficult their collection with the

dredge. The large valves are a substrate

for Calyptraea chinensis, and a shelter for

many other species.

Some other species such as Cerasto-

derma glaucum and Scrobicularia plana

were transported from the estuary of

Barbate which is close to B10 where they

were found.

B: Micromolluscs list: A total of 25

species of micromolluscs (Table IV) were

determined in the sample from April

1994. A few species (e. g. Gibbula magus,

Chamelea striatula, Tapes rhomboides) are

the juveniles of species otherwise listed

in the macrofauna, but most of them are

species of which the adult size is small,

and which would be totally missed in

the 1 mm sieve.

A few species, some of which are

illustrated on Figure 4, are very abun-

dant, and these belong to species which

are usually deemed to be rare because

they are found preferently on this kind

of bottom, where micromolluscs are not

easy to collect. The best represented

families of microgastropods were Pyra-

midellidae (5 sp.), Skeneidae (3 sp) and

106

BARRA 10

35 (1.423)

30 (31.434)

70 (32.896)

BARRA 16 TOTAL %

202) 2 (143) 5 (283) 2,87 (0,24)

58 (3.261) 99:(11.429) — 58,89(9,92)

44 (14.905) — 65(103.350) — 37,35(89,74)

2 (36) 2(27) 2 (95) 1,14 (0.082)

1(1) 1(3) 3 (7) 1,72 (0.006)

107 (18.339) 174:(115.164) 100100)

Caecidae (3 sp.). The best represented

family of microbivalves was Montacuti-

dae (3 sp.). As for the number of indivi-

duals, the larger numbers correspond to

the supposedly rare chiton Leptochiton

cimicoides, the gastropods Dikoleps nitens,

Pusillina inconspicua-and Retusa mami-

llata, and to the bivalve Limatula subauri-

culata. The small cerithiid Bittium subma-

millatum, also found in the fraction over

1 mm, was quite abundant.

In addition to these species which

are dominant in their size class, we

found several species which are noto-

riously rare elsewhere and could be

recovered here in moderate numbers,

hence are not so rare on this kind of

substrate. The gastropod Retrotortina

fuscata was found alive for the first time

in our study and reported on by GOFAS

AND WARÉN (1998). The recently descri-

bed bivalve Notolimea clandestina, descri-

bed from a few specimens only, is well

represented in our samples and could be

observed with brooded juveniles inside.

The highest overall abundance and

species richness of micromolluscs was

registered in the samples from R12, with

the lowest percentage of organic mater,

and the lowest in B10, with the greatest

percentage of organic matter (Table 1).

Analysis of the Community

A: Dominance and Frequency: In spite

of the high number of collected species,

the molluscan taxocoenosis is domina-

ted by few species. The main dominant

species were quite similar among the 4

RUEDA E7 42.: Molluscan community from bioclastic bottoms in the Strait of Gibraltar

Figure 2. Species of the family Trochidae from Bay of Barbate. A: Calliostoma sp. 1, 8.5 mm; B:

Calliostoma sp. 2., 7.0 mm; C: Gibbula guttadauri, 5.5 mm; D: Jujubinus dispar, 3.8 mm; E: Juju-

binus dispar, 6.3 mm. Dimensions are for shell height.

Figura 2. Especies de la familia Trochidae presentes en la bahía de Barbate. A: Calliostoma sp. 1, 8,5

mm; B: Calliostoma sp. 2., 7,0 mm; C: Gibbula guttadauri, 5,5 mm; D: Jujubinus dispar, 3,8 mm;

E: Jujubinus dispar, 6,3 mm. Las medidas indicadas corresponden a la altura.

sampled stations. There were 8 to 13

species with dominance values higher

than 1 % in R12, R16 and B16, but only 5

species had dominance values higher

than 1 % in the station B10 (Table V).

Chamelea striatula was the most domi-

nant species in all sample stations: in

some months it reached abundance of

5000 to 13000 individuals per sample.

The strong settlement of juveniles of this

species occurred during Spring months

of the first year of survey (1994), then it

persisted as a dominant species for a

limited time in R16, and until the end of

the studied period in B10. The higher

amount of mud and percentage of the

organic matter in the sediment from this

latter station could have favoured the

development of this species.

If we take into account the 20 first

dominant species, it is possible to find

some differences between stations. The

most different composition of dominant

species occurs in B10, where 5 species

(Acanthocardia tuberculata, Nassarius reti-

culatus, Nassarius mutabilis, Nassarius ela-

tus and Donax venustus) are typical of

shallow sand bottom communities.

Their dominant presence in comparison

with the other stations indicates simila-

107

Iberus, 18 (1), 2000

rity with communities of well sorted

fine sand (PÉRES AND PICARD, 1964; GLÉ-

MAREC, 1969; GARCÍA RASO, LUQUE,

TEMPLADO, SALAS, HERGUETA, MORENO

AND CALVO, 1992). This is further sup-

ported by the constant occurrence of

high numbers of Ophiura texturata La-

marck, 1816 and of some individuals of

Echinocardium cordatum (Pennant, 1777).

In R12 some species with affinities for

coarse sand bottoms are listed in the

first 20 dominant species: Bittium subma-

millatum, Laevicardium crassum (with the

highest abundance in the 4 stations),

Turritella turbona, Gibbula guttadauri.

Some other animals collected frequently

in this station, such as Branchiostoma lan-

ceolatum (Pallas, 1766) and Echinocyamus

pusillus (Múller, 1776), are common in

coarse sand bottom communities (FORD,

1923; CABIOCH, 1968; GLÉMAREC, 1969).

The species occurring in a high fre-

quency throughout the 2 years in B10

were different from those of the other

three stations (Table VI), and were

mainly the species with affinity for fine

and shallow sandy bottoms listed with

the dominance data. Species of nassa-

rids (Nassarius reticulatus, N. mutabilis

and N. elatus) and some bivalves such as

Spisula subtruncata, Nuculana pella and

Acanthocardia tuberculata were recorded

in all monthly samples during two

years. Nevertheless, other species recor-

ded with a high frequency in the other

stations, such as Mesalia varia, Corbula

gibba, Digitaria digitaria and Laevicardium

crassum, were also important for the

community structure of B10.

Aplysia fasciata Poiret, 1789 was

recorded in spring and summer months

from the first year and not in samples

from the second year, probably due to a

decrease on the production of algae in

the bay.

B: Faunal affinity between sampling

points: The values of qualitative and

quantitative similarity indices between

sampling stations are shown in Table

VII and Figure 5. A single dendrogram

was found for each index with the

UPGMA reconstructions. From the qua-

litative viewpoint, according to Jac-

card's as well as to Baroni-Urbani and

Buser's indices, the points Retín 12, Re-

tín 16 and Barra 16 are significantly si-

milar (p < 0.01). The lowest similarity in

qualitative terms was found between

the station B10 and the other stations,

whereas a higher similarity is found bet-

ween the two deeper stations R16 and

B16. The quantitative analysis according

to BRAY AND CURTIS (1957) also showed

high similarity between R16 and B16,

where a high number of individuals of

Chamelea striatula were collected. Assu-

ming the invasive character of the settle-

ment of this species in this kind of com-

munities, the same index was computed

also without the data for this species.

The influence of the flood of Chamelea

brings a bias so as to increase the simila-

rity between Barra 10 and other stations

(Fig. 5C), whereas the quantitative data

without Chamelea are consistent with the

qualitative indices (Fig. 5D).

DISCUSSION

Taxonomic remarks: The species of

the genus Calliostoma (Fig. 2) are proble-

matic in the area of transition between

(Right page) Figure 3. Species of the family Nassariidae from Bay of Barbate. A: Nassarius pygmaens,

11 mm; B: Nassarius vaucheri, 13 mm; C: Nassarius reticulatus, juvenile, 13 mm; D: Nassarius incras-

satus, 11 mm; E: Nassarius tingitanus, 9 mm (specimen collected in Tarifa); E: Nassarius heynemanni,

13 mm; G: Nassarius mutabilis, 12 mm; H: Nassarius elatus, 14 mm. Dimensions are for shell height.

(Página derecha) Figura 3. Especies de la familia Nassaridae presentes en la bahía de Barbate. A: Nassa-

rius pygmaeus, 11 mm; B: Nassarius vaucheri, 13 mm; C: Nassarius reticulatus, juvenil, 13 mm; D:

Nassarius incrassatus, 11 mm; E: Nassarius tingitanus, 9 mm (ejemplar recolectado en Tarifa); F: Nas-

sarius heynemanni, 13 mm; G: Nassarius mutabilis, 12 mm; H: Nassarius elatus, 14 mm. Las medidas

corresponden a la altura.

108

RUEDA ET 4Z.: Molluscan community from bioclastic bottoms in the Strait of Gibraltar

109

Iberus, 18 (1), 2000

Table IV. Species of micromolluscs collected in the survey, mainly in the samples of april 1994.

Numbers in brackets denote specimens collected only in thanatocenosis, >100 denotes abundant

species with more than 100 live-collected specimens in the sample.

Tabla IV. Especies de micromoluscos recolectadas en los muestreos, principalmente en abril de 1994. Los

numeros entre parentesis denotan ejemplares encontrados sólo en tanatocenosis, >100 indica especies

abundantes con más de 100 ejemplares colectados en las muestras.

Retin 12 Retín 16 Barra 10 Barra 16

POLYPLACOPHORA

Leptochitonidae

Leptochiton cimicoides (Monterosato, 1879). >100 3 - 1

GASTROPODA

Scissurellidae

Scissurella costata d'Orbigny, 1824 - - - 3

Skeneidae

Skenea serpuloides (Montagu, 1808) 10

Dikoleps nitens (Philippi, 1844) >100

Dikoleps pruinosa (Chaster, 1896) 93

Trochidae

Gibbula magus (Linné, 1758) 12

Rissoidae

Pusillina inconspicua [Alder, 1844) >100

Obtusella intersecta (Wood, 1857) (1)

Caecidae

Caecum trachea (Montagu, 1808) 12

Caecum cuspidatum Chaster, 1896 2

Caecum clarki¡ Carpenter, 1858 1

Vanikoridae

Macromphalina disciformis (Granata Grillo, 1877) - - - 4

Cerithiidae

Bittium pusillum (de Rayneval 8 Ponzi, 1854) 80

Turritellidae

Turritella turbona Monterosato, 1877 (2) -

Turritella communis Risso, 1826 - (2)

Mesalia varia (Kiener, 1887) - (2)

Eulimidae

Vitreolina sp. 3 - - 2

Fasciolariidae

Fusinus pulchellus (Philippi, 1844) - 1

Pyramidellidae

Evalea divisa (J. Adams, 1797) (5)

Odostomia conspicua Alder, 1850 (2) - -

Odostomia unidentata (Montagu, 1803) - - - l

Brachystomia eulimoides [Hanley, 1844) 12 l - 4

Brachystomia sp. 2 - -

Omalogyridae

Retrotortina fuscata Chaster, 1896 20

Ebalidae

Ebala pointeli (de Folin, 1868) 3+(10) 1

110

RUEDA ET 4L.: Molluscan community from bioclastic bottoms in the Strait of Gibraltar

Table IV. Continuación.

Tabla IV. Continuation.

Retusidae

Retusa mamillata (Philippi, 1836)

Cylichnidae

Cylichna crossei B.D.D., 1886

Oxynoidae

Lobiger sp.

BIVALVIA

Nuculidae

Nucula recondita Gofas 4 Salas, 1996

Glycymeridae

Glycymeris glycymeris (Linné, 1758)

Pectinidae

Chlamys sp.

Anomiidae

Anomia ephippium (Linné, 1758)

Limidae

Limatula subauriculata (Montagu, 1808)

Notolimea clandestina Salas, 1994

Leptonidae

Hemilepton nitidum (Turton, 1822)

Kelliidaee

Kellia suborbicularis (Montagu, 1803)

Montacutidae

Mysella bidentata (Montagu, 1803)

Tellimya ferruginosa (Montagu, 1808)

Montacuta goudi van Aartsen, 1996

Astartidae

Goodalia triangularis (Montagu, 1803)

Digitaria digitaria (Linné, 1758)

Cardiidae

Parvicardium scabrum (Philippi, 1844)

Matridae

Spisula subtruncata [da Costa, 1778)

Tellinidae

Tellina pusilla Lovén, 1846

Semelidae

Ervilia castanea (Montagu, 1803)

Veneridae

Tapes rhomboides (Pennant, 1777)

Gouldia minima (Montagu, 1803)

Chamelea striatula (da Costa, 1778)

Corbulidae

Corbula gibba [Olivi, 1792)

Thraciidae

Thracia sp.

Retin 12

>100

(2)

(1)

>100

(1)

38 4

Retin 16 Barra 10 Barra 16

>100

111

Iberus, 18 (1), 2000

Mediterranean Sea and Atlantic Ocean.

Atlantic forms usually have a heavier

sculpture of spiral cords than Mediterra-

nean ones, and this variation in sculp-

ture obscures the delimitation of the

species. Calliostoma zizyphinum was

easily recognized by its broader and less

conical shape, although some variability

occurs between individuals. Calliostoma

sp. 1 is close to the Mediterranean

species Calliostoma conulus (Linné, 1758),

from which it differs by the presence of

heavier ribs. Calliostoma sp. 2 resembles

Calliostoma laugieri (Payraudeau, 1826),

but a detailed comparison of proto-

conchs shows a higher number of gra-

nulated whorls in Calliostoma sp. 2.

The genus Chauvetia is well repre-

sented in the area nearby Strait of

Gibraltar with 10 to 15 species, and its

taxonomy is difficult. Chauvetia sp. from

our study has a white band in the shell

and resembles Chauvetia crassior Odhner,

1923. However, the latter was described

from the Canary Islands and has direct

development, so that it is doubtful

whether the same species is present in

the Strait of Gibraltar. Other species like

Chauvetia decorata Monterosato, 1889,

described from Morocco, also show

white bands, and this character may

even not be constant within a single

species.

Some other taxonomical or nomen-

clatural problems are related with the

Bela species, which are in need of revi-

sion. In our samples some of them were

named as Bela sp. 1 and sp. 2. Nomen-

clatural problems were found for the

species known in the literature as Bela

striolata Risso, 1826, a usage which is not

correct considering that the type speci-

men belongs to the genus Rissoina d'Or-

bigny, 1840. The next available name

could be Bela smithi Forbes, 1844, but the

description is not clear and the type

material is lost. We have followed the

incorrect usage, as revising this question

is beyond the scope of. the paper.

According to OLIVER AND COSEL

(1993), Anadara poli (Mayer, 1868) is not

the same species than the Miocene Arca

diluvii Lamarck, 1805: the Recent species

has a lower number of ribs (24-28) and a

less median umbo. All individuals of

Aequipecten opercularis were smaller and

more elongate than the usual Atlantic

form, and can be grouped in the form

audouinii. Although A. audouinii (Pay-

raudeau, 1826) is currently regarded as a

a synonym of A. opercularis, a reevalua-

tion of their relationships is needed in

order to clarify if this form is a different

species. Chamelea striatula (more Atlan-

tic) and C. gallina (more Mediterranean)

are two closely related species which

live sympatrically in the Southern part

of Iberian peninsula. BACKELJAU,

BOUCHET, GOFAS AND DE BRUYN (1994)

justified the separation of the two

species using allozyme electrophoresis.

C. gallina is common in shallow and

sandy bottoms in our latitudes, but C.

striatula prefers muddy and deeper

(Right page) Figure 4. Micromolluscs from the soft bottom in Bay of Barbate. A: Bittium subma-

millatum, height 3.8 mm; B: detail of the microsculpture of Bittium submamillatum; C: proto-

conch of Bittium submamillatum; D: Pusillina inconspicua, height 1.6 mm; E: Caecum cuspidatum,

length 2.15 mm; F: detail of the microsculpture of Caecum cuspidatum; G: Retusa mamillata,

height 1.85 mm; H: Leptochiton cimicoides, dorsal view, length 2.0 mm; l: Leptochiton cimicoides,

lateral view, length 1.65 mm; J: detail of the girdle of Leptochiton cimicoides, showing the two types

of spicules. Scale bars 100 pm.

(Página derecha) Figura 4. Micromoluscos de los substratos blandos de la bahía de Barbate. A: Bittium

submamillatum, 4/tura 3,8 mm; B: detalle de la microescultura de Bittium submamillatum; C: proto-

concha de Bittium submamillatum; D: Pusillina inconspicua, altura 1,6 mm; E: Caecum cuspida-

tum, longitud 2,15 mm; F: detalle de la microescultura de Caecum cuspidatum; G: Retusa

mamillata, altura 1,85 mm; H: Leptochiton cimicoides, vista dorsal, longitud 2,0 mm; I: Lepto-

chiton cimicoides, vista lateral, longitud 1,65 mm, J: detalle del cinturón de Leptochiton cimicoides

mostrando los dos tipos de espículas. Escalas 100 ym.

112

RUEDA ET AL.: Molluscan community from bioclastic bottoms in the Strait of Gibraltar

113

Iberus, 18 (1), 2000

Table V. The 20 most dominant species in each of the sampled stations.

Tabla V. Las 20 especies con mayor índice de dominancia en cada estación.

Retín 12

Chameleo striatula

Corbula gibba

Gouldia minima

Bittium submamillatum

Digitaria digitaria

Calyptraea chinensis

Loevicardium crassum

Spisula subtruncata

Modiolus adriaticus

Turritella turbona

Turritella communis

Gibbula guttadauri

Clausinella fasciata

Mesalia varia

Nuculana pella

Pandora incequivalvis

Aporrhais pespelicani

Tapes rhomboides

Bela laevigata

Flexopecten flexuosus

52.47

19.77

4.50

2.59

2.50

1.88

1.69

1.62

1.36

1.05

1.03

0.70

0.63

0.62

0.61

0.59

0.44

0.34

0.31

Retín 16

Chamelea striatula

Corbula gibba

Gouldia minima

Colyptraea chinensis

Digitaria digitaria

Turritella communis

Tapes rhomboides

Turritella turbona

Nuculana pella

Modiolus adriaticus

bibbula magus

Spisula subtruncata

Mesalia varia

Anomia ephippium

Bela loevigata

flexopecten flexuosus

Bittium submamillatum

Pandora inaequivalvis

Pandora pinna

Aporrhais pespelicani

65.29

9.69

4.84

3.23

2.96

2.47

1.76

1.00

0.78

0.72

0.69

0.60

0.50

0.40

0.36

0.34

0.33

0.29

0.28

Barra 16

Chamelea striatula

Corbula gibba

Gouldia minima

Digitaria digitaria

Colyptraea chinensis

Turritella communis

Mesalia varia

Gibbula magus

Nuculana pella

Tapes rhomboides

Spisula subtruncata

Modiolus adriaticus

Ocinebrina edwardsi

Anomia ephippium

Calliostoma sp. ]

Loevicordium crassum

Bela loevigata

Pandora inaequivalvis

Nassarius pygmaeus

Pandora pinna

47.14

27.34

1.66

1.15

4.33

3.52

1.77

1.47

1.31

1.22

1.03

0.94

0.80

0.77

0.74

0.72

0.58

0.56

Borra 10

Chamelea striatula

Digitaria digitaria

Corbula gibba

Spisula subtruncata

Nuculana pella

Pandora incequivalvis

Gouldia minima

Aconthocardia tuberculata

Nassarivs reticulatus

Loevicardium crassum

Nassarius mutabilis

Nassarius elatus

Donax venustus

Bela loevigata

Mesalia varia

Modiolus adriaticus

Colyptraea chinensis

Collista chione

Nassarius pygmaeus

Turritella communis

Table VI. Species with a frequency of more than 75% in each of the sampled stations.

Tabla VI. Especies con frecuencia superior a los 75% en cada una de las estaciones.

Retín 12

Laevicardium crassum

Digitaria digitaria

Clausinella fasciata

Nuculana pella

Corbula gibba

Gouldia minima

Collista chione

Aporrhais pespelecani

Spisula subtruncata

Mesalia varia

Chamelea striatula

Bolinus brandaris

114

100%

> 90%

> 85%

> 15%

Retín 16

Corbula gibba

Gouldia minima

Digitaria digitaria

Turritella communis

Nuculana pella

Colyptraea chinensis

Mesalia varia

Bela loevigata

Chamelea striatula

Bolinus brandaris

Tapes rhomboides

Nodiolus adriaticus

Spisula subtruncata

100%

> 95%

> 90%

> 80%

> 15%

Borra 16

Corbula gibba

Mesalia varia

Digitaria digitaria

Nuculana pella

Colliostoma sp 1

Chameleo striatula

Spisula subtruncata

Colyptraea chinensis

Ocenebra erinacea

Gouldia minima

bibbula magus

Ocinebrina edwardsi

Loevicardium crassum

Turritella communis

Tapes rhomboides

Pitar rudis

Fusinus pulchellus

100%

> 95%

> 90%

> 85%

> 80%

> 15%

Barra 10

Spisula subtruncata

Nuculana pella

Aconthocardia tuberculata

Nassarius reticulatus

Nasarius mutabilis

Nassarivs elatus

Digitaria digitaria

Mesalia varia

Corbula gibba

Laevicardium crassum

19.19

3.86

3.38

3.07

1.17

0.98

0.91

0.85

0.84

0.77

0.74

0.61

0.38

0.36

0.34

0.32

0.31

0.19

0.18

0.14

100%

> 20%

> 85%

> 15%

RUEDA £7 4£.: Molluscan community from bioclastic bottoms in the Strait of Gibraltar

Table VII. Values of affinity indexes between sampling stations, qualitative (Jaccard and Baroni-

Urbani and Buser) and quantitative (Bray and Curtis, with and without Chamelea striatula).

Tabla VII. Valores de los índices de de afinidad entre estaciones de muestreos, cualitativos (Jaccard y

Baroni-Urbani y Buser) y cuantitativo (Bray y Curtis, con o sin los datos para Chamelea striatula).

Jaccard R12

R12 1.000

R16

B10

B16

Baroni-Urbani and Buser R12

R12 1.000

R16

B10

B16

Bray and Curtis R12

R12 1.000

R16

B10

B16

Bray and Curtis (Sin C.s.) R12

R12 1.000

R16

B10

B16

bottoms with a higher amount of

organic matter.

Species richness: Values of species

richness in the Bay of Barbate are com-

parable with those obtained in other

studies from soft bottom communities

of molluscs, although sometimes higher.

No information about species richness

per month from detritical bottoms com-

munities of molluscs have been found in

the literature, but some information for

communities for other soft bottoms is

available. Species richness of Molluscs

in soft bottoms from the North Sea is

low (10 to 15 sp.) (ELEFTHERIOU AND

BASFORD, 1989; KUNITZER, 1990), but

there is an increase in the English

Channel (CABIOCH, GENTIL GLACON

AND RETIERE, 1977). In Northern Spain,

species richness values from 15 to 25

R16 B10 BiI6

0.496 0.432 0.510

1.000 0.296 0.536

1.000 0.393

1.000

R16 B10 B16

0.617 0.595 0.029

1.000 0.447 0.059

1.000 OLIVA

1.000

R16 B10 B16

0.667 0.579 0.742

1.000 0.839 0.593

1.000 0.433

1.000

R16 B10 B16

0.701 0.434 0.760

1.000 0.414 0.823

1.000 0.449

1.000

have been recorded in soft bottom com-

munities of molluscs (SÁNCHEZ MATA,

MORA, GARMENDIA, AND LASTRA, 1993;

GARMENDIA, SÁNCHEZ MATA AND

MORA, 1996). In Mediterranean coasts

Salas (1984) registered values for species

richness of molluscs from 20 to 30 in dif-

ferent kinds of soft bottoms of Málaga

bay (Alboran Sea). The highest values of

species richness were recorded (around

40 species per sample) in fine sand

bottoms with low percentage organic

matter. APARICI SEGUER AND GARCÍA-

CARRASCOSA (1996) recorded values of

species richness between 4 to 11 per

sample in the soft bottoms of Chafarines

islands (close to the Mediterranean area

of Morocco), but such low values pro-

bably reflect incomplete sampling.

APARICI SEGUER, ROWLAND, TAYLOR AND

GARCÍA CARRASCOSA (1996) found 10-20

INS

Iberus, 18 (1), 2000

species per sample in the soft bottoms

(fine sand) from the gulf of Valencia at

depths of 15 meters. Values of species

richness per month comparable to those

found in this study have been registered

in communities from hard bottoms

(TRONCOSO, URGORRI AND OLABARRÍA,

1996), from sea grass beds (HERGUETA,

1996; LEDOYER, 1966 a, b) or the calcare-

ous algae Mesophyllum lichenoides

(Lemoine) (HERGUETA, 1996).

Characterisation of the community:

The species living sympatrically in the

Bay of Barbate form communities with

more components than in other areas. In

the Mediterranean Sea similar commu-

nities were referred to as “Biocénose des

fonds meubles instables (MI) “ and

“Biocénose des fonds detritique du large

(DL) ” (PÉRES AND PICARD, 1964). These

have a similar faunistic compositions,

although species richness is higher in

Barbate bay. In our study these commu-

nities occurred in the infralittoral level

(25 m), whereas in the Mediterranean

they occur in the circalittoral, deeper

than in the Atlantic.

The community from B10 is also

similar to the “Biocénose des sables fins

bien calibrés (SFBC) “ (PÉRES AND

PICARD, 1964), but there is a high

influence of faunistic components from

the previously mentioned communities.

In R12 some similarities with the “Biocé-

nose des sables grossiers et des fins gra-

viers (SGFG) “ (PÉRES AND PICARD,

1964) are found, although the instability

of the bottoms due to strong currents

changes the composition of the commu-

nity through the year.

CABIOCH (1968) found similar and

comparable communities in the English

channel, named as “Peuplements des

sédiments fins a Abra alba et Corbula

gibba” and “Biocoenose du Maerl”. In

these communities a mixture of in- and

epifauna occurred over a heterogeneous

sediment. The depth in which these

communities occurred was similar to

those from Barbate, within the infralito-

ral. However, the biogeographical diffe-

rences result in that some species have

been replaced in the Strait of Gibraltar.

116

We could not trace in the literature any

reference to communities in the Mauri-

tanian region, so that a comparison was

not possible.

Structure of the community. Domi-

nance and Frequency: High values of

dominance were more common among

bivalves than among gastropod species.

This reflects their high abundance in

soft bottoms, conforming large popula-

tions. Gastropods are less numerous

than bivalves in soft bottoms samples.

In order to correct this bias, we also took

into account the frequency of the species

in the samples. It is important to take

into account both figures, because some

species with a marked seasonality may

show high abundance during a short

period of time, and thus have a low fre-

quency although they are important ele-

ments of the community.

The highest dominance was for the

species Chamelea striatula, although it

showed lower percentages of presence

than other species. This was caused by a

massive and successful recruitment of

this species in these bottoms in the first

sampling year, although no recruitment

occurred in the second year. Such strong

settlement has been registered for the

same species along the Scottish coasts

by ANSELL (1961). At the beginning of

this study Chamelea striatula was found

in some samples with low abundance.

After spring 1994 this species was domi-

nant in the four sample stations, alt-

hough the population decreased in

some stations during 1995. At the end of

the sampled period, large population of

Chamelea striatula was only present in

B10 which is close to the harbour and

the estuary. This species is common in

fine sand bottoms from shallow coasts

(3-20 m) in the North Sea (Muus, 1973;

DUINEVELD AND BELGERS, 1994). In the

Strait of Gibraltar area, Chamelea striatula

is found living sympatrically with Cha-

melea gallina which is very common in

fine sand bottoms from shallow shores

(2-15 m) in the Mediterranean Sea. C

striatula is usually found in mud

bottoms with high percentages of

organic matter in southern Spanish

RUEDA ET 4£.: Molluscan community from bioclastic bottoms in the Strait of Gibraltar

Baroni-Urbani

8 Buser index

0.3 Er

l Jaccard's index 05 E

0.6 F

R16 B16 R12 B10

0.6 Bray-Curtis index

with C. striatula

without C. striatula

R16 B10 R12 B16 R16 B16 R12 B10

Figure 5. Dendrograms representing affinity of sampling points, according to the qualitative

indices (above) and quantitative (below). Algorithm is UPGMA.

Figura 5. - Dendrogramas de afinidad de los puntos muestreados en base a los valores de los índices cua-

litativos (Superior) y cuantitativos (Inferior). Algoritmo de aglomeración UPGMA.

coasts and never reaches high domi-

nance in near-shore sandy bottoms. The

persistence of this species may have

been favoured by the cleaning and dred-

ging of the channel in the harbour of

Barbate during our sampling period,

which turned more muddy the sedi-

ments nearby.

Other molluscs of these detritic

bottom communities showed high fre-

quencies and dominance. Large popula-

tions of Corbula gibba were collected, but

this is a rather ubiquitous species, com-

monly found together with Turritella

communities on fine sandy bottoms

with large pieces of graves and pebbles

(Hrs-BRENKO, 1981).

Digitaria digitaria is a typical species

of coastal detritic bottoms from North

Spanish coasts (ORTEA, 1977; BESTEIRO,

TRONCOSO, PARAPAR, SALVINI-PLAWEN

AND URGORRI, 1990). Among the gastro-

pods, some of the most typical species

for detritic bottoms are turritellids. In

this location Mesalia varia is an impor-

tant component of the community from

Barbate and also one of the West African

representatives.

In station B10, a mixed community

occurs. Species from detritic bottom

communities (Digitaria digitaria, Mesalia

varia, Corbula gibba) have a high fre-

quency and dominance but some

species from communities of well sorted

fine sand (Nassarius species, Acanthocar-

dia tuberculata, Spisula subtruncata)

showed also high dominances and the

highest frequencies. The community

from this latter sample station seems to

be a transition between both. Communi-

ties of well sorted fine sands are known

in other points from European coasts: in

the North Sea (ForD, 1923; CABIOCH,

1968), in the Mediterranean Sea (SPADA,

SABELLI AND MORANDI, 1973; SALAS,

1984; GARCÍA RASO ET AL., 1992). In

Barbate bay this community shows a

higher species richness than other com-

munities from European coasts. Three

species of Nassariidae (Nassarius reticu-

latus, Nassarius mutabilis and Nassarius

elatus) were found in B10 with high fre-

quency, while Nassarius pygmaeus and

Nassarius vaucheri occurred with low fre-

quency. This kind of community in the

Mediterranean Sea holds usually only

two species of Nassariids (N. mutabilis

and N. reticulatus) and in the North Sea

N. reticulatus and N. pygmaeus. Other

species like Mactra stultorum and Chame-

lea gallina were not found as main com-

ponents in this sample point but they

show high abundance in similar com-

munities from the Mediterranean Sea

(SPADA ET AL., 1973; SALAS, 1984; GARCÍA

RASO ET AL., 1992).

117

Iberus, 18 (1), 2000

Zoogeographical notes: The area of

the Strait of Gibraltar has been said to

act as a barrier for many marine species

including molluscs (EkMAN, 1953). For

some Mediterranean species (Gibbula

guttadauri, Nassarius mutabilis, Naticarius

hebraeus) there is a barrier which makes

them less common or absent in the

Atlantic waters even nearby the Strait of

Gibraltar. Conversely, a few common

species of the North Atlantic waters

such as Spisula elliptica are not found, or

rarely found, in Mediterranean waters.

On the other hand, the area of the

Strait can also be regarded as one where

species of quite different zoogeographi-

cal affinity will concur (PALLARY, 1907;

SPADA AND MALDONADO, 1974; RUEDA

AND SALAS, 1998). The fauna from tro-

pical and temperate West African areas

is represented in Barbate by Mesalia

varia, Epitonium joly1, Nassarius elatus,

Nassarius heynemanni, Nassarius vaucheri

and Gari pseudoweinkauffi, among

others. These species share a northern

limit along the coasts of South Portugal

(to the North) and in the Alboran Sea

(in the Mediterranean). Most Mediterra-

nean species do occur in the Ibero-

Moroccan gulf, at least to Cape St.

Vincent and sometimes further north.

This results in an increase of the

number of species from the different

biogeographical regions, and Barbate is

a locality where their distribution

ranges overlap. This trend is well illus-

trated by the distribution patterns of

the seven species of Nassariids (Fig. 6)

which were collected sympatrically in

this bay.

The fauna of molluscs found in this

study includes mostly species which are

widespread in the Northeast Atlantic

and Mediterranean. Among 27 species

with a distribution restricted to one of

EKMAN's (1953) three regions, there is a

high percentage of similarity (48%) with

typical Mediterranean fauna as found

along the Italian coast. Nine species are

shared with the so-called Mauritanian

region (mostly Morocco), according to

the information of PASTEUR-HUMBERT

(1962 a, b) and of NICKLES (1950, 1955)

This represents a high similarity (33%).

118

This percentage of similarity could be

higher if the information available about

molluscs from North-West Africa were

more complete. Five species with a

mainly North Atlantic distribution are

present in Barbate bay, which represents

18%. This percentage resulted as a com-

parison with the British fauna (TEBBLE,

1966; GRAHAM, 1988) and indicates less

affinity than the other two regions.

It is important from a biogeographic

viewpoint to point out the presence of

endemic molluscan species in the Strait

of Gibraltar area. Recently, several

species of endemic gastropods have

been described from this area and the

data have been summarized by GOFAS

(1999). Nevertheless, most of the

endemic component are linked to rocky

shore in intertidal of very shallow envi-

ronments. It is interesting to note the

presence of Nassarius tingitanus (Pallary,

1901) on hard bottoms very near the

sampling area, but closer to the shore.

This species (Fig. 3) shows a typical

endemic distribution restricted to the

Strait of Gibraltar.

In our samples, we found large

numbers of the endemic gastropod Juju-

binus dispar, which occurs in a restricted

area from Tangier to Ceuta (North

Africa) and was only cited in European

coasts by VAN AARTSEN ET AL. (1984) in

Algeciras Bay (South Spain). Another

rather well documented endemic com-

ponent is the skeneid Dikoleps pruinosa,

originally described from Tangier,

where it is now very rare due to the

extension of the harbour.

Other species which occur in the

Barbate samples have once been

thought to be endemic of the Gibraltar

area but are now known to occur on

other subtidal current-swept gravel

bottoms. The small bivalve Notolimea

clandestina Salas, 1994 (SALas, 1994),

which is common on the bottoms of

Barbate is also known to occur near

Lampedusa, in Sicily Channel (Italy).

The same pattern is found for the rare

skeneid Parviturbo fenestratus, originally

described from Tangiers, and now

reported from Adventure Bank, Sicily

Channel (WARÉN, 1992).

RUEDA £7 A£.: Molluscan community from bioclastic bottoms in the Strait of Gibraltar

ÑN. incrassatus

N. pygmaeus

N. reticulatus

N. elatus

N. heynemanni

N. tingitanus

N. vaucheri

Figure 6. Range of the different species of Nassariidae found in Bay of Barbate.

Figura 6. - Distribución geográfica de las diferentes especies de Nasáridos presentes en la bahía de

Barbate.

To summarize, the results of this

study show that the bay of Barbate sup-

ports a rich soft bottom fauna of

molluscs, which is basically the same in

three of the sampled stations. The high

species richness recorded in these

bottoms may be influenced by three

kinds of factors:

(1) Environmental factors. The bottom

contains a mixture of hard and soft com-

ponents in the sediments. The deeper

stations (25 m) have more large bioclas-

tic material (fragments of shells) deposi-

ted over a fine sand bottom. This hetero-

geneity of the substrates induces a

diversification of the micro-habitats

119

Iberus, 18 (1), 2000

(FRONTIER AND PICHOD-VIALE, 1991;

DEWARUMEZ, DAVOULT, SANVICENTE

ANORVE AND FRONTIER 1992), and con-

sequently a diversification of the fauna.

The occurrence of large-size particles

favours the settling and development of

sessile epifaunal molluscs such as

Anomia ephippium, Modiolus adriaticus,

Calyptraea chinensis, and some mobile

pectinids as Flexopecten flexuosus,

Chlamys varia and Aequipecten species. In

B10 with a lower amount of bioclastic

material there was a decrease in the

abundance of these species but an incre-

ase of others. The mixture of these with

fine sand favours well established popu-

lations of infaunal molluscs, mainly

bivalves as Gouldia minima, Digitaria

digitaria or Spisula subtruncata.

(2) The sampling method. The survey

spanned a longer period of time than in

other reviewed researchs. This allowed

to record changes in the composition of

the community such as occurred with C.

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120

striatula, and to collect species with occa-

sional patterns of occurrence (Aplysia fas-

ciata and Philine species), or locally rare

(Cymatium corrugatum, Atrina fragilis, Pte-

ria hirundo, Spisula elliptica), as well as ju-

veniles in the recruitment season.

(3) The macrogeographic pattern. The

confluence of species from different bio-

geographical areas has been discussed

above, and accounts for much of the

species richness in the area.

ACKNOWLEDGEMENTS

This study was made within the

project DGICYT PB92-0415 supported

by the Spanish Government. We are

grateful to Dr. Raimundo Real for his

useful comments on the indices of af-

finity; and the use of the computer

program for the significance test of Jac-

card's index. Ana S. López is thanked

for help in the laboratory.

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123

O Sociedad Española de Malacología Iberus, 18 (1): 125-131, 2000

Rhopalomenia aglaopheniae (Kowalevsky y Marion, 1887) (Mollusca,

Solenogastres, Rhopalomeniidae), presente en la costa norte de la

Península Ibérica *

Rhopalomenia aglaopheniae (Kowalevsky and Marion, 1887)

(Mollusca, Solenogastres, Rhopalomeniidae), present in the north

coast of Iberian Peninsula

Oscar GARCÍA-ÁLVAREZ*, Victoriano URGORRI* y Eco. Javier CRISTOBO**

Recibido el15-IX-1999. Aceptado el24-XT-1999

RESUMEN

En la actualidad, dentro del proyecto FAUNA IBERICA, se está realizando el estudio de

los solenogastros de las costas ibéricas. En este trabajo se describe una especie casi des-

conocida para las aguas ibéricas, Rhopalomenia aglaopheniae (Kowalevsky y Marion,

1887), que solo había sido mencionada para aguas peninsulares en el Cap de Creus por

Pruvot en 1891.

ABSTRACT

A study on the solenogastres of the Iberian coast is currently underway within the project

FAUNA IBÉRICA. This paper offers a description of a species which is practically unknown

in Iberian waters, Rhopalomenia aglaopheniae (Kowalevsky and Marion, 1887), and

which has only been cited in the waters of the Iberian Peninsula on the Cap de Creus by

Pruvot in 1891.

PALABRAS CLAVE: Rhopalomenía algaopheniae, Mollusca, Solenogastres, Península Ibérica.

KEY WORDS: Rhopalomenia algaopheniae, Mollusca, Solenogastres, Iberian Peninsula.

INTRODUCCIÓN

Aunque los Moluscos Solenogastros

no constituyen un grupo zoológico raro,

ya que su presencia se extiende por

todos los fondos marinos, desde la costa

hasta las grandes profundidades, sí son

una clase poco conocida. Las informa-

ciones sobre su biología son escasas y

los datos sobre su diversidad y biogeo-

grafía son aún pobres y sobre todo desi-

guales. Esto se pone de manifiesto en las

aguas europeas, donde las costas de la

Península Ibérica muestran un vacío

muy notable en el conocimiento de esta

fauna (SALVINI-PLAWEN, 1997).

* Laboratorio de Zooloxía Mariña. Departamento de Bioloxía Animal. Facultade de Bioloxía. Universidade de

Santiago de Compostela.15706 Santiago de Compostela. baoscarOusc.es / bavitucoOusc.es

** Laboratorio de Biodiversidade e Recursos Mariños. Instituto de Acuicultura. Universidade de Santiago de

Compostela.15706 Santiago de Compostela. España. bafjcrisCVusc.es

1 Contribución científica del Instituto de Acuicultura n* 006/99

125

Iberus, 18 (1), 2000

j Península ñ

o Ibérica E

alo 6 Ñ >

Figura 1. Localización de la estación de recolección.

Figure 1. Map showing the location of the sampling station.

Fruto de las investigaciones, que,

dentro del proyecto FAUNA IBÉRICA, se

vienen realizando en el Atlántico y

Mediterráneo peninsulares, se recogió

en las costas cantábricas españolas el

solenogastro Rhopalomenia aglaopheniae

(Kowalevsky y Marion, 1887). Esta es

una especie ampliamente distribuida

tanto en el Mediterráneo, donde se

conoce del Sur de Grecia, Golfo de

Nápoles y Golfo de León (KOwALEVSKY

Y MARION, 1887; MALUQUER, 1917;

NIERSTRASZ Y STORK, 1940; PRUVOT,

1891; SALVINI-PLAWEN, 1972, 1997),

como en el Atlántico, donde existen

varias citas en las costas británica e

irlandesa, así como de Roscoff (GARS-

TANG, 1896; JONES, 1956; JONES Y BAXTER,

1987; SALVINI-PLAWEN, 1997; SEAWARD,

1992). En cambio, a excepción de la

única cita de PrRUVOT (1891) en Cap de

Creus, la Península Ibérica es una

laguna de conocimiento en cuanto a la

distribución europea de esta especie.

Por todo ello, nos parece de interés

acompañar esta nueva localización de

Rhopalomenia aglaopheniae, con una des-

cripción de los principales caracteres

anatómicos de la especie, a los que se

acompaña la reconstrucción de su orga-

126

nización interna anterior y posterior, así

como de fotografías y dibujos del

animal, de algunos cortes y de sus tipos

espiculares.

MATERIAL Y MÉTODOS

El ejemplar estudiado procede de la

Estación 165-A de la campaña de mues-

treo FAUNA Il, realizada en junio de

1991, a profundidades entre 30 y 1000

m., en las costas del N y NW de la

Península Ibérica, desde la zona orien-

tal de Guipúzcoa hasta las Islas Cíes,

incluyendo el Banco de Galicia. La Esta-

ción 165-A se localiza al oeste del Cabo

Peñas, norte de España (Fig. 1) (43? 43'

18” N - 43? 43' 46” N; 05% 55' 51” O - 05?

56" 37” O) en un fondo de roca y pie-

dras a una profundidad de 122-124 m.

El ejemplar tenía unas dimensiones de

12,2 mm de largo por 1,1 mm de ancho

y se conserva seccionado en cortes

seriados.

El ejemplar fue fijado y conservado

en alcohol al 70%. Se separaron peque-

ños trozos de cutícula de la parte media

del cuerpo y del surco ventral, para

obtener una representación de los dis-

GARCÍA-ÁLVAREZ ET AL.: Rhopalomenia aglaopheniae en el norte de la Península Ibérica

Figura 2. A: fotografía de Rhopalomenia aglaopheniae (Kowalevsky y Marion, 1887); B: microfoto-

grafía de una espícula acicular; C: espículas aciculares; D: espícula del surco pedio.

Figure 2. A: photograph of Rhopalomenia aglaopheniae (Kowalevsky and Marion, 1887); B: microp-

hotograph of acicular spicules; C: acicular spicules; D: spicules alongside the pedal groove.

tintos tipos espiculares. Estas piezas se

trataron con hipoclorito sódico al 5%

durante 12 horas para el aislamiento de

las espículas; posteriormente se lavaron

en agua destilada, se secaron en una

estufa a 40 “C y se montaron con resina

sintética. Para el estudio anatómico, los

RESULTADOS

ejemplares fueron descalcificados en

una solución de EDTA durante 12 horas,

se cortaron en parafina en series trans-

versales de 10 ym de sección. Se tiñó en

Azan de Heidenhain y se realizó la

reconstrucción anatómica a partir de los

cortes seriados.

Orden CAVIBELONIA Salvini-Plawen, 1978

Familia RHOPALOMENIIDAE Salvini-Plawen, 1978

Género Rhopalomenia Simroth, 1893

Rhopalomenia aglaopheniae (Kowalevsky y Marion, 1887)

Proneomenia aglaopheniae Kowalevsky y Marion, 1887 (denominación original), Rhopalomenia eisigi

Thiele, 1894.

Descripción

Habitus: Animal muy enrollado

sobre sí mismo, con el cuerpo liso, sin

abultamientos (Fig. 2 A). Surco ventral

bien visible. Color en alcohol amarillo.

Manto: Cutícula de 100 a 140 um de

grosor, con papilas pedunculadas y con

el extremo distal grueso. Las espículas

se disponen en varias capas estrecha-

mente entrelazadas. Son huecas, de

127

Iberus, 18 (1), 2000

Figura 3. Organización esquemática de la parte anterior del cuerpo de Rhapalomenia aglaopheniae

(Kowalevsky y Marion, 1887). At: órgano sensitivo atrial; Bg: ganglio bucal; Cg: ganglio cerebral;

Dc: ciego dorsal; Ma: manto; Mg; intestino; Ph: faringe; Pp: foseta pedia; Rs: saco radular; Vfg:

órgano glandular ventral de la faringe; Vg: ganglio ventral; A, B: cortes en sección correspondien-

tes a las líneas A, B.

Figure 3. Schematic organizaltion of the anterior body of Rhopalomenia aglaopheniae (Kowalevsky

and Marion, 1887). At: atrial sense organ; Bg: buccal ganglion; Cg: cerebral ganglion; Dc: dorsal

caecum; Ma: mantle; Mg: midgut; Ph: pharynx; Pp: pedal pit; Rs: radular sac; Vfg: ventral foregut

glandular organ; Vg: ventral ganglion; A, B: cross-section corresponding to lines A, B.

forma acicular, de hasta 150 um de lon-

gitud, levemente arqueadas, más anchas

en su parte central y con los extremos

romos (Fig. 2B, C). En el surco ventral se

sitúan escamas con forma de hoja de

cuchillo de hasta 75 ym de longitud

(Fig. 2D).

Surco pedio: Comienza en una foseta

ciliada (Fig. 3), situada debajo de la

faringe, que se comunica con el exterior

por una estrecha abertura. En el surco

pedio se encuentra un pliegue ciliado

que entra en la cavidad paleal.

Cavidad paleal: La cavidad paleal es

pequeña, no tiene pliegues respiratorios

y se comunica con el exterior por una

pequeña abertura ventro-terminal. El

conducto de desove desemboca, a través

de un orificio genital impar, en la parte

rostro-central de la cavidad paleal. Dor-

128

salmente al mismo se sitúa el ano. No

presenta ni espículas abdominales, ni

espículas copulatrices (Fig. 4).

Organos de los sentidos: El atrio pre-

senta en sus paredes frontal y laterales

numerosas papilas pequeñas e indivi-

dualizadas (Fig. 3). Dorsalmente tiene

dos pliegues ciliados que continúan

hasta la parte posterior del atrio. Posee

un solo órgano dorsoterminal, situado

en posición terminal, en el extremo pos-

terior del cuerpo (Fig. 4).

Aparato digestivo: La cavidad bucal

está separada del atrio por un pliegue,

continuándose en una faringe estrecha y

larga, que presenta ventralmente un

saco radular. No tiene rádula. En el saco

radular (Fig. 3) desemboca lateralmente

el par de órganos glandulares de la

faringe. Estos órganos glandulares, en

GARCÍA-ÁLVAREZ ET AL.: Rhopalomenia aglaophenzae en el norte de la Península Ibérica

250 um

Figura 4. Organización esquemática de la parte posterior del cuerpo de Rhopalomenia aglaopheniae

(Kowalevsky y Marion, 1887). Dso: órgano sensitivo dorsoterminal; Es: espermatozoides; Ht:

corazón; ; Ma: manto; Mg; intestino; Ov: óvulos; Pc: cavidad paleal; Pd: pericardioducto; Pr: peri-

cardio; Re: recto; Sc: comisura suprarrectal; Sd: conducto de desove; Sr: receptáculo seminal; A, B:

cortes en sección correspondientes a las líneas A, B.

Figure 4. Schematic organization of the posterior body of Rhopalomenia aglaopheniae (Kowalevsky

and Marion, 1887). Dso: dorsoterminal sense organ; Es: sperm; Ht: heart; Ma: mantle; Mg: midgut;

Ov: eggs; Pc: pallial cavity; Pd: pericardioduct; Pr: pericardium; Re: rectum; Sc: supra-rectal commis-

sure; Sd: spawning duct; Sr: seminal receptacle; A, B: cross-section corresponding to lines A, B.

su parte posterior, pertenecen al tipo A

(SALVINI-PLAWEN, 1978), ya que están

formados por un conducto muy largo en

el que se abren los folículos glandulares

subepiteliales, discurriendo ventral-

mente al intestino medio hasta la mitad

del cuerpo (Fig. 3, 3B); mientras que en

la parte anterior, cada órgano glandular

de la faringe, es del tipo C (SALVINI-

PLAWEN, 1978) con forma de una

burbuja de gran tamaño (Fig. 3, 3A).

Este par de burbujas se sitúan ventral-

mente al ciego rostral del intestino

medio y latero-dorsalmente a la faringe.

La faringe continúa durante un trecho

hasta desembocar por su parte dorsal en

el intestino medio. En esta misma zona

del intestino medio, parte rostralmente

un ciego dorsal, que llega hasta la parte

anterior del cuerpo (Fig. 3, 3A). El intes-

tino medio presenta en toda su longitud

numerosos constricciones seriadas en

sus paredes laterales y ventral. El recto

discurre bajo el pericardio (Fig. 4B) y se

abre en el ano, que desemboca dorsal-

mente en la parte rostral de la cavidad

paleal (Fig. 4).

Sistema nervioso: El ganglio cerebral es

aplanado dorso-ventralmente, estando

situado ventralmente al ciego rostral del

intestino medio y dorsalmente a la

cavidad bucal (Fig. 3). El par de ganglios

ventrales se encuentran dorsalmente a la

foseta pedia, y a ambos lados del saco

radular están el par de ganglios bucales

(Fig. 3). En la parte terminal del cuerpo,

cerca del ano, está la comisura suprarrec-

tal (150 ym de longitud) (Fig. 4).

Aparato circulatorio: El corazón,

situado en la pared dorsal del pericar-

122%)

Iberus, 18 (1), 2000

dio, tiene forma tubular (Fig. 3). Las

células sanguíneas son de dos tipos: gra-

nulocitos redondeados con un diámetro

de 7,5 ym, y eritrocitos alargados y sin

gránulos de hasta 20 um de longitud.

Aparato reproductor: El par de

gónadas se extienden sobre el intestino

medio. El ejemplar examinado las pre-

sentaba llenas de óvulos y espermatozoi-

des (Fig. 4A). Las gónadas se unen con el

pericardio a través de un par de gonope-

ricardioductos. El pericardio es volumi-

noso (Fig. 4B), posteriormente se conti-

núa en dos pericardioductos, que se

curvan y se dirigen anteriormente hasta

unirse con los conductos de desove (Fig.

4). Posee un par de receptáculos semina-

les situados dorsalmente a los conductos

de desove (Fig. 4, 4B), que se encontra-

ban llenos de espermatozoides. Estos

receptáculos no parecen ser órganos par-

ticulares, sinó que cada uno es solamente

una parte encorvada del conducto de

desove (SALVINI-PLAWEN, 1972). Los dos

conductos de desove tienen las paredes

glandulares (Fig. 4B) y se fusionan en un

único conducto, que desemboca impar,

ventralmente en la pared rostral de la

cavidad paleal (Fig. 4).

Distribución: Sur del Peloponeso

(fuera del cabo Matapan/Tainaron)

(Grecia) (SALVINI-PLAWEN, 1972, 1997);

DISCUSIÓN

El ejemplar estudiado pertenece al or-

den Cavibelonia por sus espículas acicu-

lares huecas, dispuestas en varias capas

dentro de una cutícula gruesa con papilas

epidérmicas. Se clasifica dentro de la fa-

milia Rhopalomeniidae, por sus órganos

glandulares ventrales de la faringe sube-

piteliales de tipo A, ausencia de rádula y

de pliegues respiratorios en la cavidad

paleal. Las características genéricas que

lo sitúan dentro de Rhopalomenia están

bien definidas en este ejemplar: la cavi-

dad bucal está separada del atrio, la de-

sembocadura del conducto de desove es

impar, presenta un órgano sensitivo dor-

soterminal y carece de espículas copula-

trices (SALVINEPLAWEN, 1967, 1978).

130

Golfo de Nápoles (Italia), asociada al

hidroideo Lytocarpia myriophyllum, a 50-

60 m de profundidad (NIERSTRASZ Y

STORK, 1940); Marsella (Francia), en

fondos rocosos, asociada al hidroideo

Lytocarpia myriophyllum, a 50-60 m de

profundidad (KOwALEVSKY Y MARION,

1887); Banyuls (Francia), en fondos

limosos, asociada al hidroideo Lytocarpia

myriophyllum, a 60-80 m de profundidad

(Pruvor, 1891); costa del Rosellón

(Francia), sobre Lytocarpia myriophyllum,

a 60-80 m de profundidad (MALUQUER,

1917); Isla Portaló (Cap de Creus,

España), sobre Lytocarpia myriophyllum,

a 80 m de profundidad (Pruvor, 1891);

Roscoff (Francia) (SALVINI-PLAWEN,

1997); Plymouth (Gran Bretaña), aso-

ciada al hidroideo Lytocarpia myriophy-

llum, a 47-57 m de profundidad (GARs-

TANG, 1896); Isla de Man (Gran Bretaña),

en fondos de arena fangosa, a 58-71 m

de profundidad (JONES, 1956); costas

este y oeste de Gran Bretaña, asociada al

hidroideo Lytocarpia (Theocarpus)

moyriophyllum, a 50 m o más de profundi-

dad (JONES Y BAXTER, 1987); costas este y

oeste de Gran Bretaña, Mar de Irlanda y

costas norte y sur de Irlanda (SEAWARD,

1992); Liberia (Africa), a 70 m de pro-

fundidad (THIELE, 1906), aunque es una

cita que necesita un nuevo examen y

confirmación (SALVINI-PLAWEN, 1997).

Esta especie, además de su descrip-

ción original de KOWALEVSKY Y MARION

(1887), fue redescrita por NIERSTRASZ Y

STORK (1940) y posteriormente SALVINI-

PLAWEN (1978) ofrece, en una tabla com-

parativa con las especies antárticas del

género, el conjunto de caracteres especí-

ficos más significativos. En nuestro

ejemplar se pueden observar todos los

rasgos que lo identifican como Rhopalo-

menia aglaopheniae (Kowalevsky y

Marion, 1887), únicamente no hemos

encontrado el botón sensitivo anterior

descrito por KOwALEVSKY Y MARION

(1887).

Rhopalomenia aglaopheniae (Kowa-

levsky y Marion, 1887), presenta una

GARCÍA-ÁLVAREZ ET AL.: Rhopalomenia aglaopheniae en el norte de la Península Ibérica

distribución, que va desde el Mediterrá-

neo oriental hasta las costas atlánticas

europeas, asociada al hidroideo Lytocar-

pia myriophyllum, a una profundidad de

47-137 m (SALVINEPLAWEN, 1990, 1997).

El ejemplar aquí estudiado fue recogido

al oeste del Cabo Peñas (Golfo de

Vizcaya, norte de España) a una profun-

didad de 122-124 m. Esta especie sólo

había sido señalada anteriormente en

aguas de la Península Ibérica en las

costas españolas mediterráneas en el

Cap de Creus (Pruvor, 1891), por lo que

esta cita representa la primera en las

costas atlánticas ibéricas.

Del género Rhopalomenia Simroth,

1893, además de Rhopalomenia aglaophe-

níae, se conocen en la actualidad otras 6

especies. Cinco de ellas son antárticas y

con un conjunto de caracteres específi-

cos bien diferenciados de Rhopalomenia

aglaopheniae (Tabla 4, pág. 159 en

SALVINEPLAWEN, 1978). La única especie

del género, próxima al entorno biogeo-

BIBLIOGRAFÍA

GARSTANG, W., 1896. On the Aplacophorous :

Amphineura of the British Isles. Proc. Mala-

col. Soc. London, 2: 123-125.

KOWALEVSKY, A. Y MARION, A., 1887. Contri-

butions a l' histoire des solénogastres ou apla-

cophores. Ann. Mus. Hist. nat. Marseille, Zool.,

3 (1): 1-77.

Jones, N.S., 1956. The fauna and biomass of

muddy sand deposit off port Erin, Isle of

Man. J. Aním. Ecol., 25: 217-252.

JONES, A.M. Y BAXTER, J.M., 1987. Mollusca: Cau-

dofoveata, Solenogastros, Polyplacophora and

Scaphopoda. Synopses of the British Fauna, 37:

1-37

LELOUr, E., 1948. Un nouveau solénogastre pro-

néoméniidae, Entonomenia atlantica g. nov., sp.

nov. Bull. Mus. Roy. Hist. Natur. Belgique, 24

(EN ETL

MALUQUER, J., 1917. Notes para l'estudi dels So-

lenogastros (Mollusca anfineures) de Cata-

lunya. Treballs Inst. catal. Hist. nat. (Barce-

lona), 3: 9-53.

NIERSTRASZ, H.F. Y STORK, H.A., 1940. Mono-

graphie der Solenogastren des Golfes von

Neapel. Zoologica (Stuttgart), 99: 1-92.

Pruvor, G., 1891. Sur l'organisation de quelques

néomeniens des cótes de France. Arch. Zool.

Exptl. Gén., sr. 2, 9: 699-810.

gráfico de Rhopalomenia aglopheniae, es

Rhopalomenia atlantica (Leloup, 1948).

Fue recogida en aguas de Tenerife (Islas

Canarias), a una profundidad de 540 m,

asociada a hidroideos de la Familia

Lafoeidae. Se diferencia de Rhopalomenia

aglaopheniae fundamentalmente porque

tiene los órganos glandulares de la

faringe muy cortos y sólo de tipo A, no

presenta saco radular y la faringe es

muy corta (LELOUP, 1948; SALVINI-

PLAWEN, 1972, 1978).

AGRADECIMIENTOS

Al Prof. Dr. Luitfried v. Salvini-

Plawen de la Universidad de Viena por

su inestimable ayuda. Este trabajo se

realizó dentro de los proyectos de inves-

tigación: Fauna Ibérica III (PB92-0121); y

las Acciones Integradas de Cooperación

Hispano-Austríaca (HU1995-0002;

HU1996-0002; HU1997-0002).

SALVINI-PLAWEN, L.v., 1967. Kritische Bemer-

kungen zum System der Solenogastres (Mo-

llusca, Aculifera). Zeitschr. zool. Syst. Evolut.-

forsch., 5 (4): 398-444.

SALVINI-PLAWEN, L.v., 1972. Revision der mo-

negassischen Solenosgastres (Mollusca, Acu-

lifera). Zeitschr. zool. Syst. Evolut.-forsch., 10 (3):

215-250.

SALVINI-PLAWEN, L.v., 1978. Antarktische und

subantarktische Solenogastres. Eine Mono-

graphie: 1898-1974. Zoologica (Stuttgart), 128:

1-315.

SALVINEPLAWEN, L.v., 1990. The status of the

Caudofoveata and the Solenogastres in the

Mediterranean Sea. Lavori S.I.M. Napoli, 23:

5-30.

SALVINI-PLAWEN, L.v., 1997. Fragmented know-

ledge on West-European and Iberian Cau-

dofoveata and Solenogastros. Iberus, 15 (2):

35-50.

SEAWARD, D.R,, 1992. Distribution of the marine

molluscs of north west Europe. Nature Con-

servancy Council, Peterborough, 1- 105.

THIELE, J., 1906. Archaeomenia prisca n. g. n. sp.

Wissensch Ergebnisse Dtsch. Tiefsee Exped. Val-

divia, 1898/99, 9: 317-324.

131

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Sinonimias

Doris limbata Cuvier, 1804, Ann. Mus. H. N. Paris, 4 (24): 468-469 [Localidad tipo: Marsella].

Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop. Car., 10: 275.

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Fretter, V. y Graham, A., 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp.

Ponder, W. F., 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. En Ponder, W. F.

(Ed.): Prosobranch Phylogeny, Malacological Review, suppl. 4: 129-166.

Ros, J., 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas.

Miscelánea Zoolgica, 3 (5): 21-51.

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INSTRUCTIONS TO AUTHORS

e Tberus publishes research papers, notes and monographs devoted to the various aspects of Malacology. Papers are manus-

cripts of more than 5 typed pages, including figures and tables. Notes are shorter papers. Monographs should exceed

50 pages of the final periodical, and will be published as Supplements. Authors wishing to publish monographs should

contact the Editor. Manuscripts are considered on the understanding that their contents have not appeared or will not

appeared, elsewhere in substantially the same or any abbreviated form.

+ Manuscripts and correspondence regarding editorial matters must be sent to: Dr. Ángel Guerra Sierra, Editor de Publi-

caciones, Instituto de Investigaciones Marinas (CSIC), C/Eduardo Cabello 6, 36208 Vigo, Spain.

» Manuscripts may be written in any modern language.

* When a paper exceeds 20 pages, extra pages will be charged to the author(s) at full cost.

+ Manuscripts must be typed double spaced (including the references, figure captions and tables) on one side on A-4

(297x210 mm) with margins of at least 3 cm. An original and two copies must be submitted. When a paper has joint

authorship, one author must accept responsability for all correspondence.

* Papers should conform the following, layout:

First page. This must include a concise but informative title, with mention of family of higher taxon when appropriatte,

and its Spanish translation. It will be followed by all authors? names and surnames, their full adress(es), an abstract (and

its Spanish translation) not exceeding 200 words which summarizes not only contents but results and conclusions, and

a list of Key Words (and their Spanish translation) under which the article should be indexed.

Following pages. These should content the rest of the paper, divided into sections under short headings. Whenever pos-

sible the text should be arranged as follows: Introduction, Material and methods, Results, Discussion, Conclusions,

Acknowledgements and References. Unusual abbreviations used in the text must be grouped in one alphabetic sequence

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+ Notes should follow the same layout, without the abstract.

* Footnotes and cross-references must be avoided. The International Codes of Zoological and Botanical Nomencla-

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this example (please note the punctuation):

Dendrodoris limbata (Cuvier, 1804)

Synonyms

Doris limbata Cuvier, 1804, Ann. Mus. H. N. Paris, 4 (24): 468-469 [Type locality: Marseille].

Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop. Car., 10: 275.

These references must not be included in the Bibliography list, except if referred to elsewhere in the text. Ifa full list

of references of the taxon is to be given immediately below it, the same layout should be followed (also excluding those

nowhere else cited from the Bibliography list).

Only Latin words and names of genera and species should be underlined once or be given in ¿talics. No word must

be written in UPPER CASE LETTERS. SI units are to be used, together with their appropriate symbols. In Spanish

manuscripts, decimal numbers must be separated with a comma (,), NEVER with a point (.) or upper comma (/).

» References in the text should be written in small letters or SMALL CAPITALS: Fretter 82 Graham (1962) or FRETTER

82 GRAHAM (1962). The first mention in the text of a paper with more than two authors must include all of them

[Smith, Jones 2 Brown (1970)], thereafter use et al. [Smith et al. (1970)]. Ifan author has published more than one

paper per year, refer to them with letters: (Davis, 1989a; Davis, 1989b). Avoid op. cit.

The references in the reference list should be in alphabetical order and include all the publications cited in the text but

only these. ALL the authors of a paper must be included. These should be written in small letters or SMALL CAPITALS.

The references need not be cited when the author and date are given only as authority for a taxonomic name. Titles of

periodicals must be given IN FULL, not abbreviated. For books, give the title, name of publisher, place of publication,

indication of edition if not the first and total number of pages. Keep references to doctoral theses or any other unpu-

blished documents to an absolute minimum. See the following examples (please note the punctuation):

Eretter, V. and Graham, A., 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp.

Ponder, W. F., 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. In Ponder, W. EF. (Ed.):

Prosobranch Phylogeny, Malacological Review, suppl. 4: 129-166.

Ros, J., 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Miscelá-

nea Zoológica, 3 (5): 21-51.

e Figures must be original, in Indian ink on draughtsman's tracing paper. Keep in mind page format and column size

when designing figures. These should be one column (57 mm) or two columns (120 mm) wide and up 194 mm high,

or be proportional to these sizes. Two columns format is recomended. It is desirable to print figures with their legend

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contrast, and should be submitted in the final size. When mounting photographs in a block, ensure spacers are of uni-

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Figure 1. Neodoris carvi. A: animal crawling; B: rinophore; C: gills.

If abbreviations are to be used in illustrations, group them alphabetically after the Legends for Figures section.

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rule, keep the number and extension of illustrations and tables as reduced as possible.

+ Manuscripts that do not conform to these instructions will be returned for correction before reviewing.

» Authors submitting manuscripts will receive an acknowledgement of receipt, including receipt date, and the date the

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* Proofs will be sent to the author for correcting errors. At this stage no stylistic changes will be accepted. Pay special

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e Fifty reprints per article will be supplied free of charge. Additional reprints must be ordered when the page proofs are

returned, and will be charged at cost. NO LATER orders will be accepted.

NS E

A ADA

el ta

ña E

rote csid

Mee A

PAN

Din aii

LA SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

Junta directiva desde el 18 de octubre de 1996

Presidente Emilio Rolán Mosquera

Vicepresidente Diego Moreno Lampreave

Secretario Luis Murillo Guillén

Tesorero Jorge J. Otero Schmitt

Avda. de las Ciencias s/n, Campus Universitario, 15706 Santiago

de Compostela, España

Editor de Publicaciones Ángel Guerra Sierra

Instituto de Investigaciones Marinas, c/ Eduardo Cabello 6, 36208

Vigo, España

Bibliotecario Rafael Araujo Armero

Museo Nacional de Ciencias Naturales, CSIC, c/ José Gutierrez

Abascal 2, 28006 Madrid, España

Vocales Eugenia María Martínez Cueto-Felgueroso

María de los Ángeles Ramos Sánchez

Francisco Javier Rocha Valdés

Gonzalo Rodríguez Casero

Jesús Souza Troncoso

José Templado González

La Sociedad Española de Malacología se fundó el 21 de agosto de 1980. La sociedad se registró como una aso-

ciación sin ánimo de lucro en Madrid (Registro N“ 4053) con unos estatutos que fueron aprobados el 12 de

diciembre de 1980. Esta sociedad se constituye con el fin de fomentar y difundir los estudios malacológicos

mediante reuniones y publicaciones. A esta sociedad puede pertenecer cualquier persona o institución interesada

en el estudio de los moluscos.

SEDE SOCIAL: Museo Nacional de Ciencias Naturales, c/ José Gutierrez Abascal 2, 28006 Madrid, España.

CUOTAS PARA 2000:

Socio numerario (en España): 5.500 ptas. (= 33,06 euros)

(en extranjero): 7.000 ptas (= 42,07 euros)

Socio estudiante (en España): 2.500 ptas. (= 15,03 euros)

(en extranjero): 3.500 ptas (= 21,04 euros)

Socio Familiar: 500 ptas. (= 3 euros)

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(en extranjero): 8.000 ptas (= 48,08 euros)

INSCRIPCIÓN: 1.000 ptas. (= 6,01 euros) además de la cuota correspondiente.

A los socios residentes en España se les aconseja domiciliar su cuota. Todos los abonos deberán enviarse al

Tesorero (dirección reseñada anteriormente) el 1 de enero de cada año. Los abonos se harán sin recargos para la

sociedad y en favor de la Sociedad Española de Malacología y no de ninguna persona de la junta directiva. Aque-

llos socios que no abonen su cuota anual dejarán de recibir las publicaciones de la Sociedad. Los bonos de ins-

cripción se enviarán junto con el abono de una cuota anual al Tesorero.

Members living in foreing countries can deduce 6 euros if paid before 15 April.

Cada socio tiene derecho a recibir anualmente los números de /berus, Reseñas Malacológicas y Noticiarios que

se publiquen.

ÍNDICE

Iberus 18 (1) 2000

SANCHEZ-TOCINO, L., OCAÑA, A. Y GARCÍA, E J. Contribución al conocimiento de los Moluscos

Opistobranquios de la costa de Granada (sureste de la Península Ibérica)

Contribution to the knowledge of the Opisthobranch Molluscs from the coast of Granada (sout-

DIAS LO ER NA A o e 1-14

SMRIGLIO, C. AND MARIOTTINI, P. Onoba olzverioi n. sp. (Prosobranchia, Rissoidae), a new gas-

tropod from the Mediterranean

Onoba oliverioi ». sp. (Prosobranchia, Rissoidae), un nuevo gasterópodo para el Mediterrá-

O A A A E E E NI 15119)

ROLAN, E. AND LUQUE, A. A. The subfamily Rissoininae (Mollusca: Gastropoda: Rissoidae) in

the Cape Verde Archipelago (West Africa)

La subfamilia Rissoininae (Mollusca: Gastropoda: Rissoidae) en el archipiélago de Cabo Verde

E A O A 21-94

RUEDA, J., SALAS, C. AND GOFAS, S. A molluscan community from coastal bioclastic bottoms in

the Strait of Gibraltar area

La comunidad de moluscos de un fondo bioclástico costero del Estrecho de Gibraltar .. 95-123

GARCÍA-ÁLVAREZ, O., URGORRI, V. Y CRISTOBO, E J. Rhopalomenia aglaopheniae (Kowalevsky y

Marion, 1887) (Mollusca, Solenogastres, Rhopalomeniidae), presente en la costa norte de la Penín-

sula Ibérica

Rhopalomenia aglaopheniae (Kowalevsky and Marion, 1887) (Mollusca, Solenogastres, Rho-

palomentidae), present in the north coast of Iberian Peninsula oo... 125-131

ISSN 0212-3010

Iberus

REVISTA DE LA

SOCIEDAD ESPAÑOLA

DE MALACOLOGÍA

Oviedo, diciembre 2000

Vol. 18 (2)

ComiITÉ DE REDACCIÓN

EDITOR

Ángel Guerra Sierra

EDITORES ADJUNTOS

Eugenia M? Martínez Cueto-Felgueroso

Francisco Javier Rocha Valdés

Gonzalo Rodríguez Casero

ComiTÉ EDITORIAL

Kepa Altonaga Sustacha

Eduardo Angulo Pinedo

Rafael Araujo Armero

Thierry Backeljau

Rúdiger Bieler

Sigurd v. Boletzky

Jose Castillejo Murillo

Karl Edlinger

Antonio M. de Frias Martins

José Carlos García Gómez

Edmund Gittenberger

Serge Gofas

Gerhard Hoszprunar

Yuri |. Kantor

Ángel Antonio Luque del Villar

María Yolanda Manga González

Jordi Martinell Callico

Ron K. 0'Dor

Tokashi Okutani

Marco Oliverio

Pablo E. Penchaszadeh

Winston F. Ponder

Corlos Enrique Prieto Sierra

Me de los Ángeles Ramos Sánchez

Paul G. Rodhouse

Joandoménec Ros i Aragones

María Carmen Salas Casanovas

Gerhard Steiner

José Templado González

Victoriano Urgorri Carrasco

Anders Warén

PORTADA DE /berus

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

Universidad del Poís Vasco, Bilbao, España

Museo Nacional de Ciencias Naturales, Madrid, España

Institut Royal des Sciences Naturelles de Belgique, Bruselas, Bélgica

The Field Museum, Chicago, Estados Unidos

Laboratoire Arago, Banyuls-sur-Mer, Francia

Universidad de Santiago de Compostela, Santiago de Compostela, España

Naturhistorisches Museum Wien, Viena, Austria

Universidade dos Acores, Acores, Portugal

Universidad de Sevilla, Sevilla, España

National Natuurhistorisch Museum, Leiden, Holanda

Universidad de Málaga, España

Zoologische Staatssammlung Múnchen, Múnchen, Alemania

AN. Severtzov Institute of Ecology and Evolution, Moscú, Rusia

Universidad Autónoma de Madrid, Madrid, España

Estación Agrícola Experimental, CSIC, León, España

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Dalhousie University, Halifax, Canada

Nihon University, Fujisawa City, Japón

Universitá di Roma “La Sapienza”, Roma, ltalia

Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina

Australian Museum, Sydney, Australia

Universidad del País Vasco, Bilbao, España

Museo Nacional de Ciencias Naturales, CSIC, Madrid, España

British Antarctic Survey, Cambridge, Reino Unido

Universidad de Barcelona, Barcelona, España

Universidad de Málaga, Málaga, España

Institut fir Zoologie der Universitút Wien, Viena, Austria

Museo Nacional de Ciencias Naturales, ESIC, Madrid, España

Universidad de Santiago de Compostela, Santiago de Compostela, España

Swedish Museum of Natural History, Estocolmo, Suecia

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

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

Iberus

ENITASON fa

JUL 122001

REVISTADELA.

SOCIEDAD ESPAÑOLA

DE MALACOLOGÍA

Vol. 18 (2) Oviedo, diciembre 2000

Iberus

Revista de la

SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

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

admiten también notas breves. /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

cuidadosamente 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 diri-

girse al bibliotecario.

Los no socios deberán ponerse en contacto con BACKHUYS PUBLISHERS, PO. Box 321,

2300 AH Leiden, The Netherlands. Tel.: +31-71-51 70 208, Fax: +31-71-51 71 856, Correo Elec-

trónico: backhuysCeuronet.nl

Los resumenes de los artículos editados en esta revista se publican en Aquatic Science

and Fisheries Abstracts (ASFA) y en el Zoological Records, BIOSIS.

Contents list published in Aquatic Science and Fisheries Abstracts and Zoological Records,

BIOSIS.

Dep. Leg. B-43072-81

ISSN 0212-3010

Diseño y maquetación: Gonzalo Rodríguez

Impresión: LOREDO, S. L. - Gijón

O Sociedad Española de Malacología ——_____—_—_—_———— lIberus, 18 (2): 1-15, 2000

Nuevos datos anatómicos y taxonómicos del género Chiton

Linnaeus, 1758 (Mollusca, Polyplacophora) en la Península

Ibérica

New anatomical and taxonomical data on the Genus Chiton

Linnaeus, 1758 (Mollusca, Polyplacophora) from the Iberian

Peninsula

Pilar CARMONA ZALVIDE?, Francisco J. GARCÍA? y Victoriano URGORRI **

Recibido el 8-VI-1999. Aceptado el 22-X1-1999

RESUMEN

Se presentan aspectos relacionados con la morfología y taxonomía de las especies

del género Chiton Linnaeus, 1758 en la Península Ibérica. Para cada una de las espe-

cies estudiadas, Ch. olivaceus Spengler, 1797, Ch. corallinus (Risso, 1826) y Ch.

phaseolinus Monterosato, 1879, se aportan datos sobre la concha, perinoto y rádula.

Finalmente se realiza una discusión sobre la sistemática de este género y otros rela-

cionados con él, y se acepta que Rhyssoplax Thiele, 1893 constituye un subgénero

de Chiton.

ABSTRACT

Aspects related to the morphology and taxonomy of the species belonging to the genus

Chiton Linnaeus, 1758 in Iberian Peninsula are presented. Anatomical data of the shell,

perinotum and radula of Ch. olivaceus Spengler, 1797, Ch. corallinus (Risso, 1826) and

Ch. phaseolinus Monterosato, 1879 are included. A systematic discussion on the genus

Chiton and other related genera is included. Rhyssoplax Thiele, 1893 ¡is proposed as a

subgenus of Chiton.

PALABRAS CLAVES: Mollusca, Polyplacophora, Taxonomía, Anatomía, Chiton (Rhyssoplax) olivaceus, Chiton

(Rhyssoplax) corallinus, Chiton (Rhyssoplax) phaseolinus.

Key WokrDs: Mollusca, Polyplacophora, Taxonomy, Anatomy, Chiton (Rhyssoplax) olivaceus, Chiton

(Rhyssoplax) corallinus, Chiton (Rhyssoplax) phaseolinus

INTRODUCCIÓN

El género Chiton Linnaeus, 1758 ha las posiciones subgenéricas y las

ocasionado controversias taxonómicas admiten como géneros. No obstante

debido a que ciertos autores no aceptan desde su descripción, se han establecido

* Departamento de Fisiología y Biología Animal, Facultad de Biología, Univ. Sevilla; Avda. Reina Mercedes, 6;

Apdo. 1095, 41080 Sevilla. (Spain). E-mail: fjgarcia

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

Compostela (Spain). E-mail: bavitucoCusc. es

Iberus, 18 (2), 2000

diferentes subgéneros en éste. PILSBRY

(1892) consideró en el género Chiton las

secciones Chiton s. s., Radsia Gray, 1847 y

Sclerochiton Carpenter 1873.

THIELE (1893), asimismo, describe

nuevos subgéneros y secciones para

Chiton: Amaurochiton, Chondroplax, Sy-

pharochiton, Triloplax, y Anthochiton.

IREDALE Y HULL (1926) no admiten los

subgéneros para Chiton y consideran

que las diferencias que presentan los

distintos grupos justificarían su posición

como géneros independientes e incluso

aportan los siguientes géneros nuevos:

Delicatoplax, Tegulaplax, Mucroquasma, a

la vez que mantienen a los géneros

Rhyssoplax Thiele, 1893, Clavarizona

Hull, 1923, Amaurochiton, Sypharochiton,

Liolophura Pilsbry, 1892, Amphitomura

Pilsbry, 1892, Acantopleura Guilding,

1829, Onithochiton Gray, 1847, Lucilina

Dall, 1882 y Schizochiton Gray, 1847.

Posteriormente, THIELE (1929) desa-

rrolla una nueva clasificación, en la que

admite subgéneros y secciones, de

manera que en el género Chiton esta-

blece los subgéneros: Chiton y Rhysso-

plax. En el primero incluye las secciones:

Chiton s. s., Amaurochiton, Diochiton,

Chondroplax, Radsia y Sypharochiton; y en

Rhyssoplax las secciones: Anthochiton,

Delicatoplax, Tegulaplax, Rhyssoplax s. s. y

Mucroquasma.

En el presente trabajo se ha reali-

zado un estudio anatómico de las espe-

cies de Chiton (Rhyssoplax) que se distri-

RESULTADOS

buyen en la Península Ibérica, Chiton

(Rhyssoplax) olivaceus Spengler, 1797,

Chiton (Rhyssoplax) corallinus (Risso,

1826) y Chiton (Rhyssoplax) phaseolinus

Monterosato, 1879. Así mismo, se ha

discutido la categoría taxonómica de

Rhyssoplax.

MATERIAL Y MÉTODOS

Los ejemplares estudiados se han

recolectado desde Lisboa (Portugal)

hasta Punta Europa (Gibraltar) y, a su

vez, se han estudiado los ejemplares del

Museo Nacional de Ciencias Naturales

de Madrid (MNCN). En el apartado de

material estudiado, se especifica la loca-

lidad, número de ejemplares, tamaño

máximo de éstos, la fecha de la recolec-

ción y la profundidad.

El material se ha capturado de forma

directa tanto en la zona mediolitoral

como infralitoral, utilizándose en esta

última equipos de inmersión autóno-

mos. Los ejemplares se relajaron con

cristales de mentol y se fijaron entre dos

portas con etanol absoluto, conserván-

dose posteriormente en alcohol 70%.

Para el estudio de las partes duras se

introdujeron los ejemplares en KOH

10%, separándose las placas, escamas y

espículas del perinoto y la rádula. La

estructura de las diferentes partes se

observó mediante microscopía electró-

nica de barrido (Philips XL-20).

Chiton (Rhyssoplax) olivaceus Spengler, 1797

Chiton olivaceus Spengler, 1797, Skrivt. Naturh. Selsk., 4: 73, pl 6 f 8a-c. [Localidad tipo: Mar Medi-

terráneo)].

Chiton squamosus Poli 1791 no Linneo 1764, Test. Utr. Sicil., 1: 8, pl 3 figs. 21, 22 [Localidad tipo:

Sicilia]

Chiton sulcatus Risso 1826, Hist. Nat, Eur. Mérd., 4: 268 [Localidad tipo: Niza]

Chiton siculus Gray 1828, Spicil. Zool., 1: 5 [Localidad tipo: Mar Mediterráneo]

Chiton polii Deshayes 1833, Exp. Sci. Morée, 3 Moll.: 132 [Localidad tipo: Peloponeso]

Chiton subdivisus Renier en Monterosato 1879, G. Sci. nat. Econ. Palermo, 14: 7

Chiton striatus Chierghini MS, en Nardo, 1847, Ipsa Chieregh. Conch.: 44 [Localidad tipo: Laguna

de Venecia]

Chiton estuarii Chiereghini MS, en Nardo, 1847, Ipsa Chieregh. Conch.: 44 [Localidad tipo: Venecia]

Chiton squammulosus Doilfus 1883, Feuille. Jeun. Nat.: 3. [Localidad tipo: Palavas, Hérault, Francia]

CARMONA ZALVIDE ET 4L.: Nuevos datos del género Chiton en la Península Ibérica

Sos

Figura 1. Chiton (R.) olivaceus. A: valva 1; B: valva IV; C: valva VII; D: articulamento de valva

intermedia; E: dientes pectinados del articulamento; F: ornamentación del tegmento; G: costillas

de la zona jugal; H: costillas de la zona pleural; I: disposición de las megaloestetas y microestetas.

Figure 1. Chiton (R.) olivaceus. A: valve [; B: valve IV; C: valve VIH; D: articulamentum of an inter-

mediate valve; E: pectinated teeth from articulamentum; F: ornamentation of tegmentum; G: jugal

area; H: pleural area; I: disposition of megalaesthetes and micraesthetes.

Iberus, 18 (2), 2000

Material estudiado: El total de ejemplares estudiados ha sido de 275. Portinho de Arrabida, Por-

tugal: 1 ej., 24 x 12 mm, IX/95 (10 m). Porto Covo, Portugal: 1 ej., 14 x 8 mm, VII /94 (8 m). Praia

do Lagos, Portugal: 2 ej., 14,5 x 6,8 mm, VI /93 (17 m). Praia da Marinha, Portugal: 6 ej. 22,4x 11,2

mm, VIII/88 (Intermareal). Almacao de Pera, Portugal: 2 ej., 16, 2x 9 mm, VIII /93 (22 m). El Arre-

cifillo, Conil, Cádiz: 2 ej., 17 x 10 mm, VI /92 (12 m). Isla del Tajo, Cádiz: 1 ej., 19 x 11 mm, VI /92

(8 m). Playa del Chorro, Cádiz: 5 ej., 28, 5 x 15 mm, VI /92 (Intermareal). Isla de Tarifa, Cádiz: 7 ej.,

16 x 8 mm, 111/91; 2 ej., 27 x 11 mm, VII/91 (Intermareal). Isla de las Palomas, Cádiz: 1 ej., 18 x 11

mum, VII/91; 1 ej., 21 x 12 mm, VIII/91; 5 ej., 25,5 x 16 mm, IX/92 (3-12m). Punta Carnero, Cádiz: 1

ej., 30 x 17 mm, V/91; 3 ej., 18 x 11 mm, VIII/91 (Intermareal). La Ballenera, Cádiz: 10 ej., 21 x 13

mm, IV/91; 1 ej., 14x 7 mm, VII/91; 1 ej., 16,5 x 10,5 mm, VI/91; 5 ej., 19 x 10 mm, VII /91; 8 ej.,

12 x 8,2 mm, IV/92 (12 m). Punta de San García, Cádiz: 1 ej., 16 x 9 mm, VIL/90; 1 ej., 10,5 x 6 mm,

VI /91; 2 ej., 8x5 mm, VI /91; 3 ej., 16 x 8,5 mm, IX/92; 12 ej. 21 x 13 mm, 1X/92 (12 m). Ensenada

de Cucareo, Cádiz: 11 ej., 28,5 x 15 mm, IX/92; 10 ej., 18 x 9, 5 mm, 1/93). Puerto de Algeciras, Cádiz:

2 ej., 18 x 11 mm, VI/91; 3 ej., 24 x 12 mm, VIL/91; 21 ej., 18 x 11 mm, vIn/91; 2 ej., 16x 9 mm,

VII/92 (10 m). Punta de Paredones, Cádiz: 3 ej., 8 x 6 mm, VIL/90; 12 ej., 22 x 13 mm, VII /91; 1 ej.,

8 x 4 mm, VIII/91 (16 m). El Rinconcillo, Cádiz: 1 ej., 15 x 9 mm, VIL/90; 3 ej., 12,5 x 8 mm, VIL/91

(25 m). Punta del Gallo del Mirador, Cádiz: 5 ej., 9 x 4,5 mm, 1/93 (3-6 m). Campamento, Cádiz: 1

ej., 17x 10 mm, VIL/90; 8 ej., 17x 9 mm, 111/91 (Intermareal). Crinavis, Cádiz: 1 ej., 15x 9 mm, VIL/90;

l ej. 19x 11 mm, 11/91; 1 ej., 18x 11 mm, VI/91;5 ej. 19x 11 mm, VII/91;3 ej. 22x 11 mm, VII /91;

l ej. 24 x 14 mm, I1/92; 53 ej., 21 x 12 mm, IX/93 (33 m). San Felipe, Cádiz: 3 ej., 25x 12 mm, VI /91;

11 ej. 17 x 11 mm, VII /91; 1 ej., 106 mm, X/91 (17 m). Punta Europa, Gibraltar: 2 ej., 22 x 10 mm,

VII /92 (20-25 m). MNCN: N* 1503 /138: 1 ej., 11 x5.5 mm, (Cabrera, Baleares) (Colección: Hidalgo).

1503 /230: 1 ej., 24 x 12 mm, 12/08/84 (La Herradura, Granada) (A. Luque). 1503/356: 1 ej., 21 x 11

mm, (Cádiz) (Colección: Hidalgo). 1503/364: 3 ej., 26 x 13 mm, (Valencia) (Colección: Hidalgo).

1503 /424: 5 ej., 35 x 18 mm, (Mahón, Menorca) (Colección: Hidalgo). 1503/425: 1 ej., 22 x 11 mm,

(Palma) (Colección: Hidalgo). 1503 /433: 2 ej., 22x 15 mm, (Pto. Pollensa, Mallorca). 1503 /434: 5 ej.,

27 x 10 mm, (Mahón, Menorca) (Colección: Azpeitia, 1408). 1503 /435: 5 ej., 37 x20 mm, (Tarifa, Cádiz)

(Colección: Azpeitia, 1408). 1503 /436: 2 ej., 24 x 13 mm, (Valencia) (Colección: Azpeitia, 1408).

Descripción: (Figs. 1 y 2). El tamaño

de los ejemplares ha variado entre 3 x 2

y 28,5 x 15 mm. El animal es ovalado,

carenado y con las valvas gruesas y con-

sistentes. Las valvas terminales y áreas

laterales ofrecen una estriación radial.

En la zona pleural presentan una serie

de costillas longitudinales muy marca-

das que disminuyen en longitud hacia la

zona jugal. El tegmento de las placas

está finamente granulado debido a la

presencia de pequeños tubérculos,

donde se sitúan las estetas. Éstas se dis-

ponen en líneas longitudinales en el

área central de las placas intermedias,

aunque algo distorsionadas en la zona

pleural debido a la presencia de las cos-

tillas. En el área lateral y valvas termina-

les, las líneas se disponen radialmente.

Las megaloestetas, se puede considerar

que se ordenan en quincunce entre las

microestetas, aunque se encuentran

insertas en las mismas líneas. El diáme-

tro medio de luz de las megaloestetas y

microestetas es de 9,92 mm (o: 1,2) y

7,08 mm (0: 1,2) respectivamente. En

una misma fila se encuentran con una

separación media de 10 mm (0: 0,84) y

entre filas de 11, 8 mm (0: 1,81).

Los aleros del articulamento ofrecen

un aspecto sólido, aunque son algo

esponjosos. Los robustos dientes se

encuentran pectinados en el margen.

Sobre el ápice se distingue el reborde del

tegmento. El seno jugal es pectinado.

Las láminas suturales son lisas, su

aspecto varía desde triangular en las

valvas II a IV, rectangular en las valvas

V a VII y trapezoidal, aunque con los

bordes redondeados, en la valva VIII. La

fórmula de las ranuras de las líneas de

inserción oscila entre 8-10/1-2/10-11. La

situación más frecuente es la presencia

de una sola hendidura en cada lado de

las placas intermedias, aunque se han

encontrado ejemplares que presentan

dos, pero sólo en un lado.

El perinoto es ancho y está consti-

tuido por escamas que ofrecen un

aspecto de piel de serpiente. Las escamas

presentan a su vez finas y leves estriacio-

nes en el borde anterior. El tamaño de las

CARMONA ZALVIDE £7 AL.: Nuevos datos del género Chíton en la Península Ibérica

Figura 2. Chiton (R.) olivaceus. A: rádula; B: dientes raquídeo y primer lateral; C: placa uncinada

del diente mayor lateral; D: diente espatulado; E: perinoto; F: disposición de las escamas dorsales;

G: disposición de las escamas ventrales; H: escamas dorsales; 1: espícula marginal.

Figure 2. Chiton (R.) olivaceus. A: radula; B: rachidian and first lateral teeth; C: uncinal plate of the

major lateral tooth; D: spatulate tooth; E: perinotum; F: disposition of dorsal scales; G: ventral scales;

H: dorsal scales; I: marginal spicules.

Iberus, 18 (2), 2000

Tabla I. Caracteres taxonómicos de las especies de Chiton de la Península Ibérica

Table 1. Taxonomical features of Chiton species in the Iberian Peninsula

Ch. olivaceus

Tamaño máximo (mm) DAS

Color Variable

Diámetro medio Megaloesteta (0) 9,92 pm (1,2)

Diámetro medio microestetas (0] 7,08 pm (1,2)

Fórmula de líneas de inserción

Longitud máxima de escamas dorsales 475 pm

Longitud máxima de escamas ventrales 120 pm

Longitud máxima de espículas marginales 125 pm

NY? de Costillas en el área pleural 4-11

escamas es variable según la región del

perinoto. Próximas a las placas se locali-

zan las de tamaño medio, aumentan

hacia la zona media del perinoto, y pró-

ximas a la zona marginal se encuentran

las más pequeñas. En un mismo ejem-

plar (de 14 mm), varía desde la zona

dorsal, media y próxima a la marginal

entre 105, 433 y 210 ym respectivamente.

El tamaño máximo de escama encon-

trado entre los ejemplares estudiados, ha

sido de 475 um de ancho y 230 um de

altura. Las escamas ventrales son más

pequeñas que las dorsales, generalmente

presentan forma rectangular, aunque

existen variaciones, de modo que

pueden aparecer algo curvadas, con un

extremo más estrecho, con bordes muy

redondeados, etc. El tamaño oscila entre

63 x 20 y 120 x 25 um en los distintos

individuos observados. En un mismo

8-10/1-2/10-11

Ch. corallinus Ch. phaseolinus

16 x 10,2 7,3x4,1

Variable Rosados, Verdosos

8, 17 pm (0,51) 7,7 pm (0,89)

8-10/1-2/10-12 8-10/1/9-12

280 pm 210 pm

80 pm 57,3 pm

112,5 pm 70 ym

3-8 23

ejemplar el rango de variación es menor,

aproximadamente entre 10 y 15 ym. En

la zona marginal presenta espículas

cónicas curvadas. El tamaño oscila en los

distintos individuos entre 65 x 25 ym y

125 x 30 ym. En un mismo ejemplar la

máxima diferencia de tamaño que se ha

encontrado ha sido de 20 um.

El diente central de la rádula es alar-

gado y bastante estrecho con el borde

flexible muy marcado, de manera que

ofrece forma de “bastón”. El primer

lateral, algo más ancho y largo que el

central, presenta una prominencia

redondeada en la parte terminal del

diente. El diente mayor marginal, exhibe

una protuberancia con forma de aleta en

la base de la placa uncinada. Ésta se

caracteriza, por no presentar cúspide.

En la Tabla I se especifican las carac-

terísticas de la especie.

Chiton (Rhyssoplax) corallinus (Risso, 1826)

Lepidopleurus corallinus Risso, 1826, Hist. Nat. Eur. Mérid., 4: 268. [Localidad tipo: Nizza]

Chiton rubicundus Costa 1829, Cat. Sist. Test. Sicil.: i, 1ii, pl 1 f. 3 [Localidad tipo: Sicilia]

Chiton pulchellus Philippi 1844 no Gray 1828, Enum. Moll. Sicil. 2: 83, pl 19 f. 14 [Localidad tipo:

Nápoles]

? Chiton scytodesma Scacchi 1836, Cat. Conch. Icon. R. Neapol.: 9 [Localidad tipo: Nápoles]

? Chiton freelandi Forbes 1844, Rep. Br. Ass. Advmt Sci.: 188 [Localidad tipo: Mar Egeo]

Chiton philippi Issel 1870, Bull. Malac. Ital. 3: 5 [Localidad tipo: Génova]

Chiton rubellus Carpenter MS in Pilsbry, 1893, no Nardo, 1847, Man. Conch. 14: 182.

Material estudiado: Se han estudiado 90 ejemplares. Isla de Tarifa, Cádiz: 1 ej., 6 x 3 mm, I!I/91; 2

ej., 13,5 x 8 mm, IV /92 (Intermareal). Isla de las Palomas, Cádiz: 1 ej., 6 x 3, 5 mm, VIII/91; 3 ej., 16

x 10,2 mm, IX/92 (3-12 m), 2 ej., 13,2 x 8,7 mm, X1/93 (9 m). La Ballenera, Cádiz: 3 ej., 13 x 9 mm,

CARMONA ZALVIDE ET AL.: Nuevos datos del género Chiton en la Península Ibérica

CA %

Pes

¿LE

E e

LAA

4 a,

Figura 3. Chiton (R.) corallinus. A: valva l; B: valva IV; C: valva VII; D: ornamentación del teg-

mento de la valva I; E: ornamentación del tegmento del área central; F: disposición de las megalo-

estetas y Microestetas.

Figure 3. Chiton (R.) corallinus. A: valve [; B: valve IV; C: valve VIII; D: tegmentum of valve I; E:

tegmentum of central area; F: disposition of megalaesthetes and micraesthetes.

Iberus, 18 (2), 2000

IV /92 (12 m). Punta de San García, Cádiz: 1 ej., 8 x 45 mm, VII/91; 1 ej., 7,5 x 5 mm, IX/92; 3 ej.,

10,5 x 5 mm, IX/92; 1 ej., 5x 3 mm, II1/93; 1 ej., 10 x 5, 5 mm, VI/93; 2 ej., 16 x 10 mm, VII /93; 1 ej.,

15,2 x 9,2 mm, VII /93; 4 ej., 15 x 9 mm, IX/93 (12 m); 5 ej., 16 x 9 mm, XII/93 (8 m); 1 ej., 12,5 x 7

mm, 1/94 (5 m); 4 ej., 12,3 x 6,2 mm, I1/94 (5 m). Ensenada de Cucareo, Cádiz: 2 ej., 10,5 x 7,5 mm,

1/93 (12 m); 1 ej., 12,5 x 7 mm, XI/93 (5 m); 2 ej., 16 x 13 mm, XII/93 (6 m); 2 ej., 14x 8 mm, 1/94 (5

m); 1 ej., 12x 6,5 mm, 11/94 (5 m). Puerto de Algeciras, Cádiz: 4 ej., 7 x 4 mm, VIII/91; 1 ej., 6x 3,5

mum, IX/91 (10 m). Punta de Paredones, Cádiz: 3 ej., 8,5 x 5 mm, VII/91; 1 ej. 3x2 mm, VIII/91 (16

m). Punta del Gallo del Mirador, Cádiz: 3 ej., 6 x 4 mm, VIII/91 (3-6 m). Campamento, Cádiz: 6 ej,

10 x 5,5 mm, 111/91 (Intermareal). Crinavis, Cádiz: 2 ej., 8,5 x 5,5 mm, VII/90 (33 m); 1 ej., 7x 3,5

mm, V/93 (9 m); 4 ej., 7x3 mm, VII/93 (9 m); 1 ej., 7x 5 mm, IX/93 (10 m); 1 ej., 6,5 x 3,8 mm, 11/94

(8 m). San Felipe, Cádiz: 2 ej., 5x 3 mm, VI/91; 4 ej., 7,5 x 4 mm, VIII /91; 1 ej., 14,5 x 9 mm, V/93;

5 ej. 6x 3,5 mm, VII/93; 1 ej., 11 x 5,8 mm, X/93 (17 m). Punta Europa, Gibraltar: 1 ej., 6 x 3 mm,

VII/93 (14 m). MCNM: 1503/72: 1 ej., 14 x 8 mm, (Canarias) (Colección: Hidalgo). 1503/234: 1 ej.,

10,5 x 6 mm, (Aguilas, Murcia) (Colección: Azpeitia, 3227). 1503/401: 1 ej., 16 x 8,5 mm, (España)

(hidalgo). 1503/402: 1 ej., 8,5 x 4,5 mm, (Mahón, Menorca) (Colección: Hidalgo). 1503 /403: 2 ej., 10

x 6 mm, (Aguilas, Murcia) (Colección: Hidalgo). Sin numerar: 5 ej., 11 x 7 mm, (Cabo Menorca)

(Colección: Hidalgo).

Descripción: (Figs. 3 y 4). El tamaño

de los ejemplares recolectados ha

variado entre 3,3 x 2,1 mm y 16 x 10,2

mm. El aspecto es ovalado, aunque algo

más estrecho en la parte caudal, con

conchas consistentes. Se encuentran

ornamentados con costillas longitudina-

les en el área central. En el tegmento se

detecta una granulación fina ocasionada

por las estetas, que se disponen en

líneas. No se puede distinguir entre

megalo y microestetas, ya que todas

ofrecen un diámetro similar con una

media de luz de 8,17 um (o: 0,51). Las

filas que tienden a formar varían, de

manera que en la placa oral, áreas late-

rales y zona postmucral se disponen

radialmente, mientras que en el área

central y zona anteromucral son de

forma longitudinal, aunque pueden

estar algo distorsionadas por la presen-

cia de las costillas. La distancia media

entre dos estetas en una misma fila y la

paralela es de 12,6 um (o: 5,91) y 10,05

ym (0: 5,36) respectivamente.

El borde de las láminas de inserción

del articulamento se encuentra pectinado

en todas las valvas, al igual que los

dientes de las placas terminales. Las apó-

fisis son lisas y están separadas por el

seno jugal. En éste se aprecia la presencia

de 4 a 6 láminas ligeramente dentadas en

el margen. La forma de las apófisis varía

de rectangular, en las valvas Il y IL, a tra-

pezoidal, de la IV a VII, aunque en todas

los márgenes aparecen redondeados. La

fórmula de hendiduras de las láminas de

inserción es 8-10/1-2/10-12. Las valvas

intermedias exhiben, generalmente, una

sola hendidura; en el caso de que pre-

sente dos, es sólo en uno de los lados.

El perinoto está constituido por

escamas romboides imbricadas, que

ofrecen un aspecto de piel de serpiente.

Las escamas se encuentran ornamenta-

das por unas finas y leves estriaciones.

El tamaño varía en un mismo ejemplar,

encontrándose las mayores en la zona

media del perinoto. Las dimensiones

varían desde 60 x 25 um a 250 x 100 ym

de anchura y altura en el mismo ejem-

plar. El tamaño máximo encontrado ha

sido de 280 um de ancho por 130 um de

alto. Las escamas ventrales son rectan-

gular, aunque pueden tener ligeras

variaciones. El tamaño ha oscilado en

los diferentes ejemplares entre 60 x 13

ym y 80 x 15 ym, con variación máxima

de 5 ym en la longitud de las escamas

en un mismo individuo. Las espículas

marginales, de forma cilíndrica con el

extremo terminal apuntado, tienen

estrías longitudinales. El tamaño varía,

en distintos ejemplares, entre 62,5 x 21

ym y 112,5 x 22,5 ym.

El diente central de la rádula es alar-

gado y bastante estrecho, con una longi-

tud máxima de 598 um. El primer

lateral es ligeramente más ancho, sobre-

pasa o iguala al central, a pesar de ser

más largo. La placa uncinada del diente

mayor lateral no presenta cúspides.

En la Tabla I se especifican las carac-

terística de la especie.

CARMONA ZALVIDE £7 AL.: Nuevos datos del género Chiton en la Península Ibérica

Figura 4. Chiton (R.) corallinus. A: rádula, B: dientes raquídeo y primer lateral; C, D: placa unci-

nada del diente mayor lateral y diente plumoso; E: cintura; E: disposición de las escamas dorsales.

Figure 4. Chiton (R.) corallinus. A: radule; B: rachidian and first lateral teeth; C, D: uncinal plate of

major lateral tooth and spatulate tooth; E: perinotum, E: disposition of dorsal scales.

Iberus, 18 (2), 2000

Chiton (Rhyssoplax) phaseolinus Di Monterosato, 1879

Chiton (Rhyssoplax) phaseolinus Di Monterosato, 1879, G. Sci. Nat. Econ. Palermo, 14: 8. [Localidad

tipo: Nápoles, Palermo].

Material estudiado: El total de ejemplares estudiados ha sido de 59. Isla de Tarifa, Cádiz: 2 ej., 5

x 3 mm, VII/90; 1 ej., 5x 2, 5 mm, II/91; 1 ej., 5x 3 mm, VII/91 (Intermareal). Punta Carnero,

Cádiz: 1 ej. 7x 4 mm, V/91; 2 ej., 6x3 mm, VII /91 (Intermareal). La Ballenera, Cádiz: 1 ej., 3x 2

mm, VIII/91 (12 m). Punta de San García, Cádiz: 3 ej., 6,5 x 3 mm, VII/91; 1 ej., 5x3 mm, 1X/92;

3 ej. 7 x 3 mm, VI/93; 1 ej., 8 x 3,5 mm, VII/93 (10 m); 6 ej., 6, 5x 4 mm, VIII/93; 1 ej., 5, 5 x 3,2

mm, X/93 (12 m); 1 ej. 6 x 3 mm, II/94 (5 m). Ensenada de Cucareo, Cádiz: 2 ej., 3 x 1,5 mm,

XI/91 (5 m); 2 ej., 5 x 2,5 mm, 1/93; 1 ej., 7 x 4 mm, IX/93 (12 m); 2 ej., 7,5 x 4 mm, X/93 (5 m); 2

ej, 9 x 3,2 mm, X1/93 (5 m); 1 ej., 9 x 4 mm, XII/93 (5 m). Puerto de Algeciras, Cádiz: 19 ej., 7x 3

mm, VI[/93; 2 ej., 3,2 x 2 mm, X/93 (10 m). Punta de Paredones, Cádiz: 1 ej., 5x 3 mm, VIII/91

(16 m). MCNM: 1503 /426: 2 ej., 12 x 6,5 mm, (Tarifa, Cádiz) (H. L. Strack). Sin numeración: 1 ej.,

10, 2x 6,5 mm, (Tánger) (Colección: Azpeitia, 5362).

Descripción: Los ejemplares son de

tamaño medio, varían entre 3,1 x 2,2 m y

7,3 x 4,1 mm. El animal es ovalado, no

carenado, redondeado, con conchas no

muy consistentes. Presenta un aspecto

granulado fino, con 2-3 costillas no muy

marcadas en las áreas pleurales. El teg-

mento ofrece un aspecto granuloso deri-

vado de las estetas. El diámetro medio de

éstas es de 7,7 ym (0: 0,89). Las estetas se

sitúan de manera que forman líneas, que

varían de disposición, en la valva cefá-

lica, áreas laterales de las intermedias y

zona postmucral, donde constituyen

líneas radiales, mientras que el área

central y anteromucral tienden a consti-

tuir líneas longitudinales, aunque se

encuentran algo perturbadas en la zona

pleural, donde aparecen las costillas. La

distancia media de las estetas en una

misma fila es de 5,6 um (0: 1,81) y de 7,9

ym (o: 2,6) entre las filas paralelas.

Los aleros del articulamento son

esponjosos. Los dientes de la valvas cau-

dales y el borde de las láminas de inser-

ción se encuentran dentados. En el seno

jugal también se aprecian láminas algo

pectinadas, aunque no tan finamente

como en las láminas laterales de inser-

ción. Por el contrario, las apófisis son

lisas, con una forma triangular en las

valvas II, III y IV, que varían a trapezoi-

dal en las placas siguientes hasta la

caudal. La fórmula de ranuras de las

líneas de inserción es 8-10/1/9-12.

El perinoto está constituido por

espículas romboidales imbricadas.

Pueden presentar leves estriaciones

longitudinales, aunque no se encuen-

tran en todas las escamas. En la zona

basal presentan una perforación, lugar

por donde probablemente se insertan

en el perinoto. En un mismo ejemplar

aparecen de diferentes tamaños y la

variación puede ser desde 63 um de

ancho y 20 um de alto, las más

pequeñas, a 200 x 40 ym las mayores.

El tamaño máximo observado ha sido

de 210 um de anchura por 53 um de

altura. Las escamas ventrales son

blancas y rectangulares, se disponen en

filas longitudinales. El tamaño es

menor que el de las dorsales, y éste

varió de 40 a 57,5 um de longitud y de

9 a 14,4 um de ancho en los distintos

ejemplares estudiados. En la zona mar-

ginal aparecen espículas de forma có-

nica, aplastadas, ligeramente cóncavas

y con el borde terminal redondeado y

más estrecho que el basal. Estas espí-

culas se encuentran adornadas con

estrías por la cara dorsal, que surgen en

la mitad de ella, dispuestas paralela-

mente e inclinadas con respecto al eje

longitudinal. El tamaño varía en los

distintos ejemplares entre 52,5 x 17,5

um y 70 x 20 um de longitud y base.

El diente raquídeo de la rádula es

muy estrecho y alargado, con una

pequeña protuberancia redondeada en

la zona terminal. El primer diente lateral

es más ancho y sobrepasa al central. El

diente mayor lateral presenta una apófi-

sis en la zona basal de la placa con

CARMONA ZALVIDE E7 AL.: Nuevos datos del género Chiton en la Península Ibérica

Figura 5. Chiton (R.) phaseolinus. A: valva 1; B: valva II; C: valva IV; D: valva VIII; E: ornamenta-

ción del área central; F: ornamentación del área lateral; G, H: disposición de las megaloestetas y

microestetas; 1: articulamento.

Figure 5. Chiton (R.) phaseolinus. A: valve I; B: valve II; C: valve IV: D: valve VIII; E: ornamenta-

tion of central area. Figure 6: ornamentation of lateral area. Figures 7 and 8: disposition of megalaest-

hetes and micraesthetes. Figure 9: articulamentum.

Iberus, 18 (2), 2000

aspecto de cilindro hueco, con un

reborde, en forma de lengúueta, en la

parte superior, que está dirigido hacia

atrás.

DISCUSIÓN

DaLL (1879) incluyó en el género

Chiton la sección Radsia Gray (1847).

PiLsBRY (1892) a su vez consideró en el

género Chiton las secciones Chiton s. s.

(especie tipo: Ch. tuberculatus Linneo,

1758), Radsia (especie tipo: Ch. barnesil

Gray, 1847) y Sclerochiton Carpenter,

1873 (sin mencionar especie tipo dicho

autor). KAAS Y VAN BELLE (1980) consi-

deran a Sclerochiton sinónimo del género

Squamopleura Nierstrasz (1906), clasifi-

cado en la subfamilia Acanthopleurinae.

THIELE (1893) describe nuevos sub-

géneros y secciones para Chiton: Amau-

rochiton (especie tipo: Ch. magnificus

Deshayes, 1827 = Ch. striatus Barnes,

1824), Chondroplax (especie tipo: Ch. gra-

nosus Frembly, 1827), Diochiton (especie

tipo: Ch. albolineatus Broderip y

Sowerby, 1829), Poeciloplax (especie tipo:

Ch. quoyi Deshayes, 1836 = Ch. glaucus

Gray, 1828), Sypharochiton (especie tipo:

Ch. pellisserpentis Quoy y Gaimard,

1835), Triloplax (especie tipo: Ch. scabri-

culus Quoy y Gaimard, 1832 = Ch. pellis-

serpentis Quoy y Gaimard, 1835),

Georgus (especie tipo: Ch. mauritianus

Quoy y Gaimard, 1835), Rhyssoplax

(especie tipo: Ch. affinis Issel, 1869),

Clathropleura (especie tipo: Ch. siculus

Gray, 1828 = Ch. olivaceus Spengler) y

Anthochiton (especie tipo: Ch. tulipa

Quoy y Gaimard, 1835).

No obstante, actualmente sólo se

consideran válidos a Chiton y Rhyssoplax

ya que tal como señalaron KAaAs Y VAN

BELLE (1980) se consideran simples sinó-

nimos de Chiton a Amaurochiton, Chon-

droplax, Diochiton, Poeciloplax, Sypharo-

chiton, Triloplax, Georgus y Anthochiton, y

de Rhyssoplax a Clathropleura.

NIERSTRASZ (1906), que sigue la clasi-

ficación de PILSBRY (1892), incluye en el

género Chiton los taxones subgenéricos

Chiton s. s., Radsia y Sclerochiton y

propone un nuevo género, Squamo-

En la Tabla 1 se especifican los carac-

teres taxonómicos de las especies de

Chiton (Rhyssoplax) de la Península

Ibérica.

pleura, considerado actualmente por

KAAS Y VAN BELLE (1980) como válido

en vez de Sclerochiton.

Posteriormente, THIELE (1909)

admite únicamente dos géneros: Chiton

y Sclerochiton. En el primero encuadra la

sección Radsia y el subgénero Clathro-

pleura, y obvia al resto de los géneros

que había descrito anteriormente

(THIELE, 1893).

BERGENHAYN (1914) sigue los crite-

rios de PILSBRY (1892), de manera que

acepta los subgéneros. Sin embargo,

IREDALE Y HULL (1926) no los admiten y

consideran que las diferencias que pre-

sentan los distintos grupos justifican su

separación como géneros independien-

tes e incluso consideran los siguientes

nuevos: Delicatoplax, Tegulaplax, Mucro-

quasma, a la vez que mantienen Rhysso-

plax, Clavarizona Hull (1923), Amaurochi-

ton, Sypharochiton, Squamopleura, Liolop-

hura Pilsbry (1892), Amphitomura Pilsbry

(1892), Acantopleura Guilding (1829),

Onithochiton Gray (1847), Lucilina Dal

(1882) y Schizochiton Gray (1847). KAAs Y

VAN BELLE (1980), de los géneros nuevos

propuestos por tales autores, admiten

como válidos a Tegulaplax y Mucro-

quasma, consideran a Delicatoplax sinó-

nimo de Chiton, y aceptan a Squamo-

pleura, Liolophura, Acantopleura y Clavari-

zona como géneros y subgéneros de la

subfamilia Acantopleurinae, y a Onitho-

chiton y Lucilina de Toniciinae. El resto

se encuentran sinonimizados, como

hemos comentado anteriormente.

THIELE (1929) desarrolla una nueva

clasificación, en la que admite subgéne-

ros y secciones, de manera que en la

subfamilia Chitoninae incluye los sub-

géneros: Chiton y Rhyssoplax. En el

primero considera las secciones: Chiton

s. s., Amaurochiton, Diochiton, Chondro-

plax, Radsia y Sypharochiton; y en Rhysso-

plax las secciones: Anthochiton, Delicato-

CARMONA ZALVIDE £7 42.: Nuevos datos del género Chiton en la Península Ibérica

Figura 6. Chiton (R.) phaseolinus. A: rádula; B: placa uncinada del diente mayor lateral y diente

plumoso; C: dientes raquídeo y primer lateral; D: disposición de las escamas ventrales; E: disposi-

ción de las escamas dorsales; F: escamas ventrales, vista dorsal; G: escama dorsal, vista ventral; H:

escama dorsal, vista lateral; I: espícula marginal; J: escamas ventrales.

Figure 6. Chiton (R.) phaseolinus. A: radula; B: uncinal plate of major lateral tooth and spatulate

tooth; C: rachidian and first lateral tooth; D: disposition of ventral scales; E: disposition of dorsal scales;

F: dorsal view of ventral scales; G: ventral view of dorsal scales; H: lateral view of dorsal scales; I: margi-

nal spicules; J: ventral scales.

Iberus, 18 (2), 2000

plax, Tegulaplax, Rhyssoplax s. s. y Mucro-

quasma.

Como se ha visto hasta ahora, puede

decirse que se han desarrollado dos

líneas en la clasificación de todos estos

taxones, en una se admiten los subgéne-

ros de Chiton, y en la otra se consideran

con valor genérico propio. La primera

opción es seguida, entre otros, por

ASHBY (1931), que admite el subgénero

Sypharochiton (considerado sinónimo de

Chiton por Kaas Y VAN BELLE, 1980);

FISCHER-PIETTE Y FRANC (en GRASSÉ,

1960), que citan como subgénero a

Amaurochiton; SMITH Y FERREIRA (1977),

que admiten a Radsía como subgénero

de Chiton; VAN BELLE (1978), que consi-

dera como subgéneros de Chiton a

Chiton s. s., Radsia, Rhyssoplax y Mucros-

quama; Kaas Y VAN BELLE (1980) añaden

al listado de VAN BELLE (1978) el subgé-

nero Tegulaplax, aunque VAN BELLE

(1983) lo considera como sinónimo de

Chiton. Por último, SABELLI et al (1990) y

DELL” ANGELO et al (1990) admiten a

Rhyssoplax como subgénero.

BULLOCK (1988) también sigue esta

línea, pero admite como subgéneros de

Chiton a Amaurochiton, Diochiton, Chon-

droplax y Chiton s. s. Por otro lado,

separa del género Chiton a Radsia, Sypha-

rochiton y Rhyssoplax, considerados con

el mismo rango taxonómico, y en el

último incluye como subgéneros a Deli-

catoplax y Mucrosquama. Por último, a

Typhlolochiton lo sinonimiza con Chaeto-

pleura.

En la otra línea, podemos a su vez,

considerar dos tendencias. En la

primera, los autores incluyen todas las

BIBLIOGRAFÍA

AsHBY, E., 1931. Monograph of the South Afri-

can Polyplacophora (chitons). Annals South

Africa Museum, 30 (1): 1-59.

BALUK, W., 1984. Additional data on Chitons

and cuttlefish from the Korytnica Clays

(Middle Miocene; Holy Cross Mountains,

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4): 281-297.

BARASH, A. Y DANIN, Z., 1977. Polyplacophora

(Mollusca) from the Eastern Mediterranean.

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HAY OS ELQUI EZ OS:

SABELLI, 1974; LAGHI, 1977; BARASH Y

DANIN, 1977; BALUK, 1984; MIFSUD et al.,

1990; SABELLI et al., 1990), y en la

segunda denominan a la especie con el

género que otros han considerado como

subgéneros (TAKI, 1962; FERREIRA, 1983;

ZEILER Y GOWLET, 1985; ScotT et al.,

1990).

En el presente trabajo se admite a

Rhyssoplax como subgénero de Chiton al

igual que lo hacen VAN BELLE (1978;

1983), KaaAS Y VAN BELLE (1980), SABELLI

et al. (1990), pues desde que se realiza-

ron las primeras clasificaciones se

encuadraba en este género, a pesar de

que ciertos autores no utilizan la catego-

ría de subgénero para denominar a las

especies que se encuentran en la Penín-

sula Ibérica (BERGENHAYN, 1931; LELOUP

Y VoLz, 1938; SABELLI, 1974; LAGHI,

1977; BARASH Y DANIN, 1977; BALUK,

1984; MIFSUD et al., 1990; SABELLI et al.,

1990).

VAN BELLE (1983) indica la siguiente

diagnosis para el subgénero Rhyssoplax:

El tamaño varía de pequeño a grande,

forma de oval a oval alargado. Valvas

carenadas, áreas distinguibles; la pleural

siempre provista de costillas más o

menos marcadas, la escultura del área

lateral y de las valvas terminales varía

de microgranular a radialmente

estriada. Fórmula de líneas de inserción:

8-9/1/10-12. Perinoto tapizado de

escamas. Las tres especies estudiadas

presentan estos caracteres, por lo que se

les ha denominado Ch. (R.) olivaceus, Ch.

(R.) corallinus y Ch. (R.) phaseolinus.

BERGENHAYN, J. R. M., 1914. Die Loricaten Von

Prof. Dr. Sixten Bocks expedition Nac Japan

und den Bonin-Inseln 1914. Kungl. Svenska Ve-

tenskapsakademiens Handlingar, 12 (1- 4): 4-57.

BERGENHAYN, J. R. M., 1930. Die Loricaten Von

Prof. Dr. Sixten Bocks Pazifik Expedition

1917-1918, mit spezieller Berhcksichtigung

der Perinotumbildungen und der chalens-

truktur. K. Svenska Vetensk, 9 (3): 1-54.

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BERGENHAYN, J. R. M., 1931- Beitruge zur Ma-

lakozoologie der Kanarischen Inseln. Arkiv

for zoologl. 23 (13): 1-28.

BULLOCK, R. C. 1988., The genus Chiton in the

New World (Polyplacophora: Chitonidae).

Veliger, 31 (3-4): 141-191.

DaLL, W. H., 1882. On the genus Chiton. Proce-

edings of United States National Museum. 279-

DELL'ANGELO, B., S. HONG Y VAN BELLE, R. A.,

1990. The chiton fauna (Mollusca: Polypla-

cophora) of Korea Part I: Suborder. Korean

Journal systematic Zoology, 6 (1): 29-56.

FERREIRA, A. J., 1983. Researches on the coast

of Somalia. The Chiton fauna (Mollusca Poly-

placophora). Monitore Zoologico Italiano, suppl

18 (9): 249-297.

FIiscHER-PIETTE, E. Y FRANC, A., 1960. Classe

des Polyplacophores. In: P. Grassé, (Ed.)

Traité de Zoologie. Paris. 1701-1728.

IREDALE, T. Y HULL, A. F. B., 1926. A mono-

graph of the Australian Loricates, -VIII. Aus-

tralia Zoology, 339-359.

Kaas, P. y VAN BELLE, R. A., 1980. Cataloge of

living chitons (Mollusca: Polyplacophora). Dr. W.

Publisher. Rotterdam. 144 pp.

LAGHI, G. F., 1977. Polyplacophora (Mollusca)

neogenici dell Appennino Settentrionale. Bo-

lletino della Paleontologica Italiana, 16 (1): 87-

115.

LeLOUP, E. Y VoLz, P., 1938. Die Chitonen (Poly-

placophoren) der Adria. Thalassia. 2 (10): 1-

MIFSUD, C., CACHIA, C. Y SAMMUT, P. M., 1990.

Note sui Poliplacofori delle isole Maltesi. La

Conchiglia, 256: 52-61.

NIERSTRASZ, H. F. 1906. Remarks on the Chito-

nidae. Tijdschrift der nederlandsche. Dierkunde

Vereeiging, (2) 10: 141-172.

PisBRY, H. A., 1892-1894. Monograph of the

Polyplacophora. En: Tryon, G. W.: Manual of

Conchology. Academy of Natural Sciences,

Philadelphia. 331 pp.

SABELLI, B. A., 1974. Origine e distribuzione

dei Poliplacophora viventi in Mediterraneo.

- Quaderni civico Staz. Idrobiologi Milano, 5:

71-78.

SABELLI. B., GAANNNUZZI-SAVELLI. R. Y BEDULLI,

D., 1990. Catalogo annotato dei Molluschi ma-

rini del Mediterraneo. Ed. Librería Naturalis-

tica Bolognese, Boloña, vol. 1, 348 pp.

Scorr, P. H., F. G. HOCHBERG Y RoTH, B., 1990.

Cataloge of recent and fossil molluscan ty-

pes in the Santa Barbara Museum of Natu-

ral History. I. Caudofoveata, Polyplacop-

hora, Bivalvia, Scaphopoda and Cephalo-

poda. Veliger 33, Suppl. 1: 1-27.

SMITH, A. G., y Ferreira, A. J. 1977., Chiton

fauna of the Galapagos Islands. Veliger 20

(2): 82-97.

TakKI, IL, 1962. A List of the Polyplacophora

from Japanese Islands and Vicinity. Japan

Journal Malacology, 22 (1): 29-53.

THIELE, J., 1893. Poliplacophora, Lepidoglossa,

Schupenziingler. En: Troschel, F. H.: Das ge-

gebiss der Schnnecken, 2: 325-336.

THIELE, J., 1909-1910. Revision des Systems der Chi-

tonen. Zoologica Stuttg. 22: 1-132.

THIELE, J., 1929. Handbuch der systematischen

Weichtierkunde. Classis Loricata. Smithsonian

Institution Libraries and The National Science

Foundation. Washington. 1992. 1-22.

VAN BELLE, R. A., 1975-1978. Sur la classifica-

tion des Polyplacophora. I-VII. Informations

de la Société belge de Malacologia, 4 (5): 121-131.

4 (6): 135-145. 5 (2): 15-42. 6 (1): 3-28. 6 (2): 35-

44. 6 (3): 65-82.

VAN BELLE, R. A., 1983. The systematic classi-

fication of the chitons (Mollusca: Polypla-

cophora). Informations de la Société belge de

Malacología. 11 (1-3): 1-178.

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Type-Specimens in the South Australian Mu-

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19 (8): 97-115.

O Sociedad Española de Malacología Iberus, 18 (2): 17-30, 2000

El género Lepidochitona Gray, 1921 (Mollusca, Polypla-

cophora) en el litoral Atlántico de la Península Ibérica

The genus Lepidochitona Gray, 1921 (Mollusca, Polyplacophora) in

the Atlantic littoral of Iberian Peninsula

Pilar CARMONA ZALVIDE* y Francisco J. GARCÍA GARCÍA*

Recibido el 14-VI-1999. Aceptado el 2-111-2000

RESUMEN

En el presente trabajo se citan las especies del Género Lepidochitona Gray, 1921 (Poly-

placophora) del litoral atlántico de la Península Ibérica con indicación de la distribución

de cada especie. Se describen además dos especies nuevas de dicho género.

ABSTRACT

The species of Lepidochitona Gray, 1921 (Polyplacophora) from the Atlantic littoral of Ibe-

rian Peninsula are cited. For each species the geographical distribution in this area is

included. Two new species are described.

PALABRAS CLAVES: Mollusca, Polyplacohora, Taxonomía, Anatomía, Lepidochitona cinerea, L. corrugata, L.

canariensis, L. simrothi, L. monterosatoi, L. iberica, L. kaasi sp. nov. y L. severianoi sp. nov., Península Ibérica.

KEY WORDS: Mollusca, Polyplacohora, Taxonomy, Anatomy, Lepidochitona cinerea, L. corrugata, L. cana-

riensis, L. simrothi, L. monterosatoi, L. iberica, L. kaasi sp. nov., L. severianoi sp nov., Iberian Peninsula.

INTRODUCCIÓN

El género Lepidochitona Gray, 1821 ha

sido ampliamente discutido desde el

punto de vista taxonómico. La contro-

versia mantenida ha estado motivada

por la clasificación de las especies en

diferentes géneros y familias. Amplios

estudios de diferentes autores han sino-

nimizado con Lepidochitona a los siguien-

tes taxones: Trachydermon Carpenter,

1864 (BERRY, 1918; WINCKWORTH, 1932;

Kaas Y VAN BELLE 1981; 1985; FERREIRA,

1982), Craspedochilus Sars, 1878 (PILSBRY,

1892; IREDALE, 1914; THIELE, 1929; KAAS

Y VAN BELLE, 1981), Middendoffia Dall,

1882 (ex Carpenter MS) (Kaas Y VAN

BELLE, 1981; FERREIRA, 1982), Cyanoplax

Pilsbry, 1892 (THIELE, 1929; VAN BELLE,

1977; Kaas Y VAN BELLE, 1981; 1985;

FERREIRA, 1982), Adriella Thiele, 1893

(THIELE, 1929), Mopaliopsis Thiele, 1893

(THIELE, 1929; Kaas Y VAN BELLE, 1981;

FERREIRA, 1982), Mopaliella Thiele, 1909

(KAas Y VAN BELLE, 1981; FERREIRA,

1982), Basiliochiton Berry, 1918 (FERREIRA,

1982), Plotochiton Berry, 1926 (VAN BELLE,

1977; FERREIRA, 1982).

En el presente trabajo se enumeran

un total de 8 especies: Lepidochitona

cinerea (Linneo, 1767), L. corrugata

(Reeve, 1848), L. simrothi (Thiele, 1902), L.

* Departamento Fisiología y Biología Animal, Facultad de Biología, Univ. Sevilla. Apdo. 1095, 41080 Sevilla.

(Spain).

Iberus, 18 (2), 2000

canariensis (Thiele, 1909), L. monterosatoi

Kaas y Van Belle, 1981, L. iberica Kaas y

Van Belle, 1981, L. kaasi sp. nov y L. seve-

rianoi sp nov., todas ellas pertenecientes

al dominio litoral del Atlántico Ibérico.

Se describen como especies nuevas

para la ciencia a Lepidochitona kaasi y

Lepidochitona severianol.

MATERIAL Y MÉTODOS

Se han estudiado los ejemplares de

las colecciones del Departamento de

Bioloxía Animal de la Universidade de

Santiago de Compostela, del Departa-

mento de Biología Animal de la Univer-

sidad de Sevilla y del Museo Nacional

de Ciencias Naturales de Madrid

(MNCN de Madrid).

Determinados ejemplares de L.

kaasi y L. severianoi se trataron con

KOH 20% al objeto de separar las

valvas, elementos del perinoto y la rá-

dula. El estudio de las partes duras se

ha realizado mediante el uso de mi-

croscopía electrónica de barrido (Phi-

lips XL-20).

RESULTADOS

Lepidochitona cinerea (Linneo, 1767)

Chiton cinereus Linneo, 1767, Syst. Nat., ed. 12: 1107.

Chiton marginatus Pennant, 1777, Brit. Zool. 4: 71. [Localidad tipo: England].

Chiton cimex Gmelin, 1791, Syst. Nat., ed. 13: 3206. [Localidad tipo: Norway]

Chiton cimicinus Spengler, 1797, Skkrivt. Naturh. Selsk. 4: 79. [Localidad tipo: Norway].

Chiton quinquivalvis Brown, 1823, Brit. Ennc. (Edinb.) 6 th ed., 6 (2): 402. [Localidad tipo: Wales].

Chiton fuscatus Brown, 1827, IM. Conch. Gr. Brit. Irel. P135 £ 17.

Chiton variegatus Philippi, 1836, Enum. Moll. Sicil. 1: 107 [Localidad tipo: Sicilia]

Lepidopleurus carinatus Leach, 1852, Syn. Moll. Gr. Brit.: 228 [Localidad tipo: Ireland]

(Non: Fleming, 1828; Brown, 1823; 1827; Sowerby, 1841 = Chiton asellus Gmelin, 1791).

Material estudiado: Se ha estudiado un total de 1. 087 ejemplares. Praia de Altar, Benquerencia:

3 ej. 6x 5,2 mm IX/83 (intermareal). Praia da Area Longa, Foz: 16 ej., 6, 3 x 4, 1 mm 11/84 (inter-

mareal). Area de San Román, Ría de Viveiro: 4 ej., 10 x 6 mm VIII/83 (intermareal). Vicedo, Ría

do Barqueiro: 22 ej., 10 x 6 mm IV /76 (intermareal). San Isidro, Ría de Cedeira: 6 ej., 8,2 x 6,1 mm

IV/76 (intermareal). Canal da ría, Ría de Ferrol: 2 ej., 2,5 x 1,5 mm V/87 (10-25 m). Enseada de

Laxe, Ría de Ferrol: 1 ej., 9,5 x 4,8 mm V/76; 3 ej., 6 x 4,4 mm VI!/87 (16 m). Enseada de Leuseda,

Ría de Ferrol: 1 ej. 5,7 x 4,7 mm VI/87 (12 m). Faro da Palma, Ría de Ferrol: 2 ej., 4,4 x 45 mm

VIII/91 (14 m). Punta da Redonda, Ría de Ferrol: 1 ej., 3,2 x 2,1 mm IX/87; 20 ej., 6,8 x 4,3 mm

XI /90; 1 ej., 7,9 x 5,7 mm VU /91; 1 ej., 4,7 x 3,5 mm VIH /91; 10 ej., 9,8 x 5,7 mm VII /91 (20 m).

Punta Piteira, Ría de Ferrol: 1 ej., 6,5 x 4,7 mm VI/91 (16 m). Enseada da Malata, Ría de Ferrol:

10 ej., 11,5 x 7,5 mm 1/75); 24 ej., 17 x 124 mm IX/76; 11 ej., 13 x 8 mm V/84; 1 ej., 79 x 6 mm

M/85; 1 ej., 7 x 4,3 mm 11/87 (intermareal-5 m). Enseada da Barca, Ría de Ferrol: 2 ej., 46 x 3,5

mm VII/87 intermareal). O Seixo, Ría de Ferrol: 5 ej., 6, 6 x 6 mm X/87 (intermareal). Maniños,

Ría de Ferrol: 4 ej., 8,2 x 5,9 mm X/87 (intermareal). O Montón, Ría de Ferrol: 1 ej., 3,8 x 33 mm

11/87 (5 m). As Pías, Ría de Ferrol: 12 ej., 5,9 x 3,8 mm I11/87; 1 ej., 6,3 x 4,1 mm 111/87 (5 m). A

Faísca, Ría de Ferrol: 5 ej., 4,6 x 4,7 mm VIII/87 (intermareal). O Couto, Ría de Ferrol: 105 ej., 10 x

5,1 mm VII1/87; 1 ej., 5,9 x 4,1 mm X/87 (intermareal). O Puntal, Ría de Ferrol: 1 ej., 319 x 4,3 mm

11/87 (intermareal). Illas Mirandas, Ría de Ares: 1 ej., 5x 3 mm IV/87 (13-16 m). Enseada de

Ares, Ría de Ares: 1 ej., 8 x 5 mm I1/86; 4 ej., 8 x 5 mm VII /86; 9 ej., 10x 6 mm 1/87; 2 ej., 10x 6,5

mm IV /87 (8 m). Seno de Pontedeume, Ría de Ares: 6 ej., 15 x 9 mm IV/76; 2 ej., 8 x 5 mm II1/87

(5 m). Punta de San Pedro, Ría de Ares: 1 ej., seco 1V/83; 5 ej., 11 x 7 mm 11/87 (intermareal).

Punta dos Curbeiros de Miño, Ría de Ares: 3 ej., 7,3 x 6,2 mm IV /83; 2 ej., 9 x 5, 5 mm 1/87 (inter-

mareal). Sada, Ría de Ares: 8 ej., 7,5 x 4,9 mm I11/76; 13 ej., 6,5 x 5,7 mm 11/78; 11 ej., 5,6 x 6,1

mm IV/83; 1 ej., 4,8 x 3,2 mm IV/87 (intermareal). Carnoedo, Ría de Ares: 18 ej., 17,4 x 8,5 mm

11/76; 1 ej., 10 x 5 mm IV /87 (intermareal). Lorbé, Ría de Ares: 3 ej., 6,5 x 3/9 mm XI/85 (interma-

CARMONA Y GARCÍA: Especies atlánticas de Lepidochitona en la Península Ibérica

real). Santa Cruz, Ría da Coruña: 5 ej., 10,6 x 6,1 mm 11/76. Punta do Cabo, Suevos: 1 ej., 3,1 x 2,8

mm IV/78 (intermareal). Praia das Cunchas, Ría de Corme e Laxe: 9 ej., 7,9 x 5,8 mm VIII/80

(intermareal). Baixo do Placer do Cabezo de Laxe: 1 ej., 6,7 x 5,7 mm VIII /80 (42 m). Punta Cabo

da Area, Ría de Corme e Laxe: 1 ej., 8,3 x 4,9 mm VII[/80 (intermareal). Fraga de Abaixo, Ría de

Camariñas: 14 ej., 8,7 x 5,4 mm II1/76 (intermareal). Punta dos Corvos, Ría de Camariñas: 1 ej.,

3,3 x 3,5 mm XII/83 (intermareal). Estorde, Ría de Corcubión: 4 ej., 9 x 6,8 mm 11/76 (interma-

real). Punta da Ameixenda, Ría de Corcubión: 2 ej., 11 x 7,1 mm I1/76 (intermareal). O Pindo, Ría

de Corcubión: 12 ej., 7,5 x 6,2 mm VII/84 (intermareal). Punta das Pedras, Ría de Muros e Noia: 7

ej, 8 x 5,7 mm 1/84 (intermareal). Punta Aguieira, Ría de Muros e Noia: 5 ej., 12 x 7 mm 111/76

(intermareal). Punta Cabeiro, Ría de Muros e Noia: 63 ej., 11, 4x 7 mm V/76 (intermareal). Punta

Sagrada, Porto do Son, Ría de Muros e Noia: 13 ej., 9,8 x 5,6 mm II1/76 (intermareal). Punta do

Castro, Baroña, Ría de Muros e Noia: 3 ej., 7,7 x 8,9 mm 11/76; 14 ej., 11,4 x 6,8 mm 11/76; 4 ej.,

3,8 x 2,5 mm 111/84. Aguiño, Ría de Arousa: 9 ej., 9,5 x 6,4 mm II1/76; 1 ej., 1,6 x 1,5 mm IV/76; 4

ej. 10, 5x7 mm VI/81; 1 ej., 7 x3,4 mm VI[/82; 3 ej., 4,2 x 3,5 mm 111/92 (intermareal); 29 ej., 8,6

x 5,2 mm. VI/94 (intermareal). As Insuas, Ribeiriña, Ría de Arousa: 2 ej., 13 x 7,6 mm V/76; 28

ej., 11 x7,3 mm VI/76 (intermareal). Vilagarcia, Ría de Arousa: 5 ej., 8,2 x 5,5 mm V/76 (interma-

real). Cambelo da Area, Ría de Arousa: 8 ej., 5,7 x 5,3 mm I1/82 (10-15 m). Cambados, Ría de

Arousa: 18 ej., 11, 7 x 8,6 mm VI/76 (intermareal). Marisma da Revolta, Ría de Arousa: 2 ej., 8,7 x

6,6 mm 11/76 (intermareal). Punta de A Lanzada, O Grove: 2 ej., 8,5 x 3,7 mm VIII/76 (22 m).

Praia de Bascuas, Ría de Pontevedra: 2 ej., 7,8 x 4,7 mm VIIL/76 (intermareal). Portonovo, Ría de

Pontevedra: 13 ej., 10,5 x 7 mm VII1/76 (intermareal). Sanxenxo, Ría de Pontevedra: 3 ej., 8,3 x 5,5

mm VIII/76 (intermareal). Punta de Campelo, Ría de Pontevedra: 17 ej., 15 x 11 mm IV/76

(intermareal). Illa de San Clemente, Ría de Pontevedra: 4 ej., 9 x 5,8 mm IV/76; 2 ej., 3,7 x 2,2 mm

111/78; 1 ej., 3,7 x 2,4 mm. Praia Ancora, Portugal: 4 ej., 6,2 x 3 mm. IX/93 (intermareal). Praia S.

Bartolomeu do Mar, Portugal: 5 ej., 4,5 x 2,5 mm IX/93 (intermareal). sposende, Portugal: 3 ej.,

3,9 x 2,8 mm. IX/93 (intermareal). Vila do Conde, Portugal: 5 ej., 4,2 x 2 mm. IX/93 (intermareal).

Aveiro, Portugal: 3 ej., 5,5 x 3,2 mm IX/93 (intermareal). Portinho de Arrabida, Portugal: 5 ej., 5,5

x 3,9 mm. X/96 (intermareal). Troia, Portugal: 1 ej., 9 x 5,5 mm V/88 (intermareal). Porto Covo,

Portugal: 1 ej., 7 x 4 mm. VIIL/93 (8 m) Arrifana, Portugal: 1 ej., 4x 2,6 mm (VII/94 (5 m). Praia

Mareta, Portugal: 10 ej., 5,5 x 3,6 mm. VII/94 (8 m). Praia do Burgao, Portugal: 6 ej., 12x7, 8 mm.

VI /93 (7 m). Praia do Lagos, Portuga): 1 ej., 3,1 x 2 mm. VI /93 (17 m). Praia da Marinha, Por-

tugal: 1 ej., 5,8 x 3,2 mm. 1V/88 (intermareal); 4 ej., 8,3 x 4,7 mm. VIII/88 (intermareal). Praia do

Hollos, Portugal: 10 ej., 6,2 x 3 mm. VIII/93 (intermareal). Mirador, Río Piedras: 1 ej., 3x 2 mm.

V/88 (intermareal). Aguas del Pino, Río Piedras: 1 ej., 6,3 x 3,8 mm VI1/87 (intermareal); 4 ej., 10

x 5,5 mm. V/92 (intermareal); 6 ej., 12 x 7 mm. III/93 (intermareal). El Portil, Río Piedras: 3 ej., 13

x 8 mm. IV/91 (intermareal). Bahía de Cádiz: 1 ej., 6,5 x 4 mm. XI/90 (intermareal). 1 ej., 7x5

mm. VI/92 (intermareal). Cabo de Trafalgar, Cádiz: 1 ej., 8,2 x 3,3 mm. IX/95 (intermareal).

Caños de Meca, Cádiz: 1 ej., 9 x 6 mm IV/94 (intermareal). Isla del Tajo, Cádiz: 2 ej., 5x 4 mm.

VII/92 (5 - 6 m). Playa del Chorro, Cádiz: 26 ej., 11 x 6 mm. VII/92 (intermareal). Isla de Tarifa,

Cádiz: 1 ej., 3x 2 mm. 11/91 (3 m); 2 ej., 7,5 x 5 mm. 1/93 (intermareal). La Ballenera, Cádiz: 4 ej,

15 x 9 mm. V/91 (intermareal). Campamento, Cádiz: 5 ej., 8 x 4 mm. II1/91 (intermareal); 9 ej., 9

x 6 mm. VIII/91 (intermareal). MNCN: N* 1503/223: 10 ej., 19 x 10,5 mm. (La Coruña) (Colec-

ción: Azpeitia, 3184). N” 1503 /225: 2 ej., 12 x 8 mm. (Tarifa, Cádiz) (Colección: Azpeitia, 3184). N*

1503/236: 4 ej., 12 x 7,5 mm. (Sangenjo, Pontevedra) (Colección: Azpeitia, 3184). N* 1503/287: 2

ej., 11 x 7 mm. (Cádiz) (Colección: Hidalgo). N” 1503/290: 5 ej., 13,5 x 7,5 mm. (Cascaes, Portugal)

(Colección: Hidalgo). N” 1503/291: 3 ej., 9 x 5 mm. (Foz, Lugo) (Colección: Hidalgo). N*

1503/295: 4 ej., 13,5 x 7 mm. (Murgados, Coruña) (Colección: Hidalgo). N* 1503/298: 2 ej., 11 x 6

mm. (Setúbal, Portugal) (Colección: Hidalgo). N” 1503/299: 1 ej., 9 x 5 mm. (Vigo) (Colección:

Hidalgo). N” 1503/301: 1 ej., 10 x 7,5 mm. (Bayona, Pontevedra) (Colección: Hidalgo). N*

1503/408: 1 ej., 11 x 7,5 mm. (Cascaes, Portugal) (Colección: Hidalgo). Sin N*: 3 ej.,. 19 x 11 mm.

(España) (Colección: Hidalgo). Sin N*: 2 ej., 13,5 x 8 mm. (Inglaterra) (Colección: Azpeitia, 3184).

Sin N*: 19 ej., 12 x 8 mm. (Isla la Toja, Pontevedra) (Colección: Azpeitia, 3184).

Referencias bibliográficas: Vigo, gal (FORBES Y HANLEY, 1853; BUCQUOY,

Coruña, sur de Portugal (MAC ANDREW, DAUTZENBERG Y DOLFUS, 1882; NOBRE Y

1849; 1850); Vigo, (PiLsBRY, 1892); Portu- BRAGA, 1942; PranI, 1980); Algeciras

Iberus, 18 (2), 2000

(CAPELLINI, 1859); Baiona (HIDALGO,

1867); Vigo, Portugal (JEFFREYS, 1865;

1880; 1882); Gibraltar y Algeciras (Mon-

TEROSATO, 1888); Vigo (PILsBRY, 1892;

MALUQUER, 1915); Algorta, Baiona,

Coruña, Mugardos, San Sebastián, San-

tander, Santurce, Vigo, Zumaya,

Cascais, Foz, Lisboa, Leca, Setúbal,

Algarve, Cádiz y Algeciras (HIDALGO,

1917); Berlengas, Cascais, Balieira, Ría

de Faro, y Olhao, (NOBRE, 1932; 1938-

1940); costas Atlánticas de la Península

Ibérica (MALATESTA, 1962; SABELLI,

1974); Costas Asturianas (ANADÓN,

1979); Galicia, Minho, Baixo Alentejo y

Algarve (KaAas Y VAN BELLE, 1981); costa

vasca (BORJA, 1983); Algarve (REIS Y

MONTEIRO, 1984); Algeciras (AARTSEN et

al, 1984); Galicia, Portugal y Andalucía

(Kaas Y VAN BELLE, 1985b); Berlenga

(PISANI, 1986); Illas Sisargas (OTERO Y

TRIGO, 1986); Ría de Arousa (OTERO Y

TRIGO, 1987); Illas Cíes, (ROLÁN ET AL,

1985); Ría de Pontevedra, Islas de Ons

(TRIGO Y OTERO, 1987); Ría de Muros

(OTERO Y TRIGO, 1989); Lourizán, Ría de

Pontevedra (PLANAS Y MORA, 1989); Ría

de Vigo (ROLÁN ET AL., 1989); Ría de

Ares y Betanzos (IRONCOSO y URRGO-

RRI, 1991); Río Piedras (ESTACIO ET AL.,

1992); Enseado do Baño (OLABARRÍA ET

AL., 1997).

Lepidochitona corrugata (Reeve, 1847)

Chiton corrugatus Reeve, 1847, Conch. Icon. 4. [Localidad tipo: Mediterráneo].

Chiton cinereus Poli, 1791, Test. Utr. Sicil. 1: 4. [Localidad tipo: Sicilia].

Lophyrus melphinctensis Poli, 1791, Test. Utr. Sicil. 1: 4. [Localidad tipo: Sicilia].

Chiton crenulatus Risso, 1826, Hist. Nat. Eur. Mérid. 4: 267. [Localidad tipo: Nizza].

Chiton caprearum Scacchi, 1836 (fide Monterosato, 1879), Cat. Conch. Regn. Napoli., 9. [Localidad

tipo: Napoli].

Chiton poli Philippi, 1836, Enum. Moll. Sicil., 1: 106. [Localidad tipo: Sicilia].

Chiton decipiens Tiberi, 1877, Boll. Malac. Ital., 3: 14

Nuttallina (Middendorffia) cinerea var. pseudorissoi Carpenter (en Pilsbry, 1893), Man. Conch. 14:

285. [Localidad tipo: Malta].

Chiton (Nuttallina) (sic!) caprearum, Scacchi, 1836 var. major Pallary, 1900, J. Conch. Paris, 48: 366

Non: Chiton cinereus Linneo, 1767 = Lepidochitona (L.) cinerea (Linneo, 1767)

Non: Chiton polii Deshayes, 1833=Chiton (Rhyssoplax) olivaceus Spengler 1797

Material estudiado: El total de ejemplares recolectados ha sido de 69. Bahía de Cádiz: 10 ej., 14 x

9 mm VI/92 (intermareal); 8 ej., 14 x 9 mm, [11/94 (intermareal). Isla de Tarifa, Cádiz: 2 e]., 3x 1,

5mm, V/91 (intermareal); 1 ej., 3x 1, 5 mm VI/91 (intermareal). Punta Carnero, Cádiz: 8 ej., 4 x

2, 5 mm. V/91 (intermareal). La Ballenera, Cádiz: 1 ej., 6 x 3, 7 mm. V/91 (intermareal). Puerto

de Algeciras, Cádiz: 4 ej., 5, 5x 4 mm, VIII /91 (6 m). Campamento, Cádiz: 1 ej., 10x 7 mm.

VIII/91 (intermareal). Crinavis, Cádiz: 2 ej., 8 x 4, 5 mm, VI/93 (10 m); 1 ej., 10,5 x 6, 2 mm,

VIT/93 (10 m). MNCN: N* 1503/94: 1 ej., 18, 2 x 12 mm. (Mahón, Menorca, Baleares) (Colección:

Hidalgo). N* 1503/227: 5 ej., 16 x 11 mm. (Alicante) (Colección: Azpeitia). N* 1503/228: 5 ej., 15. 5

x 9 mm. (Cartagena, Murcia) (Colección: Azpeitia, 413). N” 1503/464: 2 ej., 14 x 10 mm. (B.

Columbreta) (Colección: Hidalgo). N” 1503/465: 4 ej., 14. 5 x 8. 5 mm. (Cartagena, Murcia)

(Colección: Hidalgo). N* 1503/466: 4 ej., 13 x 8. 5 mm. (Valencia) (Colección: Hidalgo). N*

1503/467: 2 ej., 11 x 8 mm. (Barcelona) (Colección: Hidalgo). Sin N*: 4 ej., 13 x 8, 2 mm. (Valencia)

(Colección: Azpeitia, 413). Sin N*: 4 ej., 11 x 7 mm. (Málaga) (Colección: Hidalgo).

Referencias bibliográficas: Portugal,

(CAPELLINI, 1858; BUCOOUY, DAUTZEN-

BERG Y DOLFUS, 1882; LOCARD, 1886);

Lagos, Ría de Faro y Olhao, Cádiz

(PILSBRY, 1892); Cádiz (MALUQUER, 1915);

Océano Atlántico Ibérico (THIELE, 1929);

Lagos, Ría de Faro, Olhao (NOBRE, 1938 -

1940); costas suratlánticas de la Península

Ibérica, (MALATESTA, 1962); Costas meri-

dionales Atlánticas españolas, (SABELLI,

1974); Algarve y Golfo de Cádiz, (KAAs Y

VAN BELLE, 1981; 1985b); Algarve (REIS Y

MONTEIRO, 1984); Algeciras (AARTSEN ET

AL. 1984); Berlenga (PISANI, 1986).

CARMONA Y GARCÍA: Especies atlánticas de Lepidochitona en la Península Ibérica

Figura 1. Lepidochitona kaasi. A: valva I; B: valva IL; C: valva IV; D: valva VIII; E: disposición de

los tubérculos en la valva 1; E: disposición de las estetas en los tubérculos en la valva I; G: disposi-

ción de los tubérculos en la zona pleural; H: disposición de las megaloestetas y microestetas.

Figure 1. Lepidochitona kaasi. A: valve l; B: valve II; C: valve IV; D: valve VIII; E: arrangement of

the tubercles on valve I; EF: arrangement of aesthetes on valve I; G: arrangement of tubercles on pleural

area; H: arrangement of megaloaesthetes and microaesthetes.

Iberus, 18 (2), 2000

Lepidochitona simrothi (Thiele, 1902)

Nutallina (Middendorffia) simrothi Thiele, 1902, Z. Wiss. Zool. 72: 287. [Localidad tipo: Azores].

Material estudiado: Se ha estudiado un total de 6 ejemplares. Praia da Marinha (Portugal): 6 ej., 9,

8x7,7 mm, IV/88 (intermareal).

Lepidochitona canariensis (Thiele, 1909)

Trachydermon canariensis Thiele, 1909, Zoologica Stuttg. 22: 15. [Localidad tipo: Tenerife].

Material estudiado: Se ha estudiado un total de 21 ejemplares. Arrifana, Portugal: 1 ej., 3, 7x2, 2

mm, VII/94 (intermareal). Praia Mareta (Sagres, Portugal): 6 ej., 4, 2 x2, 9 mm, VII/94 (8 m). Isla

Tarifa, Cádiz: 4 ej., 4, 8x3, 2 mm, VI/91 (intermareal). La Ballenera, Cádiz: 2 ej., 6, 5x4 mm, V/91

(intermareal); 1 ej., 7, 2 x 4 mm, IX/91 (12 m). Puerto de Algeciras, Cádiz: 1 ej., 3,8 x 2, 2 mm,

VIII/91 (6 m). Crinavis, Cádiz: 3 ej., 6, 1 x 4, 1 mm, 111/93 (10 m); 1 ej., 4, 5x3 mm, V/93 (10 m); 1

ej, 4x2, 5 mm, VI/93 (10 m); 1 ej., 4x2 mm, IX/93 (10 m).

Lepidochitona monterosatoi Kaas y Van Belle, 1981

Lepidochitona (Mopaliella) sp. Van Belle, 1977, Inf. Soc. belg. Malac. 5 (2): 15-42.

Material estudiado: Se han estudiado un total de 50 ejemplares: Praia da Area Longa, Foz: 4

ej., 4, 2x2, 5 mm, II/84 (intermareal). Portocelo, Viveiro: 2 ej., 3, 5x2, 7 mm, VIII/83 (inter-

mareal). Santa Cruz, Ría da Coruña: 1 ej., 4, 6x3, 4 mm, 1/84 (intermareal). Punta do Cabo,

Suevos: 1 ej., 4x2 mm, IV/78 (intermareal). Punta Chan, Malpica: 2 ej., 4, 5x3 mm, XI/83

(intermareal). Praia das Cunchas, Ría de Corme e Laxe: 1 ej., 4, 2x2, 8 mm, VIII /80 (interma-

real). Punta dos Corvos, Ría de Camariñas: 1 ej., 4x2, 5 mm, I1/83 (intermareal). Illa da Creba,

Ría de Muros e Noia: 4 ej., 4, 4x2, 7 mm, XI/74 (1-4 m). Punta Aguieira, Ría de Muros e Noia:

l ej., 3, 7x2, 1 mm, XI /74 (intermareal). Punta Sagrada, Ría de Muros e Noia: 6 ej., 5x3 mm,

[11/76 (intermareal). Aguiño, Ría de Arousa: 1 ej., 6x4 mm, III/92 (intermareal); 1ej., 2, 8x1, 5

mm, XII/83 (intermareal); 4 ej., 4, 5x2, 3 mm, VI/94 (intermareal). Marisma da Revolta, Ría

de Arousa: 1 ej., 4x2, 5 mm, 111/76 (intermareal). Praia Mexiloeira, Ría de Arousa: 2 ej., 3, 5x2,

5 mm, X/76 (2 m). Illa de San Clemente, Ría de Pontevedra: 1 ej., 4, 2x2, 6 mm, IV/76 (inter-

mareal). Monteagudo, Illas Cíes: 1 ej., 4, 5x3, 4 mm, II1/83 (intermareal). Enseada do Lago,

Illas Cíes: 4 ej., 5x3 mm, 111/83 (intermareal). Faro, Illas Cíes: 1 ej., 7x5 mm V/84 (interma-

real). Canido, Ría de Vigo: 1 ej., 1, 7x1, 1 mm, XI1/74; (intermareal); 3 ej., 7, 5x5, 5 mm, IX/76

(intermareal); 1 ej., 5x3 mm, IV/84 (intermareal). Praia Mareta, Portugal: 1 ej., 3, 8x2 mm,

VII[/94 (5 m). Isla Tarifa, Cádiz: 1 ej., 4, 8x3 mm, VI/91 (intermareal). La Ballenera, Cádiz: 2

ej., 5, 5x4 mm, V/91 (intermareal). Puerto de Algeciras, Cádiz: 3 ej., 7x5 mm, VIII/90 (6 m).

El Rinconcillo, Cádiz: 1 ej., 3, 5x2 mm, VII/90 (3m). Crinavis, Cádiz: 1 ej., 6x3, 5 mm, IV/93

(11 m).

Lepidochitona iberica Kaas y Van Belle, 1981

- Material estudiado: Se han estudiado dos ejemplares. SW da Illa da Arousa, Ría de Arousa: 1 ej.,

2 x 1,5 mm, 111/82 (8-28 m). Museum Leiden-Mollusca (paratipo) N1 5538512 (Ría de Arousa,

VI /1962, 2,5 x 1 mm, 15 m).

Referencias bibliográficas: Ría de Galicia (ROLÁN, 1989; OTERO Y TRIGO,

Arousa (KAas Y VAN BELLE, 1981); 1989).

CARMONA Y GARCÍA: Especies atlánticas de Lepidochitona en la Península Ibérica

Figure 2. Lepidochitona kaasí. A: rádula; B: dientes raquídeo y primer lateral; C: placa uncinada del

diente mayor lateral; D: diente plumoso; E: corpúsculos dorsales; F: espícula marginal, vista

ventral; G: espícula marginal, vista dorsal; H: escama ventral.

Figure 2. Lepidochitona kaasi. A: radula; B: rachidian and first lateral teeth; C: uncinal plate of the

major lateral to0th; D: spatulate tooth; E: dorsal corpuscles; E: ventral view of the marginal spicules; G:

dorsal view of the marginal spicules; H: ventral scales.

Iberus, 18 (2), 2000

)

Lepidochitona kaasi spec. nov.

Material estudiado: Se ha estudiado un total de 25 ejemplares: Isla Tarifa, Cádiz: 22 ej., 3,5x 2 mm,

VI/91 (intermareal); 3 ej., 2,9 x 1,7 mm, VI /95 (intermareal).

Se ha designado como holotipo el ejemplar de 3,0 x 1,5 mm, procedente de la Isla las Palomas de

Tarifa (localidad tipo, 36? 01' 8” N; 05 36' 22” O), que ha sido depositado en el Museo Nacional de

Ciencias Naturales de Madrid con número de registro MNCN 15.03/486 y los restantes constitu-

yen la serie paratípica, que se encuentra en la colección de del Departamento de Fisiología y Bio-

logía Animal de la Universidad de Sevilla.

Derivatio nomíinis: La especie ha sido denominada como Lepidochitona kaasi, dedicada al Doctor

Piet Kaas, recientemente fallecido, y que dedicó la mayor parte de su vida al estudio de los poli-

placóforos de todo el mundo.

Diagnosis: Ejemplares de hasta 3,5 x 2

mm. La coloración es parda, a veces con

manchas blancas. El aspecto es ovalado,

las valvas son redondeadas y no carena-

das con el ápice muy marcado. La orna-

mentación está constituida por gránulos

toscos redondeados. El perinoto es estre-

cho, con bandas alternas claras y oscuras.

Descripción (Figs. 1, 2): La valva cefá-

lica muestra el borde anterior semicircu-

lar y el posterior triangular, con un

ángulo muy obtuso (Fig. 14). La pen-

diente de la valva tiende a ser convexa.

El tegmento ofrece leves surcos radiales

que recorren la valva hasta el margen.

Las valvas intermedias son triangulares,

en forma de V, con el borde anterior

cóncavo (Fig. 1C) a excepción de la valva

II (Fig. 1B) que tiende a ser recto, los late-

rales son ligeramente redondeados, y los

márgenes posteriores concurren de

forma casi recta en el ápice. Las áreas

laterales sobresalen ligeramente de la

central. La valva caudal (Fig. 1D) es de

menor tamaño que la cefálica, el margen

anterior tiende a ser recto y el posterior

semicircular. El mucro, poco marcado, se

encuentra en disposición central con una

pendiente ligeramente convexa. Todas

las valvas se caracterizan por presentar

un reborde muy marcado del tegmento

en la zona posterior de la cara ventral.

El tegmento ofrece una ornamenta-

ción constituida por toscos gránulos dis-

puestos en quincunce, con alguna varia-

ción en las distintas áreas de las valvas.

En la valva cefálica, zona postmucral de

la caudal y áreas laterales de las interme-

dias tienden a formar cadenas semicircu-

lares concéntricas (Figs. 1E, F), mientras

que en las áreas centrales y anteromucral

se disponen en líneas que forman V con-

céntricas (Fig. 1 G). En los gránulos se

dispone una megaloesteta central de 5

um (o: 0,38) de diámetro y en círculo

concéntrico las microestetas de 1,3 um

(o: 0,23) de diámetro (Fig. 1H). Entre los

gránulos también se disponen de forma

azarosa estas microestetas.

El articulamento es blanco aunque

puede variar a pardo en la zona jugal. Los

aleros son muy esponjosos y los dientes

algo ondulados. Las apófisis son lisas, de

forma triangular con los bordes algo

redondeados en las valvas intermedias.

Presenta entre 8 y 10 líneas de inserción

en la valva cefálica, una en las interme-

dias y ninguna en la caudal, ofreciendo 8

ó 9 dientes dirigidos hacia delante.

El perinoto se caracteriza por la

alternancia de bandas blancas y pardas.

Está constituido por corpúsculos no

imbricados, cilindrocónicos curvados,

con una serie de costillas en ambas

caras, que se disponen desde la mitad

del corpúsculo hasta el borde apical. Su

número varía entre 4 y 6 (Fig. 2E). El

tamaño oscila entre 52 y 65 ym de altura

y 20 y 30 um de base. Las escamas ven-

trales ofrecen una forma cónica de 28

mm de altura y 10 mm de diámetro (Fig.

2H). El fleco marginal está formado por

espículas cilíndricas con el borde termi-

nal afilado, de un tamaño de 105 ym de

longitud y 25 um de diámetro. Se carac-

terizan por presentar estrías longitudi-

nales en la cara dorsal (Fig. 2G) de la

espícula y lisa por los laterales y la cara

ventral (Fig. 2H).

Las branquias se clasifican como

holobranquia abanales. Se han contabili-

zado hasta 14 branquias.

CARMONA Y GARCÍA: Especies atlánticas de Lepidochitona en la Península Ibérica

Eigure 3. Lepidochitona severianoi. A: valva B; 2 valva Il; C: valva IV; D: valva V; E: valva VIII; E:

disposición de los tubérculos en la valva I; G: disposición de los tubérculos en la zona jugal; H: dis-

posición de los tubérculos en la zona pleural; I: disposición de las megaestetas y microestetas.

Figure 3. Lepidochitona severianoi. A: valve l; B: valve II; C: valve IV: D: valve V; E: valve VII; E:

arrangement of the tubercles on valve I; G: arrangement of the tubercles on the jugal area; H: arrange-

ment of the tubercles on the pleural area; [: arrangement of megaesthetes and microesthetes.

Iberus, 18 (2), 2000

El diente radular raquídeo es alar-

gado, con un borde terminal flexible. El

primer lateral es más estrecho y de una

longitud similar al central, aunque no lo

sobrepasa por encontrarse insertado a

un nivel más bajo (Figs. 2A, B). El diente

mayor lateral presenta tres puntas de

sobresale ligeramente la central (Fig.

2C). El diente plumoso que sobrepasa al

diente mayor lateral, presenta el borde

pectinado (Fig. 2D).

Biología: Solamente se han recolec-

tado ejemplares en pozas de roquedos

de la zona intermareal de la Isla de

tamaños prácticamente iguales, aunque Tarifa.

Lepidochitona severianoi spec. nov.

Material estudiado: Se ha estudiado un total de 8 ejemplares: Caños de Meca, Cádiz: 1 ej. 9 x 5,2

mm, IX/93 (Intermareal). Islas de las Palomas, Cádiz: 1 ej., 8 x 5 mm, IX/92 (12 m). Punta Carnero,

Cádiz: 1 ej. 4x2, 4 mm, V/91 (Intermareal). La Ballenera, Cádiz: 1 ej., 4x 2,5 mm, V/91 (Interma-

real). 1 ej., 7, 8x 4 mm, IX/91 (12 m). Crinavis, Cádiz: 1 ej., 8x5 mm, V/93 (10 m); 1 ej. 46 x 2,5

mum, VI/93 (10 m); 1 ej. 5,5 x 3,5, X/93 (10 m).

Se ha designado como holotipo al ejemplar de 7,8 x 4 mm recolectado en la bahía de Algeciras (loca-

lidad tipo) que ha sido depositado en el Museo de Ciencias Naturales de Madrid con número de

registro MNCN 15.03/487 y los restantes constituyen la serie paratípica que se encuentra en la

colección de del Departamento de Biología Animal de la Universidad de Sevilla.

Derivatio nominis: La especie ha sido denominada como Lepidochitona severianoi dedicada a D.

Severiano Carmona Cuñales.

Diagnosis: Ejemplares de talla media,

de hasta 9 mm de longitud y 5,2 mm de

anchura. La coloración es rosada con

matices anaranjados muy pálidos y

manchas pardas en la zona central de

las valvas. La forma es ovalada, con las

valvas redondeadas y las áreas laterales

son patentes y elevadas, aunque nor-

malmente se encuentran muy erosiona-

das. La ornamentación la forman gránu-

los redondeados de aspecto tosco, dis-

puestos en quincunce. Puede ofrecer de

3 a 4 costillas longitudinales en las áreas

pleurales. Presenta 8 ó 9 surcos radiales

en la valva cefálica y hasta 3 en las áreas

laterales. La valva caudal es de menor

tamaño. El aspecto del perinoto es gra-

nulado, constituido por corpúsculos no

imbricados y lisos, entre los cuales

sobresalen espículas aisladas o en pena-

chos de dos o tres. El fleco marginal es

poco patente.

Descripción (Figs. 3, 4): La valva cefá-

lica ofrece el borde anterior en forma de

un tercio de círculo (Fig. 3A). El margen

posterior es triangular. La pendiente ori-

ginada es fuertemente convexa. Se

observan de 8 a 9 surcos radiales muy

marcados, dispuestos desde el ápice

hasta el margen anterior. La valva

segunda es triangular (Fig. 3B). El resto

de las valvas intermedias son rectangu-

lar, con el margen anterior ligeramente

convexo, los laterales curvados y el pos-

terior tiende a ser recto pues el ápice

generalmente está erosionado (Figs. 3C,

D). Las áreas laterales están fuertemente

elevadas con respecto al área central y

presentan surcos radiales obsoletos, en

número de 2 a 3. En las áreas pleurales,

pueden presentar hasta 4 costillas longi-

tudinales patentes, aunque no en todas

las valvas. La valva caudal, de menor

tamaño que la cefálica, presenta el borde

anterior algo convexo y el posterior

semicircular (Fig. 3E). El mucro se posi-

ciona de forma anterocentral y es poco

prominente. La pendiente que origina el

mucro es convexa.

La ornamentación de las valvas está

constituida por toscos tubérculos redon-

deados dispuestos en quincunce en la

zona jugal (Fig. 3G) y con tendencia a

formar cadenas convergentes hacia el

margen anterior en la zona pleural (Fig.

3H), perdiendo de esta forma la identi-

dad de tubérculo redondeado. En las

áreas laterales y valvas terminales se

observa una tendencia a constituir

cadenas concéntricas (Fig. 3F).

CARMONA Y GARCÍA: Especies atlánticas de Lepidochitona en la Península Ibérica

Figura 4. Lepidochitona severianoí. A, B: rádula; C: placa uncinada del diente mayor lateral y diente

plumoso; D: dientes raquídeo y primer lateral; E: disposición de los corpúsculos dorsales; F: dispo-

sición de las escamas ventrales; G: espículas dorsales; H: corpúsculos dorsales; 1: espículas margina-

les; J: escama ventral.

Figure 4. Lepidochitona severianoi. A, B: radula; C: uncinal plate of major lateral tooth and spatulate

tooth; D: rachidian and first lateral teeth; E: arrangement of dorsal corpuscles; F: arrangement of

ventral spicules; G: dorsal spicules; H: dorsal corpuscles; I: marginal spicules; J: ventral scales.

Iberus, 18 (2), 2000

El diámetro de los gránulos es de

86,99 um (0: 5,92). En el centro se sitúa

una megaloesteta de 9,3 um (0: 0,5) de

diámetro. Las microestetas se disponen

en círculos concéntricos alrededor de la

megaloesteta, con un diámetro medio

de 3,4 ym (0: 0,83) (Fig. 3D).

El articulamento es de color blanco.

Los aleros son esponjosos y los dientes

fuertes y ligeramente ondulados, dirigi-

dos hacia el centro en la valva caudal.

En las valvas intermedias y cefálica se

observa el reborde del tegmento de la

zona del ápice sobre la articulamento.

La forma de las apófisis varía desde

triangulares en las valvas más anteriores

a trapezoidal en las posteriores. Las

líneas de inserción son patentes en todas

las valvas, la fórmula es 8/1/9.

El perinoto ofrece bandas alternas de

tonos claros y pardo anaranjado. Está

constituido por corpúsculos lisos con

forma de “almohadillas”, con los bordes

proximal y terminal redondeados y los

laterales rectos, aunque pueden estar

algo curvadas en posición convexa

(Figs. 4E, H). El diámetro oscila entre 15

y 20 um y la longitud de 25 a 50 um.

Entre ellas, se encuentran penachos de

espículas dorsales curvadas y lisas, que

pueden alcanzar tamaños de 200 um

(Fig. 4G). Las escamas ventrales son de

DISCUSIÓN

A las dos especies, Lepidochitona kaasi

y lepidochitona severianoi se les ha clasi-

fica dentro del género Lepidochitona por

manifestar las características propias

que definen KaAAs Y VAN BELLE (1981),

distinguiéndose de Dendrochiton y Spon-

gioradsia por no presentar los apéndices

córneos ramificados en el perinoto.

Al comparar a Lepidochitona kaasi con

otras especies de Lepidochitona del Atlán-

tico Oeste, Sur Africa, Noroeste y

Noreste del Pacífico, se comprueba que

ninguna presenta las valvas intermedias

en forma de V, característica marcada de

los ejemplares de esta especie.

Con respecto a las especies del

Atlántico Oriental, por su aspecto

externo se puede confundir en un prin-

aspecto triangular y lisas (Figs. 4E, J). Su

tamaño varía entre 27 y 40 um de altura

y de 12 a 15 um de ancho. El fleco mar-

ginal es poco patente, estando formado

por espículas cilíndricas con el borde

terminal agudo y estriadas dorsalmente

por 5 costillas longitudinales. El rango

de longitud es de 80 a 85 um.

Las branquias se clasifican como

merobranquia abanal. El numero de

branquias en cada lado ha oscilado entre

10 y 14.

El diente radular raquídeo presenta

forma rectangular, con una lengúeta fle-

xible terminal estrecha (Figs. 4A, B). El

primer lateral (Fig. 4D), a pesar de no

sobrepasar al diente central, ofrece una

mayor longitud, y es más estrecho que

el diente raquídeo. El diente mayor

lateral es tricúspide; la punta central

sobresale en los dientes de nueva forma-

ción, mientras que en los más antiguos

se encuentran al mismo nivel. El diente

plumoso exhibe el borde terminal pecti-

nado (Fig. 4C).

Biología: Se ha recolectado en zonas

rocosas con abundancia de algas calcá-

reas. Algunos ejemplares se han encon-

trado cubiertos, casi en su totalidad, por

este tipo de algas. Su distribución bati-

métrica se extiende desde la zona inter-

mareal hasta 10 m de profundidad.

cipio con L. corrugata, L monterosatoi, L.

simrothi y L. iberica, descartándose de L.

cinerea, L. canariensis, L. furtiva, L. strom-

felti y L. severiano! porque éstas presen-

tan las valvas intermedias claramente

rectangulares. Al estudiar los corpúscu-

los calcáreos dorsales del perinoto, se

determina que por la forma son diferen-

tes a los de L. piceola. De igual manera,

se puede distinguir de L. simrothi, ya

que la especie en cuestión no presenta

espículas calcáreas entre los corpúsculos

del perinoto y el tamaño de las espículas

marginales y escamas ventrales es

menor.

Al desmontar las valvas, se observa

que no presenta líneas de inserción en la

valva VIII, lo que la diferencia de L.

CARMONA Y GARCÍA: Especies atlánticas de Lepidochitona en la Península Ibérica

iberica y L. monterosatoi. Comparándola

con L. corrugata, a pesar de ser diferente

la forma de las valvas intermedias, se

distingue por el número de costillas de

los corpúsculos dorsales del perinoto y

el tamaño de éstos, por el nivel de inser-

ción de las branquias y disposición y

tamaños de los dientes raquídeos y

primer lateral de la rádula.

Comparando Lepidochitona severianoi

con las especies de Lepidochitona del

Atlántico Oeste, Sur Africa y Noroeste y

Noreste del Pacífico, se diferencia

porque las que presentan costillas en la

zona pleural también presentan estria-

dos los corpúsculos del perinoto.

Con respecto a las especies del

Atlántico Oriental, partimos de la base

que ninguna de ellas presenta costillas

en la zona pleural y los corpúsculos del

perinoto son estriados en todas. No obs-

tante por su aspecto externo tosco se

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O Sociedad Española de Malacología ——_—_—_—— Iberus, 18 (2): 31-39, 2000

Descripción de Trochoidea (Xerocrassa) roblesíi spec. nov.

(Mollusca, Gastropoda, Hygromiidae) de la Comunidad

Valenciana (España)

Description of Trochoidea (Xerocrassa) roblesi spec. nov. (Mollusca,

Gastropoda, Hygromiidae) from “Comunidad Valenciana” (Spain)

Alberto MARTÍNEZ-ORTÍ*

Recibido el 23-XII-1999. Aceptado el 20-1X-2000

RESUMEN

Se describe una nueva especie de Hygromiidae, Trochoidea (Xerocrassa) roblesi spec.

nov., endemismo valenciano que se ha recolectado en varias localidades de la Sierra Cal-

derona, en la provincia de Valencia. Se caracteriza por su concha pequeña, parduzca con

flamulaciones blanquecinas y abundante pilosidad. Genitalia con la porción masculina y el

conducto de la bolsa copulatriz de gran longitud. Se compara con otras especies de la

misma familia como Trochoidea (Xerocrassa) penchinati (Bourguignat, 1868), T. (X.) ripa-

curcica (Bofill, 1886), T. (X.) salvanae (Fagot, 1886) y Microxeromagna armillata (Lowe,

1852), con las que guarda alguna similitud conquiológica o de la genitalia.

ABSTRACT

A new species from Hygromiidae, Trochoidea (Xerocrassa) roblesi spec. nov., is descri-

bed, a Valencian endemism which has been collected in various localities of the Sierra

Calderona, in Valencia (Spain). lt is characterised by ¡ts small, brown shell with white stre-

aks and abundant hair. Genitalia with the masculine section and copulatrix bursa's duct in

great length. The Trochoidea (X.) roblesi spec. nov. is compared to other species of the

same family such as, Trochoidea (X.) penchinati (Bourguignat, 1868), T. (X.) ripacurcica

(Bofill, 1886), 7. (X.) salvanae (Fagot, 1886) and Microxeromagna armillata (Lowe,

1852), with those which have some conquiological or genitalial similarity.

PALABRAS CLAVE: Mollusca, Hygromiidae, 7rochoidea roblesi, especie nueva, Comunidad Valenciana, España.

KEY WORDS: Molluscs, Hygromiidae, Trochoidea roblesi, new species, “Comunidad Valenciana”, Spain.

INTRODUCCIÓN

Durante los últimos 10 años el autor en el este de la Península Ibérica. Uno

ha realizado un intensivo muestreo por de los resultados obtenidos destacables

toda la Comunidad Valenciana, con el ha sido el hallazgo de una nueva

propósito de estudiar la malacofauna especie, perteneciente al género Trochoi-

terrestre presente en esta región, situada dea Brown, 1827 (MARTÍNEZ-ORTÍ, 1999).

* Museu Valencia d “Historia Natural. Passeig de la Petxina, 15. E-46008 Valencia. Comunidad Valenciana

(España). E-mail: alberto.martinezOuv.es

Iberus, 18 (2), 2000

Este género se caracteriza por presen-

tar una variabilidad conquiológica tan

acusada que no permite realizar una des-

cripción a nivel genérico satisfactoria. En

su genitalia destaca la presencia de un

aparato estimulador constituido por dos

sacos sin dardos, dispuestos a ambos lados

de la vagina, con sus bases contiguas O

alejadas, incluyendo todas las posiciones

intermedias, y dos a cuatro glándulas

mucosas simples o bifurcadas dispuestas

alrededor de la vagina (PUENTE, 1994).

En la Comunidad Valenciana se han

hallado 12 especies atribuidas al género

SISTEMÁTICA

Trochoidea, tres del subgénero Trochoidea

s. str. y nueve de Xerocrassa Montero-

sato, 1892 (MARTÍNEZ-ORTÍ Y ROBLES,

1998) siguiendo los criterios de FORCART

(1976). Así, el nuevo taxon se asigna a

Xerocrassa por la ausencia de apéndice

atrial.

Dado el número elevado de espe-

cies del subgénero Xerocrassa presentes

en la Comunidad Valenciana, el nuevo

taxon se ha comparado con las especies

que presentan características conquioló-

gicas O del aparato reproductor simi-

lares.

Género Trochoidea Brown, 1827

Subgénero Xerocrassa Monterosato, 1892

Trochoidea (Xerocrassa) roblesi spec. nov.

Localidad tipo: Náquera: La Fonteta, (263V), 305YJ2292, 260 m, 25 ejemplares y 19 conchas.

Material tipo: Holotipo depositado en el Museu Valencia d Historia Natural (MVHN) de Valen-

cia con N* 401-A; 36 paratipos depositados en el MVHN (20 ejemplares y 16 conchas), N* 401-B; 2

paratipos (ejemplares) en el Museo Nacional de Ciencias Naturales de Madrid con N* 15.05 /32055;

3 paratipos (2 ejemplares y una concha) en el Nationaal Natuurhistorisch Museum de Leiden

(Holanda) con N* 59157; 2 paratipos (conchas) en el Museo de Geología de la Universitat de Valen-

cia con N* 5751. Todos ellos proceden de la misma localidad que el holotipo.

Otras localidades: Olocau: Barranco de Olocau (258V), 305YJ1197, 260 m, 1 concha; Náquera: Fuente

del Salt (271V), 305YJ2293, 280 m, 1 ejemplar y 2 conchas. Serra: Fuente del Berro (262V), 305YJ1698,

4 conchas.

Etimología: Especie dedicada al Doctor Fernando Robles Cuenca por su larga trayectoria en el

estudio de los moluscos valencianos y por sus consejos y ayuda para llevar a cabo mi proyecto de

Tesis doctoral.

Diagnosis: Concha pequeña, de

coloración parduzca con pequeñas fla-

mulaciones blanquecinas y recubierta

por una pilosidad diminuta y abun-

dante. La genitalia está caracterizada

por una gran longitud del conjunto

pene, epifalo y flagelo, y del conducto

de la bolsa copulatriz.

Descripción:

Concha (Figs. 1-4): dextrógira, sub-

globosa cónica de pequeño tamaño, con

4 a 5 vueltas de espira, de crecimiento

regular, con una ligera quilla en la

última vuelta. Protoconcha formada por

1 1/4 vueltas, con flamulaciones y pre-

sencia de una diminuta pilosidad menos

abundante que en la teloconcha.

Abertura oval, con el peristoma

interrumpido, simple, cortante y sin re-

borde interno. Ombligo pequeño, entre

1,20 y 1,50 mm de diámetro, parcial-

mente oculto por la reflexión del peris-

toma, que posee una anchura máxima

de 1 mm, dejando ver el interior del

ombligo.

La superficie de la teloconcha posee

una ligera y fina costulación. Coloración

parduzca con numerosas flamulaciones

blanquecinas de pequeño tamaño, sobre

todo en la zona más próxima al ápice y

que forman bandas a lo largo de toda la

concha en ambas caras, si bien en la

umbilical se forma un reticulado. Pre-

senta una abundante pilosidad consti-

tuida por pelos cortos, curvados en su

MARTÍNEZ-ORTÍ: Trochoidea roblesi spec. nov. de la Comunidad Valenciana (España)

6 y 1 mm

| 1 mm

Figuras 1-6. Holotipo de Trochoidea (Xerocrassa) roblesi spec. nov. 1a-c: Concha (4,77 mm h; 7,52

mm 0); 2: protoconcha; 3: teloconcha; 4a-c: dibujo de la concha; 5: aparato genital; 6: papila penial.

Figures 1-6. Holotype of Trochoidea (Xerocrassa) roblesi spec. nov. 1a-c: Shell (4.77 mm h; 7.52 mm

1D); 2: protoconch; 3: teloconch; 4a-c: drawing of the shell; 5: genital system; 6: penial papilla.

Iberus, 18 (2), 2000

«sed. riada e A A A .

Figuras 7-10. Rádula del Holotipo de Trochoidea (Xerocrassa) roblesí spec. nov. (M.E.B.) 7: dientes

central y laterales; 8: diente central; 9: dientes marginales; 10: diente marginal.

Figures 7-10. Holotype radula of lrochoidea (Xerocrassa) roblesi spec. nov. (S.E.M). 7: central and

lateral teeth; 8: central tooth; 9: marginal teeth; 10: marginal tooth.

extremo más distal, que se presentan ali-

neados siguiendo las estrías colabrales.

Las dimensiones oscilan entre 3,73 y

4,89 mm de altura y 6,66 y 7,84 mm de

diámetro. Holotipo, 4,77 mm de altura y

7,52 mm de diámetro.

Aparato genital (Figs. 5, 6): El esquema

general de la genitalia, cuyo estudio ana-

tómico se ha realizado sobre siete ejem-

plares, es similar al de las restantes espe-

cies de Trochoidea (Xerocrassa) (Fig. 5).

En el atrio, corto, aparece un ensan-

chamiento de la pared en la zona

enfrentada al pene, que se presenta en

algunas otras especies del género Tro-

choidea y que posiblemente pueda estar

relacionado con la eversión del pene, ya

que únicamente se ha observado en

ejemplares que ya han copulado.

Vagina desde 2,2 hasta 3,05 mm de

longitud, en cuya zona media o en la

más próxima a la inserción con el con-

ducto de la bolsa copulatriz se insertan

los dos sacos del dardo, a ambos lados

relativamente distanciados, curvados,

rudimentarios, de 1,35 mm de longitud

máxima, con un engrosamiento basal y

carentes de dardo. El oviducto libre es

corto, curvado, de 1 a 1,2 mm, formando

un fuerte pliegue.

Las glándulas mucosas, que se pre-

sentan insertadas en la vagina, se

reúnen en cuatro troncos independien-

tes. Se han encontrado seis glándulas,

dos de ellas bifurcadas y con una longi-

tud máxima de 2,9 mm.

El conjunto pene, epifalo y flagelo

tiene una longitud considerable. El pene

proporcionalmente pequeño, tiene una

longitud entre 2,65 y 3,55 mm. En su

interior se presenta una papila penial

corta, cónica, en ocasiones curvada, de

1,25 mm de longitud y con la abertura

subapical (Fig. 6). El epifalo presenta

una longitud comprendida entre 6,25 y

10,5 mm y el flagelo varía entre 5,5 y

10,05 mm. El conducto deferente pre-

senta una longitud entre 7,60 y 10,75

MARTÍNEZ-ORTÍ: Trochoidea roblesi spec. nov. de la Comunidad Valenciana (España)

16 e

Figuras 11-16. Conchas. 11a-c: Trochoidea (Xerocrassa) penchinati (3,09 mm h; 6,19 mm (DJ), Tibi, Les

Casetes, No 89A; 12: teloconcha de 7. (X.) penchinati (M.E.B., No 89A); 13a-c: T (X) ripacurcica (5,7

mm h; 10,6 mm 0), Fredes, Barranco del Salt, N9 205C; 14a-c: T (X.) salvanae (4,54 mm h; 6,88 mm

(1D), Utiel, ermita Virgen del Remedio, No 299V; 15a-c: M. armillata (2,7 mm h; 5,16 mm 9), Ayora,

Meca, No 169V; 16: teloconcha de M. armillata (M.E.B., Ne 169V), colección Martínez-Ortí.

Figures 11-16. Shells. 11a-c: Trochoidea (Xerocrassa) penchinati (3.09 mm h; 6.19 mm 0D), Tibi,

Les Casetes, N* 894; 12: teloconch of Y. (X.) penchinati (S.E.M., N* 894); 13a-c: Y. (X.) ripacurcica

(5.7 mm h; 10.6 mm 0D), Fredes, Bco. del Salt, N* 205C; 14a-c: T. (X.) salvanae (4.54 mm h; 6.88

mm 0D), Utiel, ermita Virgen del Remedio, N* 299V; 15a-c: M. armillata (2.7 mm h; 5.16 mm O),

Ayora, Meca, N* 169V; 16: teloconch ofM. armillata (S.E.M.), Martínez-Ortí collection.

Iberus, 18 (2), 2000

Figuras 17-23. Aparato genital de: 17. Trochoidea (Xerocrassa) penchinati. Tibi: Cabezo de la

Alcocha, N* 202. 18. Papila penial de 7. (X.) penchinati, No 202. 19. T (X.) ripacurcica. Eredes:

Barranco del Salt, N2 205C. 20. Papila penial de 7; (X.) ripacurcica. 21. T. (X.) salvanae. Aras de

Alpuente: ermita Santa Catalina, No 267V. 22. Papila penial de 7! (X.) salvanae, N* 267V. 23. M.

armillata. La Loberuela: entrada, N“ 189V. (e= 1 mm).

Figures 17-23. Genital system of: 17. Trochoidea (Xerocrassa) penchinati. Tibi. Cabezo de la Alcocha,

No 202. 18. Penial papilla of T. (X.) penchinati, No 202. 19. T. (X.) ripacurcica. Fredes: Barranco del

Salt, No 205C. 20. Penial papilla of T. (X.) ripacurcica. 21. T (X.) salvanae. Aras de Alpuente: ermita

Santa Catalina, No 267V. 22. Penial papilla of T. (X.) salvanae, N* 267V. 23. M. armillata. La Lobe-

ruela: entry, No 189V. (b= 1 mm).

mm. Músculo retractor del pene corto,

inferior a 2 mm de longitud.

El conducto de la bolsa copulatriz

presenta una gran longitud, llegando a

alcanzar entre 11,4 y 15,25 mm. La bolsa

copulatriz, sin espermatóforo en su inte-

rior, es de aspecto redondeado y pre-

senta unas dimensiones de 1,60 mm de

ancho por 2,05 mm de alto. Sin embargo,

con espermatóforo presente ésta puede

deformarse hasta alcanzar 2,35 mm de

ancho y 2,50 mm de longitud.

Otros caracteres: El cuerpo es de color

blanquecino sin pigmentación aparente.

MARTÍNEZ-ORTÍ: Trochoidea roblesi spec. nov. de la Comunidad Valenciana (España)

Figuras 24, 25. Localización geográfica de 24* Trochoidea (Xerocrassa) roblesi spec. nov. (puntos), T.

(X.) ripacurcica (circulo vacío), T. (X.) penchinati (aspas), T. (X.) salvanae (triángulos); 25: M. armi-

llata. (círculos llenos: nuevas localidades; círculos vacíos: localidades citadas; círculos semivacíos: ambas).

Figures 24, 25. Geographic location in the “Comunidad Valenciana” of 24: Trochoidea (Xerocrassa) roblesi

spec. nov. (points), T. (X.) ripacurcica (empty circle), Y. (X.) penchinati (crosses), T. (X.) salvanae (trian-

gles); 25: M. armillata (full circles: new localities; empty circles: cited localities; halfempty: both).

Pie de tipo holópoda. Presenta el músculo

retractor del ommatóforo derecho inde-

pendiente del aparato genital.

La mandíbula del holotipo es de tipo

odontognato, con 0,50 mm de anchura,

arqueada y con once costillas.

La rádula del holotipo presenta 1,70

mm de longitud y 0,50 mm de anchura

(Figs. 7-10) con la siguiente fórmula ra-

dular: 13M+12L+C+15L+16M. Como se

puede observar la fórmula radular del ho-

lotipo es asimétrica. La hemirrádula de-

recha está constituida por 25 dientes de los

cuales 12 corresponden a dientes laterales

y 13 a marginales. La fila 15 presenta dien-

tes de morfología anómala. La hemirrádula

izquierda está constituida por 31 dientes

de los cuales 15 corresponden a los late-

rales y 16 a los marginales. La fila 27 pre-

senta dientes de morfología anómala.

Distribución geográfica y hábitat:

Trochoidea (Xerocrassa) roblesi spec. nov.

es un endemismo valenciano que vive

en la Sierra Calderona, comarca de El

Camp de Túria (provincia de Valencia),

donde se ha recogido en varias localida-

des próximas entre sí (Fig. 24).

Iberus, 18 (2), 2000

Vive en ambientes xerófilos, como

pinadas y matorral mediterráneo, y en

una Ocasión se recogió una concha en un

barranco con curso de agua, junto al

municipio de Olocau, probablemente

arrastrada. Los ejemplares vivos se cap-

turaron semienterrados entre la pinocha

(hojarasca) y debajo de pequeñas pie-

dras, a altitudes desde los 260 hasta los

560 m. Se ha encontrado conviviendo

con Trochoidea (Xerocrassa) murcica (Gui-

rao in Rossmássler, 1854).

Ninguna de las especies con las que

se podría confundir conquiológica o

anatómicamente se ha recolectado junto

a T. (X.) roblesi spec. nov., siendo M.

armillata la especie más extendida por la

región, mientras que las otras tres apare-

cen en áreas más concretas y reducidas

(MARTÍNEZ-ORTÍ, 1999) (Figs. 24-25).

Conservación: Trochoidea (Xerocrassa)

roblesi spec. nov. es una especie poco

común en la Comunidad Valenciana que

se extiende por un área muy reducida

de su territorio, la Sierra Calderona. Se

conoce viva en pocas localidades, por lo

que es recomendable realizar un mayor

número de prospecciones para intentar

encontrar nuevas poblaciones. Las prin-

cipales amenazas para esta especie son

el auge urbanístico en la zona, los incen-

dios, vertederos, carreteras y canteras.

Por todo ello, T. (X.) roblesi spec. nov.

debe considerarse como especie vulne-

rable, según la IUCN-1994, y el autor

pretende realizar las gestiones oportu-

nas para incluirla en el catálogo de espe-

cies amenazadas de España.

Discusión: Desde el punto de vista

conquiológico, las especies más parecidas

a T. (X.) roblesi spec. nov. son Trochoidea

(Xerocrassa) penchinati (Bourguignat,

1868) (Fig. 11) y Microxeromagna armillata

(Lowe, 1852) (Fig. 15), ya que ambas pre-

sentan una pilosidad diminuta y abun-

dante por toda la concha (Figs. 12 y 16), y

unas dimensiones que entran en el rango

de variabilidad de T. (X.) roblesi spec. nov.

De ellas difiere principalmente por las

numerosas flamulaciones blanquecinas

que presenta por toda la concha, el

menor aquillamiento de la última vuelta

y el ombligo más reflejado. Las genitalias

de estas dos especies, son claramente dis-

tintas de T. (X.) roblesi spec. nov.; T. (X.)

penchinatí presenta una menor longitud

de los conductos de la parte masculina y

del conducto de la bolsa copulatriz (Fig.

17) y una papila penial con morfología

distinta (Fig. 18), mientras que M. armi-

llata presenta un aparato estimulador

constituido por un saco del dardo, con un

dardo en su interior, y un saco accesorio a

un lado de la vagina, y dos glándulas

mucosas bifurcadas que se insertan en el

mismo lado que los sacos (MANGANELLI

Y GIUsTI, 1988; PUENTE, 1994; PUENTE Y

ALTONAGA, 1995) (Fig. 23).

En cuanto al aparato genital, Trochoi-

dea (Xerocrassa) ripacurcica (Bofill, 1886),

Trochoidea (Xerocrassa) salvanae (Fagot,

1886) y T. (X.) roblesi spec. nov. presentan

un esquema general similar (Figs. 5, 19,

21). Las principales diferencias halladas

entre las tres especies se basan en las dife-

rentes medidas obtenidas del pene,

epifalo, flagelo y la bolsa copulatriz y su

relación entre ellas. El flagelo es largo en

todas ellas. El epifalo es de mayor longi-

tud en T. (X.) roblesi spec. nov. y T. (X.)

ripacurcica que en T. (X.) salvanae, mien-

tras que el conducto de la bolsa copula-

triz es de mayor longitud en T. (X.) roblesi

spec. nov. que en las otras dos especies.

Además, las papilas peniales de las tres

especies presentan distinta morfología

(Figs. 6, 20, 22). Por otra parte, las caracte-

rísticas morfológicas de la concha de

estas especies permiten diferenciarlas cla-

ramente (Figs. 1-4, 13 y 14). Las dimen-

siones máximas de los ejemplares valen-

cianos de T. (X). ripacurcica son 5,97 mm

de altura y 10,08 mm diámetro, mayores

que las de las otras dos especies. Las

medidas obtenidas por Faci (1991) para

esta especie en Aragón son 7,0 mm de

altura y 12,25 mm de diámetro. Las

dimensiones máximas encontradas para

T. (X.) salvanae en la Comunidad Valen-

ciana son de 6,0 mm de altura y 8,60 mm

de diámetro, mientras que para T. (X.)

roblesí spec. nov. son 4,89 mm de altura y

7,834 mm de diámetro. Además, la presen-

cia de pilosidad en la concha y una colo-

ración característica permiten que 7. (X.)

roblesi spec. nov. pueda ser fácilmente

diferenciada de las otras dos especies.

MARTÍNEZ-ORTÍ: Trochoidea roblesi spec. nov. de la Comunidad Valenciana (España)

AGRADECIMIENTOS

Al Dr. E. Gittenberger (Nationaal Na-

tuurhistorisch Museum, Leiden), Chus

del Valle y Eulalia García (Museu de Zo-

ologia de Barcelona) y al Dr. Óscar So-

riano y Miguel Villena (Museo Nacional

de Ciencias Naturales de Madrid) por la

cesión de algunas muestras de las especies

tratadas en este trabajo. También agrade-

BIBLIOGRAFÍA

Faci, G., 1991. Contribución al conocimiento de di-

versos moluscos terrestres y su distribución en la

Comunidad Autónoma Aragonesa. Tesis Doc-

toral (inédita). Universidad de Zaragoza. 787

ar L., 1976. Die Cochlicellinae und He-

licellinae von Palástina und Sinai. Archiv fur

Molluskenkunde, 106 (4-6): 123-189.

MANGANELLL G. Y GIUSTI, F., 1988. A new Hy-

gromiidae from the italian Apennines and no-

tes on the genus Cernuella and related taxa

(Pulmonata: Helicoidea). Bolletino Malacolo-

gico, 23 (11-12): 327-380.

MARTÍNEZ-ORTÍ, A., 1999. Moluscos terrestres

testáceos de la Comunidad Valenciana. Tesis

doctoral (inédita). Univ. Valencia. 743 pp.

MARTÍNEZ-ORTÍ, A. Y ROBLES, F., 1998. El Sub-

género Xerocrassa Monterossato, 1892 (Gas-

tropoda, Pulmonata, Hygromiidae) en la Co-

munidad Valenciana. XII Congreso Nacional

de Malacología. Málaga. C. Salas Ed.: págs. 24-

DO,

cer al Dr. Gittenberger, a la Dra. M* Teresa

Aparicio (Museo Nacional de Ciencias

Naturales de Madrid) y al Dr. Benjamín

Gómez (Universidad del País Vasco) por

sus Observaciones sobre esta nueva espe-

cie. Asimismo, al Dr. Agustín Tato por su

ayuda en la realización de las imágenes en

el microscopio electrónico de barrido.

PUENTE, A. L, 1994. Estudio taxonómico y bioge-

ográfico de la superfamilia Helicoidea Rafines-

que, 1815 (Gastropoda:Pulmonata:Stylomma-

tophora) de la Península Ibérica e Islas Baleares.

Tesis Doctoral (inédita). Universidad del País

Vasco. 1.037 pp.

PUENTE, A. L Y ALTONAGA, K., 1995. Estudio

morfológico y corológico de dos especies

conquiológicamente similares, Helicella cons-

purcata (Draparnaud, 1801) y Microxeromagna

armillata (Lowe, 1852), en la Península Ibérica

e Islas Baleares (Pulmonata: Helicoidea: Hy-

gromiidae). Zoologica baetica, 6: 121-148.

O Sociedad Española de Malacología ——_—_—_—_—_—_——— Iberus, 18 (2): 41-50, 2000

Plaxiphora mercatoris Leloup, 1936 (Polyplacophora:

Mopaliidae) de Isla de Pascua, Chile

Plaxiphora mercatoris Leloup, 1936 (Polyplacophora: Mopaliidae)

from Easter Island, Chile

Cecilia OSORIO RUIZ!, María E. RAMÍREZ CASALI?, Ana M. MORA

TAPIA? y Marco VEGA PETOKVIC'

Recibido el 6-IV-2000. Aceptado el 13-X-2000

RESUMEN

Se analizaron 588 ejemplares de P. mercatoris por tallas y sexos, entre 6 y 57 mm. de longi-

tud total. P. mercatoris presenta sexos separados, sin diferencias sexuales externas. No se

observaron hermafroditas. La proporción entre machos y hembras fue de 1,04: 0,89. Los

machos se diferenciaron sexualmente a los 15,6 mm y las hembras a los 18,2 mm de longi-

tud; las tallas máximas registradas para ambos sexos fueron de 53 y 57 mm respectiva-

mente, P. mercatoris es una especie endémica y exclusiva de isla de Pascua, que se distri-

buye verticalmente en el intermareal medio al inferior de los sistemas rocosos de la isla. P

mercatoris se presenta con densidades entre 1 y 8 individuos/m?. La flora acompañante de

este quitón presentó una dominancia de las algas calcáreas Mesophyllum siamense, Amphi-

roa yendoi y otras Corallinaceae. La fauna acompañante está constituida por las especies

Chthamalus belyiaevi, Cypraea caputdraconis, Echinometra insularis, Nodilittorina pyrami-

dalis pascua, Nerita morio, Polychaeta, Cnidaria (Antozoa), Briozoa y otros.

ABSTRACT

A total 588 specimens of Plaxiphora mercatoris were analyzed, in terms of sizes and sex,

registering sizes between 6 and 57 mm. P. mercatoris presents separate sexes, with no

external sexual differences; no hermaphrodites were observed. The proportion of males

versus females was 1.04: 0.89. Males differentiate sexually at 15.6 mm and females at

18.2 mm, and the maximum sizes registered were 53 mm and 57 mm respectively. P mer-

catoris is distributed only along the perimeter of the island. lts vertical distribution in the

rocky system is from mid to low intertidal with densities between 1 and 8 individuals/m?.

The accompanying flora of this chiton presented some algae, calcareous algae being

dominant: Mesophyllum siamense, Amphiroa yendoi and other Corallinaceae. The accom-

panying fauna is made by following species Chtamalus belyiaevi, Cypraea

caputdraconis, Echinometra insularis, Nodilittorina pyramidalis pascua and Nerita morio.

PALABRAS CLAVES: Plaxiphora mercatoris, Polyplacophora, población, distribución, densidades, fauna y flora

asociada, Isla de Pascua, Océano Pacífico.

KEY WORDS: Plaxiphora mercatoris, Polyplacophora, population, distribution, densities, fauna and flora asso-

ciated Easter Island, Pacific Ocean.

! Departamento de Ciencias Ecológicas, Universidad de Chile, Casilla 653, Santiago. E-mail: cosorioGuchile.cl

2 Laboratorio de Botánica, Museo Nacional de Historia Natural, Casilla 787, Santiago Chile. E-mail: mrami-

rezQmnhn.cl

Iberus, 18 (2), 2000

INTRODUCCIÓN

Los poliplacóforos constituyen un

grupo de invertebrados marinos que ha-

bitan especialmente las zonas rocosas

intermareales de gran parte de las costas

del mundo (VAN BELLE, 1983). Por su

abundancia y hábitos alimentarios, estas

especies son estructuradoras y/o modi-

ficadoras de las comunidades de algas

del sistema intermareal (Prercey, 1987;

GLYNN, 1970; OTAÍZA Y SANTELICES,

1985; PEÑA, ZUÑIGA Y RODRIGUEZ, 1987),

forman parte de la dieta de peces, aves y

estrellas de mar (OTwaY, 1994); y tam-

bién han constituido históricamente un

recurso alimenticio para el hombre.

Existen antecedentes de su consumo

por los habitantes en la Isla de Pascua,

en la Polinesia, en Oriente, en las costas

de Chile Continental y en Cape Banks,

Australia (OSORIO, ATRIA Y MANN, 1979;

Orway, 1994). Poca importancia se ha

dado al conocimiento de la biología y

ecología de este grupo pese a su relativa

importancia en algunos ecosistemas.

El conocimiento científico de la

única especie registrada para la Isla de

Pascua (o Rapa Nui), Plaxiphora mercato-

ris Leloup, 1936 está limitado sólo su

taxonomía. Se ha recolectado en las loca-

lidades de Hanga Piko, Haka Ea, Hotu

Iti, Otuu, Hanga Pukura, Ovahe, y

Anakena (REDHER, 1980).

Al igual que Isla de Pascua, en Sa-

moa, Marquesas, Tahití y Mangareva,

los poliplacóforos se conocen con el

nombre común de “mama” cuyo signi-

ficado en lengua pascuense es “bostezo”

(Atan, com. pers). En Rapa Nui, los gru-

pos de isleños más tradicionales creen

que estos moluscos son un alimento

muy nutritivo para los niños. Por otra

parte, su extracción es parte de la trans-

ferencia de conocimientos entre padres e

hijos. Esta actividad es una práctica co-

mún, donde participa la familia en la ex-

tracción de organismos marinos, para

alimentarse y obtener materia prima

para su artesanía, permitiendo además

la enseñanza del medio marino. Los

“mamas” son preferidos como alimento

a Otros moluscos, por ser grandes y más

fáciles de comer que los caracoles Ne-

rita morio (Sowerby, 1833) y Nerita lire-

llata Redher (1980). Se comen crudos o

cocidos en agua. Los análisis químicos

ratifican que son una buena fuente de

proteína (14,3%) y de minerales (3,6%),

con un bajo contenido graso (2,6%) una

humedad de 77,1% y un 2,4% de extrac-

tivo no nitrogenado (hidratos de car-

bono y otros) (Masson com. pers).

Los escasos antecedentes biológicos

sobre representantes del Género Plaxip-

hora en el mundo, se limitan a los trabajos

de BRANDANI, FAEDO Y PENCHAZADEH

(1974), quienes estudian la estructura por

tallas y sexo, densidad específica y la epi-

biosis de P. aurata del intermareal de Mar

del Plata, Argentina. En la ultima década

sólo conocemos los trabajos de GALVEZ

(1991) sobre la hipomería en P. fernandezi;

de OrwaY (1994) quien describe la ecolo-

gía poblacional de Plaxiphora albida (Blain-

ville, 1825) en Australia y LÓPEZ Y

TABLADO (1997) quiénes estudian aspec-

tos del crecimiento y producción en P.

aurata en Quequen, Argentina.

Las características especiales de ais-

lamiento geográfico, de este único y

endémico representante de la Clase

Polyplacophora en Isla de Pascua, y el

uso que los isleños hacen de ella, hicie-

ron particularmente interesante este

estudio, cuyos objetivos son caracterizar

la especie basados en aspectos biológi-

cos como la morfogravimetría, la estruc-

tura poblacional y la sexualidad, ade-

más de caracterizar su hábitat y distri-

bución en el intermareal.

El conocimiento de la fauna de Isla

de Pascua es muy escaso, debido princi-

palmente a su lejanía y a la dificultad

para trabajar en sus costas, por lo cual la

información de este trabajo es valiosa y

amerita ser conocida por la comunidad

científica, datos que serán útiles en

futuras investigaciones.

MATERIAL Y MÉTODOS

El estudio fue realizado en Isla de

Pascua (2710'S, 109%20'0). Las mues-

tras proceden de diferentes localidades

OSORIO ET AL.: Plaxiphora mercatoris en Isla de Pascua, Chile

109%25'

== HANGA ROA

NES HANGA — VINAPU

OCEANO

PACIFICO

HAKA EA

10920" 109%15'

ANAKENA

PAPA TI

OVAHE

LA PEROUSE 27:05

d (—MOTU ARIKI

PRAURE TUU

AKAHANGA

o 1000 Km

ISLA DE

PASCUA

Figura 1. Mapa de Isla de Pascua y su localización en el océano Pacífico. Las localidades señaladas

con un circulo corresponden a lugares de recolección de muestras de Plaxiphora mercatoris.

Figure 1. Eastern Island in the Pacific Ocean. Sampling stations of Plaxiphora mercatoris are indicated

by an open circle.

del intermareal rocoso (Fig. 1), y fueron

recolectadas durante los períodos de

marea baja, en distintas fechas desde

1991 a 1994 (Tabla I). El muestreo consi-

deró una muestra de un mínimo de 50

ejemplares, número que en algunas

oportunidades no se alcanzó. Estos

ejemplares se mantuvieron extendidos y

congelados sobre una base de madera

por algunas horas, finalmente fueron

fijados en formol al 10% diluido en agua

de mar y trasladados al laboratorio en

Santiago para su análisis.

El análisis poblacional para determi-

nar la estructura de talla y la sexualidad

de la especie se hizo sobre un total de

588 ejemplares, que incluían los anterio-

res. La determinación de sexos se realizó

después de la disección, basado en la

observación de frotis gonádico y/o por

observación del color de las gónadas,

verde para hembras y rojo para machos.

En un total de 278 ejemplares, de los

588 recolectados, se midió a cada ejem-

plar la longitud y ancho máximos consi-

derando el cinturón, con un calibre 0,01

mm precisión. También se registró el

peso total de cada ejemplar con una

balanza Sartorius de 0,02 g de precisión.

La cobertura de algas y la densidad

de las especies, se cuantificó en terreno

mediante dos transectos extendidos

desde un punto máximo superior hasta

el mínimo de marea, el día 20 de Sep-

tiembre de 1993. Se cuantificó la densi-

dad y cobertura usando cuadrantes reti-

culados de 0,25 m? y de 0,50 m”. La des-

cripción del hábitat se hizo basándose

en la identificación de muestras de la

flora y fauna acompañante, recolectadas

en los transectos indicados.

Para describir la morfometría de ma-

chos y hembras se usaron diagramas de

dispersión y un análisis de regresión sim-

ple. En los cálculos se utilizó el programa

Microsof Excel para Windows 95 para

describir las relaciones longitud - ancho y

longitud - peso. La selección del modelo

Iberus, 18 (2), 2000

Tabla IL. Plaxiphora mercatoris de Isla de Pascua. Localidad de recolección, fecha, número de indivi-

duos por sexo y tipo de análisis realizado. Q: químico; M: morfométrico; G: gravimétrico; Med*:

medidos ¿n situ. E

Table I. Plaxiphora mercatoris of Easter Island. Sampling station, date, number of individual, sex and

type of analysis. Q: chemical; M: morphometric; G: gravimetric; Med. *: measured in situ.

Localidad Fecha Número de individuos Tipo análisis

Total Machos Hembras Indeterminados

Prai Ahure 13.03.91 53 28 19 o) MG

Prai Ahure 23.06.91 43 19 24 M, G

Prai Ahure 22.09.91 15 8 Ys M, G

Prai Ahure 10.10.91 23 14 8 1 M, 6

Motu Ariki 19.05.92 55 2 23 M, G

One Tea 21.11.92 35 23 12 M, 6

Tahai 22.09.93 58 24 28 o) M, 6

La Perouse 22.09.93 90 M,Q

Papa lti 19.09.93 A Med*

Vaihu 19.02.94 9 S) 3 1 MG

Vaihu 18.03.94 44 2 ZA MG

Vaihu 27.04.94 54 28 26 M, 6

Vaihu 09.05.94 IZ 25 27 M, 6

Total 588 229 198 14

de regresión se basó en el coeficiente de

determinación 1? para los modelos lineal,

exponencial y potencial (CANAVOS, 1996).

Además se compararon las relaciones

longitud - ancho y longitud - peso entre

machos y hembras, a través de la prueba

de Chow (GUJARATI, 1997).

Para describir la estructura de la

población, se usó el método tradicional

de Petersen del análisis de frecuencias

de tallas (MIRANDA, 1967). Para describir

la distribución por sexos se utilizaron

frecuencias acumuladas expresadas en

porcentajes de machos y hembra y pro-

porción sexual (HERNÁNDEZ, FERNÁN-

DEZ Y BAPTISTA, 1996).

RESULTADOS

A. Parámetros Morfológicos: Las

relaciones entre la longitud total (LT)

con el ancho total (AT) y el peso total

(PT) fueron estudiadas en 278 ejempla-

res (142 machos y 136 hembras).

Dado que no se encontró diferencias

significativas entre sexos (FAT v/s LT:

2,66; gl=2.274; p< 0,05 y F PT v/s LT:

0,52; gl=2.274; p< 0,05), machos y

hembras fueron analizados conjunta-

mente para cada relación. La relación

potencial en ambos casos fue la que

mejor se ajusta a los datos (Figs. 2 y 3):

AT = 2,426 * LT 0,6286, 1”: 0,4861

PT = 0,0013 * LT 2,3916, r”: 0,7582

B. Análisis de la población: Plaxip-

hora mercatoris presenta sexos separados,

sin diferencias sexuales externas. De los

435 ejemplares analizados (Tabla 1), 192

correspondieron a hembras (44,1%) y

229 fueron machos (52,6%) y en 14 ejem-

plares (3,2%) no se pudo determinar el

sexo por presentar gónadas incoloras,

pese a realizarse frotis. La proporción

entre machos y hembras, fue 1,04: 0,89.

La talla mínima a la cual se reconoció el

sexo fue de 15,6 mm en machos y 18,2

mm en hembras. La talla máxima obser-

vada en machos fue de 53 mm, mientras

que las hembras alcanzan tallas de hasta

57 mm. Los individuos indeterminados

presentaron tallas entre 12 mm y 47 mm.

De 11 muestreos, en ocho oportunidades

OSORIO ET AL.: Plaxiphora mercatoris en Isla de Pascua, Chile

N NN

ANCHO (mm)

15 25 35 45 55 65

LONGITUD (mm)

25 3

PESO TOTAL (gr)

LONGITUD (mm)

Figuras 2, 3. Relación longitud - ancho (2) y longitud total y peso total (3) en Plaxiphora mercato-

ris de Isla de Pascua.

Figures 2, 3. Length - width (2)and total length vs total weight (3) relationship of Plaxiphora mercato-

ris from Easter Island.

predominaron los machos, y en tres las

hembras (Tabla ID).

El rango de tallas estudiado osciló

entre 13 y 57 mm a excepción de 3 indi-

viduos del área de Papa Iti (19/09/93)

que registraron tallas inferiores a 13

mm. Aún cuando existe una gran dife-

rencia en el número de individuos por

muestra (9 a 90), el promedio de las

tallas tiene poca variación, (Fig. 4). Los

promedios de tallas variaron entre 30 y

44 mm y solo la muestra 3 (22/09/91)

tiene un promedio bajo (23,35mm). Se

observó una tendencia a la disminución

de tallas en junio 1991 (muestra 2), sep-

tiembre 1991 (muestra 3) y septiembre

de 1993 (muestra 9).

En las localidades de Prai Ahure

(1991) y Vaihu (1994), se realizó un

muestreo continuo de cuatro meses de

duración, donde se observó una distri-

bución normal de tallas de la población.

En Prai Ahure (Fig 5A), el rango de

tallas estuvo entre 11 y 57 mm de longi-

tud total. En cada uno de los meses se

encontró una moda principal y otras

secundarias. La moda principal parece

desplazarse hacia tallas menores al

pasar el tiempo y sólo en el último mes

aumenta con relación al anterior. En

Vaihu (Fig. 5B), las tallas de la población

estuvieron entre 15 y 55 mm de longitud

total. Se encontró una moda principal en

cada uno de los meses casi sin desplaza-

miento de moda, registrándose una talla

media, de 35 mm entre febrero y mayo

de 1994.

C. Descripción del hábitat: Los

ejemplares de P. mercatoris se encontra-

ron adheridos a grandes rocas en áreas

semi expuestas con suaves a pronuncia-

das pendientes y en pozas del interma-

real. Su distribución se observa desde el

intermareal medio al intermareal infe-

rior. Se adhieren por medio de su pie al

sustrato, de superficies casi lisas o algo

rugosas, en grietas o fisuras estrechas,

ajustados a su talla; en paredes vertica-

les, laterales u horizontales al oleaje.

La flora acompañante de este Poli-

placóforo consistió principalmente en

algas calcáreas como (Mesophyllum sia-

mense (Foslie) Adey, Amphiroa yendoi

Borgesen y otras Corallinaceae). Tam-

bién se encontraron otras algas como

Hypnea cenomyce J. Ag., Ulva sp, Entero-

morpha sp., Lobophora variegata (Lamour)

Wom., Hincksia mitchelliae (Harv.) Silva,

Colpomenia sinuosa (Roth) Derb. y Sol.,

Hydroclathrus clathratus (Bory) Howe y

Cladophora socialis Kuetz., representadas

escasamente en cobertura, cuya distri-

bución vertical y abundancia en porcen-

taje se indican en la Tabla II

La fauna acompañante a P. mercato-

ris está constituida preferentemente por

un conjunto de organismos como Chtha-

malus belyiaevi Zevina y Kurshakova,

1973; Cypraea caputdraconis Melvill,

1888; Echinometra insularis Clark, 1972;

Nodilittorina pyramidalis pascua Rosewa-

ter, 1970; Nerita morio (Sowerby,1833),

Polychaeta, Cnidaria (Antozoa), Brio-

zOa y otros.

Iberus, 18 (2), 2000

LONGITUD (mm)

13-03-91

23-06-91

22-09-91

10-10-91

19-05-92

21-11-92

22-09-93

19-09-93

19-02-94

18-03-94

27-04-94

09-05-94

Figura 4. Promedio de tallas y desviación estándar por muestra en Plaxiphora mercatoris, Isla de

Pascua. En el recuadro se indica el número de individuos controlados por muestra.

Figure 4. Mean length and standard deviation per sample in Plaxiphora mercatoris of Easter Island.

boxes: number of individuals in each sample.

D. Distribución vertical y abundan-

cia de P. mercatoris: En Tahai, en un

frente semiexpuesto se ubicaron los

transectos realizados (Fig. 1, Tabla II).

En el muestreo, realizado el 20/09/93,

se observó la presencia de P. mercatoris

desde los 32 cm del máximo de altura

de las mareas, hacia el intermareal infe-

rior. Los ejemplares se encuentran por

lo general aislados, con una densidad de

l ind/m?, ocasionalmente se observa-

ron 3 o más ejemplares juntos. También

tenemos constancia de la presencia de 4

ejemplares, de pequeña talla (<2 cm) en

el interior de marmitas (orificios concá-

VOS) marinas, bajo el erizo Echinometra

insularis y junto a turbelarios. Como

resultado de este análisis es posible

reconocer que Plaxiphora mercatoris tiene

densidades entre 1 ind/m? (20/09/93

Tahai) a 8 ind/m? (22/09/91 Prai

Ahure).

E. Cobertura: Se midió la cobertura

de los organismos dominantes en la

localidad de Tahai (20/09/93), en una

extensión de superficie de playa de 9,25

m2 y desde el punto de marea máxima,

denominado Estación 0 (Tabla II). Entre

las Estaciones O y 1, que equivalen a un

desplazamiento de 3,75 m horizontales

y 0,77 m de altura intermareal, las dos

especies dominantes son Nodilittorina

pyramidalis pascua (12%) y Chthamalus

belyiaevi (22%).

Entre la Estación 1 y la Estación 2,

que equivale a un desplazamiento de 3,0

m horizontales hacia el mar y 0,12 m de

altura de marea, los organismos domi-

nantes son Chthamalus belyiaevi (41% en

el cuadrante 21), las algas Ulva sp (60%

cuadrante 25) e Hypnea cenomyce (45%

cuadrante 24). En las áreas inferiores,

entre las Estaciones 2 a 3, aumenta la

cobertura de otras algas como Lobophora

variegata (40% cuadrante 33), Hincksia

mitchelliae (58% cuadrante 35) y en el

cuadrante 36, las algas H. mitchelliae

(19%) y Amphiroa yendo! (14%).

DISCUSIÓN

La talla máxima registrada en P. mer-

catoris (57 mm de longitud) es seme-

jante a lo indicado por REDHER (1980),

quien da una longitud máxima de 60

mm, medida estimada para un ejemplar

parcialmente doblado (USNM 756279),

y superior a lo señalado por LELOUP

(1936), quien dio una longitud máxima

de 31 mm, basado en cuatro ejemplares.

La estructura de la población de P.

mercatoris en Isla de Pascua mostró tres

grupos modales, semejantes a lo regis-

trado por LÓPEZ Y TABLADO (1997) en

Plaxiphora aurata para las costas de

Argentina. Similar observación realiza

GLYNN (1970) para tres especies de poli-

placóforos tropicales donde existirían

OSORIO ET AL.: Plaxiphora mercatoris en Isla de Pascua, Chile

Tabla II. Distribución vertical de Plaxiphora mercatoris en el intermareal de Isla de Pascua. Cober-

tura de algas y presencia de fauna asociada en la localidad de Tahai (20/09/93, 10:50 a 15:30 h).

Anp: Anphiroa yendoni Borgesen; Cla: Cladophora socialis Kutzing; Cirr: Chthamalus belyiaevi

Zevina y Kurshakova, 1973; Col: Colpomenia sinuosa(Roth)Derb. y Sol.; Cor: Coralinaceae; Ech:

Echinometra insularis Clarck, 1972; Ent: Enteromorpha sp.; Hin: Hincksia mitchelliae (Harv.) Silva;

Hyd: Hydroclathrus clathratus (Bory) Howe; Hyp: Aypnea cenomyce Agardh; Lob: Lobophora varie-

gata (Lamour) Wom.; Mes: Mesophyllum siamense (Foslie) Adey; Nod: Nodilittorina pyramidalis

pascua Rosewater, 1970; Pla: Plaxiphora mercatoris Leloup,1936; Ulv: Ulva sp. X: presencia de P

mercatoris en áreas paralelas. Cuadrantes de 25 cm?.

Table IT. Vertical distribution of Plaxiphora mercatoris on the rocky intertidal of Easter Island. Algal cove-

rage and presence of associated fauna in Tahai, (20/09/93, 10:50 to 15:30 h). Anp: Anphiroa yendoni

Borgesen; Cla: Cladophora socialis Kutzing; Cirr: Chthamalus belyiaevi Zevina and Kurshakova, 1973; Col:

Colpomenia sinuosa (Roth) Derb. and Sol.; Cor: Coralinaceae; Ech: Echinometra insularis Clarck, 1972;

Ent: Enteromorpha sp.; Hin: Hincksia mitchelliae (Harv.) Silva; Hyd: Hydroclathrus clathratus (Bory)

Howe; Hyp: Hypnea cenomyce Agardh; Lob: Lobophora variegata (Lamour) Wom.; Mes: Mesophyllum

siamense (Foslie) Adey; Nod: Nodilittorina pyramidalis pascua Rosewater, 1970; Pla: Plaxiphora merca-

toris Leloup, 1936; Ulv: Ulva sp. X: presence ofP. mercatoris in adjacent areas. Squares has 25 cor.

Cuadrante Estación ESPECIES

N?2 N* Nod Cirr Hyp Cor Ulv Ent Pla Lob Mes Hin Col Anp Ech Hyd Cla

Na o o o INE o O adoro lero Ne loo

1 0 1

2 o)

3 3

4 12

5 0

o 1

YA 0

8 0

9 2

10 0

11 0

12 1 8

13 3

14 10

15 1 22

16 1 30

17 10

18 14 3 1

19 3 16

20 ONES

21 41 AA

22 12 10 43 1

23 12 33 Xx

24 ARAS 50 Xx

25 60 Xx

26 $) 18 23 Xx 1

2 17 8 Xx

28 2 20 19 Xx 32

29 3 3 Xx

30 Xx

31 Xx 24

32 8 10 A 50 4 4

33 5 x 40 Y

34 4 Xx lo) 30 al

35 S) 0 Xx IA

36 1 Xx 19 14

Iberus, 18 (2), 2000

[0))

Frecuenc

OÍ Lo

40 .

A 20)

o a

Ss $

Na N

Intervalo de Talla

(mm)

46-51 /

Frecuencia

22-27

34-39

Intervalo de Talla

(mm)

E -

re o

23-jun

: 13-mar Fecha de muestreo

2/-Apr

18-mar

Fecha de muestreo

19-feb

Figura 5. Distribución de tallas de la población de Plaxiphora mercatoris de Isla de Pascua, durante

cuatro muestreos realizados en 1994 en dos localidades diferentes. A. Prai Ahure; B. Vaihu.

Figure 5. Size distribution of the Plaxiphora mercatoris population found in two different sampling

stations of Easter Island during a period of four month, 1994. A. Prai Ahure; B. Vaihu.

tres grupos de edades, compuestos por

animales jóvenes, o menores a 2 años,

con un rápido reemplazo de los ejem-

plares, esto explicaría que las poblacio-

nes se mantienen en el tiempo con tallas

mas O menos constantes.

La población de chitones de Prai

Ahure muestra un desplazamiento de

las modas con reducción de sus tallas

entre marzo, junio y septiembre. Esto

indicaría reclutamiento a la población y

ausencia de individuos de mayor

tamaño. En octubre nuevamente desa-

parecen los juveniles y aumenta el

número de ejemplares de tallas mayores

(Fig. 5A).

El bajo número de ejemplares supe-

riores a 40 mm en Isla de Pascua,

(octubre, septiembre y junio 1991; sep-

tiembre 1993; marzo y abril de 1994),

podría estar relacionado con la extrac-

ción de este recurso en tallas superiores

a 40 mm, (Obs. per.; González, com.

pers.).

En Plaxiphora mercatoris predominan

los machos, lo que coincide con otras

especies del grupo, GLYNN (1970) indica

que la relación machos:hembras en

Chiton stokesi Broderip, 1832, es de

1,46:1,0, en Acanthozostera gemmata

(Blainville, 1825), es de 1,52:1,0 y en

Chiton marmoratus Gmelin, 1791, es de

1,1:1,0. Orway (1994) encontró una rela-

ción de 2:1 para Onithochiton quercinus

(Gould, 1846), mientras que en Plaxi-

phora albida el mismo autor encontró una

proporción (1,0:0,99) entre ambos sexos

que no difiere significativamente. Por

OSORIO ET AL.: Plaxiphora mercatoris en Isla de Pascua, Chile

otra parte BRANDANI ET AL. (1974)

indican para P. aurata, un 48% de

machos y un 52% de hembras.

Microscópicamente la diferenciación

de sexos en P. mercatoris ocurre tempra-

namente en ambos sexos, a partir de los

15,6 mm de longitud en machos y a los

18,2 mm en hembras, en el primer año

de vida. BRANDANI ET AL. (1974) indica

para P. aurata una talla de 12 mm de lon-

gitud. También GLYNN (1970), encuentra

que la madurez sexual se logra en el

primer año, en las tres especies de poli-

placóforos tropicales que ha estudiado.

Los estudios que describen el hábitat

de otras especies de Plaxiphora.son coin-

cidentes con lo encontrado para P. mer-

catoris. SAITO Y OKUTANI (1991) mencio-

nan que Plaxiphora integra (Taki, 1954) y

Plaxiphora kamehamehae

Bertsch, 1979, viven en áreas con gran

cantidad de algas; LOPEZ Y TABLADO

(1997) indican que Plaxiphora aurata vive

en paredes verticales sombreadas del

intermareal sin embargo mencionan que

faltan completamente sobre sustratos

horizontales.

La densidad de P. mercatoris (1 a 8

individuos /m?), fue ligeramente inferior

a los datos publicados para otros poli-

placóforos. LOPEZ Y TABLADO (1997)

indican para Plaxiphora aurata, densida-

des de 7,3 y 11,8 indiv/m?* y un prome-

dio de 9,5 indiv /m? en Quequen, Argen-

tina. Para la misma especie, BRANDANI

ET AL. (1974) registran valores entre 5 a

20 indiv/m?, en Mar del Plata. GLYNN

(1970) indica para Achantopleura sp. 1 a 8

indiv/m?, y para Chiton sp. 1 a 22

indiv / m?.

En contraste a lo que ocurre general-

mente en los sistemas intermareales, P.

mercatoris no es abundante en Isla de

Pascua. Entre los factores que pueden

causar esto, cabe destacar la acción

antrópica ya que, continuamente son

recolectados por los isleños (González,

com. per.). Por otra parte, la acción de

fenómenos naturales ocasionales,

también colabora a disminuir la densi-

dad de P. mercatoris. Por ejemplo

semanas antes del muestreo de septiem-

bre de 1993, ocurrieron mareas diurnas

extremadamente bajas (Tabla de mareas

Ferreira y

de la Costa de Chile, 1993) que coinci-

dieron con altas temperaturas, lo que

ocasionó una elevada mortalidad en la

especie; los “mamas” se desprendían

fácilmente con una pequeña presión de

los dedos, y durante los días posteriores

al fenómeno se observaron muchos

ejemplares muertos en las rocas (Paka-

rati, com. per.). El efecto de la deshidra-

tación es probablemente importante;

BOYLE (1970), describe que en Sypharo-

chiton pelliserpentis Quoy y Gaimard,

1835, la deshidratación es un factor

importante que afecta la densidad de la

población. El autor determinó que la

especie tolera pérdidas del 75% del con-

tenido de agua, antes de ocurrir una

mortalidad del 50% de la población.

GLYNN (1970) observó mortalidades

naturales como resultado de una pro-

longada exposición a bajas mareas

diurnas, desprendimiento por impacto

del oleaje, abrasión y probablemente

depredación de aves y peces.

Así, P. mercatoris constituye actual-

mente un recurso explotado por los

isleños, cuya población parece mante-

nerse sin llegar a estar sobre-explotada

ya que no se registra una disminución

en la talla media de la población.

Aunque su densidad se mantiene baja,

tanto por la mortalidad natural como

por la antrópica, la población se man-

tendría estable gracias a un rápido

recambio de individuos.

AGRADECIMIENTOS

Los autores agradecen la ayuda reci-

bida en las diferentes etapas del trabajo

a las siguientes personas: Sras: G.

Acosta, M. Bustos , E. Elgueta, Sres. H.

Atan y F. Rocha . A la antropóloga L.

González por informaciones sobre las

costumbres de isleños. A la Dra. L.

Masson por los datos del análisis

químico. A los Sres. O Gálvez, Dr. J.

López Gappa y Dr. P. Penchazadeh, por

el material bibliográfico. Este estudio

fue financiado por el proyecto N*3638-

9312, del Departamento de Investiga-

ción y Desarrollo de la Universidad de

Chile.

Iberus, 18 (2), 2000

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REDHER, H. A,, 1980. The marine mollusks of

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SAITO, H. Y OKUTANI T., 1991. Taxonomy of Ja-

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Plaxiphora. The Veliger, 34(2): 172-194.

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O Sociedad Española de Malacología —___—__———— Iberus, 18 (2): 51-76, 2000

The shallow-water Rissoidae (Mollusca, Gastropoda) of the

Azores and some aspects of their ecology

Los Rissoidae (Mollusca, Gastropoda) de las Azores y algunos aspec-

tos de su ecología

Sérgio P. ÁVILA*

Recibido el 11-VIT-2000. Aceptado el 16-X-2000

ABSTRACT

A critical review of the distribution of the Rissoidae on the islands of the Azorean Archipe-

lago is made, on the basis of bibliographic data as well as newly-collected samples.

Twenty three taxa of Rissoidae are given to the Azores: twelve endemic species, three res-

tricted to the Azores and Madeira/Selvagens archipelago and two with wider distribution.

Of the remaining taxa, one is an unidentified species of Setia and five are records not

confirmed by this study. The assemblage of Rissoidae associated with heterogeneous

algae on a rocky shore on the northern coast of Sáo Miguel Island, Azores, is also descri-

bed. Aspects of community structure [species composition, abundance and zonation) were

studied and a multispecies analysis conducted using clustering techniques. Some com-

ments are also made regarding the Rissoidae speciation that has occurred in the Azores

and its relation to the main sea-surface circulation in this area of the Atlantic Ocean.

RESUMEN

Se realiza una revisión crítica de la distribución de los Rissoidae de las Azores, en base a

datos bibliográficos y a muestras recientes. Se han encontrado 23 táxones de Rissoidae:

12 especies endémicas, 3 restringidas a las Azores y a Madeira/Salvajes y otras dos con

una distribución mayor. De las especies restantes, una es una especie sin identificar de

Setia y otras 5 son citas no confirmadas en el presente estudio. Se describe también la

asociación de Rissoidae con algas en una costa rocosa del norte de Sáo Miguel (Azores).

Se incluyen datos sobre la estructura de la comunidad (composición específica, abundan-

cia y zonación), así como un análisis multiespecífico empleando técnicas de cluster. Se

comenta también la especiación que ha tenido lugar en las Azores con la famila Rissoi-

dae y su relación con las principales corrientes superficiales en esta zona del Atlántico.

KEY WORDS: Rissoidae, Azores, taxonomy, ecology, geographic range.

PALABRAS CLAVE: Rissoidae, Azores, taxonomía, ecología, distribución geográfica.

INTRODUCTION

The marine Rissoidae of the Azores ZENBERG (1889) and NOBRE (1924; 1930),

were studied by DROUÉT (1858), DAUT- who provided annotated check-lists.

* Seccáo de Biologia Marinha and CIRN. Departamento de Biologia, Universidade dos Agores. Rua da Máe de

Deus, 9500 Ponta Delgada. Azores - Portugal avilaWalf.uac.pt

Iberus, 18 (2), 2000

During the scientific expeditions made

by the Prince of Monaco to the Azores,

25 species of Rissoidae were described

from deep-water samples, most of them

Alvania (11 species) (DAUTZENBERG,

1889). Using SEM techniques, two new

species were described by AARTSEN

(1982a; 1982b; 1982c; 1982d) and Amati

(1987), from material collected by the

Prince of Monaco. With the same metho-

dology (SEM photos of the protoconch

and of the microsculpture of the body

whorl), MOOLENBEEK AND FABER (1987)

revised the genus Manzonia in the Maca-

ronesian islands, identifying a single

species from the Azorean archipelago

(Manzonia unifasciata Dautzenberg,

1889).

As a result of the scientific expedi-

tions organized by the Department of

Biology of the University of the Azores

to some of the islands (e.g.: “Gra-

ciosa/88”, “Flores/89”, “Santa Maria e

Formigas 1990” and “Pico/1991”) and

also the scientific expedition “Acores

89”, organized by the Department of

Oceanography and Fisheries (DOP/UA)

of the University of the Azores, several

check-lists (some of them not yet publis-

hed) have allowed preliminary reports

on the geographical distribution of Ris-

soidae species on the islands of the

Azores (AZEVEDO AND MARTINS, 1989;

AZEVEDO, 1990; AZEVEDO AND GOFAS,

1990; ÁVILA AND AZEVEDO, 1996; ÁVILA

AND AZEVEDO, 1997; ÁviLa, 1998;

ÁVILA, AZEVEDO, GONCALVES, FONTES

AND CARDIGOS, 1998; ÁVILA, AZEVEDO,

GONCALVES, FONTES AND CARDIGOS, in

press).

During the “I International Works-

hop of Malacology” held at Vila Franca

do Campo (Sao Miguel island), GOFAS

(1989; 1990) refers to 11 species of Ris-

soidae from the Azorean littoral (Alvania

angioyi Van Aartsen, 1982, A. cancellata

(Da Costa, 1778), A. mediolittoralis Gofas,

1989, A. poucheti Dautzenberg, 1889, A.

sleursi (Amati, 1987), Botryphallus ovum-

muscae (Gofas, 1990), Cingula trifasciata

(Adams, 1798), Crisilla postrema (Gotas,

1990), Manzonia unifasciata (Dautzen-

berg, 1889), Rissoa guernei Dautzenberg,

1889 and Setia subvaricosa Gofas, 1990) to

which we must add Alvania formicarum

Gofas 1989, a species endemic to Formi-

gas and Santa Maria (GOFaAs, 1989;

1990).

Samples taken by the author from

several locations in the Azores revealed

another species at Sáo Miguel island

that was formerly reported by Gofas

(1990) to be restricted to the islands of

the central and western groups 1.e.,

Onoba moreleti Dautzenberg, 1889.

AZEVEDO AND GOFAS (1990) recorded a

species of Setia from Flores. This species

was later found by ÁVILA ET AL. (1998)

at Pico and Sáo Miguel. A new species

of Alvania, described by Hoenselaar and

Goud (1998) as A. internodula, was also

collected from Formigas by Ávila and

AZEVEDO (1997). The revision of the

material of the CANCAP expeditions

(1976-1986) has confirmed some species

and described a few others to the Azores

(Hoenselaar and Goud, 1998).

The Rissoidae is one of the best

represented families of shallow-water

marine molluscs in the Azores, with 8

genera and 18 confirmed taxa, of which

12 species are considered as endemic

(MOOLENBEEK AND FABER, 1987; GOFAS,

1989; 1990; KNUDSEN, 1995; HOENSE-

LAAR AND GOUD, 1998).

This study had three main objecti-

ves: to undertake a taxonomic revision

of Rissoidae in the Azores, to identify

any island to island endemisms, that is

species restricted to some of the islands

and to describe the zonation of the Ris-

soidae on the Azorean shores.

MATERIAL AND METHODS

A bibliographic analysis was made,

in order to compile published informa-

tion about the shallow-water Rissoidae

of the Azores (intertidal to a depth of

about 50m). The synonymy and the dis-

tribution of the species, by islands, was

also annotated. A table with the distri-

bution of the Rissoidae species, by

islands, was constructed and multiva-

riate analysis was performed on the

data obtained (Bray-Curtis similarity

index/UPGMA as well as MDS). The

ÁvILA: The shallow-water Rissoidae of the Azores and some aspects of their ecology

* VESTERN

GROUP

Flores ñ

Graciosa

Faial 5)

Pico

31 W 30 Y 29 Y

Figure 1. Map of the Azores Archipelago.

Figura 1. Mapa del archipiélago de las Azores.

gastropod classification follows PONDER

AND LINDBERG (1997). CLEMAM (Check

List of European Marine Mollusca) data-

base classification was quite useful for

synonymy.

Protoconchs of almost all the

Azorean Rissoidae were photographed

with a SEM, as well as other detailed

structures of the shells (e.g. microsculp-

ture of the last whorl and protoconch,

shell aperture and the whole shell).

The zonation of the Rissoidae was

examined for Sáo Miguel, which is

located in the eastern group of islands of

the archipelago (Fig. 1). Several dives

were performed in July 1996 at Porto da

Baleia, a former whaling ramp boat,

located at Sáo Vicente (Capelas) on the

north coast of Sáo Miguel (Fig. 2). In the

selected zone, a 400m long transect was

done, from the intertidal zone to a depth

of 30m (Fig. 3). Quadrates of 50x50 cm,

placed on algae covering the rocky subs-

tratum were scrapped, and the material

collected put into labeled cotton draws-

tring bags. Three replicates were obtai-

ned from eight chosen deptbs, ¡.e., 3.5m,

5.1m, 8m, 12m, 13.6m, 16.3m, 22m and

26.8m. In the laboratory, each of the

replicates was washed several times and

the animals removed from the algae by

CENTRAL|GROUP

a Sao Jorge

28” W

40% N

39 N

38 N

Sao Miguel

EASTERN | Formigas

GROUP

Santa Maria

islets

37 N

25” W

26" W

27. W

pouring the washing water through a

sieve tower with decreasing mesh sizes

(Imm, 0.5mm and 0.25mm). Samples

were then labeled and preserved in 70%

ethanol. The molluscs from the 1mm

mesh were sorted and the Rissoidae

identified and counted.

Multispecies analysis between all the

samples were conducted using ordina-

tion techniques. Prior to the multivariate

analysis, the absolute values of the

counts were transformed, in order to

standardize the data and ensure that the

multivariate ordination would not be

determined by the most abundant

species (CLARKE AND AINSWORTH, 1993).

Absolute counts were transformed by

double square root transformation,

which weights the abundant species and

is advisable when a Bray-Curtis

measure is used as a similarity coeffi-

cient in further steps (FIELD, CLARKE

AND WARWICK, 1982).

Triangular matrices of similarities

between every pair of samples were

then computed from transformed data

of absolute counts, using the Bray-

Curtis coefficient (FIELD ET AL., 1982;

CLARKE AND AINSWORTH, 1993). The

similarity matrices were subjected to

clustering by an hierarchical agglomera-

Iberus, 18 (2), 2000

Lat 37% 50” N

A Long 25” 50” W

Figure 2. Sáo Miguel island. SVC - Sáo Vicente.

Figura 2. Isla de Sáo Miguel. SVC - Sáo Vicente.

tive method employing group-average

linking (UPGMA).

Data analysis were undertaken

using the PRIMER (Plymouth Routines

in Multivariate Ecological Research) set

of programs developed and tested by

Plymouth Marine Laboratory.

All the material is deposited at the

reference collection of the Department

of Biology of the University of the

Azores (DBUA), unless otherwise

stated.

RESULTS

0 kl

Abbreviations used in text:

DBUA: marine molluscs reference col-

lection of the Department of Biology

of the University of the Azores.

MCM(HN): Museu Carlos Machado

/ História Natural, Ponta Delgada,

Azores.

MNHN: Muséum National d Histoire

Naturelle, Paris (Malacologie).

NNM: Nationaal Natuurhistorisch

Museum, Leiden.

Phylum MOLLUSCA

Class GASTROPODA

Subclass ORTHOGASTROPODA

Superorder CAENOGASTROPODA

Order SORBEOCONCHA

Suborder HYPSOGASTROPODA

Superfamily RISSOOIDEA

Family RISSOIDAE

Alvania Risso, 1826

Alvania abstersa Van der Linden and Van Aartsen, 1994

References to the Azores:

Alvania obsoleta Van der Linden, 1993: 79-82.

Alvania abstersa Van der Linden and Van Aartsen, 1994: 2; Hoenselaar and Goud, 1998:71.

Occurrence: Pico(Lajes do Pico), Ter-

ceira (Porto Martins), Sáo Jorge (Faja da

Caldeira), Sao Miguel (Lagoa and Mos-

teiros), Santa Maria (VAN DER LINDEN,

1993: 80). Azores (CANCAP expeditions)

(HOENSELAAR AND GOUD, 1998: 71).

DBUA 726.

Comments: although more common

just below the intertidal, it may appear

to a depth of 35m.

Dimensions: up to 3.3 mm long,

1.7mm wide.

Geographic distribution: endemic to

the Azores.

ÁviLa: The shallow-water Rissoidae of the Azores and some aspects of their ecology

Transect (m)

0 50 100 150 200

Depth (m)

250 300 350 400 450

Figure 3. Transect performed at Porto da Baleia (Sáo Vicente, Capelas), north coast of Sio Miguel

island, Azores.

Figura 3. Transecto realizado en Porto da Baleia (Sáo Vicente, Capelas), costa N de la isla de Sáo

Miguel, Azores.

Alvania angioyi Van Aartsen, 1982

References to the Azores:

Alvania (Alvinia) watsoni Schwartz, in Watson, 1873: Dautzenberg, 1889: 51.

Alvania watsoni (Schwartz MS) Watson, 1873: Bullock, Turner and Fralick, 1990: 45.

Alvania angioyi Van Aartsen, 1982: Azevedo and Martins, 1989: 69; Gofas, 1990: 112; Azevedo and

Gofas, 1990: 85; Ávila, 1996: 27; Ávila and Azevedo, 1996: 106; Ávila and Azevedo, 1997: 326;

Bullock, 1995: 16; Knudsen, 1995: 140; Hoenselaar and Goud, 1998: 72.

Alvania angioyi Van Aartsen, 1982d: Azevedo 1991b: 44.

Occurrence: Sáo Miguel, Faial (Bay of

Horta, -15 to -20 m, Stn. 103, “Hironde-

lle”, 1887), Pico (-1287 m) (DAUTZEN-

BERG, 1889: 51), Graciosa (AZEVEDO AND

MARTINS, 1989: 69), Flores (Santa Cruz, -

20m) (AZEVEDO AND GOFAs, 1990: 85),

Sáo Miguel (BULLOCK ET AL., 1990: 45);

Sáo Miguel (Vila Franca (-24m) and

Morro das Capelas (“Biacores” 1971

expedition); Lagoa (-10 to -22m); Fetei-

ras (-15 to -22m); Ponta da Galera (inter-

tidal and -13 to -18m); Ponta Delgada;

Capelas (intertidal); llhéu de Vila

Franca; Ponta da Pirámide (-13m)), Faial

(Horta, -3m; -7m; Monte da Guia, -20m,

“Biacores” 1971), Terceira (Pedra

Furada, Angra do Heroísmo; Praia da

Vitória), Flores (“Pr. Alice” st. 703, 1896;

Santa Cruz, -20m) and Formigas (west

coast, -l6m, “Biacores” 1971) (GOFas,

1990: 112), Sáo Miguel (Caloura and

Ribeirinha) (AZEVEDO, 1991la: 22), Santa

Maria (Vila do Porto, Ponta do Marvao)

(AZEVEDO, 1991b: 44), Sáo Miguel (Mhéu

de Vila Franca) (BuLLock, 1995: 16;

KNUDSEN, 1995: 140), Pico (intertidal)

(ÁviLa, 1996: 27), Pico (ÁVILA AND

AZEVEDO, 1996: 106), Formigas islets

(ÁVILA AND AZEVEDO, 1997: 326) and

Azores (CANCAP expeditions) (HOEN-

SELAAR AND GOUD, 1998: 72).

DBUATI94A73, 188,227, 2/4 9217,

281, 335, 340, 343, 350, 352, 353, 35), 372,

374, 379, 394, 398, 400, 407, 410, 412, 462,

493, 496, 499, 556, 560, 564, 568, 571, 574,

579, 666, 675, 715, 719.

MCM(HN)7, 9.

Comments: this species is closely

related to specimens from Madeira and

is also similar to Alvania oranica (Pallary,

1900) from Ceuta (Gorras, 1990). It

occurs to a depth of 35m, but is commo-

nest in the first 10m. Although Gofas

raises the hypothesis of its existence at

Madeira archipelago, its presence was

not confirmed in the samples from the

CANCAP expeditions (HOENSELAAR

AND GOUD, 1998: 72).

Dimensions: 1.8mm long, 1.1 mm

wide.

Geographic distribution: endemic to

the Azores.

Iberus, 18 (2), 2000

Alvanta beani (Hanley in Thorpe, 1844)

References to the Azores:

Rissoa calathus Forbes and Hanley, 1858: Mac Andrew, 1856: 121.

Alvania (Turbona) reticulata (Montagu, 1803): Simroth, 1888.

Occurrence: Azores (-10 to -90m)

(MAC ANDREW, 1856: 121). Sáo Miguel

(Ponta Delgada) (SimrOoTH, 1888).

Comments: this record needs to be

confirmed. Its occurrence in the Azores

is possible and, if so, it lives predomi-

nantly at depths >50m and I have only

a few samples collected by scuba

diving at these depths. However, in the

significant amount of samples

collected at the Azores by the

CANCAP expeditions, not a single

specimen was found (HOENSELAAR

AND GOUD, 1998).

Dimensions: 3.5mm long, 2.0mm

wide (FRETTER AND GRAHAM, 1978).

Geographic distribution: Norway to

the Mediterranean, Azores (?) and

Canary Islands (FRETTER AND GRAHAM,

1978; ROLÁN, 1984).

Alvanta cancellata (Da Costa, 1778)

References to the Azores:

Rissoa crenulata Michaud, 1832: Mac Andrew, 1856: 148.

Rissoa (Alvania) cancellata Da Costa: Watson, 1886: 592.

Alvania laxa Dautzenberg and Fischer, 1896: 62-63, pl. 19, figs. 10,11.

Alvania cancellata Da Costa: Dautzenberg, 1889: 49.

Alvania cancellata (Da Costa, 1778): Nobre, 1924: 80; 1930: 57; Morton, 1967: 36; Azevedo, 1990: 59;

Gofas, 1990: 104; Azevedo and Gofas, 1990: 85; Azevedo, 1991a: 21; 1991b: 44; Ávila, 1996: 27;

Ávila and Azevedo, 1997: 326; Hoenselaar and Goud, 1998: 73.

Occurrence: Azores (MAC ANDREW,

1856: 148); Faial (-823 to -914m)

(WATSON, 1886: 592), Faial (Horta, -15 to

-20m, Stn. 103, “Hirondelle”, 1887), Pico

(-1287m) and Sáo Miguel (DAUTZEN-

BERG, 1889: 49), Azores (NOBRE, 1924: 80;

1930: 57), Sao Jorge (Velas) (MORTON,

1967: 36), Pico (Baía de Sao Pedro, Lajes

do Pico) (AZEVEDO, 1990: 59), Flores

(Santa Cruz) (AZEVEDO AND GOFAS,

1990: 85), Sao Miguel (Ponta Delgada;

Vila Franca do Campo, -10m, -24m;

Morro das Capelas (“Biacores” 1971

expedition); Feteiras, -15 to -22m; Ponta

da Galera, -13 to -18m, -20m, Lagoa, -10

to -22m; Ponta da Pirámide, -13m), For-

migas (east coast, -16m), Terceira (Ponta

de Sao Diogo, Pedra Furada - Angra do

Heroísmo), Flores (Santa Cruz, -20m)

and Pico (-1287m) (GOFas, 1990: 104),

Sao Miguel (infralittoral of Ribeirinha)

(AZEVEDO, 1991a: 21), Santa Maria (Vila

do Porto, Ilhéu da Vila) (AZEVEDO,

1991b: 44), Sáo Miguel (Ilhéu de Vila

Franca) (KNUDSEN, 1995: 141), Pico (sub-

5Ó

tidal) (ÁviLa, 1996: 27), Formigas islets

(ÁvILA AND AZEVEDO, 1997: 326) and

Azores (CANCAP expeditions) (HOEN-

SELAAR AND GOUD, 1998: 73).

DIU MS MS, 197, 220),

274, 281, 341, 350, 379, 394, 395, 405, 408,

410, 411, 415, 421, 422, 438, 441, 446, 448,

459, 489, 493, 496, 499, 500, 555, 558, 561,

569, 570, 574, 579, 605, 608, 609, 614, 658,

659, 660, 661, 662, 665, 666, 667, 670, 672,

OACI OS

Comments: it occurs from low tide le-

vel to a depth of 45m (SALDANHA, 1995),

but its presence at low depths is rare

(GRAHAM, 1988; GOFaS, 1990). It is detri-

tivorous (GRAHAM, 1988). The popula-

tions in the Azores are conspecific with

those on European mainland, their pro-

toconchs matching exactly (KNUDSEN,

1995). GOrAS (1990) also states their

conspecificity with the populations of

Madeira and the Canary Islands, be-

cause of external similarities and the

existence of a multispiral protoconch,

denoting a planktotrophic development.

ÁVILA: The shallow-water Rissoidae of the Azores and some aspects of their ecology

Dimensions: 3.7 mm long, 2.4 mm

wide.

Geographic distribution: Atlantic,

Azores, English Channel and North Sea

(CAMPBELL, 1994), Madeira (NOBRE,

1889, 1937; Goras, 1990), British isles,

Mediterranean, Canary islands and

Cape Vert Islands (KNUDSEN, 1995). Sáo

Tomé island (FERNANDES AND ROLÁN,

1993).

Alvantia cimex (Linnaeus, 1758)

References to the Azores:

Rissoa granulata Philippi, 1836: Mac Andrew, 1856: 121.

Occurrence: Azores from shores

(dead) (MAC ANDREW, 1856: 121).

Comments: DAUTZENBERG (1889),

raised doubts about the specific status

of the specimen(s) identified by Mac

Andrew. 1 agree with him and

consider that this record needs to be

confirmed.

Alvanta formicarum Gofas, 1989

References to the Azores:

Alvania formicarum Gofas, 1989: 40-41; Hoenselaar and Goud, 1998: 72.

Occurrence: Formigas and Ilhéu de

Sao Lourenco (Santa Maria) (GOFAS,

1989:40-41).

DBUA 332, 335, 338, 340, 341, 342,

343, 345, 348, 350, 352, 353, 355, 359.

Comments: the zonation of this

species is not known. However, living

specimens have been collected from 15

to 43m depth.

Dimensions: 2.4mm long, 1.4mm

wide.

Geographic distribution: restricted to

Formigas islets and Santa Maria

island.

Alvania internodula Hoenselaar and Goud, 1998

References to the Azores:

Alvania sp.: Ávila and Azevedo, 1997: 326.

Alvania internodula Hoenselaar and Goud, 1998: 83.

Occurrence: Formigas (ÁVILA AND

AZEVEDO, 1997: 326). Azores (CANCAP

expeditions: Sta. 5033, 33m/1 specimen,;

Sta. 5039, 43m/2; Sta. 5040, 41-47m/25;

Sta. 5091, 33m/7; Sta. 5098, 40m/1; Sta.

5113, 45m/12; Sta. AZO.022, at shore/1)

(HOENSELAAR AND GOUD, 1998: 83).

DBUA 336, 338.

Comments: in their check-list of the

shallow-water marine molluscs of Formi-

gas, ÁVILA AND AZEVEDO (1997) indica-

ted the presence of a new species of Alva-

nia in 6 lots of the DBUA collection:

DBUA 332, 335, 336, 338, 350, 355. Later

work at the SEM level revealed the exis-

tence of a species already described as A.

internodula Hoenselaar and Goud, 1998.

Also, only the specimens in the DBUA

336 and 338 lots were correctly assigned

to this new species, all other specimens

being Alvania angioyi Van Aartsen, 1982d.

The shells of the young specimens of Al-

vania internodula resemble Alvania angioyi

Van Aartsen, 1982d, but the adults are

quite different, with stronger knobs in the

whorls and with deeper sutures.

Additional description: Protoconch sculp-

tured with 5-6 marked spiral ribs, the in-

terstices covered with numerous very

small nodules, not aligned. In the second

whorl of the teleoconch, the intermediate

3 ribs are more prominent than the others,

Iberus, 18 (2), 2000

the same happening in the body whorl.

The crossings of spiral ribs and costae pro-

duce nodules, that are stronger in the in-

termediate 3 ribs. The ribs located in the

anterior part of the body whorl are quite

smooth. Inside the outer lip there are 8-9

faint denticles (Fig. 8: A-J).

Dimensions:2.3mmlong, 1.32mm wide.

Geographic distribution: restricted to

the Formigas islets, Azores.

Alvania mediolittoralis Gofas, 1989

References to the Azores:

Alvania mariae (D'Orbigny): Dautzenberg, 1889: 49.

Rissoa (Alvania) reticulata Montagu var. mariae D'Orbigny: Nobre, 1924: 81.

Alvania (Turbona) reticulata (Montagu, 1803): Martins, 1980: 17 (misidentification, A.M.F. Martins,

pers. comm).

Alvania mediolittoralis Gofas, 1989: Gofas, 1989: 39; Azevedo and Martins, 1989: 69; Azevedo,

1990: 59; Azevedo and Gofas 1990: 85; Gofas, 1990: 110-112; Azevedo, 1991a: 21; 1991b: 44;

Ávila, 1996:27; Hoenselaar and Goud, 1998: 91.

Occurrence: Sao Miguel and Pico (-

1287m) (DAUTZENBERG, 1889: 49), Sáo

Miguel (Ponta Delgada; Praia do Rosto

do Cao), Terceira, Pico and Graciosa

(NOBRE, 1924: 81; 1930: 57), Sao Miguel

(Atalhada, Lagoa) (MARTINS, 1980: 17),

Graciosa (mediolittoral of Porto Afonso

and Santa Cruz; infralittoral of Baía da

Folga) (AZEVEDO AND MARTINS, 1989:

69), Flores (Santa Cruz) (AZEVEDO AND

GOFAS, 1990: 85), Pico (Baía de Sáo

Pedro, Lajes do Pico) (AZEVvEDO, 1990:

59), Sao Miguel (Caloura, -4m; Vila

Franca, -9m, -24m “Biacores” 1971 expe-

dition; Ponta da Galera, intertidal;

Capelas, intertidal; Água d'Alto, interti-

dal; Calheta - Ponta Delgada, intertidal),

Faial (Horta, -7m), Terceira (Praia da

Vitória; Cais da Silveira; Pedra Furada -

Angra do Heroísmo); Flores (Santa Cruz)

(GOFAs, 1989: 39; 1990: 110), Sáo Miguel

(mediolittoral of Caloura; infralittoral of

Caloura and Ribeirinha) (AZEVEDO,

1991a: 21), Santa Maria (Vila do Porto,

Ponta do Marváo) (AZEVEDO, 1991b: 44),

IIhéu de Vila Franca (BULLOCK, 1995: 16),

Pico (mediolittoral) (ÁviLA, 1996: 27) and

Azores (CANCAP expeditions) (HOEN-

SELAAR AND GOUD, 1998: 91).

DBUA 124, 188, 193, 197, 229, 240,

274, 442, 444, 445, 446, 448, 449, 450, 451,

452, 453, 455, 456, 457, 458, 459, 460, 461,

462, 471, 473, 474, 475, 476, 483, 486, 489,

490, 492, 493, 496, 499, 500, 551, 553, 558,

560, 561, 564, 565, 566, 568, 570, 571, 574,

579, 614, 632, 659, 661, 662, 663, 665, 666,

COTAS SAO:

Comments: common in sheltered

places, especially under rocks. Sometimes

present in large numbers in the intertidal

zone, together with Fossarus ambiguus (Lin-

naeus, 1758) and Cingula trifasciata

(Adams, 1798) (GOFAs, 1990). It is similar

to Alvania manzonía (Nordsieck, 1972) from

the Canary Islands and Selvagens, and

resembles also Alvania leacocki (Watson,

1873), from Madeira (GOFAS, 1989).

Dimensions: up to 2.7 mm long, 1.5

mm wide.

Geographic distribution: Azores and Ma-

deira archipelago (CANCAP expeditions,

Sta. 1.D48, 0-22m/1 specimen, Sta.1.K14,

at shore /1 and Sta. 1.K16, at shore /2) (Ho-

ENSELAAR AND GOUD, 1998: 91).

Alvania poucheti Dautzenberg, 1889

References to the Azores:

Alvania poucheti Dautzenberg, 1889: 49-50; Bullock et al., 1990: 45; Gofas, 1990: 108; Morton and

Britton, 1995: 70; Knudsen, 1995: 141; Ávila, 1996: 27; Ávila and Azevedo, 1996: 106; Ávila and

Azevedo, 1997: 326; Ávila et al., 1998: 497; Hoenselaar and Goud, 1998: 99.

Alvania poucheti var. cingulifera Dautzenberg, 1889: 50.

ÁVILA: The shallow-water Rissoidae of the Azores and some aspects of their ecology

Occurrence: Sáo Miguel (DAUTZEN-

BERG, 1889: 49-50), Sao Miguel (Ponta da

Galera; Queimada, Água d'Alto; Mostei-

ros; llhéu de Vila Franca) (BULLOCK ET AL.,

1990: 45), Sao Miguel (Capelas, -24m

(“Biacores” 1971 expedition); Feteiras, -15

to -22m; Ponta Delgada, -10 to -20m;

Lagoa; Ilhéu de Vila Franca, -1m; Ponta

da Pirámide, -13m; Calheta, Ponta

Delgada; Ponta da Galera, -13 to -18m),

Faial (Horta, -17 and -20m), Terceira (Praia

da Vitória; Angra do Heroísmo, Pedra

Furada) (GoFas, 1990: 108), Sio Miguel

(off Vila Franca do Campo) (MORTON AND

BRITTON, 1995: 70), Sáo Miguel (Ilhéu de

Vila Franca) (KNUDSEN, 1995: 141), Pico

(intertidal) (ÁviLA, 1996: 27), Pico (ÁVILA

AND AZEVEDO, 1996: 106), Formigas islets

(ÁVILA AND AZEVEDO, 1997: 326), Flores

(ÁVILA ET AL., 1998: 497) and Azores

(CANCAP expeditions) (HOENSELAAR

AND GOUD, 1998: 99).

DBUA 119, 173, 240, 350, 352, 353, 355,

447, 465, 493, 499, 500, 556, 563, 570, 631, 666.

MCM(HN) 3, 11, 107, 108.

Comments: usually associated with

brown algae. It may occur at 20m depth,

but is commonest in the first 10m.

Dimensions: up to 2.2mm long,

1.3mm wide.

Geographic distribution: endemic to

the Azores.

Alvanía sleursi (Amati, 1987)

References to the Azores:

(?) Rissoa (Alvania) hispidula Monterosato: Watson, 1886: 593 (misidentification).

Alvania hirta Monterosato: Dautzenberg and Fischer, 1896: 456 (misidentification).

Manzonia sleursi Amati, 1987: 25-30.

Alvania sleursi (Amati, 1987): Gofas, 1990: 107; Knudsen, 1995: 142; Ávila, 1996: 27; Ávila and

Azevedo, 1996: 106; Ávila and Azevedo, 1997: 326.

Occurrence: Faial (-823 to -914 m)

(WATSON, 1886: 593), Banco Princesa

Alice (st. 46, -1385 m) (DAUTZENBERG

AND FISCHER, 1896: 456), Sáo Miguel

(Ponta da Galera, -7 to -8m, -13 to -18m

and -20m; Vila Franca, -24m; Morro das

Capelas, -15 to -20m,; Lagoa, -10 to -22m;

Feteiras, -15 to -22m; Ponta da Pirámide,

-13m; Ilhéu de Vila Franca, -1m), Pico (-

1287m), Terceira (Angra do Heroísmo,

Pedra Furada; Praia da Vitória), Flores

(Santa Cruz, -40m) (GOFAas, 1990: 107), Il-

héu de Vila Franca (KNUDSEN, 1995: 142),

Pico (intertidal) (ÁviLa, 1996: 27), Pico

(ÁvILa AND AZEVEDO, 1996: 106) and

Formigas islets (ÁVILA AND AZEVEDO,

1997: 326).

DBUA 173, 335, 340, 341, 342, 343,

350, 352, 353, 355, 446, 448, 458, 459, 493,

496, 499, 500, 666, 667, 719.

MCM(HN) 40.

Comments: occurs from the intertidal

to 45m depth, being more abundant on

rocky shores, between -10 to -20m. HOEN-

SELAAR AND GOUD (1998) reported this spe-

cies to Selvagens archipelago (CANCAP

expeditions, Sta. 3070, 645m depth / 8 speci-

mens; Sta. 3072, 830m / 3; Sta. 3087, 322m/8,

with all specimens strongly eroded).

Dimensions: up to 2.5 mm long, 1.6 mm

wide.

Geographic distribution: Azores and Sel-

vagens archipelago (HOENSELAAR AND

GOUD, 1998: 103).

Alvania tarsodes (Watson, 1886)

References to the Azores:

Rissoa (Alvania) tarsodes Watson, 1886: 595, pl. XLIV, fig. 2.

Alvania tarsodes (Watson, 1886): Bouchet and Warén, 1993: 642; Hoenselaar and Goud, 1998: 106.

Occurrence: Azores, from 35m depth to

620m (HOENSELAAR AND GOUD, 1998: 106).

Comments: although first reported to

be a bathyal species (480-1385m depth,

Iberus, 18 (2), 2000

BOUCHET AND WARÉN, 1993), HOENSE-

LAAR AND GOUD (1998) have found spe-

cimens in some shallow samples of the

CANCAP expeditions (Sta. 5033, 33m/3

specimens; Sta. 5039, 43m/8; Sta. 5040,

41-47m/8; Sta. 5050, 55m/3; Sta. 5096,

52m/4; Sta. 5100, 55m/1 and Sta. 5113,

45m/1).

Dimensions: 2.2mm long, 1.3mm

wide (WATSON, 1886).

Geographic distribution: restricted to

the Azores.

Botryphallus Ponder, 1990

Botryphallus ovummuscae (Gofas, 1990)

References to the Azores:

Peringtella nitida Brusina: Dautzenberg, 1889: 53.

Cingula (Peringiella) nitida (Brusina) Monterosato: Martins, 1980: 5.

“Peringiella” sp.: Azevedo and Gofas, 1990: 85.

“Peringiella” ovummuscae (Gofas, 1990): Gofas, 1990: 119-121, fig. 11.

Botryphallus ovummuscae (Gofas, 1990): Ávila, 1996: 27; 1998: 466; Ávila and Azevedo, 1996: 106.

Occurrence: Faial (-15 to -20m) and Sáo

Miguel (DAUTZENBERG, 1889, p. 53), Ter-

ceira (Poca dos Frades, Silveira; Caminho

de Baixo, Sáo Mateus) e Sáo Miguel (Água

d'Alto; Pópulo; Atalhada, Lagoa) (MaAr-

TINS, 1980, pp. 9-16), Flores (Santa Cruz,

mediolittoral) (AZEVEDO AND GOFAS, 1990:

85), Sao Miguel (Ponta da Galera, interti-

dal; Calheta, Ponta Delgada), Terceira

(Porto Martins), Flores (Santa Cruz, up-

per intertidal zone), (GOFAS, 1990, p. 120),

Pico (intertidal) (ÁviLaA, 1996: 27; 1998: 466)

and Pico (ÁVILA AND AZEVEDO, 1996: 106).

DBUA 209, 493, 499, 500, 659, 661,

662, 665, 666, 715.

Comments: according to GOFAS

(1990), there are related species in the

Straits of Gibraltar (Peringiella epidaurica

Brusina, 1886), at Madeira and also on

the Canary Islands (in this last archipe-

lago there are two species similar to the

Azorean one).

Dimensions: up to 1.3 mm long, 0.7

mm wide.

Geographic distribution: endemic to

the Azores.

Cingula Fleming, 1828

Cingula ordinaria Smith

References to the Azores:

Cingula ordinaria Smith: Chapman, 1955: 803.

Occurrence: Faial (Feteira, mid-tide)

(CHAPMAN, 1955: 803).

Comments: this species was pro-

bably misidentified. Most probably, it

represents Cingula trifasciata (J. Adams,

1800), the only representative of this

genus that lives at the Azores Archipe-

lago.

Cingula trifasciata (J. Adams, 1800)

References to the Azores:

Rissoa (Cingula) cingillus Montagu, 1803: Mac Andrew, 1856: 148.

Cingula cingillus Montagu, 1803: Dautzenberg, 1889: 52; Knudsen, 1995: 143.

Rissoa (Cingula) cingillus Montagu, 1803: Nobre, 1924: 80; 1930: 57.

Cingula (Cingula) cingillus (Montagu, 1803): Martins, 1980: 5; Lemos and Viegas, 1987: 65.

Cingula trifasciata (Adams, 1798): Azevedo and Gofas, 1990: 85.

ÁVILA: The shallow-water Rissoidae of the Azores and some aspects of their ecology

Cingula trifasciata, (Adams, 1800): Gofas, 1990: 97-134; Bullock, 1995: 9-55; Ávila, 1996: 27 Ávila

and Azevedo, 1997:326.

Occurrence: Azores, at shore (Mac

Andrew, 1856: 122, 148), Sáo Miguel

(DAUTZENBERG, 1889: 52), Sáo Miguel

(Ponta Delgada), Faial (Horta), Terceira

(Angra do Heroísmo), Graciosa, Pico and

Sáo Jorge (Calheta; Velas) (NOBRE, 1924: 80;

1930: 57), Sáo Jorge (Velas) (MORTON, 1967:

36), Terceira (Poca dos Frades, Silveira;

Fanal, Sáo Pedro, Angra do Heroísmo;

Caminho de Baixo, Sáo Mateus) and Sao

Miguel (Água d'Alto; Calheta, Ponta

Delgada; Pópulo; Atalhada, Lagoa)

(MARTINS, 1980: 9-17), Sáo Miguel (Vila

Franca do Campo: intertidal) (LEMOS AND

VIEGAS, 1987: 65), Flores (mediolittoral of

Santa Cruz) (AZEVEDO AND GOFAS, 1990:

85), Sáo Miguel (Ponta da Galera, interti-

dal; Água d'Alto, intertidal) (GOFAS, 1990:

119), Sáo Miguel (Mhéu de Vila Franca)

(BuLLock, 1995: 9-55), Pico (intertidal)

(ÁviLa, 1996: 27) and Formigas islets

(ÁVILA AND AZEVEDO, 1997: 326).

DBUA 128, 205, 240, 352, 442, 445,

448, 449, 457, 460, 461, 470, 474, 475, 489,

490, 496, 499, 500, 659, 632, 660, 661, 662,

663, 665, 666, 667.

Comments: NOBRE (1924; 1930) states

that this species is common in littoral

debris. It is extremely common in shelte-

red places, especially under rocks

(ÁVILA AND AZEVEDO, 1997). This

species is detritivorous and usually

occurs from the upper limit of barnacles

(Chthamalus stellatus) to a few meter's

depth (GRAHAM, 1988). It has non-

planktotrophic development (KNUDSEN,

1995) and because of this, GOFAs (1990)

has some doubts about its conspecificity

with populations in Europe. The speci-

mens collected by the author at Lajes do

Pico, have a wide range of external

color, from almost black to light-brown

(pers. obs.).

Dimensions: 3.9 mm long, 2.1 mm

wide.

Geographic distribution: Bay of Biscay

to the West coast of Norway, the English

Channel, Azores (GRAHAM, 1988; POPPE

AND GOTO, 1991; HAYWARD, WIGHAM

AND YONOW, 1995; KNUDSEN, 1995),

Madeira (NOBRE, 1937), Berlenga (Portu-

gal) (BURNAY, 1986).

Crisilla Monterosato, 1917

Crisilla postrema (Gotas, 1990)

References to the Azores:

Setia abjecta (Watson, 1873): Dautzenberg, 1889: 52.

Setia picta (Jeffreys, 1867): Dautzenberg, 1889: 53.

Alvania (Crisilla) sp.: Azevedo and Gofas, 1990: 85.

Alvania (Crisilla) postrema Gotas, 1990: 114.

Alvania postrema Gofas, 1990. Azevedo, 1991b: 44; Ávila and Azevedo, 1997: 326; Hoenselaar and

Goud, 1998: 99.

Occurrence: Azores (colec. G. Doll-

fus), Sao Miguel and Faial (-15 to -20 m)

(DAUTZENBERG, 1889: 53), Flores (Santa

Cruz) (AZEVEDO AND GOFAS, 1990: 85),

Sáo Miguel (Capelas, -12m; llhéu dos

Mosteiros, -3 to -5m; Morro das Capelas,

intertidal and at -29m (“Biacores” 1971

expedition); Ponta da Galera, intertidal

and from -13 to -18m; Ponta Delgada, -

10 to -20m, Ilhéu de Vila Franca; Ponta

da Pirámide, -13m; Calheta, Ponta Del-

gada), Faial (Horta, -3m, -7m and -17m),

Terceira (Praia da Vitória; Angra do He-

roísmo, Pedra Furada), Flores (Santa

Cruz, intertidal pool), Formigas (-16m)

(GoFas, 1990: 114-115), Caloura and Ri-

beirinha (AZEVEDO, 199la: 22), Santa

Maria (Vila do Porto, llhéu da Vila,

Ponta do Marváo) (AZEVEDO, 1991b: 44),

Sao Miguel (Ilhéu de Vila Franca) (Bu-

LLOECK, 1995: 16), Pico (intertidal)

(ÁviLa, 1996: 27), Pico (ÁVILA AND AZE-

VEDO, 1996: 106) and Formigas islets

(ÁVILA AND AZEVEDO, 1997: 326).

Iberus, 18 (2), 2000

DBUANRIA73 11887198 2748277;

340, 350, 351, 352, 353, 355, 359, 447, 462,

465, 470, 472, 492, 496, 499, 500, 545, 564,

632, 670.

Comments: this is an uncommon

species. It occurs from the low-tide level

to a depth of 20m. Two specimens were

recently found at Madeira (CANCAP

expeditions, Sta. 4.K27, at shore) (HOEN-

SELAAR AND GOUD, 1998: 99).

Dimensions: 1.5mm long, 0.9mm wide.

Geographic distribution: Azores and

Madeira (HOENSELAAR AND GOUD, 1998:

99)

Manzonia Brusina, 1870

Manzonia unifasciata Dautzenberg, 1889

References to the Azores:

Manzonia costata J. Adams, 1797 var. ex coloure: unifasciata: Dautzenberg, 1889: 51, pl. II, fig.10.

Manzonia costata J. Adams var. ex coloure: bifasciata: Dautzenberg, 1889: 51, pl. IL, fig. 9.

Manzonia costata J. Adams var. ex coloure: luteola: Dautzenberg, 1889: 51.

Manzonia aurantiaca (Watson, 1873): Dautzenberg, 1889: 52.

Manzonia costata (Adams, 1797). Pico (Nobre, 1924: 80; 1930: 56).

Alvania (Manzonia) crassa (Kanmacher, 1798): Morton, 1967: 36.

Manzonia aurantiaca (Watson, 1873): Nordsieck, 1972: 176, pl. R VI fig. 2.

Alvania (Manzonia) costata (Adams): Martins, 1980: 5, 16.

Manzonia unifasciata Dautzenberg, 1889: Moolenbeek and Faber,1987, p.26, fig. 42; Azevedo and

Martins, 1989: 69; Azevedo, 1990: 59; Azevedo and Gofas, 1990: 85; Gofas, 1990: 116, figs. 9; 59-

64; Azevedo, 199la: 22; Ávila, 1996: 27; Ávila and Azevedo, 1997: 326.

Manzonia crassa (Kanmacher, 1798) (misidentification?): Bullock et al., 1990: 45.

Manzonia unifasciata (Dautzenberg, 1889): Azevedo, 1991b: 44.

Alvania crassa (Kanmacher, 1798) (misidentification?). Knudsen, 1995: 141.

Occurrence: Sáo Miguel (Ponta Del-

gada), Faial (Horta), Pico, Graciosa and

Terceira (Angra do Heroísmo) (DAUT-

ZENBERG, 1889: 51-52), Sáo Miguel

(Ponta Delgada), Faial (Horta), Pico,

Graciosa and Terceira (Angra do Hero-

ísmo) (NOBRE, 1924: 80; 1930: 56) Sáo

Jorge (Velas) (MORTON, 1967: 36), Pico (-

1276m) and Sao Miguel (NORDSIECK,

1972: 176), Sáo Miguel (Brejela, Atal-

hada, Lagoa) (MARTINS, 1980: 5, 16),

Graciosa (mediolittoral of Fonte da

Areia, Porto Afonso and Santa Cruz. In-

fralittoral of Baía da Folga and Carapa-

cho) (AZEVEDO AND MARTINS, 1989: 69),

Faial (Monte da Guia) (AZEVEDO, 1990:

59), Flores (Santa Cruz) (AZEVEDO AND

GOFAS, 1990: 85), Sao Miguel (Ponta da

Galera; Queimada, Água d'Alto; Mostei-

ros; Calheta, Ponta Delgada; Ilhéu de

Vila Franca) (BULLOCK ET AL., 1990: 43,

45), Sáo Miguel (Caloura, -4m; Vila

Franca do Campo, -24m; Ilhéu de Vila

Franca do Campo; Morro das Capelas, -

29m (“Biacores” 1971 expedition); Cape-

las, intertidal; Lagoa, intertidal; Calheta,

Ponta Delgada, 0 to -1m; Ponta da Ga-

lera, -13 to -18m), Faial (Horta, -3m, -

20m), Terceira (Porto Martins; Praia da

Vitória; Pedra Furada-Angra do Hero-

ísmo); Flores (Santa Cruz, intertidal)

(GOFAs, 1990: 116), Sáo Miguel (medio-

littoral of Caloura. Infralittoral of Ca-

loura and Ribeirinha) (AZEVEDO, 199la:

22), Santa Maria (Vila do Porto, llhéu da

Vila, Ponta do Marváo) (AZEVEDO,

1991b: 44), Sáo Miguel (Ilhéu de Vila

Franca) (BULLOCK, 1995: 16; KNUDSEN,

1995: 142), Pico (intertidal) (ÁviLa, 1996:

27) and Formigas islets (ÁVILA AND AZE-

VEDO, 1997: 326).

DBUA 173, 188, 266, 273, 274, 281,

332, 338, 340, 341,346, 390/1392,1999%

355442, 443, 445, 446, 449, 451, 452, 462,

470, 471, 475, 476, 486, 492, 493, 496, 499,

500, 556, 571, 574, 579, 657, 660,661, 662,

665, 666, 667, 670, 715, 719.

MCM(HN) 1, 75.

Comments: from low-tide level to -

20m. This species has a quite variable

ÁviLa: The shallow-water Rissoidae of the Azores and some aspects of their ecology

color pattern (pers. obs.). The diameter

of the protoconch (340 ym, KNUDSEN,

1995: 142, fig. 3B), seems to indicate that

M unifasciata has non-planktotrophic

development (KNUDSEN, 1995). Manzo-

nia crassa, a species that exists on the

Portuguese mainland (e.g.: Berlenga, see

BURrNAy, 1986: 27; BULLOCK ET AL., 1990),

Rissoa costata, reported from the Canary

Islands (MAC ANDREW, 1852: 5) and

Alvania costata, reported from Madeira

by NOBRE (1937: 45) have also been

reported from the Azores. 1 believe that

all of them were misidentified with

Manzonia unifasciata, which is endemic

to the Azores and is the only representa-

tive of this genus in the Azorean Archi-

pelago.

Dimensions: 2.5 mm long, 1.2 mm

wide.

Geographic distribution: endemic to

the Azores.

Onoba Adams H. and A,., 1854

Onoba moreleti Dautzenberg, 1889

References to the Azores:

Onoba moreleti Dautzenberg, 1889: 52; Moolenbeek and Hoenselaar, 1987: 154; Ávila et al. in press.

Occurrence: Pico, Faial and Flores

(ÁVILA ET AL., 1998: 498), Faial, Horta

bay (Stn. 193) (-20m) (MOOLENBEEK AND

HOENSELAAR, 1987: 154), Sáo Miguel

(ÁVILA ET AL., in press).

DBUA 181, 410, 411, 500, 556, 666,

748.

Comments: this is a rare species of the

Azorean littoral. AARTSEN, MENKHORST

AND GITTENBERGER (1984) were surpri-

sed to find specimens of Onoba moreleti

at the Bay of Algeciras (Southern Spain),

but this species was later described as

Onoba josae by MOOLENBEEK AND HOEN-

SELAAR (1987). Its presence was also

reported from Graciosa, Canary Islands

(collection M. C. Fehr-de Wal) by

AARTSEN ET AL. (1984) but, once again, it

was a different species, described as

Onoba manzoniana by ROLÁN (1987).

According to MOOLENBEEK AND FABER

(1987) this species is Manzonia manzo-

niana (Rolán, 1987). Formerly thought to

be restricted to the western and central

groups of islands of the Azores, recent

sorting of material collected at Sáo

Miguel island (DBUA 748 - Capelas,

north coast, 14m depth) has revealed

that O. moreleti also occur in the eastern

group.

Dimensions: 2.6mm long, 1.3mm

wide.

Geographic distribution: endemic to

the Azores.

Rissoa (Fréminville, ms.) Desmarest, 1814

Rissoa guerini Récluz, 1843

References to the Azores:

Cingula costulata Alder, 1844: Chapman, 1955: 803.

Occurrence: Faial (Feteira, mid tide)

(CHAPMAN, 1955: 803). ;

Comments: a dubious record. This

species has not since been cite from the

Azores yet, although its distribution

ranges from the British isles to the Cana-

ries (FRETTER AND GRAHAM, 1978; POPPE

AND GOTO, 1991). Probably, Chapman

misidentified specimens of Rissoa guer-

nei Dautzenberg, 1889 for his species

Cingula costulata.

Dimensions: 6.0 mm long, 3.0 mm

wide (FRETTER AND GRAHAM, 1978).

Geographic distribution: British isles to

Portugal and the Canaries (FRETTER AND

GRAHAM, 1978; POPPE AND GOTO, 1991).

Iberus, 18 (2), 2000

Rissoa guernei Dautzenberg, 1889

References to the Azores:

Rissoa guernei Dautzenberg, 1889: 47-48, pl. 3, figs. la, b; Azevedo and Gofas, 1990: 85; Bullock

et al., 1990: 45; Gofas, 1990: 100; Azevedo, 1991a: 21; 1991b: 44; Bullock, 1995: 16; Knudsen,

1995: 140; Ávila, 1996: 27; Ávila and Azevedo, 1996: 106.

Rissoa obesula Dautzenberg, 1889: 48, pl. 3, figs. 2a, b.

Rissoa jousseaumei Dautzenberg and Fischer, 1896: 60-61, pl. 19, fig. 9 fide Gofas, 1990, p. 99.

Moniziella moniziana azorica Nordsieck, 1972: 173, pl. R V, fig. 28.

Occurrence: Sao Miguel and Faial (-15

to -20 m) (DAUTZENBERG, 1889: 47-48), Pico

(-1287m) (DAUTZENBERG, 1889: 48, pl. 3,

figs. 2a, b), Sao Miguel (-1385m) (DAuT-

ZENBERG AND FISCHER, 1896: 60-61, pl. 19,

fig. 9), Sao Miguel (Ponta Delgada) (NORrD-

SIECK, 1972: 173, pl. R V, fig. 28), Graciosa

(mediolittoral of Porto Afonso; infralitto-

ral of Baía da Folga) (AZEVEDO AND MAR-

TINS, 1989: 69), Pico (Lajes do Pico) (AZE-

VEDO, 1990: 59), Flores (Faja Grande; Santa

Cruz) (AZEVEDO AND GOFAS, 1990: 85), Sao

Miguel (Ponta da Galera; Queimada, Água

d'Alto; Mosteiros; Calheta, Ponta Delgada;

Théu de Vila Franca; Porto do Ilhéu, Vila

Franca do Campo) (BULLOCK ET AL.,1990:

43, 45), Flores (infralittoral of Faja Grande

and Piscina of Ponta Delgada) (NETO AND

AZEVEDO, 1990: 96, 98), Sáo Miguel (Vila

Franca (“Biacores” 1971 expedition); Ponta

Delgada, (-10 to -20m); Ponta da Galera

(intertidal); Capelas (intertidal); Vila Franca

(0 to -5 m); Mlhéu de Vila Franca (0 to -1

m); Calheta, Ponta Delgada (intertidal);

Ponta da Pirámide (-13m) (GOFas, 1990:

100), Sao Miguel (mediolittoral of Caloura;

infralittoral of Caloura and Ribeirinha)

(AZEVEDO, 1991la: 21), Santa Maria (Vila

do Porto, Ponta da Malbusca, Ilhéu da Vila,

Ponta do Marváo) (AZEVEDO, 1991b: 44),

Sáo Miguel (Ilhéu de Vila Franca) (Bu-

LLOCK, 1995: 16; KNUDSEN, 1995: 140), Pico

(intertidal) (ÁviLa, 1996: 27) and Pico

(ÁVILA AND AZEVEDO, 1996: 106).

DBUA 188, 190, 193, 195, 220, 240, 274,

281, 442, 443, 448, 451, 452, 459, 460, 462,

468, 470, 471, 472, 473, 475, 492, 493, 496,

499, 500, 551, 554, 556, 565, 566, 568, 570,

571, 574, 579, 632, 661, 662, 666, 667, 719.

Comments: feeds on detritus and on

epiphytics algae (GrRAHAM, 1988). Accor-

ding to GOFAS (1990) this species is

sexual dimorphic. It occurs from the

low-tide level to -8m. PONDER (1985)

states that the genus Rissoa has pelagic

larvae, being restricted to the Mediterra-

nean and north-eastern Atlantic. Rissoa

guernei however, is a direct development

species and one may hypothesize that

the ancestral of this species probably

lost its planktotrophic veliger larvae

after colonizing the Azores.

Dimensions: up to 2.3 mm long, 1.3

mm wide.

Geographic distribution: endemic to the

Azores, even though it is closely related

to Macaronesian/European species

(GOFAS, 1990).

Setia H and A. Adams, 1852

Setia sp.

References to the Azores:

Setia sp. Azevedo and Gofas, 1990: 85; Ávila et al., 1998: 496.

Occurrence: Flores (Santa Cruz, -20m)

(AZEVEDO AND GOFAS, 1990: 85), Pico

and Flores (ÁVILA ET AL., 1998: 496).

DBUA 274, 276, 277, 281, 446, 449,

478, 496, 499, 662.

Comments: the small dimensions of

this species has probably led to its being

overlook in samples. The sorting of

samples with a mesh size of 0.5mm will

help to clarify its zonation and its geo-

graphical distribution.

Dimensions: 1.0mm long, 0.8mm wide.

Geographic distribution: Flores, Pico

and Sao Miguel.

ÁVILA: The shallow-water Rissoidae of the Azores and some aspects of their ecology

Setia pulcherrima (Jeffreys, 1848)

References to the Azores:

Cingula pulcherrima (Jeffreys, 1848): Bullock et al., 1990: 45; Knudsen, 1995: 143-144.

Occurrence: Sáo Miguel (Ponta da

Galera; Queimada, Água d'Alto; Mostei-

ros; Calheta, Ponta Delgada; Ilhéu de Vila

Franca; Porto do Ilhéu, Vila Franca) (BUL-

LOCK ET AL., 1990: 45), Sáo Miguel (Ilhéu

de Vila Franca) (KNUDSEN, 1995: 143-144).

Comments: this record needs to be

confirmed. This species is reported to

live on rocky shores, among fine weeds

at the low tide (FRETTER AND GRAHAM,

1978). However in all the samples

collected from such places in the Azores,

I have never found this species. It might

be a misidentification for Setia subvari-

cosa Gofas, 1990.

Dimensions: 1.2 mm long, 0.8 mm

wide (FRETTER AND GRAHAM, 1978).

Geographic distribution: Azores (?).

North to the Channel islands (FRETTER

AND GRAHAM, 1978).

Setia quisquiliarum Watson, 1886

References to the Azores:

Setia quisquiliarum Watson, 1886: Dautzenberg, 1889: 53.

“Rissoa” quisquiliarum Watson, 1886: Gofas, 1990: 103.

Occurrence: Sao Miguel (DAUTZEN-

BERG, 1889: 53). Off Faial (38? 38' N, 28*

28" 30 W, in 730-910 m), Terceira

(GOFas, 1990).

Comments: GOFAS (1990) states that

this species is restricted to the central

group of islands, but DAUTZENBERG

(1889) quotes it from Sáo Miguel, in the

eastern group.

Dimensions: 1.4mm long, 0.9mm wide.

Geographic distribution: Faial, Terceira

and Sao Miguel.

Setia subvaricosa Gofas, 1989

References to the Azores:

Setia abjecta Watson, 1873: Dautzenberg, 1889: 52.

Setia subvaricosa Gofas, 1989: Azevedo, 1990: 58; Gofas, 1990: 102-104; Ávila, 1996: 27; Ávila and

Azevedo, 1996: 106; Ávila and Azevedo, 1997: 326; Ávila et al., 1998: 496.

Occurrence: Faial (-15 a -20 m)

(DAUTZENBERG, 1889: 52), Faial (Monte

da Guia) (AZEVEDO, 1990: 58), Sáo

Miguel (Ilhéu de Vila Franca, intertidal;

Capelas, intertidal; Feteiras, -15 m;

Lagoa, -10 to -22 m; Ponta da Galera,

intertidal; Ponta da Pirámide, -13 m;

Calheta, Ponta Delgada, intertidal), Ter-

ceira (Praia da Vitória, Pedra Furada -

Angra do Heroísmo), Flores (Santa

Cruz, -20 m) (GOFAs, 1990: 102-103), Pico

(intertidal) (ÁviLA, 1996: 27), Pico

(ÁVILA AND AZEVEDO, 1996: 106), Formi-

gas islets (ÁVILA AND AZEVEDO, 1997:

326), Flores (ÁVILA ET AL., 1998: 496).

DBUA 176, 188, 193, 195, 223, 274,

281, 332, 335, 336, 338, 343, 345, 350, 352,

355, 447, 451, 462, 465, 467, 471, 481, 496,

499, 500, 545, 557, 564, 571, 574, 660, 662,

666.

Comments: this is an uncommon

species. The outer lip of the adult shell

of S. subvaricosa is thicker than that in

the other species of Setia from the

Mediterranean and the Atlantic

(GOFAs, 1990).

Dimensions: up to 1.4 mm long, 0.8

mm wide.

Geographic distribution: endemic to

the Azores.

Iberus, 18 (2), 2000

Table I. Distribution of the Rissoidae on the islands and islets of the Azorean Archipelago.

Tabla 1. Distribución de los Rissoidae en las islas e islotes del archipiélago de las Azores.

western group central group eastern group

Flores Pico Súo Jorge Faial Graciosa Terceira Súo Miguel Santa Maria Formigas

Alvania abstersa 1 | | 1

Alvania angioyi l 1 1 1 1 1

Alvania cancellata | l 1 1 1 1 |

Alvania formicarum

Alvania internodula

Alvania mediolittoralis |

Alvania poucheti 1

Alvania sleursi 1

Botryphallus ovummuscae 1

Cingula trifasciata l

Crisilla postrema

Manzonia unifasciata

Onoba moreleti

Rissoa guernei

Setia quisquiliarum

Setia subvaricosa 1 1

Setia sp. 1 l

Total number of taxa 13 14 4 13 6 12 15 8 10

— al ll o a A

A o a al al o o A A

a dd a o a al al al a a

—

Table IL. Morphometry of the Rissoidae of the Azores. 4Wp: number of protoconch whorls; 4Wt:

number of teleoconch whorls; Di p: diameter of the protoconch; 1 protoconch 1; II: protoconch 2;

L: total length of the shell; W: total breadth of the shell (based on own data; WATSON, 1886;

GOFAS, 1990; KNUDSEN, 1995; HOENSELAAR AND GOUD, 1998).

Tabla 1. Morfometría de los Rissoidae de las Azores. 4Wp: número de vueltas de la protoconcha; 4W!t:

número de vueltas de la teleoconcha; Di p: diámetro de la protoconcha; I protoconcha 1; II: protoconcha

2; L: longitud total de la concha; W: anchura total de la concha (basado en datos propios; WATSON,

1886; GOFAS, 1990; KNUDSEN, 1995; HOENSELAAR AND GOUD, 1998).

RISSOIDAE +Wp FWt Di p (m) L (mm) W (mm)

Alvania abstersa Van der Linden and Van Aartsen, 1994 1,5 3.5 300-400 DINSA

Alvania angioyi Van Aartsen, 1982 1.5 3.25 -3.75 283.3 -292.3 1.2-1.8 0.7-1.1

Alvania cancellata (Da Costa, 1778) El 3,5 ENLAZA

11: 1.5 Il: 375.0 — 440.0

Alvania formicarum Gotas, 1989 1.25 3 366.7 2.4 1.4

Alvania internodula Hoenselaar and Goud, 1998 1.25 3,5 310.0-333.3 20-23 11-13

Alvania mediolittoralis Gotas, 1989 1.25 3.25 -3.75 294.1 - 304.0 22-27 13-15

Alvania poucheti Dautzenberg, 1889 1.25 3.25 363.4 - 383.4 18-22 1.0-13

Alvania sleursi (Amati, 1987) 1.25 3.75-4 358.3 - 400.0 22-25 15-16

Alvania tarsodes (Watson, 1886) ? ? ? 2.2 1.3

Botryphallus ovummuscae (Gofas, 1990) 1.25 3 222.2 11-13 0.6-07

Cingula trifasciata (Adams, 1798) 20-25 4 500.0 IMAN

Crisilla postrema (Gotas, 1990) 1.25 3 ? 1.4-1.5 0.8-0.9

Manzonia unifasciata (Dautzenberg, 1889, 1.25 4 304.3 - 347.8 20-25 1.0-1.2

Onoba moreleti Dauizenberg, 1889 1.25 3 322.7 - 333.3 1.9-2.6 09-13

Rissoa guernei Dautzenberg, 1889 1.25 4 258.8 -281.3 19-23 11-13

Setia sp. 0.6 2.0-2.5 200.0 - 213.0 0.8-1.0 0.6-0.8

Setia quisquiliarum Watson, 1886 ? 3 ? 1.4 0.

Setia subvaricosa Gotas, 1990 1.25 3 238.5 1.1-1.4 0.7-0:8

ÁVILA: The shallow-water Rissoidae of the Azores and some aspects of their ecology

40 50 60 70 80

Formigas

Faial

Flores

Pico

Sáo Miguel

Terceira

Graciosa

Santa Maria

Sáo Jorge

90 100

Figure 4. Bray-Curtis/non transformed presence/absence data / UPGMA for the Rissoidae of the

Azores.

Figura 4. Análisis Bray-Curtis/no transformado de presencialausencia data / UPGMA de los Rissoidae

de las Azores.

RESULTS

At the present status of our know-

ledge, 23 taxa of Rissoidae are given to

the Azores: 12 are endemic species, three

are restricted to the Azores and

Madeira /Selvagens archipelago, two

have a wider distribution, one is an uni-

dentified species of Setia and there are

five records that were not confirmed by

this study. The presence / absence of Ris-

soidae species on the islands of the

Azores, based on the literature and new

data, is shown in Table I. Classification

techniques used for this table, resulted in

Figure 4. Sáo Miguel, Pico and Flores,

clustered at more than the 95% similarity

level, form a consistent group, to which

Faial (92%) and Terceira (85%) are also

joined. Formigas, clustered at 65%,

seems to be different from the first group

of islands in terms of the Rissoidae

species. Sáo Jorge is the last island to

cluster, at only the 44% similarity level.

Morphometry: The largest Rissoidae

present on Azorean littoral are Alvania

cancellata (3.7 x 2.4 mm) and Cingula tri-

fasciata (3.9 x 2.1 mm), whereas the sma-

llest are Setia sp. (1.0 x 0.8 mm), Botryp-

hallus ovummuscae (1.3 x 0.7 mm) and

Setiía subvaricosa (1.4 x 0.8 mm). Almost

all protoconchs have 1 */4 whorls,

Alvania cancellata being the exception

with 2 */2 whorls. The smallest proto-

conchs belong to Setia sp. and S. subvari-

cosa (200 to 238,5 um). Alvania cancellata,

with a multispiral protoconch, has the

largest (protoconch I= 120 yum; proto-

conch Il= 440 um). The number of the

teleoconch whorls range from 3 to 4 in

all rissoids (Table ID).

Zonation: Nine species of Rissoidae

were found at Sáo Vicente, Capelas, on

the north coast of Sáo Miguel, Azores. A

total of 1,564 specimens were counted,

on the 24 collected quadrates of 50 x 50

cm, Manzonía unifasciata Dautzenberg,

1889 being the most abundant with a

total of 631 individuals, whilst Setia sp.

and S. subvaricosa Gofas, 1990 uncom-

mon species, (11 and 13 specimens, res-

pectively) (Table ID).

The zonation of the Rissoidae at Sáo

Vicente, Capelas, seems to indicate the

Iberus, 18 (2), 2000

Table IM. Rissoidae collected in July 1996 at Sáo Vicente-Capelas, north coast of Sao Miguel (all

specimens larger than 1mm).

Tabla III. Rissoidae recogidos en julio de 1996 en Sáo Vicente-Capelas, costa norte de Sáo Miguel (todos

los especimenes mayores de Imm).

Depth (m) 3.5 3.5 EL

A. angioyi 0) 6 27 27

A. cancellata | 0 1 |

A. poucheti 2 4 3 3

A. sleursi 16 19 28 18

C. postrema 0 0 8 0

M. unifasciata 10 11 28 87

R. guernei 16 14 4] 16

S. subvaricosa 1 0 2 2

Setia sp. 1 1 1 2

TOTAL 53 55 139 156

5.1

197

existence of common species at shallow

depths (from low tide level to -10 m),

such as Rissoa guernei, Manzonta unifas-

ciata and Alvania angioyi, and species

more abundant from 15 m down, such

as Alvania sleursi, A. cancellata and A.

poucheti (Figs. 5 and 6).

By clustering the stations, at the 60%

similarity level, two groups appear. The

first one, with the highest number of sta-

tions and with the exception of replicates

20 and 21 (22 m depth), contains stations

in shallow/medium depths. The second

group, with the exception of stations 10

(12 m depth) and 14 (13.6 m depth) are

all medium / high depths (Fig. 7).

DISCUSSION

It seems evident that there is some

island to island endemism, as suspected

by GOFAS (1990), Alvania formicarum and

A. internodula being restricted to the eastern

group of islands (Sáo Miguel, Santa Maria

and Formigas islets). Pico, Faial, Flores

and Sáo Miguel, the best studied islands,

are almost identical in the composition of

the Rissoidae, with the exceptions of Setia

quisquiliarum (not found yet at Flores and

Pico), Alvania abstersa (not found at Flores

and Faial) and Setía sp. (not found at Faial).

There is a clear distinction between the

Rissoidae of Formigas islets and the remai-

5.1 8 8 8 12 127812

4 6 15 16 0 1 9

1 8 o) l 1 3 4

0 0 Y) 3 0 | 1

23 11 15 20 6 5 11

0 0 0 4 0 0 0

16 51 ISSO 0 5 13

11 8 7 9 0 0 1

0 0 2 0 0 0 2

0 0 0 1 0 l 2

35 SA O NAS y IAS

ning islands of the Azores. In fact, A. inter-

nodula is restricted to these islets and A.

mediolittoralis and Rissoa guernei, common

species in the other islands, do not occur

at Formigas. The importance of the For-

migas islets as a Nature Reserve is there-

fore reinforced by the results of this study.

Santa Maria, Sao Jorge and Graciosa

must be considered as outsiders in this

biogeographic puzzle, as long as the

number of samples and the quality of

them is not increased (Table I and Fig.

4). As for Setia sp., it may have been

overlooked in some samples because of

1ts small size.

The abundance of the Rissoidae in

the littoral of the Azores seems to be

variable. AZEVEDO (1991) found that Cri-

silla postrema (= Alvania postrema) (Ímedio-

littoral) and Rissoa guernel (infralittoral)

were the most abundant species associa-

ted with macroalgae in two sites at Sáo

Miguel island (Caloura / south coast and

Ribeirinha /north coast). At Lajes do Pico

(rocky intertidal conditions), the most

abundant species is Cingula trifasciata,

which may reach densities of 32,500 spe-

cimens / m? (ÁviLa, 1998). In this study,

Manzonia unifasciata and Alvania sleursi

are revealed to be the most abundant

species, both in the infralittoral. Only

long term and seasonal studies will

answer this apparent discrepancy

between mine and Azevedo's data.

ÁVILA: The shallow-water Rissoidae of the Azores and some aspects of their ecology

Table HI. Continuación.

Tabla UI. Continuation.

13.6 136 13.6 163 16.3 16.3

0 1 0 4 0 59

4 2 2 3 0 2

2 1 0 2 0 11

20 A7 8 16 6 11

0 0 0 0 0 5

18 8 3 14 2 2

0 0 0 1 0 0

0 0 0 0 0 0

Ad 59 13 41 8 39

With the sole exception of Alvania

cancellata none of the other species of

Rissoidae in the Azores has a long plank-

totrophic larval development, because

their protoconchs are bigger than about

200 um (VERDUIN, 1982; 1985) (Table ID.

However, studies on other marine Proso-

branchs (e.2. Turridae) have demonstra-

ted that a paucispiral protoconch (as

seen in the majority of the Azorean Ris-

soidae) must not be interpreted as evi-

dence for lacking a planktonic phase

(SHIMEK, 1986; BOUCHET, 1990). Nevert-

heless, if there is a planktonic phase, it

must be of small duration, but of high

importance to the dispersal of the

species within the archipelago.

The similarity between the shells of

Alvania cancellata and A. sleursi was

pointed out by Goras (1990), who

thought the latter species could have

speciated from the former, by losing the

planktotrophic phase. On the other

hand, Alvania abstersa, A. formicarum and

A. mediolittoralis are so similar in their

protoconchs as well as in their teleo-

conchs, that we may hypothesize their

relation with a common ancestor.

MAC ANDREW (1854: 49), stated that

the marine molluscs of the Azores,

Madeira and Canary Islands were

«closely related to that of the old conti-

nent, notwithstanding that the prevai-

ling set of currents is from America».

22 22 26.8 26.8 26.8 TOTAL

3 0 0 0 3 160

19 6 2 1 1 71

16 2 0 1 0 59

106 13 4 4 Y A55

0 0 0 0 0 18

19 3 0 0 1 631

0 0 0 0 0 143

0 0 0 0 0 13

0 0 0 0 0 11

163 24 o o) 12 1,564

The surface currents in the Northern

Atlantic, especially the Gulf Stream,

have been studied in detail during this

century (ISELIN, 1936; GOULD, 1985;

FIALHO AND BARROS, 1988; KLEINE AND

SIEDLER, 1989; ALVES, 1990; 1992;). All

studies indicate that the surface currents

are mainly from West to the East, that is,

from America to Europe.

The larvae of Alvania formicarum pro-

bably did not reach the islands of the

central and western groups because of the

main direction of surface currents in the

Azores. The same may be true for A. inter-

nodula, but the scarcity of data on this last

species, does not allow for a stronger con-

clusion. Additional samples must be taken

at Formigas, in order to determine the

zonation of A. internodula. If it becomes

apparent that it usually occurs at depths

of about 45 m (as is the case at Formigas

islets), this species may be common on the

other islands of the Azores, but not found

yet, because no representative samples

have been collected by me at depths

greater than 30m. However, this species

was not found in any of the CANCAP

samples (ranging from 33 to 47 m depth),

so we have some evidence that it may be

restricted to Formigas.

A non-planktotrophic species may

be distributed over a large area if there

is another plausible means of transport.

It is likely that those species of Rissoidae

Iberus, 18 (2), 2000

3.5 5.1 8 12

US SS SI

Rissoa guernel

26.8 Depth (m)

13.6 16.3 22

14 ANgioyl

0) 5.1 8 12

13.6 16.3 22

26.8 Depth (m)

Figure 5. Common Rissoidae species collected at shallow depths (from low tide level to -10m)

collected at Sáo Vicente, Capelas, Sio Miguel, Azores (O: maximum; 0: average; U: minimum

values).

Figura 5. Especies de Rissoidae comunes en aguas someras (del nivel de bajamar hasta -10m) recogidas

en Sáo Vicente, Capelas, Sáo Miguel, Azores (O: máximo; O: media; U: mínimo).

living in the first few meters of water

(e.g. Alvania angioyi, Manzonia unifasciata

and Rissoa guernei) may drift from island

to island on “rafts” of algae provided by

heavy seas breaking into the shore.

There are 231 confirmed species of

shallow-water molluscs on the littoral of

the Azores (ÁvILA, 2000). Only 16 spe-

cies (6.9%) are amphi-Atlantic species, in

contrast to the 181 species (78.4%) shared

with the western Mediterranean, or the

147 species (63.6%) shared with Madeira

(with the Desertas and Selvagens). Main-

land Portugal and Canary Islands, share

144 (62,3%) and 137 species (59,3%), res-

pectively, with the Azores, whereas Saint

Helena (5,2%) and Ascension island

(5,6%) share only a small number of spe-

cies. A total of 112 species (48,5%) occurs

simultaneously in the Azores, Madeira

and the Canary Islands, and 53 species

(22,9%) occur in all the Macaronesian Ar-

ÁVILA: The shallow-water Rissoidae of the Azores and some aspects of their ecology

Alvania sleursi

100

30) 5.1 8 12

13.6 16.3 22

Dd) P= == === === === ===

Alvania cancellata

Do) el 8 12

13.6 16.3 22

26.8 Depth (m)

Figure 6. Common Rissoidae species collected at high depths (-15 to -30m) collected at Sáo

Vicente, Capelas, Sao Miguel, Azores (O: maximum; 0: average; U: minimum values).

Figura 6. Especies de Rissoidae comunes en aguas profundas(-15 a -30 m) recogidas en Sáo Vicente,

Capelas, Sáo Miguel, Azores (O: máximo; O: media; Ú: mínimo).

chipelagos (ÁVILA, 2000). Of the 231 re-

ported species, 19 are endemic (8,2%)

and of these, 13 (5,6%) are endemic Ris-

soidae (ÁVILA, 2000). The dominance of

the Rissoidae in the littoral fauna of the

Azores, is one piece of evidence that

supports its higher similarity with that

of Europe (GOFASs, 1990).

An understanding of the coloniza-

tion and subsequent speciation that has

occurred in the molluscan fauna of the

Azores, with special emphasis on the

Rissoidae, clearly merits further rese-

arch. Samples of plankton should be

taken in the three groups of islands, in

order to evaluate the dispersal capabili-

ties of the endemic Rissoidae, especially

the most problematic species (Alvania

formicarum, A. internodula, Onoba moreleti

and the undescribed Setia). Deeper

samples should also be collected in

order to respond to the questions raised

by the so far apparent restricted range of

Alvania internodula.

Iberus, 18 (2), 2000

102 RO PO A

de Depth (m) Quadrats

20 35) 1-3

11 al 4-6

14 8.0 7-9

15 12.0 10-12

2 13.6 13-15

SS 16.3 16-18

19 22.0 19-21

5 26.8 22-24

$ + + + + + 4 2

40 50 60 70 80 90 100

Figure 7. Rissoidae at Sáo Vicente, Capelas, Sio Miguel, Azores. Double Square Root/Bray-

Curtis/UPGMA.

Figura 7. Rissoidae de Sáo Vicente, Capelas, Sáo Miguel, Azores. Doble Raíz/Bray-Curtis/UPGMA.

ACKNOWLEDGEMENTS with the SEM photos and to Robert

Moolenbeek, by the generous help with

I am grateful to Frias Martins (Uni- bibliography and comments on the

versity of the Azores) for the support manuscript. I specially acknowledge the

given during the last three years and to useful comments of Winston Ponder

Brian Morton (University of Hong and an anonymous referee.

Kong), whose comments have greatly This study was supported by grants

improved a first draft of this paper. l am PRAXIS XXI BM/20521/99, PRAXIS

also indebted to Jorge Medeiros (Uni- XXI/BIC/2788/96 and PRAXIS/2/2.1/

versity of the Azores) for his assistance BIA/169/94.

(Right page) Figure 8. Alvania internodula Hoenselaar and Goud, 1998. A: shell (DBUA 338/19-

3) 2.3 x 1.3 mm; B, C: protoconch (lateral and frontal view); D: microsculpture of protoconch; E:

microsculpture of 1* post-larval whorl; F: microsculpture of body whorl; G: aperture of the shell;

H: shell (DBUA 338/19-4), 2.0 x 1.1 mms; I: shell (DBUA 338/31-4), 1.6 x 0.9mm,; J: shell of

juvenile (DBUA 338/31-2), 1.1 x 0.9 mm.

(Página derecha) Figura 8. Alvania internodula Hoenselaar y Goud, 1998. A: concha (DBUA

338/19-3) 2,3 x 1,3 mm; B, C: protoconcha (vistas lateral y frontal); D: microescultura de la protocon-

cha; E: microescultura de la primera vuelta postlarvaria; EF: microesculptura de la vuelta del cuerpo; G:

apertura de la concha; H: shell (DBUA 338/19-4), 2,0 x 1,1mm, I: concha (DBUA 338/31-4), 1,6 x

0,9mm; J: concha de juvenil (DBUA 338/31-2), 1,1 x 0,9mm.

ÁVILA: The shallow-water Rissoidae of the Azores and some aspects of their ecology

Iberus, 18 (2), 2000

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Iberus, 18 (2): 77-97, 2000

A peculiar high-tidal molluscan assemblage from a

Madeiran boulder beach

Una peculiar comunidad de moluscos del nivel superior de la marea

en una playa de cantos rodados de Madeira

Emilio ROLÁN* and José TEMPLADO**

Recibido el 7-VILI-2000. Aceptado el 23-X-2000

ABSTRACT

The molluscs living at the higher intertidal level in a protected area of boulders in Madeira

are described. Fourteen species of molluscs were found in this peculiar habitat, the gastro-

pods Littorina striata, Melaraphe neritoides, Assiminea cf. grayana, Paludinella littorina, Caecum

armoricum, Caecum clarkii, Truncatella subcylindrica, Botryphallus epidauricus, Odostomia

microeques (parasitizing the former species), Ovatella aequalis, Auriculinella bidentata,

Pedipes pedipes, Pseudomelampus exiguus, and the bivalve Lasaea rubra. Their abundance

and distribution assemblages in this habitat are noted and they are compared with the molluscs

found in similar habitats in other areas of the Northeastern Atlantic and Mediterranean.

RESUMEN

Se describen los moluscos hallados en el nivel superior de la marea en una zona de bloques

y cantos rodados de la isla de Madeira. Se hallaron catorce especies de moluscos en este

hábitat tan peculiar, los gasterópodos Littorina striata, Melaraphe neritoides, Assiminea cf.

grayana, Paludinella littorina, Caecum armoricum, Caecum clarkii, Truncatella subcylin-

drica, Botryphallus epidauricus, Odostomia microeques (parasitando la especie anterior),

Ovatella aequalis, Auriculinella bidentata, Pedipes pedipes, Pseudomelampus exiguus y el

bivalvo Lasaea rubra. Se aportan datos sobre la abundancia de todas estas especies y su

distribución dentro de este hábitat. Por último, se compara esta comunidad de moluscos con

las halladas en hábitats similares en otras zonas del Atlántico nordeste y del Mediterráneo.

KEY WORDS: Madeira, boulder beach, high-tidal molluscan assemblage, Littorinidae, Assimineidae, Caecidae,

Truncatellidae, Rissoidae, Pyramidellidae, Ellobiidae, Kelliidae.

PALABRAS CLAVE: Madeira, playa de cantos rodados, comunidad de moluscos supralitoral, Littorinidae,

Assimineidae, Caecidae, Truncatellidae, Rossoidae, Pyramidellidae, Ellobiidae, Kelliidae.

INTRODUCTION

Faunas associated with boulder

beachs were described by MORTON

(1975) in New Zealand. He pointed out

that the molluscs that lives at the high-

* Cánovas del Castillo 22, 36202 Vigo, Spain

tidal level in such places form a distinc-

tive ecological grouping. These mollus-

can communities are characterised by

species of both marine and terrestrial

** Museo Nacional de Ciencias Naturales (CSIC), José Gutiérrez Abascal 2, 28006 Madrid, Spain

Dd,

Iberus, 18 (2), 2000

Figure 1. Schematic representation of the sampling site. 1: the upper level formed by cobbles of

about 15-25 cm, occasionally receiving sunlight; 2: among and under them, there was another layer

of scarcely smaller stones of about 9-15 cm, never exposed to sunlight; 3: other small stones without

any contact with sand; 4: smaller ones mixed with some coarse sand formed a layer of about 6-8 cm

thick; 5: bed of gravel and coarse sand mixed with very small stones; A: high tide level.

Figura 1. Esquema del lugar de muestreo. 1: nivel superior formado por cantos rodados de unos 15-25

cm, que reciben ocasionalmente la luz del sol; 2: entre ellos y por debajo hay otro nivel con piedras lige-

ramente más pequeñas (9-15 cm), que nunca están expuestas a la luz del sol; 3: otras piedras más peque-

ñas sin contacto alguno con arena; 4: otras piedras pequeñas mezcladas con arena compacta formando

un estrato de unos 6-8 cm de espesor; 5: nivel de grava y arena compactada con pequeñas piedras; A:

nivel de la marea alta.

groups, which form a mixed assem-

blage. According to this author espe-

cially common here are species of the

caenogastropods families Assimineidae

and Caecidae, and primitive pulmona-

tes of the family Ellobiidae. PONDER

(1990) studied a close related habitat in

the Strait of Gibraltar, but in a some-

what lower level. He pointed out that

“these habitats, long neglected by bio-

logist, clearly deserve much closer

attention”.

During a short visit to Madeira

Island in Octuber of 1993 we had the

opportunity to find a surprisingly

diverse molluscan assemblage, quite

similar to that described by MORTON

(1975), in such seemingly inhospitable

habitat. These molluscs and their abun-

dance and distribution within this

habitat are here described.

The small molluscs of Madeira were

firstly studied by MANZONI (1868a,

1968b), WATSON (1873, 1891, 1898), and

in more recent times by NORDSIECK AND

GARCÍA-TALAVERA (1979), VERDUIN

(1984, 1988), MOOLENBEEK AND FABER

(1987), PALAZZI (1988) and MOOLENBEEK

AND HOENSELAAR (1989, 1998), among

others. All these publications are mainly

lists, inventories or description of new

species, and most of them are based on

dead material. In the other hand, the

Ellobids from Madeira were studied

firsly by WOLLASTON (1878) and in

recent times by MARTINS (1995, 1996,

1999).

MATERIAL AND METHODS

The Madeiran coast is very steep,

and exposed to an intense wave action.

In most part of its coast only boulder

beachs are found. In October of 1993 a

small area of about 2 x 2 m was sampled

in Funchal, near the Club Naval. The

sampling site was located at the corner

of a small bay, where wave-rounded

boulders and large cobbles were over a

ROLÁN AND TEMPLADO: High-tidal molluscan assemblage from a Madeiran boulder beach

Table L. Species found in Madeira and their abundance and habitat where they predominated. 1:

upper part of big boulders; 2: amongst and under boulders; 3: under cobbles without any contact

with the sand layer. ; 4: amongst and under small stones in contact with the sand layer. ; 5: in the

sand under cobbles; +: 1 - 10 specimens; ++: 11 - 50 specimens; +++: 51-150 specimens; ++++:

more than 150 specimens.

Tabla I. Especies encontradas en Madeira, su abundancia y nivel en el que eran predominantes. 1:

parte superior de bloques grandes; 2: entre y bajo bloques; 3: bajo guijarros sin ningún contacto

con la capa de arena; 4: entre y bajo pequeñas piedras en contacto con la arena; 5: en la arena bajo

guijarros; +: 1 - 10 especímenes; ++: 11 - 50 especímenes; +++: 51-150 especímenes; ++++: más de

150 especímenes.

Species 1 2 3 4 5

L. striata +

M. neritoides +

A. cf. grayana +++ de

P. littorina + eri

C. clarkii A

C. armoricum Joc

T. subeylindrica + su

B. epidauricus + A

O. microeques ai

O. aequalis + dnd +

A. bidentata se EN

P. pedipes ES

P. exiguus +

L. rubra +

coarse sand and gravel layer, among

rocks (Fig. 1). It was a moderately

stable, shady place, protected against

the direct impact of the waves. The sea-

water was received slowly braked by its

filtration through the gravel. Thus, the

main ecological factors of the small area

sampled were the high humidity, and

permanently low levels of light and

temperature. Some decaying algal

wrack and plant debris can be found

under stones, which constitute the food

for most of the animals inhabiting there.

No macroscopic seaweeds were observ-

ed in this habitat.

Material was collected in the high

tide spring level by direct observation

with frontal lens. Also, some cobbles

were cleaned in a box with sea-water,

and samples of the coarse sand and

gravel placed under boulders and

stones were taken in order to be studied

later under magnification.

The samples obtained cannot be

treated as a valid quantitative estimates.

Therefore, the number of specimens

given might be interpreted as a general

picture of abundance and distribution of

each species.

In order to make comparisons, a

very similar habitat and level was

sampled in other localities, two located

in the Atlantic coast of NW Spain (Cies

Islands, Ría de Vigo, June, 1997, and

Ribadeo, Lugo, June, 1998), and another

in the Mediterranean (Los Escullos,

Almería, SE Spain, September 1996).

Some specimens from the Muséum

Nationale d'Histoire Naturelle of Paris

(MNHN) (loaned by Serge Gofas),

coming from Ceuta, Azores and Canary

Islands were used for comparison.

Voucher material of all the species

recorded has been deposited in the

Museo Nacional de Ciencias Naturales

of Madrid.

JE)

Iberus, 18 (2), 2000

Table II. Species found in similar habitat in Madeira and in several localities of the Spanish coasts

(Ribadeo, Cies Islands, in NW Spain, and Los Escullos, SE Spain).

Tabla 11. Especies presentes en hábitats similares en Madeira y varias localidades del litoral español

(Ribadeo, Islas Cies, en el NO de España, y Los Escullos, SE de España).

Species in NW SPAIN

L. saxatilis

P. littorina

C. trifasciata

T. subcylindrica

Species in SE SPAIN

M. neritoides

P. littorina

C. armoricum

T. subcylindrica

B. epidauricus

Genera Species in MADEIRA

Littorina L. striata

Melaraphe M. neritoides

Assiminea A. cf. grayana

Paludinella P. littorina

Ca C. armoricum

C. clarkii

Cingula

Truncatella T. subcylindrica

Botryphalus B. epidauricus

Odostomia O. microeques

Ovatella O. aequalis

Myosotella

Auriculinella A. bidentata

Pedipes P. pedipes

Pseudomelampus P. exiguus

Lasaea L. rubra

RESULTS

Description of the habitat: We consid-

ered the following levels in the sampling

site (Fig. 1), with independence of the big

stones which were around the place:

1- the upper level was formed by

cobbles of about 15-25 cm, occasionally

receiving sunlight;

2- among and under them» there was

another layer of scarcely smaller stones of

about 9-15 cm, never exposed to sunlight;

some algal debris were deposited here;

3- under these, other small stones

without any contact with sand;

4- amongst and under small stones,

smaller ones mixed with some coarse sand

formed a layer of about 6-8 cm thick.

5- under this layer there was a bed of

gravel and coarse sand mixed with very

small stones.

M. myosotis M. myosotis

A. bidentata A. bidentata

P. exiguus P. exiguus

L. rubra

All these levels were at the higher

intertidal level. Probably, the sea-water

could reach the upper cobbles when the

sea was strong, but normally the waves

arrived to this place very attenuated.

The cobbles and stones under boulders

were always humid and they were not

hardly heated by the sun.

The molluscan species found in this

habitat lived very close one to another

but occupied different levels or niches.

Most of them were found very close

because there were only about 25 cm

from the higher to the lower level.

Species found: The species found in

Madeira and its relative abundance and

location within this habitat are shown in

Table I. The Table II shows the species

found in similar habitat in Madeira, NW

Spain, and SE Spain.

(Right page) Figures 2-8. Assíminea cf. grayana, Madeira. 2: shell; 3-5: protoconch; 6: detail of the

suture; 7-8: radula.

(Página derecha) Figuras 2-8. Assiminea cf. grayana, Madeira. 2: concha; 3-5: protoconcha; 6: detalle

de la sutura; 7-8: rádula.

ROLÁN AND TEMPLADO: High-tidal molluscan assemblage from a Madeiran boulder beach

Iberus, 18 (2), 2000

Family LITTORINIDAE Gray, 1840

Littorina striata King and Broderip, 1832

Material studied: 8 specimens from Madeira.

Habitat: Found in the upper level

amongst big boulders.

Remarks: This species is known from

Azores, Madeira, Canarias, Cabo Verde

and Sáo Tomé islands. Our specimens

were found in the upper part of the

studied area, on the rocks. Most of them

were not adults. Some of them had

tubercles in the upper part of the first

whorls, even in one specimen the tuber-

cles reached the last whorl. This pattern

is frequent in the same species in Cape

Verde Islands. No other differences were

appreciated between Madeiran and

Canarian populations of this species. A

detailed study on patterns of shell varia-

tion in this species along Macaronesia

was done by DE WOLF ET AL. (1998).

Melaraphe neritoides (Linné, 1758)

Material studied: 7 specimens from Madeira, 20 specimens from Almería.

Habitat: Found in the upper level on

the surface of big boulders.

Remarks: The species ranges from

Scandinavia to the Mediterranean. The

shells from Madeira had the same

characteristics that the ones found in

the European mainland coasts popula-

tions.

Family ASSIMINEIDAE H. and A. Adams, 1856

Assiminea cf. grayana Fleming, 1828 (Figs. 2-8, 30)

Material studied: 104 specimens from Madeira.

Description: Shell (Fig. 2) small, solid,

globose-conical, with spire scarcely

pointed. Surface smooth, only with

growth lines and a subsutural spiral

groove (Fig. 6), not clearly appreciated

in first whorls. Suture slowly depressed.

Protoconch (Figs. 3-5) smooth, with

somewhat more than one spiral whorl.

Its nucleus measured 130 um in diame-

ter

Animal white, with short and con-

tractile cephalic tentacles, showing the

eyes close to their tip. The anterior part

of the foot is rounded and bilobulated

(Fig. 30). Black pigmentation was very

constant in the external apical part of

the tentacles. A black or grey spots in

the head, between the tentacles, can be

also present. In some specimens a quite

dark first whorls can be observed by

transparence.

Radula (Figs. 7-8) with a narrow and

elongate rachidian tooth, which presents

a prominent central cusp, with one or

two at each side. There are also three

small cusps near its base at each side,

being less prominent the more basal.

The lateral teeth have a spoon-like form,

(Right page) Figures 9-12. Paludinella littorina, Madeira. 9: shell; 10-11: protoconch; 12: micros-

culpture of the protoconch. Figure 13. Caecum armoricum, apex (Madeira). Figure 14. Caecum

clarki, apex (Madeira).

(Página derecha) Figuras 9-12. Paludinella littorina, Madeira. 9: concha; 10-11: protoconcha; 12:

microescultura de la protoconcha. Figura 13. Caecum armoricum, ápice (Madeira). Figura 14.

Caecum clarki, ápice (Madeira).

ROLÁN AND TEMPLADO: High-tidal molluscan assemblage from a Madeiran boulder beach

Iberus, 18 (2), 2000

with some cusps at their margin. The

marginal internal are very similar to the

lateral, being also spoon-like, somewhat

wider, with smaller and more numerous

cusps.

Habitat: Found living under boulders

in humid parts, without contact with the

sand, but very near to this layer.

Remarks: In a first glance we identi-

fied this species as A. grayana Fleming,

1828. But after a most detailed study we

had some doubts because FRETTER AND

GRAHAM (1978) noted spiral lines in the

protoconch of this taxon, which are not

present in our shells. They referred also

a protoconch of two whorls, but our

shells have only somewhat more than

one (using the method of VERDUIN,

1984). We have tried to examine the pro-

toconch of specimens of populations of

this species recorded in the Ría de Arosa

by CADÉE (1968) and ROLÁN (1987), but

all the shells had the protoconch eroded.

A. grayana ranges from the Atlantic

coast of Europe to the Mediterranean

Sea. Other forms of the genus Assiminea

has been observed in some areas of West

Africa down to Angola. ÁVILA (1998,

2000) used the name Assiíminea eliae

Paladilhe, 1875 for the specimens of the

Azores. Until a detalied review of all

these forms is done, we prefer to keep

the specimens from Madeira under the

current name A. grayana, following the

nomenclature proposed by the

CLEMAN checklist.

Paludinella littorina (delle Chiaje, 1828) (Figs. 9-12, 29)

Material studied: 62 specimens from Madeira; 30 from Almería; 35 from Cies Islands; 15 from

Ribadeo.

Description: Shell (Fig. 9) small,

globose, glossy, semitransparent. Proto-

conch (Figs. 10-12) with a huge nucleus

and one whorl and little more; its

surface is covered by a fine microsculp-

ture of irregular granulations. There is a

distinc boundary with the teleoconch.

Animal milky-white, with cephalic ten-

tacles very short, flat, semitriangular,

with subapical eyes (Fig. 29). A pinkish

area can be observed by transparence

between the tentacles.

Habitat: P. littorina was living under

boulders near or in contact with the

sand layer.

Remarks: FRETTER AND GRAHAM

(1978) noted that P. littorina has a

protoconch of 1 3/4 smooth whorls.

The photograph presented by them

(FRETTER AND GRAHAM, 1978, fig. 130)

show a protoconch that seems to have

less than one whorl. In fact, in our

material there is only a little more than

1/2 whorl after a wide nucleus (using

the method of VERDUIN, 1984). The

surface of the protoconch of our mate-

rial is covered by irregular fine sculp-

ture and was never smooth, as in the

protoconch showed by FRETTER AND

GRAHAM (1978, p. 149, fig. 130).

Perhaps, the shells studied by these

authors had eroded protoconchs or

they were not studied at enough

magnification. We have compared the

Madeiran shells with those from SE

Spain, and their microsculpture (Fig.

12) is similar in both populations. Thus

we consider to be the same species

both Madeiran and Mediterranean

specimens.

This species occurs throughout the

Mediterranean Sea and along the

Eastern Atlantic coast, from Madeira to

the southern coast of the British Isles.

(Right page) Figures 15-16. Truncatella subcylindrica, protoconch, Madeira. Figures 17-20. Botryp-

hallus epidauricus., Madeira. 17: shell; 18: protoconch; 19-20: radula.

(Página derecha) Figuras 15-16. Truncatella subcylindrica, protoconcha, Madeira. Figuras 17-20.

Botryphallus epidauricus., Madeira. 17: concha; 18: protoconcha; 19-20: rádula.

ROLÁN AND TEMPLADO: High-tidal molluscan assemblage from a Madeiran boulder beach

Iberus, 18 (2), 2000

Family CAECIDAE Gray, 1850

Caecum armoricum de Folin, 1869 (Fig. 13)

Caecum incomptum (Monterosato, 1884).

Material studied: 22 specimens from Madeira; 9 specimens from Almería.

Description: A complete description

of the shell of this species can be seen in

the papers of VAN AARTSEN AND HOEN-

SELAAR (1984) and HOEKSEMA AND

SEGERS (1993). The specimens studied

did not differ from these descriptions.

Animal white.

Habitat: Found buried in the sand,

under boulders and stones.

Remarks: PONDER (1990) pointed out

that this species (as C. incomptum) is

very abundant in the intertidal gravel

habitat in the Strait of Gibraltar. Our

specimens are quite similar to those

mentioned by this author. Caecum armo-

ricum occurs throughout the Mediterra-

nean Sea and along the Eastern Atlantic

coast, from Canary Island to Azores and

Northern France. Its distribution and

systematic had been discused by

AARTSEN AND HOENSELAAR (1984) and

HOEKSEMA AND SEGERS (1993).

Caecum clarkiú Carpenter, 1858 (Fig. 14)

Material studied: 1 living specimen from Madeira.

Habitat: Buried in sand under cobbles.

Remarks: C. clarkii was originally

described from Canary Islands and

widespread throughout the European

Atlantic coasts and the Mediteranean

Sea.

Family TRUNCATELLIDAE Gray, 1840

Truncatella subcylindrica (L., 1767) (Figs. 15-16, 28)

Material studied: 39 specimens from Madeira; 109 from Almería; 12 from Vigo; 45 from Ribadeo.

Description: Shells of Madeiran speci-

mens were very similar to those from

the Mediterranean populations, with

numerous axial ribs, but some speci-

mens were smooth. Protoconch smooth

(Figs. 15-16). Animal white. Anterior

end of the foot with two flat enlarge-

ments. Tentacles not very long, cylindri-

cal with the eyes at their bases (Fig. 28).

Habitat: Amongst vegetal debris and

cobbles.

Remarks: The characteristics of the

habitat and shells are quite similar to

those of the European populations.

Family RISSOIDAE Gray, 1847

Botryphallus epidauricus (Brusina, 1866) (Figs. 17-27)

Material studied: 570 specimens from Madeira; 409 from Almería, more than 500 from Canary

Islands (MNHN), and more than 500 from Ceuta (MNHN).

Description: Shell (Fig. 17) minute,

tall-spired, almost cylindrical, smooth,

whitish, with a dome-shaped apex.

Aperture pyriform, slightly widening,

peristome continuous and outer lip

smooth. Protoconch (Fig. 18) with about

1 smooth whorl. Teleoconch with about

3 whorls.

ROLÁN AND TEMPLADO: High-tidal molluscan assemblage from a Madeiran boulder beach

Figures 21-27. Botryphallus epidauricus, Madeira. 21: sketch of one specimen showing by transpa-

rence part of the intestinal tract; 22, 23: details of the head-foot; 24-27: penis of several males.

Figure 28. Sketch of the head-foot of Truncatella subcylindrica, Madeira. Figure 29. Sketch of the

head-foot of Paludinella littorina, Madeira. Figure 30. Sketch of the head-foot of Assiminea cf.

grayana, Madeira. Figure 31. Sketch of the head-foot of Pseudomelampus exiguus, Madeira. Figure

32. Sketch of the head-foot of Pedipes pedipes, Madeira. Figure 33. Sketch of the head-foot of

Odostomia microeques, Madeira. Figure 34. Live position of Odostomia microeques On its host,

Botryphallus epidauricus, Madeira.

Figuras 21-27, Botryphallus epidauricus, Madeira. 21: esquema de un ejemplar mostrando por trans-

parencia parte del tracto intestinal; 22, 23: detalles de la parte anterior del animal; 24-27: penes de

algunos ejemplares. Figura 28. Esquema de la parte anterior del animal de Truncatella subcylindrica,

Madeira. Figura 29. Esquema de la parte anterior del animal de Paludinella littorina, Madeira.

Figura 30. Esquema de la parte anterior del animal de Assiminea cf. grayana, Madeira. Figura 31.

Esquema de la parte anterior del animal de Pseudomelampus exiguus, Madeira. Figura 32. Esquema

de la parte anterior del animal de Pedipes pedipes, Madeira. Figura 33. Esquema de la parte anterior

del animal de Odostomia microeques, Madeira. Figura 34. Posición habitual de Odostomia microe-

ques sobre su hospedador, Botryphallus epidauricus, Madeira.

Iberus, 18 (2), 2000

Figures 35-37. Shells of Odostomia microeques, Madeira. 35: Holotype (MNCN); 36-37: paratypes

(CER and MNHN).

Figuras 35-37. Conchas de Odostomia microeques, Madeira. 35: holotipo (MNCN); 36-37: parati-

pos (CER y MNAN).

Animal (Figs. 22, 23) translucent

white, with cephalic tentacles thin and

elongated. The eyes lie at their bases. Pro-

podium with a conspicuous, triangular,

and opaque-white pedal gland. Snout

with two pronounced distal lobes. Intes-

tine visible by transparence throughout

the last whorl of the shell, with a slight

curvature (Fig. 21). Males with a large

and flower-like penis (Figs. 24-27), also

visible by transparence, bearing several

(from 11 to 16) digitiform appendages

(apocrine glands, sensu Ponder, 1990).

Operculum thin, translucent, paucispiral

with a eccentric nucleus.

Radula (Figs. 19-20) taenioglossan,

with a raquidian tooth having the cutting

edge with 9 cusps (the central more

elongated and the others smaller towards

the periphery). Lateral teeth with 9 cusps

also, one of the central more prominent

and the others decreasing in size at both

sides. Marginal teeth with many small

cusps of the same size in their distal edge.

Habitat: Some specimens were found

under cobbles and stones that were per-

manently humid and in contact with the

sand, but most of them were buried in

the sand.

Remarks: At first we suspected that

our specimens belonged to an undescri-

bed species, due its short protoconch

(meaning a direct development) and the

long distance between Madeira and the

Mediterranean Sea. Besides, the

drawing of the penis of B. epidauricus

showed by PONDER (1990, fig. 8C) upon

material from the Strait of Gibraltar

appeared to be rather different from our

drawings. But later, we had the oppor-

tunity to study samples of a population

of B. epidauricus from Los Escullos

(Almería, SE Spain) and we found that

the shell and the male genitalia in speci-

mens of this population were identical

to those from Madeira, and therefore

belong to the same species. Perhaps the

differences between our observations

and the Ponder drawings are due to the

fact that we have studied alcohol preser-

ved material and he probably examined

living specimens.

We have also studied two similar

forms from the Canary Islands (MNHN)

ROLÁN AND TEMPLADO: High-tidal molluscan assemblage from a Madeiran boulder beach

Figures 38-41. Ovatella aequalis, Madeira. 38: shell; 39-40: protoconch; 41: radula.

Figuras 38-41. Ovatella aequalis, Madeira. 38: concha; 39-40: protoconcha; 41: rádula.

of different size, about 1. 5 mm hight the

larger one, and about 1. 0 the smaller.

The bigger form is almost identical with

the Madeiran specimens and probably is

the species named by MANZONI (1868a)

Rissoa balteata, as VERDUIN (1988) and

GOFAS (1990) suspected. The smaller

form from the Canary Island, has a shell

quite similar, but the males lack the

characteristic penis of the genus Botryp-

Iberus, 18 (2), 2000

hallus and it might be an undescribed

species of the genus Peringíella.

Two similar species have been descri-

bed in recent years: B. ovummuscae

(Gofas, 1990) from Azores and B. tuber

(Rolán, 1991) from Cape Verde islands,

both described under the genus Peringie-

lla. According to the results of this study,

the type species of this genus, B. epidauri-

cus, occurs along the Mediterranean Sea,

Atlantic coast of Spain and Portugal, and

Madeira and Canary Islands.

Family PYRAMIDELLIDAE Gray, 1840

Odostomia microeques Rolán and Templado, 1999 (Figs. 33-37)

Material studied: 22 specimens from Madeira.

Description: Shell (Figs. 35-37)

minute, oval-cylindrical, very fragile.

Spire from 2 to 3 spiral whorls, slightly

rounded, with suture impressed, and

very evident prosoclines growth lines.

Apex blunt, dome-shaped, with the pro-

toconch of type C of 206 um, emerging

and showing less than one spiral whorl.

Aperture oval and somewhat pyriform.

Columela slightly curved. Peristoma

continuous. Umbilicus absent. Columel-

ar lip not showing any fold externaly,

but a slight fold begin somewhat inner

and continues internally well developed.

Animal translucent white, with short

and divergent cephalic tentacles,

slightly tapering distally but the distal

end is expanded into a spheric tip (Fig.

33). Eyes small and placed very close

behind the base of the tentacles. Opercu-

lum with a perpendicular line to the

columela.

Habitat: Found just with its host,

Botryphallus epidauricus, in the sand

under stones.

Remarks: O. microeques is the smallest

pyramidellid gastropod known. It was

not observed during the first time in the

collecting site due its very small size.

Some samples of the sand taken under

the cobbles was carried to the laboratory

for examination under magnification.

During this study the presence of some

tiny shells, smaller than those of Botryp-

hallus epidauricus, were observed, which

was abundant in the sample. Curiously,

it was observed that these specimens

were placed on the shell of B. epidauricus

(Fig. 34). Many times, we took off the

specimens of O. microeques from the

shells of B. epidauricus, but immediately,

they looked for another specimen to go

up it again. In oposition, they rejected

the shells of other species that lived in

the same habitat. This convinced us that

O. microeques parasites B. epidauricus.

The closest species to O. microeques is

Odostomia megerlei (Locard, 1886), but

the latter is clearly bigger (see com-

ments in PEÑAS AND ROLÁN, 1999).

Family ELLOBIDAE Pfeiffer, 1854

Ovatella aequalis (Lowe, 1832) (Figs. 38-41)

Material studied: 260 specimens from Madeira.

(Right page) Figures 42-47. Protoconchs of some European species of Ellobiidae. 42: Auriculinella

bidentata, Cies Islands; 43: A. bidentata, Almería; 44-45: Myosotella myosotis, Cies Islands; 46: M.

myosotis, Almería; 47: Ovatella firminii, Cies Islands.

(Página derecha) Figuras 42-47. Protoconchas de algunas especies europeas de Ellobiidae. 42: Auriculi-

nella bidentata, /5/as Cies; 43: A. bidentata, Almería; 44-45: Myosotella myosotis, /slas Cies; 46: M.

myosotis, Almería; 47: Ovatella firminii, /slas Cies.

ROLÁN AND TEMPLADO: High-tidal molluscan assemblage from a Madeiran boulder beach

Iberus, 18 (2), 2000

Description: Shell (Fig. 38) oval-conic,

elongated, somewhat solid, brown. Pro-

toconch (Figs. 39-40) smooth, with about

one whorl. Teleoconch of about 6 whorls.

Body whorl about 70% of the total

height. First whorls with fine spiral stria-

tion and narrow and spaced prosocline

axial ribs. There is a deep subsutural

furrow in which sharp periostracal hairs

inserted at each axial rib. Suture not

depressed. Aperture oval-elongated with

two columelar folds and two prominent

parietal teeth. Animal whitish to pale-

yellowish. Radula (Fig. 41) with many

teeth, all quite similar, disposed in rows.

Habitat: Below and under cobbles of

middle size, in humid places, not close

to the sand layer.

Remarks: MARTINS (1995, 1999) has

compared the Acorean O. vulcani

(Morelet, 1860) with the Mediterranean

O. firminii (Payraudeau, 1826) and the

Madeiran O. aequalis, concluding, for the

recognition all of them as valid species,

often previously considered as

synonyms. According to this author, the

sculpture of the protoconch and the first

whorls of the teloconch seems to be very

important as specific characters in this

group. To show the differences with

other close European species we present

the protoconchs of Myosotella myosotis

(Draparnaud, 1801) (Figs. 44-46) from

Cies Islands and Almería, and Ovatella

firmini (Payraudeau, 1826) (Fig. 47)

from the Cies islands.

Auriculinella bidentata (Montagu, 1808) (Figs. 42, 43, 48-50)

Material studied: 39 specimens from Madeira; 21 from Almería; 12 from Ribadeo; 15 from Cies

Islands.

Description: Shell (Fig. 48) oval-conic

elongated, smooth, withish, with blunt

apex, very similar to the European popu-

lations studied. Protoconch (Figs. 42, 43,

49, 50) smooth with about half whorl.

The teleoconch begins with axial proso-

cline striation which cross fine spiral

threads. This sculpture almost desappe-

ars in subsequent whorls. Aperture oval,

elongated with a prominent parietal

tooth and a curved columelar fold below.

Animal white, sometimes with small

dark areas at the tip of the cephalic tentacles.

Habitat: Found amongst and under

cobbles of middle size, in humid

places.

Remarks: The specimens from Ma-

deira are almost identical to the Euro-

pean populations examined, showing

the specimens from Almería the typical

sculpture of the first whorl of the teleo-

conch (Figs. 49-50), and being less evi-

dent in the shells of the Cies Islands

(Fig. 42). This species widespread from

the British Isles to the Mediterrnean,

Azores and Madeira.

Pedipes pedipes (Bruguiere, 1789) (Figs. 32, 51-54)

Pedipes afer (Gmelin, 1791)

Material studied: 5 specimens from Madeira.

Description: Shell (Fig. 51) globose,

solid, brown, with low spire and very

large body whorl, that averaging near the

90% of shell length. Protoconch (Figs. 52-

54) very small, with less than one whorl.

The teleoconch begins with spiral and pro-

socline axial sculpture, the later desap-

pearing in the following whorls. Aperture

ovate, widely rounded at base, about 70%

of the length of body whorl. There are two

strong columellar teeth, and one strongest

parietal tooth very elongated.

Animal (Fig. 32) white with grey

pigmentation towards the tip of the cep-

halic tentacles. Anterior part of the foot

bilobulated by a central incision. Snout

ROLÁN AND TEMPLADO: High-tidal molluscan assemblage from a Madeiran boulder beach

500 um

100 um

Figures 48-50: Auriculinella bidentata, Madeira. 48: shell; 49-50: protoconch.

Figuras 48-50: Auriculinella bidentata, Madeira. 48: concha; 49-50: protoconcha.

short and cephalic tentacles contractile,

somewhat elongated when extended.

Habitat: Found amongst boulders in

the upper level of the collecting site.

Remarks: This species is known from

Azores, Madeira, Canary, Cape Verde

and Sao Tomé Islands (FERNANDES AND

ROLÁN, 1993; ÁVILA, 2000).

Pseudomelampus exiguus (Lowe, 1832) (Figs. 31, 55-57)

Material studied: 2 specimens from Madeira; 12 from Almería; 20 from Ribadeo.

Description: Shell (Fig. 55) ovoid,

solid, pinkish-brown, body whorl more

than 90% of total shell hight. Protoconch

(Fig. 56) smooth, heterostrophic, with its

spiral axis perpendicular to the axis of

the shell, and partly covered with first

whorl of teleoconch. Microsculpture of

irregular spiral striae (Fig. 57). Aperture

elongate with one columellar tooth and

two parietal theeth. Animal (Fig. 31)

whitish, foot bilobulated anteriorly, cep-

halic tentacles short, somewhat flattened,

with the eyes in the middle of their bases.

Habitat: Found amongst big stones in

the upper part of the bottom.

Remarks: The lectotype of this species

was figured by MARTINS (1996, fig. 180),

being Madeira the type locality.

Iberus, 18 (2), 2000

Family KELLIDAE Forbes and Hanley, 1848

Lasaea rubra (Montagu, 1803)

Material studied: 2 specimens from Madeira; more than 200 specimens in Cies Islands.

Remarks: L. rubra is an extremely

common species, frequently associated

with some of the previously mentioned

DISCUSSION

The under-boulder molluscs assem-

blage studied in a protected place of the

upper level of the tide and its distribu-

tion in different levels in Madeira has

some resemblance with that described

by MORTON (1975) in New Zealand

(dominated by species of Ellobiidae,

Assimineidae and Caecidae). Four of the

fourteen species we found clearly domi-

nated in this habitat in Madeira. Botryp-

hallus epidauricus in the lowest level, on

the sand under rocks, Paludinella litto-

rina in a somewhat higher layer,

amongst small stones in contact with

sand. Ovatella aequalis and Assiminea cf.

grayana dominated in an upper level,

under boulders without contact with

sand. Other species frequent in this

habitat were Caecum armoricum, Trunca-

tella subcylindrica and Odostomia microe-

ques in the same layer that B. epidauricus,

and Auriculinella bidentata, living togeth-

er with Paludinella littorina. The other

species found were scarce. The ellobids

Pedipes pedipes and Pseudomelampus

exiguus, and the littorinids Littorina

striata and Melaraphe neritoides are

typical supralittoral species, which

sometimes can be found in the highest

level of the habitat studied. Lasaea rubra

is very common in tufts of coralline

algae and lichens (Lichina sp.) high in

the intertidal zone and only isolated

specimens can be found in boulder

beach (BULLOCK ET AL., 1990). Last,

species in other areas, sometimes in

high number. In the studied community

its presence was scarce.

Caecum clarkii prefer lower shore (mid-

littoral or sublittoral) but is capable of

penetrating the upper levels when con-

ditions are favourable (PONDER, 1990).

Only two of the species found are

apparently endemic of Madeira: Ovatella

aequalis and Odostomia microeques. The

latter might be present in other areas

where its host occurs, but it might be

over looked due its minute size (smaller

than 1 mm). Ovatella aequalis is replaced

by its related species Myosotella myosotis

(Draparnaud, 1801) in the European

coasts and by O. vulcani (Morelet, 1860)

in Azores. All the other species, but Lit-

torina striata and Pedipes pedipes, are also

present in the European mainland, both

in the Atlantic and Mediterranean

coasts. Ten of the species found in

Madeira occurs also in Azores and eight

in Canary Islands.

The most abundant species, Botryp-

hallus epidauricus, widespreads along the

Atlantic and Mediterranean coasts of

Europe and also in Canary Islands. It is

replaced in Azores (northward) and in

Cabo Verde Island (southward) by the

related species B. ovummuscae and B.

tuber, respectively.

PONDER (1990) studied a similar ha-

bitat in the Strait of Gibraltar, but in a so-

mewhat lower level. He studied an inter-

tidal gravel beach at Ceuta, and its up-

per level coincides with the lowest one

studied by us. In both localities (Funchal

(Right page) Figures 51-54. Pedipes pedipes, Madeira. 51: shell; 52-54: protoconch. Figures 55-57:

Pseudomelampus exiguus, Madeira. 55: shell; 56: protoconch; 57: microsculpture.

(Página derecha) Figuras 51-54. Pedipes pedipes, Madeira. 51: concha; 52-54: protoconcha. Figuras

55-57: Pseudomelampus exiguus, Madeira. 55: concha; 56: protoconcha; 57: microescultura.

ROLÁN AND TEMPLADO: High-tidal molluscan assemblage from a Madeiran boulder beach

Iberus, 18 (2), 2000

in Madeira, and Ceuta in the Strait of Gi-

braltar) this level is dominated by the

same species: Botryphallus epidauricus.

The small molluscs found in this

peculiar habitat in Madeira seem to

form a distinctive ecological grouping

which have close parallels in Euro-

pean/North African mainland and in

other Macaronesian Islands. Some

species of this grouping are present in

this habitat in all areas of the temperate

NE Atlantic, and other are replaced by

close related species according to the

geographical area.

The species found in this habitat in

Madeira and in some localities of the

Spanish coasts (Ribadeo and Cies

Islands, NW Spain, and Los Escullos, SE

Spain) are included in Table II. Four

species have been found in this habitat

in all localities sampled: P. littorina, T.

subcylindrica, A. bidentata and P. exiguus.

In Madeira this molluscan assemblage is

particularly diverse. Six of the species

found here, including the two endemic

of Madeira, were not found in the other

localities sampled. Some of them are

present in these localities, but they did

not find in this habitat. The more remar-

kable peculiarity of this molluscan

assemblage in Madeira is the presence

of an parasitic pyramidellid gastropod

(O. microeques). In contrast, only two of

the species collected in this habitat in

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