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Iberus 


Vol. 28 (1) 


REVISTA DE LA 
SOCIEDAD ESPAÑOLA 
DE MALACOLOGÍA 


Oviedo, junio 2010 


Iberus 
Revista de la 
SOCIEDAD ESPAÑOLA DE MALACOLOGÍA 


Comité DE REDACCIÓN (BOARD OF EDITORS) 
EDITOR DE PUBLICACIONES (EDITOR=-IN-CHIEF) 
Serge Gofas Universidad de Málaga, España 


DIRECTOR DE REDACCIÓN (EXECUTIVE EDITOR) 
Gonzalo Rodríguez Casero Apdo. 156, Mieres del Camino, Asturias, España 


EDITORA EJECUTIVA (MANAGING EDITOR) 
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EDITORES ADJUNTOS (ASSOCIATE EDITORS) 


Francisco Javier Conde de Saro Embajada de España, Japón 

Benjamín Gómez Moliner Universidad del País Vasco, Vitoria, España 

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

Emilio Rolán Mosquera Universidad de Vigo, Vigo, España 

José Templado González Museo Nacional de Ciencias Naturales, ESIC, Madrid, España 
Jesús S. Troncoso Universidad de Vigo, Vigo, España 


Comité EDITORIAL (BOARD OF REVIEWERS) 


Kepa Altonaga Sustacha Universidad del País Vasco, Bilbao, España 

Eduardo Angulo Pinedo Universidad del País Vasco, Bilbao, España 

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

Thierry Backeljau Institut Royal des Sciences Naturelles de Belgique, Bruselas, Bélgica 
Rijdiger Bieler The Field Museum, Chicago, Estados Unidos 

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

Jose Castillejo Murillo Universidad de Santiago de Compostela, Santiago de Compostela, España 
Karl Edlinger Noturhistorisches Museum Wien, Viena, Austria 

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

José Carlos García Gómez Universidad de Sevilla, Sevilla, España 

Gonzalo Giribet de Sebastián Harvard University, EE.UU. 

Edmund Gittenberger National Natuurhistorisch Museum, Leiden, Holanda 

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

Gerhard Haszprunar Zoologische Staatssammlung Múnchen, Múnchen, Alemania 

Yuri |. Kantor A.N. Severtzov Institute of Ecology and Evolution, Moscú, Rusia 
María Yolanda Manga González Estación Agrícola Experimental, CSIC, León, España 

Jordi Martinell Callico Universidad de Barcelona, Barcelona, España 

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

Tokashi Okutani Nihon University, Fujisawa City, Japón 

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

Pablo E. Penchaszadeh Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina 
Winston F. Ponder Australian Museum, Sydney, Australia 

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

Me de los Ángeles Ramos Sánchez Museo Nacional de Ciencias Naturales, CSIC, Madrid, España 
Francisco Javier Rocha Valdés Instituto de Investigaciones Marinas, CSIC, Vigo, España 

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

Joandoménec Ros ¡ Aragones Universidad de Barcelona, Barcelona, España 

María Carmen Salas Casanovas Universidad de Málaga, Málaga, España 

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

Victoriano Urgorri Carrasco Universidad de Santiago de Compostela, Santiago de Compostela, España 
Anders Warén Swedish Museum of Natural History, Estocolmo, Suecia 


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


Iberus 


ys A IN Y 00 TIA 
(Curs 19 2010 m0) 


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REVISTA DE LA 
SOCIEDAD ESPANOLA 
DE MALACOLOGIA 


Vol. 28 (1) Oviedo, junio 2010 


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: Serge Gofas, Editor de Publicaciones, Departamento de Bio- 
logía Animal, Universidad de Málaga, Campus de Teatinos, s/n, 29071, Málaga, España. 

Los trabajos se entregarán por triplicado (original y dos copias). Se recomienda a los autores leer 
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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, 
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Dep. Leg. B-43072-81 
ISSN 0212-3010 
Diseño y maquetación: Gonzalo Rodríguez 


Impresión: LOREDO, S. L. - Gijón 


O Sociedad Española de Malacología —___—_—_———  Iberus, 28 (1): 1-11, 2010 


Reproduction du mollusque envahisseur Corbicula flumi- 
nea (O. E Miiller, 1774) (Bivalvia: Corbiculidae) et recru- 
tement et distribution de ses juvéniles dans un canal de la 
région toulousaine (France) 


Reproducción del molusco invasor Corbicula fluminea (O. E Múller, 
1774) (Bivalvia: Corbiculidae) y reclutamiento y distribución de sus 
juveniles en un canal la región de Toulouse (France) 


Charles DUBOIS et Alberto MARTÍNEZ-ORTÍ* 


Recibido el 13-V-2009. Aceptado el 6-X-2009 


RÉSUMÉ 


Les données recueillies sur un cycle d'un an (mars 1993 — mars 1994) concernant l'incu- 
bation des larves et le recrutement des ¡juvéniles du Corbicula fluminea, bivalve d'origine 
asiatique introduites en Europe, sont présentées pour deux habitats différents (milieu du 
canal et herbiers) d'un méme site du Canal Latéral, pres de Toulouse. La présence de 
larves incubées au niveau des hémibranchies internes des adultes est observée durant 5 
mois: de mi-mai á mi-octobre, ou deux périodes d'incubation se dégagent. Le nombre de 
larves incubées s'éléve aá plusieurs milliers et se trouve proportionnel a la taille de la 
coquille des adultes. Le recrutement des ¡uvéniles de taille comprise entre 1,5 et 2,5 mm 
s'observe du mois d'aoút au mois de juin de l'année suivante, ou l'on peut également dis- 
tinguer deux périodes: une premiere s'étalant d'aoút á novembre et une seconde débutant 
en décembre jusqu'a juin. Selon l'habitat considéré, ces périodes de recrutement ont une 
importance relative trés différentes: la premiere (fin été — automne], quasi-inexistante au 
niveau des herbiers, est importante dans la zone profonde. Inversement, lors de la 
seconde (hivers — début printemps), le nombre de ¡juvéniles par rapport au nombre 
d'adultes est plus fort au niveau des herbiers. Cette différence dans le recrutement suggére 
une migration ou une dispersion de la part de ces petits individus, notamment durant la 
saison chaude ou de nombreux bateaux naviguent, créant des remous importants. 


ABSTRACT 


Data obtained over a cycle of one year (March 1993 — March 1994) regarding the incu- 
bation of the larvae and the recruitment of the juveniles of Corbicula fluminea, bivalve of 
Asian origin introduced in Europe, are shown for two different habitats (middle part of the 
channel and vegetated area) of the same area of the Canal Latéral, near Toulouse 
(France). The presence of incubated larvae in the inner demibranchs of the adults has 
been observed during five months: from mid May to mid October, in which there are two 
different periods of incubation. The number of incubated larvae amounts to several thou- 
sands and is proportional to the shell size of the adults. The recruitment of the juveniles of 
size between 1.5 and 2.5 mm is observed between August and June of the following 


* Departament de Zoologia. Facultat de Ciencies Biológiques. Universitat de Valencia. c/. Dr. Moliner, 50. E- 
46100 Burjassot (Valencia, Espagne) et Museu Valencia d'Historia Natural. 


Iberus, 28 (1), 2010 


year, where two periods can also be distinguished: the first one from August to November 
and the second one from December until June. According to the considered habitat, these 
periods of recruitment have a very different importance: the first one [end of summer — 
autumn), almost non-existent at the vegetated level, is important in the deep zone. On the 
contrary, in the second one (winter — beginning of spring), the number of juvenile in rela- 
tion to the number of adults is higher in the vegetated area. This difference in the recruit- 
ment suggests a migration or dispersion of these small individuals, particularly during 
the warm season where a large number if boats are sailing, which can create important 
eddies in the water column. 


RESUMEN 


Se muestran los datos obtenidos durante un ciclo de un año (marzo 1993 — marzo 1994) 
sobre la incubación de las larvas y el reclutamiento de los juveniles de Corbicula flumi- 
nea, bivalvo de origen asiático introducido en Europa, se muestran en dos hábitats dife- 
rentes (centro del canal y zona lateral con vegetación) de un mismo lugar del Canal Laté- 
ral, cerca de Toulouse (Francia). Se ha observado la presencia de larvas incubadas en las 
hemibranquias internas de los ejemplares adultos durante cinco meses: desde mitad de 
mayo a mitad de octubre, existiendo dos periodos diferentes de incubación. El número de 
larvas incubadas es de varios miles y es proporcional al tamaño de la concha de los adul- 
tos. El reclutamiento de los juveniles, de tamaño comprendido entre 1,5 y 2,5 mm, se 
observa entre los meses de agosto y junio del año siguiente, donde se pueden igualmente 
distinguir dos periodos: el primero desde agosto a noviembre y el segundo desde diciem- 
bre hasta junio. Según el hábitat considerado, estos periodos de reclutamiento tienen una 
importancia relativa muy diferente: el primero [final del verano — otoño), casi inexistente 
al nivel de la zona de vegetación, es importante en la zona profunda. Inversamente, en el 
segundo periodo (invierno — comienzo de la primavera), el número de ¡juveniles en rela- 
ción al número de adultos es más elevado en la zona de vegetación. Esta diferencia en el 
reclutamiento sugiere una migración o una dispersión de parte de estos pequeños indivi- 
duos, especialmente durante la estación cálida, cuando numerosos barcos navegan por el 
canal creando importantes remolinos en la columna de agua. 


INTRODUCTION 


Au début des années 1980, la pré- 
sence du bivalve envahisseur Corbicula 
fluminea (O.F. Múller, 1774) (Fig. 1A), 
d'origine asiatique, fut signalée en 
Europe occidentale pour la premiere fois 
(MOUTHON, 1981). A l'heure actuelle, de 
nombreux cours d'eau européens héber- 
gent ce mollusque, qui fut décrit comme 
une véritable peste aux Etats-Unis dont 
la colonisation remonte aux années 1930 
(Isom, 1986): le fleuve Rhin aux Pays- 
Bas (BI] DE VAATE ET GREIDANUS-KLAAS, 
1990), en Allemagne (BI DE VAATE, 1991; 
DEN HARTOG, VAN DEN BRINK ET VAN 
DER  VELDE, 1992), en Belgique 
(SWINNEN, LEYNEN, SABLON, DUVIVIER 
ET VANMAELE, 1998), en Pologne (DOMA- 
GALA, LABECKA, PILECKA-RAPACZ ET 


MIGDALSKA, 2004; LABECKA, DOMAGALA 
ET PILECKA-RAPACZ, 2005), en Répu- 
blique Tcheque (BERAN, 2006) et au 
Royaume Uni (BAKER, CLARKE ET 
HOWwLETT, 1999; ALDRIDGE ET MULLER, 
2001). En France les fleuves Dordogne, 
Loire, Garomne, Rhóne, Seine, 1'Adour, 
le Lot, le Canal du Midi et le Canal 
Latéral, le Tarn et 11 Aude (MOUTHON, 
1981; GIRARDL, 1989; TOURENO. ET PUJOL, 
1991; GRUET, 1992; DuBo1s, 1995a, 1995b; 
DuBoIs Er TOURENO, 1995; DUBOIS ET 
ORIGNY, 1998; MOUTHON, 2001a, 2001b, 
2003). En Espagne, la riviere Miño, le 
Guadalquivir, l'Ebre, le Guadiana et le 
Piedras sont également colonisées 
(PÉREZ-QUINTERO, 1990, 2008; ARAUJO, 
MORENO ET RAMOS, 1993), ainsi que l'es- 


DuBoIs Er MARTÍNEZ-ORTÍ: Reproduction du Corbicula fluminea 


tuaire du Tage, le Duero et les affluents 
de Guadiana au Portugal (MOUTHON, 
1981; ARAUJO ET AL., 1993; PÉREZ-QUIN- 
TERO, 2008). ARAUJO ET AL. (1993) jugent 
que seul C. fluminea colonise les cours 
d'eaux francais et espagnols. 

C. fluminea incube les larves au 
niveau des hémibranchies internes des 
adultes (Fig. 1B) jusqu'a leur libération 
en pleine eau ou elles se dispersent par 
flottaison (PRETZANT ET CHALERMWAT, 
1984; Dubo1Is, 1995a, 1995b). 

L'objet de cette publication s'inscrit 
dans l'étude de la dynamique des 
populations et dans l'analyse des 
modalités de dispersion a l'échelle sai- 
sonniere de C. fluminea, menées dans le 
Canal Latéral, au nord de Toulouse 
(FRANCE) (SITE DE LALANDE). DUBOIS ET 
TOURENO (1995) ont réalisé une étude 
préliminaire sur la zone profonde du 
méme Canal toulousaine qui nous 
continuons et nous présentons ici les 
données recueillies en 1993-94 concer- 
nant la reproduction et le recrutement 
des juvéniles des deux sous-popula- 
tions benthiques: celle habitant les 
bords du canal, colonisé par des macro- 
phytes immergés et celle habitant le 
sédiment de la zone profonde; ainsi que 
la répartition inter et intra-habitat 


MATÉRIEL ET MÉTHODES 


Le Canal Latéral, qui relie Toulouse 
a Bordeaux (France) a une largeur 
d'une vingtaine de metres et une pro- 
fondeur de 2,5 m au centre du lit. Les 
bords, sur une largeur d“environ 1,5 m, 
sont peu profonds (0,5 m a 0,8 m) et sont 
colonisés d'herbiers de macrophytes 
immergés (Vallisneria spiralis L.). 

Les 10 premiers centimetres de sédi- 
ment sont composés de vase, tres enri- 
chie en débris organiques (feuilles en 
décomposition, essentiellement de pla- 
tanes, qui bordent le canal). Plus profon- 
dément, le sédiment est un limon argi- 
leux  progressivement réduit (sur 
environ 15 cm), reposant sur une couche 
d'argile imperméable. 

La vitesse du courant, relativement 
constante, s'éleve a 7 cm/s en moyenne. 


La température a la mi-journée varie 
entre 6 *C (janvier) et 25 *C (aoút); la 
température maximale relevée étant de 
DIE. 

Pour l'étude de la reproduction nous 
avons prélevé dans les herbiers, tous les 
15 jours, 20 a 30 individus adultes [dont 
la longueur antéro-postérieure maxi- 
male L est supérieure a 10 mm, d'apres 
ALDRIGE ET MC MAHON (1978)], de 
maniere a suivre le nombre de larves 
incubées dans les hémibranchies 
internes (Fig. 1B). Pour cela, les spéci- 
mens sélectionnés ont été fixés au 
formol (a 10 %), puis disséqués: une 
hémibranchie interne par individu a été 
prélevée et délicatement lacérée afin de 
recueillir les larves. Celles-ci sont alors 
maintenues en suspension dans un 
volume d'eau distillée. Des échantillons 
(en général 5) de 4 ml sont prélevés (a 
lVaide d'une seringue graduée), les 
larves dénombrées et le nombre total 
incubé est estimé. De cette maniere, l'er- 
reur standard (SE= y (s?/n) avec s?= 
variance, n= nombre d'échantillons) par 
rapport a la moyenne n'excede jamais 
5% pour un adulte gravide. 

Pour le suivi de la structure de la 
population, des prélevements quantita- 
tifs ont été effectués a l'aide d'une benne 
de type Eckman (surface= 225 cm) 
pénétrant a une profondeur d'environ 
10 cm dans le sédiment. A chaque cam- 
pagne de terrain (une fois par mois), 6 
échantillons de 2 bennes ont été récoltés 
dans les herbiers et 8 échantillons de 4 
bennes le long d'un transect dans la 
zone profonde (sauf cas particuliers, cf 
Tableau 1). Au laboratoire, les individus 
sont triés du matériel retenu et mesurés 
(L= longueur antéropostérieure maxi- 
male, a 0,1 mm pres) sous une loupe 
binoculaire pour L<10 mm (grossisse- 
ment x10, précision: 0,05 mm) ou avec 
un pied-a-coulisse pour L>10 mm (pré- 
cision: 0,02 mm). Les plus petits spéci- 
mens considérés ont une taille de 1,5 
mm, et nous prendrons en compte dans 
ce travail que des juvéniles de taille 
comprise entre 1,5 et 2,5 mm. 


RÉSULTATS 


Iberus, 28 (1), 2010 


Figure 1. Corbicula fluminea du Canal Latéral (Toulouse, France). A. Coquille (24,2 mm de 
largeur). B. Hémibranchie interne avec des larves. 

Figura 1. Corbicula fluminea del Canal Latéral (Toulouse, Francia). A. Concha (24,2 mm longitua). 
B. Hemibranquia interna mostrando las larvas incubadas. 


Incubation des larves 

La période d'incubation des larves 
dans les hémibranchies internes des 
adultes dure 5 mois, de la mi-mai a la 
mi-octobre (Fig. 2). Nous pouvons dis- 
cerner 3 pics de fort taux d'incubation 
(Tl= proportion d'adultes incubant des 


larves, en %): un premier couvrant le 
mois de juin (TI maximal= 90,5%) et les 
deux suivants (TI maximal= 80,0% et 
66% respectivement) qui couvrent les 
mois d'aoút et septembre (Fig. 2). 

Sur lensemble de la période d'incu- 
bation, le nombre de larves est tres 


DUBOIS ET MARTÍNEZ-ORTÍ: Reproduction du Corbicula fluminea 


Tableau 1. Densité des juvéniles (1,5 — 2,5 mm) aux différentes dates de prélevement. Ne= nombre 
d'échantillons; Nb= nombre de bennes pour un échantillon; dens= densité en ind./m?; SE= erreur 


standard relative (en %). 


Tabla 1. Densidad de juveniles (1,5 — 2,5 mm) en diferentes periodos de recolección. Ne= número de mues- 
tras; Nb= número de dragas por muestra; dens= densidad en ind./m?; SE= error estándar relativo (en %). 


ZONE PROFONDE 
Date Ne Nb dens SE (9) 
23.09:73 4 5 120 26,3 
20.04.93 8 4 178 11558 
18.05.93 8 4 143 218 
22.06.93 8 4 46 16,1 
20.07.93 8 4 0 0,0 
23.08.93 6 4 254 22,6 
20.09.93 8 4 340 16,4 
18.10.93 8 4 413 14,4 
ANOS 8 4 182 16,6 
17.01.94 8 4 265 14,4 


variable. Le nombre moyen par hémi- 
branchie de larves trouvées dans un 
exemplaire adulte de taille moyenne est 
de 2114 (Fig. 3). Cependant, le maximum 
observé est proportionnel a la taille de 
l'individu mature considéré. De sorte que 
nous avons pu établir la relation: 


Nm= - 8568 + 691 L (mm) (1) 


ou: 

Nm= nombre maximal théorique de 
larves par hémibranchie interne. 

L (mm)= taille (en mm) de l'adulte 
considéré. 

Parmi les adultes gravides, nous 
avons alors calculé le “taux moyen de 
remplissage hémibranchial” (TRHg) a 
chaque date de prélevement. Nous défi- 
nissons TRHg comme la moyenne, sur 
l'ensemble des individus gravides d'un 
prélevement, du pourcentage observé 
par rapport au nombre maximal théo- 
rique: 


TRHg=1/n) (Ni / Nmi) 


i=1 


avec: 


HERBIERS 

Date Ne  Nb dens SE(%) 
23.03.93 l 5 2027 

30.04.93 5 2 876 1117 
28.05.93 6 2 1178 1153 
28.06.93 6 2 159 19) 
21.07.93 6 2 3) 46,1 
30.08.93 6 2 48 49 7 
28.09.93 6 2 385 31,5 
26.10.93 6 2 122 38,9 
2993 6 2 81 5,2 
ZONE 6 Í 215 28,5 
02.02.94 6 2 911 23,6 


n= nombre d'adultes gravides 1 
observé. 

Ni= nombre observé de larves incu- 
bées par hémibranchie interne de 
l'adulte i. 

Nmi= nombre maximal théorique de 
larves incubées par hémibranchie 
interne de l'adulte i, obtenu d'apres (1). 

Pour un prélevement, nous pouvons 
considérer que le taux moyen de rem- 
plissage hémibranchial est indépendant 
de la structure fréquence-taille de 
l'échantillon (la proportion Ni/Nmi est 
indépendante de la taille, contrairement 
a Ni), si bien que les valeurs TRHg sont 
immédiatement comparables entre elles 
(Fig. 3), et peuvent étre directement 
appliquées a la population (dont la 
structure est différente de celle des 
échantillons). 

Le taux moyen de remplissage 
TRHg connaít une évolution similaire a 
celle du taux d'incubation Tl, avec deux 
périodes qui se dégagent (Fig. 2): une 
premiere couvrant le mois du juin 
(MRS timaximale "65,5. ..70), et une 
seconde comprenant plusieurs pics, de 
juillet (TRHg maximal= 54,9%) a début 


Iberus, 28 (1), 2010 


TAUX (%) 


Nombre (N) de larves incubees 


2 
s0— INCUBATION 
SN 
7 40 
Í 30— E 
ES 20 Á 2 TRHt= TI. TRHg A 
10 sE 
a LN 
RECRUTEMENT 
600 
.- 
y 
A A Herbiers JP Zone profonde 
Z 450 1 4 
O / 
OS 
== 
A a 
5 300 ss 
Y NN 
Z / M 
% 
150 id Ñ 
/ 
ca / 
a , B 
A ss 
EN ¿e? 
0 
4 03 04 05 06 07 08 09 10 11 12 01 02 03 
1993 1994 


MOIS 


Figure 2. Evolution temporelle du taux d'incubation (TI, en %) et Pindice de remplissage hémi- 
branchial des adultes gravides (PRHg, en %). Figure 3. Nombre de larves incubées en fonction de 
la taille des adultes gravides. Les carrés noirs représentent les valeurs maximales observées, á partir 
desquelles fut établie la régression linéaire. Figure 4. A: Evolution temporelle de Pindice de rem- 
plissage hémibranchial de la population totale d'adultes (TRHt, en %); B: Evolution temporelle 
du nombre de juvéniles de la premiére classe de taille (1,5-2,5 mm) rapportée au nombre 
d'adultes, au niveau des herbiers et de la zone profonde. 

Figura 2. Evolución temporal de la tasa de incubación (TL, en %) y el índice de rellenado hemibran- 
quial de los adultos grávidos (TRHg, en %). Figura 3. Número de larvas incubadas respecto al tamaño 
de los adultos grávidos. Los cuadros negros representan los valores máximos observados, a partir de los 
cuales se estableció la regresión lineal. Figura 4. A: Evolución temporal del índice de rellenado hemi- 
branquial de la población total de adultos (IRHt, en %); B: Evolución temporal del número de juveni- 
les de la primera clase de tamaño (1,5-2,5 mm) en función del número de adultos, a nivel de la vegeta- 


ción y de la zona profunda. 


DUBOIS Er MARTÍNEZ-ORTÍ: Reproduction du Corbicula fluminea 


Tableau H. Parametres de la relation puissance de Taylor: S?= aXP. Dindice d'agrégation “b” est 
donné avec un intervalle de confiance de 95%. R*= coefficient de variation; p= niveau de probabi- 
lité du test de Student. 

Tabla II. Parámetros de la relación de Poder de Taylor: S*= aX?. El índice de agregación “b” se da con 
un intervalo de confianza del 95%). R*= coeficiente de variación; p= nivel de probabilidad del test de 


Student. 


Habitat 0 R p 
Zone profonde 0,415 1,886 + 0,288 0,954 < 0,005 
Herbiers 120179 1,355 + 0,407 0,826 < 0,005 


octobre oú TRHg diminue progressive- 
ment. 

Le décalage que l'on observe entre 
les deux courbes résulte de l'asynchro- 
nisme dans la reproduction entre les 
individus. 

Ceci rend difficile l'estimation du 
temps d'incubation des larves, mais on 
peut considérer cependant qu'il est infé- 
rieur ou égal au pas de temps choisi -15 
jours— puisqu'on observe de grosses varia- 
tions entre certaines dates successives. 

Le produit de ces deux courbes (Fig. 
4A) donne l'évolution du taux moyen 
de remplissage hémibranchial de la 
population entiére d'adultes (gravides 
et non-gravides: THRt= TI x TRHg). 


Recrutement et distribution des 
juvéniles 

Nous donnons l'évolution de la 
densité benthique des plus petits juvé- 
niles considérés: dont la taille est com- 
prise entre 1,5 et 2,5 mm, dans la zone 
profonde du canal et au niveau des her- 
biers a Vallisneria (Fig. 4B). 

La densité des adultes (L>10 mm) 
est tres différente selon l'habitat: elle 
s'éleve, en moyenne sur l'ensemble des 
prélevements, a 78 + 5 ad./m? en zone 
profonde et a 359 + 32 ad./m* au niveau 
des herbiers. Ainsi, nous exprimons 
(Fig. 4B) la densité des juvéniles en 
nombre pour 100 adultes (juv./100 ad.) 
de maniére a comparer les évolutions 
du recrutement dans les deux habitats 
(les densités exprimées en nombre par 
m? sont données Tableau D). 

Dans la zone profonde, on observe 
trois pics de recrutement d'importance 


décroissante. Le premier pic (octobre) 
correspond au recrutement des juvéniles 
issus de la premiere période d'incuba- 
tion (juin). Ce recrutement débute des le 
mois d'aoút et s'étale jusqu'au mois de 
novembre. Les pics suivants (janvier et 
avril) correspondent au recrutement 
issu de la seconde période d'incubation 
(mi-juillet — fin septembre) et s'étale de 
décembre á juin (Fig. 4B). 

Au niveau des herbiers, le recrute- 
ment des juvéniles issus de la premiere 
période d'incubation s'étend de fin sep- 
tembre a début octobre, et est tres peu 
marqué  (densité maximale= 107 
juv./100 ad.) par rapport á celui observé 
en zone profonde (densité maximale= 
529 juv./100 ad., avec une amplitude 
plus forte). Le recrutement correspon- 
dant a la seconde période d'incubation 
apparait début janvier et s'étale jus- 
qu'au mois de juin. Cette phase de 
recrutement possede a peu pres la 
méme amplitude que celle observée en 
zone profonde, mais elle est cette fois-ci 
d'intensité supérieure (les densités 
maximales s'élevent á 565 juv./100 ad. 
au niveau des herbiers et 340 juv./100 
ad. en zone profonde) (Fig. 4B). 

Nous donnons les erreurs standards 
obtenues par rapport aux densités 
moyennes des juvéniles considérés (SE 
en %) a chaque prélevement (Tableau 1). 
Au niveau des herbiers, ces valeurs sont 
en moyenne plus fortes et augmentent 
(globalement) lorsque la  densité 
diminue, ce qui n'est pas le cas en zone 
profonde. 

De maniere a savoir si ces variations 
traduisent une variation de la réparti- 


Iberus, 28 (1), 2010 


tion au sein d'un méme habitat (au 
cours du cycle) et comparer les résultats 
entre herbiers et zone profonde, nous 
avons appliqué la loi de puissance de 
TAYLOR (1961): 5:23 aX ou is esta 
variance et X la moyemne. 

Cette loi générale s'applique á une 
grande gamme de taille de surface 
d'échantillonnage  (PALOHEIMO ET 
VUKOv, 1976; TAYLOR, WoIwOD ET 
PERRY, 1978). L'exposant b traduit le 
degré de contagion et peut donc étre 
assimilé a un indice d'agrégation : 

lorsque b= 1, la distribution est au 
hasard. 

lorsque b< 1, il y a tendance vers une 
distribution uniforme  (quand  b 
diminue). 

lorsque b> 1, il y a tendance vers une 
distribution contagieuse (quand b aug- 
mente). 

Le terme constant a est plus difficile 
a interpréter et dépend des conditions 
d'échantillonnage (DOWNING, 1979). 
Cependant, toutes choses égales par 
ailleurs, le terme a est d'autant plus fort 
que SE est important (c'est-á-dire que S? 
est grand devant X). 

Cette analyse (Tableau II) nous 
permet de conclure que, pour les deux 
habitats, la répartition des juvéniles est 
constante au cours de l'année puisque 
les régressions puissances sont statisti- 
quement significatives (p< 0,005). Aussi, 
ces répartitions sont contagieuses (b> 1 
dans les deux cas) et le degré de conta- 
gion est plus fort en zone profonde (b= 
1,886) qu'au niveau des herbiers (b= 
1,355). Cependant, il semble que 
l'échantillonnage des juvéniles considé- 
rés soit de moins bonne qualité dans les 
herbiers que dans la zone profonde, au 
regard de la constante a (Tableau Il), ce 
qui réduit la puissance de l'affirmation 
précédente. 


DISCUSSION 


D'apres la littérature, nous savons que 
C. fluminea est une espece hermaphrodite 
simultanée. Les spermatozoides sont émis 
dans le milieu aquatique, mais la fécon- 
dation a lieu, au niveau des hémibranchies 


internes. Les zygotes s'y développent, puis 
sont libérés au premier stade juvénile avec 
une taille d'environ 220 um (BRITTON ET 
MORTON, 1979; KRAEMER ET GALLOWAY, 
1986; RAJAGOPAL, VAN DER VELDE ET BJJ 
DE VAATE, 2000). 

Deux périodes de reproduction, c'est- 
a-dire d'émission des spermatozoides, ont 
été observées chez cette espece: l'une au 
printemps — début été, lorsque la tempé- 
rature augmente sensiblement; l'autre a 
la fin de l'été — automne lorsque la tem- 
pérature, alors forte et stable, commence 
a diminuer. Plus précisément, c'est la sper- 
matogénese qui est amorcée par ces varia- 
tions de température, alors que l'ovogé- 
nese est continue tout au long de l'année 
(avec un ralentissement en hiver) chez les 
individus adultes (KRAEMER ET AL., 1986). 

Notre étude sur le suivi du taux d'in- 
cubation et du taux moyen de remplissage 
hémibranchial permet de distinguer en 
effet deux périodes: l'une, intense, durant 
le mois de juin et l'autre s'étalant de mi- 
juillet a début octobre, présentant plu- 
sieurs pics, ou le nombre de larves incu- 
bées décroit progressivement (Fig. 2). Par 
contre, la relation avec la température n'est 
pas claire puisque la seconde période d'in- 
cubation est amorcée en juillet, alors que 
les températures maximales sont atteintes 
au mois d'aoút. 

Cependant ces deux périodes sont tres 
proches dans le temps, si bien que leur 
distinction est délicate, compte tenu, de 
plus, de l'asynchronisme entre les indivi- 
dus. Aussi, le nombre et la distinction des 
périodes d'incubation dépendent du pas 
de temps choisi pour les prélevements (15 
jours dans notre étude) par rapport au 
temps d'incubation. 

En laboratoire, KING, LANGDON ET 
COUNTS (1986) et KORNIUSHIN ET GLAU- 
BRECHT (2003) ont observé que les larves 
sont libérées apres 4 a 5 jours d'incuba- 
tion. Par contre, ENG (1979) estime, au 
cours d'une étude en milieu naturel, que 
la durée d'incubation s'éleve a 1 mois, 
mais doit dépendre des conditions envi- 
ronnementales. D'apres nos données, il 
est clair que cette durée est inférieure a 1 
mois, et probablement inférieure a 15 
jours. Ainsi, la premiere période d'incu- 
bation dériverait d'une phase d'émis- 


DuBoIs Er MARTÍNEZ-ORTÍ: Reproduction du Corbicula fluminea 


sion de spermatozoides, alors que la 
seconde résulterait de plusieurs phases 
d'émission, avec un asynchronisme 
marqué (Figs. 3 et 4A), ce qui est 
conforme aux observations de KRAEMER 
ET GALLOWAY (1986). 

En ce qui concerne le recrutement des 
juvéniles considérés (1,5-2,5 mm), la com- 
paraison des courbes de densité (Fig. 4B) 
indique que les maxima — en octobre en 
zone profonde et en mars dans les herbiers 
- sont de méme ordre de grandeur: res- 
pectivement 529 et 565 juv. / 100 ad., c'est- 
a-dire que l'on retrouve au mieux5a6 
juvéniles de cette classe de taille par adulte, 
alors que des milliers de larves sont incu- 
bées puis libérées. Ce nombre de juvéniles 
libérés par adultes a été estimé dans un lac 
américain (Lake Arlington, Texas) a 387 par 
jour durant la premiere période de repro- 
duction et a 320 par jour durant la seconde 
(ALDRIDGE ET MC MAHON, 1978). 

Aussi, ces courbes montrent cette 
opposition dans l'importance relative 
des deux phases de recrutement: la pre- 
miere phase est quasi-inexistante au 
niveau des herbiers et tres intense en 
zone profonde; la seconde phase, de 
forte amplitude, est moins intense dans 
la zone profonde que dans les herbiers. 
L'apparition tardive  (mi-décembre 
environ) et l'étendue de cette seconde 
phase de recrutement s'explique par la 
faible croissance individuelle annuelle 
(DUBOIS ET TOURENO, 1995). 

Il est probable que l'opposition de 
phase observée soit le résultat de migra- 
tion Ou dispersion des juvéniles, dont 
certains mécanismes sont connus. Nous 
savons en effet que les petits individus 
peuvent développer un filament de type 
« byssus » leur permettant de s'attacher, 
notamment a des particules qui peuvent 
étre mises en suspension dans l'eau 
(KRAEMER, 1979). Aussi, ces organismes 
ont la capacité de sécréter un filament 
muqueux, conduit par le siphon exha- 
lant, qui leur permet de flotter en pleine 
eau (en abaissant la densité corporelle). 
Cette sécrétion est stimulée, en aqua- 
rium, par le courant a partir de vitesses 
de 10 a 20 cm/s (PREZANT ET CHALER- 
WATT, 1984). 


La vitesse du courant dans le Canal 
Latéral est certes faible (7 cm/s en 
moyenne), mais en saison estivale, un 
grand nombre de bateaux de plaisance 
naviguent, créant un remous important 
qui peut étre a l'origine d'une disper- 
sion (Neck, 1986). Enfin, il a également 
été mis en évidence une forte capacité 
de la part de C. fluminea a se mouvoir 
sur le substrat dans un courent artificiel: 
250 cm/h pour des individus de taille 
inférieure á 2 mm (KRAEMER, 1979). 

Les résultats concernant la réparti- 
tion intra-habitat indiquent que celle-ci 
est constante tout au long de l'année 
aussi bien dans les herbiers qu'en zone 
profonde, en dépit des variations de 
densité dues au recrutement. Les indices 
de contagion calculés (Tableau II) sont 
du méme ordre de grandeur que ceux 
obtenus par DOWNING (1979) a partir de 
couples de données (S?, X) tirées de la 
littérature: b= 1,496 (a= 7,87) pour 
divers bivalves (138 données); b= 1,462 
(a= 5,24) pour les invertébrés benthiques 
en général (1462 données). 

Cependant, il semble que le degré 
d'agrégation soit plus important en 
zone profonde (b= 1,886) qu'au niveau 
des herbiers (b= 1,355). Nous avons en 
effet observé (non quantifié) que le sédi- 
ment récolté n'était pas toujours homo- 
gene (pour la granulométrie ou la 
teneur en débris organiques) le long 
d'un transept dans la zone profonde, ce 
qui peut augmenter le degré de conta- 
gion. A l'inverse, les zones a herbiers 
sont tres homogenes quant a la nature 
du sédiment et la densité de Vallisneria. 


REMERCIEMENTS 


Au Dr. Rafael Araujo du Museo 
Nacional de Ciencias Naturales de 
Madrid pour la révision critique du 
manuscrit et au Dr. Jean Noél Tourenq 
du Laboratoire d'Hydrobiologie a la 
Université Paul Sabatier de Toulouse, 
ainsi que pour son aide sur le terrain. 
Aux techniciens de la Section de Micro- 
scopie Electronique du S.C.S.I.E. de 
l'Universitat de Valencia, pour leur aide 
a lutilisation du M.E.S Hitachi S- 4100. 


Iberus, 28 (1), 2010 


RÉFÉRENCES BIBLIOGRAPHIQUES 


ALDRIDGE D.W. Er Mc MAHON R.F. 1978. 
Growth, fecundity and bioenergetics in a 
natural population of the Asiatic freshwater 
clam Corbicula manilensis Philippi, from North 
Central Texas. Journal of Molluscan Studies, 44: 
49-70. 

ALDRIDGE D.C. ET MULLER $. 2001. The Asiatic 
clam, Corbicula fluminea, in Britain: current sta- 
tus and potential impacts. Journal of Con- 
chology, 37(2): 177-183 

ARAUJO R., MORENO D. ET RAMOS M.A. 1993. 
The Asiatic clam Corbicula fluminea (Múller, 
1774) (Bivalvia: Corbiculidae) in Europe. 
American Malacological Bulletin, 10: 39-49. 

BAKER R., CLARKE K. Er HOWLETT D. 1999. The 
Asiatic clam Corbicula fluminea (Múller) new 
to the U.K. Transactions of the Norwich Natu- 
ralists” Society, 31: 70-76. 

BERAN L. 2006. Spreading expansion of Corbi- 
cula fluminea (Mollusca: Bivalvia) in the Czech 
Republic. Heldia, 6(3-4):187-192. 

Bi] DE VAATE A. 1991. Colonization of the Ger- 
man part of the river Rhine by the Asiatic 
clam, Corbicula fluminea Múller, 1774 (Pele- 
cypoda, Corbiculidae). Bulletin Zoólogisch 
Museum Universiteit van Amsterdam, 13: 13-16. 

Bi] DE VAATE A. ET GREJDANUS-KLASS M. 1990. 
The Asiatic clam, Corbicula fluminea (Múller, 
1774) (Pelecypoda, Corbiculidae), a new im- 
migrant in the Netherlands. Bulletin Zoólogisch 
Museum Universiteit van Amsterdam, 12: 173- 
WT 

BRITTON J.C. Er MORTON B. 1979. Corbicula in 
North America: the evidence reviewed and 
evaluated. In: J.C. Britton (ed.), Proceedings 
First International Corbicula Symposium, Texas 
Christian University, Fort Worth, Texas: 249-287. 

DEN HARTOOG C., VAN DEN BRINK F.W.B. ET 
VAN DER VELDE G. 1992. Why was the inva- 
sion of the River Rhine by Corophium 
curvispinum and Corbicula species so suc- 
cessful?. Journal of Natural History, 26: 1121- 
1129. 

DOMAGALA J., LABECKA A.M., PILECKA-RAPACZ 
M. ET MIGDALSKA B. 2004. Corbicula fluminea 
(O.F. Múller, 1774) (Bivalvia, Corbiculidae) 
a species new to the polish malacofauna. Fo- 
lia Malacologica, 12(3):145-148. 

DOWNING J.A. 1979. Aggregation, transforma- 
tion and the design of benthos sampling pro- 
grams. Journal of the Fisheries Research Board 
of Canada, 36. 1454-1463. 

Dubols C. 1995a. Biologie et démo-écologie d'une 
espece invasive, Corbicula fluminea (Mollusca: Bi- 
valvia) originaire d'Asie: Etude in situ (Canal La- 
téral a la Garonne, France) et en canal expéri- 
mental. These de Doctorat. Université de Tou- 
louse. 1-169 pp. Toulouse. 


10 


DuborIs C. 1995b. Corbicula fluminea: un mol- 
lusque opportuniste. Revue Agence de l'Eau 
Adour-Garonne, 63: 13-16. 

Dunors C. Er TOURENQ J.N. 1995. Étude préli- 
minaire dynamique des populations de Cor- 
bicula fluminea (Bivalvia: Corbiculidae) dans 
la zone profonde d'un canal de la région tou- 
lousaine (France). Hydroécologie appliquée, 7(1- 
2): 19-28.. 

Dubois C. ET ORIGNY R. 1998. Corbicula flumi- 
nea (O.F. Múller, 1774). Un mollusque bi- 
valve venant d'Asie, trouvé dans le Cher. 
Bulletin de la Société des Sciences Naturelles de 
Touraine, 1997: 49-59, 

ENG L.L. 1979. Population dynamics of the Asi- 
atic clam, Corbicula fluminea (Muller), in the 
Concrete-lined Delta Mendota Canal of Cen- 
tral California. In: J.C. Britton (ed.), Proceed- 
ings First International Corbicula Symposium, 
Texas Christian University, Forth Worth, Texas: 
39-68. 

GIRARDI H. 1989. Deux bivalves d'eau douce ré- 
cents pour la faune francaise (Mollusca, Bi- 
valvia). Bulletin de la Société d'Étude des Sciences 
Naturelles du Vaucluse, 2: 87-93. 

GRUET Y. 1992. Un nouveau mollusque bivalves 
pour notre région: Corbicula sp. (Heterodonta: 
Sphaeriacea). Bulletin de la Société des Sciences 
Naturelles de l'Ouest de la France, 14: 37-43. 

Isom B.G. 1986. Historical review of Asiatic 
clam (Corbicula) invasion and biofouling of 
waters and industries in the Americas. In: J.C. 
Britton (ed.), Proceedings 2"* Int. Corbicula Sym- 
posium, American Malacological Bulletin, spe- 
cial edition 2: 1-5. 

KING C.A., LANGDON C.J. Er COUNTS C.L. 1986. 
Spawning an early development Corbicula 
fluminea (Bivalvia: Corbiculidae) in laboratory 
culture. American Malacological Bulletin, 4: 81- 
88. 

KRAEMER L.R. 1979. Juvenile Corbicula: their 
distribution in the Arkansas River benthos. 
In: J.C. Britton (ed.), Proceedings First Inter- 
national Corbicula Symposium, Texas Christian 
University, Fort Worth, Texas: 89-97. 

KRAEMER L.R. ET GALLOWAY M.L. 1986. Larval 
development of Corbicula fluminea (Múller) 
(Bivalvia: Corbiculacea): an appraisal of its 
heterochrony. American Malacological Bul- 
letin, 4: 61-71. 

KRAEMER L.R., SWANSON C., GALLOWAY M. ET 
KRAEMER R. 1986. Biological basis of behav- 
iour in Corbicula fluminea U. Funtional mor- 
phology of reproduction and development 
and review of evidence for self-fertilitation. 
In: J.C. Britton (ed.), Proceedings of 2”* Inter- 
national Corbicula Symposium, American Mala- 
cological Bulletin, special edition 2: 193-201. 


DuBoIs Er MARTÍNEZ-ORTÍ: Reproduction du Corbicula fluminea 


LABECKA A.M., DOMAGALA J. ET PILECKA-RA- 
PACZ M. 2005. First record of Corbicula flu- 
minea (O.F. Miller, 1774) (Bivalvia: Corbi- 
culidae) in Poland. Folia Malacologica, 13(1): 
25-27. 

KORNIUSHIN A.V. ET GLAUBRECHT M. 2003. 
Novel reproductive modes in freshwater 
clams: brooding and larval morphology in 
Southeast Asian taxa of Corbicula (Mollusca, 
Bivalvia, Corbiculidae). Acta Zoologica, 84: 
2935310: 

MOUTHON J. 1981. Sur la présence en France et 
au Portugal de Corbicula (Bivalvia: Corbicu- 
lidae) originaire d'Asie. Basteria, 45: 109-116. 

MOUTHON J. 2001a. Life cycle and population 
dynamics of the Asian clam Corbicula flu- 
minea (Bivalvia: Corbiculidae) in the Saone 
River at Lyon (France). Hydrobiología, 452(1- 
3): 109-119. 

MOUTHON J. 2001b. Life cycle and population 
dynamics of the Asian clam Corbicula flu- 
minea (Bivalvia: Corbiculidae) in Rhóne River 
at Creys-Malville (France). Archiv fiir Hydro- 
biologie, 151(4): 571-589. 

MOUTHON J. 2003. Longitudinal and temporal 
variations of density and size structure of 
Corbicula fluminea (Bivalvia) populations in 
the Saóne and Rhóne rivers (France). Annales 
de Limnologie, 39(1): 15-25. 

NECK R.W. 1986. Corbicula in public recreation 
waters of Texas: habitat spectrum and clam- 
human interactions. In: J.C. Britton (ed.), Pro- 
ceedings 2”* International Corbicula Symposium, 
American Malacological Bulletin, 2: 179-184. 

PALOHEIMO J.E. ET VUKOV A.M. 1976. On mea- 
sures of aggregation and indices of contagion. 
Mathematical Biosciences, 30: 69-97. 


PÉREZ-QUINTERO J.C. 1990. Primeros datos so- 
bre la presencia de Corbicula fluminea Múller 
(Bivalvia, Corbiculidae) en España. Biome- 
tría. Scientia Gerundensis, 16(1): 175-182. 

PÉREZ-QUINTERO J.C. 2008. Revision of the dis- 
tribution of Corbicula fluminea (Múller 1744) 
in the Iberian Peninsula. Aquatic Invasions, 
3(3): 355-358. 

PREZANT R.S. ET CHALERMWAT K. 1984. Flota- 
tion of the bivalve Corbicula fluminea as a 
means of dispersal. Science, 225: 1491-1493. 

RAJAGOPAL S., VAN DER VELDE G. ET BI] DE VA- 
ATE A. 2000. Reproductive biology of the 
Asiatic clams Corbicula fluminalis and Cor- 
bicula fluminea in the river Rhine. Archiv fir 
Hydrobiologie, 149(3): 403-420. 

SWINNEN F., LEYNEN M., SABLON R., DUVIVIER 
L. Er VANMAELE R. 1998. The Asiatic clam 
Corbicula (Bivalvia: Corbiculidae) in Belgium. 
Bulletin de l'Institut Royal des Sciences Naturelles 
de Belgique, Biologie 68: 47-53. 

TAYLOR L.R. 1961. Aggregation, variance and 
the bivalve Corbicula fluminea as a means of 
dispersal. Science, 225: 1491-1493. 

TAYLOR L.R., WoIiwobD I.P. ET PERRY J.N. 1978. 
The density-dependence of special behav- 
iour and the rarity of randomness. Journal of 
Animal Ecology, 47: 383-406. 

TOURENO J.N. ET PUJOL J.L. 1991. Recherches sur 
la bio-écologie de Corbicula fluminea. Étude bib- 
liographique et observations dans la Garonne. 
Moyen de lutte pour en limiter les nuisances 
dans les circuits d'irrigation du Sud-Ouest. DDA 
du lot et Garonne. 1-312 pp. Toulouse. 


11 


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O Sociedad Española de Malacología Iberus, 28 (1): 13-22, 2010 


La Arqueomalacología: una introducción al estudio de los 
restos de moluscos recuperados en yacimientos arqueológi- 
COS 


Archaeomalacology: an introduction to the analysis of shellfish 
remains from archaeological sites 


Víctor BEJEGA GARCÍA, Eduardo GONZÁLEZ GÓMEZ DE AGUERO 
y Carlos FERNÁNDEZ RODRÍGUEZ* 


Recibido el 21-IX-2009. Aceptado el 23-11-2010 


RESUMEN 


La presencia de moluscos en yacimientos arqueológicos genera la necesidad de desarro- 
llar una metodología de estudio. Para ello, la Arqueomalacología nace como disciplina 
dentro de la Arqueozoología. A través del estudio de los diferentes elementos de los 
moluscos, aplicando diversas metodologías analíticas, podemos conocer una serie de 
aspectos importantes de la vida en las sociedades del pasado. La alimentación, las estra- 
tegias de recolección, la economía, la temperatura del mar, el comercio, el uso de ador- 
nos son algunos de los temas que pueden ser estudiados a través de ésta disciplina arque- 
ológica. 


ABSTRACT 


The presence of shellfish remains in archaeological sites has prompted the development of 
a methodology for their study. With this purpose, archaeomalacology emerges as a disci- 
pline within archaeozoology. A number of important aspects of life in ancient societies 
may be known by studying shellfish middens using analytical methods. The origin of food 
supplies, gathering strategies, economy, temperature of the sea water, trade, use of orna- 
ments and so on, are all subjects approached by a discipline such as archaeomalacology. 


INTRODUCCIÓN 


La Arqueomalacología es la rama de 
la Arqueozoología encargada del 
estudio de los restos de moluscos apare- 
cidos en contextos arqueológicos. Sus 
inicios se remontan al siglo XIX, como 
respuesta a la atención prestada a las 
importantes concentraciones de molus- 
cos de origen antrópico y cronología 
prehistórica presentes principalmente 
en las costas noroccidentales europeas, a 
las que se denominó con el término 


danés de kjoekkenmoeddings (literalmente 
“restos de cocina”). El estudio de estos 
depósitos supuso el desarrollo de la 
Arqueomalacología, cuyo objetivo es, 
como hemos señalado, el de analizar e 
interpretar los moluscos presentes en los 
yacimientos arqueológicos. 

En la Península Ibérica, los estudios 
arqueomalacológicos tienen sus inicios a 
finales del siglo XIX, como resultado de 
los hallazgos de depósitos de conchas 


* Universidad de León. Área de Prehistoria. Facultad de Filosofía y Letras. Campus de Vegazana s/n. 24007. 


IS 


Iberus, 28 (1), 2010 


en dos zonas geográficas diferenciadas: 
una localizada en torno al río Muge, en 
Portugal, y la segunda vinculada al río 
Sella, en Asturias. En ambos casos se 
trataba de grandes acumulaciones de 
moluscos marinos (denominadas conche- 
ros), adscritas al Mesolítico, que en el 
caso del Sella servirán para definir un 
determinado periodo cultural desarro- 
llado a inicios del Holoceno, el Astu- 
riense. El estudio de estos depósitos se 
ve motivado por dos aspectos: la infor- 
mación potencial que pueden aportar 
acerca del modo de vida de los grupos 
humanos que los originaron y la exce- 
lente capacidad de conservación que 
aportan a otros tipos de evidencias 
materiales. 

No obstante aquellos primeros epi- 
sodios, la Arqueomalacología no se con- 
solidará como parte habitual en los estu- 
dios arqueológicos hasta la década de 
los 70 del siglo XX. A partir de ese 
momento, se va a producir una evolu- 
ción metodológica que permite ampliar 
la información obtenida de los análisis 
de los concheros, destacando en este 
aspecto el trabajo que constituye la Tesis 
Doctoral de MORENO NUÑO (1994). 

Como hemos señalado, la Arqueo- 
malacología se ocupa del estudio de los 
restos de moluscos (incluyendo ocasio- 
nalmente crustáceos y equinodermos) 
que se documentan en los yacimientos 
arqueológicos, ya sea como elementos 
aislados o en acumulaciones (conche- 
ros). 

La aparición de moluscos aislados 
en contextos arqueológicos suele ser un 
hecho bastante habitual, tanto en yaci- 
mientos prehistóricos como históricos. 
Por otra parte, estos restos no son exclu- 
sivos de yacimientos costeros, no resul- 
tando extraordinario encontrarlos en 
sitios alejados del litoral, de manera 
especial -aunque no como norma taxa- 
tiva- a partir de época romana. La docu- 
mentación de estas evidencias, y espe- 
cialmente en el caso de restos alterados 
intencionalmente, no siempre debe rela- 
cionarse con una finalidad alimenticia, 
sino que han podido también utilizarse 
como material constructivo o incluso 
estar relacionados con la ornamentación 


14 


u otras actividades vinculadas con el 
ámbito ritual o simbólico. La presencia 
de colgantes, collares y otros elementos 
de adorno realizados con moluscos es 
conocida ya desde el Paleolítico Supe- 
rior, con especial relevancia de algunas 
especies como es el caso de Columbella 
rustica (Linné, 1758) (TABORIN, 1993a; 
1993b). 

El principal problema que se plantea 
con relación a estos materiales es su 
recuperación durante el proceso de 
excavación. En muchas ocasiones única- 
mente se recogen, en el mejor de los 
casos, los más llamativos o los mejor 
conservados; a lo que debe añadirse que 
la ausencia de un proceso de cribado 
provoca que fácilmente pasen desaperci- 
bidos este tipo de evidencias. 

El término conchero, como traducción 
del danés “kjoekkenmoedding”, es utili- 
zado por primera vez en España, según 
VEGA DEL SELLA (1923), por Aquilino 
Padrón en el Boletín de la Sociedad Geo- 
gráfica de 1877 refiriéndose a los depósi- 
tos de conchas existentes en las Cana- 
rias. No obstante, hay que tener pre- 
sente que un conchero no está formado 
exclusivamente por conchas de molus- 
cos, sino que puede incluir otros 
muchos tipos de materiales tanto orgá- 
nicos como inorgánicos (ictiofauna, 
macrofauna, cerámica, industria lítica, 
metal,...), con un muy buen estado de 
conservación, incluso aun cuando se 
trate de suelos ácidos, debido a la capa- 
cidad aislante del carbonato cálcico de 
los moluscos. Objeto de debate ha sido 
el porcentaje de conchas presente en un 
depósito necesario para su considera- 
ción o no como un conchero (una revi- 
sión del tema en BEJEGA GARCÍA, 2009a; 
2009b), aceptándose como valor medio 
un 30-50% de la composición total 
(MEIGHAN, 1980; BOWDLER, 2006). 

Estos depósitos (Figura 1) suelen 
corresponderse con basureros formados 
como resultado de la deposición conti- 
nuada de las partes duras de los molus- 
cos que han sido utilizados como ali- 
mento, aun cuando también pueden 
albergar algún elemento de esta misma 
naturaleza que haya tenido otra función. 
Así, en ocasiones existen problemas a la 


BEJEGA GARCÍA ET AL.: La Arqueomalacología 


es d 


ES 


hora de asignar alguno de los restos a 
una categoría tafonómica concreta, ya 
que su presencia puede reflejar causas 
bien diferenciadas (alimento, intrusión, 
adorno,...). 

Del mismo modo, también se han 
propuesto diferentes criterios a la hora 
de clasificar los tipos de concheros, 
basados en aspectos tales como la crono- 
logía (MEIGHAN, 1980), la densidad de 
materiales (STEIN, 1992; CLAASSEN, 1998) 
o la morfología (DUPONT, 2008), siendo 
este último el que consideramos más 
adecuado para definir las estructuras 
que habitualmente venimos estudiando 
(Tabla 1, Figura 2). 


METODOLOGÍA DE ANÁLISIS 


A la hora de realizar un estudio 
arqueomalacológico el primer aspecto 
básico a tener en cuenta es la aplicación 
de una metodología adecuada a las par- 


Figura 1. Conchero de un castro Galaico-romano ($ 


a e Lugo) 
Figure 1. Shell Miden of Galaico-roman hillfort (San Cibrao, Lugo, Spain) 


ticularidades del depósito, ya desde el 
proceso de recuperación de muestras. Es 
sin duda un requisito indispensable 
para la obtención de la mayor cantidad 
de información posible. Este proceso 
analítico se puede dividir en tres fases 
generales: 

Excavación y muestreo: en esta fase se 
obtiene el objeto de estudio. Durante la 
excavación se debe prestar especial 
atención a la aparición de moluscos ais- 
lados, así como a su correcta documen- 
tación estratigráfica. En caso de existir 
un conchero, es recomendable la aplica- 
ción de diferentes sistemas de muestreo 
(BEJEGA GARCÍA, 2009a; 2009b), desta- 
cando, por ser los más habituales, el de 
columnas y el selectivo. 

Análisis de las muestras: fase analítica 
propiamente dicha, que engloba básica- 
mente cuatro procesos: cribado (Figura 
3), triado, cuantificación y biometría. El 
cribado consiste en el lavado de las 
muestras utilizando un tamiz (en 


5 


Iberus, 28 (1), 2010 


Tabla I. Nomenclatura aplicada a los distintos tipos de conchero, según DUPONT (2008). 
Table I. Nomenclature applied to different types of'shell deposits, following DUPONT (2008). 


Depósitos en positivo: 


Conchero: depósito en relieve con un volumen superior a dos 
metros cúbicos. 


Depósito de conchas: depósito en relieve con un volumen infe- 
rior a dos metros cúbicos. 


Nivel o estrato de conchas: depósito con débil relieve (inferior 
a 10 cm desde el suelo) formado por conchas esparcidas. 


nuestro caso, teniendo en cuenta el tipo 
de evidencias que pueden recuperarse, 
solemos utilizar cribas con diámetro de 
malla no superior a 1 mm). El triado 
consiste en un proceso de selección de 
los restos conforme a la identificación 
taxonómica. Finalmente, se realiza una 
cuantificación y se toman una serie de 
valores: peso y biometría (siempre que 
se conserven las dimensiones máximas 
originales de altura, anchura y longi- 
tud). Asimismo, durante esta fase 
también se procede a la asignación de 
restos a las diferentes Categorías de 
Fragmentación (MORENO NUÑO, 1994; 
GUTIÉRREZ ZUGASTI, 2005), que serán la 
base para el cálculo de abundancias. 

Estimación de Abundancias: para 
conocer la importancia de cada una de 
las diferentes especies de moluscos en 
un conchero, se aplican tres Estimadores 
de Abundancias, con las adaptaciones 
necesarias para el cálculo de este tipo de 
evidencias arqueológicas (MORENO 
NUÑO, 1994; GUTIÉRREZ ZUGASTI, 2008; 
2005; BEJEGA GARCÍA, 2009a; 2009b): 
Número Mínimo de Individuos (N MI), 
Número de Restos (NR) y Peso. Los 
datos obtenidos por estos estimadores 
son básicos para realizar la posterior 
interpretación de los componentes 
malacológicos del conchero. 

Al tratarse de restos arqueológicos 
las partes blandas del animal no se han 
conservado, lo que indudablemente difi- 
culta una correcta identificación taxonó- 
mica. Este hecho se hace más evidente 
en aquellas especies que, aun dispo- 
niendo de ejemplares vivos, presentan 


1ó 


Depósitos en negativo: 


Nivel o Estrato de conchas en negativo: depósito con un 
relieve inferior a 10 cm desde el suelo. 


Nivel o Estrato de conchas en negativo de hábitat: depósito de 
menores dimensiones ubicado en zona habitada 


Depósito de conchas en hoyo de almacenamiento u otro 


dificultades para una asignación especí- 
fica, como las del género Patella. Estas 
deben ser identificadas en base a la mor- 
fología de la concha, no pudiendo consi- 
derar los criterios diagnósticos reconoci- 
dos en la rádula, la coloración u otros 
elementos clarificadores para la diferen- 
ciación interespecífica, tal y como 
señalan diferentes autores (p.e. CABRAL 
Y SILVA, 2003). Así mismo, los altos 
índices de fragmentación que presentan 
habitualmente estas piezas dificultan su 
identificación, impidiendo frecuente- 
mente, por ejemplo, la diferenciación 
entre Solen marginatus (Pulteney, 1799) y 
Ensis siliqua (Linné, 1758), o entre 
Mytilus galloprovincialis (Lamarck, 1819) 
y Muytillus edulis (Linné, 1758). Por este 
motivo, en arqueomalacología suele uti- 
lizarse la identificación por asociación, 
consistente en realizar una asignación 
específica de determinados restos con 
caracteres diagnósticos que sólo permi- 
ten su identificación a niveles taxonómi- 
cos supraespecíficos, mediante su aso- 
ciación con restos que sí son asignables 
a una especie concreta (MORENO NUÑO, 
1994). 


INFORMACIÓN POTENCIAL DE 
LOS ESTUDIOS ARQUEOMALACO- 
LÓGICOS. 


Con el fin de poder determinar el 
uso que han tenido los moluscos en un 
yacimiento, se realiza una evaluación 
tafonómica de los restos. Los grupos 
tafonómicos se establecen siguiendo dos 


BEJEGA GARCÍA ET AL.: La Arqueomalacología 


Depósito en relieve sobre el sustrato 


SÍ 
Depósito en relieve Depósito de escaso relieve, 
<10 em. de altura formado 
Mo por conchas dispersas 
< 2 


Conchero Depósito Nivel 


de conchas de conchas 


Figura 2. Tipos de conchero (DUPONT, 2006) 


Figure 2. Types of shell middens (DUPONT, 2006) 


criterios: el agente que deposita los 
restos (antrópico o no antrópico) y, en su 
caso, la función para la que fueron reco- 
lectados (GUTIÉRREZ ZUGASTI, 2005; 
2008). Su adscripción a cada uno de los 
diferentes grupos definidos sigue unos 
criterios generales que MORENO NUÑO 
(1994) adaptó de los establecidos por 
GAUTIER (1987): 

Restos alimenticios: aquellos que apa- 
rentemente han sido objeto de consumo 
por los humanos. 

Restos alterados: los que presentan 
alteraciones antrópicas para la confec- 
ción de utensilios, adornos... o aquellos 
con modificaciones naturales, pero cuya 
morfología ha propiciado su uso por los 
humanos. 

Restos de funcionalidad antrópica desco- 
nocida: restos recolectados una vez el 
animal está ya muerto y que no presen- 
tan ningún tipo de modificación que nos 
permita intuir su función. 

Restos intrusivos: los no acumulados 
intencionalmente por el ser humano. Se 
pueden diferenciar dos grupos: 

a) Intrusivos penecontemporáneos: 
depositados al mismo tiempo de forma- 
ción del depósito. 


No 


Depósito de fosa Depósito en fosa 


En hábitat En fosa 


Nivel de 


Nivel de h Depósito de 
conchas en fosa *Onchas conchas en 
en fosa de fosa 
habitación 


b) Intrusivos posteriores: introduci- 
dos con posterioridad a la formación del 
depósito. 

En la mayoría de las ocasiones los 
restos de moluscos se adscriben a la 
primera de las categorías taxonómicas 
citadas, la de restos alimenticios. No obs- 
tante, la información que podemos 
obtener mediante un estudio arqueoma- 
lacológico es muy diversa, abarcando 
planteamientos mucho más amplios que 
una mera enumeración de especies (GON- 
ZÁLEZ GÓMEZ DE AGUERO, 2009a; 2009b): 


Información alimenticia 

Como hemos señalado con anteriori- 
dad, los restos de moluscos recuperados 
en un yacimiento arqueológico se identi- 
fican mayoritariamente con desechos 
alimenticios (al igual que suele suceder 
con otros ecofactos, como macrofauna, 
ictiofauna o semillas). La importancia 
real de los moluscos en la dieta no es 
fácil de evaluar: un mayor volumen de 
conchas no implica un mayor aporte 
cárnico, ya que la proporción de carne 
de un molusco no se corresponde con el 
tamaño de la concha, influyendo un 
gran número de factores. Además hay 


WZ 


Iberus, 28 (1), 2010 


: > : S : da iS 
Figura 3. Proceso de cribado de una muestra 
Figure 3. Screening process with a 0.8 mm mesh 


a 


que tener en cuenta que su consumo 
está condicionado por toda una serie de 
valores sociales (gustos, modas, tabúes, 
etc.). Por otro lado, la proporción exis- 
tente entre las especies de moluscos que 
componen la muestra nos ofrece infor- 
mación sobre el tipo de sustratos explo- 
tados, y consecuentemente podemos 
evaluar las posibles áreas de captación y 
las técnicas de recolección utilizadas, 
tanto en el caso de moluscos marinos 
como en el de moluscos fluviales y 
terrestres (CALLAPEZ, 2002; STEIN, 1992). 

Los resultados del análisis biométrico 
pueden reflejar diferentes tendencias. 
Así, por ejemplo, una rápida reducción 
en las tallas de los individuos normal- 
mente apunta a una sobreexplotación del 
medio, ya sea debida tanto a un periodo 
de carestía como a una incipiente presión 
sobre el entorno (SHACKLETON, 1980; 
DAvIÉS, 2008), aunque también puede 
estar indicando la incidencia de la reco- 
lección en una zona concreta. 


18 


| 


pe 


con una malla AO Saa 7 


Información comercial 

La presencia de moluscos marinos 
en yacimientos arqueológicos de interior 
o la aparición de especies procedentes 
de ecosistemas muy distintos a los del 
entorno inmediato de un yacimiento 
litoral determinado suponen un proceso 
antrópico de traslado. 

Así, es frecuente la aparición de 
restos de moluscos en yacimientos 
romanos del interior, tales como Asturica 
Augusta (Astorga), Lucus Augusti (Lugo) 
o el campamento de la Legio VII en 
León, por citar algunos ejemplos en el 
noroeste de la Península Ibérica. Este 
tipo de evidencias nos indica la existen- 
cia de redes de comercio capaces de 
abastecer de moluscos a poblaciones 
relativamente alejadas del mar, respon- 
diendo a una demanda vinculada a los 
grupos sociales con mayor poder adqui- 
sitivo y que se especifica en una serie de 
especies entre las que cabe destacar, 
para época romana, Ostrea edulis (Linné, 


BEJEGA GARCÍA ET AL.: La Arqueomalacología 


Hierro 


Figura 4. Concha de Ostrea edulis procedente del castro de Neixón (Boiro, A Coruña) de la 11 Edad de 


Figure 4. Shell of Ostrea edulis from Neixón hillfort of 2" Iron Age (Boiro, A Coruña, Spain) 


1758) (Figura 4), pero también otras 
como Ruditapes decussatus (Linné, 1758), 
Osilinus lineatus (da Costa, 1778), etc. 

A la existencia de estas redes de 
comercio deberíamos sumar el desarro- 
llo de sistemas de conservación y trans- 
porte de los moluscos que permitiesen 
su consumo, evitando su deterioro. En 
este sentido, se ha venido planteando 
una doble hipótesis: la utilización de 
métodos de conserva, tales como la 
salazón, o el transporte de los animales 
vivos, sea en contenedores con agua O 
bien en recipientes sin agua aprove- 
chando la capacidad de algunas espe- 
cies para resistir varios días mediante la 
almacenada en su interior tras cerrar las 
valvas. 


Información paleoclimática 

La sensibilidad de los moluscos a los 
factores abióticos y bióticos hace que las 
diferentes especies puedan asociarse a 
ecosistemas muy concretos, cuyas carac- 
terísticas de temperatura y salinidad del 
agua, en el caso de los moluscos 
marinos, son conocidas. Partiendo del 
principio de que las especies no han 
variado sus preferencias de hábitat 
(SPARKS, 1980), podemos suponer que 
estos moluscos son representantes de un 
paleoclima determinado, lo que permite 
reconstruir las condiciones del medio en 
el que fueron recolectados, información 
a su vez de interés para valorar las con- 
diciones de vida de las sociedades del 
pasado. 


19 


Iberus, 28 (1), 2010 


De este modo, la existencia en los 
yacimientos gallegos del cambio de Era 
(siglos 1 a.C.- I d.C.) de especies como 
Stramonita haemastoma (Linné, 1758) que 
actualmente no se encuentran en las 
costas gallegas, nos indica un descenso 
de la temperatura de unos 2,5-3” C entre 
ese momento y la actualidad (RODRÍ- 
GUEZ LÓPEZ Y FERNÁNDEZ RODRÍGUEZ, 
1996). 

Igualmente, los moluscos terrestres 
son un claro indicador de las condicio- 
nes climáticas de la época de formación 
del yacimiento, así como de las caracte- 
rísticas del ecosistema que lo rodeaba 
(SPARKS, 1980). La presencia de especies 
como Pomatias elegans (Muller, 1774), 
por ejemplo, indica un entorno húmedo, 
de temperaturas suaves y con suelos 
calcáreos. 

Uno de los principales métodos apli- 
cados en la interpretación paleoclimá- 
tica es la comparación entre la distribu- 
ción actual de las especies y su presen- 
cia en los depósitos arqueológicos 
(GUTIÉRREZ ZUGASTI, 2005; 2008). Sin 
embargo, este tipo de análisis no resulta 
totalmente definitorio, debido a que 
gran cantidad de especies son euriter- 
mas y eurohalinas, siendo necesario 
complementarlo con otros estudios. 

Un sistema más preciso para la 
reconstrucción paleoclimática se basa en 
el análisis de los isótopos de oxígeno 
(016, 017 y O18) presentes en las 
conchas de los moluscos, cuya propor- 
ción varía en función de las fases cálidas 
y frías. Sin embargo, a pesar de tratarse 
de un análisis que ofrece una fiabilidad 
contrastada a la hora de definir grandes 
episodios climáticos, las dificultades 
inherentes al mismo, motivadas por la 
fluctuación de la temperatura del agua y 
su composición isotópica, limitan su 
aplicación ante eventos de menor 
entidad (SHACKLETON, 1980; STEIN, 1992; 
CLAASSEN, 1998). 


Estacionalidad 

La diferencia en el grosor de las 
líneas de crecimiento correspondientes a 
cada ciclo estival e invernal, especial- 
mente en los bivalvos, permite diferen- 
ciar cada uno de estos ciclos. Así, las 


20 


líneas más gruesas son propias de perío- 
dos estivales, con una mayor cantidad 
de alimento y unas mejores condiciones 
del agua, frente a las de menor espesor 
propias de ciclos invernales. 

En consecuencia, el análisis de las 
líneas de crecimiento teóricamente 
permite identificar la época de recolec- 
ción de los moluscos, pudiendo estable- 
cer si se trata de un recurso anual o esta- 
cional, lo que a su vez tiene implicacio- 
nes relativas a las estrategias de aprovi- 
sionamiento de las comunidades 
humanas y la importancia que estos 
recursos tendrían en su sistema econó- 
mico. 


Otros tipos de información 

Los moluscos recuperados en los 
yacimientos arqueológicos también 
pueden aportar información relacionada 
con otro tipo de actividades humanas 
no asociadas con la alimentación. Por 
ejemplo, la utilización de los murícidos 
para la elaboración de tintes (obtención 
del color púrpura) se tradujo en la exis- 
tencia de una actividad industrial, refle- 
jada en el establecimiento de factorías 
dedicadas a los procesos de recolección 
y transformación de estos moluscos. 

Otro tipo de evidencias indirectas 
que reflejan el uso y la importancia que 
tuvieron los moluscos para determina- 
das sociedades del pasado son las 
impresiones de determinadas especies, 
especialmente Cerastoderma edule (Linné, 
1758), como método de decoración de la 
cerámica, dando lugar a la denominada 
“cerámica cardial”. 

Por otro lado la existencia de colgan- 
tes, pulseras y otros objetos de adorno 
fabricados a partir de conchas, o la pre- 
sencia de las partes duras de los molus- 
cos en determinados contextos como los 
funerarios pueden ser evidencia del 
valor simbólico y/o ritual que también 
han recibido en ocasiones estos materia- 
les. 

En relación con los moluscos no 
marinos presentes en los yacimientos 
arqueológicos, aun cuando su trata- 
miento analítico de cuantificación no 
varía con respecto a aquellos, sí lo hace 
su interpretación. En el caso de los 


BEJEGA GARCÍA ET AL.: La Arqueomalacología 


moluscos fluviales, debe prestarse espe- 
cial atención a los ecosistemas caracte- 
rísticos de cada especie presente, ya que 
pueden indicarnos zonas de recolección 
y ayudar a valorar la intencionalidad o 
no de su recogida. Incluso la presencia 
de moluscos terrestres y micromoluscos 
puede ser indicativa de las condiciones 
de formación de un depósito de molus- 
cos fluviales o marinos, evidenciando la 
cantidad de tiempo que el mismo per- 
maneció expuesto hasta ser enterrado. 


CONCLUSIONES 


Los restos de moluscos presentes en 
los yacimientos son una fuente de infor- 
mación básica, tanto biológica como 
socio-cultural. El estudio sistemático de 
los depósitos con conchas permite 
obtener información sobre las estrate- 
gias de abastecimiento y la dieta de las 
poblaciones humanas, pero además son 
un elemento indispensable para estu- 
diar otros aspectos, tanto socio-econó- 
micos como paleoambientales. 

Uno de los errores que deben evi- 
tarse es el de interpretar la diversidad 
biológica de un periodo concreto a 
partir de las evidencias de origen arque- 
ológico. Los moluscos reflejan la presen- 
cia de determinadas especies en el 
medio en un momento concreto, pero no 
pueden considerarse como poblaciones 


BIBLIOGRAFÍA 


BEJEGA GARCÍA V. 2009a. Composición y me- 
todología de análisis de concheros aplicada 
a los castros litorales gallegos. Actas de las 1 
Jornadas de Jóvenes en Investigación Arqueoló- 
gica (J[A): Dialogando con la cultura material 
(Madrid, 2008), Tomo 1: 247-254. 

BEJEGA GARCÍA V. 2009b. El aprovechamiento de 
los recursos marinos en el Castro Grande de O 
Neixón (Boiro, A Coruña): un análisis arqueo- 
malacológico. Tesina de Licenciatura. Uni- 
versidad de León. Inédita. 

BOWDLER S. 2006 Mollusks and other shells. 
En Balme J. y Paterson A. (Eds.): Archaeol- 
ogy in Practice. A Student Guide to Archaeo- 
logical Analyses. Blackwell Publishing: 317- 
39 


biológicas, ya que los concheros son 
acumulaciones antrópicas en las que 
van a resultar muy importantes los fac- 
tores socio-culturales. De este modo la 
mayor presencia de Patella sp. en 
muchos castros galaicos puede que no 
esté reflejando la mayor abundancia de 
esta especie, sino una preferencia ali- 
menticia, bien por motivos sociales 
(simbolismo, gusto, tabú...) o económi- 
cos (fácil recolección, mayor rendi- 
miento, etc.). 

Sin duda alguna, para la correcta 
validación de estos estudios es necesaria 
una metodología de excavación, mues- 
treo e identificación muy precisa, ya que 
lo contrario supondría inevitablemente 
una pérdida de información. 

Pese a que la Arqueomalacología 
tiene un largo desarrollo en el tiempo 
como disciplina arqueológica, ha sido 
poco utilizada e incluso se ha infrautili- 
zado en muchas ocasiones. Actualmente 
está adquiriendo importancia debido 
tanto al avance metodológico de la dis- 
ciplina, así como por el conocimiento de 
la información potencialmente obtenible 
mediante su aplicación. El aprovecha- 
miento de los recursos marinos por las 
comunidades del pasado y su evolución 
a lo largo del tiempo es un aspecto a 
tener presente si queremos conocer y 
definir de forma correcta y global los 
modos de vida de aquellos grupos 
humanos. 


CABRAL J.P., COELHO F. Y DA SILVA A. 2003. 
Morphometric analysis of limpets from an 
Iron-Age shell midden found in northwest 
Portugal. Journal of Archaeological Science, 30: 
817-829. 

CALLAPEZ P. 2002. A malacofauna críptica da 
Gruta do Caldeirao (Tomar, Portugal) e as 
faunas de gastrópodes terrestres do Plisto- 
cénico superior e Holocénico da Extrema- 
dura portuguesa. Revista Portuguesa de Ar- 
queología, Vol 5 n*2 : 5-28 

CLAASSEN C. 1998. Shells. Cambridge Manuals 
in Archaeology, Cambridge University Press. 
266 pp. 

Davies P. 2008. Snails. Archaeology and Land- 
scape change. Oxbow Books, Oxford. 199 pp. 


21 


Iberus, 28 (1), 2010 


DUPONT C. 2006 La malacofaune de sites mé- 
solithiques et néolithiques de la facade at- 
lantique de la France: Contribution a l'eco- 
nomie et a l'identité culturelle des groupes 
concernés. British Archaeological Reports, In- 
ternational Series, 1571, v+439 pp. 

GAUTIER A. 1987. Taphonomic Groups: How 
and Why? Archaeozoologia, 1 (2): 47-52. 

GONZÁLEZ GÓMEZ DE AGUERO E. 2009a. Re- 
sultados obtenidos en el análisis de un con- 
chero: el caso de los castros litorales gallegos. 
Actas de las I Jornadas de Jóvenes en Investiga- 
ción Arqueológica (JLA): Dialogando con la cul- 
tura material (Madrid, 2008). Madrid: 255- 
262. 

GONZÁLEZ GÓMEZ DE AGUERO E. 2009b. El 
marisqueo en los castros galaico-romanos de la 
costa cantábrica gallega: el conchero de Punta 
Atalaya (San Cibrao, Cervo, Lugo). Tesina de 
Licenciatura. Universidad de León. Iné- 
dita. 

GUTIÉRREZ ZUGASTI I. 2005. La explotación de 
moluscos en la cuenca baja del río Asón (Canta- 
bria, España) a inicios del Holoceno (10.000- 
5.000 B.P.) y su importancia en las comunida- 
des humanas del Aziliense y del Mesolítico. Tra- 
bajo de Investigación de Doctorado. 
Universidad de Cantabria. Inédito. 

GUTIÉRREZ ZUGASTI I. 2008. La explotación de 
moluscos y otros recursos litorales en la región 
cantábrica durante el Pleistoceno Final y el Ho- 
loceno Inicial. Tesis Doctoral. Edición CD. 
Universidad de Cantabria. 

MEIGHAN C.W. 1980. Los moluscos como res- 
tos de alimentos en los sitios arqueológicos. 
En Brothwell D. y Higgs E. (Coord.): Ciencia 
en Arqueología. Fondo de Cultura Económica, 
México: 427-434. 


22 


MORENO NUÑO R. 1994. Análisis arqueomalaco- 
lógicos en la Península Ibérica. Contribución me- 
todológica y biocultural. Tesis Doctoral. Uni- 
versidad Autónoma de Madrid. Inédita. 

RODRÍGUEZ LÓPEZ C. Y FERNÁNDEZ RODRÍGUEZ 
C. 1996. Una aproximación al estudio de los 
yacimientos castreños del litoral galaico: di- 
mensiones ambientales y económicas. En Ra- 
mil Rego P., Fernández Rodríguez C. y Ro- 
dríguez Guitián (Coord.): Biogeografía Pleis- 
tocena-Holocena de la Península Ibérica. Xunta 
de Galicia: 363-375. 

SHACKLETON NJ. 1980. Los moluscos marinos 
en Arqueología. En Brothwell D. y Higgs E. 
(Coord.): Ciencia en Arqueología. Fondo de 
Cultura Económica, México: 418-426. 

SPARKS B.W. 1980. Los moluscos no marinos en 
la Arqueología. En Brothwell D. y Higgs E. 
(Coord.): Ciencia en Arqueología. Fondo de 
Cultura Económica, México: 406-417. 

STEIN K.J. (Ed.). 1992. Deciphering a Shell Mid- 
den. Academic Press, xix + 375 pp. 

TABORIN Y. 1993a. La parure en coquillage au 
Paléolithique. Gallia Préhistoire, supplément 
29, Centre National de la Recherche Scienti- 
fique. 544 pp. 

TABORIN Y. 1993b. Traces de faconnage et 
d'usage sur les coquillages perforés. Traces 
et fonction: les gestes retrouvés. Vol. 50. Collo- 
que International de Liége. Editions Études 
et Recherches Archéologiques de Université 
de Liege, 50(2): 255-267. 

VEGA DEL SELLA R.E. Conde de la 1923. El As- 
turiense. Nueva industria preneolítica. Memorias 
de la Comisión de Investigaciones Paleontológi- 
cas y Prehistóricas, 32 (serie Prehistórica 27), 
Museo Nacional de Ciencias Naturales, Ma- 
drid. 


O Sociedad Española de Malacología Iberus, 28 (1): 23-60, 2010 


Chauvetia 
Neogastropoda, Buccinidae) del área ibero-marroquí, con 


Las especies del género (Gastropoda, 


descripción de cuatro especies nuevas 


The species of the genus Chauvetía (Gastropoda, Neogastropoda, 
Buccinidae) in the Ibero-moroccan area, with the description of four 
new species 


Serge GOFAS* y Joan Daniel OLIVER** 


Recibido el 25-111-2010. Aceptado el 22-IV-2010 


RESUMEN 


Se revisan las especies del género Chauvetia Monterosato, 1884 encontradas en Andalu- 
cía, el sur de Portugal y Marruecos. En esta área se reconocen doce especies, cuatro de 
las cuales se describen como nuevas, dos con localidad tipo en el estrecho de Gibraltar y 
dos con localidad tipo en el noroeste marroquí. El estrecho de Gibraltar alberga la diver- 
sidad más alta para el género en Europa, apareciendo hasta ocho especies en la misma 
muestra. 


ABSTRACT 


The species belonging to the genus Chauvetia Monterosato, 1884 are revised for the area 
including Andalucía, southern Portugal and Morocco. Twelve species are recognized in 
this area, of which four are described as new, two with a type locality in the Strait of 
Gibraltar and two with a type locality in northwestern Morocco. The Strait of Gibraltar 
holds the highest diversity for the genus in Europe, with up to eight species potentially 
occurring in the same sample. 


INTRODUCCIÓN 


El género Chauvetia Monterosato, 
1884 comprende pequeños gasterópo- 
dos incluidos en la familia Buccinidae, 
cuya taxonomía resulta particularmente 
dificultosa debido al elevado número de 
especies y a la escasa diferenciación 
entre éstas. El género se conoce exclusi- 
vamente en el Mediterráneo y en la 
parte adyacente del Atlántico, entre el 
Canal de la Mancha y Senegal. La 
mayor diversidad de especies se conoce 
hasta la fecha entre Senegal (14 especies 
en el área de Dakar, OLIVER Y ROLÁN, 


2008) y las costas del Sáhara (9 especies, 
OLIVER Y ROLÁN, 2009). 

Existen varios trabajos dedicados a 
la taxonomía de este género en el Medi- 
terráneo, entre ellos los de TIBERI (1868), 
NORDSIECK (1976), MICALI (1999) y HER- 
GUETA, LUQUE Y TEMPLADO (2002). Sin 
embargo, son aún muchos los proble- 
mas taxonómicos sin aclarar. En el pre- 
sente trabajo se revisan las especies del 
entorno ibero-marroquí, con particular 
atención a las especies del estrecho de 
Gibraltar. 


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


** Alcorisa, 83-12C, E-28043 Madrid. 


Ss 


Iberus, 28 (1), 2010 


MATERIAL Y MÉTODOS 


El material estudiado para esta revi- 
sión incluye el recolectado por los 
autores en el área de estudio, así como 
el existente en las colecciones del 
Muséum National d'Histoire Naturelle 
de París (UNHN) y el Museo Nacional 
de Ciencias Naturales de Madrid 
(MNCN). Una parte sustancial de los 
datos usados para este trabajo se deriva 
de tres campañas organizadas por el 
MNHN. Los datos de las localidades se 
recogen en la Tabla L. 

La campaña Ceuta 1986 se desarrolló 
en mayo de 1986, y durante la misma se 
muestrearon una decena de puntos 
entre el nivel de bajamar y 40 m, situa- 
dos alrededor de la península de Ceuta, 
desde La Almadraba (Mediterráneo) 
hasta Benzú (Estrecho), principalmente 
mediante el uso de una aspiradora sub- 
marina de aire comprimido. Al llegar a 
puerto, las muestras eran inmediata- 
mente tamizadas en agua de mar y 
observadas bajo una lupa binocular, lo 
que posibilitó la observación en vivo de 
muchas especies, entre ellas nueve de 
Chauvetia. Para las especies de pequeño 
tamaño se realizó un dibujo del animal 
vivo, identificado por un número en la 
etiqueta del ejemplar correspondiente. 
El material recolectado está repartido 
entre el MNHN y el SMNH. 

La campaña Algarve 1988 se desa- 
rrolló en mayo de 1988 en la bahía de 
Sagres, próxima al extremo surocciden- 
tal de la Península Ibérica, y en junio del 
mismo año en la zona de la Ría Formosa 
y en algunos puntos cerca de Tavira, en 
la parte oriental de la costa del Algarve. 
Los métodos de muestreo eran básica- 
mente los mismos, y también se realiza- 
ron dibujos de los animales vivos. En la 
zona de Sagres, las especies de Chauvetia 
estaban representadas por un número 
de individuos mayor de lo habitual, 
mientras el género no se encontró en el 
muestreo de las lagunas de Ría 
Formosa, a pesar de realizarse un 
esfuerzo de muestreo similar. En esta 
campaña se pudieron observar en vivo 
cinco especies, todas ellas también pre- 
sentes en el material de Ceuta. 


24 


El material procedente de los bancos 
submarinos de Gorringe y Ampere, en 
el que se halló una especie de Chauvetia, 
fue recolectado durante la campaña Sea- 
mount 1 llevada a cabo en el B/O “Le 
Noroít” en 1987. 

Se ha examinado material proce- 
dente de diversas colecciones deposita- 
das en el MNCN como las colecciones 
de Hidalgo (1913), Azpeitia (1934) y 
Cobos (2000). En algunos casos se indica 
que estas colecciones incluyen otras 
como las de Gavala, Boscá o Sierra. 
También se ha estudiado material del 
MNCN aportado por José Templado y 
material procedente de las Campañas 
Fauna Ibérica 111 (1994) y IV (1996). 

Se estudió el material de la colección 
del segundo autor (JDO), procedente de 
material arrojado a las playas o de sedi- 
mentos recogidos a poca profundidad 
(entre 0 y 20 m). El material de Alborán 
se ha obtenido a partir de sacos de sedi- 
mentos depositados en su momento en 
el MNCN, y procede de las redes de 
barcos de pesca de coral rojo del puerto 
de Almería, que faenaron entre 1984 y 
1986 con el arte de la “barra italiana” 
entre 80 y 200 metros de profundidad 
(PEÑAS, ROLÁN, LUQUE, TEMPLADO, 
MORENO, RUBIO, SALAS, SIERRA Y GOFAS, 
2006). 

El material examinado se discrimina 
en el apartado correspondiente de cada 
especie, distinguiendo entre ejemplares 
recogidos vivos (ej.) y conchas vacías 
(c.); para cada lote se dan las dimensio- 
nes de los ejemplares adultos (con el 
labio formado) más pequeño y mayor; 
los juveniles se indican con j. Las proto- 
conchas fueron medidas bajo la lupa 
binocular equipada de un micrómetro 
ocular, a un aumento de 40x, en vista 
apical, según las pautas indicadas en la 
Figura 1. 

Los ejemplares destinados a ser foto- 
grafiados en microscopía electrónica de 
barrido fueron limpiados en un baño de 
lauril sulfato de sodio (un detergente de 
pH neutro) al 10%, sometidos a una 
breve (pocos segundos) exposición a 
ultrasonidos en agua, montados en los 
portaobjetos con un pegamento conduc- 
tor y metalizados con oro. Las observa- 


GOFAS Y OLIVER: Las especies de Chauvetía en el área ibero-marroquí 


Tabla I. Localidades de Ceuta, del Algarve y de los bancos lusitanos muestreadas en las campañas 
del MNHN, y de la costa mediterránea ibérica muestreadas en las campañas del MNCN (Fauna 


Ibérica). 


Table 1. Localities from Ceuta, Algarve and the lusitanian banks sampled during MNHAN campaigns, 
and those from the Iberian mediterranean coast sampled during MNCN campatgns (Fauna Ibérica). 


Localidad 
Campaña Ceuta 1986 


Benzú 

Playa Benítez 

Punta Almina 

Punta del Saudiño 
Ensenada de Sarchal 
Ensenada de La Almadraba 


Campaña Algarve 1988 


Sagres, Ponta da Baleeira 
Sagres, Baía da Baleeira 
Sagres, Boía da Baleeira 
Sagres, Boía de Sagres 
Sagres, Pontal dos Corvos 
Sagres, Ponta dos Caminhos 
Tavira, Pedra do Barril 
Tavira, Cabanas 


Campaña Seamount 1 


Banco Gorringe, DWOA4 
Banco Gorringe, DWO5 
Banco Gorringe, DWO6 
Banco Gorringe, DWO8 
Banco Gorringe, DEO9 
Banco Gorringe, DW32 
Banco Gorringe, DW33 
Banco Ampere, DW92 
Banco Ampere, DE98 


Fauna Ibérica 


Punta Foradada (Mallorca) 

Cabo Nati (Menorca) 

Punta Galera (1. Espardell, Ibiza) 
Bleda Mayor (Ibiza) 

Cala Eubarco (Ibiza) 

Placer Barra Alta N. (Columbretes) 
Norte Isla Columbrete Grande 

Isla de Alborán 


Coordenadas 


a SA 
35" 54,6" N— 05* 20,0 W 
35" 54,1” N—05* 16,5 W 
35* 54,1 N— 05* 18,0 W 
39 09,4 N=05* 17,0 W 
d0 02 0 Ni 057 19,00 


37* 00,3" N— 08* 55,5" W 
37* 00,7” N— 08* 55,0" W 
37* 00,8" N— 08* 55,0" W 
36% 59,0 N—08* 56,3 W 
3101/3007 50,3W 
37" 02,0" N—08* 52,0" W 
37 02,4 N—0739,7'W 
06 NOS 


36" 32 N—11% 34 W 
30:32 N= 11530 W 
36” 30'N—118 38" W 
36" 28'N—-119 377'W 
30 91 N=117 30 W 
30 JN T1P 35 
36" 31" N—11% 34W 
OSI 12053 We 
SOS IN ZAS 


39494 0237: E 
40* 43'N—03* 49' E 
38 47'N-01*28'E 
39 50,N=01"09'E 
39 04"N:01221'E 
39" 49" N—00* 32' E 
39 54N—01*41'E 
35 58 N—02* 58" W 


Hábito Profundidad (m) 
Infralitoral rocoso 0-4 
Cascajo bioclástico 15-25 
Acantilado rocoso 25-40) 
Acantilado rocoso 17-35 
Infralitoral rocoso 0-3 
Fondo blando enfangado 20-36 
Bajo de acantilado rocoso 17-23 
Fondo rocoso 3-15 
Intermareal rocoso 
Cascajo bioclástico 20-30 
Bajo de acantilado rocoso 17-22 
Cascajo bioclástico 23-33 
Fondo rocoso enfangado 25 
Fondo rocoso enfangado 14 
Piedras, bioclastos y rodolitos 93-96 
Fondo con bioclastos 180 
Fondo de bioclastos con ofiuras 250 
Fondo endurecido con bioclastos gruesos 470-485 
Fondo con arena bioclástica 350-360 
Fondo con rocas y algas 54-62 
Fondo con rocas y algas 35-70 
Fondo con arena bioclástica gruesa y rodolitos 117-129 
300-325 
Cascajo bajo un extraplomo 24 
Arena y cascajo entre Posidonia 31 
Sedimento 35 
Detrítico costero 45 
Roca con coralígeno y detrítico 44 
Arena gruesa y restos calcáreos 32 
Coralígeno rocoso, detrítico 4] 
Fondo rocoso 35-37 


25 


Iberus, 28 (1), 2010 


ciones y micrografías se realizaron en un 
microscopio electrónico de barrido JEOL 
JSM-840, ampliado con un sistema de 
adquisición digital de imágenes. 

Las descripciones están basadas en 
el material examinado de la zona de 
estudio, por lo que pueden no tener en 
cuenta elementos de variabilidad que se 
manifiesten fuera de esta zona. Las 
especies asignadas al género Chauvetia 
presentan una gran homogeneidad mor- 
fológica, por lo que se hace primero una 
diagnosis de los caracteres genéricos 
compartidos entre todas las especies, 


PARTE TAXONÓMICA 


que no serán repetidos en las descripcio- 
nes de cada especie. 


Abreviaturas 


CINZ: Código Internacional de Nomen- 
clatura Zoológica. 

Col.: colección. 

JDO: Joan Daniel Oliver. 

SG: Serge Gofas. 

SMN H: Swedish Museum of Natural 
History, Estocolmo 

USNM: National Museum of Natural 
History (Smithsonian). 


Chauvetia Monterosato, 1884 


Chauvetia Monterosato, 1884. Nom. Gen. Spec.: 137. [Nomen novum para Nesaea Risso, 1826; especie 
tipo: Nesaea mamillata Risso, 1826, ver notas nomenclaturales]. 

Nesaea Risso, 1826 (non Lamarck, 1812, nec Leach, 1814). Hist. Nat. Eur. Mérid., iv: 211. [Especie 
tipo por designación subsiguiente de Bucquoy, Dautzenberg y Dollfus, 1883: Nesaea mamillata 


Risso, 1826, ver notas nomenclaturales]. 


Lachesis Risso, 1826 (non Daudin, 1803, nec Wood, 1804). Hist. Nat. Eur. Mérid., iv: 223. [Especie 
tipo por monotipia: Lachesis mamillata Risso, 1826]. 

Donovania Bucquoy, Dautzenberg y Dollfus, 1883 (non Donovania Leach, 1814). Moll. Roussillon, 
vol. 1: 85, 112. [Nomen novum para Lachesis Risso, 1826 y Nesaea Risso, 1826]. 

Folineaea Monterosato, 1884. Nom. Gen. Spec.: 136. [Especie tipo por designación subsiguiente de 
Crosse, 1885: Buccinum lefebvrii Maravigna, 1840]. 

Adansonia Pallary, 1902. J. Conchyl. 50: 13. [Nomen novum para Folineaea Monterosato, 1884, non 


Folinia Crosse, 1868]. 


Syntagma lredale, 1918. Proc. Malac. Soc. London, 13: 34-35. [Nomen novum para Donovania 


Bucquoy, Dautzenberg y Dollfus 1883]. 


Chauvetiella Nordsieck, 1968. Eur. Gehiuseschn.: 137. [Especie tipo por designación original: Chau- 


vetiella vulpecula (Monterosato, 1884)|. 


Donovaniella Nordsieck, 1968. Eur. Geháuseschn.: 136. [Especie tipo por designación original: 


Donovaniella minima (Montagu, 1803)|. 


Diagnosis: La concha es fusiforme, 
sólida, con espira alta y un tamaño 
generalmente comprendido entre 5 y 10 
mm en los adultos (aunque la mayor 
especie conocida, Chauvetia gigantissima 
Oliver y Rolán, 2009, de la plataforma 
continental del Sáhara, alcanza hasta 20 
mm). La protoconcha consta por lo 
general de poco más de una vuelta, 
cuyo núcleo hemisférico alcanza alrede- 
dor de la mitad del diámetro de la 
vuelta siguiente. La escultura de la pro- 
toconcha puede incluir cordoncillos 
espirales aplanados, costillas axiales o 
una combinación de ambos. La transi- 


26 


ción con la teleoconcha es difícil de 
apreciar; se considera el punto en el que 
las costillas axiales, después de aproxi- 
marse entre sí al final de la primera 
vuelta, se vuelven a distanciar. Un poco 
antes de ese paso, todavía pueden apre- 
ciarse claramente los cordoncillos espi- 
rales de la protoconcha y se inician los 
primeros cordones de la teleoconcha. 

La teleoconcha tiene una escultura 
de cordones espirales y costillas axiales, 
pudiendo variar el aspecto y el número 
de estos elementos. Generalmente, hay 
entre 3 y 5 cordones espirales en las 
vueltas de la espira y 15-20 en la última 


GOFAS Y OLIVER: Las especies de Chauvetía en el área ibero-marroquí 


Figura 1. Medidas tomadas en las protoconchas. 1: diámetro máximo, medido entre el medio de la 
zona de transición y el borde opuesto de la protoconcha; 2: diámetro del núcleo, 3: diámetro de la 
primera media vuelta, ambos medidos en un eje perpendicular al máximo diámetro de la proto- 
concha. Figura 2. Vista parcial de la rádula de Chauvetia mamillata (Risso, 1826), especie tipo del 


género. Ejemplar de Sagres 17-23 m. 


Figure 1. Measurements taken on the protoconchs. 1: maximum diameter, measured between the transition 
zone and the opposite side of the protoconch; 2: diameter of the nucleus; 3: diameter of the first half+wbhorl, 
both measured along an axis perpendicular to the maximum diameter. Figure 2. Partial view of the radula 
of Chauvetia mamillata (Risso, 1826), type species of the genus. Specimen from Sagres 17-23 m. 


vuelta, siendo los que cubren el canal 
sifonal mucho más finos que los de la 
parte adapical de la última vuelta. La 
terminación de las costillas forma 
amplias ondulaciones en la sutura. La 
abertura es ovalada, continuada en el 
lado abapical por un canal sifonal muy 
corto y ancho. En los ejemplares 
adultos, el labio está engrosado por su 
lado externo, pero esta variz no está 
separada de la parte previa de la última 
vuelta por ninguna discontinuidad. 

El animal es relativamente pequeño 
con relación a la concha, con un pie alar- 
gado, truncado por delante y redonde- 
ado en su extremo posterior, que se 
estrecha progresivamente hacia atrás. La 
parte posterior del pie está cubierta por 
un Opérculo ovalado bien desarrollado, 
con núcleo subterminal. La cabeza 
carece de morro y es estrecha y bifur- 
cada, con los ojos situados lateralmente 


en cada rama y los tentáculos cefálicos 
delgados. El sifón extendido es cilín- 
drico y relativamente largo. El color de 
fondo del animal es blanco o amari- 
llento con reflejos irisados, y puede 
tener un patrón de pigmentos negros 
cuya presencia y configuración es 
propia de cada especie. 

La rádula se ha estudiado en ejem- 
plares del Algarve de Chauvetia mami- 
llata (Fig. 2). Es muy pequeña con rela- 
ción al tamaño del animal, unos 200 um 
de largo por menos de 40 um de ancho, 
y cada hilera de dientes consta de un 
diente central y un par de dientes latera- 
les. El diente central es casi cuadrado, 
con una cúspide axial, bastante estrecha, 
aguzada y situada en el mismo plano 
que la base. Los dientes laterales son el 
doble de anchos que los centrales y su 
borde lleva tres cúspides muy fuertes, 
curvadas, la interna más pequeña y la 


zi 


Iberus, 28 (1), 2010 


externa más grande. Por lo que se 
conoce en esta especie (véase también 
HERGUETA ET AL., 2002) y otras (C. 
brunnea en WOODWARD, 1899; varias 
especies africanas en PEÑAS Y ROLÁN, 
2008) es de poca utilidad para la discri- 
minación de especies afines y no se ha 
estudiado sistemáticamente en este 
trabajo. 

La puesta se conoce en el caso de C. 
mamillata (HERGUETA ET AL., 2002, y 
nuestro material de Sagres) y consta de 
una pequeña (menos de 1 mm) cápsula 
semitransparente en forma de copa, 
sujeta al sustrato por un pedúnculo muy 
corto. La parte superior está truncada, 
limitada por un reborde marcado, y fun- 
ciona como una tapadera que se abre en 
el momento de la eclosión. Esta puesta 
contiene un solo huevo, que se desarro- 
lla en su interior hasta formar una pro- 
toconcha de algo más de una vuelta. 
Este tipo de puesta y de protoconcha 
indica un desarrollo directo sin fase 
pelágica, pero, a la vez, estas cápsulas 
muy ligeras y generalmente fijadas a las 
algas constituyen un potente medio de 
dispersión pasiva en el caso de que las 
algas sean arrancadas por el oleaje y 
floten a la deriva. Considerando la 
homogeneidad morfológica que se 
observa entre todas las especies, consi- 
deramos que este tipo de desarrollo 
directo se puede inferir para las demás 
especies estudiadas. 

Notas nomenclaturales: La nomencla- 
tura supraespecífica de este grupo tiene 
una historia de una complejidad poco 
común. En ocasiones, se ha incluido este 
género en la subfamilia Donovaniinae 
Casey, 1904 (= Lachesinae L. Bellardi, 
1877). Ambos nombres son inválidos 
según el artículo 39 del CINZ, al ser los 
géneros tipo homónimos más recientes; 
sin embargo, al no estar en uso, no 
procede establecer un nombre de susti- 
tución basado en un nombre genérico 
válido. 

En cuanto a la nomenclatura gené- 
rica, Risso (1826) no deja claro cuáles 
son los caracteres que diferencian Lache- 
sis de Nesaea, ni hace ninguna compara- 
ción entre los dos géneros; además, 
existen ciertas incoherencias entre las 


28 


diagnosis genéricas y las de las especies 
incluidas. Una de las diferencias litera- 
les entre las dos diagnosis genéricas está 
en la espira, que se describe como muy 
elevada en Lachesis y moderadamente 
elevada en Nesaea, si bien esta diferencia 
no es obvia en las figuras. La otra dife- 
rencia literal atañe a la descripción del 
refuerzo del labio (“peritrema” en el 
vocabulario de Risso), ausente en Lache- 
sis y sencillo y ligeramente desviado 
hacia dentro en Nesaea; en ambos casos 
se trataría de los caracteres de los juve- 
niles de las figuras 65 y 67 de Risso 
(1826). Sin embargo, tanto en las diagno- 
sis de Nesaea granulata como de N. mami- 
llata este autor indica pliegues (dentícu- 
los) en el lado izquierdo (interno) del 
peritrema (labio), lo cual solamente 
aparece en su figura 69 de N. mamillata, 
así como en el lectotipo (ARNAUD, 1978; 
OLIVER Y ROLÁN, 2009: figuras 1-2) de 
esta especie. 

TIBERI (1868) incluye una diagnosis 
revisada para cada uno de estos dos 
géneros, pero añade confusión al tener 
un concepto de Nesaea granulata obvia- 
mente distinto al de Risso (1826). Las 
diferencias literales entre las diagnosis 
genéricas se resumen a una espira bas- 
tante elevada, con vueltas algo conve- 
xas, sutura poco profunda y labio senci- 
llo (es decir, sin variz) en Lachesis, y 
espira moderadamente elevada, con 
vueltas redondeadas, sutura profunda y 
labio varicoso en el lado externo en 
Nesaea. Quizás sea más aclaratorio el 
reparto de especies que hizo MONTERO- 
SATO (1884), teniendo a mano la colec- 
ción de Tiberi que había comprado, y 
usando Donovania (= Lachesis) para el 
grupo de D. mamillata, y Chauvetia (= 
Nesaea) para C. granulata sensu Tiberi, o 
sea las especies de forma más tosca. Esta 
distinción carece de importancia, dado 
que Donovania se considera sinónimo de 
Chauvetia. 

Existe, además, un problema nomen- 
clatural acerca de la especie tipo de 
Chauvetia. La designación por MONTE- 
ROSATO (1884) de Buccinum candidissi- 
mum Philippi, 1836 es inválida, al no tra- 
tarse de una de las especies original- 
mente incluidas por Risso (1826) en el 


GOFAS Y OLIVER: Las especies de Chauvetía en el área ibero-marroquí 


género sustituido Nesaea. Crosse (1885) 
designó formalmente “Nesaea granulata 
Tiberi”, lo cual se debe entender como 
una referencia al concepto que TIBERI 
(1868) tuviera de Nesaea granulata. 
Siendo así, deja de ser una de las espe- 
cies originalmente incluidas en Nesaea y, 
por lo tanto, no se puede aplicar el artí- 
culo 69.2.4 del CINZ sobre especies tipo 
con identificación errónea, ni aceptar 
este acto como designación válida. Sin 
embargo, la designación anterior de 
Donovania minima (Montagu, 1803) como 
especie tipo de Donovania por parte de 
BUCQUOY, DAUTZENBERG Y DOLLFUS 
(1883) se aplica también a Nesaea (y por 
consecuencia, a Chauvetia) en virtud del 
artículo 67.8 del CIZN y cumple con los 
requisitos establecidos en el artículo 
69.2.2 al incluir en la sinonimia de D. 
minima una, y una sola (N. mamillata), de 
las especies originalmente incluidas en 
Nesaea. El propio género Donovania 
siendo nomen novum simultáneamente 


para dos géneros distintos (Nesaea y 
Lachesis), escapa a las especificaciones 
del CINZ; de cualquier modo la cues- 
tión es irrelevante, puesto que este 
nombre también está preocupado. 

El lectotipo (ARNAUD, 1978) de 
Nesaea granulata Risso, 1826 es un ejem- 
plar juvenil de color uniforme que asig- 
namos a C. mamillata, con localidad tipo 
implícita en los alrededores de Niza. Sin 
embargo, BUCQUOY ET AL. (1883) y MON- 
TEROSATO (1884), siguiendo el criterio de 
TIBERI (1868), consideran que N. granu- 
lata es una especie extremadamente rara 
y muy distinta, e indican una distribu- 
ción en el sur de Italia (golfo de 
Nápoles), las costas norteafricanas y el 
Adriático. Siendo así, nos parece verosí- 
mil la sugerencia por parte de MICALI 
(1999: 57) de que Chauvetia granulata 
sensu Tiberi sea lo mismo que C. ven- 
trosa Nordsieck, 1976, un endemismo 
centromediterráneo que no se encuentra 
en las costas ibéricas. 


Chauvetia mamillata Risso, 1826 (Figs. 2-14, 23-26, 97-98) 


Nesaea mamillata Risso, 1826. Hist. Nat. Eur. Mérid., iv: 223-224, lám. 5, fig. 69. [Localidad tipo: 
Alpes-Maritimes, Francia]. 

Nesaea granulata Risso, 1826. Hist. Nat. Eur. Mérid., iv: 223, lám. 5, fig. 67. [Localidad tipo: Alpes- 
Maritimes, Francia]. 

Lachesis mamillata Risso, 1826. Hist. Nat. Eur. Mérid., iv: 211, lám. 5, fig. 65. [Localidad tipo: Alpes- 
Maritimes, Francia]. 

Fusus turritellatus Deshayes, 1835. Exp. Sc. Morée, Moll.: 174, lám. 19 figs. 28, 45. [Localidad tipo: 
Peloponeso, Grecia]. 

Buccinum minimum Philippi, 1836 (non Montagu, 1803). En. Moll. Siciliae, vol. 1: 222. [Localidad 
tipo: Palermo y Catania, Sicilia]. 

Donovania minima var. attenuata Bucquoy, Dautzenberg y Dollfus, 1883. Moll. Roussillon, vol. 1: 
113-114. 

Donovania minima var. submamillata Bucquoy, Dautzenberg y Dollfus, 1883. Moll. Roussillon, vol. 
as lám 15 't1e 30. 

Donovania minima var. insignis Bucquoy, Dautzenberg y Dollfus, 1883. Moll. Roussillon, vol. 1: 114. 


Material tipo: Nesaea mamillata: lectotipo (Arnaud, 1978), MNHN 6600 y 4 paralectotipos, MNHN 
5599; Nesaea granulata: lectotipo (Arnaud, 1978), MNHN 6601; no se han encontrado los tipos de 
Lachesis mamillata, pero fueron examinados junto con los dos anteriores por Monterosato (1872: 48), 
quién los consideró coespecíficos. 

Material estudiado: Baleares: Punta Foradada (Mallorca), 4 c. + 393. (6,1 x 2,8 mm), MNCN; Santa 
Ponca ( Mallorca), 6 c. (6,8 x 2,8 mm) + 2j., MNCN; Menorca (Mahón), 8 c. (5 morfotipo no mamilar, 
7,2 x 2,7 mm; 3 morfotipo mamilar, 6,0 x 2,5 mm) + 1j., col. Hidalgo, MNCN; Menorca (cabo Natí), 
2 c. (7,2 x 3/0 mm) + 1j., MNCN,; Ibiza (Cala Eubarco), 1 c. (6,0 x 2,6 mm) + 25j., MNCN; Ibiza 
(Punta Galera), 1 c. (7,5 x 2,9 mm) + 9j., MNCN. Levante ibérico: Columbretes (Placer de la Barra 
Alta), 2 c. (morfotipo mamilar, 6,5 x 2,8 mm), MNCN; Norte Columbrete Grande (47 m), 33., MNCN; 
Valencia, 17 c. (11 morfotipo mamilar, 6,0 x 2,3 — 7,3 x 3,0 mm, 6 morfotipo no mamilar, 6,3 x 2,9 


29 


Iberus, 28 (1), 2010 


mm) + 4 j., col. Rosselló en col. Azpeitia, MNCN; Valencia, 3 c. (morfotipo no mamilar, 6,3 x 2,5 
mm), col. Boscá en col. Azpeitia, MNCN; Cullera (15 m); 80 c. + 200 j. (todas morfotipo mamilar, 
5,7 x 2,4 — 7,3 x 3,0 mm), col. JDO; Denia (arrojadas a la playa), 140 c.+ 100 j. (morfotipo mamilar 
40 c. +403., 5,8 x 2,4 -— 7,5 x 3,2 mm, morfotipo oscuro 50 c. + 30j., 5,6 x 2,6 — 7,1 x 3,4 mm, morfo- 
tipo con bandas 50 c. + 303., 5,5 x 2,3 — 7,0 x 2,6 mm), col. JDO; Jávea (arrojadas a la playa), 11 c. + 
173. (3 ej. morfotipo mamilar; 11 morfotipo con bandas, 5,1 x 2,0 mm — 6,0 x 2,6 mm + 14j.), col. 
JDO; cabo de Palos, 27 c. (15 morfotipo mamilar, 5,3 x 2,3 — 7,1 x 3,2 mm + 14 j.; 12 morfotipo no 
mamilar, 5,8 x 2,4 — 6,6 x 2,7 mm), MNCN, cabo de Palos, 43 c., (morfotipo mamilar, 5,4 x 2,4 — 6,9 
x 3,7 mm), col. Hidalgo, MNCN,; cabo de Palos, 5 c. (5,0 x 2,3- 5,7 x 2,6 mm), col. Azpeitia, MNCN; 
Cartagena (morfotipo no mamilar), 2 c. (6,2 x 2,5 mm), col. Hidalgo, MNCN. Mar de Alborán y 
estrecho de Gibraltar: Cabo de Gata, Punta de Loma Pelada (15 m), 3 j. (morfotipo mamillar); 
Almería, 10 c. (morfotipo con bandas, 7,3 x 3,2 —7,9 x 3,4 mm), col. A. Sierra, MNCN; Almería (40 
m), 506 c. + 22. (486 morfotipo con bandas, 4,8 x 2,3- 6,4 x 2,7 mmy 40 c. + 22j. morfotipo mamilar, 
6,1 x2,6-7,9 x 3,5 mm), col. A. Sierra en col. Cobos, MNCN; Dorsal de la isla de Alborán (35 58' 
N, 02” 58" W), 15 c. + 45j. (morfotipo de Alborán, 5,5 x 2,5-7,5 x 3,1 mm), MNCN; La Herradura 
(Granada), 2 ej. (morfotipo con bandas, 7,3 x 3,1 — 7,4 x 3,1 mm), col. SG, MNHN; Calahonda 
(Málaga), 21 c. (morfotipo con bandas, 5,3 x 2,3 — 6,5 x 2,9 mm), col. SG 1978, MNHN. Sotogrande 
(Cádiz), bajamar, 13 ej. (5,5 x 2,4 - 6,3 x 2,7 mm) + 34 c. (5,4 x 2,3 — 6,6 x 2,9 mm), col. SG, MNHN; 
bahía de Algeciras, 4 c. + 13. (morfotipo con bandas, 6,6 x 2,9 -7,1 x 3,4 mm), MNCN, Tarifa, Torre 
de la Peña, 32 c. (5,8 x 2,6-7,7 x 3,4 mm) + 2j., col. 5SG, MNHN; Cádiz, 12 c. (morfotipo con bandas, 
5,6 x 2,5 6,5 x 2,9 mm) col. Hidalgo, MNCN; Cádiz, 5 c. (morfotipo con bandas, 5,1 x 2,5-7,2 x 3,1 
mm), col. Azpeitia, MNCN; M' diq (Marruecos), 27 c. + 4 ej. (10 morfotipo algo más claro, resto mor- 
fotipo con bandas; 5,2 x 2,3 — 6,4 x 2,6 mm), col. SG, MNHN; Ceuta, Almadraba, 20-36 m, 3 c. (4,9 
x 2,2 — 6,3 x 2,8 mm); Ceuta, Sarchal, 0-3 m, 4 ej. (morfotipo con bandas, 5,9 x 2,8 — 6,5 x 3.0 m) + 10 
j.; Ceuta, Punta Almina, 25-40 m, 107 ej. + c. (42 morfotipo claro, 6,2 x 2,8 —7,5 x 3,3 mm; 65 mor- 
fotipo con bandas, 5,9 x 2,9 mm - 7,3 x 3,1 mm); Ceuta, puerto, 4 ej. (1 morfotipo claro, resto con 
bandas; 6,9 x 2,4 - 7,5 x 2,8 mm); Ceuta, Punta del Saudiño, 10 ej. (1 morfotipo con bandas, resto 
morfotipo claro; 6,1 x 2,6 — 7,1 x 2,9 mm, ejemplar dibujado); Ceuta, Playa Benítez, 15-25 m, 2 c. 
(7,0 x 3,0 -7,4 x 3,3 mm); Benzú, infralitoral rocoso, 87 c. (35 morfotipo claro, 5,3 x 2,4-7,0 x 3,0 
mm, 52 morfotipo con bandas oscuras, 5,5 x 2,5 mm —7,3 x 3,1 mm) + 4 ej. (morfotipo oscuro, 5,6 
x 2,5 — 6,4 x 2,8 mm) + 4j.; Punta Cires, 1 ej., col. SG, MNHN (morfotipo claro, 6,8 x 2,7 mm) + 1 
ej.j.; Tánger, 4 c., col. SG, MNHN (morfotipo claro, 5,5 x 2,4- 6,8 x 2,9 mm). Portugal: Tavira, Pedra 
do Barril, 12 c. (5,8 x 2,6 — 6,8 x 2,8 mm); Tavira, Cabanas, 2 ej. + 2 c. (7,0 x 2,7 - 7,3 x 2,9 mm); 
Sagres, Baía da Baleeira, bajamar, 72 ej. (5,5 x 2,4 — 6,4 x 2,6 mm) + 20 j.; Sagres, Baía da Baleeira 
3-15 m, Sagres, Baía da Baleeira 3-15 m, 60 ej. (5,2 x 2,2 — 6,6 x 2,7 mm) + 10j.; Sagres, Ponta da 
Baleeira, 17-23 m, 451 ej. (315 morfotipo con bandas oscuras 6,0 x 2,3 - 7,2 x 2,8 mm, 136 morfo- 
tipo affinis 5,6 x 2,3 — 6,8 x 2,7 mm) + 316 j.; Sagres, Pontal dos Corvos, 5 ej. (6,3 x 2,6 —- 7,0 x 2,7 
mm); Sagres, Ponta dos Caminhos, 14 c. (6,4 x 2,7 - 6,8 x 2,8 mm) + 4 j. Banco Gorringe: Seamount 
1, DWO04, 100 c.; DWO5, 15 c.; DWO0€, 7 c.; DWO8, 2j.; DWO9, 6 c.; DW32, 1 ej. + 13.; DW 'S33, 5 ej. (5,7 
x 2,6 mm) + 2j. Banco Ampere: “Seamount 1” DW92, 10 c.; DW98, 4 c. 


Descripción: Concha de hasta 7,7 vuelta se añade una escultura de 4-5 


mm (generalmente, menos de 7,5 mm), 
con 5!/2-6 vueltas de espira convexas, 
con la sutura bastante marcada. 
Protoconcha de 600-650 um de diá- 
metro máximo (núcleo: 300-350 um, 
primera media vuelta: 500-550 um). 
Desde el mismo núcleo aparece una 
escultura de cordoncillos espirales apla- 
nados, de grosor similar a los interespa- 
cios, en el fondo de los cuales se apre- 
cian diminutas laminillas transversales 
paralelas a las líneas de crecimiento; hay 
20-22 cordoncillos al final de la primera 
media vuelta. En la segunda media 


30 


costillas axiales algo sigmoideas y opis- 
toclinas, de anchura similar a los espa- 
cios que las separan, sobre las cuales los 
cordoncillos discurren sin interrupción. 
Teleoconcha con ornamentación 
formada por cordones espirales, el doble 
de anchos que sus interespacios, y por 
costillas axiales amplias sobre las cuales 
los cordones discurren sin discontinui- 
dad e incluso se ensanchan un poco en 
la parte más saliente de la costilla. En la 
primera vuelta aparecen normalmente 
cuatro cordones espirales (a veces 3, en 
poblaciones del Estrecho y de Alborán); 


GOFAS Y OLIVER: Las especies de Chauvetía en el área ibero-marroquí 


el cordón subsutural se desdobla en las 
vueltas siguientes. En la última vuelta, 
hay un total de 15-17 cordones, de los 
cuales 4 ó 5 continúan los de la penúl- 
tima vuelta, los 5 medios son de un 
grosor equivalente, y el grosor de los 
siguientes cordones decrece marcada- 
mente hacia el canal sifonal. Las costillas 
axiales son suaves y de igual anchura 
que los interespacios y se reparten de 
forma aleatoria, sin estar sistemática- 
mente alineadas entre una vuelta y otra. 
En la penúltima vuelta se aprecian 11-14 
costillas axiales (hasta 15 en algunos 
ejemplares del Algarve), algo flexuosas 
y prosoclinas. La última vuelta alcanza 
en torno al 60% de la altura total, y la 
abertura en torno al 40%. 

Exterior del labio engrosado en los 
ejemplares adultos, con los cordones 
que persisten, aunque atenuados, sobre 
la variz externa. En el interior del labio 
se observan, en los ejemplares muy 
adultos, hasta 6-7 dentículos poco pro- 
nunciados, alargados hacia el interior de 
la abertura. 

El color de la concha es variable. En 
el lectotipo de Nesaea mamillata y en las 
demás poblaciones mediterráneas, hasta 
un límite occidental que se sitúa en la 
provincia de Almería, aparece un patrón 
de color amarillento con las interseccio- 
nes de cordones y costillas de un color 
castaño oscuro (Figs. 3 y 5, “morfotipo 
mamilar” en material examinado). En 
toda la zona de estudio, el patrón más 
frecuente es un color castaño claro con 
los cordones resaltados por bandas cas- 
tañas más oscuras, incluso sobre el exte- 
rior del labio (Figs. 4 y 6). En la isla de 
Alborán, en algunas localidades del 
Estrecho (Figs. 7, 8) y en los bancos lusi- 
tanos (Fig. 14), este patrón puede estar 
muy contrastado, con los intervalos 
entre cordones hasta casi blancos. En la 
zona del Estrecho, aparecen con fre- 
cuencia ejemplares de un color amari- 
llento claro uniforme (Figs. 9, 10). Final- 
mente, en algunos ejemplares de Sicilia, 
incluido el posible sintipo de Chauvetia 
affinis, se observa un patrón de color con 
las intersecciones de los cordones y de 
las costillas más claras sobre un fondo 
castaño. 


Animal con una pigmentación negra 
superficial, uniforme o formando 
densas manchas negras sobre fondo 
blanco; el extremo anterior y la suela del 
pie, el espacio que rodea los ojos y la 
base del sifón son más claros. 

Distribución: Mediterráneo; Atlán- 
tico, común en el banco Gorringe y el 
litoral del suroeste ibérico desde el cabo 
de San Vicente hasta el estrecho de 
Gibraltar; esporádico en la costa de 
Marruecos (MICALI, 1999), el banco 
Ampere y Canarias (PEÑAS Y ROLÁN, 
2009) 

Notas: Desde el punto de vista 
nomenclatural, si se consideran coespe- 
cíficas Lachesis mamillata, Nesaea granu- 
lata y N. mamillata, fue SCACCHI (1836: 
12) quien actuó como primer revisor, 
utilizando el nombre Murex granulatus y 
citando los otros dos nombres como 
sinónimos. Sin embargo, el uso del 
nombre C. mamillata está tan generali- 
zado que es indeseable un cambio, sobre 
todo si tenemos en cuenta que los usos 
posteriores del nombre C. granulata se 
entienden generalmente en el sentido de 
TIBERI (1868). 

La forma mediterránea habitual- 
mente identificada como Chauvetia 
mamillata se caracteriza por un patrón 
de color en el que las intersecciones de 
los cordones con las costillas están mar- 
cadas por manchas subcuadradas de 
color castaño oscuro, sobre un fondo 
amarillento claro. Este patrón no se ha 
encontrado jamás al oeste de la provin- 
cia de Almería. Existen patrones de tran- 
sición que BUCQUOY ET AL. (1883) distin- 
guieron como variedad submamillata. En 
esta revisión nos atenemos a la conclu- 
sión de HERGUETA ET AL. (2002), de que 
hay formas con bandas castañas conti- 
nuas o de color castaño uniforme, sin 
que por ello se puedan reconocer espe- 
cies distintas. En todo el estrecho de 
Gibraltar, las formas de color amari- 
llento pálido uniforme son frecuentes, 
aunque coexisten con las formas con 
bandas castañas. En estos ejemplares 
amarillentos, el albinismo de la concha 
no se extiende al animal, que sigue mos- 
trando el patrón con pigmentos negros 
superficiales característico de la especie. 


31 


Iberus, 28 (1), 2010 


En los bancos submarinos de 
Gorringe y Ampere, al oeste de Portugal 
y de Marruecos respectivamente, existen 
poblaciones que asignamos a Chauvetia 
mamillata y que se parecen mucho a las 
formas con bandas del estrecho de 
Gibraltar. Son algo más pequeñas, con 
un tamaño muy uniforme de unos 6 mm 
y el patrón de coloración del animal es 
negro, como en los ejemplares del Estre- 
cho. Los ejemplares vivos se han reco- 
gido en la zona fótica, en la parte más 
somera del Banco Gorringe y las nume- 
rosas conchas recogidas hasta unos 500 
metros de profundidad fueron probable- 
mente transportadas desde zonas más 
someras. El aislamiento de los bancos, 
junto al tamaño necesariamente 
pequeño de estas poblaciones, son cir- 
cunstancias que propician su divergen- 
cia evolutiva, pero, de momento, las 
diferencias observadas no pasan de lo 
que se puede atribuir a la variación 
intraespecífica. 

En las muestras de los alrededores 
de Sagres, en el extremo sudoccidental 
de la Península Ibérica y muy cerca del 
límite norte de distribución, la especie 
es extremadamente abundante y varia- 
ble y se encuentra en las mismas mues- 
tras que C. brunnea, C. retifera, C. denti- 
fera spec. nov. y C. taeniata spec. nov. En 


muchos ejemplares el número de costi- 
llas es más elevado (hasta 15 en la 
última vuelta, frente a 11-13 habitual- 
mente, Figs. 11, 12), y éstas son menos 
flexuosas. En este aspecto se aproximan 
a lo descrito por MONTEROSATO (1889) 
para su Chauvetia affinis “con reticulado 
áspero como una lima”; sin embargo, 
existen transiciones con el tipo morfoló- 
gico más habitual (Fig. 13) y entra en 
nuestro concepto de C. mamillata del sur 
ibérico. OLIVER Y ROLÁN (2009, figs. 6-9) 
ilustran ejemplares de Sicilia identifica- 
dos por Monterosato como C-. affinis y 
enviados a Locard, actualmente conser- 
vados en el MNHN. Al contrario de la 
tendencia habitual de C. mamillata en el 
Mediterráneo, estos ejemplares sicilia- 
nos tienen las intersecciones de los cor- 
dones y las costillas más claras que el 
color de fondo, con lo que no se puede 
descartar que se trate de una especie 
distinta. En los ejemplares portugueses, 
el color más oscuro se sitúa, como es 
habitual en C. mamillata, sobre los cordo- 
nes, aunque poco contrastado. 

MICALI (1999) distingue a nivel espe- 
cifico entre Chauvetia mamillata y C. 
turritellata e ilustra bajo este último 
nombre tres animales vivos de Aci 
Trezza (Sicilia) con un patrón de 
manchas negras mucho más disperso 


(Página derecha) Figuras 3-14. Chauvetia mamillata (Risso, 1826), principales variaciones de la 
concha en la zona de estudio. 3, 4: patrón con las intersecciones de cordones y costillas de un color 
castaño oscuro, exclusivo del Mediterráneo, y patrón con bandas castañas, Denia (6,4 y 6,2 mm); 
5, 6: los mismos patrones de color, Almería (6,0 y 5,9 mm); 7, 8: patrón con bandas castañas sobre 
los cordones, habitual en el mar de Alborán, Estrecho y Atlántico, Punta Almina (Ceuta), 32-40 m 
(6,4 mm); 9, 10: patrón de color muy pálido, frecuente en el Estrecho, Punta del Saudiño (6,7 
mm); 11, 12: ejemplar de Sagres, con escultura más fina al modo de Chauvetia affinis (5,8 mm); 
13: ejemplar de la misma localidad, con la escultura más habitual en C. mamillata del mar de 
Alborán y del Estrecho; 14: ejemplar del Banco Gorringe, Seamount 1, DW33, 55-70 m (5,7 
mm). 

(Right page) Figures 3-14. Chauvetia mamillata (Risso, 1826), most common variants of the shells in 
the study area. 3, 4: pattern with dark brown intersections of cords and ribs, found only in the Medite- 
rranean, and pattern with brown bands, Denia (6.4 and 6.2 mm); 5, 6: the same colour patterns, 
Almería (6.0 and 5.9 mm); 7, 8: colour pattern with brown bands over the cords, usual in the Alborán 
Sea, the Straits and the Atlantic, Punta Almina 32-40 m (6.4 mm); 9, 10: very pale colour pattern, 
frequent in the Straits, Punta del Saudiño (6.7 mm); 11, 12: specimen from Sagres, with a sculpture in 
the mode of Chauvetia affinis (5.8 mm); 13: specimen from the same locality, with sculpture more as 
usual in C. mamillata from Alborán Sea and the Straits; 14: specimen from Gorringe Bank, “Seamount 


1” DW33, 55-70 m (5.7 mm). 


32 


GOFAS Y OLIVER: Las especies de Chauvetia en el área ibero-marroquí 


Iberus, 28 (1), 2010 


Figuras 15-18. Chauvetia brunnea (Donovan, 1804). 15: ejemplar de lle de Sein, Bretaña (4,7 
mm); 16: ejemplar de La Toja, Galicia (4,0 mm); 17, 18: ejemplar de Baia da Baleeira Sagres, Por- 
tugal (5,5 mm); nótese el aspecto liso del exterior de la variz. Figuras 19-22. Chauvetia maroccana 
spec. nov. 19-21: holotipo de El Jadida (4,2 mm); 22: ejemplar de Essaouira (4,8 mm). 

Figures 15-18. Chauvetia brunnea (Donovan, 1804). 15: specimen from lle de Sein, Brittany (4.7 
mm); 16: specimen from La Toja, Galicia (4.0 mm); 17, 18: specimen from Baia do Martinhal, Sagres, 
Portugal (5.5 mm); note the smooth appearance of the exterior of the varix. Figures 19-22. Chauvetia 
maroccana spec. nov. 19-21: holotype from El Jadida (4.2 mm); 22: specimen from Essaouira (4.8 


mm). 


que en los ejemplares del sur de la 
Península Ibérica. Dicho autor no cita 
esta última especie en España, aunque sí 
en el sur de Marruecos, Canarias (se 
trata de la forma referida por OLIVER Y 
ROLÁN, 2009 a Chauvetia affinis) y 
Senegal. 

En el material estudiado del área ibero- 
marroquí, a pesar de una indiscutible 


34 


variabilidad, no hemos encontrado crite- 
rios que permitan distinguir más de una 
especie en lo que tratamos aquí como C. 
mamillata. No obstante, se debe de tener 
en cuenta que la localidad tipo de Fusus 
turritellatus se sitúa en el Peloponeso, en 
la cuenca oriental del Mediterráneo y que 
la revisión de dicho taxón debe contar con 
material procedente de esta zona. 


GOFAS Y OLIVER: Las especies de Chauvetía en el área ibero-marroquí 


Chauvetia brunnea (Donovan, 1804) (Figs. 15-18, 27-30, 99-100) 


Buccinum minimum Montagu, 1803. Test. Brit.: 247, lám. 8, fig. 2. [Localidad tipo: Devon, Gran 
Bretaña; preocupado por Buccinum minimum Berckenhout, 1795; Buccinum minimum Turton, 
1802]. 

Buccinum brunneum Donovan, 1804. Brit. Sh. vol. 5: lám. 179, fig. 2. y explicación de la lámina (no 
paginada) [Localidad tipo: Cornwall, Gran Bretaña]. 

Fusus subnigris Brown, 1827. III. Conch.: 5, lám. 48, figs. 58-59. 

Buccinum rubrum Potiez y Michaud, 1838. Catal. Mus. Douai, : 381, lám. 32, fig. 17-18. 

Lachesis minima var. pallescens Jeffreys, 1867. Brit. Conch. vol. iv: 313. 


Material estudiado: Canal de la Mancha y Bretaña: Guernesey, Rocquaine Bay (49* 27” N, 02* 39 
W), bajamar, 2 ej. (4,4 x 1,9 mm, ejemplar dibujado), col. SG, MNHN,; Roscoff, “les Cochons Noirs”, 
20 m, 1 ej. (4,6 x 2,1 mm), col. SG, MNHN; Ploumanac'h, en Corallina en bajamar, 5 j., col. SG, 
MNHAN; Ile de Sein, bajamar, 2 ej. (4,3 x 2,1 -4,7 x 2,1 mm) + 5j.; Lampaul-Ploudalmézeau, bajamar, 
23., col. SG, MNAHN; Baie de Bertheaume 20-30 m, 17 c. (3,6 x 1,8 - 4,7 x 2,1 mm), col. SG, MNHN. 
Asturias: Muros, 1 ej. (4,7 x 2,1 mm, ejemplar dibujado). Galicia: La Toja, 1c. (4 x 1,8 mm), col. JDO. 
Portugal: Sagres, Ponta da Baleeira 17-23 m, 52 ej. (4,3 x 2,3 -5,6 x 2,1 mm); Sagres, Baia da Baleeira, 
bajamar, 10 ej. (4,5 x 2,0— 5,3 x 2,3 mm); Sagres, Baia da Baleeira, 3-15 m, 1 ej. (5,0 x 2,0 mm); Sagres, 
Ponta dos Caminhos, 4 c. (4,5 x 2 - 4,8 x 2,1 mm); Sagres, Pontal dos Corvos, 1 ej. (5,0 x 2,1 mm); 
Tavira, Pedra do Barril, 3 ej. (5,4 x 2,4 — 5,7 x 2,4 mm); Tavira, Cabanas, 1 ej. (5,7 x 2,4 mm) + 1j. 
Estrecho de Gibraltar y mar de Alborán: Calahonda (Málaga), 1 ej. (5,2 x 2,3 mm), col. SG 1978, 
MNHN, Sotogrande (Cádiz), bajamar, 2 ej. (morfotipo claro, 4,9 x 2,0 - 5,6 x 2,5mm) +2 c. (4,2 x 
1,9 - 4,9 x 2,2 mm), col. SG, MNHN,; Tarifa, Torre de la Peña, 1 c. (5,2 x 2,2 mm), col. SG, MNHN; 
Ceuta, Sarchal, 0-3 m, 4 ej. (4,2 x 1,8 — 5,0 x 2,1 mm); Ceuta, Punta Almina, 25-40 m, 4 ej. (conchas 
muy claras, 5,0 x 2,2 — 5,2 x 2,2 mm, ejemplar dibujado); Ceuta, Benzú, 04 m, 1 ej. (5,0 x 3,2 mm, 


ejemplar dibujado). 


Descripción: Concha de hasta 5,7 mm 
(generalmente, menos de 5 mm), con 
cuatro a cinco vueltas de espira conve- 
xas, con la sutura bastante marcada. 

Protoconcha de 500-550 um de diá- 
metro máximo (núcleo: 260-300 um, 
primera media vuelta: 400-450 um). El 
núcleo es inicialmente casi liso, luego 
aparece una escultura de cordoncillos 
aplanados y desiguales, más anchos que 
los interespacios, en el fondo de los 
cuales se aprecian diminutas laminillas 
transversales paralelas a las líneas de 
crecimiento. En algunos ejemplares bien 
conservados, los cordoncillos se mani- 
fiestan desde el núcleo; hay 20-25 cor- 
doncillos al final de la primera media 
vuelta. En la segunda media vuelta 
aparece una escultura de costillas 
axiales algo sigmoideas y opistoclinas, 
de anchura similar a los espacios que las 
separan, y sobre las cuales los cordonci- 
llos tienden a atenuarse. 

Teleoconcha con ornamentación 
formada por cordones espirales, el doble 
de anchos que sus interespacios, y por 
costillas axiales amplias, sobre las cuales 


los cordones discurren sin discontinui- 
dad o incluso se ensanchan un poco en 
la parte más saliente de la costilla. En la 
primera vuelta aparecen normalmente 
cuatro cordones, que persisten en las 
vueltas sucesivas de la espira; el cordón 
subsutural está a veces algo desdoblado 
al final. En la última vuelta hay un total 
de 17-18 cordones, de los cuales cuatro 
continúan los de la penúltima vuelta, los 
5 medios son de un grosor equivalente y 
los que cubren el canal sifonal son mar- 
cadamente más finos. Las costillas 
axiales son 9-10 en la penúltima vuelta, 
casi ortoclinas, algo más estrechas que 
los interespacios y con tendencia a aline- 
arse aproximadamente entre una vuelta 
y otra. La última vuelta alcanza el 61% 
de la altura total, mientras que la aber- 
tura ocupa en torno al 42%. 

Exterior del labio muy engrosado en 
los ejemplares adultos, sobre todo en su 
porción adapical, que suele formar una 
variz en la cual los cordones están extre- 
madamente atenuados o incluso llegan 
a desaparecer, siendo la parte adapical 
completamente lisa. En el interior del 


0) 


Iberus, 28 (1), 2010 


labio se observan, en los ejemplares 
muy adultos, hasta 6-7 dentículos poco 
pronunciados, alargados hacia el inte- 
rior de la abertura. 

El color de la concha es castaño uni- 
forme. Animal de color amarillento uni- 
forme, con puntitos blancos brillantes. 

Distribución: Desde el Canal de la 
Mancha hasta el estrecho de Gibraltar, 
sin citas comprobadas para el Medite- 
rráneo más al este del extremo occiden- 
tal del mar de Alborán. Esta distribución 
un tanto extraña la tiene también el ris- 
sóido Setia pulcherrima (Jeffreys, 1848) y 
el pulmonado Otina ovata (Brown, 1827). 

Notas: Coincidimos con MONTERO- 
SATO (1884: 136) y con HERGUETA ET AL. 
(2002) en considerar que esta especie 
atlántica con localidad tipo en Gran 
Bretaña es distinta a la que predomina 
en el Mediterráneo occidental (C. mami- 
llata) y que se ha confundido habitual- 
mente con ella (BuUCQOUOY ET AL. 1883, 
entre otros). 

Chauvetia brunnea es más pequeña 
que la especie mediterránea, superando 


poco los 5 mm, tiene menos costillas 
axiales en la última vuelta (9-10), el lado 
adapical externo de la variz del labio es 
liso y brilloso, sin que se marque la con- 
tinuación de los cordones de la última 
vuelta y la concha suele ser monocroma; 
en C. mamillata, los cordones, y even- 
tualmente sus bandas oscuras, se conti- 
núan, aunque atenuados, sobre la parte 
externa del labio. El animal de C. 
brunnea es siempre amarillento con pun- 
titos blancos, sin ninguna marca negra 
por muy oscura que sea la concha, 
mientras que es negro liso o densamente 
moteado de negro en C. mamillata, 
también en los individuos con concha 
clara. 

Las muestras de los alrededores de 
Ceuta contienen unos pocos ejemplares 
que se caracterizan por un color muy 
claro, amarillento, de la concha. El 
animal de estos ejemplares carece com- 
pletamente de pigmentos negros y se 
ajusta al patrón observado en el Atlán- 
tico europeo; por esta razón, se conside- 
ran como pertenecientes a C. brunnea. 


Chauvetia maroccana spec. nov. (Figs. 19-22, 31-34) 


Material tipo: Holotipo [ej., 4,2 x 1,9 mm], col. SG 1991 (MNHN 22868) y 10 paratipos (5 ej., MNHN 
22869, 5 ej. MNCN 15.05/53585), todos de la localidad tipo. 

Material estudiado: Marruecos: Temara, bajamar, 3 ej. (4,5 x 1,9 - 4,7 x 2,1 mm) + 7 j.; El Jadida, 
bajamar, 43 ej. (3,9 x 1,8 - 4,9 x 2,1 mm; incluye material tipo) + 28j. Essaouira, bajamar, 41 ej. + 40 


j. (4,2 x 1,9 - 5,1 x 2,3 mm). 


Localidad tipo: El Jadida, Marruecos (33* 15,1” N — 08” 29,7” W, intermareal). 
Etimología: El nombre alude a la distribución geográfica de la especie en la costa marroquí. 


Descripción: Concha de hasta 5,1 mm 
(generalmente menos de 5 mm), con 
unas cuatro vueltas de espira convexas, 
con la sutura bastante marcada. 

Protoconcha de 500-550 um de diá- 
metro máximo (núcleo: 260-300 um, 
primera media vuelta: 400-450 um). 
Desde el núcleo aparece una escultura 
de cordoncillos aplanados, desiguales, 
más estrechos que los interespacios o 
iguales a ellos, en el fondo de los cuales 
se aprecian diminutas laminillas trans- 
versales paralelas a las líneas de creci- 
miento; hay 20-25 cordoncillos al final 
de la primera media vuelta. En la 
segunda media vuelta aparece una 


36 


escultura de costillas axiales algo sig- 
moideas y opistoclinas, de anchura 
similar a los espacios que los separan, 
sobre las cuales los cordoncillos tienden 
a atenuarse. 

Teleoconcha con ornamentación 
formada por cordones espirales altos, 
más estrechos o igual de anchos que sus 
interespacios, y por costillas axiales 
amplias, sobre las cuales los cordones 
discurren sin discontinuidad. En la 
primera vuelta aparecen cuatro cordo- 
nes, que persisten en las vueltas sucesi- 
vas de la espira y están nítidamente 
delimitados de los interespacios. En la 
última vuelta hay un total de 18-20 cor- 


GOFAS Y OLIVER: Las especies de Chauvetía en el área ibero-marroquí 


Figuras 23, 34. Protoconchas en vista lateral y apical, todas a la misma escala. 23-24: Chauvetia 
mamillata, Punta de Loma Pelada, Almería; 25, 26: Chauvetia mamillata, Punta da Baleeira, 
Sagres; 27, 28: Chauvetia brunnea, Tarifa; 29, 30: Chauvetia brunnea, Baia da Baleeira, Sagres; 31- 
34: Chauvetia maroccana spec. nov., El Jadida. 

Figures 23, 34. Protoconchs in lateral and apical views, all to scale. 23-24: Chauvetia mamillata, 
Punta de Loma Pelada, Almería; 25, 26: Chauvetia mamillata, Punta da Baleeira, Sagres; 27, 28: 
Chauvetia brunnea, Tarifa; 29, 30: Chauvetia brunnea, Bay of Baleeira, Sagres; 31-34: Chauvetia 
maroccana spec. n02., El Jadida. 


3/7 


Iberus, 28 (1), 2010 


dones, de los cuales cuatro continúan 
los de la penúltima vuelta, los 5 medios 
son de un tamaño equivalente y los que 
cubren el canal sifonal son marcada- 
mente más finos. Las costillas axiales 
son 9-10 en la penúltima vuelta, relati- 
vamente rectas, equivalentes a los inte- 
respacios y con tendencia a alinearse 
aproximadamente entre una vuelta y 
otra. La última vuelta alcanza menos del 
60% de la altura total, mientras que la 
abertura ocupa en torno al 40%. 

Exterior del labio engrosado en los 
ejemplares adultos, formando una variz 
en la cual los cordones se continúan, 
aunque algo atenuados. En el interior 
del labio se observan, en los ejemplares 
adultos, hasta 5-6 dentículos poco pro- 
nunciados, alargados hacia el interior de 
la abertura. 

El color de la concha es castaño 
oscuro uniforme. Animal de un color 
amarillento uniforme, con puntos de 
color blanco brillante. 

Distribución: Esta especie se ha 
encontrado hasta ahora sólo en las 
muestras procedentes de tres localida- 
des de la costa atlántica marroquí, en 
lavados de algas del piso intermareal. 

Notas: Se parece mucho a Chauvetia 
brunnea por el tamaño y el color del 


animal, pero se diferencia claramente 
por tener hasta una vuelta menos en la 
teleoconcha, un perfil de espira menos 
puntiagudo en un tamaño comparable y, 
sobre todo, en la forma de los cordones 
espirales, que son de igual anchura o 
más estrechos que los interespacios 
(compárense las figuras 27 y 29 con las 
figuras 31 y 33) y ello, desde las prime- 
ras vueltas. Los cordones espirales están 
nítidamente delimitados y, a diferencia 
de C. brunnea, se continúan sobre la 
parte externa del labio. Chauvetia maroc- 
cana forma abundantes poblaciones en 
el piso intermareal de dos de los tres 
lugares donde fue recolectada, siendo 
allí uno de los moluscos más abundan- 
tes, mientras C. brunnea suele ser poco 
abundante, incluso en localidades como 
Sagres, donde son abundantes otras 
especies del género. 

La especie marroquí tiene también 
un cierto parecido con Chauvetia borgesi 
Oliver y Rolán 2009, de Canarias, pero, 
con un tamaño comparable, esta última 
se diferencia por tener cinco cordones 
espirales en las primeras vueltas de la 
teleoconcha, que son más anchos que los 
interespacios; el animal de C. borgesi se 
diferencia por tener el sifón pigmentado 
de negro. 


Chauvetia procerula (Monterosato, 1889) (Figs. 35-41, 42-45) 


Donovania procerula Monterosato, 1889. J. Conchyl. 37: 116. [Localidad tipo: Casablanca, Marrue- 
cos y Argel, Argelia]. 


Material tipo: No mencionado por Micali (1999) entre las Chauvetia de la colección Monterosato en 
Roma. 

Material examinado: Levante ibérico: Islas Columbretes (Placer de la Barra Alta), 2 c. (7,9 x 2,9 
mm), MNCN,; Denia (arrojadas a la playa), 2 c. (6,5 x 2,7 — 8,3 x 3,5 mm) + 6j., col. JDO; Cullera 
(15 m) 3 c. (6,8 x 2,7 — 7,3 x 3,0 mm), col. JDO; Jávea (arrojada a la playa), 1j., col. JDO. Mar de 
Alborán y estrecho de Gibraltar: isla de Alborán, 5 c. (7,1 x 3,0 — 7,9 x 3,1 mm) + 35 j., col. JDO; 
Rincón de la Victoria (Málaga), 1 c. (8,3 x 3,3 mm), col. SG, MNHN; Benalmádena costa (Málaga), 
2 ej. (6,8 x 2,7 —7,0 x 2,8 mm), col. SG; Sotogrande (Cádiz), 1 j.; Tarifa, Torre de la Peña, 1 c. (7,3 x 
3,0 mm); Barbate, 29 m, 1j., col. SG; Cabo de Trafalgar (36” 08” N, 06” 01” W), 4j., MNCN; Cádiz, 22 
c. (7,2 x 3,1 - 8,5 x 3,7 mm) + 4j., col. Azpeitia MNCN; M'dig (200 m), 1 c. (7,3 x 3,1 mm), col. “Al 
Mounir”, MNHN,; Ceuta, Punta Almina, 25-40 m, 3 ej. (7,2 x 3,0 - 8,2 x 3,4 mm) + 2j.; Ceuta, playa 
en el puerto, 2 c. (8,1 x 3,1 y 8,4 x 3,2 mm); Ceuta, Punta del Saudiño, 1 ej. (7,2 x 3,0 mm, ejemplar 
dibujado); Tánger, 12 c. (6,8 x 2,9 - 8,1 x 3,3 mm), coll. SG, MNHN. Portugal: Tavira, Pedra do 
Barril, 33 c. (6,7 x 2,8 — 8,7 x 3,6 mm) + 34 c.j.; Algarve (36" 58,0” N, 08% 55,6" W, 65 nl (AE 
3,2 mm) + 1j., col. Péres, Gautier, Vacelet 7,8,1957, MNHN,; Sagres, Baía da Baleeira, 3-15 m, 36 ej. 
(6,6 x 2,7 - 9,4 x 3,6 mm, ejemplar dibujado); Sagres, Ponta da Baleeira, 17-23 m, 31 ej. (7,7 x 3,2 — 
9,7 x 3,6 mm, ejemplar dibujado) + 30j.; Sagres, Pontal dos Corvos, 5 ej. (8,0 x 3,3 - 8,9 x 3,5 mm); 


38 


GOFAS Y OLIVER: Las especies de Chauvetía en el área ¡ibero-marroquí 


Figuras 35-41. Chauvetia procerula (Monterosato, 1889). 35, 36: ejemplar de Benalmádena, 
Málaga (6,8 mm); 37: ejemplar de Denia (7,2 mm); 38, 39: ejemplar con el patrón de color con 
banda blanca, Benzú, Ceuta (7,2 mm); 40: ilustración de PALLARY (1902, fig. 14), correspondiente 
al concepto que dicho autor tenía de Chauvetia decorata Monterosato, 1889, Tánger; 41: otro 
ejemplar con banda blanca, Cádiz (7,7 mm). 

Figures 35-41. Chauvetia procerula (Monterosato, 1889). 35, 36: specimen from Benalmádena, 
Málaga (6.8 mm); 37: specimen from Denia (7.2 mm); 38, 39: specimen with a colour pattern with a 
white band, Benzú, Ceuta (7.2 mm); 40: illustration in PALLARY (1902, fig. 14) matching this authors 
concept of Chauvetia decorata Monterosato, 1889, Tangiers; 41: another specimen with a white band, 
Cádiz (7.7 mm) 


39 


Iberus, 28 (1), 2010 


Sagres, Ponta dos Caminhos, 9 c. (7,8 x 3,2 — 8,8 x 3,6 mm); entre Faro y Sagres, redes de pesca, 2 
ej. (subadultos, 6,5 x 2,5—7,3 x 3,0 mm, ejemplares dibujados); entre Salema y Praia da Luz, redes 
de pesca —70 m, 10 ej. (morfotipo claro, 7,3 x 3,0 - 8,6 x 3,4 mm). Patrón de color con banda blanca: 
Punta Almina (Ceuta), 25-40 m, 1 c. (7,2 x 2,6 mm); Ceuta, Punta del Saudiño, 2 ej. (7,0 x 2,9 - 7,1 
x 3,1 mm, ejemplar dibujado); Ceuta, Benzú, 12 c. 6,4 x 2,7 -7,7 x 3,1 mm), col. SG, MNHN; Tánger 
(arrojadas a la playa), 7 c. (6,0 x 2,7 — 6,8 x 2,8 mm), col. SG, MNHN,; Asilah, 1 j., col. SG, MNHN; 


Mohammedia, 2 c. (6,7 x 2,6 mm), col. SG, MNHN. 


Descripción: Concha de hasta 9,7 mm 
(generalmente menos de 8 mm), con 
6-6! / 2 vueltas de espira convexas, con la 
sutura bastante marcada. 

Protoconcha de 600-650 um de diá- 
metro máximo (núcleo: 350-400 um, 
primera media vuelta: 500-550 um). 
Desde el mismo núcleo aparece una 
escultura de cordoncillos aplanados y 
anchos, bien visibles en microscopía 
óptica, desiguales entre sí, mucho más 
anchos que los interespacios, en el fondo 
de los cuales se aprecian diminutas 
laminillas transversales paralelas a las 
líneas de crecimiento; hay 15-16 cordon- 
cillos al final de la primera media 
vuelta. En la segunda media vuelta se 
añade una escultura de costillas axiales 
algo sigmoideas y opistoclinas, de 
anchura progresivamente mayor hasta 
el principio de la teleoconcha; en esta 
parte, los cordoncillos espirales se 
vuelven más altos o pasan a ser más 
estrechos que los interespacios. 

Teleoconcha con ornamentación 
formada por cordones espirales, de 
anchura equivalente a sus interespacios, 
y por costillas axiales amplias sobre las 
cuales los cordones discurren sin discon- 
tinuidad e incluso se vuelven más gruesos, 
formando tubérculos a modo de perlas. 
En la primera vuelta aparecen cuatro cor- 
dones espirales; el cordón subsutural se 
desdobla en la tercera vuelta, de modo que 
hay 5 cordones en la penúltima vuelta. En 
la última vuelta, hay un total de 19-20 cor- 
dones, de los cuales 5 continúan los de la 
penúltima vuelta, los 5 medios son de un 
tamaño equivalente y, en la parte abapi- 
cal, los cordones decrecen progresiva- 
mente en tamaño. Las costillas axiales son 
altas y más estrechas que los interespacios 
y tienden a alinearse entre vueltas sucesi- 
vas. En la penúltima vuelta se aprecian 9- 
10 costillas axiales algo prosoclinas. La 
última vuelta alcanza el 57% de la altura 


40 


total, mientras que la abertura ocupa en 
torno al 39%. 

Exterior del labio muy engrosado en 
los ejemplares adultos con los cordones 
continuados, aunque algo atenuados, 
sobre la variz externa. El perfil de la 
última vuelta está marcadamente estre- 
chado en su tercio abapical alrededor 
del canal, que así aparece más promi- 
nente. En el interior del labio se obser- 
van, en los ejemplares adultos, hasta 7-8 
dentículos poco pronunciados, alarga- 
dos hacia el interior de la abertura. 

El color de la concha es habitual- 
mente castaño uniforme, en algunos 
casos amarillento claro; en ningún caso 
los cordones se destacan con un color 
distinto a los interespacios. En el estre- 
cho de Gibraltar y en la costa atlántica 
marroquí aparecen ejemplares con la 
protoconcha casi blanca, la teleoconcha 
castaña en la mitad adapical de las 
vueltas y en la parte abapical de la 
última vuelta, con una zona blanca en la 
mitad abapical de las vueltas de espira, 
continuada como una banda en la mitad 
de la última vuelta. 

El animal es de color negro uni- 
forme, tendiendo a veces a más pálido 
en la parte anterior del pie y en los ten- 
táculos cefálicos; una zona descolorida 
bajo el opérculo, donde éste hace con- 
tacto con el pie. 

Distribución: Mediterráneo; Atlán- 
tico, común en el litoral del suroeste 
ibérico desde el cabo de San Vicente 
hasta el estrecho de Gibraltar; esporá- 
dico en la costa de Marruecos (MONTE- 
ROSATO, 1889; MICALI, 1999) y Canarias 
(OLIVER Y ROLÁN, 2009). La cita en 
Senegal de MiIcaLI (1999) se da por 
dudosa, puesto que la especie no fue 
confirmada en el extenso material estu- 
diado por OLIVER Y ROLÁN (2008). 

Notas: Los cordoncillos de la proto- 
concha en esta especie son marcada- 


GOFAS Y OLIVER: Las especies de Chauvetia en el área ibero-marroquí 


Figuras 42-47. Protoconchas en vista lateral y apical. 42, 43: Chauvetia procerula, con patrón de 
color uniforme, isla de Alborán y Denia. 44, 45: Chauvetia procerula, con banda blanca, Benzú. 
46, 47: Chauvetia recondita, Denia e isla de Alborán. 

Figures 42-47. Protoconchs in lateral and apical views, all to scale. 42, 43: Chauvetia procerula, with 
a uniform colour pattern, Alborán Island and Denia. 44, 45: Chauvetia procerula, with a white 
band, Benzú. 46, 47: Chauvetia recondita, Denia and Alborán Island. 


mente más anchos que en C. mamillata; 
la teleoconcha difiere por tener las costi- 
llas axiales más altas y los cordones no 
más anchos que los interespacios. El 
perfil de la última vuelta está más mar- 
cadamente estrechado alrededor del 
sifón que en las demás especies. 

El color de la concha suele ser 
castaño uniforme, sin formar bandas 
sobre los cordones. Las formas con 
banda blanca del estrecho de Gibraltar y 
de Marruecos fueron identificadas por 
PALLARY (1902) con Chauvetia decorata 
Monterosato, 1889. Este patrón de color 
de la concha existe en al menos cuatro 
especies de la zona y, teniendo en 
cuenta que Pallary se correspondía con 
Monterosato durante su estudio de la 
fauna de Marruecos, podríamos pensar 


que su interpretación (Fig. 40) ha de ser 
la correcta. Sin embargo, cuesta aceptar 
que MONTEROSATO (1889), en el auge de 
su trayectoria malacológica, describiese 
en el mismo trabajo dos variantes de 
color bajo nombres genéricos (Donovania 
y Chauvetia) distintos, por lo que identi- 
ficamos C. decorata con otro taxón des- 
crito más adelante. 

Estos ejemplares con banda blanca 
coinciden con los demás de C. procerula 
en la protoconcha con cordoncillos 
anchos, así como en el color completa- 
mente negro del animal. Por esta razón, 
consideramos  tentativamente estos 
ejemplares dentro de la variabilidad de 
C. procerula, a falta de otros caracteres 
diferenciales que se puedan correlacio- 
nar con el color de concha. 


41 


Iberus, 28 (1), 2010 


MICALI (1999: 58) figuró ilustró un 
animal vivo de Aci Trezza (Sicilia) atri- 
buido a C. procerula. Sin embargo, este 
animal se diferencia por un color del 
cuerpo amarillento con puntos brillan- 
tes, completamente distinto de lo obser- 
vado en los ejemplares del sur de la 


Península Ibérica. Cabe la posibilidad 
de que sea una especie distinta, pero 
hay que ser prudente en la interpreta- 
ción de este carácter, puesto que hemos 
observado ejemplares con la pigmenta- 
ción negra atenuada O ausente en 
algunas muestras del Algarve. 


Chauvetia recondita (Brugnone, 1873) (Figs. 46-47, 48-51, 103) 


Lachesis recondita Brugnone, 1873. Misc. Malac.: 10, fig. 15. [Localidad tipo: Palermo, Sicilia, fósil 
del Plio-Pleistoceno]. 

Lachesis vulpecula Monterosato, 1874. J. Conchy]l.: 276. [Localidad tipo: Capo San Vito, Sicilia]. 

? Chauvetia vulpecula attenuata Nordsieck, 1976. [Preocupado por Donovania minima var. attenuata 
Bucquoy, Dautzenberg y Dollfus, 1883]. 


Material tipo: Dos sintipos de Lachesis vulpecula en MNHN, col. Locard ex Monterosato (4,8 x 2,2 
- 5,2 x 2,3 mm). Los tipos de L. recondita podrían estar en la col. Jeffreys del USNM. 

Material estudiado: Islas Baleares: Ibiza (Bleda Mayor), 4 j., MNCN,; Ibiza (Cala Eubarco), 2 j., 
MNCN. Levante ibérico: Denia, 10 c. (5,3 x 2,1 — 6,4 x 2,6 mm) + 13j., col. JDO; Cullera, 5 c. + 23. 
(5,9 x 2,7 — 5,8 x 2,2 mm), col. JDO; Columbretes 1c. +1 j., MNCN. Sicilia: Banco Skerki (37 53,6' 
N, 10” 48,6' E, 113 m), 1 c. (4,8 x 2,1 mm), leg. Taviani 12-1996, Museo di Zoologia, Bologna. Mar 
de Alborán y estrecho de Gibraltar: Dorsal de la isla de Alborán (35” 58' N, 02” 58' W), 43., MNCN; 
isla de Alborán, 16 c. (5,5 x 2,7 — 6,5 x 2,9 mm) + 50j., col. JDO; Cádiz, 1 c. (5,6 x 2,5 mm), col. Azpei- 
tia, MNCN; Ceuta, Punta Almina, 25-40 m, 1 ej. +5 c. (4,5 x 2,3 - 5,5 x 2,5 mm, ejemplar dibujado); 
Ceuta, Almadraba 20-36 m, 2 c. (5,9 x 2,7 mm); M'diq (35” 41,5 N, 05” 12,0” W, 200 m), 1 c. (4,7 x 
2,2 mm), col. “Al Mounir”, MNHN; Marbella (Málaga), 1 j.; Estrecho de Gibraltar [sin más preci- 
sión], 3j. Portugal: Algarve (36” 58,0” N, 08% 55,6' W, 65 m), 1 c. (5,2 x 2,4 mm), leg. Péres, Gautier, 


Vacelet 7-8-1957, MNHN,; Tavira, 15 ej. (3,7 x 1,8 — 5,8 x 2,4 mm). 


Descripción: Concha de hasta 6,5 mm, 
con 5*/2-6 vueltas de espira convexas, 
con la sutura muy marcada. 

Protoconcha de 620-650 um de diá- 
metro máximo (núcleo: 350-400 um, 
primera media vuelta: 500 um). Desde el 
mismo núcleo aparece una escultura de 
cordoncillos altos, bien visibles en micros- 
copía óptica, de anchura equivalente a los 
interespacios, en el fondo de los cuales se 
aprecian diminutas laminillas transver- 
sales paralelas a las líneas de crecimiento; 
hay 16-18 cordoncillos al final de la 
primera media vuelta. Desde el inicio de 
la segunda media vuelta se añade una 
escultura de costillas axiales algo sigmoi- 
deas y opistoclinas, altas, más estrechas 
que los espacios que las separan, sobre las 
cuales continúan los cordoncillos. 

Teleoconcha con ornamentación 
formada por cordones espirales altos, algo 
más estrechos que sus interespacios, y por 
costillas axiales sobre las cuales los cor- 
dones discurren formando tubérculos a 


42 


modo de perlas. En la primera vuelta apa- 
recen cuatro cordones espirales que se 
mantienen a lo largo de la espira. En la 
última vuelta, hay 14-15 cordones, de los 
cuales 4 continúan los de la penúltima 
vuelta, los 5 medios son de un tamaño 
equivalente y, en la parte abapical, los cor- 
dones decrecen algo en tamaño. Las cos- 
tillas axiales son altas y más estrechas que 
los interespacios y se distribuyen aleato- 
riamente entre vueltas sucesivas. En la 
penúltima vuelta se aprecian 9-10 costillas 
axiales algo prosoclinas. La última vuelta 
alcanza el 60% de la altura total, mientras 
que la abertura ocupa en torno al 40%. 

Exterior del labio muy engrosado en 
los ejemplares adultos, con los cordones 
continuados sobre la variz externa. En el 
interior del labio se observan, en los 
ejemplares adultos, hasta 6-7 dentículos 
poco pronunciados, alargados hacia el 
interior de la abertura. 

El color de la concha es castaño, en 
algunos casos amarillento claro, frecuen- 


GOFAS Y OLIVER: Las especies de Chauvetía en el área ¡ibero-marroquí 


Figuras 48-51. Chauvetia recondita (Brugnone, 1873). 48, 49, ejemplar de Punta Almina, Ceuta 
(5,2 mm); 50, 51: ejemplares de la isla de Alborán (5,5 y 4,5 mm). 
Figures 48-51. Chauvetia recondita (Brugnone, 1873). 48, 49, specimen from Punta Almina, Ceuta 
(5.2 mm); 50, 51: specimens from Alborán Island (5.5 and 4.5 mm). 


temente con una línea más oscura a lo 
largo de los cordones espirales. El animal 
es de color negro moteado, tendiendo a 
más pálido en la parte anterior del pie, la 
base del sifón y los tentáculos cefálicos. 


Distribución: Conocida en el Medite- 
rráneo desde la costa oriental de Sicilia 
hasta el mar de Alborán y, en el Atlán- 
tico, desde el cabo de San Vicente hasta 
Marruecos. 


Chauvetia candidissima (Philippi, 1836) (Figs. 52-54, 63-64) 


Buccinum candidissimum Philippi, 1836. Enum. Moll. Siciliae, vol. 1: 222, lám. 11, fig. 18. [Localidad 


tipo: Catania, Sicilia]. 


Material estudiado: Sicilia: Localidad sin precisar, 1 c. (9,6 x 4,4 mm), col. Petit, MNHN; Messina, 
1 ej. (9,0 x 3,8 mm), col. A. Peñas. Estrecho de Gibraltar: Ceuta, Benzú, 20 c. (8,4 x 3,7 - 9,1 x 3,9 
mm); Cádiz, 1 c. (9,4 x 3,9), col. Gavala en col. Azpeitia, MNCN. Marruecos: Asilah, Oued el Helou, 
13., col. SG, MNHN; Mohammedia, 2 c. (8,5 x 3,7 - 8,6 x 3,8 mm), col. SG, MNHN. 


Descripción: Concha de hasta 9,1 mm, 
sólida, con unas seis vueltas de espira 
poco convexas y una sutura moderada- 
mente marcada. 

Protoconcha de 820 um de diámetro 
máximo (núcleo: 450-500 um, primera 
media vuelta: 625 um). El núcleo es liso, 
y al final de la segunda media vuelta 
aparece una escultura de costillas 
axiales algo sigmoideas y opistoclinas; 
no se aprecia escultura espiral. 

Teleoconcha con ornamentación 
formada por cordones espirales, el doble 


de anchos que sus interespacios, y costi- 
llas axiales de poco relieve que forman 
nódulos cuadrangulares aplanados, a 
modo de adoquines, al cruzarse con los 
cordones. En la primera vuelta aparecen 
dos cordones espirales; un tercer cordón 
aparece en la tercera vuelta por desdo- 
blamiento del cordón subsutural; este 
cordón adapical se vuelve a ensanchar y 
se desdobla de nuevo en la última 
vuelta. En la última vuelta, hay unos 15 
cordones, de los cuales 4 continúan los 
de la penúltima vuelta, los 5 medios son 


A3 


Iberus, 28 (1), 2010 


Figuras 52-54. Chauvetia candidissima (Philippi, 1836). 52, 53, ejemplar de Benzú, Ceuta (8,6 


mm); 54, ejemplar de Messina, Sicilia, col. A. Peñas (9,0 mm). 
Figures 52-54. Chauvetia candidissima (Philippi, 1836). 52, 53, specimen from Benzú (8.6 mm); 54, 
specimen from Messina, Sicilia, col. A. Peñas (9.0 mm). 


de tamaño equivalente y los que cubren 
el canal sifonal son algo más delgados y 
dejan de ser granulosos. La terminación 
de las costillas no se manifiesta apenas 
sobre la sutura. En la penúltima, así 
como en la última vuelta, se aprecian 22- 
24 costillas axiales algo prosoclinas. La 
última vuelta alcanza en torno al 60% de 
la altura total, mientras que la abertura 
ocupa en torno al 42%. 

Exterior del labio engrosado en los 
ejemplares adultos con los cordones 
continuados. En el interior del labio se 
observan, en los ejemplares adultos, 
hasta 6-7 dentículos alargados hacia el 
interior de la abertura. 

El color de la concha es blanco. El 
animal no se ha observado. 

Distribución: Sicilia, Malta, Argelia 
(MICALI, 1999) y estrecho de Gibraltar; 
las citas de Senegal corresponden a 
otras especies (véase OLIVER Y ROLÁN, 
2008). 

Notas: Chauvetia tenuisculpta (Daut- 
zenberg, 1890) es un endemismo de 


44 


Senegal, que se distingue por tener 
unas costillas axiales muy fuertes desde 
la primera media vuelta de la protocon- 
cha. Las citas de dicha especie para el 
estrecho de Gibraltar (MICALI, 1999; 
OLIVER Y ROLÁN, 2008, con dudas) nos 
parecen dudosas y tal vez estén basadas 
en ejemplares de C. candididissima del 
Estrecho, como los que describimos 
aquí. Estos ejemplares del Estrecho se 
parecen más, por su forma general, a C. 
tenuisculpta, pero tienen la mayor parte 
de la protoconcha lisa, como los ejem- 
plares de C. candidissima de Sicilia. Sin 
embargo, los ejemplares de Sicilia que 
hemos visto se diferencian por tener las 
vueltas mucho más convexas y, sobre 
todo, los cordones mucho más marca- 
dos en el exterior del labio. Es posible 
que la forma del Estrecho sea una 
especie distinta, pero haría falta un 
seguimiento de su variabilidad a lo 
largo de las costas norteafricanas, si es 
que allí existe, antes de llegar a una 
conclusión. 


GOFAS Y OLIVER: Las especies de Chauvetia en el área ibero-marroquí 


Chauvetia lefebvrii (Maravigna, 1840) (Figs. 55-58, 65-68) 


Buccinum lefebvrii Maravigna, 1840. Rev. Zool. Soc. Cuvier.: 325. [Localidad tipo: Aci Trezza, 
Sicilia]. 

Chauvetia lefebvrei auct. [Ortografía subsiguiente incorrecta, iniciada por Locard, 1892: 72]. 

Fusus granulatus Calcara, 1839, non Nesaea granulata Risso, 1826. Ric. Malac.: 16, fig. 10. [Localidad 
tipo: Palermo, Sicilia]. 

Buccinum folineae, sensu Philippi, 1844, non Murex folineae Delle Chiaje, 1828. En. Moll. Sic. vol. 2: 
189, lám. 27, fig. 10. 

Lachesis areolata Tiberi, 1868. J. Conchy]l.: 73. [Localidad tipo: Sicilia]. 

? Folinia retifera var. glomulus Monterosato, 1889. J. Conchyl., 37: 117. [Localidad tipo: Casablanca]. 

? Folinia retifera var. labrosa Monterosato, 1889. J. Conchyl., 37: 117. [Localidad tipo: Casablanca]. 

Chauvetia pellisphocae sensu Nordsieck, 1976, non Pleurotoma pellisphocae Reeve, 1845. 


Material estudiado: Sicilia: Capo dei Mulini, 10 c. (5,6 x 2,7 — 6,4 x 2,8 mm), MNHN,; Isola Lachea, 
1 c. (6,0 x 3,7 mm), MNHN. Argelia: Argel, 9 c. (5,8 x 2,8 — 6,6 x 3,1 mm), col. Hidalgo, MNCN; 
Argel, 5 c. (5,7 x 2,8-—7,4 x 3,3 mm), col. Azpeitia, MNCN; Argel, 3 c. (7,2 x 3,2 -7,5 x 3,2 mm), col. 
Pallary, MNHN,; Dellys, 4 c. (6,8 x 3,1 mm), col. Pallary, como L. folineae det. Dautzenberg, MNHN. 
Baleares: Ibiza (Punta Galera), 2 protoconchas, MNCN; Mallorca (Punta Foradada), 2 c. (deterio- 
radas) + 2 fragmentos + 2 protoconchas, MNCN; Mar de Alborán y estrecho de Gibraltar: La Herra- 
dura (Granada), 50 ej., MNCN; Mijas (Málaga), 4 ej., MNCN; Algeciras, 30 c., MNCN; Getares 
(Cádiz), 2 c. (6,5 x 3,1 mm -7,8 x 3,5 mm), col. SG, MNHN; Tarifa, 4 c. (8,5 x 3,6 — 9,0 x 3,9 mm), 
col. Gavala en col. Azpeitia, MNCN,; Barbate, 1 c. (6,8 x 3,1 mm), col. SG; Ceuta, Benzú, 0-4 m, 3 
ej. (9,0 x 3,8 mm - 10,0 x 4,0 mmy ejemplar dibujado); Ceuta, Benzú, 24 c. (8,9 x 3,7 - 10,3 x 4,3 mm), 
col. SG, MNHN. Forma de color castaño uniforme: Marruecos: Asilah, 3 c. (6,8 x 3,2 — 7,0 x 3,5 mm), 
col. SG, MNHN; Temara, 33 ej. (5,8 x 2,8 - 8,5 x 3,7 mm) + 15j., col. SG, MNHN; Temara, 18 c. (8,5 
x 3,8 mm) MNCN; Mohammedia, 2 c. (8,9 x 4,0 mm), col. SG, MNHN; Casablanca, 7 c. (arrojadas 
a la playa), col. Rigotard 1917, MNHN,; El Jadida, 52 ej. (8,6 x 3,7 - 10,1 x 4,4 mm) + 12j., col. SG, 
MNHN, Essaouira, 60 ej. (8,0 x 3,7 - 9,4 x 4,0 mm) + 29j., col. SG, MNHN,; Essaouira 6 c. + 1j. (8,2 
x 3,7 mm), MNCN. Estrecho de Gibraltar: Tarifa, 34 c. (7,9 x 33 — 11,1 x 4,3 mm) + 4j. + 3 proto- 


conchas, col. Gavala en col. Azpeitia, MNCN. 


Descripción: Concha de hasta 11,1 
mm, sólida, con unas seis vueltas de 
espira poco convexas, con la sutura 
moderadamente marcada. 

Protoconcha de 900 um de diámetro 
máximo (núcleo: 500 um, primera media 
vuelta: 750 um). El núcleo es liso o casi 
liso; desde la primera media vuelta se 
pueden apreciar unos cordoncillos espi- 
rales muy débiles, de tamaño irregular, 
alternando gruesos y finos. En la 
segunda media vuelta aparece una escul- 
tura de costillas axiales algo sigmoideas y 
opistoclinas, de anchura similar a los 
espacios que las separan, sobre las cuales 
se continúan los cordoncillos. 

Teleoconcha con ornamentación 
formada por cordones espirales, el doble 
de anchos que sus interespacios, y costi- 
llas axiales de poco relieve que forman 
nódulos cuadrangulares aplanados, a 
modo de adoquines, al cruzarse con los 
cordones. La terminación de las costillas 


repercute un poco sobre la sutura, que 
es algo canaliculada. En la primera 
vuelta aparecen tres cordones espirales; 
un cuarto cordón aparece en la tercera 
vuelta por desdoblamiento del cordón 
subsutural; este cordón adapical se 
vuelve a ensanchar y se desdobla de 
nuevo en la última vuelta. En la última 
vuelta, hay 17-18 cordones, de los cuales 
4 continúan los de la penúltima vuelta, 
los 5 medios son de tamaño equivalente 
y los que cubren el canal sifonal decre- 
cen marcadamente en grosor. En la 
penúltima vuelta, así como en la última, 
se aprecian 20-24 costillas axiales algo 
prosoclinas. La última vuelta alcanza el 
56-58% de la altura total, mientras que 
la abertura ocupa en torno al 40%. 
Exterior del labio muy engrosado en 
los ejemplares adultos, particularmente 
en su parte adapical; cordones muy ate- 
nuados en su parte externa. En el inte- 
rior del labio se observan 4-5 dentículos, 


45 


Iberus, 28 (1), 2010 


Figuras 55, 56. Chauvetia lefeburii (Maravigna, 1840), con patrón de color con nódulos oscuros, 
habitual en el Mediterráneo y en el estrecho de Gibraltar, Benzú, Ceuta (9,2 mm). Figuras 57, 58. 
Chauvetia cf. lefebvriz, forma de color castaño uniforme propia de la costa atlántica de Marruecos, 
Temara (8,2 mm). 
Figures 55, 56. Chauvetia lefebvrii (Maravigna, 1840), the colour pattern with dark nodes, usual in the 
Mediterranean and Strait of Gibraltar, Benzú (9.2 mm). Figures 57, 58. Chauvetia cf. lefebvrii, with pat- 
tern of continuousdark bands over the cords, as found on the Atlantic coast of Morocco, Temara (8.2 mm). 


de los cuales el adapical es más pronun- 
ciado, y el abapical constituye el borde 
externo del canal sifonal. 

El color de la concha es blanco 
rosáceo con puntos de color castaño 
rojizo en la intersección de los cordones 
espirales con las costillas axiales; estos 
puntos son más pálidos en las primeras 
vueltas y al principio tienden a confluir 
a lo largo de los cordones. En la última 
vuelta, se dibujan bandas castañas sobre 
el borde externo del labio, en continui- 
dad con las filas espirales de puntos. 

El animal es blanquecino con puntos 
de color blanco lácteo. 

Distribución: Conocida en el Medite- 
rráneo desde la costa oriental de Sicilia 
hasta el mar de Alborán, y el estrecho de 
Gibraltar (forma con nódulos oscuros) y 
en la costa atlántica de Marruecos 
(Chauvetia cf. lefebvrii, forma de color 
castaño uniforme). No hemos encon- 
trado esta especie en el abundante mate- 


46 


rial del Algarve, y la cita para la ría de 
Vigo (ROLÁN, 1983) corresponde en 
nuestra opinión a C. retifera. 

Notas: La sinonimia de esta especie 
fue establecida ¡por MONTEROSATO 
(1884) teniendo en mano la colección de 
Tiberi. De ello se entiende que TIBERI 
(1868), al considerar Buccinum lefeborii 
una sinonimia de Lachesis mamillata, 
volvió a describir la verdadera C. lefebv- 
rii como especie nueva (L. areolata). 

La descripción anterior se aplica a 
poblaciones del Mediterráneo y de las 
costas del Estrecho. En la costa atlántica 
de Marruecos se encuentra una forma 
de color castaño uniforme que MICALI 
(1999) considera una variedad de C. reti- 


fera. En nuestra opinión, la protoconcha, 


el perfil de la espira, la configuración de 
los tubérculos a modo de adoquines 
desde las primeras vueltas y los cuatro 
cordones situados sobre la abertura en 
la última vuelta se ajustan más a C. 


GOFAS Y OLIVER: Las especies de Chauvetía en el área ibero-marroquí 


Figuras 59-62. Chauvetia retifera (Brugnone, 1880). 59, 60: ejemplar de Benzú, Ceuta (9,4 mm); 
61, 62: ejemplar de Sagres, 3-15 m (9,5 mm). 

Figures 59-62. Chauvetia retifera (Brugnone, 1880). 59, 60: specimen from Benzú (9.4 mm); 61, 62: 
specimen from Sagres, 3-15 m (9.5 mm). 


lefeburii que a C. retifera. A esta forma de distinta, alguno de los nombres pro- 
coloración se le puede aplicar, si se puestos por MONTEROSATO (1889) como 
demostrase que se trata de una especie variedades de C. retifera. 


Chauvetia retifera (Brugnone 1880) (Figs. 59-62, 70-72, 104) 


Lachesis retifera Brugnone, 1880. Bull. Soc. Malac. Ital.: 111, lám. 1, fig. 6. [Localidad tipo: Giannet- 
tello, cerca de Caltanisetta, Sicilia; fósil plio-pleistocénico]. 

Lachesis dolioliformis Monterosato, 1884. Nom. Gen. Spec.; 137. [Nombre introducido en la sinoni- 
mia de Chauvetia retifera y nunca usado como válido; por lo tanto, no disponible en nomencla- 
tura]. 

Folinta retifera var. lirifera Monterosato, 1889. J. Conchyl.: 117. 

Donovania (Adansonia) pellisphocae sensu Pallary, 1920, non Pleurotoma pellisphocae Reeve, 1845. 

Chauvetia elongata Nordsieck y Talavera, 1979. [Localidad tipo: sur de Gran Canaria, probable- 
mente un error, según Oliver y Rolán, 2009: 152]. 


Material estudiado: Galicia: Bayona: 1 c., (7,8 x 3,1 mm), MNCN. Portugal: Peniche, 1 c. (arrojada 
a la playa, 10,5 x 4,2 mm), col. SG, MNHN,; Sagres, Punta da Baleeira, 17-23 m, 31 ej. (8,4 x 3,7 — 
10,0 x 3,7 mm) + 6 j.; Sagres, Ponta dos Caminhos, 23-33 m, 4 c. (9,0 x 3,7 - 11,0 x 3,7 mm); Sagres, 
3-15 m, 1 c. (9,7 x 3,8 mm); Sagres, Pontal dos Corvos, 2 c. (8,7 x 3,8 — 9,3 x 3,7 mm); Sagres, Praia 
do Martinhal, bajamar, 1 c. (10,3 x 3,7 mm); Salema, pesca de arrastre, 1 ej. (dibujado); Tavira, Pedra 
do Barril, 13 c. (8,6 x 3,5 — 10,6 x 3,7 mm); Tavira, Cabanas, 3 c. (10,5 x 3,8 mm). Mar de Alborán y 
estrecho de Gibraltar: cala Higuera (Almería), 1 c. (sin protoconcha), col. Cobos, MNCN,; Benal- 
mádena (Málaga), 1 c. (10,4 x 4,0 mm), col. SG; Tarifa, isla de Tarifa, 2 ej. (10,0 x 3,8 mm) + 1j., col. 
SG; Tarifa, Torre de la Peña, 1 c., col. SG, MNHN; Tarifa, 20 c. (8,9 x 3,7 — 12,5 x 4,4 mm), col. Gavala 


47 


Iberus, 28 (1), 2010 


en col. Azpeitia, MNCN; Barbate, 2 c. (8,8 x 3,5 mm - 9,8 x 3,8 mm), col. SG; Cádiz, La Cortadura, 
2 c. (8,0 x 3,4 mm), col. SG, MNHN,; Cádiz, 1 c. (8,4 x 3,4 mm), col. Gavala en col. Azpeitia, MNCN; 
Ceuta, Punta Almina, 25-40 m, 4 c. (8,6 x 3,5 - 8,6 x 3,6 mm); Ceuta, Benzú, 0-4 m, 2 ej. (9,6 x 3,6 
mm -— 10,3 x 3,8 mm; ejemplar dibujado); Ceuta, Benzú, 46 c. (8,5 x 3,5 - 11,2 x 3,8 mm), col. SG, 
MNHN. Marruecos: Tánger, muelle Este del puerto, 14 ej. + c. (9,0 x 34 — 11,5 x 44 mm) + 3j., 
MNHN, Asilah, 3 c. (10,0 x 3,7 mm), MNHN; Temara, 5 ej. (9,2 x 3,5 — 9,9 x 3,8 mm) +5 ]., col. SG, 
MNHN; Temara, 5 c. (8,1 x 3,5 — 9,9 x 3,7 mm) + 2j., MNCN; Casablanca 9 c. (arrojadas a la playa), 
col. Rigotard 1917, MNHN,; Essaouira, 7 ej. (8,7 x 3,4 — 9,7 x 3,5 mm) + 2j., col. SG, MNHN,; Essa- 


ouira, 1j., MNCN. 


Descripción: Concha de hasta 12,5 
mm, sólida, con unas seis vueltas de 
espira altas, poco convexas, con la 
sutura moderadamente marcada. 

Protoconcha de 850 um de diámetro 
máximo (núcleo: 450 um, primera media 
vuelta: 650 um). El núcleo es liso o casi 
liso; desde la primera media vuelta 
aparece una escultura de costillas 
axiales muy fuertes, de anchura similar 
a los espacios que las separan, sin 
ningún rastro de escultura espiral. 

Teleoconcha con ornamentación 
formada por cordones espirales, el doble 
de anchos que sus interespacios, y costi- 
llas axiales de muy poco relieve que 
forman nódulos cuadrangulares aplana- 
dos, a modo de adoquines, al cruzarse 
con los cordones. La terminación de las 
costillas se manifiesta un poco sobre la 
sutura, que es algo canaliculada. En la 
primera vuelta aparecen tres cordones 
espirales; en la tercera vuelta aparece un 
cuarto cordón por desdoblamiento del 
cordón subsutural; este cordón adapical 
se vuelve a ensanchar y se desdobla de 
nuevo en la penúltima vuelta. En la 
última vuelta, hay 17-18 cordones, de 
los cuales 5 continúan los de la penúl- 
tima vuelta, los 4 medios son de grosor 
similar, y los que cubren el canal sifonal 
decrecen en grosor. En la penúltima 
vuelta, así como en la última, se apre- 
cian 20-24 costillas axiales algo prosocli- 
nas. La última vuelta alcanza el 56-58% 
de la altura total, mientras que la aber- 
tura ocupa algo menos del 40%. 

Exterior del labio muy engrosado en 
los ejemplares adultos, particularmente 
en su parte adapical; cordones muy ate- 
nuados en su parte externa. En el inte- 
rior del labio se observan 4-5 dentículos, 
de los cuales el adapical es más pronun- 
ciado y está bastante separado del 


48 


extremo de la abertura, y el abapical 
constituye el borde externo del canal 
sifonal. 

El color de la protoconcha es blanco, 
el de la teleoconcha es blanco rosáceo o 
amarillento, más pálido en las primeras 
vueltas, con bandas de color castaño 
rojizo que cubren completamente los 
cordones espirales sin interrumpirse en 
los intervalos y se continúan sobre el 
borde externo del labio. 

El animal es blanquecino, con 
algunos puntos de color blanco lácteo 
en la parte alta del cuerpo. 

Distribución: Mediterráneo, sola- 
mente en el mar de Alborán; Atlántico, 
desde Galicia hasta Marruecos. No se ha 
encontrado en Canarias en el extenso 
material examinado por Oliver y Rolán 
(2009). 

Notas: Esta especie fue citada por 
PALLARY (1902, 1920) con el nombre 
erróneo de Chauvetia pellisphocae (Reeve, 
1845) pero, según Maes (1983), este 
nombre específico corresponde a una 
especie del Caribe, perteneciente al 
género Crassispira (superfamilia Conoi- 
dea) y superficialmente parecida. Chau- 
vetia retifera se parece mucho a C-. lefebv- 
rit y coexiste con ella en gran parte de su 
área de distribución. Las dos especies 
tienen en común una concha gruesa con 
vueltas poco convexas, el tamaño relati- 
vamente grande de la protoconcha, el 
color blanquecino del animal, y la exis- 
tencia de dentículos fuertes en la aber- 
tura. Chauvetia retifera se distingue, sin 
embargo, por tener un perfil más alto, 
por el dentículo adapical del labio más 
apartado de lo alto de la abertura y por 
la protoconcha que carece de microes- 
cultura espiral y tiene fuertes costillas 
axiales desde la primera media vuelta. 
En este último aspecto, se parece a la 


GOFAS Y OLIVER: Las especies de Chauvetía en el área ibero-marroquí 


Figuras 63-72. Protoconchas en vista lateral y apical. 63, 64: Chauvetia candidissima, Benzú, 
Ceuta; 65: Chauvetia lefebvrii, forma con nódulos oscuros, Cádiz; 66, 67: Chauvetia lefebvriz, 
forma con nódulos oscuros, La Herradura, Granada. 68, 69: Chauvetia cf. lefebvrii, forma con 
color uniforme, Temara. 70-72: Chauvetia retifera, La Herradura, Granada. 

Figures 63-72. Protoconchs in lateral and apical views. 63, 64: Chauvetia candidissima, Benzú; 65: 
Chauvetia lefebvrii, form with dark nodes, Cádiz; 66, 67: same, La Herradura. 68, 69: Chauvetia cf. 


lefebvrii, with uniform colour pattern, Temara. 70-72: Chauvetia retifera, La Herradura. 


especie senegalesa C. tenuisculpta, pero 
ésta difiere por su color completamente 
blanco. El color con bandas castañas 
continuas en los cordones espirales es 
otro carácter distintivo en las costas 1bé- 


ricas, pero en el litoral marroquí existe 
este mismo patrón de color también en 
formas que asignamos a C. cf. lefebvrii, 
manteniéndose las diferencias citadas en 
la protoconcha y el perfil de la concha. 


49 


Iberus, 28 (1), 2010 


Chauvetia dentifera spec. nov. (Figs. 73-76, 85-88, 105) 


Material tipo: Holotipo [ej., 6,9 x 3,2 mm], MNHN 22872, y 10 paratipos (5 ej. MNHN 226873, 5 ej. 
MNCN 15.05/53587), todos de la localidad tipo. 

Material estudiado: Mar de Alborán y estrecho de Gibraltar: Mijas Costa (Málaga), 1 c. + 1j., col. 
A. Peñas; Sotogrande (Cádiz), 1 ej. (8,0 x 3,5 mm), col. SG, MNHN,; Barbate, 1 c. j., col. SG; Cádiz, 
10c.+1j. (7,1 x3,1-—7,8 x 3,9 mm), col. Azpeitia, MANCN; M dig, 1 c. (5,5 x 2,8 mm), col. SG, MNHN; 
Ceuta, La Almadraba, 1 c.j.; Ceuta, Punta Almina, 25-40 m, 32 ej. y c. (6,7 x 3,2 — 7,8 x 3,5 mm) y 
18 j.; Ceuta, puerto, 1 c. (11,0 x 319 mm); Ceuta, Benzú, 16 c. (5,6 x 2,7 — 6,3 x 3,0 mm), col. SG, 
MNHN. Portugal: Cascais, B /O “Faial” P3 (38 24,1" N, 09% 14,1! W, 47-50 m), 5 c. (6,0 x 3,0-7,0 x 
3,3 mm), col. Péres, Gautier, Vacelet, 5-8-1957, MNHN,; Sines, 1 c. (arrojada a la playa) (7,2 x 3,0 
mm), col. Locard, MNHN,; Sagres, Ponta da Baleeira, 17-23 m, 148 ej. (6,5 x 3,1 -7,8 x 3,4 mm) + 28 
j.; Sagres, Baia da Baleeira, 3-15 m, 7 ej. (7,3 x 3,2 - 8,3 x 3,5 mm) + 1j.; Sagres, Ponta dos Caminhos, 
20 c. (7,3 x 3,3 - 8,2 x 3,4 mm); Sagres, Pontal dos Corvos, 16 ej. + 2j. (6,8 x 3,2 - 8,0 x 3,0 mm); 
Tavira, Pedra do Barril, 23 ej. (6,4 x 3-7,7 x 3,3 mm) + 4j.; Tavira, Cabanas, 1 ej. (7,8 x 3,3 mm) + 
2 j. Marruecos: Asilah, 1 c.j., col. SG, MNHN; Rabat, “Vanneau” sta. 37, 33 59 N, 07” 50” W, 155 
m, 6 c. (5,5 x 2,8 — 6,2 x 2,9 mm) + 3j., MNHN,; Safi, 10 km sur ciudad, 3 c. (8,0 x 3,4 mm), col. J. de 
Lepiney, MNHN. Sáhara: 22” 35' N, 16” 58' W, 1c;23*05' N, 16” 25” W, 2 c. Mauritania: Cabo Blanco, 


80 m, 1 c. (fotografía en Oliver y Rolán, 2009: fig. 24). 
Localidad tipo: Punta Almina, Ceuta (35* 54,1" N — 05? 16,5" W, 25-40 m). 
Etimología: el nombre específico alude al dentículo labial propio de esta especie. 


Descripción: Concha fusiforme algo 
pupoide, sólida, con 5!*/2-6 vueltas de 
espira y hasta 7,8 mm. Las primeras 
vueltas son regularmente convexas, con 
una sutura bastante marcada, la penúl- 
tima y la última tienen el máximo de 
convexidad cerca de la sutura y la parte 
media más aplanada. 

Protoconcha con algo más de una 
vuelta, diámetro máximo de 650-700 um 
(núcleo: 380-400 um, primera media 
vuelta: 530-550 um). El núcleo tiene cor- 
doncillos espirales muy tenues, casi ina- 
preciables con microscopía óptica, desi- 
guales entre sí, con interespacios de 
aspecto poroso a mucho aumento; a 
partir de la segunda media vuelta 
aparece una escultura de costillas 
axiales algo sigmoideas y opistoclinas, 
de anchura similar a los espacios que las 
separan. En los espacios se puede apre- 
ciar la continuación de los cordoncillos 
espirales muy débiles y de tamaño irre- 
gular, alternando gruesos y finos. 

Teleoconcha con ornamentación 
formada por cordones espirales y costi- 
llas axiales. En la primera vuelta apare- 
cen cuatro cordones espirales bien marca- 
dos, los dos adapicales algo más estre- 
chos; a partir de la tercera vuelta, el 
cordón subsutural se desdobla, mientras 
que los cordones abapicales se ensanchan 


50 


y aplanan, siendo en esta parte mucho 
más anchos que los interespacios. En la 
última vuelta los cordones de la parte 
media están tan aplanados que llegan a 
ser indistintos; la parte abapical está 
separada del resto de la vuelta por un 
ligero surco y lleva 15-16 cordones algo 
más patentes, decreciendo algo en 
tamaño al acercarse al canal sifonal. En la 
penúltima vuelta se aprecian 11-12 costi- 
llas axiales, y en la parte final de la última 
vuelta las costillas se van atenuando. 

La última vuelta alcanza el 60% de la 
altura total, mientras que la abertura 
ocupa en torno al 40% 

Abertura oval, con un canal sifonal 
muy corto y ancho. Labio engrosado en 
los ejemplares adultos, particularmente 
en su parte adapical; el lado externo 
forma una variz suave, no delimitada 
del resto de la vuelta. En el interior del 
labio de los ejemplares completamente 
adultos, se observan 4-5 dentículos poco 
marcados, alargados y situados muy 
adentro de la abertura; sin embargo, el 
interior de la abertura de los ejemplares 
subadultos es frecuentemente liso. La 
parte abapical del labio forma en su 
borde un dentículo saliente, situado en 
la terminación del surco externo de la 
última vuelta (Fig. 76) y apuntando en 
la dirección de crecimiento del borde 


GOFAS Y OLIVER: Las especies de Chauvetía en el área 1ibero-marroquí 


Figura 73-76. Chauvetia dentifera spec. nov. 73, 74, holotipo de Punta Almina, Ceuta, 32-40 m 
(6,9 mm); 75, 76, ejemplar de Sotogrande (8,0 mm). 
Figure 73-76. Chauvetia dentifera spec. nov. 73, 74, holotype from Punta Almina, Ceuta, 32-40 m 
(6.9 mm); 75, 76, specimen from Sotogrande (8.0 mm). 


El color de la concha es castaño en la 
mitad adapical de las vueltas y en la 
parte abapical de la última vuelta, con 
una zona amarillenta en la mitad abapi- 
cal de las vueltas de espira, continuada 
como una banda central en la última 
vuelta. 

El animal es blanquecino con puntos 
de color blanco lácteo. 

Distribución: Mediterráneo, sola- 
mente en la costa occidental de Málaga; 
Atlántico, desde el sur de Portugal hasta 
el cabo Blanco. 

Notas: Esta especie ha sido confun- 
dida (MICALI, 1999) con Chauvetia crassior 
(Odhner, 1932), descrita de la plataforma 
insular de Gran Canaria. El holotipo de 
C. crassior (véase OLIVER Y ROLÁN 2009: 
116-117, fig. 19-20), conservado en el 
SMNH, es subadulto y el carácter del 
dentículo labial puede no estar expresado 
por esta razón; sin embargo, muestra una 


diferencia considerable en la configura- 
ción de las costillas axiales, que son 
menos numerosas, más altas y más sepa- 
radas en lo que sería la penúltima vuelta. 
Los ejemplares adultos de Chauvetia cras- 
sior de Gran Canaria, ilustrados en 
OLIVER Y ROLÁN (2009, figs. 21-23) no 
llegan a desarrollar un dentículo labial 
aunque el labio esté muy engrosado. 

El dentículo labial observado en esta 
especie es un rasgo que se conoce entre 
numerosas especies de Caenogastro- 
poda (VERMEI, 2001), la mayoría de 
ellas, así como los casos más extremos, 
pertenecientes a la familia Muricidae. Se 
considera este rasgo como convergente 
en distintos linajes y relacionado con el 
comportamiento predador. Se señala 
aquí por primera vez en el género Chau- 
vetia, aunque de momento no se conoce 
el uso que el animal pueda hacer de este 
dentículo. 


ON 


Iberus, 28 (1), 2010 


Chauvetia taeniata spec. nov. (Figs. 77-80, 89-90) 


Material tipo: Holotipo [ej., 7,9 x 3,6 mm], MNHN 22870, y 10 paratipos (5 ej. MNHN 22871, 5 ej. 


MNCN 15.05/53586), todos de la localidad tipo. 


Material estudiado: Estrecho de Gibraltar: Cádiz, 10 c. + 23. (6,9 x 3,4- 9,0 x 3,7 mm), col. Gavala 
en col. Azpeitia, MNCN; Ceuta, La Almadraba, 3 c. (8,5 x 3,9 mm); Ceuta, Punta Almina, 25-40 m, 
24 ej. +c. (7,2 x 3,4 -7,9 x 3,6 mm; ejemplar dibujado); Ceuta, Benzú, 5 c. (6,5 x 3,3 -7,6 x 3,3 mm). 
Tánger, 1 c. (7,8 x 3,3 mm), vol. SG, MNHN. Portugal: Sagres, Ponta da Baleeira, 17-23 m, 20 ej. (7,2 
x 3,2 —7,8 x 3,6 mm) + 9j.; Sagres, Ponta dos Caminhos, 6 c. + 1j. (7,6 x 3,5-8,1 x 3,5 mm); Sagres, 
Pontal dos Corvos, 5 ej. (8,2 x 3,6 - 8,5 x 3,7 mm); entre Faro y Sagres, redes de pesca, 1j.; Tavira, 
Pedra do Barril, 21 ej. (7,0 x 3,2 - 7,8 x 3,6 mm) + 12j. 

Localidad tipo: Punta Almina, Ceuta (35 54.1” N — 05? 16.5” W, 25-40 m). 

Etimología: Del latín taenia, venda o cinta, aludiendo a la forma aplanada de los cordones. 


Descripción: Concha  fusiforme, 
sólida, con 5!/2-6 vueltas y hasta 8,5 
mm. Las primeras vueltas son poco pero 
regularmente convexas, con una sutura 
bastante marcada, y crecen regular- 
mente en diámetro sin formar una 
espira cirtoconoide. 

Protoconcha con algo más de una 
vuelta, diámetro máximo de 750-800 um 
(núcleo: 400-420 um, primera media 
vuelta: 550-600 um). Desde el mismo 
núcleo aparece una escultura de cordon- 
cillos aplanados y anchos, muy desigua- 
les entre sí, mucho más anchos que los 
interespacios, en el fondo de los cuales 
se aprecian diminutas laminillas trans- 
versales irregulares; hay 15-18 cordonci- 
llos al final de la primera media vuelta. 
Al final de la segunda media vuelta se 
añade una escultura de costillas axiales 
estrechas algo sigmoideas y opistocli- 
nas, de anchura progresivamente mayor 
hasta el principio de la teleoconcha; en 
esta parte, los cordoncillos espirales se 
vuelven más altos y pasan a ser más 
estrechos que los interespacios. 

Teleoconcha con ornamentación for- 
mada por cordones espirales y costillas 
axiales. En la primera vuelta de la teleo- 
concha aparecen cuatro cordones espira- 
les bien marcados, aumentando algo en 
tamaño hacia el ápice; a partir de la ter- 
cera vuelta, el cordón subsutural se des- 
dobla, de modo que hay cinco cordones 
en la penúltima vuelta. En la última 
vuelta, los cordones de la parte media se 
ensanchan y aplanan, siendo en esta par- 
te mucho más anchos que los interespa- 
cios; en la parte abapical, los cordones 
pasan a ser más fuertes y más estrechos 


2 


que los interespacios, decreciendo algo 
en tamaño al acercarse al canal sifonal; 
hay en total 25-30 cordones en la última 
vuelta. En la penúltima vuelta se apre- 
cian 9-10 costillas axiales muy marcadas, 
y en la parte final de la última vuelta las 
costillas se van atenuando. 

La última vuelta alcanza el 60% de la 
altura total, mientras que la abertura 
ocupa algo más del 40% 

Abertura oval, engrosada en los 
ejemplares adultos, con canal sifonal 
relativamente corto y ancho. Labio 
grueso en los ejemplares adultos, parti- 
cularmente en su parte adapical; el lado 
externo forma una variz suave, no deli- 
mitada del resto de la vuelta, sobre la 
cual los cordones se atenúan o casi desa- 
parecen. En el interior del labio de los 
ejemplares completamente adultos, se 
observan 9-10 dentículos poco marca- 
dos, alargados y situados muy adentro 
de la abertura; sin embargo, el interior 
de la abertura de los ejemplares suba- 
dultos es frecuentemente liso. 

Protoconcha castaña muy clara; tele- 
oconcha castaña en la parte adapical de 
las vueltas, con una zona blanca en los 
dos tercios abapicales de las vueltas de 
espira, continuada como una banda 
ancha central en la última vuelta. 

El animal es blanquecino con puntos 
de color blanco lácteo. 

Distribución: Suroeste de Portugal y 
estrecho de Gibraltar. 

Notas: Esta especie tiene una proto- 
concha muy parecida a la de C. proce- 
rula, aunque algo más grande. La teleo- 
concha, sin embargo, es muy distinta y 
se caracteriza por la tendencia de los 


GOFAS Y OLIVER: Las especies de Chauvetia en el área ibero-marroquí 


Figuras 77-80. Chauvetia taeniata spec. nov. 77-79: holotipo de Punta Almina 32-40 m (7,9 mm); 
80: ejemplar de Cádiz (8 mm). 
Figures 77-80. Chauvetia taeniata spec. nov. 77-79: holotype from Punta Almina 32-40 m (7.9 mm); 
80: specimen from Cádiz (8 mm). 


cordones a hacerse aplanados y anchos, 
hasta casi desaparecer en la última 
vuelta. El patrón de color es parecido al 
de C. dentifera spec. nov. y C. decorata y 
los estadios juveniles podrían confun- 
dirse, pero los juveniles de C. dentifera 
son proporcionalmente más anchos y 
toscos, reflejando el perfil de espira algo 


cirtoconoide de ésta, y la protoconcha 
de C. dentifera tiene cordoncillos mucho 
más tenues, y el doble en número. Chau- 
vetia decorata tiene una protoconcha 
marcadamente más pequeña y las tres 
especies se separan con claridad en las 
localidades, como Punta Almina, donde 
ocurren juntas. 


Chauvetia decorata Monterosato, 1889 (Figs. 81-84, 91-94) 


Chauvetia decorata Monterosato, 1889. J. Conchy]l.: 117. [Localidad tipo: Casablanca, Marruecos]. 


Material estudiado: Estrecho de Gibraltar: Barbate, 1 c.j., col. SG; Cádiz 1 c. (7,2 x 3,1 mm), col. 
Gavala en col. Azpeitia, MNCN; Ceuta, La Almadraba, 1 c. (6,5 x 2,9); Ceuta, Punta Almina, 25-40 
m, 14 ej. y c. (5,9 x 3,2 — 6,8 x 3,3 mm); Ceuta, Benzú, 1 c. (6,6 x 3,2 mm). Marruecos: Safi, 10 km. 


sur ciudad, 2 c. (7,2 x 3,0 mm). 


Descripción: Concha fusiforme algo 
pupoide, sólida, con 5-5 */2 vueltas de 
espira y hasta 7,2 mm, y vueltas regular- 
mente convexas con una sutura bastante 
marcada. 

Protoconcha con aproximadamente 
una vuelta, diámetro máximo de 600-620 
um (núcleo: 330-350 um, primera media 
vuelta: 480-500 um). El núcleo es promi- 
nente y tiene cordoncillos espirales bas- 
tante marcados, aplanados, algo más 


anchos que los interespacios, cuyo fondo 
forma una fila de diminutos puntos exca- 
vados. Al final de la segunda media 
vuelta aparece una escultura de costillas 
axiales algo sigmoideas y opistoclinas, de 
anchura similar a los espacios que las 
separan. En los espacios se pueden apre- 
ciar unos cordoncillos espirales que con- 
tinúan los del núcleo. 

Teleoconcha con ornamentación 
formada por cordones espirales y costi- 


9 


Iberus, 28 (1), 2010 


Figuras 81-84. Chauvetia decorata Monterosato, 1889. 81-83, ejemplar de Punta Almina, Ceuta, 
32-40 m (6,0 mm); 84, ejemplar de Cádiz (7,2 mm). 

Figures 81-84. Chauvetia decorata Monterosato, 1889. 81-83, specimen from Punta Almina 32-40 m 
(6.0 mm); 84, specimen from Cádiz (7.2 mm). 


llas axiales. En la primera vuelta de la 
teleoconcha aparecen cuatro cordones 
espirales, los dos abapicales bien marca- 
dos, los adapicales (subsuturales) menos 
prominentes o casi fusionados; a partir 
de la tercera vuelta, los cordones subsu- 
turales se afirman, de modo que hay 
cuatro cordones casi iguales en la penúl- 
tima vuelta; esta escultura persiste en la 
última vuelta, que presenta en total 15- 
16 cordones que decrecen algo en 
tamaño al acercarse al canal sifonal. En 
la penúltima vuelta se aprecian 11-13 
costillas axiales, que persisten en la 
última vuelta; a su paso por las costillas 
axiales, los cordones forman tubérculos 
redondeados, a modo de perlas. 

La última vuelta alcanza el 65% de la 
altura total, mientras que la abertura 
ocupa en torno al 45%. 

Abertura oval, engrosada en los 
ejemplares adultos, con un canal sifonal 
muy corto y ancho. Labio grueso en los 
ejemplares adultos, particularmente en 
su parte adapical; el lado externo forma 
una variz distinta, aunque no delimi- 
tada del resto de la vuelta, sobre la cual 
los cordones están muy atenuados, y 
son casi inapreciables. En el interior del 
labio de los ejemplares completamente 


54 


adultos se observan 4-5 dentículos poco 
marcados, alargados y situados muy 
adentro de la abertura; sin embargo, el 
interior de la abertura de los ejemplares 
subadultos es frecuentemente liso. El 
borde del labio en vista lateral forma 
una curvatura suave, sin proyección ni 
dentículo. 

Protoconcha castaña clara; teleocon- 
cha castaña en la mitad adapical de las 
vueltas y en la parte abapical de la 
última vuelta, con una zona blanca en la 
mitad abapical de las vueltas de espira, 
continuada como una banda en la mitad 
de la última vuelta. 

Animal desconocido. 

Distribución: Costa atlántica de 
Marruecos y estrecho de Gibraltar. 

Notas: Esta especie es muy parecida 
a C. taentata spec. nov., pero su espira es 
más ancha y los cordones se mantienen 
en toda la superficie y no tienden a apla- 
narse en las últimas vueltas. Los juveni- 
les se distinguen bien, en las poblacio- 
nes simpátricas de Ceuta, por tener la 
protoconcha más pequeña y las vueltas 
no tan altas en proporción, regular- 
mente convexas (no aplanadas con un 
ángulo en la periferia) y con los cordo- 
nes adapicales pequeños, en lugar de 


GOFAS Y OLIVER: Las especies de Chauvetia en el área ibero-marroquí 


Figuras 85-96. Protoconchas en vista lateral y apical, todas a la misma escala. 85-88: Chauvetia 
dentifera, Punta Almina, Ceuta, 32-40 m; 89-90: Chauvetia taeniata, Punta Almina, Ceuta, 32-40 
m; 91-94: Chauvetia decorata, Punta Almina, Ceuta, 32-40 m; 95-96: Chauvetia balgimae, frente a 
Rabat, 355 m. 

Figures 85-96. Protoconchs in lateral and apical views, all to scale. 85-88: Chauvetia dentifera, Punta 
Almina, Ceuta, 32-40 m; 89-90: Chauvetia taeniata, Punta Almina, Ceuta, 32-40 m; 91-94: Chauvetia deco- 
rata, Punta Almina, Ceuta, 32-40 m; 95-96: Chauvetia balgimae, off Rabat, 355 m. 


99 


Iberus, 28 (1), 2010 


Figuras 97-105. Esquemas de los patrones de coloración de los animales vivos. 97: Chauvetia 
mamillata, Sagres, 5 m; 98: C. mamillata, Punta Almina, Ceuta, 32-40 m; 99: C. brunnea, Sagres, 
5 m; 100: C. brunnea, Punta Almina, Ceuta, 32-40 m; 101: C. procerula con color de concha uni- 
forme, Sagres, 5 m; 102: C. procerula con banda blanca, Punta del Saudiño; 103: C. recondita, 
Punta Almina, Ceuta, a 32-40 m; 104: C. retifera juvenil, frente a Salema, Algarve; 105: C. denti- 
fera, Sagres, 20 m. 

Figures 97-105. Sketches of colour patterns in the living animals. 97: Chauvetia mamillata, Sagres, 5 
m; 98: C. mamillata, Punta Almina, 32-40 m; 99: C. brunnea, Punta Almina, 32-40 m; 100: C. 
brunnea, Sagres, 5 m; 101: C. procerula with uniform shell colour, Sagres, 5 m; 102: C. procerula 
with white band, Punta del Saudiño; 103: C. recondita, Punta Almina, 32-40 m; 104: juvenile C. 
retifera, off Salema, Algarve; 105: C. dentifera, Sagres, 20 m. 


cuatro cordones casi iguales desde el 
principio. Entre las cuatro especies que 
presentan el patrón de color con banda 
blanca, ésta nos parece la que mejor se 
ajusta a la breve diagnosis proporcio- 
nada por MONTEROSATO (1889), sobre 


9/0 


todo porque la incluye en el género 
Chauvetia (en su criterio, junto con C. 
granulata sensu Tiberi, 1868 y con C. 
recondita), y no en Donovania (junto con 
C. procerula, descrita en el mismo artí- 
culo). 


GOFAS Y OLIVER: Las especies de Chauvetía en el área ibero-marroquí 


Figuras 106-108. Chauvetia balgimae spec. nov., holotipo de BALGIM DR82, frente a Rabat, 355 


m (6,3 mm). 


Figures 106-108. Chauvetia balgimae spec. nov., holotype from BALGIM DR82, off Rabat, 355 m 


(6.3 mm). 


Chauvetia balgimae spec. nov. (Figs. 95-96, 106-108) 


Material tipo: Holotipo [ej., 6,3 x 2,9 mm], MNHN 22874 y 9 paratipos (4 ej. MNHN 22875, 5 ej. 


MNCN 15.05/53587), todos de la localidad tipo. 


Material estudiado: El material tipo y Balgim Sta. DR81 (33? 46' N, 08? 30” W), 309 m, 1 ej. 
Localidad tipo: Frente a Rabat, Marruecos (33? 45' N — 08* 32” W, 355 m), Balgim St. DR82. 
Etimología: Del nombre de la campaña Balgim (Bentos del AtLántico, Glbraltar y Mediterráneo), 


en la cual la especie fue recolectada. 


Descripción: Concha fusiforme, no 
muy sólida, con 4! /2-5 vueltas de espira 
y hasta 6,8 mm, y vueltas muy conve- 
xas, con una sutura bastante marcada. 

Protoconcha con aproximadamente 
una vuelta, diámetro máximo de 850- 
900 um (núcleo: 500 um, primera media 
vuelta: 700 um). El núcleo es redonde- 
ado y tiene cordoncillos espirales muy 
tenues, casi inapreciables con microsco- 
pía óptica, desiguales entre sí, con inte- 
respacios relativamente lisos; desde la 
primera media vuelta aparece una 
escultura de numerosas costillas axiales 
algo sigmoideas y opistoclinas, de 
anchura similar a los espacios que las 


separan. En los espacios se puede apre- 
ciar la continuación de los cordoncillos 
espirales, que están atenuados en la 
parte más saliente de las costillas. 
Teleoconcha con ornamentación 
formada por cordones espirales y costi- 
llas axiales. En la primera vuelta de la 
teleoconcha aparecen cuatro cordones 
espirales, de los que uno (subsutural) 
está algo apartado de la sutura; en este 
intervalo surge un quinto cordón a partir 
de la segunda vuelta. Esta escultura per- 
siste en la última vuelta, que presenta en 
total unos 20 cordones que decrecen mar- 
cadamente en grosor al acercarse el canal 
sifonal. Las costillas axiales son algo sig- 


34 


Iberus, 28 (1), 2010 


moideas, ligeramente más estrechas que 
los interespacios; en la penúltima vuelta 
se aprecian 17-19 costillas axiales, que se 
atenúan algo al final de la última vuelta. 
A su paso por las costillas axiales, los cor- 
dones forman pequeños tubérculos 
redondeados. 

La última vuelta alcanza el 62% de la 
altura total, mientras que la abertura 
ocupa en torno al 42% 

Abertura ovalada, con un canal 
sifonal largo para el género, y ancho. 
Labio en los ejemplares adultos con el 
lado externo formando una variz estre- 
cha y elevada, situada algo separada del 
borde, que es fino y cortante; los cordo- 
nes están muy atenuados, y son casi ina- 


CONCLUSIONES 


En este trabajo consideramos doce 
especies válidas del género Chauvetia en 
el litoral ibero-marroquí, situando en el 
estrecho de Gibraltar y en el sur de la 
Península Ibérica la máxima diversidad 
específica del género en Europa. En 
algunos puntos del Estrecho, como 
Punta Almina o Benzú, cerca de Ceuta, 
se han podido recolectar hasta ocho 
especies en la misma muestra. En otras 
regiones del Mediterráneo, contando 
incluso con especies que no se encuen- 
tran en el área aquí estudiada, pueden 
encontrarse hasta siete especies en el 
entorno de Sicilia, que es el otro “punto 
caliente” de la riqueza especifica en el 
Mediterráneo. El máximo absoluto de 
riqueza específica, sin embargo, sigue 
centrado en la península del cabo Verde, 
en Senegal, donde Oliver y Rolán (2008) 
han encontrado 14 especies, seguido de 
cerca por la plataforma sahariana, con 
una docena de especies (Oliver y Rolán, 
2009). Luego, más al sur, el género es 
muy escaso en África occidental, exis- 
tiendo alguna representación hasta 
Costa de Marfil, pero con un número de 
especies muy bajo. 

El género Chauvetia no está represen- 
tado en el Atlántico occidental, lo que es 
explicable considerando el tipo de desa- 
rrollo sin fase planctónica que se deduce 
de la protoconcha de todas las especies 


58 


preciables sobre la variz. En el interior 
del labio de los ejemplares completa- 
mente adultos no se aprecian dentículos. 
El borde del labio en vista lateral es sig- 
moideo. 

Color de la concha y del animal 
completamente blanco. 

Notas: La protoconcha de esta 
especie se parece mucho a la de Chauve- 
tia edentula Oliver y Rolán, 2009, descrita 
de una profundidad similar en el banco 
sahariano y en la isla de Gran Canaria. 
Sin embargo, la teleoconcha es marcada- 
mente diferente, siendo C. edentula 
mucho más sólida, con la última vuelta 
proporcionalmente más compacta y la 
abertura más pequeña. 


estudiadas del género. Tampoco está 
representado en las islas Azores, 
poniendo así de manifiesto que las dis- 
tancias transoceánicas son infranqueables 
por sus representantes. No siempre las 
especies con desarrollo directo intracap- 
sular son incapaces de colonizar sitios 
remotos, mediante, por ejemplo, la flota- 
ción de cápsulas ovígeras, pero parece 
evidente que, en el caso de Chauvetia, la 
capacidad de dispersión es escasa. 

Esta capacidad limitada de disper- 
sión hace probablemente que las espe- 
cies de Chauvetia sean propensas a dife- 
renciarse en poblaciones locales con 
escaso intercambio genético con otras 
poblaciones de la especie. Quizás ésta 
sea una razón para que existan tantos 
problemas taxonómicos sin resolver. En 
este trabajo hemos puesto de manifiesto 
algunos de ellos y es obvio que aún 
faltan datos a lo largo de las costas nor- 
teafricanas y en la cuenca oriental para 
que se pueda llegar a una revisión taxo- 
nómica completa del género. También es 
de esperar que los caracteres de secuen- 
cias moleculares aporten nuevos ele- 
mentos decisivos cuando se conozcan 
para estas especies. 

Las especies litorales incluidas en 
este estudio están todas claramente vin- 
culadas a fondos rocosos o, a lo sumo, a 
fondos colindantes de cascajo biógeno. 


GOFAS Y OLIVER: Las especies de Chauvetia en el área ibero-marroquí 


Quizá por ello no se ha encontrado 
ningún ejemplar de Chauvetia en el 
amplio muestreo realizado en las 
lagunas del Algarve, en los alrededores 
de Faro y Olháo, a pesar de la presencia 
de extensas praderas de fanerógamas 
ricas en especies y de un esfuerzo de 
muestreo equivalente al desplegado en 
la zona de Sagres. Por la misma razón, 
los tramos costeros con litoral arenoso y 
lagunas, como los del suroeste ibérico o 
del noroeste marroquí, podrían actuar 
como barreras que convierten en islas 
los tramos de costa rocosa intercalados y 
propician la diferenciación de las pobla- 
ciones. 


AGRADECIMIENTOS 


La campaña Ceuta 1986 fue el fruto 
de un esfuerzo conjunto del MNHN y 
de la Universidad de Sevilla (José Carlos 


BIBLIOGRAFÍA 


ARNAUD P. 1978. Révision des taxa malacolo- 
gliques méditerranéens introduits par An- 
toine Risso. Annales du Muséum d'Histoire 
Naturelle de Nice, 5: 101-150. 

BUCQUOY E., DAUTZENBERG P. Y DOLLFUS G. 
1882-1886. Les mollusques marins du Roussil- 
lon. Tome ler. Gastropodes. Paris, J. B. Bail- 
liere et fils. 570 pp., 66 lám. [pp. 85-135, pl. 
11-15, febrero 1883] 

CROSSE H. 1885. Nomenclatura generica e 
specifica di alcune conchiglie mediterra- 
nee, pel Marchese di Monterosato [reseña 
bibliográfica]. Journal de Conchyliologie, 33: 
139-142. 

HERGUETA H., LUQUE A. Y TEMPLADO J. 2002. 
On the taxonomy and biology of Chauvetia 
mamillata (Risso, 1826) (Gastropoda: Bucci- 
nidae) in south East Spain. Bollettino Mala- 
cologico, supplemento 4: 135-146. 

HORRO J. Y ROLÁN E. 2004. Las especies de 
Chauvetia de Galicia (Mollusca, Muricoidea). 
Noticiario SEM, 41: 27-30. 

LOCARD A. 1892. Les coquilles marines des cótes 
de France. J. B. Bailliere et fils, Paris, 384 pp. 

MAgs V.O. 1983. Observations on the syste- 
matics and biology of a turrid gastropod as- 
semblage in the British Virgin Islands. Bul- 
letin of Marine Science, 33 (2): 305-335. 

MICALI P. 1999. Note sulle specie di Chauvetia 
dell' Atlantico nord-orientale. Bollettino Ma- 
lacologico, 34 (5-8): 53-68. 


García Gomez), la del Algarve (1988) y 
Seamount 1 (1987) fueron dirigidas por 
Philippe Bouchet (MNHN), y las cam- 
pañas Fauna Ibérica III (1994) y IV 
(1996), realizadas en el Proyecto Fauna 
Ibérica (DGICYT PB92-0121) fueron diri- 
gidas por Ángel Guerra, del Instituto de 
Investigaciones Marinas (Consejo Supe- 
rior de Investigaciones Científicas). 

Las fotografías de microscopio elec- 
trónico de barrido fueron realizadas en 
su mayoría en los servicios centrales de 
apoyo a la investigación de la Universi- 
dad de Málaga, con la ayuda de Grego- 
rio Martín Caballero; algunas de ellas se 
realizaron en el laboratorio de microsco- 
pía electrónica del Museo Nacional de 
Ciencias Naturales por Laura Tormo y 
Marta Furió. 

Los autores también agradecen a 
Ángel Luque (Universidad Autónoma, 
Madrid) una minuciosa revisión del 
manuscrito y sugerencias valiosas. 


MONTEROSATO T. A. D1 1872. Notizie intorno alle 
conchiglie mediterranee. Palermo, Michele 
Amenta, 61 pp. 

MONTEROSATO T. A. DI 1884. Nomenclatura ge- 
nerica e specifica di alcune conchiglie mediterra- 
nee. Palermo, Virzi, 152 pp. 

MONTEROSATO T. DI 1889. Coquilles marines 
Marocaines. Journal de Conchyliologie, 37: 20- 
40, 112-121. 

NORDsIECK F. 1976. Familia Buccinidae. Il ge- 
nere Chauvetia Monterosato, 1884 nei mari 
d'Europa. La Conchiglia, 89-90: 3-7. 

OLIVER J. D. Y ROLÁN E. 2008. Las especies del 
género Chauvetia (Gastropoda, Neogastro- 
poda) del área de Dakar, Senegal, Africa oc- 
cidental, con la descripción de diez especies 
nuevas. Iberus, 26 (2): 133-175. 

OLIVER J. D. Y ROLÁN E. 2009. Las especies de 
Chauvetia Monterosato, 1884 (Mollusca, Ne- 
ogastropoda) de Canarias y el área Oeste 
Africana de Mauritania y Sahara. Iberus, 27 
(2): 113-154. 

PALLARY P. 1902. Liste des mollusques testacés 
de la baie de Tanger. Journal de Conchyliolo- 
core pUriS9, plsl 

PALLARY P. 1920. Exploration scientifique du Ma- 
roc organisée par la Société de Géographie de Pa- 
ris et continuée par la Société des Sciences Na- 
turelles du Maroc. Deuxieme fascicule. Mala- 
cologie (1912). Larose, Rabat y Paris, 108 pp., 
1 lám., 1 mapa. 


SY 


Iberus, 28 (1), 2010 


PEÑAS A., ROLÁN E., LUQUE A.A., TEMPLADO J., 
MORENO D., RUBIO F., SALAS C., SIERRA A. Y 
GOFAS S. 2006. Moluscos marinos de la isla 
de Alborán. Iberus, 24 (1): 23-151 

Risso A. 1826. Histoire naturelle des principales pro- 
ductions de l'Europe méridionale et particuliere- 
ment de celles des environs de Nice et des Alpes 
Maritimes, vol. 4. Paris, Levrault, iv + 439 
pp., 12 láms. 

SCACCHI A. 1836. Catalogus conchyliorum regni 
Neapolitani. Neapoli [Napoles], Typis Filiatre- 
Sebetii, 18 pp., 1 lám. 


60 


TIBERI N. 1868. Des testacés de la Méditerranée 
qui doivent étre compris dans les genres La- 
chesis et Neseea de Risso. Journal de Conchy- 
liologie, 16: 68-81, lám. 5. 

VERMEEJ G. 2001. Innovation and evolution at 
the edge: origins and fates of gastropods 
with a labral tooth. Biological Journal of the Lin- 
nean Society, 72: 461-508. 

WOODWARD S.P. 1899. Some account of the sy- 
nonymy and affinities of Donovania minima 
(Mont.). Proceedings of the Malacological Society 
of London, 3 (3): 235-238. 


O Sociedad Española de Malacología Iberus, 28 (1); 61-66, 2010 


Record of the largest specimen of neon flying squid 
Ommastrephes bartramii (Cephalopoda: Ommastrephidae) 


Registro del mayor ejemplar de pota saltadora Ommastrephes bar- 
tramii (Cephalopoda: Ommastrephidae) 


Ángel GUERRA*, Graham J. PIERCE**, María Begoña SANTOS***, Ángel 
E GONZÁLEZ*, Gema HERNÁNDEZ-MILIAN***, Carmela POR- 
TEIRO*** and Baltasar PATIÑO*** 


Recibido el 26-XI1-2009. Aceptado el 23-IV-2010 


ABSTRACT 


We report a record of the largest known specimen of Ommastrephes bartramii. lt was an 
almost mature female, with dorsal mantle length (ML) of 1020 mm and weighing around 
35 kg, caught by a Spanish fishing vessel in October 2007 on a surface long-line in the 
Eastern Tropical Pacific. Growth increments on a statolith indicate an age of 492-512 
days. The stomach was full and contained beaks of ommastrephid squids and of a small 
pelagic octopus [Japetella spp.), and remains of mackerel [possibly bait from the long-line) 
and unidentified fish. 


RESUMEN 


En este trabajo se presenta información sobre el mayor ejemplar de Ommastrephes bar- 
tramii descrito hasta la actualidad. Se trata de una hembra casi madura, cuya longitud 
dorsal del manto es de 1020 mm y con un peso total de aproximadamente 35 kg, captu- 
rada en octubre de 2007 por un pesquero español utilizando palangre de superficie en el 
Pacífico Tropical Este. Los incrementos de crecimiento observados en un estatolito indican 
una edad comprendida entre 492 y 512 días. El estómago estaba lleno y contenía picos 
de potas [omastréfidos) y de un pulpo pelágico pequeño [Japatella spp.), así como restos 
de caballas (posiblemente cebo del palangre) y de otros peces que no pudieron ser identi- 
ficados. 


INTRODUCTION 


The neon flying squid Ommastrephes 
bartramii (LeSueur, 1821) is distributed 
worldwide in subtropical and temperate 
oceanic waters (ROPER, SWEENEY AND 
NAUEN, 1984; NeEsis, 1987; DUNNING, 
1998). It has supported major jig and 


surface driftnet fisheries in the North 
Pacific since about 1974, and its life 
history in this area is well known 
(BOWER AND ICHIL, 2005; IcHií, MAHAPA- 
TRA, OKAMURA, AND OKADA 2006). It 
occurs in the South Pacific where SST 


* Instituto de Investigaciones Marinas (CSIC), Vigo, Spain. E-mail: angelguerraCiim.csic.es 
** Centro Oceanográfico de Vigo, Instituto Español de Oceanografía, P.O. Box 1552, 36200 Vigo, Spain and 
Institute of Biological and Environmental Sciences, Zoology Building, University of Aberdeen, Tillydrone 


Avenue, AB24 2TX Aberdeen, U.K. 


*** Centro Oceanográfico de Vigo, Instituto Español de Oceanografía, P.O. Box 1552, 36200 Vigo, Spain. 


61 


Iberus, 28 (1), 2010 


Figure 1. Worldwide distribution of Ommastrephes bartramii (AquaMaps, GBIF OBIS) and 


capture location (O) of this record. 


Figura 1. Distribución mundial de Ommastrephes bartramii (Aqua Maps, GBIF OBIS) y localiza- 


ción del lugar de captura del ejemplar (O). 


ranges from - 12” to 26” C and is rarely 
caught in cooler waters. Adjacent to the 
continental slope of western South 
America, O. bartrami is replaced in 
waters warmer than 15” C by the jumbo 
squid Dosidicus gigas (YATSU AND 
YAMASHIRO, 1999; ZUEV, NIGMATULLIN, 
CHESALIN, AND NESIS, 2002). The Sub- 
tropical Convergence (SC) is the south- 
ern boundary of its distribution in the 
South Pacific, generally at about 40-502 
S. However, the SC zone is highly vari- 
able from year to year and shows signif- 
icant seasonal latitudinal shifts, occur- 
ring as far south as 52” S (DUNNING, 
1998). 

Although this species was identified 
as a potential fishery resource in the late 
1970s, there is still no commercial 
exploitation in the Southern hemisphere 
(DUNNING, 1998). NIGMATULLIN, 
SHCHETINNIKOV AND  SHUKHGALTER 
(2009) sampled 60 specimens of this 
species, to study the diet and parasites, 
by jigging in the south-eastern Pacific in 
the early 19805. The animals ranged 
from 16 to 39 cm mantle length (ML). 


62 


The largest specimen recorded to date 
was a female of 800 mm mantle length 
weighing 20-25 kg caught in Argen- 
tinean waters (DUNNING, 1998). In this 
paper we describe a new record of a 
larger specimen caught in the SE Pacific, 
including information on age, maturity 
and stomach contents. 


MATERIAL AND METHODS 


The present specimen was caught by 
the Spanish fishing vessel Nuevo Monte 
Ventoso, 10/10/07, on a surface long-line 
at 21” S, 88” W (FAO area 87, Figure 1). It 
was frozen on board and transported to 
Vigo (NW Spain). It was defrosted at 
room temperature, dissected and mea- 
sured (see Figure 2) We sampled 
stomach contents, one statolith (the 
other was not located), beaks, sucker 
rings, mantle and ovary tissue. 

The method applied for ageing 
involved mounting the statolith on a 
microscope slide, using Crystalbond, 
with the anterior concave side upper- 


GUERRA £7 AL.: Record of the largest specimen of Ommastrephes bartramii 


PS 


Figure 2. Ommastrephes bartramii, view of the mouth, arms and anterior ventral mantle margin of 


the specimen. 


Figura 2. Ommastrephes bartramii, vista de la boca, brazos y margen ventral anterior del manto del 


ejemplar. 


most. The statolith was ground, first on 
the anterior surface, then turned over 
and ground on the posterior surface. 
The statolith was then turned over so 
that the anterior surface was uppermost. 
This grinding of both surfaces in the 
sagittal plane results in the production 
of a relatively thin statolith section. 
Increments were determined along the 
axis of maximum statolith growth with 
a NIS Elements D 2.30 image analysis 
system interfaced with a Nikon com- 
pound microscope (400x magnification). 
Counts were obtained semi-automati- 
cally: putative increments were detected 
automatically by computer software 
from an enhanced image but final iden- 
tification of increments was carried out 
manually. Increments were not clearly 
identifiable near the outer margin of the 
ground surface, and the number of 
increments missed in this area was esti- 


mated by extrapolation from the adja- 
cent area (GONZÁLEZ, DAWE, BECK AND 
PÉREZ, 2000). 

Stomach contents (consisting of 
semi-digested flesh and hard remains) 
were washed through a sieve (mesh size 
0.335 mm) and all identifiable prey 
remains (e.g. fish otoliths, bones and 
cephalopod beaks) extracted and trans- 
ferred to 75% alcohol. Fish hard parts 
were later dried. Beaks, otoliths and 
bones were identified to the lowest pos- 
sible taxon using guides (e.g. CLARKE 
1986, HARKÓNEN 1986; BoscH1, Fis- 
CHBACH AND IORIO, 1992; SMALE, 
WATSON AND HECHT, 1995; WArT, PIERCE 
AND BOYLE, 1997; TUSET, LOMBARTE, AND 
Assis, 2008) and reference material held 
by the authors. Original prey size was 
calculated from standard measurements 
(lower rostral length for squid beaks) 
using published regressions (CLARKE, 


63 


Iberus, 28 (1), 2010 


Table I. Main measurements (mm for linear measurements, g for weights). 
Tabla I. Principales medidas del ejemplar (en mm las morfométricas y en g los pesos). 


Linear dimensions mm 
Dorsal montle lengih 1020 
Ventral mantle length 970 
Ventral mantle width 315 
Fin length 500 
Fin width OS 
Arm R] length 580 
Arm R3 length 630 
Left tentacle 1020 


1986). In the case of the fish remains, the 
otoliths found were fragmented. Of the 
intact fish bones identifiable to family 
we were able to use the dentary length 
to estimate the approximate size, using 
an unpublished regression. 


RESULTS AND DISCUSSION 


The specimen was a female, with 
dorsal mantle length (ML) of 1020 mm 
and weighing around 35 kg. It was 
almost mature, with mature oocytes 
passing through the oocyte chamber, 
and was mated (see Table 1). Repeat 
readings of growth increments on the 
one statolith located indicate an age of 
492-512 days. 

Age data from the North Pacific 
suggest a 1-year life cycle whereas this 
specimen was already 16-17 months old. 
However, similar discrepancies between 
reported age and the known seasonality 
of the life cycle are known from other 
squids, e.g. Loligo spp. (see GONZÁLEZ, 
OTERO, GUERRA AND PIERCE, In Press). 
BOWER AND IcHIH1 (2005) reported two 
seasonal cohorts in the North Pacific. In 
the loliginid squid Loligo forbesi, BOYLE, 
PIERCE AND HASTIE (1995) proposed that 
individual growth rate determines 
whether an animal becomes a winter or 
summer spawner, and individuals from 
summer and winter breeding seasons 
might thus be of mixed age. Thus an 
apparently annual spawning and 
recruitment cycle is not necessarily 


64 


Weights J 
Total weight Approx. 35000 
Nidamental gland weight 250 (1 out of 2) 
Oviduct gland weight 150 (1 out of 2) 
Ovary weight 500 
Digestive gland weight 950 
Stomach contents weight 950 


inconsistent with some animals living 
for up to 2 years. CHEN AND CHIU (2003) 
recorded maximum mantle lengths of 
527 mm for Ommastrephes bartramil in 
the North Pacific between September 
and December, which would be consis- 
tent with the present specimen 
approaching spawning readiness in 
October. 

The stomach was full (containing 
9508 of food remains, a mixture of semi- 
digested flesh and hard prey remains). 
Table II summarises the information on 
prey found in the stomachs, including 
reconstructed lengths and weights. 
Remains included beaks of ommas- 
trephid squids (as well as fragments of 
gladius and cephalopod flesh), verte- 
brae, dentaries and a broken otolith 
from a fish of the family Scombridae, 
probably a species of mackerel and 
remains of unidentified fish (broken 
otoliths, bones, a post-temporal, and 
numerous dermal scutes). An intact 
specimen of alfonsino (Beryx sp., proba- 
bly B. decadactylus) was recovered from 
the mantle cavity of the squid (it had 
not been eaten). 

It was not possible to identify the 
ommastrephid beaks to species since 
beaks from species in this family are 
very similar but it is likely that they 
belong to the same species, since canni- 
balism is known to be quite common in 
cephalopods (IBÁÑEZ AND KEYL, 2010). 
The other identified remains of 
cephalopods belonged to the genus 
Japetella. With the exception of Japetella, 


GUERRA ET AL.: Record of the largest specimen of Ommastrephes bartramii 


Table II. Prey species found in the stomachs of the specimen of Ommastrephes bartramií taken by a 
long-liner in the south-eastern Pacific. For all prey species, number of beaks/other remains, estima- 
ted number of individuals (N) and estimated prey length (mm) and weight (g) are indicated. 

Tabla 11. Especies de presas encontradas en el estómago de Ommastrephes bartramii capturado con un 
palangre en el Pacífico sureste. Para todas las especies se indica el número de picoslotros restos, el número 


de individuos estimados (N) y una estimación de la longitud de la presa (mm) y su peso (2). 


Cephalopod prey Importance 

Family Species Remains N Length (mm) Weight (g) 
Ommastrephidae Unidentified 3 lower + 2 upper beaks, gladius, flesh 3 259-301 536-728 
Bolitaenidae Japatella sp. 1 lower + 1 upper beaks | d á 

Fish prey 

Family Species 

Scombridae Scomber spp. Bones +1 otolith / 359-385 393-493 
Unidentified Unidentified broken otoliths, bones, dermal scutes 


which is a small pelagic octopus (speci- 
mens of Japetella recorded from pygmy 
sperm whale stomachs in Hawaii aver- 
aged 8 g in body weight, West et al. 
2009), the estimated size of the prey 
taken by the squid ranged from 259 to 
385 mm in length and 390 to 720 g. The 
occurrence of mackerel in the diet is at 
first sight surprising since mackerel is a 
shelf species, but it may have been used 
as bait on the long-line (although 
remains of more than 1 individual 
mackerel were found in the stomach 
and bait fish are normally spaced out 
along a long-line). 

There is little information on the diet 
of the species, NIGMATULLIN, SHCHE- 
TINNIKOV AND SHUKHGALTER (2009) 
reported on the stomach contents of 60 
specimens of O. bartramil taken by 
hand-jigging in the southeast Pacific. 
These squid had taken a wide variety of 
prey, mainly myctophid fish (Symbo- 
lophorus, Myctophum and Hygophun) 
and cephalopods (Onychoteuthidae and 
Enoploteuthidae). The authors also 
noted the presence in the stomachs of 
beaks of the family Ommastrephidae 
(including some remains of O. bar- 
tramii). Myctophid fish and squid of the 
families Onychoteuthidae and Enoplo- 


teuthidae were also found to be the 
main prey in a sample of 315 O. bar- 
tramii from off Hawaii (PARRY, 2006). No 
myctophid remains were found in the 
stomach of the present specimen. 

The main preys of this species in the 
Northwest Pacific were myctophids. 
Secondary important prey items 
included onychoteuthid and gonatid 
squids. The study was done using drift- 
nets offshore but Engraulis japonicus and 
Carangidae (as well as other species that 
could be found in shelf waters) were 
reported in the diet in low numbers. 
Chnages in feeding habits of the neon 
flying squid were found in relation to 
their seasonal south-north migrations 
and diel vertical displacements (WATAN- 
ABE, KUBODERA, ICHIl AND KAWAHARA, 
2004). 


ACKNOWLEDGEMENTS 


We would like to thank the skipper 
and crew of the Nuevo Monte Ventoso 
and Rafael Bañón Díaz who identified 
the fish found inside the squid mantle. 
G.J. Pierce was supported by the 
ANIMATE project (MEXC-CT-2006- 
042337). 


Ó5 


Iberus, 28 (1), 2010 


BIBLIOGRAPHY 


BoscH1 E., FISCHBACH C. AND JORIO M. 1992. 
Catálogo ilustrado de los crustáceos estom- 
atópodos y decápodos marinos de Argentina. 
Frente Marítimo, 10: 56-57. 

BOWER J.R. AND IcHn T. 2005. The red flying 
squid (Ommastrephes bartramii): A review of 
recent research and the fishery in Japan. Fishe- 
ries Research 76, 39-55. 

BOYLE P.R., PIERCE G.J. AND HASTIE L.C. 1995. 
Flexible reproductive strategies in the 
squid Loligo forbesi. Marine Biology, 121: 
501-508. 

CHEN C.-S. AND CHIU T.S. 2003. Variations of 
life history parameters in two geographical 
groups of the neon flying squid, Ommas- 
trephes bartramii, from the North Pacific. Fish- 
eries Research, 63: 349-366. 

CLARKE M.R. 1986. A handbook for the identifica- 
tion of Cephalopod beaks. Clarendon Press. Ox- 
ford. 273 p. 

DUNNING M. 1998. An overview of the fish- 
eries biology and resource potential of Om- 
mastrephes bartramii (Cephalopoda: Ommas- 
trephidae) in the Southern Hemisphere. In 
Okutani, Y. (Ed.): Contributed papers to In- 
ternational Symposium on Large Pelagic 
Squids. Japan Marine Fishery Resources Re- 
search Center, Tokyo: 65-76. 

GONZÁLEZ A.F., DAWE E.G., BECK P.C. AND 
PÉREZ J.A.A. 2000. Bias associated with sta- 
tolith-based methodologies for ageing squid: 
a comparative study on Illex ¡llecebrosus 
(Cephalopoda: Ommastrephidae). Journal of 
Experimental Marine Biology and Ecology, 244: 
161-180. 

GONZÁLEZ A.F, OTERO J., GUERRA A., PIERCE G. 
J. (in press) Age, growth and mortality of 
Loligo vulgaris wild planktonic paralarvae in 
the Ría de Vigo (NE Atlantic Ocean). ICES 
Journal of Marine Science. 

HARKÓNEN T. 1986. Guide to the otoliths of bony 
fishes of the Northeast Atlantic. Daubin ApsS. 
Sweden. 256 p. 

IBÁÑEZ CH.M. AND KEYL F. 2010. Cannibalism 
in cephalopods. Reviews in Fish Biology and 
Fisheries, 20: 123-136. 

IcHhu T., MAHAPATRA K., OKAMURA H. AND 
OKADA Y. 2006. Stock assessment of the au- 
tumn cohort of neon flying squid (Ommas- 
trephes bartramii) in the North Pacific based 
on past large-scale high seas drifnet fishery 
data. Fisheries Research, 78: 286-297. 

Nesis K.N. 1987. Cephalopods of the world. 
Squids, cuttlefishes, octopuse, and allies. 
T.F.H. Publications, Neptune City. 351 p. 


66 


NIGMATULLIN C.M., SHCHETINNIKOV A.S. AND 
SHUKHGALTER O.A. 2009. On feeding and 
helminth fauna of neon flying squid Om- 
mastrephes bartramil (Lesueur, 1821) 
(Cephalopoda: Ommastrephidae) in the 
southeastern Pacific. Revista de Biología Ma- 
rina y Oceanografía, 44: 227-235. 

PARRY M. 2006. Feeding behaviour of two om- 
mastrephid squids Ommastrephes bartramii 
and Sthenoteuthis oualaniensis off Hawaii. Ma- 
rine Ecology Progress Series, 318: 229-235. 

ROPER C.F.E., SWEENEY M.J. AND NAUEN C.E. 
1984. FAO species catalogue. Cephalopods of the 
world. An annotated and illustrated catalogue of 
species of interest to fisheries. FAO Fisheries 
Synopsis, 3 (125): 277 p. 

SMALE M.J., WATSON G. AND HECHT T. 1995. 
Otolith atlas of Southern African marine fishes 
(Ichthyological monographs). J.L.B. Smith In- 
stitute of Ichthyology, 1, 253 pp. 

TuseT V.M., LOMBARTE A. AND ASsIS C.A. 2008. 
Otolith atlas for the western Mediterranean, 
north and central eastern Atlantic. Scientia 
Marina, 72: 1-203. 

WATANABE H., KUBODERA T., IcHt T. AND 
KAWAHARA $. 2004. Feeding habits of neon 
flying squid Ommastrephes bartramii in the 
transitional region of the central North Pa- 
cific. Marine Ecology Progress Series, 266: 173- 
184. 

WATT J., PIERCE G.J. AND BOYLE P.R. 1997. A 
guide to the identification of North Sea fish 
using premaxillae and vertebrae. Co-opera- 
tive Research Report No 220, International 
Council for the Exploration of the Sea, 231 p. 

WesT K.L., WALKER W.A., BAIRD R.W., WHITE 
W., LEVINE G., BROWN E. AND SCHOFIELD D. 
2009. Diet of pygmy sperm whales (Kogia 
breviceps) in the Hawaiian Archipelago. Ma- 
rine Mammal Science, DOI: 10.1111 /3.1748- 
7692.2009.00295.x 

Y ATSU A. AND YAMASHIRO C. 1999. Report of the 
Kaiyo Maru Cruise for study on the resources of 
tivo ommastrephid squids, Dosidicus gigas and 
Onmmastrephes bartrami, in the Pacific Ocean, 
during September 11 - December 24, 1997. Fish- 
eries Agency of Japan, 206 p. 

ZUEV G.V., NIGMATULLIN CH.M., CHESALIN 
M.V. AND NEsIs K.N. 2002. Main results of 
long-term worldwide studies on tropical nek- 
tonic oceanic squid genus Sthenoteuthis: an 
overview of the soviet investigations. Bulle- 
tin of Marine Science, 71: 1019-1060. 


O Sociedad Española de Malacología 


Iberus, 28 (1): 67-72, 2010 


A new species of Candidula (Gastropoda, Hygromiidae) 


from central Portugal 


Una nueva especie de Candidula (Gastropoda, Hygromiidae) de 


Portugal 


Geraldine A. HOLYOAK and David T. HOLYOAK* 


Recibido el 19-11-2010. Aceptado el 23-IV-2010 


ABSTRACT 


A new species Candidula coudensis is described from Vale da Couda (Almoster, Leiria), in 
central Portugal. lt differs from other species of the genus in combining a large shell with sharp 
peripheral keel and coarse radial ribs with a penis bearing a long flagellum. The new species 
lives together with Candidula cf. belemensis (Servain, 1880) on rocky limestone slopes. 


RESUMEN 


Se describe la nueva especie Candidula coudensis del Vale da Couda (Almoster, Leiria), 
en la región central de Portugal. Difiere de otras especies del género por presentar una 
concha grande con fuerte quilla carenal y gruesas costillas radiales, y un pene provisto de 
un largo flagelo. La nueva especie vive junto con Candidula cf. belemensis [Servain, 


1880) en pendientes calcáreas rocosas. 


INTRODUCTION 


During fieldwork in the limestone 
hills above Vale da Couda (district of 
Leiria, formerly Beira Litoral), central 
Portugal on 31st January 2008 an unfami- 
liar “helicellid” (Hygromiidae) was 
found. Because of its light-coloured, 
sharply keeled shell with strong radial 
ribs it was initially thought to be Can- 
didula setubalensis (L. Pfeiffer, 1850), 
known only as an endemic of the Serra 
da Arrábida near Setúbal (district of 
Setúbal, Portugal). Subsequent studies 


TAXONOMIC PART 


have revealed that the genital anatomy of 
the snail from Vale da Couda is typical of 
Candidula in having only a single large 
dart sac as in C. setubalensis (GITTEN- 
BERGER, 1985) but it differs from that 
species in the much longer flagellum on 
its penis. Since the shells of C. setubalensis 
also differ in several characters from 
those of the snail from Vale da Couda the 
latter is described here as a new species. 
A fuller review of the species of Candidula 
in Portugal is in preparation. 


Family HYGROMIIDAE Tryon, 1866 
Subfamily HYGROMIINAE Tryon, 1866 
Tribe Helicellini Ihering, 1909 


* Quinta da Cachopa, Barcoila, 6100-014 Cabegudo, Portugal. 


67 


Iberus, 28 (1), 2010 


Genus Candidula Kobelt, 1871 


Type species (by absolute tautonymy): Helix candidula Studer, 1818 = Candidula unifasciata (Poiret, 
1801). 


PROSCHWITZ AND  RIPKEN (2001), 
BANK, GROH AND RIPKEN (2002) and 
FALKNER, BANK AND  PROSCHWITZ 
(2001). 


Remarks: Allocation of the genus 
Candidula to the Tribe Helicellini in the 
Hygromiidae follows BANK, BOUCHET, 
FALKNER, GITTENBERGER, HAUSDORFE, 


Candidula coudensis spec. nov. (Figs. 1-7) 


Type locality: Vale da Couda, by N348 road SE. of Almoster (district of Leiria, Portugal), 298 
NE498099, ca 390 m alt., herb-rich grassland and low shrubs by road. 

Type material: Holotype (Figs 1-5; in the BM, reg. no. 20100177) from type-locality collected 12 
Dec. 2009, leg. G.A. Holyoak; body in spirit and dry shell kept separately (holotype and several 
paratypes kept alive until 16 Jan. 2010). 

Paratypes: 4 dry shells (BM, reg. no. 20100178) and 4 in spirit (3 shell and body, 1 body only, BM, 
reg. no. 20100179), 57 dry shells, 2 shells containing bodies kept in spirit and 5 bodies in spirit with 
dry shells kept separately (Collection ofG.A. Holyoak). Additional paratypes from near type-local- 
ity, all in Collection of G.A. Holyoak: 10 shells, 1 body in spirit (used for Figs. 6, 7) and dry shell 
kept separately, collected 31 Jan. 2008 (limestone crags and slopes, 295 NE498099); 31 shells, 1 body 
in spirit and dry shell kept separately, collected 10 June 2009 (low limestone crags and slopes, 295 
NE498099); 44 shells, collected 12 Dec. 2009 (scrub-covered limestone hillside with low walls and 
scree, 295 NE497096). 

Etymology: The specific epiphet coudensis is an adjective derived from the name of the type-local- 
ity at Vale da Couda. The generic name Candidula has been created as a feminine noun, so the ter- 


mination of the epiphet coudensis is therefore feminine in agreement. 


Description: Adult shell (Figs. 1-3) 
dextral, strongly compressed above, 
with low convex to low-conical spire of 
5/4-5% flattened whorls with shallow 
sutures and sharp slightly raised keel at 
periphery. Umbilicus */5 - */7 width of 
shell, symmetrical, exposing parts of 
several whorls of spire internally, open 
or slightly overlapped by peristome. 
Mouth slightly oblique, oval but some- 
what flattened above and below with 
prominent keel at periphery, thin peris- 
tome and white internal rib. Shell 
opaque, pale cream to light brown 
(fading to whitish) with very variable 
broad to narrow bands of dull brown to 
blackish-brown that are variously dis- 
tinct, fused, blotched or interrupted, 
sometimes almost lacking on underside 
of shell. Shell surface not or slightly 
glossy, with radial ribs which are pale 
on top; on lower whorls ribs become 
strong, evenly spaced, with each rib 
conspicuously raised and thickened at 
the peripheral keel; on underside of 


68 


shell the ribs reach the umbilicus, within 
which they are reduced to rows of papil- 
lae; microsculpture of fine spiral parallel 
grooves is often present on body whorl, 
especially on underside. The protoconch 
appears smooth at x30 magnification; 
there is no trace of hairs or hair-pits on 
the upper whorls. 

Exposed parts of body of living 
animal (Figs. 4, 5) light to rather dark 
grey, with some brown suffusion and 
blackish foot fringe. In detail, the skin 
tubercles are variably suffused with 
brown and outlined in dark grey. 

Genital anatomy (Figs. 6, 7); descrip- 
tion based on dissection of four individ- 
uals; “proximal” and “distal” in the fol- 
lowing account refer to the position in 
relation to the genital orifice. The right 
ommatophore retractor muscle runs 
free, ¡i.e. it does not cross between penis 
and vagina. Flagellum long (longer than 
epiphallus), slender, tapering, curved; 
epiphallus slightly shorter than penis; 
penial retractor muscle attached to prox- 


HOLYOAK AND HOLYOAK: Á new species of Candidula from central Portugal 


Figures 1-5. Candidula coudensis spec. nov. holotype, shell 11.6 mm in width (BM reg. no. 


20100177). 1-3: adult shell; 4, 5: holotype photographed alive. 
Figuras 1-5. Candidula coudensis spec. nov. holotipo, anchura de la concha 11,6 mm (BM n* 
20100177). 1-3: concha adulta; 4, 5: holotipo fotografiado en vivo. 


imal part of epiphallus; penis thin- 
walled, internally with conical verge 
(having small simple apical pore) 
descending from distal part of penis 
(p2) into proximal part of penis (p1). 
Free oviduct moderately long. Dart sac 
(stylophore) single, moderately large, 
muscular, united with wall of vagina for 
about three-quarters of its length, enter- 
ing proximal part of vagina through 
conical papilla. Digitiform  glands 
(“mucus glands”) two, arising from 
vagina just distal to its union with dart- 
sac complex, each gland divided near 
base into two or three short branches. 
Spermatheca (bursa-copulatrix) cylin- 
drical-conical, lying on spermoviduct; 
spermatheca duct rather short and wide. 

Dimensions: holotype 11.6 x 5.0 mm, 
90 adult paratypes 8.8-11.4 x 4.7-5.4 mm. 

Distribution and habitat: C. coudensis 
has been found only in a range less than 
one kilometre in total extent, near Vale 


da Couda, by the N348 road SE. of 
Almoster (district of Leiria), Portugal. 
This range overlaps the edges of three 
different 10-km squares of the U.T.M. 
grid (NE40, NE41 and NE51). Candidula 
cf. belemensis lives in the same areas, 
and also at several other sites within a 5 
km radius on the same limestone hills 
where C. coudensis was not found. 
Within its restricted range C. coudensis 
occurs at ca 380-390 m altitude, over 
Mesozoic limestone that is exposed in 
crags facing east and north and also as 
scattered rocks and in stone walls. The 
largest numbers were found living on 
12th December 2009, resting above the 
ground on herbs, grasses and low 
bushes on almost flat, rather open, dis- 
turbed ground near the roadside (11 
living C. coudensis were found, with at 
least 11 living C. cf. belemensis in similar 
sites in the same small area). Single C. 
coudensis were also found living under 


69 


Iberus, 28 (1), 2010 


go 


Figures 6, 7. Candidula coudensis spec. nov. Anatomy of genitalia of a paratype (in collection of 
G.A. Holyoak). 6: general view of anatomy of proximal genitalia; 7: view of vagina and dart sac 
from other side. Abbreviations, dg: digitiform gland(s); ds: dart sac; e: epiphallus; f: flagellum; go: 


genital orifice; pl: proximal part of penis; p2: distal part of penis; r: penial retractor muscle; s: 


spermatheca; sd: spermathecal duct; v: vagina. 


Figuras 6, 7. Candidula coudensis spec. nov. Anatomía de genitales de un paratipo (en colección de 
G.A. Holyoak). Fig. 6 is Vista anatómica general de la parte proximal del tracto genital, Fig. 7 vista de 
la vagina y del saco del dardo desde el lado opuesto. Abreviaturas, de: glándula(s) digitiforme(s); ds: saco 
del dardo; e: epifalo; f: flagelo; go: orificio genital; pl: parte proxima del pene; p2: parte distal del pene; 


r: músculo retractor del pene; s: espermateca; sd: conduto de la espermatecas; v: vagina. 


limestone rocks near the sparsely vege- 
tated base of the east-facing crags on 
31st January 2008 and 10th June 2009. 

Remarks: Among Iberian Helicellini 
the presence of only a single moderately 
large clearly visible dart sac is a distinc- 
tive feature of Candidula, although an 
externally unnoticeable “accessory sac” 
lacking a dart may also be present 
alongside it and sometimes also a trace 
of an atrophied dart sac within the 
opposite wall of the vagina (e.g. GITTEN- 
BERGER, 1985; HAUSDORFE, 1988; ALONSO, 
IBÁNEZ AND HENRÍQUES, 1996; ScHI- 
LEYKO, 2006). 

A review of Portuguese species of 
Candidula by GITTENBERGER (1993) 


7O 


recognised six species, largely from shell 
characters. Of these, C. intersecta (Poiret, 
1801) and C. gigaxii (L. Pfeiffer, 1848) 
have wide ranges in western Europe, 
whereas the other four are apparently 
endemic to Portugal (C. setubalensis 
known only from Serra da Arrábida; C. 
belemensis known from districts of Faro 
(the Algarve) and Setúbal, and possibly 
(pers. obs.) northwards to the district of 
Leiria; C. olisippensis (Servain, 1880) 
known locally from the Algarve north- 
wards to the district of Porto (formerly, 
Douro Litoral); C. codía (Bourguignat, 
1859) known only from the Algarve, 
where additional localities were 
reported by MENDES SIMOES, 2006). 


HOLYOAK AND HOLYOAK: A new species of Candidula from central Portugal 


C. coudensis differs from all of these 
except C. setubalensis in having a sharply 
keeled shell. However, the flagellum in C. 
setubalensis is much shorter (less than half 
length of epiphallus: GITTENBERGER, 
1985) than in C. coudensis (in which it is 
longer than epiphallus: Fig. 6). Their 
shells also differ, comparisons based on 
37 adult or nearly adult shells of C. se- 
tubalensis (in Collection of G.A. Holyoak) 
revealing that they are on average small- 
er (largest shell 9.7 x 4.6 mm), with even 
less convex whorls (so spire often lower, 
mouth more elliptical, peripheral keel 
even sharper), and umbilicus larger (av- 
eraging ca 1/5 of shell width) and expos- 
ing more of spire internally; the sculpture 
and coloration of the shell are similar. All 
of the five other species of Candidula ac- 
cepted for Portugal by GITTENBERGER 
(1993) have depressed-globular shells 
with a rounded body whorl. In all five of 
these species the sculpture of radial ribs 
is weaker than in C. coudensis, although it 
is approached by C. codia, which other- 
wise differs markedly in its globular shell 
with domed spire. 

Elsewhere, the genus Candidula has 
been reported over a wide range from 
Ireland eastwards to southernmost 
Sweden and southwards to Fuerteven- 


ACKNOWLEDGEMENTS 


The authors want to thank T.L. 
Blockeel for advice on nomenclature 


BIBLIOOGRAPHY 


ALONSO M.R., IBÁÑEZ M. AND HENRÍQUES F.C. 
1996. Candidula ultima (Mousson, 1872) (Mo- 
lIlusca, Pulmonata, Hygromiidae), a nice case 
of adaptive convergence. Journal of Concho- 
logy, 35: 455-465. 

APARICIO M.T. 1982. Observations on the ana- 
tomy of some Helicidae from central Spain. 
Malacologia, 22 (1-2): 621-626. 

BANK R.A., BOUCHET P., FALKNER G., GITTEN- 
BERGER E., HAUSDORF B., PROSCHWITZ T. von 
AND RIPKEN T.E.J. 2001. Supraspecific clas- 
sification of European non-marine Mollusca 
(CLECOM Sections I + ID. Heldia, 4 (1-2): 77- 
128. 


tura (Canary Islands) in the west and 
the Balkan Peninsula in the east, with 
approximately 23 recognised species 
(KERNEY AND CAMERON, 1979; HAUS- 
DORE, 1988, 1991; MANGANELLI AND 
GiusTI, 1988; PUENTE, 1994; ALONSO, 
IBÁNEZ AND HENRÍQUES, 1996). Except 
for C. setubalensis, as discussed above, 
no other species currently assigned to 
the genus has a sharply keeled shell. 
However, a relatively long penial flagel- 
lum, exceeding one-half of the length of 
the epiphallus, has been described for 
the Spanish endemic species C. campor- 
roblensis (De Fez, 1944) and C. rocandioi 
(Ortiz de Zárate, 1950) (APARICIO, 1982). 
C. coudensis apparently reaches 
reproductive maturity in winter. A 
paratype with mature genitalia (Figs 6- 
7) was collected on 31st January 2008. 
Five of the snails collected on 12th 
December 2009 were kept alive indoors 
and these laid three clusters of eggs on 
16th-18th January 2010. The clusters 
were of 36, 64 and 77 eggs respectively, 
laid in loosely-adherent rounded-conical 
heaps, with a few single eggs scattered 
elsewhere in the box. Individual eggs 
were spherical, 1.0-1.3 mm in diameter, 
more or less translucent, whitish, or 
sometimes with a slight yellow tinge. 


and the referees for helpful sugges- 
tions. 


BANK R.A., GROH K. AND RIPKEN T.E.J. 2002. Ca- 
talogue and bibliography of the non-marine 
Mollusca of Macaronesia. In: Falkner, M., 
Groh, K. and Speight, M.C.D. (Eds), Collec- 
tanea Malacologica. Festschrift fiir Gerhard Falk- 
ner. Hackenheim: ConchBooks. Pp. 89-235. 

FALKNER G., BANK R.A. AND PROSCHWITZ “TT. 
VON 2001. Check-list of the non-marine mo- 
lluscan species-group taxa of the states of 
northern, Atlantic and central Europe (CLE- 
COM D). Heldia, 4 (1-2): 1-76. 


7 


Iberus, 28 (1), 2010 


GITTENBERGER E. 1985. The taxonomic status of 
Xeroplexa Monterosato, 1892 (Pulmonata: 
Helicidae: Helicellinae), a surprise. Iberus, 5: 
59-62. 

GITTENBERGER E. 1993. Digging in the graveyard 
of synonymy, in search of Portuguese species 
of Candidula Kobelt, 1871 (Mollusca: Gas- 
tropoda Pulmonata: Hygromiidae). Zoolo- 
gische Mededelingen, Leiden, 67 (17): 283-293. 

HAUSDORF B. 1988. Zur Kenntnis der systema- 
tischen Beziehung einiger Taxa der Helice- 
llinae Ihering 1909 (Gastropoda: Hygromii- 
dae). Archiv fúr Molluskenkunde, 119 (1-3): 9- 
37: 

HAUSDORF B. 1991. Uber zwei Candidula-Arten 
von der súdlichen Balkanhalbinsel (Gastro- 
poda: Hygromiidae). Archiv fir Mollusken- 
kunde, 120 (4-6): 119-129. 

KERNEY M.P. AND CAMERON R.A.D. 1979. Land 
snails of Britain and north-west Europe. Har- 
perCollins, London. 


da 


MANGANELLI G. AND GIUSTI F. 1988. Notulae 
malacologicae, 38. A new Hygromiidae from 
the Italian Apenines and notes on the genus 
Cernuella and related taxa (Pulmonata: He- 
licoidea). Bollettino Malacologico, 23 (11-12): 
327-380 [for 1987]. 

MENDES SIMOES J.M. 2006. Notas sobre a dis- 
tribuicáo geográfica de Candidula codia (Bour- 
guignat, 1859) (Gastropoda, Pulmonata: Hy- 
gromiidae) em Portugal. Portugala, 7: 3-4. 

PUENTE A.I. 1994. Estudio taxonómico y biogeo- 
gráfico de la superfamilia Helicoidea Rafinesque, 
1815 (Gastropoda: Pulmonata: Stylommatop- 
hora) de la Península Ibérica e Islas Baleares. 
Doctoral thesis (unpublished). Bilbao, 970 


pp- 

SCHILEYKO A.A. 2006. Treatise on recent te- 
rrestrial pulmonate molluscs, Part 14, Heli- 
codontidae, Ciliellidae, Hygromiidae. Rut- 
henica, Supplement 2: 1905-2047. 


O Sociedad Española de Malacología 


Iberus, 28 (19: 73-78, 2010 


A new species of Gibbula (Mollusca, Archaegastopoda) 


from Namibia 


Una nueva especie de Gibbula (Mollusca, Archaegastopoda) de 


Namibia 


Emilio ROLÁN* and Michael L. ZETTLER** 


Recibido el 14-XII-2009. Aceptado el 24-IV-2010 


ABSTRACT 


A new species is described from Namibian waters and compared with the morphologi- 


cally close species from the study area. 


RESUMEN 


Se describe una nueva especie de Namibia comparándola con las especies morfológica- 


mente similares del área de estudio. 


INTRODUCTION 


The Namibian molluscan fauna is 
very far from being well known. Only 
few works with information about the 
molluscs of this area are known (e.g. 
PENRITH AND KINSLEY, 1970a, 1970b). 
Usually, it is thought that this fauna is 
mostly influenced by that of South 
Africa (KILBURN AND RIPPEY, 1982, 
STEYN AND Lussi, 1998) due to the pres- 
ence of an oceanic current from the 
south towards the north that obstructs 
the dispersion of the West African 
fauna. 


MATERIAL AND METHODS 


The Leibniz Institute for Baltic Sea 
Research from Rostock-Warnemiinde, 
Germany, sampled the macrozooben- 
thos along the northern Namibian coast 
in 2008 with the research vessel “Maria 
Sybilla Merian”. Triplicate benthic 
samples were taken with a 0.1 m? van 


Veen grab at each station. Additional 
dredge hauls using a net mesh size of 5 
mm) were taken for collection of larger, 
mobile or rare species. All samples were 
sieved through a 1-mm? screen and 
animals were preserved on board in 4% 
buffered formaldehyde. Sorting proce- 
dures were conducted at the laboratory 
with a stereomicroscope with 10-40x 
magnification. For more information on 
the study area and abiotic characteristics 
at the sampling station see ZETTLER, 
BOCHERT AND POLLEHNE (2009). 

In this material, several samples of a 
species with conical form were found; 
since it was apparently undescribed, 
the purpose of the present paper is to 
name it. 


Abbreviations 
MHNS Museo de Historia Natural de la 


Universidad de Santiago de Com- 
postela, (coll. E. Rolán), 


* Museo de Historia Natural, Campus Universitario Sur, 15782 Santiago de Compostela, Spain. 
** Leibniz-Institute for Baltic Sea Research, Seestr. 15, 18119, Rostock, Germany. 


AS 


Iberus, 28 (1), 2010 


IOW Institute for Baltic Sea Research, 
Warnemimnde, 

MNCN Museo Nacional de Ciencias 
Naturales, Madrid, 


SYSTEMATICS 


WMC collection of Werner Massier, 
Swakopmund 

ZMB Museum fúr Naturkunde of the 
Humboldt University, Berlin, 


Family TROCHIDAE Rafinesque, 1815 
Subfalily TROCHINAE Rafinesque, 1815 
Genus Gibbula Risso, 1826 


Gibbula massieri spec. nov. (Figs. 1-14) 


Type material: Holotype (Figs. 1-3) in ZMB (Moll. 107743). Paratypes in the following collec- 
tions: MNCN (1, n* 15.05/53490, Fig. 5); IOW (3, IOW-NA-0001-3, Figs. 4, 7-9); MHNS (1); WMC 


(3, Figs. 6-9). 


Other material examined: One specimen was destroyed for radular study; 25 juveniles and some 


fragments (I0W). 


Type locality: Offshore Namibia south of the River Kunene, 30 m depth, 17.390” S 11.724 E (5-03- 
2008). Shell deposits of the brachiopod Discinisca tenuis (Sowerby, 1847). 
Etymology: The specific name is after Werner Massier, from Swakopmund, Namibia, for his coop- 


eration in the study of the material. 


Description: Shell (Figs. 1-9) conical, 
solid, with spiral striation and axial 
threads. Protoconch (Figs. 10-11) with 
about one smooth whorl and a diam- 
eter of about 90 um. The beginning of 
the teleoconch has 4 spiral cords, sepa- 
rated by deep interspaces. The subse- 
quent whorls have a straight profile 
and a similar number of cords, the last 
whorl having between 5 and 6 cords 
down to the periphery, which is well 
angled. Below it, towards the base, 
there are 7-8 cords, clearly separated, 
and closer near the umbilical 
infundibulum. In the  interspaces 
between the cords there are numerous 
small very close prosocline threads. 
Aperture rectangular, columella slightly 
prosocline, straight, external border 
sharp, slightly undulating as a result of 
the endings of the spiral cords. The 
colour of the first whorls (corre- 
sponding to the juvenile stages) is 
brown on the spiral cords and white on 
its interspaces; in the subsequent 
whorls, axial dark blotches appear, 
alternating with white or light colour. 
On the base, the cords usually are alter- 
nate brown and white or cream, but 
this is very irregular. 


74 


Dimensions: holotype 6.5 mm in 
height by 6.1 mm in diameter. The 
paratypes are of a similar size. 

The animal, examined in specimens 
preserved in alcohol, is whitish with iso- 
lated dark lines on the head. There are 
three evident epipodial tentacles on 
each side. Operculum (Fig. 12) rounded, 
fine, semitransparent and multispiral, 
with a central nucleus. 

Radula (Figs. 13, 14) rather typical of 
the genus: rachidian tooth with a 
reduced shaft, which has only a few 
small cusps; the lateral teeth 1-5 are 
similar in form, increasing slightly in 
size outwards; all of them have a 
pointed apex and several lateral cusps 
(up to 6), the external one larger. The 
marginal ones number about 30 on each 
side, the 5-6 more internal ones are 
larger and have 1-2 lateral cusps; going 
up to the external ones, their size dimin- 
ishes while the number of their lateral 
cusps increases. 

Distribution: 
Namibia. 

Remarks: The placement of this 
species in the genus Gibbula is based on 
the relatively small size of the shell, its 
conical form, lack of columellar denticle, 


Only known from 


ROLÁN AND ZETTLER: A new species of Gibbula from Namibia 


Figures 1-9. Gibbula massieri spec. nov. 1-3: holotype, 6.5 mm (ZMB); 4-9: paratypes; 4: 5.4 mm 
(IOW); 5: 5.5 mm (MNCN); 6: 5.0 mm (WMOC); 7,9: 6.4 mm (1OW); 8: 5.0 mm (IOW). 
Figuras 1-9. Gibbula massieri spec. nov. 1-3: holotipo, 6,5 mm (ZMB); 4-9: paratipos; 4: 5,4 mm 
(LOW); 5: 5,5 mm (MNCN); 6: 5,0 mm (WMC); 7, 9: 6,4 mm (1I0OW); 8: 5,0 mm (I0W). 


Iberus, 28 (1), 2010 


Figures 10-14. Gibbula massieri spec. nov. 10-11: protoconch; 12: operculum; 13-14: radula. 
Figuras 10-14. Gibbula massieri spec. nov. 10-11: protoconcha; 12: opérculo; 13-14: rádula. 


7Ó 


ROLÁN AND ZETTLER: A new species of Gzbbula from Namibia 


presence of an umbilicus, existence of 
three epipodial tentacles at each side of 
the body, and radula with a narrow 
shaft and reduced cusp in the rachidian 
tooth. 

Among the South African species of 
small size we can separate the new 
species from the following; 

Cantharidus suarezensis (P. Fischer, 
1878) is lighter in colour, the umbilicus 
narrower. 

Gibbula beckeri Sowerby, 1901, is 
more depressed, the whorls convex and 
slightly shouldered, the sculpture is 
weaker, the colour is also a combination 
of dark and light blotches but with a 
smaller pattern. 

Gibbula zonata (Wood, 1828) is more 
depressed, with convex whorls, spiral 
cords in low number, and a wider 
umbilicus. 

Gibbula cicer (Menke in Philippi, 
1844) has convex whorls, prominent and 
less numerous spiral cords, the umbili- 
cus usually closed. 

Gibbula fultoni (Sobwerby, 1889), 
according to the description in BARNARD 
(1963), has a conical form similar to that 
of the present new species but it has a 


ACKNOWNLEDGEMENTS 


We thank Dr. U. Struck (Berlin), Dr. 
V. Mohrholz (Rostock) U. Hehl 
(Rostock), M. Rómer (Hamburg) and C. 
Berg (Rostock) for assistance during 
sampling at RV Maria S. Merian. We 
wish also to thank Dr. Ralf Bochert and 
I. Glockzin (both Rostock) for analysis 


BIBLIOGRAPHY 


BARNARD R. H. 1963. Contribution to the knowl- 
edge of South African Marine Mollusca. Part 
IV. Gastropoda: Prosobranchiata: Rhipi- 
doglossa, Docoglossa, Tectibranchiata. Poly- 
placophora. Solenogastres. Scaphopoda. An- 
nals of the South African Museum, 47 (2): 201- 
360. 

DAUTZENBERG P. 1910. Contribution a la faune 
malacologique de l' Afrique occidentale Actes 
de la Société Linnénne de Bordeaux, 44: 1-174, 
pl. 1-4 


completely or almost closed umbilicus. 
The shell is also larger. 

Among the Angolan species, com- 
parison must be made with: 

Jujubinus fulgor Gofas, 1991, which 
has an apparently similar shell, but is 
more solid, smaller, without any umbili- 
cus and having a columellar lengthen- 
ing on its lower part. 

The comparison with species from 
other areas: 

Gibbula verdensis Rolán and Tem- 
plado, 2001 has a similar form and size, 
but the colour is usually lighter, the 
white blotches are very small and the 
umbilicus is reduced to a small furrow. 
It is endemic to the Cape Verde islands. 

Gibbula joubini Dautzenberg, 1910 is 
smaller (about 4.5 mm) the peripheral 
border is more rounded, the dark brown 
blotches are most frequently opistocline, 
the aperture is relatively smaller. This is 
an endemism from Senegal which has 
not been recorded from other countries. 

The species recorded by GOFas, 
PINTO AFONSO AND BRANDAO (1985) 
from Angola as Gibbula aff. joubini, may 
be the species here described or other 
similar (material not examined). 


of benthic samples in the laboratory. To 
Jesús Méndez of the Centro de Apoyo 
Científico y Tecnológico a la Investi- 
gación (CACTI) of the University of 
Vigo for the SEM photographs. And 
António A. Monteiro for English correc- 
tions to the manuscript. 


GOFAS S. 1991. Un nouveau Jujubinus d'An- 
gola. Apex, 6 (1): 21-24. 

GOFAS S., PINTO AFONSO J. AND BRANDAO M. 
[1985]. Conchas e moluscos de Angola. Uni- 
versidade Agostinho Neto / Elf Aquitaine 
Angola, 144 pp. 

KILBURN R. AND RIPPEY E. 1982. Sea Shells of 
Southern Africa. Macmillan South Africa, Jo- 
hanesburg. 249 pp. 


YN 


Iberus, 28 (1), 2010 


PENRITH M. L. AND KENSLEY B. 1970a. The cons- STEYN D. G. AND Lussi M. 1998. Marine Shells 
titution of the intertidal fauna of rocky sho- of South Africa. Ekogilde Publishers, Hartbe- 
res of South West Africa. Part I. Lúderitz- espoort. 264 pp. 
bucht. Cimbebasia, 1: 191-239. ZETTLER M.L., BOCHERT R. AND POLLEHNE F. 

PENRITH M. L. AND KENSLEY B. 1970b. The cons- 2009. Macrozoobenthos diversity in an oxy- 
titution of the intertidal fauna of rocky sho- gen minimum zone off northern Namibia. 
res of South West Africa. Part II. Rocky Point. Marine Biology, 156: 1949-1961. 


Cimbebasia, 1: 241-268. 

ROLÁN E. AND TEMPLADO J. 2001. New species 
of Trochidae (Mollusca, Gastropoda) from the 
Cape Verde Archipelago. Iberus, 19 (2): 41-55. 


78 


O Sociedad Española de Malacología Iberus, 28 (1):79-89, 2010 


New information on the Caribbean Rissoina (Gastropoda, 
Rissoidae) of the group R. sagraiana-cancellata with the 
description of a new species 


Nueva información sobre las Ríssoina del Caribe (Gastropoda, 
Rissoidae) del grupo de R. sagraiana-cancellata, con la descripción 


de una nueva especie 


Emilio ROLÁN* and Raúl FERNÁNDEZ-GARCÉS** 


Recibido el 2-X11-2009. Aceptado el 27-IV-2010 


ABSTRACT 


Five species of the genus Rissoina from Cuba are characterized and illustrated with scan- 
ning electron micrographs: Rissoina sagraiana, R. pulchra, R. labrosa, R. redferni and R. 
cancellina spec. nov. Rissoina cancellata is here placed in synonymy of R. sagraiana. 


RESUMEN 


Se caracterizan e ilustran con micrografías electrónicas de barrido cinco especies del 
género Rissoina de Cuba: Rissoina sagraiana, R. pulchra, R. labrosa, R. redferni y R. can- 
cellina spec. nov. Se considera Rissoina cancellata como un sinónimo de R. sagraiana. 


INTRODUCTION 


Some of the species of the genus Ri- 
ssoina in the Caribbean are poorly known, 
sometimes only from bibliographic refer- 
ences or from just a few figures. Even 
species recently described lack good pho- 
tographs of details, such as the protoconch 
or microsculpture; this hinders the perfect 
knowledge of the species as well as the 
comparison with other species of the group, 
sometimes forcing other researchers to face 
problems about synonymy or validity. 

Some species of this group were 
described for the Caribbean in a previ- 
ous century: D'ORBIGNY (1842), PHILIPPI 
(1847), ADAMS (1850), etc. 

PONDER (1985) revised the family 
Rissoidae at the generic and subgeneric 
levels, including among others the sub- 


genus Phosinella Mórch, 1876 based on 
one of the species here studied. 

The taxonomy of the species treated 
herein has been subject to disagreement. 
DESJARDIN (1949) revised the Cuban 
species of Rissoina and considers R. pulchra 
as a synonym of R. cancellata, followed in 
this by PONDER (1985). Conversely DE 
JONG AND COOMANS (1988) consider that 
R. pulchra is a synonym of R. sagraiana. 
Other works on the Caribbean malaco- 
logical fauna have recorded and repre- 
sented species of this group: WARMKE AND 
ABBOTT (1961), ABBOTT (1974), DE JONG 
AND COOMANS (1988), Díaz MERLANO AND 
PUYANA HEGEDUS (1994), Rios (1994), 
REDFERN (2001), ESPINOSA AND ORTEA 
(2002), LEE (2009), among others. 


* Museo de Historia Natural, Campus Universitario Sur, 15782 Santiago de Compostela, Spain. 
** Centro de Estudios Ambientales de Cienfuegos (CEAC), División de Gestión Ambiental (DGA), calle 17, 


esquina Ave. 46, Cienfuegos, Cuba. 


a 


Iberus, 28 (1), 2010 


In the present paper, we present 
information on some characters of five 
Caribbean species of Rissoina which are 
morphologically very close, although the 
protoconchs show enough characters for 
differentiation. The previous papers on 
this group by the authors were ROLÁN 
AND FERNÁNDEZ-GARCÉS (2009a, 2009b). 


Abbreviations 


BMNH The Natural History Museum, 
London 

CFG coll. Fernández Garcés, Cienfuegos 

CHL collection of Harry G. Lee, Florida 


SYSTEMATICS 


IES Intituto de Ecología y Sistemática, 
La Habana 

IGH Instituto de Oceanología, La 
Habana 

MCZ Museum of Comparative Zoology, 
Harvard 

MHNS Museo de Historia Natural, San- 
tiago de Compostela 

MNCN Museo Nacional de Ciencias 
Naturales, Madrid 

MNHN Museum national d'Histoire 
naturelle, Paris 

ZMB Zoologisch Museum, Berlin 

s shell 

] juvenile 


Superfamily RISSOIDEA J. E. Gray, 1847 
Family RISSOIDEA J. E. Gray, 1847 
Subfamily RISSOININAE Stimpson, 1865 


Genus Rissoina d'Orbigny, 1840 


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


Rissoina sagraiana d'Orbigny, 1842 (Figs. 1A-F) 


Rissoina sagra d'Orbigny, 1842. Hist. Nat. Cuba, Atlas pl. 12, figs 4,5. 
Rissoina sagraiana d'Orbigny, 1846. Hist. Nat. Cuba, vol. 2: 25. 
Rissoina cancellata Philippi, 1847. Zeitsch. Malak., 1846: 127. 


Type material: R. sagraiana: Lectotype, selected in original lot of syntypes by Ángel Luque and 
hereby designated (Fig. 1A-C) and 3 paralectotypes in BMNH. R. cancellata: Holotype in ZMB (coll. 


Pfeiffer, 2334), 5.8 mm (see Remarks). 


Other material examined: Cuba: 12 s, 2j, Cayo Carenas, Cienfuegos Bay, 5-10 m (CFG); 20 s, 16 j, 
Rancho Luna, Cienfuegos, 12 m (CFG); 12 s, Faro Luna, Cienfuegos, 40 m (CFG); 8 s, 3j Mangles 
Altos, Cienfuegos, 8 m (CFG); 5 s, Rancho Club, Cienfuegos Bay, 2 m (CFG); 10 s, Rancho Luna, 
Cienfuegos, 10-55 m (MHNS); 10 s, Faro de los Colorados, Cienfuegos, 20 m (MHNS). Nicaragua: 


1 s, 1 f, Cayo Witties, 20 m (MHNS). 


Description: Shell (Figs. 1A-C, 1E): see 
the original descriptions (D'ORBIGNY, 
1846; PHILIPPL, 1847). Protoconch (Figs. 
1D, 1F) with about 2 */2 - 2 9/4 whorls, 
sometimes almost 3 whorls, and with a 
diameter of about 390-400 um. It is 
smooth but on the last whorl (or starting 
a little before) an evident spiral thread 
can be observed at the middle of the pe- 
riphery. At the beginning of the teleo- 
conch there are two spiral cords in the 
middle of the whorl; in the following 


80 


whorls, a new spiral cord appears be- 
tween the two previous ones. The spiral 
cords are 6-7 on the last whorl, plus 
some smaller ones near the base. 
Holotype of 5.8 mm, other shells 
from Cuba are slightly larger. 
Distribution: This species is well 
known from many works from all the 
Caribbean: WARMKE AND ABBOTT (1961) 
from Puerto Rico; VOKES AND VOKES 
(1983) from Campeche and Yucatan; DE 
JONG AND GCOOMANS (1988) from 


ROLÁN AND FERNÁNDEZ-GARCÉS: Caribbean Rissoina of the group R. sagraiana-cancellata 


Figure 1. Rissoina sagraiana d'Orbigny, 1842. A-C: lectotype, 5.5 mm (BMNH); D: protoconch of 
the lectotype; E: shell from Cienfuegos, Cuba, 5.7 mm; F: Protoconch, same shell as E. 

Figura 1. Rissoina sagraiana d'Orbigny 1842. A-C: lectotipo, 5,5 mm (BMNAH); D: protoconcha del 
lectotipo; E: concha de Cienfuegos, Cuba, 5,7 mm; E: Protoconcha, misma concha que E. 


Curacao, Aruba and Bonaire; DÍAZ REDFERN (2001) from Abaco (Bahamas) 
MERLANO AND PUYANA HEGEDUS (1994) with a good representation of the proto- 
from Colombia; Rios (1994) from Brasil; conch; LEE (2009) from Northeast 


8 | 


Iberus, 28 (1), 2010 


Florida. Cuba is the type locality and 
also we have material from Nicaragua. 

Remarks: Examining the material from 
ZMB we found one lot (ZMB, 2334) with 
only one shell of 5.8 mm from Cuba, which 
is labelled as R. cancellata Philippi (coll. 
Pfeiffer), in good condition, with multi- 
spiral protoconch, with a spiral cordlet on 
the last whorl. We have no doubt that this 
shell is the holotype of R. cancellata because 
the locality and collector coincide with the 
indications given by PHILIPPI (1847) in the 
original description. It seems to be the 
same species that we usually call R. sagra- 
lana with the characters previously men- 
tioned in the description. 

Rissoina sagraiana appears in the liter- 
ature as the oldest species name for this 
morphologically similar group, despite 
the indication by DESJARDIN (1949) listing 
the name Rissoina sagraiana d'Orbigny, as 
from 1853. The species was first published 
by D'ORBIGNY (1842) under the name Ris- 
soina sagra in the caption of the plate and 


thereby made available under the provi- 
sions of article 12.2.7 of the International 
Code of Zoological Nomenclature; the 
complete description was published years 
later (1846) under the name R. sagraiana 
and the publication in parts was com- 
pleted only in 1853 (dates as stated in 
BOUCHET AND ROCROI, 2005). Therefore 
there are two different spellings of the 
name, but according to article 32.5.1.1 of 
the ICZN, correction of the spelling of a 
name in a later part of a work published 
in parts constitutes an evidence for an 
inadverted error and therefore, the species 
must be cited as Rissoina sagraiana d'Or- 
bigny, 1842 having priority over Rissoina 
cancellata Philippi, 1847. 

This species is figured under the 
name R. cancellata by PONDER (1985: 83, 
fig. 135) who considered it as a synonym 
of R. pulchra C. B. Adams, 1850), the type 
species of subgenus Phosinella Mórch, 
1876. The protoconch shown therein is 
typical of Rissoina sagraiana. 


Rissoina pulchra (C. B. Adams, 1850) (Figs. 2A-E) 


Rissoa pulchra C. B. Adams, 1850. Cont. Conch., 7: 114. 


Type material: Lectotype (Fig. 2A) in MCZ (156423)1(C. B. Adams coll., Acc. 1173), designated by 
CLENCH AND TURNER (1950) by inference of a holotype. 

Other material examined: Cuba: 1 s, Faro Luna, Cienfuegos 40 m (CFG); 1 s, Rancho Luna, 20 m 
(MHNS); 1 s, Faro de los Colorados, 35 m (MHNS); 1 s, Rancho Club, 10 m (all in Cienfuegos Bay); 


1 j, Maria la Gorda, 30 m (MHNS). 


Description: See original description 
(ADAMS, 1850 and CLENCH AND TURNER, 
1950). The shell (Figs. 3A-B) seems to be 
very similar to that of R. cancellata even 
in the protoconch, which is smaller (Fig. 
3C), also multispiral, with about 2 !/2 
whorls and a diameter of 270 um. Under 
magnification it seems that this proto- 
conch lacks any spiral sculpture. 

Dimensions: the lectotype is 5.7 mm. 
Shells from Cuba may be a little larger 
(up to 7.0 mm). 

Distribution: From Abaco (Bahamas) 
by REDFERN (2001) showing a proto- 
conch. “Jamaica” in the original descrip- 
tion (type locality). 

Remarks: This is the type species of the 
subgenus Phosinella Mórch, 1876 (SD 


82 


Nevilll, 1885). PONDER (1985) considered 
R. pulchra C. B. Adams, 1850 as a synonym 
of R. cancellata and used the latter name 
because Rissoa pulchra is a primary 
homonym of Rissoa pulchra Forbes, 1844, 
described from the Aegean Sea. As already 
mentioned the shell figured in PONDER 
(1985) and presumably also the specimens 
used for anatomy are R. sagraiana = can- 
cellata rather than R. pulchra. The taxon 
Rissoa pulchra Forbes, 1844 has never been 
mentioned since its description, and it is 
uncertain to which species it refers. Con- 
versely, R. pulchra Adams, 1850 has been 
recorded sometimes during the last 50 
years: NOWELL-USTICKE (1959), ABBOTT 
(1974), Díaz MERLANO AND PUYANA 
HEGEDUS (1984), PONDER (1985), DE JonG 


ROLÁN AND FERNÁNDEZ-GARCÉS: Caribbean Rissoina of the group R. sagraiana-cancellata 


Figure 2. Rissoina pulchra (C. B. Adams, 1850) 


¿JAS Lectotype, )./ min, MCZ (156423). B: shell, 


5.6 mm, Cuba (MHNS); C: protoconch, same shell as B; D-E: protoconch of the lectotype. 
Figura 2. Rissoina pulchra (C. B. Adams, 1850). A: Lectotipo, 5,7 mm, MCZ (156423). B: concha, 
5,6 mm, Cuba (MHNS); C: protoconcha, misma concha que B; D-E: protoconcha del lectotipo. 


AND COOMANS (1988), Rios (1994), 
REDFERN (2001). Although this is proba- 
bly short of the 25 publications required 
to make it a nomen protectum under the 
provisions of ICZN, we also consider 
current usage in leading databases such 


as Malacolog <http:/ /www.malacolog. 
org/>, the World Register of Marine 
species <www.marinespecies.org> and 
the Encyclopedia of Life <www.eol.org>. 
For these reasons we propose to keep 
using the name R. pulchra C. B. Adams. 


83 


Iberus, 28 (1), 2010 


Table I. Schematic comparison of the shells of the Rissoina sagraíana group. 
Tabla I. Comparación de las conchas del grupo de Rissoina sagraiana. 


species protoconch whorls pm diameter protoconch 
sagraiana 21-24 390-400 
pulchra a 270 
cancellina 2 320-330 
labrosa 2 400 

redferni 11/ 380 


CLENCH AND TURNER (1950) men- 
tioned and figured the holotype of this 
taxon: as Adams (1850) did not designate 
a holotype, the previous mention may be 
considered as a lectotype designation. 

REDFERN (2001) represented the three 
species sagratana-cancellata-pulchra and 
described the protoconchs with the dif- 


spiral cord in protoconch 


spiral cords at first teleoconch whorl 


yes 2 
no 2 
yes 3 
yes 2 small, joint 
no 3 


ference of their being keeled or not. 
According to him, R. pulchra differs from 
R. sagraiana and R. cancellata (actually, from 
the species described herein as R. cancel- 
lina spec. nov.) in having a wider proto- 
conch without any spiral cord. We agree 
with this difference. The details of this 
comparison are given in Table 1. 


Rissoina cancellina spec. nov. (Figs. 3A-E) 


Type material: Holotype (Fig. 3A) in MNCN (15.05/53590). Paratypes: ZMB (1, n* 115039) (Fig. 3B) 
labelled as Rissoina cancellata Phil. v. pulchra C. B. Adams, Jamaica, coll. Paetel; MNHN (1), MHNS 
(5), IES (1), CFG (10). 

Other material examined: Cuba: 8 s, 3j, Cayo Carenas, Cienfuegos Bay, 5-10 m (CFG); 10 s, Rancho 
Luna, 20-40 m (CFG); 11 s, 3j, Faro de los Colorados, Cienfuegos, 15 m (CFG); 2 s, La Concha, Cien- 
fuegos Bay, 3 m (CFG); 2 ej, Guajimico, Cienfuegos, 3 m (CFG); 10 s, 8j, Rancho Luna, Cienfuegos, 
10-55 m (MHNS); 3 s, Faro de los Colorados, Cienfuegos, 20 m (MHNS); 2 s, 3 j, Bajo de Sancho 
Pardo, 4 m (MHNS); 1 s, Cuba, ZMB (coll. Dunker, 115036). Saint Thomas: 2 s, ZMB (115038). West 
Indies: 7 s, ZMB (coll. Paetel, 115037). Nicaragua: 5 s, 5j, Cayo Miskitos, 25 m (MHNS). 

Type locality: Cienfuegos, Cuba. 

Etymology: From the Latin word cancelli “lattice gate” alluding to the crossed sculpture, and also due 
to the similarity with Rissoina cancellata, taxon which now is considered in synonymy of R. sagraiana. 


Description: Shell (Figs. 3A-B) ovoid 
elongate, pointed, solid and whitish. 
Protoconch (Figs. 3C, 3D) almost cylin- 
drical and a little depressed, with 
between 320-330 um, and about 2 
whorls, the first one smooth and the 
second with a spiral cord at the middle. 
The beginning of the teleoconch has 
immediatly three spiral cords. The sub- 
sequent whorls also have 3-5 spiral 
cords crossed by axial ribs. These ribs 
are 10-13 in the first whorls and 16-19 on 
the last one. On the body whorl the 
cords are 9-10. On the subsutural area 
(Fig. 3E) there are about five very small 
spiral threads. Under high magnifica- 


84 


tion (Fig. 3F) numerous micro perfora- 
tions can be seen on the surface of the 
shell. Aperture ovoid with a typical 
depression of the genus near the base. 

The holotype is 9.3 mm. Other shells 
are slightly smaller. 

Distribution: Probably this species 
has a large distribution in the 
Caribbean, but usually it has been 
recorded under different names, so that 
confirmation is needed in order to avoid 
possible confusion with other similar 
species. REDFERN (2001, as R. cancellata) 
figured the protoconch. 

Remarks: The holotype is a shell in 
good condition and with a good proto- 


ROLÁN AND FERNÁNDEZ-GARCÉS: Caribbean Rissoina of the group R. sagraiana-cancellata 


Figure 3. Rissoina cancellina spec. nov. A: holotype, 5.7 mm, Cuba (MNCN); B: paratype, 8.5 
mm, ZMB (115039); C: protoconch of the holotype; D: protoconch, Cayo Miskitos, Nicaragua; 
E, F: microsculpture of the holotype. 

Figura 3. Rissoina cancellina spec. nov. A: holotipo, 5,7 mm, Cuba (MNCN); B: paratipo, 8,5 mm, 
ZMB (115039); C: protoconcha del holotipo; D: protoconcha, Cayo Miskitos, Nicaragua; E, EF: micro- 
escultura del holotipo. 


85 


Iberus, 28 (1), 2010 


Figure 4. Rissoina labrosa Schwartz, 1860. A: Shell, 11.0 mm, Cienfuegos, Cuba; B, C: Protoconch. 
Figura 4. Rissoina labrosa Schwartz, 1860. A: Concha, 11,0 mm, Cienfuegos, Cuba; B, C: Protoconcha. 


conch (Fig. 2A) measuring 9.3 mm 
(ZMB, 115039) and labelled as Rissoina 
cancellata Phil. v. pulchra C. B. Adams, 
Jamaica, coll. Paetel. 

In the material in the ZMB, there are 
several shells which can belong to this 
species. One of them (ZMB, 115036) 
from Cuba, coll. Dunker, has a shell 
whose protoconch presents excrescences 
on its upper part and is hence difficult 
to observe. Six shells more in this lot 
have lost the protoconch and are badly 
eroded. Another lot (ZMB, 115037) 
includes a probable R. cancellina but 
eroded and without protoconch. One 


more (ZMB, 115038) may also be the 
same species but is much eroded. 

R. cancellina spec. nov. must be com- 
pared with the following; 

R. sagraiana has a protoconch with 
more whorls (2 */2 - 2 9/4) and the first 
whorls of the teleoconch have only two 
spiral cords (see Table 1). 

R. pulchra has a protoconch with more 
whorls, lacking any spiral cord, while in 
the beginning of the teleoconch there are 
only two cords on the first whorls. 

R. redferni has a protoconch with less 
whorls (1 */2) but wider, and without 
any spiral cord. 


Rissoina labrosa Schwartz, 1860 (Figs. 4A-B) 


Rissoina labrosa Schwartz, 1860. Uber... Die Gattung Rissoina, pl. 7, fig. 58. 


Rissoina sheaferi Mc Ginty, 1962: 42, pl. 3, fig. 5. 


Type material: Supposedly in the Vienna Museum. Not examined. 
Material examined: Cuba: 1 s, Cayo Carenas, Cienfuegos Bay, 10 m (CEG); 1 s, Faro de los Colorados, 
Cienfuegos, intertidal (CFG); 3 j, Rancho Luna, Cienfuegos, 12 m (CFG); 1 s, Faro Luna, Cienfuegos, 


86 


ROLÁN AND FERNÁNDEZ-GARCÉS: Caribbean Ríissozna of the group R. sagraiana-cancellata 


Figure 5. Rissoina redferni Espinosa and Ortea, 2002. A: shell, 6.4 mm, Egmond Key, Florida 
(CHL); B: shell, 4.8 mm, Cienfuegos, Cuba; C: shell, 5.8 mm, Cayo Miskitos, Nicaragua; D, E: 
protoconch, shell from Florida; F: protoconch, shell from Cuba; G: microsculpture of the proto- 
conch, same shell as D; H, I: microsculpture of the teleoconch, shell from Florida. 

Figura 5. Rissoina redferni Espinosa y Ortea, 2002. A: concha, 6,4 mm, Egmond Key, Florida (CHL); 
B: concha, 4,8 mm, Cienfuegos, Cuba; C: concha, 5,8 mm, Cayo Miskitos, Nicaragua; D, E: protocon- 
cha, concha de Florida; F: protoconcha, concha de Cuba; G: microescultura de la protoconcha, misma 
concha que D; H, I: microescultura de la teleoconcha, concha de Florida. 


87 


Iberus, 28 (1), 2010 


20 m (MHNS); 2 s, Rancho Luna, Cienfuegos, 22-54 m (MHNS); 1 s, Cable Inglés, Cienfuegos, 20 m 
(CEG); 5 s, 6j, Bajo de Sancho Pardo, 3-5 m (MHNS); 2 s, 5 j, María la Gorda, 30 m (MHNS). Mexico: 
4 s, Puerto Morelos, Quintana Roo, Yucatán (MHNS). Nicaragua: 3 s, Cayo Los Muertos, 15 m (MHNS). 


Description: See SCHWARTZ (1860). 
The protoconch has a little more than 
2 whorls and has a cylindrical pro- 
file. 

Distribution: Known from Cuba 
(type locality), Curacao, Aruba and 
Bonaire (DE JONG AND COOMANS, 1988). 
Nicaragua and Mexico, in the present 
work. 


Remarks: This shell may be consi- 
dered somewhat different from the 
others included in the group, and was 
previously mentioned as being larger 
and more pointed, with a straight 
profile, with the suture scarcely marked, 
and the protoconch cylindrical and ele- 
vated. We figure a shell (Fig. 4A) and 
protoconch (Figs. 4B-C) for comparison. 


Rissoina redferni Espinosa and Ortea, 2002 (Figs. 5A-H) 
Rissoina redferni Espinosa and Ortea, 2002. Avicennia, 15: 142. 


Type material: Holotype (IGH) not examined. 

Other material examined: Cuba: 25 shells, Cienfuegos Bay, 35 m (MHNS); 3 s, Punta Tamarindo, 
Cienfuegos, 20 m (MHNS); 5 s, de Sagua la Grande, Villa Clara, 3-10 m (CFG); 4 s, Batabanó, Habana, 
5-7 m (CFG). Florida: 4 s, Egmond Key (CHL). Nicaragua: 3 s, Cayo Miskitos, 6 m (MHNS); 7 s, 


Cayo Los Muertos, 12 m (MHNS); 5 s, Cayo Witties, 20 m (MHNS). 


Description: Shell (Figs. 5A-C): see Es- 
PINOSA AND ORTEA (2002). This descrip- 
tion is short in some aspects referring to 
micro sculpture: the protoconch (Figs. 
5D-E) has 1 */4 whorls which have small 
tubercles spirally aligned, more evident 
near the end (Fig. 5G). The nucleus has 
about 160 um; the first half whorl mea- 
sures about 270 um, and the protoconch 
about 380 um. The micro sculpture of the 
teleoconch (Figs. 5H-I) presents spiral 
lines formed by a rough surface and un- 
der very high magnification shows small 
pits. The comparison of the material 
from Cuba and from the Bahamas did 
not show meaningful differences. 

Distribution: Known from Bahamas 
(REDFERN, 2001), Nicaragua and Cuba 


ACKNOWLEDGEMENTS 


The authors thank Jesús Méndez 
and Inés Pazos of the Centro de Apoyo 
Científico y Tecnológico a la Investi- 
gación (CACTI) of Vigo University for 
the SEM micrographs of the shells. 
Thanks also to Matthias Glaubrecht 
(ZMB) for sending type material for ex- 


88 


(type locality: Playa Flamenco, Cayo 
Coco). 

Remaks: We consider R. redferni a valid 
species, different from all similar species 
previously mentioned. In the original 
description comparison with R. pulchra 
had not been made; this latter has a more 
reticulated shell, with more numerous 
spiral ribs, the protoconch being larger 
and with more whorls; R. cancellina spec. 
nov. has a similar shell, but its proto- 
conch is larger and has 2 */4 whorls 
instead of 1 */4. R. sagraiana has a proto- 
conch with 2 */2 - 2 9/4 whorls. On the 
other hand, comparison with R. fenestrata 
is not necessary, as the shells and proto- 
conchs are rather different (see ROLÁN 
AND FERNÁNDEZ-GARCÉS, 2009b). 


amination; to Adam Baldiger (MCZ) for 
sending photographs of the type of R. 
pulchra and Amelia MacLellan (BMNH) 
for the information and the photographs 
of the types from the type material of R. 
sagraiana; to Antonio A. Monteiro for the 
English language revision. 


ROLÁN AND FERNÁNDEZ-GARCÉS: Caribbean Ríissozna of the group R. sagraiana-cancellata 


BIBLIOGRAPHY 


ABBOTT]J.T. 1974. American Seashells. Van Nos- 
trand Reinhold Company, New York. 663 
Pp 291pls: 

ADAMS C.B. 1850. Descriptions of supposed 
new species of marine shells which inhabit 
Jamaica. Contributions to Conchology, 7: 109- 
123: 

BOUCHET P. AND ROCROI J.P. 2005. Classifica- 
tion and nomenclator of gastropod families. 
Malacologia, 47(1-2): 1-357. 

CLENCH W.J. AND TURNER R. D. 1950. The west- 
ern Atlantic marine mollusks described by C. 
B. Adams. Occasional Papers on Mollusks 1(15): 
233-403. 

DESJARDIN M. 1949. Les Rissoina de l'Ile de 
Cuba. Journal de Conchyliologie, 89: 193-208, 
pIs59=0! 

DíAz MERLANO J.M. AND PUYANA HEGEDUS M. 
1994. Moluscos del Caribe Colombiano. Un ca- 
tálogo ilustrado. Colciencias-Fundación Na- 
tura-Invemar. Santafé de Bogotá. 291 pp 

ESPINOSA J. AND ORTEA J. 2002. Descripción de 
cuatro nuevas especies de la familia Rissoi- 
nidae (Mollusca: Gastropoda). Avicennia, 15: 
141-146. 

JONG K.M. DE AND COOMANS H.E. 1988. Marine 
gastropods from Curacao, Aruba and 
Bonaire. Studies on the Fauna of Curacgao and 
other Caribbean Islands, 69. 261 pp, 47 pls. 

LEE H.G. 2009. Marine shells from Northeast 
Florida. Jacksonville Shell Club, Jacksonville. 
204 pp. 

NEvILL G. 1885. Hand list of Mollusca in the In- 
dian Museum, Calcutta. Part Il. Gastropoda, 
Prosobranchia-Neurobranchia (contd.). Gov- 
ernment Printer, calcutta, x + 306 pp. 

MÓRCH O.A.L. 1875-1877. Synopsis mollusco- 
rum marinorum Indiarum occidentalium im- 
primis insularum danicarum. Malakozoolo- 
gische Blátter, 22: 142-184 [1875], 23: 45-58, 
87-143 [1876], 24: 14-66, 93-123 [1877]. 

NOwELL-USTICKE G.W. 1959. A check-liat of the 
marine shells of St. Croix and Virgin Islands. 
Christiansted, St. Croix, 90 pp. 


ORBIGNY A. D”, 1841-1853. Mollusques. In: R. de 
la Sagra, Histoire physique, politique et nature- 
lle de l'1le de Cuba. Arthus Bertrand, Paris.[Vol- 
ume 2 pp. 1-112, issued 1846 and Atlas pl. 1- 
28 issued 1842, fide Bouchet and Rocroi, 
2005]. 

PHILIPPI R.A. 1847. Testaceorum novorum cen- 
turia. Zeitschrift fúr Malakozoologie, 4: 113- 
128. 

PONDER W.F. 1985. A review pf the Genera of 
the Rissoidae (Mollusca: Mesogastropoda: 
Rissoacea). Records of the Australian Museum, 
suppl. 4: 1-221. 

REDFERN C. 2001. Bahamian seashells. A thou- 
sand Species from Abaco, Bahamas. Bahami- 
ansehlls.com Inc. Boca Raton, 280 pp, 124 

ls. 

Rios E.C. 1994. Seashells of Brazil. Fundacáo Ci- 
dade do Rio Grande, Rio Grande, 345 pp, 
113 pls. 

ROLÁN E. AND FERNÁNDEZ-GARCÉS R. 2009a. 
Description of a new species of Rissoina 
(Prosobranchia, Rissoidae) from Bermuda. 
Gloria Maris, 48 (1): 17-21. 

ROLÁN E. AND FERNÁNDEZ-GARCÉS R. 2009b. 
New information on Cuban Rissoina (Mol- 
lusca: Rissoidae) 1. Rissoina fenestrata and R. 
vanderspoeli. Novapex, 10 (2): 59-64. 

SCHWARTZ VON MOHRENSTERN G. 1860. Uber 
die Familie der Rissoiden und insbesondere 
die Gattung Rissoina. Denkschriften der Math- 
ematisch-Naturwissenschaftlichen Classe der 
Kaiserlichen Akademie der Wissenschaften, 19: 
71-188, 11 pls. 

VOKES H.E. AND VOKES E.H. 1983. Distribution 
of shallow-water marine Mollusca, Yucatan 
Peninsula, Mexico. Mesoamerican Ecology In- 
stitute Monograph 1, Middle American Re- 
search Institute Publication, 54: viii, 183 pp, 
50 pls. 

WARMKE G.L. AND ABBOTT J.T. 1961. Caribbean 
Seashells. Livingstone Publishing Co., Penn- 
sylvania. 348 pp. 


89 


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Iberus, 28 (14: 91-96, 2010 


A new Manzoniía (Gastropoda: Rissoidae) from northwes- 


tern Morocco 


Una nueva Manzonía (Gastropoda, Rissoidae) del Noroeste de 


Marruecos 


Serge GOFAS* 


Recibido el 10-11-2010. Aceptado el 30-IV-2010 


ABSTRACT 


A new species of the family Rissoidae is described from the upper slope of Northwestern 
Morocco. Some characters of this species would relate it to the genus Manzonia whereas 
others are shared with species currently assigned to the genus Alvania (subgenus Alvinia). 
Some possible diagnostic characters of Manzonia are discussed. 


RESUMEN 


Se describe una nueva especie de la familia Rissoidae, de la parte superior del talud con- 
tinental del Noroeste de Marruecos. Algunos caracteres de esta especie sugieren su inclu- 
sión en el género Manzonia, mientras otros se asemejan a especies que se clasifican 
actualmente en el género Alvania [sugénero Alvinia). Se comentan algunos caracteres 
posiblemente diagnósticos del género Manzonia. 


INTRODUCTION 


Small gastropods belonging to the 
family Rissoidae are a prominent part of 
the littoral, shelf and upper bathyal mol- 
luscan faunas in the Mediterranean, 
temperate Western Europe and are par- 
ticularly species-rich in the north-east 
Atlantic archipelagos: Canary Islands 
and Madeira (vAN AARTSEN, 1981; 
ROLÁN, 1987; MOOLENBEEK AND FABER 
1987; MOOLENBEEK AND HOENSELAAR 
1989; ROLÁN AND FERNANDES, 1990; 
AMATI, 1992; HOENSELAAR AND GOUD, 
1998; SEGERS AND SWINNEN, 2002), the 
Azores (GorFas 1990; BOUCHET AND 
WARÉN 1993; HOENSELAAR AND GOUD 
1998), and the Cape Verde Islands 
(ROLÁN 1987; MOOLENBEEK AND ROLÁN 
1988). There are currently 317 species of 


this family recognized as valid for the 
Mediterranean and Northeast Atlantic 
south to 28” N (CLEMAM database 
<http: / / www.somali.asso.fr / clemam/> 
searched April, 2010) and this inventory 
may be near completion. However, the 
species in this family are particularly 
prone to evolve towards the loss of 
planktotrophic larval development, 
therefore giving rise to “pairs” of related 
species where the non-planktotrophic 
species is derived and tends to have a 
restricted geographic range (OLIVERIO, 
1994; 1996). For this reason, more dis- 
coveries of new species are to be 
expected. 

This paper provides a description of a 
species encountered in the straits of 


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


Correspondence: sgofasuma.es 


91 


Iberus, 28 (1), 2010 


Gibraltar, presently known only from 
two specimens but definitely distinct 
from those previously known in the area. 


SYSTEMATICS 


The species is tentatively assigned to the 
genus Manzonia, and the definitive char- 
acters of Manzonía are discussed. 


Genus Manzoniía Brusina, 1870 


Type species: Turbo costatus J. Adams, 1797, by original designation (= Turbo crassus Kanmacher, 
1798; non Turbo costatus von Salis Marschlins, 1793) 


Manzonia alexandrei n. sp. (Figs. 1-7) 


Type material: Holotype MNHN 22876 (shell, 1,8 x 1.1 mm) from “Balgim” sta. DW57, and 1 
paratype MNHN 22877 (specimen, 1.5 x 1.0 mm) from a sample collected in the vicinity of strait 
of Gibraltar, from fishermen, 1971. Only known from the type specimens. 

Type locality: off NW Morocco, 35* 42” N, 06? 35” W, 548 m. 

Etymology: The species is dedicated to my son Alexandre 


Description. Shell with a moderately 
high spire, adults up to 1.8 x 1.1 mm. 
Protoconch of 1.25 convex whorls, with 
a sculpture of six rather flat, beaded 
spiral cords, narrower than the inter- 
spaces; the nodes on the cords extend 
towards the interspaces in which they 
determine faint axial folds. Teleoconch 
of 2 */4 to 2 */2 convex whorls, angu- 
lated at a distance abapically from 
suture so as to make the whorls shoul- 
dered. Axial sculpture of flexuous, 
narrow and very elevated folds, highest 
at the shoulder of whorls, fading out 
prior to reaching the suture which is not 
at all undulated, and reaching quite far 
towards the abapical surface of the body 
whorl. Spiral sculpture of very flat 
cords, broader than the interspaces, 
bearing irregular, very faint grooves 
along spiral lines forming irregular pits; 
interspaces with those grooves much 
more definite, separating ca. 15 raised 
spiral threads; the spiral cords are more 
elevated towards the abapical half of the 
body whorl, below the line prolonging 
the suture. The base of the body whorl 
with a rather strong and elevated spiral 
cord, situated very close to a small 
umbilical chink and separated from the 
rest of the whorl by a distinct spiral 
depression. Outer lip opisthocline, bent 
in its adapical part and thickened exter- 
nally by a broad rim sloping gently 


za 


towards the adjacent surface of body 
whorl; smooth inside. Inner lip rather 
thin, slightly detached from the preced- 
ing whorl except in the centre of the 
parietal area. Shell colour white. 

Remarks: This species seems to be 
living on hard substrate in rather deep- 
water, which would explain its rarity, or 
difficulty of collection. The specimen 
from the strait of Gibraltar was retrieved 
from a large stone riddled with cavities, 
together with many specimens of 
Alvanta zylensis Gofas and Warén 1982 
and other gastropods. 

The new species features a combina- 
tion of character states which are not 
found together in any of the species 
described in the eastern Atlantic or 
Mediterranean. The protoconch with 
nodose spirals differs drastically from the 
pattern seen in most species of Manzonia 
with paucispiral protoconch from Mac- 
aronesia (see MOOLENBECK AND FABER, 
1987) or from the Lusitanian seamounts 
(see GOFAS, 2007) where the cords, if any, 
are smooth. It is reminiscent of the kind 
of protoconch seen in Alvania weinkauffi 
Weinkautff, 1868 (see PONDER, 1985, fig. 
102B) and in its sibling Alvania fischeri 
(Jeffreys, 1884) but there, the nodes are 
even more separated, looking under high 
magnification like small patches aligned 
over a minute spiral thread and pasted 
over the underlying surface. 


GOFAS: A new Manzonía from nothwestern Morocco 


Figures 1-7. Manzonia alexandrei n. sp. 1: holotype from off NW Morocco, 548 m, apertural view 
(actual size 1.8 mm); 2: protoconch of the holotype; 3: detail of microsculpture of the holotype; 4: 
paratype from near the strait of Gibraltar, apertural view (actual size 1.5 mm); 5: protoconch of 


the paratype; 6, 7: detail of microsculpture of the paratype. 
Figures 1-7. Manzonia alexandrei ». sp. 1: holotipo, frente a la costa NO de Marruecos, 548 m, vista 


apertural (tamaño real 1,8 mm); 2: protoconcha del holotipo; 3: detalle de la microescultura del holo- 
tipo; 4: paratipo, cerca del estrecho de Gibraltar, vista apertural (tamaño real 1,5 mm); 5: protoconcha 
del paratipo; 6, 7: detalle de la microescultura del paratipo. 


93 


Iberus, 28 (1), 2010 


Figures 8, 9. Details of microsculpture of Manzonía, comparable to Figures 3 and 7. 8: Manzonia 
crassa (Kanmacher, 1798), specimen from Benalmádena, southern Spain; 9: Manzonia arata Gofas, 
2007, shell from Ampere seamount. 
Figuras 8, 9. Detalles de la microescultura de Manzonia, comparables con las Figuras 3 y 7. 8: Manzo- 
nia crassa (Kanmacher, 1798), ejemplar de Benalmádena, sur de España; 9: Manzonia arata Gofas, 
2007, concha del banco Ampere . 


The most similar species with respect 
to characters of the teleoconch may be 
Manzonia arata Gofas, 2007, described 
from the Ampere seamount off western 
Morocco, which shares the general 
outline and aspect of the spiral sculpture. 
The latter is nevertheless clearly different 
in having the ribs protruding against the 
suture so as to make it strongly undu- 
lated; the completely different proto- 
conch, keeled with indistinct and smooth 
spirals, may indicate that they are unre- 
lated. There is also a superficial resem- 
blance with Frigidoalvania  thalassae 
Bouchet and Warén, 1993, described from 
deep water of Bay of Biscay, but the latter 
differs in being much larger and in 
having more numerous and less raised 
axial ribs. The protoconch of E. thalassae is 
radically different, being rather conical in 
shape with faint spiral striae. 

The generic placement in Manzonia 
is backed by the general build of the 


94 


shell and the presence of a spiral depres- 
sion around the base of the body whorl. 
Nevertheless it must be stressed that the 
separation between the genera Manzo- 
nia, Alvanta s.l. and Onoba is not at all 
clearcut. Manzonta, as currently recog- 
nized, may form a monophyletic group 
around its type species Manzonia crassa. 
Its diagnostic character states (MOOLEN- 
BEEK AND FABER, 1987) include (1) the 
presence of a “duplicate” peristome 1. e. 
a strong outer rim on which the sculp- 
ture of the outer surface of the body 
whorl is usually continued, and an inner 
rim continuing the inner surface of the 
aperture, (2) the presence of a spiral de- 
pression and / or stronger spirals on the 
abapical part of body whorl, and (3) a 
peculiar microsculpture where the 
larger spirals bear spirally aligned rows 
of minute (ca. 1 um) pits whereas the in- 
tervening spaces are furnished with 
minute, raised spiral lamellae, and still 


GOFAS: A new Manzon:a from nothwestern Morocco 


more minute axial threads abutting be- 
tween these. The value of this latter 
character has been treated as doubtful 
by BOUCHET AND WARÉN (1993: 657) 

The microsculpture seen in Manzonia 
alexandrei n. sp. is not as characteristic as 
that seen in the type species and in 
many Macaronesian coastal species. It is 
nevertheless not structurally different, 
and differs from Manzonia crassa (figure 
8) in that the spiral elements in the 
grooves are more closely packed 
together and the pits are not so neatly 
demarcated. However such a structure 
can be seen to be very similar in Manzo- 
nia arata, which from other characters 
can be assumed to be phylogenetically 
related to Manzonia lusitanica Gotas 
2007, M. crispa (Watson, 1897) and to the 
type species. 

PONDER (1985: 48) advocated that 
the subgenus Alvinia Monterosato, 1884 
(with Alvania weinkauffi as type species) 


BIBLIOGRAPHY 


AMATI B. 1992. On a new species of Manzonia 
from Selvagens Islands, (Gastropoda, Pro- 
sobranchia, Rissoidae). Publicacóes Ocasionais 
da Sociedade Portuguesa de Malacología, 16: 9- 
pas 

BOUCHET P. AND WARÉN A. 1993. Revision of 
the Northeast Atlantic bathyal and abyssal 
Mesogastropoda. Bollettino Malacologico, sup- 
plement 3: 579-840. 

GOFASS. 1990. The littoral Rissoidae and Ana- 
bathridae of Sáo Miguel, Azores. Acgoreana, 
supplement: 97-134. 

GOFAS S. 2007. Rissoidae (Mollusca: Gas- 
tropoda) from northeast Atlantic sea- 
mounts. Journal of Natural History, 41 (13-16): 
779-885 

HOENSELAAR H.J. AND GOUD J. 1998. The Ri- 
ssoidae of the Cancap expeditions, I: the 
genus Alvania Risso, 1826 (Gastropoda Proso- 
branchia). Basteria, 62: 69-115. 

MOOLENBEEK R.G. AND FABER M.J. 1987. The 
Macaronesian species of the genus Manzonia. 
De Kreukel, 23 (1): 1-16, pl. 1; 23 (2-3): 23-31; 
23 (10): 166-179, pl. 2-3. 

MOOLENBEEK R.G. AND ROLÁN E. 1988. New 
species of Rissoidae from the Cape Verde Is- 
lands (Mollusca, Gastropoda) part 1. Bul- 
letin, Zoólogisch Museum, Universiteit van Am- 
sterdam, 11 (14): 121-126. 


should belong to Manzonia rather than 
to Alvania, based on shared characters 
such as the presence of a duplicated 
peristome, of stronger spiral cords on 
the abapical part and the radula with 
numerous cusps on the inner side of the 
laterals. This view is also supported by 
the fact that Alvinia has a simple trian- 
gular metapodial tentacle like Manzonia, 
and unlike Alvania s. str. (see PONDER, 
1985: 39) which has a bundle of separate 
metapodial tentacles. 

The combination of characters seen 
in Manzonía alexandrei n. sp. further sup- 
ports Ponder's view that some species 
which lack the typical Manzonia- 
microsculpture may nevertheless be 
more closely related to Manzonia than to 
Alvania s. str. More independent charac- 
ters, including those based on molecular 
sequences, are needed to assess the 
value of this morphological character to 
be held as diagnostic of Manzonia. 


MOOLENBEEK R.G. AND HOENSELAAR H.J. 1989. 
The genus Alvania on the Canary Islands and 
Madeira (Mollusca, Gastropoda) part 1. Bul- 
letin, Zoólogisch Museum, Untversiteit van Am- 
sterdam, 11 (27): 215-228. 

OLIVERIO M. 1994. Developmental vs. genetic 
variation in two Mediterranean rissoid com- 
plexes. Journal of Molluscan Studies, 60: 461- 
465. 


OLIVERIO M. 1996. Contrasting developmental 
strategies and speciation in N.E. Atlantic 
prosobranchs: preliminary analysis. En: Ori- 
gin and evolutionary radiation of the Mollusca 
(Taylor, J. ed.), Oxford University Press, pp. 
261-266. 

PONDER W. 1985. A review of the genera of the 
Rissoidae (Mollusca: Mesogastropoda: Ri- 
ssoacea). Records of the Australian Museum, 
supplement 4: 1-221. 

ROLÁN E. 1987. El género Manzonia Brusina, 
1870 en el Archipiélago de Cabo Verde. Pub- 
licagoes Ocasionais da Sociedade Portuguesa de 
Malacología, 9: 27-36. 

ROLÁN, E. 1987. Aportaciones al estudio de 
los Risoaceos de las Islas Canarias: I. Des- 
cripción de tres especies nuevas. Publi- 
cacóes Ocasionais da Sociedade Portuguesa de 
Malacología, 8: 1-4. 


95 


Iberus, 28 (1), 2010 


ROLÁN E. AND FERNANDES F. 1990. Tres nuevas VAN AARTSEN J.J. 1981. Manzonia overdiepi, a 
especies del género Manzonia (Mollusca, new marine gastropoda (Rissoidae) from Ca- 
Gastropoda) para la costa occidental de nary and Madeira Is. La Conchiglia, 15 (168- 
Africa. Publicacoes Ocasionais da Sociedade Por- 169): 6-7. 


tuguesa de Malacología, 15: 63-68. 

SEGERS W. AND SWINNEN F. 2002. Manzonia ba- 
calladoi, a new species from the Madeiran ar- 
chipiélago. Gloria Maris, 40 (6): 120-125. 


96 


O Sociedad Española de Malacología —__———  Iberus, 28 (1): 97-114, 2010 


Notes on the benthic macrofauna of Agadir Bay (Atlantic 
Morocco) 


Datos sobre la macrofauna bentónica de la Bahía de Agadir 
(Marruecos Atlántico) 


Abdellatif MOUKRIM*, José Enrique GARCÍA RASO** and Serge GOFAS** 


Recibido el 16-IV-2010. Aceptado el 30-IV-2010 


ABSTRACT 


The macrofauna collected in 25 dredge hauls from subtidal soft bottoms [6-25 m) in 
Agadir Bay, Southern Morocco, is listed. Seventy species are identified, chiefly Mollusca 
but also some decapods, echinoderms, and polychaetes. The results show four main 
assemblages with a similarity above 50%. The community of soft bottoms is quite homoge- 
neous within the bay and is comparable to the community of fine sands with Chamelea 
striatula - Mactra corallina as described by Glémarec (1969, 1973) and with the shallow 
well calibrated fine sandy bottoms (SFBC) or “terrigenous” fine sandy bottoms (Pérés and 
Picard 1964; ledoyer, 1968). Towards the deeper part there is an increase of Ophiura 
texturata together with an increase in mud content. Á more significantly different commu- 
nity is associated to bottoms with rocks in the northern part of the bay. Almost half of the 
species identified belong to the temperate European (Atlanto-Mediterranean) faunal 
province whereas one quarter belong to the tropical West African province. Despite the 
modest sampling effort, there are five species of molluscs [Nassarius goreensis, N. argen- 
teus, Cuna gambiensis, Tellina rubicincta, Pandora oblonga) new to the Moroccan fauna, 
indicating the need for a more thorough survey in the area. 


RESUMEN 


Se presenta una lista de las especies de la macrofauna recolectada en 25 dragados en 
fondos blandos sublitorales (6-25 m) de la Bahía de Agadir, Sur de Marruecos. Se identifi- 
caron 70 especies, principalmente moluscos, pero también decápodos, equinodermos y 
poliquetos. Los resultados de este estudio muestran cuatro grupos de especies con una 
similitud superior al 50%. La comunidad de fondos blandos es bastante homogénea den- 
tro de la bahía y se corresponde con la comunidad de arenas finas con Chamelea striat- 
ula - Mactra corallina descrita por Glémarec (1969, 1973) y con la de arenas finas bien 
calibradas poco profundas (SFBC) o arenas finas terrígenas [Péres and Picard 1964; 
Ledoyer, 1968). Hacia la parte más profunda se aprecia una mayor abundancia de 
Ophiura texturata, así como un incremento de la proporción de fango. La comunidad más 
diferente se encuentra en un fondo con rocas en la parte norte de la bahía. Aproximada- 
mente la mitad de las especies pertenecen a la provincia faunística europea templada 
(Atlanto-Mediterránea) y cerca de una cuarta parte a la provincia tropical del Oeste 
Africano. A pesar de lo limitado del muestreo, se citan cinco especies de moluscos nuevas 
para la fauna de Marruecos (Nassarius goreensis, N. argenteus, Cuna gambiensis, Tellina 
rubicincta, Pandora oblonga), lo que pone de manifiesto la necesidad de una prospec- 
ción más intensa en este área. 


* Laboratoire Eaux et Environnement, Faculté des Sciences, B.P. 28S, Agadir (Morocco). 
** Departamento de Biología Animal, Facultad de Ciencias, E-29071 Málaga (Spain). 


37 


Iberus, 28 (1), 2010 


INTRODUCTION 


The benthic fauna of the Atlantic 
coast of Morocco is known quite ade- 
quately for bathyal environments 
sampled by deep-sea expeditions (see 
review in SALAS, 1996 and GARCÍA RASO, 
1996) or for the intertidal communities 
accessible from the shore. The subtidal 
environments in between have received 
little attention and their species compo- 
sition remains largely unknown. Previ- 
ous data are mainly based on a sam- 
pling programme conducted in the 
years 1924-1925 with the trawler 
“Vanneau”, following the foundation of 
the Institut Scientifique Chérifien, 
Morocco's Natural History Museum in 
1923 (LIOUVILLE, 1930). However, only a 
small part of the material obtained was 
eventually studied. 

The geographical situation of the 
Bay of Agadir falls mid-way within the 
so-called “Mauretanian region” of 
EKMAN (1953). Thus, it is important to 
determine how much of its fauna 
belongs to the European temperate 
province, as does most of Atlantic 
Morocco, and how much belongs to the 
tropical West African province. The bay 
marks the end to the north of a very 
long stretch of coastline with a broad 
shallow continental shelf clad with sedi- 
ments, and bordered by long, straight 
sandy beaches exposed to heavy surf. 
Northwards, Cape Ghir sets a sharp 
physiographic limit to a predominantly 
rocky shore with a narrow platform; a 
comparable sandy shore resumes only 
much further to the north along the 
coastal plains of Bouregreg and Sebou in 
Northern Morocco. Thus, this physical 
boundary may contribute to sharpen the 
northwards boundary of any West 
African elements likely to occur in the 
area. The area of Agadir is also coinci- 
dent with a major area of upwelling 
(MITTELSTAED, 1983), which makes it 
potentially species-rich. 

The growing concern for conserva- 
tion and control of water quality has 
triggered new interest for the benthos in 
nearshore environments close to the 
large urban agglomeration of Agadir. 


98 


The purpose of this paper is to present 
some results regarding the benthic 
macrofauna (chiefly Mollusca and 
Decapoda) of the soft bottoms which 
constitute most of the bay, and to draw 
some preliminary conclusions regarding 
benthic communities and their biogeo- 
graphic setting. 


MATERIAL AND METHODS 


A set of samples (Figure 1) was col- 
lected in May 1999, using a rectangular 
dredge with an opening of 50 cm, 
geared with a net of 10 mm mesh. The 
dredge was towed by a boat at a speed 
of approximately one knot, for 15 
minutes, parallel to shore at each posi- 
tion on a transect. Three transects were 
sampled perpendicular to shore, in the 
vicinity of Oued Souss, which was the 
target area for an environmental impact 
study, and within each transect four 
samples were taken at 10, 15, 20 and 25 
m depth. Two other transects and some 
samples were taken with the same gear 
at the northern end of the beach (D), 
inside the harbour (E), and in front of 
the suburb of Anza (FE), just northwest to 
the harbour. In each of the transects D 
and E, samples were taken, at 6, 10, 15, 
20 and 25 m. Samples were sorted 
immediately upon return to the lab, and 
the animals preserved in 70” ethanol for 
further determination. 

The literature used for species iden- 
tification was not specific to the area. 
The fauna of both Western Europe and 
West Africa has been considered. 
PASTEUR HUMBERT (1962a, b), POPPE AND 
GOTO (1991, 1993) and the unpublished 
manuscript on West African Bivalvia by 
Rudo von Cosel (MNHN) were the basic 
references for the Mollusca; MONOD 
(1956), FOREST AND GANTES (1960) and 
ZARIQUEY ÁLVAREZ (1968), among others 
for the decapods, TORTONESE (1965) for 
the echinoderms. The polychaetes were 
identified by Patrick Gillet (of Institut 
d'Ecologie Appliquée, Angers, France). 

Sediment samples were taken by 
SCUBA diving along the three transects 
of Oued Souss (A: 30” 18' N, B: 30” 20” N 


MOUKRIM ET 42.: Notes on the benthic macrofauna of Agadir Bay (Atlantic Morocco) 


CES E 
S.v 20, 
P* d'Anz 
29 
Transect F 


28 24 


19 


1 
Transect Cf 22 0d g 
S.v.R hs 


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Bag ea EA] PAE JA 20 PJ JS (O CA RA O A A E DUE JE FE FE JE JE A E JA A SC E FA E (O A PA A a REO FA A IO LA A DA 


h 25 dl 
pS Transect B e , 


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4 


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7 . TA "DNS 
1 A Z 


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TransectE|* |, ese 


22 ¿ 
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22  Transect D 18 
24 : 
25 ? 
Ñ 


1 py -3 
nn —HE_ ——_ 5 5 ¿e 
TE. CA AA A 
PAS A A 


3025 


Figure 1. Map of the Bay of Agadir, with location of the sampling stations. 
Figura 1. Mapa de la bahía de Agadir, con la ubicación de las estaciones de muestreo. 


and C: 30” 22” N), in order to determine 
granulometry using a column of stan- 
dard sieves. 

PRIMER software (CLARKE AND 
WARWICK 1994, CLARKE AND GORLEY, 
2001) was used in the study of the struc- 
ture of the communities and their 
spatial variations using qualitative data 
without transformation and standard- 
ization. Possible significant differences 
between transects and depths were 
assessed using the ANOSIM routine. 
Samples were grouped according to 
their faunal composition using the Bray- 
Curtis similarity index. In the cluster the 
mode group average was selected. The 
samples were then plotted with a non- 
metric multidimensional scaling (MDS 
routine) and, to know the level of contri- 


bution of the different species, the 
SIMPER routine was applied. 


RESULTS 


The list of all collected species 
amounts to 70 (Table I). There is an 
increase in species richness with dis- 
tance from shore. Molluscs and 
decapods are the best represented 
groups, whereas polychaetes probably 
were undersampled due to the kind of 
dredge used, which skims the superfi- 
cial part of the sediment. 

The sediments are essentially fine 
sands in transects A and C (range of 
median 0.17-0.32 mm). The pelite 
content is very low nearshore and 


97 


Iberus, 28 (1), 2010 


Table I. List of species collected in the dredgings. The * denotes species shared with “sables fins a 
Venus gallina - Mactra corallina” and “sables fins envasés 4 Pharus legumen - Ophiura texturata” or 
denoted as characteristic of fine sand communities by Glemarec (1969). W: West african species. 
The abundance of species is coded (1) 1-2 specimens, (2) 3-5 specimens, (3) 6-10 specimens, (4) 


over 10 specimens. 


Transect 302 18' N Transect 302 20' N 
depth (m) AO OS 


MOLLUSCA GASTROPODA 

Bela sp. 

Bivetiella cancellata (Linné, 1758) W Za] 

Bullia miran (Bruguiére, 1792) W A eS DA 
Calyptraea chinensis (Linné, 1758) 
Clanculus kraussi (Philippi, 1846) 
Cabestana dolaria (Linné, 1767) 
Cymbivm cucumis Róding, 1798 
Epitonium turtonis (Turton, 1819) 
Euspira fusca (de Blainville, 1845) l 

Euspira pulchella (Risso, 1826) l 

Marginella glabella (Linné, 1758) W 

Mesalia varia (Kiener, 1844) W DES | 

Mitrello sp. l 

Nassarius argenteus (Marrat, 1877) W 

Nassarius elatus (Gould, 1845) W Mal vaa] 
Nassarius goreensis (von Maltzan, 1884) W 

Nassarius incrassatus (Stróm, 1768) 

Nassarius reticulatus (Linné, 1758) AO 

Nassarius vaucheri (Pallary, 1906) W l l l 
Ocenebra brevirobusta Hovart, 2000 

Philine aperta (Linné, 1758) l 
Stramonita haemastoma (Linné, 1758) 
Solatia piscatoria (Gmelin, 1791) W 

Tectonatica sagraiana (d'Orbigny, 1844) W l 
Volvarina sp. W 


=== 


MOLLUSCA BIVALVIA 

Anomia ephippium (Linné, 1758) l 

Abra alba (Wood, 1802) E Il ad: la 
Chamelea striatula (da Costa, 1778) bl | 
Corbula gibba (Olivi, 1792) 2 2 
Cuna gambiensis Nicklés, 1955 W 

Donax vittatus (da Costa, 1778) ss 2 

Dosinia lupinus (Linné, 1758) E (a | 

Ensis ensis (Linné, 1758) 
Gari fervensis (Gmelin, 1791) 
Macoma cumana (Costa, 1829) st | 

Mactra coralina (Linné, 1758) ÉS l Zap dE 


100 


MOUKRIM £7 4Z.: Notes on the benthic macrofauna of Agadir Bay (Atlantic Morocco) 


Tabla 1. Lista de especies recolectadas en los dragados. El * indica especies que se han citado para las 
comunidades denominadas “sables fins a Venus gallina - Mactra corallina” y “sables fins envasés a 
Pharus legumen - Ophiura texturata”, o indicadas como características de comunidades de arena fina 
por Glemarec (1969). W: Especies Oeste-Africanas. La abundancia de las especies esta anotada como (1) 
1-2 ejemplares, (2) 3-5 ejemplares, (3) 6-10 ejemplares, (4) más de 10 ejemplares. 


Tronsect 302 22' N North of Beach Harbour Off Anza 
IAS A IS Orba E 
2 
l l 
A E IAE AN 4] O A O E 
l l 
la l 
l 
| 
1 
l 
¡A 
| de A E 4 2 DS 
| 
l ] 
2 l 3 3 | 24 24 
74 
4 2 EA 
e AA LA IE O e ROS OS 
l 4 4 1 l 
Z 
l 
l 
1 1 l 
l (A 
2 
In 1 
157008] lia A 4 3 DA 
1 do DEL AI LL 
2 | A 
l 
l dial 1 4 aaró 
Es] 2 4 ls 
l 1 l | A 
l 
Ica l 4 
2 1 l 1 DS MA] 


101 


Iberus, 28 (1), 2010 MOUKRIM £7 AL.: Notes on the benthic macrofauna of Agadir Bay (Atlantic Morocco) 


? 
Table 1. List of species collected in the dredgings. The * denores species shared with “sables fins á Tabla 1. Lista de especies recolectadas en los dragados. El * indica especies que se han citado para las 
Venus gallina - Mactra corallina” and “sables fins envasés a Pharus legumen - Ophiura texturata”. or comunidades denominadas "sables fins a Venus gallina - Mactra corallina” y “sables fins envasés á 
denoted as characteristic of fine sand communities by Glemarec (1969). W: West african species. Pharus legumen - Ophiura texturata”, o indicadas como características de comunidades de arena fina 
The abundance of species is coded (1) 1-2 specimens, (2) 3-5 specimens, (3) 6-10 specimens, (4) por Glemarec (1969). W: Especies Oeste-Africanas. La abundancia de las especies esta anotada como (1) 
over 10 specimens. | 1-2 ejemplares, (2) 3-5 ejemplares, (3) 6-10 ejemplares, (4) más de 10 ejemplares. 

Transect 30* 18' N Transect 302 20' N | Tronsect 30* 22' N North of Beach Horbour Of Anza 

depth (m) 10 15 20 25 10 15 20 25 9 10 15 20 25 6 10 15 20 25 6-6 6 10 15 20 25 
MOLLUSCA GASTROPODA , 
Belo sp. 2 
Bivetiello concelloto (Linné, 1758) W 2 ? l 1 
Bullio miran (Bruquiere, 1792) W E PA E, E 4 3 4 2.4. 4 2 4 1 4.1 1 2 4 
Colyptroeo chinensis (Linné, 1758) ) 1 1 
Clonculus kroussi (Philippi, 1846) W | 2 1 
Cobestono doloria (Linné, 1767) W ? 1 
Cymbium cucumis Róding, 1798 W 1 | 
Epitonium turtonis (Turton, 1819) p 1 
Euspiro fusca (de Bloinville, 1845) 1 1 
Euspira pulchella (Risso, 1826) 1 ) 1 
Morginello globello (Linné, 1758) W | 1 
Mesolia varia (Kiener, 1844) W 2.4 3 1 y 25 4 4 1 1.5405 4 2 3 2 
Mitrello sp. 1 1 
Nossarius orgenteus (Morrot, 1877) W h 1 1 
Nossorivs elotus (Gould, 1845) W A | JA 2 1 3 3 110.4.4 
Nossorivs goreensis (von Maltzon, 1884) W j 2 
Nassorivs incrassatus (Stróm, 1768) 4 2 META 
Nossarius reficulatus (Linné, 1758) LA] 3.41 2 4 4 4 2 3 4 
Nossorius voucheri (Pollary, 1906) W 1 1 p 1 4 4 1 1 
Ocenebra brevirobusta Hovort, 2000 2 
Philine aperto (Linné, 1758) 1 ' 1 
Stramonita hoemastomo (Linné, 1758) 1 
Solo piscotorio (Gmeln, 1791) " PA 2 ? 
Tectonotico sagraiona (d'Orbigny, 1844) W 1 1 1.1 
Volvarina sp. W A 2 
MOLLUSCA BIVALVIA 1 
Anomia ephippivm (Limné, 1758) 1 | 1 1 1 
Abro olbo (Wood, 1802) E A A 3 1 A 4 3 WT 
Chomeleo striatula (da Costo, 1778) E 1 | a | DAA 
Corbulo gibbo (Olivi, 1792) 2 2 a 2 12 1óe-4 
Cuno gombiensis Nicklés, 1955 W | 1 
Donox vittotus (da Costo, 1778) E 2 1 1 yA 1 4 de] 
Dosinia lupinus (Linné, 1758) * ATA 1 | 1057] 2 4 1 
Ensis ensis (Linné, 1758) á » 1 1 1 1 dei 
Gori fervensis (Gmelin, 1791) ] 1 
Mocomo cumnano (Costo, 1829) 1041 1 ' 1 1 4 
Macro coralina (Lim, 1758) * 1 22 ñ 2 : ( 1 2.3 Dei 


100 101 


Iberus, 28 (1), 2010 


Table I. Continuation. 
Tabla I. Continuación 


Modiolus stultorum (Jousseaume, 1893) W (ne 


Transect 30% T8”N Transect 30% 20"N 
depth (m) O lo 200 925 A 


Nuculana bicuspidata (Gould, 1845) W 3 1 

Pandora oblonga (Sowerby, 1830) W 

Pandora inaequivalvis (Linné, 1758) 1 l 

Parvicardium exiguum (Gmelin, 1791) 

Pharus legumen (Linné, 1758) de O E, AM TON 
Phaxas pellucidus (Pennant, 1777) dx l 
Scacchia zorni van Aartsen 8. Fehr-de Wal, 1985 

Sinupharus combieri (Fischer-Piette 8. Nicklés, 1946) W 

Spisula subtruncata (da Costa, 1778) á l l 

Tellina tenvis (da Costa, 1778) Ís 1 

Tellina rubicincta Gould, 1845 W l 

Tellina fabula Gmelin, 1791 ES A MecoAN] 
Thracia papyracea (Poli, 1791) ña ] 

Thyasira flexuosa (Montagu, 1803) 2 


MOLLUSCA CEPHALOPODA 
Sepietta oweniana (d'Orbigny, 1839) 2 
Sepiola rondeleti Leach, 1817 2 


CRUSTACEA DECAPODA 

Diogenes pugilator (Roux, 1829) ñ 3 3 4 

Liocarcinus cf. holsatus (Fabricius, 1798) E LAS AAA 
Liocarcinus depurator (Linné, 1758) 

Macropodia rostrata (Linné, 1761) 2 

Philocheras trispinosus (Hailstone, 1835) á Sl 

Polybivs henslowii Leach, 1820 1 | 

Scyllarus arctus (Linné, 1758) 1 


PYCNOGONIDA 
Nymphon sp. l 


ECHINODERMATA 
Echinocardium sp. l 
Ophivra texturata Lamarck, 1816 ; 40 5d l AA 


ANNELIDA POLYCHAETA 

Cirratulus filiformis Keferstein, 1862 

Diopatra neapolitana Delle Chiaje, 1841 IA LL 
Lugia pterophora (Ehlers, 1864) 

Magellona papillicornis Miller, 1858 

Nepthys sp l 


102 


MOUKRIM £7 42.: Notes on the benthic macrofauna of Agadir Bay (Atlantic Morocco) 


Table I. Continuation. 
Tabla I. Continuación 


ranse 3 


10 


arbour z0 
CAOS OS 
Ii MD SA 
OS 
3 
lol l ¡dl 
l 
Al E PA 
3 IP 
2 
LAO l 
4 
3 
4 A AN 
2 
l 
19] 
O 
l 
la A O 
l 
280 de] 


103 


Iberus, 28 (1), 2010 MOUKRIM £7 AZ.: Notes on the benthic macrofauna of Agadir Bay (Atlantic Morocco) 


Table I. Continuation. e Table I. Continuation. 
Tabla I. Continuación | Tabla I. Continuación 
| 


Modiolus stultorum (Jousseaume, 1893) W A ' 
Trance 30r TEN Tromsed SO ZO | —Trnsed SONT Ñ Nor of Beach Harbour UA 

depth (m) 10 15 20 25 10 15 20 25 | 10 15 20 25 6 10 15 20 25 6 6» 6 10 15 20 25 

Í 

> 
Nuculona bicuspidata (Gould, 1845) W 3 1 1.41] DL AA 
Pandora oblongo (Sowerby, 1830) W » 4 
Pandora inaequivalvis (Linné, 1758) 1 1 | 1 4.43 1 
Porvicordium exiguum (Gmelin, 1791) » 3 
Phorus legumen (Linné, 1758) * IAEA IZ | A INIA 11 1 O 
Phoxos pellucidus (Pennont, 1777) ú l ) 
Scocchia zomi von Aortsen 8. Fehr-de Wal, 1985 | 1 1 
Sinuphorus combieri (FischerPiette 8. Nickles, 1946) W i 1 
Spisulo subtruncato (da Costa, 1778) si 1 1 | 201 4 1 A 
Tellino fenvis (da Costo, 1778) $ 1 ( 2 3 1423 
Tellina rubicincto Gould, 1845 W 1 P 
Tellino fobulo Gmelin, 1791 á 201 1 | 1 1 1 AA] 1 
Thracio popyracea (Poli, 1791) Ed 1 Ñ 2 2 4 
Thyosira flexuoso (Montagu, 1803) 2 ' 
MOLLUSCA CEPHALOPODA | 
Sepietta oweniana (d'Orbigny, 1839) 2 p 
Sepiolo rondeleti Leach, 1817 2 | 

, 
CRUSTACEA DECAPODA 
Diogenes pugilotor (Roux, 1829) > 3 3 4 ? 4 3 2 3 
Liocorcinus ct. holsatus (Fabricius, 1798) dl ES) LANA | MEN EA | 4 SI 2 
Liocarcinus depurator (Linné, 1758) » 2 
Mocropodia rostrato (Limné, 1761) 2 | 1 0) 
Philocheras trispinosus (Hailstone, 1835) * Sl ? PEA 
Polybivs henslowii Leach, 1820 1 1 1 al 11 1 
Seyllarus arctus (Linné, 1758) 1 E 

| 
PYCNOGONIDA ? 
Nymphon sp. 1 | 
ECHINODERMATA h 
Echinocordium sp. 1 y 1 Deal 
Ophiura texturata Lamorck, 1816 4 4 1 4 4 | 4 4 DAA 2 4 
ANNELIDA POLYCHAETA h 
Cirratulus filiformis Keferstein, 1862 Ú 1 
Diopatra neapolitana Delle Chioje, 1841 AAA MIA | 1 304 1 3 TEN VA SES 
Lugia pterophora (Ehlers, 1864) 1 
Mogellono popillicornis Miller, 1858 2 1 
Nepthys sp 1 A] 

> 
102 103 


D- A 


Iberus, 28 (1), 2010 


Figure 2 A-E. Gastropods from Agadir Bay. A: Nassarius vaucheri (Pallary, 1901), Agadir beach, 6 
m (height 12 mm); B: Nassarius argenteus (Marrat, 1877), inside harbour, 15 m (height 7.4 mm); 
C: Nassarius reticulatus (Linné, 1758), Agadir beach, 6 m (height 16 mm); D: Nassarius goreensis 
(von Maltzan, 1884), off Anza, 20 m (height 9.3 mm); E: Bela cf. zonata (Locard, 1892), Agadir 
beach, 6 m (height 10.5 mm). Figure 2 E Decapod Crustacean: Liocarcinus cf. holsatus (Fabricius, 
1798), off Oued Souss, 15 m (breadth of carapace 36 mm). 

Figura 2 A-E. Gasterópodos de la bahía de Agadir. A: Nassarius vaucheri (Pallary 1901), playa de 
Agadir, 6 m (altura 12 mm); B: Nassarius argenteus (Marrat, 1877), en el puerto, 15 m (altura 7,4 
mm); C: Nassarius reticulatus (Linné, 1758), playa de Agadir, 6 m (altura 16 mm); D: Nassarius 
goreensis (von Maltzan, 1884), frente a Anza, 20 m (altura 9,3 mm); E: Bela cf. zonata (Locard, 
1892), playa de Agadir, 6 m (altura 10,5 mm). Figura 2 E Crustáceo Decápodo: Liocarcinus cf. hol- 
satus (Fabricius, 1798), frente al Oued Souss, 15 m (ancho del caparazón 36 mm). 


increases to ca. 10% at 20 m. The values 
in transect B, under the influence of 
Oued Souss, show a higher content of 
pelites at shallower depths (up to 30% at 
10 m depth). The northernmost tran- 
sects, off the harbour and E contain also 
some rocky outcrops. 


104 


Taxonomic and  biogeographic 
notes on some species 
MOLLUSCA GASTROPODA 


Family Turritellidae 

Turritellids are represented by one 
species of Mesalia, the same one that 
extends northwards to the Ibero-Moroc- 


MOUKRIM £7 4Z.: Notes on the benthic macrofauna of Agadir Bay (Atlantic Morocco) 


YN 5 


5 ») 


Figure 3. Cuna gambiensis Nickles, 1955 from Agadir Bay, Loa “Vanneau” sta. 118 (309 40” N, 


090 55" W, 20 m). A: Outside view of the left valve of a live-collected specimen; note the hydro- 
zoan on posterior edge (actual height of shell 3.0 mm). B. inside view of the same valve with dried 
soft parts inside, note the four juveniles in prodissoconch stage incubated in the pallial cavity. C- 
D. Inside of left and right valve of another specimen (actual size 2.9 mm). 

Figura 3. Cuna gambiensis Vickles, 1955 de la bahía de Agadir, campaña del “Vanneau” est. 118 (30% 
40" N, 09 55" W 20 m). A: Vista externa de la valva izquierda de un ejemplar recolectado vivo; nótese el 
hidrozoo asentado en el borde posterior (altura de la concha 3,0 mm); B: vista interna de la misma valva 
con partes blandas desecadas; nótense los cuatro juveniles en estadio de prodisoconcha, incubados en la 
cavidad paleal. C-D. interior de las valvas izquierda y derecha de otro ejemplar (altura 2,9 mm). 


can Gulf and to the Alboran Sea and 
southwards to Senegal. There has been 
considerable confusion regarding the 
taxonomy of this species, often misiden- 
tified as the Senegalese endemic Mesalia 
brevialis (Lamarck, 1822) (e.g. PASTEUR- 
HUMBERT, 1962a: 39). We agree with 
PALLARY (1900) and MARCHE-MARCHAD 
(1981) in considering that this is a dis- 
tinct species, correctly named Mesalia 
varia (Kiener, 1844). 


Family Nassariidae 

This family is represented in our 
material by seven species, of which two 
are new to the Moroccan fauna. Nas- 
sarids are well represented in West 
Africa (see Adam and Knudsen, 1984) 
and are mostly scavengers. Bullia miran 
is one of the largest species and is a con- 
spicuous element of the assemblage. It is 
a West African species, already men- 
tioned from Agadir Bay by PALLARY 
(1920: 37, as Dorsanum miran) and 
PASTEUR-HUMBERT (1962a). 

Nassarius vaucheri (Fig. 2A) and Nas- 
sarius argenteus (Fig. 2B) are two very 
similar species, the former endemic to 
the Ibero-Moroccan gulf and the latter 
West African, which could be suspected 


of being geographical subspecies one of 
the other. The occurrence at Agadir of N. 
argenteus documents a range extension 
into Morocco, and supports their treat- 
ment as separate species. 

Nassarius goreensis (Fig. 2D) is also a 
West African species new to the Moroc- 
can fauna. Besides this occurrence, there 
are occasional findings as far north as 
Asilah and Tangiers (36” N; specimens 
in Muséum National  d'Histoire 
Naturelle, Paris, collected by S.G.). 

Nassarius reticulatus (Fig. 2C) con- 
versely, is a member of the temperate 
West European fauna which reaches 
here its southern limit. 


Family Muricidae 

Ocenebra brevirobusta is a Moroccan 
endemic, formerly identified as Ocenebra 
torosa (Lamarck, 1816) or Ocenebra eri- 
naceus (Linné, 1758) (see PALLARY, 1920; 
PASTEUR-HUMBERT, 1962a). HOUART 
(2000) described it as a new species, 
from material collected at Essaouira, 
some 100 km north of Agadir. 


Family Turridae 


There is one species of Bela (Fig. 2E) 
which resembles both morphologically 


1109 


Iberus, 28 (1), 2010 


Figure 4 A-F. Bivalves from Agadir Bay. A, B: Macoma cumana (Costa, 1829), right valve, South of 
Oued Souss, 15 m (length 20 mm); C, D: Tellina rubicincta Gould, 1845, right valve, off Anza, 25 
m. (length 22 mm); E, F: Donax vittatus (da Costa, 1778), left and right valves, off Anza, 10 m 


(length 21 mm). 


Figura 4 A-E Bivalvos de la bahía de Agadir. A, B: Macoma cumana (Costa, 1829), valva derecha, 
sur de Oued Souss, 15 m (longitud 20 mm); C, D: Tellina rubicincta Gould, 1845, valva derecha, 
frente a Anza, 25 m. (longitud 22 mm); E, F: Donax vittatus (da Costa, 1778), valvas izquierda y 


derecha, frente a Anza, 10 m (length 21 mm). 


and in habitat the Mediterranean species 
Bela zonata (Locard, 1892), but there is 
such confusion in the species-level tax- 
onomy of European Bela that we prefer 
not to venture a specific name. 


MOLLUSCA, BIVALVIA 

Family Condylocardiidae 

Cuna gambiensis (Fig. 3) is a small 
bivalve living in algal turf and thus 
linked to hard bottom. A few specimens 


106 


were collected in the transect off Anza, 
but there are many specimens from 
Agadir collected in the years 1920 by G. 
Dollfus with R/V “Vanneau” (unpub- 
lished material in Muséum National 
d'Histoire Naturelle, Paris). This is a 
tropical West African species, new to the 
Moroccan fauna. Live-taken specimens 
showed a small hydrozoan growing on 
the posterior edge of the valve, and, like 
other species of the Condylocardiidae 


MOUKRIM £7 42.: Notes on the benthic macrofauna of Agadir Bay (Atlantic Morocco) 


Figure 5 A-D. Pandora spp. from Agadir Bay. A-B: Pandora inaequivalvis (Linné, 1758), Agadir 


beach, 6 m (length 24 mm); C-D: Pandora oblonga (Sowerby, 1830), Agadir beach, 6 m (length 14 


mm). 


Figure 5 A-D. Pandora spp. de la bahía de Agadir. A-B: Pandora inaequivalvis (Linné, 1758), Agadir 
beach, 6 m (length 24 mm); C-D: Pandora oblonga (Sowerby, 1830), Agadir beach, 6 m (length 14 mm). 


(SALAS AND ROLÁN, 1990; SALAS AND 
COSEL, 1991) it was found brooding 
juveniles. 


Family Pharidae 

The sand-dwelling razor shells are 
represented by four species. It is note- 
worthy that the genus Ensis, represented 
by three species in comparable sandy 
beaches of the Ibero-Moroccan Gulf, is 
here only represented by the smaller 
(and usually rarer) Ensis ensis. The West 
African Sinupharus combieri was already 
recorded (BELLON-HUMBERT, GLÉMAREC 
AND GOFEAS, 1975), but the record of the 
European Phaxas pellucidus is a range 
extension, the known southern limit 
being Tangiers. 


Family Tellinidae 

Macoma cumana (Fig. 4A-B) has its 
type locality in the Western Mediter- 
ranean and is reported to have a West 
African range (von Cosel, unpublished 
data), but its distribution is quite puz- 
Zling, being replaced in the Alboran Sea 
by the very similar Macoma melo 
(Sowerby, 1866). The West African 


species Tellina rubicincta (Fig. 4C-D) is 
new to the Moroccan fauna, the previ- 
ously known northern limit being in 
Mauritania (DAUTZENBERG, 1910). It 
strongly resembles the European T. dis- 
torta Poli, 1791 but is larger, slenderer 
and with more distinct comarginal 
lamellae towards the posterior end. 


Family Donacidae 

Donax vittatus was quite frequent in 
the sandy bottom of the shallower part 
of the transects. The specimens from 
Agadir (Fig. 4E-F) resemble the Mediter- 
ranean D. venustus (Poli, 1791) in having 
sometimes three broad radial bands, but 
should be assigned to D. vittatus on the 
basis of having striae which are not 
restricted to the part of the shell 
between the posterior angle and the 
posterior margin. The relationships of 
these species are still unsettled (TIRADO 
AND SALAS, 1999). 


Family Veneridae 

BACKELJAU, BOUCHET, GOFAS AND DE 
BRUYN (1994) have shown that the 
Atlantic populations formerly called 


107 


Iberus, 28 (1), 2010 


Venus gallina correspond to Chamelea 
striatula, whereas the real Chamelea 
gallina is restricted to the Mediterranean 
and southwestern Iberian Peninsula. 
This is an important point because the 
Atlantic species is the eponym of 
“sables fins a Venus gallina-Mactra coral- 
lina” of GLEMAREC (1969), to be cor- 
rected as Chamelea striatula-Mactra coral- 
lina. lt is noteworthy that C. striatula is 
displaced towards more muddy facies 
where both species are sympatric, 
whereas, on the European West coast, it 
occupies facies of fine sand which are 
similar to the biotope of C. gallina in the 
Mediterranean. 


Family Pandoridae 

The shallow samples of the Agadir 
beach yield, in the same habitat, both 
the European Pandora inaequivalvis (Fig. 
3D), here at its southern limit (range 
extension, previously known only from 
Essaouira northwards) and the West 
African P. oblonga (Fig. 3E), which is 
here recorded as new to the Moroccan 
fauna. Pandora oblonga is smaller, has a 
distinctly more convex left valve and 
more concave right valve, and is less 
rostrated posteriorly. 


CRUSTACEA, DECAPODA 

The genus Diogenes (Diogenidae) is 
represented in West Africa by a species 
complex (FOREST, 1961), but examina- 
tion of the very large sample from 
Agadir bay suggests that only one 
species, D. pugilator, is present. The 
genus Liocarcinus (Portunidae) is repre- 
sented by two species, the widespread 
L. depurator, and another one of doubtful 
identification (Fig. 2F), tentatively 
assigned to L. cf. holsatus, although the 
contour of the carapace resembles some- 
what the Mediterranean L. vernalis 
(Risso, 1826). Nevertheless, specimens 
from Agadir lack the characteristic 
velvety surface of the latter. Material 
from intermediate localities on the 
Moroccan coast is needed to decide 
whether this is a geographical variation 
of one of these two European species, or 
if a third species must be considered. 
However, this group of species is in 


108 


revision (d'Udekem d'Acoz personal 
communication). 


ANNELIDA: POLYCHAETA 

There is a species of the genus Diopa- 
tra (family Onuphidae) which is abun- 
dant in all the transects of the bay. This 
may be D. marocensis, described from 
similar bottoms south of Safi (PAXTON, 
FADLAOUI AND LECHAPT, 1995). The 
number of Polychaete species in the 
samples is low, but this may be a bias 
due to our sampling gear which does 
not dig deeply into the sediments. 


Characterization of the assem- 
blages 

The results of the aggregation and 
ordination analyses (MDS) using pres- 
ence-absence data are shown in Figure 
6, in which four main groups with a 
similarity above 50% are defined (stress 
value 0.18). The analysis of the grouped 
“stations” by transect shows significant 
difference (ANOSIM global test, p = 
0.003), with highest values between the 
F and the A, B and C transects (pairwise 
tests, p = 0.008). In this way, the material 
collected off Anza and inside the 
harbour contain, in addition to the other 
assemblages, several species normally 
associated with rocky  substrates. 
Among these are Clanculus kraussi and 
Nassarius incrassatus, found off Anza, 
and Stramonita haemastoma, Ocenebra bre- 
virobusta, and Cymatium doliarium col- 
lected inside the harbour. The harbour 
appears comparatively species-rich (34 
species), which can result both from the 
presence of hard substrates along the 
piers, and from the shelter of the piers 
which maintains low wave action. 

Along the depth gradient a low sig- 
nificance differences have been found 
(ANOSIM Global test, p = 0,04). 

A semi-quantitative analysis shows 
similar results (by transects: global test, 
p = 0.002), with highest values between 
the F and the A, B and C transects (pair- 
wise tests, p = 0.008); by depth: global 
test, p = 0,01). 

The average similarity within the dif- 
ferent transects was around 40.1 (A) — 
53.5% (E) (SIMPER) and between 9 (B) to 


MOUKRIM £7 ALz.: Notes on the benthic macrofauna of Agadir Bay (Atlantic Morocco) 


soft a 


Stress: 0.18 
A25 


mixed with ES 
hard bottoms 


Figure 6. Multi-dimensional scaling (MDS) of the samples. The similarity matrix is based on pres- 
ence/absence of the species and the Bray-Curtis similarity coefficient. The contours highlight 
groups of samples which cluster with a similarity coefficient above 50%. The samples are identified 
by a letter (A: radial of 30918"N, B: radial of 30920'N, C: radial of 30%22'N, D: North of Beach; 


E: Harbour, E: off Anza), and their depth. 


Figura 6. Ordenación multidimensional (MDS) de las muestras. La matriz de similitud esta basada en 
la presencialausencia de las especies y en el coeficiente de similitud de Bray-Curtis. Los contornos señalan 
grupos de muestras que se juntan con un coeficiente superior a 50%. Las muestras están identidficadas 
por una letra (A: radial de 3018'N, B: radial de 3020'N, C: radial de 3022'N, D: Norte de la 
playa; E: Puerto, EF: frente a Anza) y por su profundidad. 


18 (F) species were necessary to get an 
accumulative contribution of 90-92 %. By 
depths, 10 (20m) to 14 (10 m) discrimi- 
nating species are necessary to get a sim- 
ilar accumulative contribution (higher 
than 90%) with an average similarity be- 
tween 35.3 (25 m) - 53.9% (15 m). 


DISCUSSION 


The species collected include some 
well known as characteristic of fine 
sands. The assemblages on soft bottoms 
are quite comparable to those reported 


by GLEMAREC (1969, 1973) from “sable 
fin a Venus gallina - Mactra corallina” and 
“sables fins envasés a Pharus legumen - 
Ophiura texturata”, where many species 
(denoted by * on Table I) are shared. 
Some species collected here are 
restricted to rather low latitudes so that 
they have not been mentioned in any of 
the classical works on benthic bionomy 
Among these, Bullia miran, Nassarius 
elatus and Mesalia varia are dominant 
and should qualify as characteristic of 
the fine sand or slightly muddy fine 
sand communities at the latitude of 
Agadir. The equivalence with the 


109 


Iberus, 28 (1), 2010 


Mediterranean community of “sables 
fins bien calibrés” SFBC (PÉRES AND 
PICARD, 1964, AUGIER, 1982) or of “ter- 
rigenous” fine sandy bottoms (Ledoyer, 
1968) is not straightforward because of 
these biogeographic differences, but 
many of the characteristic Mediter- 
ranean species are here replaced in the 
Atlantic Morocco by ecological vicari- 
ants: Nassarius mutabilis (Linné, 1758) by 
Bullia miran, Nassarius  pygmaeus 
(Lamarck, 1822) by N. goreensis, Chame- 
lea gallina (Linné, 1758) by C. striatula, 
Tellina pulchella Lamarck, 1818 by T. 
rubicincta and Neverita josephina Risso, 
1826 by other Naticids. The same 
happens with the decapods, because the 
characteristic species of the SFBC com- 
munity of the Mediterranean and south- 
ern Spain are Diogenes pugllator, 
Philocheras trispinosus and Liocarcinus 
vernalis (PÉRES AND PICARD 1964, 
LEDOYER, 1968, (GARCÍA MUÑOZ, 
MANJÓN-CABEZA AND GARCÍA RASO, 
2008) the latter replaced by L. cf. holsatus 
in this study. Such habitats are listed 
with very little detail, as sublittoral 
sands (code A5.2) in the European 
Nature Information System (EUNIS) 
classification  (DAvIES, MoOss AND 
O'HILL, 2004). 

The more differentiated assemblage 
found is that associated with sand and 
rocky substrate, which also shows the 
highest specific richness as usual in 
other areas (GARCÍA MUÑOZ ET AL, 
2008). 

The assemblages also respond to a 
bathymetric gradient: towards the 
deeper part of the transects, there is a 
set of species that prefer slightly muddy 
sands, the most noteworthy is Ophiura 
texturata. Nevertheless, the community 
is quite homogeneous along the shore, 


even where facing the rocky area at 
Anza. This can be explained by the very 
flat topography of the sublittoral part of 
the bay, in which the sandy bottom 
swamps the rocky outcrops even north 
of the harbour. Also, the mobility of the 
macrofauna explains that some species 
are found across several neighbouring 
transects on the same kind of bottoms. 
The same happens in  sublittoral 
bottoms of the Alborán Sea, where the 
decapod assemblage of coarse sandy 
bottoms (with high values of organic 
material) presented practically the same 
discriminating species as the neighbour- 
ing assemblage of muddy fine sandy 
bottoms, but with different quantitative 
contribution of the species (GARCÍA 
MUÑOZ ET AL, 2008). 

A macrobenthic assemblage from 
fine sand bottoms in a depth range 16- 
40 m off Sidi Boulbra (31% 52 to 31* 58' 
N, some 160 km northwards, in the 
province of Safi), was described by FAD- 
LAQUI (1994) in an environmental study 
for a conventional thermic power plant. 
This was assigned to “sédiments fin 
plus ou moins envasés a Abra alba- 
Diopatra brevicirris”, referring to a classi- 
fication of benthic communities derived 
from THORSON (1957), but is basically 
similar to the assemblages referred 
herein. We suspect that the unidentified 
“terebrid” (FADLAOUL, 1994: 58) reported 
there with a frequency of 80% is Bullia 
miran, since there are no terebrids in 
Morocco and shells of Bullia are mor- 
phologically very terebrid-like. 


Biogeographic notes 

Among the 70 species collected, 20 
(28.5 %) have a predominantly tropical 
distribution and can be considered as 
belonging to the West African zoogeo- 


(Right page) Figure 7. Plot of the latitudinal ranges of the molluscan species known from Agadir 
Bay, ordered by southern (0) and thence by northern (ME) distributional limits; ranges in the 
southern hemisphere not shown and not taken into account for ordering. 

(Página derecha) Figura 7. Representación de la extensión latitudinal de las especies conocidas en la 
Bahía de Agadir, ordenadas por límite sur (0) y luego por limite norte (WM) de distribución; no se mues- 
tran, ni se tienen en cuenta para la ordenación, las distribucuiones en el hemisferio sur. 


110 


80 


BA IA UA PU AA UTA 


MS 

o 

0) 

E 

O - 

> = North Cape 

9 ' 

S | 

= | 

+ p 

< | 

o +S. Norway 

as | 

ES 

5 E 

< a] Channel 

E. 2 | 

3 1) Bayof 

S D |Biscay 

E 5 | 

O O) 

z O 

E LS) eze 

E = | Gibraltar 
Y) | 

Q 2 | 

E £ 

3 sE 

6 O 

Q = 

== 59) 

a == 

o = a CapBlanc 

E 

Z 

OS Dakar 

E 

kh 

z 

5) 

e) 

2 


10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 
Species 


1 


Iberus, 28 (1), 2010 


graphic region. This percentage is consis- 
tent with the general characterization of 
the area, which belongs essentially to the 
Atlanto-Mediterranean province of 
EKMAN (1953). A survey of the shelled 
Mollusca, for which good distributional 
data are available, has been made using 
the same unpublished dataset as in 
GOFAS (1999) for latitudinal ranges (Fig. 
7). There are ca. 150 species of molluscs 
known from Agadir bay or from a stretch 
of coastline which includes Agadir 
(PALLARY, 1920, PASTEUR-HUMBERT, 
1962a,b and this study). Among these, 85 
(53%) are temperate Lusitanian species 
which reach their southern limits at 
Agadir (30 N) or at Cap Blanc (21*” N), 
whereas 41 (26%) are West African 
species which reach their northern limit 
at Agadir, within Morocco or in the Ibero- 
Moroccan Gulf (up to 36” N). The remain- 
der are species with a broad temperate 
and tropical range, some of which (e.g. 
Venus verrucosa Linné, 1758) extend to 
both hemispheres. 

The tropical element, however, is 
overrepresented among the Mollusca 
from the soft bottoms (i.e. the three 
southernmost transects). Among the 40 
species collected, 15 (37.5%) belong to 
the West African fauna. This trend is 
particularly noteworthy among the gas- 
tropods, there being 10 of 16 species 
(62%) which are tropical. This may be a 
consequence of the overwhelming 
occurrence of soft bottoms along the 
tropical West African coast, to which a 
large number of species have become 
adapted. Thus, soft bottom littoral com- 
munities have a larger pool to draw 
from to the South than to the North. 

There are five species (Nassarius 
goreensis, N. argenteus, Cuna gambiensis, 
Tellina rubicincta, Pandora oblonga) which 
are new to the Atlantic coast of Morocco 
and also to the area covered by 


BIBLIOGRAPHY 


ADAM W. AND KNUDSEN J. 1984. Révision des 
Nassariidae (Mollusca: Gastropoda Proso- 
branchia) de l'Afrique Occidentale. Bulletin 
de l'Institut Royal des Sciences Naturelles de 
Belgique, 55 (9): 1-95, 5 pl. 


112 


CLEMAM (Check List of European 
Marine Mollusca) and ERMS (European 
Register of Marine Species, Costello et 
al., eds. 2002) checklists. This is consid- 
erable taking into account the compara- 
tively small sampling effort and high- 
lights the need for a more thorough 
faunal survey of this area. The number 
of recorded continental shelf species 
(156 Mollusca) is also very low com- 
pared to other areas (Roscoff, in the 
Western Entrance of the English 
Channel: 420 species; Strait of Gibraltar, 
ca. 1000 species), and it can be specu- 
lated that an accurate sampling should 
at least duplicate this number. 


AKNOWLEDGEMENTS 


This work would not have been pos- 
sible without RAMSA, the authority in 
charge of water supply and sewage for 
the Agadir urban area. We owe to the 
collaboration of the authorities of the 
Harbour of Agadir (ODEP) the possibil- 
ity to use the towboat “Argoub” and the 
pilot boat for the dredgings, and extend 
our special thanks to the Captain for his 
care in manoevring a very small dredge 
with a very powerful boat. Special 
thanks are also expressed to the 
Muséum National d'Histoire Naturelle 
(Laboratoire BIMM), Paris, of which the 
extensive library and collections were 
indispensable to this work. We are 
indebted to Dr. Rudo von Cosel 
(MNHN) for useful advice about the 
bivalves and to Dr. Patrick Gillet (Uni- 
versité d'Angers) for collaborating with 
the determination of polychaete species. 
We also thank Dr. Carmen Salas (Uni- 
versity of Málaga) for her interesting 
remarks and Pr. B. Moumni (Faculty of 
Science, Tangiers) for help with the 
granulometry of sediment. 


AUGIER H. 1982. Inventatre et classification des bio- 
cenoses marines bentiques de la Mediterranée. 
Conseil de l'Europe, Collection Sauvegarde 
de la Nature, 25 Estrasburgo, 59 pp. 


MOUKRIM £7 42.: Notes on the benthic macrofauna of Agadir Bay (Atlantic Morocco) 


BACKELJAU T., BOUCHET P., GOFAS S. AND DE 
BRUYN L. 1994. Genetic variation, systemat- 
ics and distribution of the venerid clam 
Chamelea gallina. Journal of the Marine Biolog- 
ical Association ofthe United Kingdom, 74: 211- 
PS 

BELLON-HUMBERT C., GLÉMAREC M. AND. GO- 
FAS S. 1975. Cultellus combieri Fischer-Piette 
et Nickles (Mollusca, Bivalvia), espece nou- 
velle pour la faune atlantique marocaine. 
Bulletin de la Société des Sciences naturelles et 
Physiques du Maroc, 55: 91-93. 

CLEMAM. Unitas Malacologica Check List of 
European Marine Mollusca. Internet site [cur- 
rent URL <http:/ /www.somali.asso.fr/ 
clemam/>, last searched April 2010]. 

CLARKE K.R. AND GORLEY R.N. 2001. PRIMER 
v5: User Manual/Tutorial. Primer-E, Plymouth. 
91 pp. and computer software package. 

CLARKE K. AND WARWICK R. 1994. Change in ma- 
rine communities: An approach to statistical 
analysis and interpretation. Natural Environ- 
ment Research Council, Plymouth, U. K., 1- 
150. 

COSTELLO M., EMBLOW C. AND WHITE R. (EDS.) 
2002. European Register of Marine Species. 
A checklist to the marine species in Europe and 
a bibliography of guides to their identification. 
Muséum National d'Histoire Naturelle, Paris, 
collection Patrimoines naturels, 50: 463 pp. 

DAUTZENBERG P. 1910. Contribution a la faune 
malacologique de l' Afrique Occidentale. Actes 
de la Société Linnéenne de Bordeaux, 64: 47-220, 
pl. 1-4. 

DavIÉS C.E., Moss D. AND O'HILL 2004. EUNIS 
Habitat classification revised 2004. European 
Topic Centre on Nature Protection and Bio- 
diversity, Paris. 307 p. 

EKMAN 5. 1953. Zoogeography of the sea. Sidgwick 
and Jackson, London, xiv + 417 pp. 

FADLAOUI 5. 1994. Contribution a l'étude de la 
structure et du fonctionnement des peuplements 
benthiques de la cóte atlantique marocaine (Ré- 
gion de Sidi Boulbra). Doctoral Thesis, Uni- 
versité de Rennes-I, 1-188. 

FOREST J. 1961. Paguridés de l'Afrique Occi- 
dentale. Atlantide Reports, 6: 203-250. 

FOREST J. AND GANTES H. 1960. Sur une collec- 
tion de Crustacés Décapodes marcheurs du 
Maroc. Bulletin du Muséum National d'His- 
toire Naturelle, 2? sér, 32 (4): 346-358. 

GARCÍA RASO J.E. 1996. Crustacea Decapoda 
(Excl, Sergestidae) from Ibero-moroccan wa- 
ters. Results of BALGIM-84 Expedition. Bul- 
letin of Marine Science, 58 (3): 730-752. 

GARCÍA MUÑOZ J.E., MANJÓN-CABEZA M.E. 
AND GARCÍA RASO J.E. 2008. Decapod crus- 
tacean assemblages from littoral bottoms of 
the Alborán Sea (Spain, west Mediterranean 
Sea): spatial and temporal variability. Scien- 
tia Marina, 72 (3): 437-449 


GLÉMAREC M. 1969. Les peuplements benthiques 
du plateau continental Nord-Gascogne. These de 
Doctorat d'Etat, Faculté des Sciences de Paris, 
167 pp. 

GLÉMAREC M. 1973. The benthic communities 
of the European North Atlantic continental 
shelf. Oceanography and Marine Biology, an 
Annual Review, 11: 263-289. 

GOFAS S. 1999. Marine molluscs with a very 
small range in the Strait of Gibraltar. Diver- 
sity and Distributions, 4: 255-266. 

HOUART R. 2000. New species of Muricidae 
(Gastropoda) from the northeastern Atlantic 
and the Mediterranean sea. Zoosystema, 22 (3): 
459-469. 

LEDOYER M. 1968. Ecologie de la faune vagile 
des biotopes méditerranéens accessibles en 
scaphandre autonome. IV. Synthese de lé- 
tude écologique. Recueil des Travaux de la 
Station Marine d'Endoume, 60 (Bulletin 44): 
1-295. 

LIOUVILLE J. 1930. Liste provisotre des stations du 
“Vanneau”, de la “Dédaigneuse” et du “Las- 
signy” sur la cóte atlantique du Maroc, de 1923 
a 1929 (Liste révisée par MM. Rob-Ph. Dollfus, 
du Muséum, et le lieutenant de vaisseau Jean 
Spitz, commandant l'unité marine a Casablanca). 
Paris, Larose, 24 pp. 

MARCHE-MARCHAD 1. 1981. Notes sur des rep- 
resentants du genre Mesalia Gray, 1847 (Gas- 
tropoda, Turritellidae) trouvés dans la 
Méditerranée. Bolllettino Malacologico, 17 (3- 
4): 41-48. 

MITTELSTAED E. 1983. The upwelling area off 
Northwest Africa. A description of phe- 
nomena related to coastal upwelling. Progress 
in Oceanography, 12: 307-331. 

MONOD T. 1956. Hippidea et Brachyura Ouest- 
Africains. Mémoires de l'Institut Francais 
d'Afrique Notre, 45: 1-674. 

PALLARY P. 1900. Coquilles marines du littoral 
du Département d'Oran. Journal de Conchyli- 
ologie, 48: 211-422, pl. 6-8 

PALLARY P. 1920. Exploration Scientifique du 
Maroc. Malacologie. Larose, Paris, and Insti- 
tut Scientifique Chérifien, Rabat. 109 p., 1 
pl., 1 map. 

PASTEUR-HUMBERT C. 1962a. Les mollusques 
marins  testacés. du. Maroc... 1. Les 
Gastéropodes. Travaux de l'Institut Scientifique 
Chérifien, sér. Zoologie, 23: 1- 245. 

PASTEUR-HUMBERT C. 1962b. Les mollusques 
marins testacés du Maroc. 2. Les Lamelli- 
branches et les Scaphopodes. Travaux de l'In- 
stitut Scientifique Chérifien, sér. Zoologie, 28: 
1-184. 

PAXTON H., FADLAOUI S. AND LECHAPT J.P. 
1995. Diopatra marocensis, a new brooding 
species of Onuphidae (Annelida: Polychaeta). 
Journal of the Marine Biological Association of 
the United Kingdom, 75, 949-955. 


US 


Iberus, 28 (1), 2010 


PÉRES J.M. AND PICARD J. 1964. Nouveau manuel 
de bionomie benthique de la Mer Méditer- 
ranée. Recueil des Travaux de la Station Marine 
d'Endoume, 47 (Bulletin 31): 1-137. 

POPPE G.T. AND GOTO Y. 1991. European 
Seashells. Volume 1 (Polyplacophora, Caudo- 
foveata, Solenogastra, Gastropoda), 352 pp. 
Christa Hemmen, Wiesbaden. 

PoPPE G.T. AND'GOTO Y. 1993. European 
Seashells. Volume II (Scaphopoda, Bivalvia, 
Cephalopoda), 221 pp. Christa Hemmen, Wies- 
baden. 

SALAS C. 1996. The Bivalves from off the South- 
ern Iberian Peninsula collected by the 
FAUNA I and BALGIM expeditions. Halio- 
tis, 25: 33-100. 

SALAS C. AND ROLÁN E. 1990. Four new species 
of Condylocardiidae from Cape Verde Is- 
lands. Bulletin du Muséum National d'Histoire 
Naturelle, (4e sér.) 12A (2): 349-363 

SALAS C. AND COSEL R. VON 1991. Taxonomy 
of tropical West African bivalves III. Four 
new species of Condylocardiidae from the 
continental shelf. Bulletin du Muséum Na- 
tional d' Histoire Naturelle (4e sér.) 13A (3-4): 
263-281 


114 


THORSON G. 1957. Bottom communities (sub- 
littoral or shallow shelf). In Hedgpeth, J.W. 
(Ed.): Treatise on Marine Ecology and Palaeoe- 
cology: 461-534. Geological Society of Amer- 
ica. 

TIRADO C. ANDSALASC. 1999. Reproduction of 
Donax venustus Poli, 1795, Donax semistriatus 
Poli, 1795 and intermediate morphotypes 
(Bivalvia: Donacidae) in the littoral of Málaga 
(Southern Spain). P.S.Z.N. Marine Ecology, 
20 (2): 111-130. 

TORTONESE E. 1965. Fauna d'Italia, VI. Echino- 
dermata. Calderini, Bologna. 422 pp. 

ZARIQUEY ÁLVAREZ R. 1968. Crustáceos decápo- 
dos ibéricos. Investigación Pesquera, 32: i-xv, 
1-510. 


NORMAS DE PUBLICACIÓN 


e La revista Iberus publica artículos de fondo, notas y monografías que versen sobre cualquiera de los aspectos 
relacionados con la Malacología. Se entiende por artículo un trabajo de investigación de más de 5 páginas de 
texto, incluidas láminas, gráficos y tablas. Las notas son trabajos de menor extensión. Las monografías son tra- 
bajos sobre un tema único, de extensión superior a las 50 páginas de la revista y que serán publicadas, si 
procede, como un suplemento de Iberus. Los autores interesados en publicar monografías deberán ponerse 
previamente en contacto con el Editor de Publicaciones. Se entiende que el contenido de los manuscritos no ha 
sido publicado, ni enviado simultáneamente a otra revista para su consideración. 

e Los manuscritos, así como toda la correspondencia relacionada con los mismos, deberán ser remitidos a: 
Serge Gofas, Editor de Publicaciones, Departamento de Biología Animal, Universidad de Málaga, Campus de 
Teatinos, s/n, 29071, Málaga, España y / o al correo electrónico <sgofasWuma.es>. 

e El texto del trabajo podrá estar redactado en español, inglés, italiano, francés o portugués. 

e Los artículos, notas y monografías deberán presentarse sobre DIN A-4, por una sola cara a doble espacio 
(incluyendo referencias, pies de figura y tablas), con al menos 3 centímetros de margen por cada lado. Los tra- 
bajos se entregarán por triplicado (original y dos copias) y se incluirá una versión en un CD, o bien remitida 
por correo electrónico, utilizando procesadores de texto en sus versiones de corrientes de Windows (pero no 
en el formato .docx de Word 2007, el habitual de Windows Vista) o Macintosh. En caso de autoría compartida, 
uno de los autores deberá hacerse responsable de toda la correspondencia. 

e Junto con el trabajo debe incluirse una lista de al menos 4 posibles revisores del mismo, sin perjuicio de los 
que el propio Editor considere oportunos. 

e Los manuscritos se presentarán de acuerdo al siguiente esquema: 

Primera página. Deberá incluir un título conciso, pero sugerente del contenido del trabajo, así como una traduc- 
ción al inglés del mismo (si el artículo no está escrito en inglés). Cuando sea preciso, deberá incluir referencia a 
familia o taxones superiores. A continuación figurarán, por este orden, el nombre y apellidos completos del 
autor o autores, las direcciones completas de los mismos, y un resumen del trabajo y su traducción al inglés. 
Dicho resumen deberá sintetizar, en conjunción con el título, tanto los resultados como las conclusiones del 
artículo; se sugiere una extensión de 100 a 200 palabras. 

Páginas siguientes. Incluirán el resto del artículo, que debe dividirse en secciones precedidas por breves encabe- 
zamientos. Siempre que sea posible, se recomienda seguir el siguiente esquema: Introducción, Material y 
métodos, Resultados, Discusión, Conclusiones, Agradecimientos y Bibliografía. Si se emplean abreviaturas no 
habituales en el texto, deberán indicarse tras el apartado de Material y Métodos. 

e Las notas breves deberán presentarse de la misma forma, pero sin resumen. 

e Deberán evitarse notas a pie de página y referencias cruzadas. Deberán respetarse estrictamente los Códigos 
Internacionales de Nomenclatura Zoológica y Botánica (últimas ediciones). Cuando un taxón aparezca por 
primera vez deberá citarse su autor y fecha de su descripción. En el caso de artículos sistemáticos, cuando se 
den las sinonimias de los taxones, éstas deberán citarse COMPLETAS, incluyendo en forma abreviada la 
publicación donde fueron descritas, y la localidad tipo si es conocida entre corchetes, según el siguiente 
esquema (préstese especial cuidado a la puntuación): 

Dendrodoris limbata (Cuvier, 1804) 

Sinonimias 

Doris limbata Cuvier, 1804, Ann. Mus. Hist. Nat. Paris, 4 (24): 468-469 [Localidad tipo: Marsella]. 

Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop.-Car., 10: 275. 

Dichas referencias no deberán incluirse en la lista de Bibliografía si es la única vez que se nombran en el texto. 
Si se incluyen una lista completa de referencias de un taxón inmediatamente tras éste, deberá seguirse el 
mismo esquema (sin incluir en Bibliografía las referencias que no se mencionen en otro lugar del texto). 

e Sólo los nombres en latín y los de taxones genéricos y específicos deberán llevar subrayado sencillo o prefe- 
rentemente ir en cursiva. En ningún caso deberá escribirse una palabra totalmente en letras mayúsculas, ni 
siquiera el Título. Las unidades a utilizar deberán pertenecer al Sistema Métrico Decimal, junto con sus correc- 
tas abreviaturas. En artículos escritos en castellano, en los números decimales sepárese la parte entera de la 
decimal por una coma inferior (,), NUNCA por un punto (.) o coma superior ('). 

e Las referencias bibliográficas irán en el texto con minúsculas o versalitas: Fretter y Graham (1962) o FRETTER 
Y GRAHAM (1962). Si son más de dos autores se deberán citar todos la primera vez que aparecen en el texto 
[Smith, Jones y Brown (1970)] empleándose et al. las siguientes veces [Smith et al. (1970)]. Si un autor ha publi- 
cado más de un trabajo en un año se citarán con letras: (Davis, 1989a; Davis, 1989b). No deberá emplearse op. 
cit. La lista de referencias deberá incluir todas las citas del texto y sólo éstas, ordenadas alfabéticamente. Se 
citarán los nombres de todos los autores de cada referencia, sea cual sea su número. Los nombres de los 
autores deberán escribirse, en letras minúsculas o Versalitas. No deberán incluirse referencias a autores 
cuando éstos aparezcan en el texto exclusivamente como autoridades de un taxón. Los nombres de las publica- 


ciones periódicas deberán aparecer COMPLETOS, no abreviados. Cuando se citen libros, dése el título, editor, 
lugar de publicación, n* de edición si no es la primera y número total de páginas. Deberán evitarse referencias 
a Tesis Doctorales u otros documentos inéditos de difícil consulta. Síganse los siguientes ejemplos (préstese 
atención a la puntuación): 

Fretter V. y Graham A. 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp. 

Ponder W.F. 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. En Ponder, W.F. 
(Ed.): Prosobranch Phylogeny. Malacological Review, suppl. 4: 129-166. 

Ros J. 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Mis- 
celánea Zoológica, 3 (5): 21-51. 

e Las gráficas e ilustraciones deberán ser originales y presentarse preferentemente en formato electrónico al 
formato de caja de la revista o proporcional a éste. Este formato es de 57 mm (una columna) o 121 mm (dos) de 
anchura y hasta 196 mm de altura, si bien se recomienda utilizar el formato a dos columnas. Es importante que 
todas las figuras sean remitidas en su formato original (por ejemplo, las fotografías en .j¡pg de alta calidad o .tif, 
las gráficas en hojas de cálculo Excel o documentos de CorelDraw), puesto que las ilustraciones insertadas en 
el manuscrito WORD son inservibles en la fase de imprenta. Las imágenes digitales deben ser formateadas en 
su tamaño de impresión con una resolución mínima de 300 ppp para imágenes en color o escala de grises y de 
600 ppp para las de blanco y negro. Considérese la reducción que será necesaria a la hora de decidir el tamaño 
de las escalas o letras en las figuras, que no deberán bajar de los 2 mm. En figuras compuestas, cada parte 
deberá etiquetarse con letras mayúsculas, el resto de las letras deberán ser minúsculas. Las escalas de dibujos y 
fotografías deberán ser gráficas, utilizando unidades del sistema métrico decimal; no deberán hacerse referen- 
cias a los aumentos de una determinada ilustración, ya que éstos cambian con la reducción. En su caso, se 
recomienda la utilización de mapas con proyección UTM. Cada figura, gráfica o ilustración deberá presentarse 
en hojas separadas y con numeración arábiga (1, 2, 3,...), sin separar “Figuras” y “Láminas”. Los pies de 
figura, en una hoja aparte, deberán acompañarse de su traducción al inglés (o español si el inglés es la lengua 
del trabajo). Utilícese el esquema siguiente: 

Figura 1. Neodoris carvi. A: animal desplazándose; B: detalle de un rinóforo; C: branquia. 

Las abreviaturas empleadas en las ilustraciones deberán incluirse en los pies de figura. 

Los autores interesados en incluir láminas en color deberán consultar con el editor y sufragar el sobrecoste con 
una contribución de 30 € por página. Por lo demás, deberán ajustarse a los mismos requisitos indicados para 
las figuras. 

Si se pretende enviar gráficas o ilustraciones en impresión de papel es imprescindible presentar originales de 
buena calidad. Las imágenes en semitonos deben estar bien contrastadas y ajustarse al tamaño definitivo de 
impresión; al componer fotografías sobre una hoja, procúrese que los espacios entre ellas sean regulares y que 
estén debidamente alineadas. Téngase en cuenta que incluir fotografías de distinto contraste en una misma 
página conlleva una pobre reproducción final. Las gráficas de ordenador deberán imprimirse con impresora 
láser sobre papel de buena calidad. 

e Las Tablas se presentarán en hojas separadas, siempre con numeración romana (1, II, II1...). Las leyendas se 
incluirán en una hoja aparte acompañándose de una traducción al inglés. Deberán evitarse las tablas particu- 
larmente complejas. Se recomienda reducir el número y extensión de ilustraciones, láminas o tablas al mínimo 
necesario. 

e Los artículos que no se ajusten a las normas de publicación serán devueltos al autor con las indicaciones de 
los cambios necesarios. 

e El Comité Editorial comunicará al autor responsable del trabajo la fecha de recepción del trabajo y la fecha 
de envío a revisión. Cada original recibido será sometido a revisión por al menos dos investigadores. El 
Comité Editorial, a la vista de los informes de los revisores decidirá sobre la aceptación o no de cada manus- 
crito. El autor recibirá en cada caso copia de los comentarios de los revisores sobre su artículo. En caso de 
aceptación, el mismo Comité Editorial, si lo considera conveniente, podrá solicitar a los autores otras modifica- 
ciones que considere oportunas. Si el trabajo es aceptado, el autor deberá enviar una copia impresa del mismo 
corregida, acompañada por una versión en un CD, utilizando procesadores de texto en sus versiones corrien- 
tes de Windows (pero no el formato .docx generado por Word 2007, el habitual de Windows Vista) o Macin- 
tosh. La fecha de aceptación figurará en el artículo publicado. 

e Las pruebas de imprenta serán enviadas por correo electrónico al autor responsable, exclusivamente para la 
corrección de erratas, y deberán ser devueltas en un plazo máximo de una semana. Se recomienda prestar 
especial atención en la corrección de las pruebas. 

e De cada trabajo se entregarán gratuitamente 50 separatas, además de una versión electrónica en formato 
«pdf. Aquellos autores que deseen un número mayor, deberán hacerlo constar al devolver las pruebas de 
imprenta, y nunca posteriormente. El coste de las separatas adicionales será cargado al autor. 


INSTRUCTIONS TO AUTHORS 


* Iberus publishes research papers, notes and monographs devoted to the various aspects of Malacology. 
Papers are manuscripts of more than 5 typed pages, including figures and tables. Notes are shorter papers. 
Monographs should exceed 50 pages of the final periodical, and may be published as Supplements. Authors 
wishing to publish monographs should contact the Editor. Manuscripts are considered on the understanding 
that their contents have not been published or simultaneously submitted for publication elsewhere. 

+ Manuscripts and correspondence regarding editorial matters must be sent to: Serge Gofas, Editor de Publica- 
ciones, Departamento de Biología Animal, Universidad de Málaga, Campus de Teatinos, s/n, 29071, Málaga, 
Spain and / or to the e-mail <sgofasuma.es>. 

e Manuscripts may be written in Spanish, English, Italian, French or Portuguese. 

e Manuscripts must be typed double spaced (including the references, figure captions and tables) on one side 
on A-4 (297x210 mm) with margins of at least 3 cm. An original and two copies must be submitted, together 
with a CD or e-mail remittance containing the article written with a current Windows (but not .docx format 
generated by Word 2007, mainly used with Windows Vista) or Macintosh word processor. When a paper has 
joint authorship, one author must accept responsibility for all correspondence. 

* The authors must include a list of at least 4 possible referees; the Editor can choose any others if appropriate. 

e Papers should conform the following layout: 

First page. This must include a concise but informative title, with mention of family of higher taxon when 
appropriate, and its Spanish translation. It will be followed by all authors” names and surnames, their full 
address(es), an abstract (and its Spanish translation) not exceeding 200 words which summarizes not only con- 
tents but results and conclusions. 

Following pages. These should content the rest of the paper, divided into sections under short headings. When- 
ever possible the text should be arranged as follows: Introduction, Material and methods, Results, Discussion, 
Conclusions, Acknowledgements and References. Unusual abbreviations used in the text must be grouped in 
one alphabetic sequence after the Material and methods section. 

e Notes should follow the same layout, without the abstract. 

e Footnotes and cross-references must be avoided. The International Codes of Zoological and Botanical 
Nomenclature must be strictly followed. The first mention in the text of any taxon must be followed by its 
authority including the year. In systematic papers, when synonyms of a taxon are given, they must be cited IN 
FULL, including the periodical, in an abbreviate form, where they were described, and the type localities in 
square brackets when known. Follow this example (please note the punctuation): 

Dendrodoris limbata (Cuvier, 1804) 

Synonyms 

Doris limbata Cuvier, 1804, Ann. Mus. Hist. Nat. 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. If a 
full list of references of the taxon is to be given immediately below it, the same layout should be followed (also 
excluding from the Bibliography list those which are not cited elsewhere). 

Only Latin words and names of genera and species should be underlined once or be given in italics. No word 
must be written in UPPER CASE LETTERS. SI units are to be used, together with their appropriate symbols. In 
Spanish manuscripts, decimal numbers must be separated with a comma (,), NEVER with a point (.) or upper 
comma ('). 

e References in the text should be written in small letters or Small capitals: Fretter and Graham (1962) or 
FRETTER AND GRAHAM (1962). The first mention in the text of a paper with more than two authors must 
include all of them [Smith, Jones and Brown (1970)], thereafter use et al. [Smith et al. (1970)]. If an author has 
published more than one paper per year, refer to them with letters: (Davis, 1989a; Davis, 1989b). Avoid op. cit. 

The references in the reference list should be in alphabetical order and 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 tax- 
onomic name. Titles of periodicals must be given IN FULL, not abbreviated. For books, give the title, name of 
publisher, place of publication, indication of edition if not the first and total number of pages. Keep references 
to doctoral theses or any other unpublished documents to an absolute minimum. See the following examples 
(please note the punctuation): 

Fretter V. and Graham A. 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp. 

Ponder W.F. 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. In Ponder W.F. 
(Ed.): Prosobranch Phylogeny. Malacological Review, suppl. 4: 129-166. 

Ros J. 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Mis- 
celánea Zoológica, 3 (5): 21-51. 


e Figures must be original and provided preferably in electronic format and adjusted to page format and 
column size. These should be one column (57 mm) or two columns (121 mm) wide and up 196 mm high, or be 
proportional to these sizes. Two columns format is recommended. It is essential that all figures be supplied in 
their original format (e.g. photographs as high-grade .jpg or as .tif files, graphs as Excel spreadsheets or Corel- 
Draw files), as the files inserted into WORD documents cannot be used for printing. Digital images must be 
given their final printing size with a resolution at least 300 dpi for colour and halftones, and at least 600 dpi for 
black / white. 

Take into account possible reduction in lettering drawings; final lettering must be at least 2 mm high. In com- 
posite drawings, each figure should be given a capital letter; additional lettering should be in lower-case 
letters. A scale line, labelled with SÍ units, must be used to indicate size; magnification ratio must be avoided 
as it may be changed during printing. UTM maps are to be used if necessary. Figures must be submitted on 
separate sheets, and numbered with consecutive Arabic numbers (1, 2, 3,...), without separating “Plates” and 
“Figures”. Legends for Figures must be typed in numerical order on a separate sheet, and a Spanish translation 
must be included. Follow this example (please note the punctuation): 

Figure 1. Neodoris carvi. A: animal crawling; B: rhinophore; C: gills. 

If abbreviations are to be used in illustrations, they should be included in the figure captions. 

Authors wishing to publish illustrations in colour should consult with the editor and will be charged a contri- 
bution of 30 € per page. They should otherwise follow the same standards as black and white prints. 

If the authors want to send Figures in printed format, it is essential to supply good quality originals. Half-tone 
images must be of good contrast, and should be submitted in the final printing size. When mounting pho- 
tographs in a block, ensure spacers are of uniform width. Remember that grouping photographs of varied con- 
trast results in poor reproduction. Computer graphics must be printed on high quality white paper with a 
laser printer. 

e Tables must be numbered with Roman numbers (1, II, IIL...) and each typed on a separate sheet. Headings 
should be typed on a separate sheet, together with their English translation. Complex tables should be 
avoided. As a general rule, keep the number and extension of illustrations and tables as reduced as possible. 

e Manuscripts that do not conform to these instructions will be returned for correction before reviewing. 

e Authors submitting manuscripts will receive an acknowledgement of receipt, including receipt date, and the 
date the manuscript was sent for reviewing. Each manuscript will be critically evaluated by at least two refer- 
ees. Based of these evaluations, the Editorial Board will decide on acceptance or rejection. Anyway, authors 
will receive a copy of the referees” comments. If a manuscript is accepted, the Editorial Board may indicate 
additional changes if desirable. Acceptable manuscripts will be returned to the author for consideration of 
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TUTION LIBRARIES 


miii 


1 5455 


ÍNDICE 
Iberus 28 (1) 2010 


DuoIs C. Er MARTÍNEZ-ORTÍ A. Reproduction du mollusque envahisseur Corbicula fuminea 
(O. E. Múller, 1774) (Bivalvia: Corbiculidae) et recrutement et distribution de ses juvé- 
niles dans un canal de la région toulousaine (France) 

Reproducción del molusco invasor Corbicula fluminea (O. E Múller, 1774) (Bivalvia: Corbi- 
culidae) y reclutamiento y distribución de sus juveniles en un canal la región de Toulouse 
(ARANCEL O O a 1-11 


BEJEGA GARCÍA V., GONZÁLEZ GÓMEZ DE AGUERO E. Y FERNÁNDEZ RODRÍGUEZ C. La Arque- 
- omalacología: una introducción al estudio de los restos de moluscos recuperados en yaci- 
mientos arqueológicos 
Archaeomalacology: an introduction to the analysis of'shellfish remains from archaeological 
SES AS AE MN. O di 13-22 
GOFAS S. Y OLIVER J.D. Las especies del género Chauvetia (Gastropoda, Neogastropoda, Buccini- 
dae) del área ibero-marroquí, con descripción de cuatro especies nuevas 
The species of the genus Chauvetia (Gastropoda, Neogastropoda, Buccinidae) in the Ibero- 
moroccan area, with the description of four new species ..ooonniccocaciccccacno 23-60 
GUERRA A., PIERCE G.J., SANTOS, M?,B., GONZÁLEZ A.F, HERNÁNDEZ-MILIAN G., PORTEIRO 
C. AND PATIÑO B. Record of the largest specimen of neon flying squid Ommastrephes bar- 
tramil (Cephalopoda: Ommastrephidae) 
Registro del mayor ejemplar de pota saltadora Ommastrephes bartramii (Cephalopoda: 
Onmastreprrda o eL. 61-66 
HOLYOAK G.A. AND HOLYOAK, D.T. A new species of Candidula (Gastropoda, Hygromiidae) 
from central Portugal 


Una nueva especie de Candidula (Gastropoda, Hygromiidae) de Portugal .......... 67-72 
ROLÁN E. AND ZETTLER M.L. A new species of Gibbula (Mollusca, Archaegastopoda) from 

Namibia A 

Una nueva especie de Gibbula (Mollusca, Archaegastopoda) de Namibia ........... 73-78 


ROLÁN E. AND RAÚL FERNÁNDEZ-GARCÉS R. New information on the Caribbean Rissoina (Gas- 
tropoda, Rissoidae) of the group R. sagraiana-cancellata with the description of a new 


species 
Nueva información sobre las Rissoina del Caribe (Gastropoda, Rissoidae) del grupo de R. 
sagralana-cancellata, con la descripción de una nueva especie ooo 79-89 


GOFAS S. Á new Manzonia (Gastropoda; Rissoidae) from nothwestern Morocco 
Una nueva Mañzonia (| Gastropoda, Rissoidae) del Noroeste de Marruecos .......... 91-96 
MOUKRIM A., GARCÍA RAso J.E. AND GOFAS S. Notes on the benthic macrofauna of Agadir Bay 
(Atlantic Moroceo) 
Datos sobre la macrofauna bentónica de la Bahía de Agadir (Marruecos Atlántico) ... 97-114 


ISSN 0212-3010 


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do! 
“Jberus 


Vol. 28 (2) 


REVISTA DE LA 
SOCIEDAD ESPAÑOLA 
DE MALACOLOGÍA 


Oviedo, diciembre 2010 


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Revista de la 
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REVISTA DE LA 
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Vol. 28 (2) Oviedo, diciembre 2010 


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SOCIEDAD ESPAÑOLA DE MALACOLOGÍA 


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Dep. Leg. B-43072-81 

ISSN 0212-3010 

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O Sociedad Española de Malacología 


Iberus, 28 (2): 1-4, 2010 


Haedropleura ryalli, a new species from Sáo Tomé Island 


(Gastropoda, Turridae) 


Haedropleura ryalli, mueva especie de la isla de Sáo Tomé 


(Gastropoda, Turridae) 


Juan HORRO*, Sandro GORI** and Emilio ROLÁN*** 


Recibido el 10-11-2010. Aceptado el 31-VIIL-2010 


ABSTRACT 


A new species of the genus Haedropleura Bucquoy, Dautzenberg and Dollfus, 1883 from 
the island of Súo Tomé, West Africa, is described and figured. 


RESUMEN 


Se describe e ilustra una nueva especie del género Haedropleura Bucquoy, Dautzenberg 
y Dollfus, 1883 recolectada en la isla de Sáo Tomé, Africa occidental. 


INTRODUCTION 


The genus Haedropleura Bucquoy, 
Dautzenberg and Dollfus, 1883 is men- 
tioned in POWELL (1966) who describes 
the shell, operculum and radula and 
refers that this genus has species from 
the European Miocene and Pliocene and 
also Recent from Europe and West 
Africa down to South Africa. 

The introduction of scuba diving as 
a sampling technique in recent times 
along several areas of the West African 
coast where it had scarcely been used 
before has led to the availability of 
material from waters deeper than previ- 
ously studied ones, bringing a number 
of interesting new species to the atten- 
tion of malacologists. That happened in 
particular in Sáo Tomé Island, a place 
characterised by the abundance of 
endemic species. 


While studying the material collected 
by the second author by scuba diving in 
that island, the authors have identified a 
new species of Haedropleura which is 
described and illustrated in this paper. 


Abbreviations: 


MNHM Muséum national d'Histoire 
naturelle, Paris 

MNCN Museo Nacional de Ciencias 
Naturales, Madrid 

MNHN Muséum national d' Histoire 
naturelle, Paris 

ZMB Zoologisches Museum, Berlin 

CJH collection Juan Horro, Vigo 

CPR collection of Peter Ryall, Maria 
Rain 

CSG collection Sandro Gori, Livorno 

s shell collected empty 


* Montero Rios 30-39, 36201 Vigo, Spain; juanhorroCtelefonica.net 
** Via Semesi, 7, 57123, Livorno, Italy; sandrogoriCfastwebnet.it 
*** Museo de Historia Natural, Campus Universitario Sur, 15782 Santiago de Compostela, Spain; 


erolanGemiliorolan.com 


Iberus, 28 (2), 2010 


TAXONOMIC PART 


Family TURRIDAE Swainson, 1840 
Subfamily CRASSISPIRINAE Morrison, 1966 
Genus Haedropleura Bucquoy, Dautzenberg and Dollfus, 1883 


Type species (o. d.): Pleurotoma septangularis Montagu, 1803 


Haedropleura ryalli spec. nov. (Figs. 1-7) 


Type material: Holotype (Figs. 1-2) in MNHN (22883). One paratype (Figs. 3, 4) in MNCN 
(15.05/53578, 1 s) from type locality. Other paratypes from Minerio Reef, Sáo Tomé, 00* 23” 
01.6” N, 067 46' 22.8” E, 41 m, in the following collections: MHNS (1 s), CSG (4 s), CJH (1 s), 


CPR (2 s). 


Other material examined: 1 fragment, Minerio, Sáo Tomé (MHNS). 
Type locality: Lagoa Azul “Fundao”, 36 m, Sáo Tomé. 
Etymology: Named after our colleague and friend Peter Ryall, who was the first to call our atten- 


tion to the singularity of these shells. 


Description: Shell (Figs. 1-6) fusiform 
rhomboid, elongate and solid with a 
high spire. Protoconch (Fig. 7) with 
about 1 */4 whorls, with a diameter 
between 500 and 650 um, placed in an 
oblique position, the nucleus not situ- 
ated at the apex. 

Teleoconch of the holotype with 4 Y 
convex whorls, each one having 7-8 
prominent and thick axial ribs which go 
from suture to suture without any sub- 
sutural rib or smooth area, and are 
mostly aligned with those of the follow- 
ing whorl. Ribs becoming thinner and 
somewhat s-shaped on the last whorl. 
Suture deep and sinuous. Last whorl 
representing 65/70% of total shell 
height. Spiral sculpture formed by very 
fine and numerous striae on the whole 
surface, which pass over the axial ribs, 
extend down to the base and cover also 
the outer lip. Aperture oval elongate, 
with an evident parietal callus and 
smooth columella; siphonal canal short 
and wide; outer lip very much thick- 
ened. Background coloration light 
cream, with a dark brown subsutural 
band below which there are irregular 
and interrupted spiral lines of the same 
colour; in the middle of the body whorl 
these lines tend to form another broad 
band; lower base and mouth almost 
white. 

Dimensions: The holotype is 5.7 mm; 
the paratypes measure from 5.6 to 6.2 mm. 


Animal: Not studied. 

Distribution: Only known from the 
two mentioned locations in Sáo Tomé 
Island, from which we suppose it to be 
endemic. So far it has not been found in 
the closeby Príncipe Island, although 
material from this island available for 
study, from similar depths, has been 
scarce so far. 

Remarks: Although the generic place- 
ment of this new species in the genus 
Haedropleura must be considered a tenta- 
tive assignation, due to the lack of soft 
parts, it seems clear from a conchologi- 
cal perspective, because it presents a 
shape and micro sculpture typical of 
this genus. 

Specially after the introduction of 
the genus Anacithara Hedley, 1922, for 
west african fauna (HORRO, RYALL AND 
ROLÁN, 2010), we are aware of the close 
relationship between these two genera. 
These can only be certainly separated 
using radular characters, according to 
KILBURN (1994) with whom we agree 
this point. It must be noted that H. ryalli 
shows the spiral sculpture of finer and 
numerous spiral striae which is typical 
of Haedropleura, instead of more distant 
incised grooves present in Anacithara 
and that it also presents a larger and dif- 
ferent protoconch. 

It must also be remembered that 
genus Bellaspira Conrad, 1868 has some- 
times been considered as a senior 


HORRO ET 4L.: Haedropleura ryalli, a new species from Sáo Tomé Island 


Figures 1-7. Haedropleura ryalli spec. nov. 1, 2: holotype, 5.7 mm, Lagoa Azul (MNHN); 3, 4: 
paratype, 5.7 mm, Lagoa Azul (MNCN); 5: paratype, 6.1 mm, Minerio (CSG); 6: paratype, 5.8 
mm, Minerio (CPR); 7: protoconch of the holotype. 

Figuras 1-7. Haedropleura ryalli spec. nov. 1, 2: holotipo, 5,7 mm, Lagoa Azul (MNHN), 3, 4: 
paratipo, 5,7 mm, Lagoa Azul (MNCN); 5: paratipo, 6,1 mm, Minerio (CSG); 6: paratipo, 5,8 mm, 
Minerio (CPR); 7: protoconcha del holotipo. 


synonym of Haedropleura, but GIAN- showing conchological and radular dif- 
NUZZI-SAVELLI AND PUSATERI (1986) ferences between both genera and con- 
have already solved this question, cluding that Bellaspira species is 


Iberus, 28 (2), 2010 


restricted to the Caribbean and Panamic 
provinces, while the appropriate genus 
for European species is Haedropleura. 
There is great confusion regarding the 
genus Haedropleura in Eastern Atlantic 
and Mediterranean waters and a general 
revision of the genus is very much 
needed, especially for West African 
waters. Recently MICALI (2010) referring 
only to Mediterranean species, shows 
three different ones and calls them H. sep- 
tangularis, H. secalina (Philippi, 1844) and 
H. flexicosta Monterosato, 1884. It is not 
the aim of this paper to start on such a 
revision, which, at least in Atlantic 
waters, would include more species and 
should include information on the types. 


ACKNOWLEDGEMENTS 


We are grateful to the Department 
of Genetics of the University of Vigo 
for their help with photography, and to 


BIBLIOGRAPHY 


GIANNUZIESAVELLI R. AND PUSATERI F. 1986. 
Ripristino validitá del taxon generico Hae- 
dropleura Monterosato in B.D.D., 1883 (Mol- 
lusca: Gastropoda). Lavori, Societa Italiana di 
Malacología, 22, Atti Congreso Palermo 13-16 
sett 1984: 163-168. 

HORRO J., RYALL P. AND ROLÁN E. 2010. 
Anacithara (Conoidea, Turridae) a new genus 
to West Africa. Gloria Maris, 49 (1): 14-22. 

KILBURN R.N. 1988. Turridae (Mollusca: Gas- 
tropoda) of southern Africa and Mozam- 
bique, Part 4. Drilliinae, Crassispirinae and 
Strictispirinae. Annals of the Natal Museum, 29 
(1): 167-320. 


However, we must state that Haedropleura 
ryalli sp. nov. is easy to separate from all 
other European and West African Hae- 
dropleura species due to its unique proto- 
conch. This does not appear on any other 
species of this genus, even in those with 
paucispiral protoconchs such as H. 
secalina and H. flexicosta mentioned by 
MICALI (2010), or such as the one figured 
by NORDSIECK (1977: 75, plate 1, n* 8) 
under the genus name Bellaspira, which 
are clearly larger, with a more flattened 
top and does not show the typical posi- 
tion of the nucleus of H. ryalli. The con- 
stant pattern of H. ryalli which is not 
found in specimens from other locations 
also confirms its valid specific status. 


Dr. S. Gofas, editor of Iberus who 
made important suggestions for this 


paper. 


KILBURN R.N. 1994. Turridae [s. 1.] (Mollusca: 
Gastropoda) of southern Africa and Mozam- 
bique. Part. 7. Subfamily Crassispirinae, sec- 
tion 2. Annals of the Natal Museum, 35: 177-228. 

MICAaLI P. 2010. Nota sul genere Haedropleura 
B.B.D., 1883 nel Mediterraneo. Malacología 
Mostra Mondiale, 67: 3-5. 

NORDSIECK E. 1977. The Turridae of the European 
Seas. Rome, La Piramide. 131 pp. 

POwELL A.W.B. 1966. The Molluscan families 
Speightiidae and Turridae. Bulletin of the 
Auckland Institute and Museum, 5: 1-184, 23 pls. 


O Sociedad Española de Malacología —__—_—_——T—  Iberus, 28 (2): 5-21, 2010 


Ultrastructural study of oogenesis in the African mussel, 


Perna perna (Bivalvia: Mytilidae) 


Estudio ultraestructural de la ovogénesis en el mejillón africano, 


Perna perna (Bivalvia: Mytilidae) 


Soumya BENOMAR*, Oum Keltoum BELHSEN**, Michel MATHIEU** 
and Abdellatif MOUKRIM* 


Recibido el 15-IV-2010. Aceptado el 7-X-2010 


ABSTRACT 


Transmission electron microscopy of female gonads in Perna perna (Mytilidae) was carried 
out to study the fine structure of the different developmental stages during oogenesis and 
to examine the functions of the follicular cells. The gonad is composed of acini, each con- 
taining oogoniae and oocytes surrounded by follicular cells, and of an interacinar connec- 
tive tissue with vesicular cells. Groups of primary oogoniae, surmounted by secondary 
oogoniae, were found along the inner wall of each acinus. Previtellogenesis of oocytes 
was characterized by a great increase of these cells in volume, the accumulation of numer- 
ous organelles, and the formation of the first yolk granules at the end of this phase. The 
period of vitellogenesis involved both autosynthetic and heterosynthetic pathways, and 
was marked by the accumulation of cortical granules and of yolk granules. Numerous lipid 
droplets (two types), several inclusions resulting from the heterosynthetic uptake of exoge- 
nous substances by pinocytosis, and droplets showing two types of electron-lucent materi- 
als in the cytoplasm of mature oocytes could also be observed. In the atretic ovocytes, a 
vacuolization progressively developed in their cytoplasm, with the subsequent rupture of 
the vitelline membrane and the release of oocyte remnants in the lumen of the acinus. At 
the end of vitellogenesis, the follicular cells became detached from the oocyte and con- 
tained numerous lipid droplets and glycogen inclusions. In P. perna, the formation of 
female gametes was mostly similar to the oogenesis described in other species of bivalvia. 
The only difference concerned the composition of the yolk in the mature oocyte, as it was 
constituted of several endogenous substances (lipoproteins mainly) and also of exogenous 
materials intaken by the oocyte via pinocytosis. 


RESUMEN 


Las gónadas femeninas en Perna perna (Mytilidae) se observaron por microscopía elec- 
trónica de transmisión para estudiar la estructura fina de las diferentes etapas de desar- 
rollo durante la ovogénesis y examinar las funciones de las células foliculares. La gónada 
está formada por acinos, cada uno de ellos conteniendo oogonias y ovocitos rodeados 
por células foliculares, y por un tejido conectivo interacinar con células vesiculares. Gru- 
pos de oogonias primarias, a los que se sobreponen oogonias secundarias, se encuentran 
a lo largo de la pared interna de cada acino. La previtelogénesis de los ovocitos se carac- 
terizó por un gran aumento de volumen en estas células, por la acumulación de 


* Laboratory Aquatic Ecosystems : Marine and Continental Field, Biology Department, Sciences Faculty, Ibn 
Zohr University, BP 8106, 80000, Agadir, Morocco. e-mail: moukrimCuniv-ibnzohr.ac.ma. 
** Laboratory Marine Biology and Biotechnology, 1. B. B. A., Caen University, France. 


Iberus, 28 (2), 2010 


numerosos orgánulos y por la formación de los primeros gránulos de vitelo al final de esta 
fase. El período de vitelogénesis implica ambas vías autosintética y heterosintética y se 
caracteriza por la acumulación de gránulos corticales y de los gránulos de vitelo. En el 
citoplasma de los ovocitos maduros, se observaron también numerosas gotitas de lípidos 
(de dos tipos), varias inclusiones resultando de la absorción heterosintética de sustancias 
exógenas por pinocitosis, así como gotitas con dos tipos de materiales electrón lúcidas. 
En los ovocitos atrésicos, una vacuolización se desarrolló progresivamente en su cito- 
plasma, con la posterior ruptura de la membrana vitelina y la liberación de los restos de 
ovocitos en el lumen de los acinos. Al final de la vitelogénesis, las células foliculares se 
desprendían del ovocito y contenían numerosas gotitas de lípidos y inclusiones de 
glucógeno. En P. perna, la formación de gametos femeninos fue muy similar a la ovogéne- 
sis descrita en otras especies de bivalvos. La única diferencia se refiere a la composición 
del vitelo en el ovocito maduro, ya que está constituido por varias sustancias endógenas 
(lipoproteínas, principalmente), así como por materiales exógenos absorbidos por el ovoc- 


ito, mediante pinocitosis. 


INTRODUCTION 


The African mussel: Perna perna, is a 
worldwide species (BERRY, 1978; HICKS , 
TUNNELL AND MCMAHON,  2001a). 
However, the studies performed on the 
biology of this mussel only concerned its 
reproductive cycles, as there was an 
intraspecific variability in the reproduc- 
tion. Indeed, in the South African popula- 
tions of P. perna, two main spawning 
periods between April and October, and 
several minor and scarcer spawning 
events after October were reported by 
BERRY (1978) and LAsiaK (1986). By con- 
trast, in other Perna populations from the 
same country, SCHURINK AND GRIFFITHS 
(1991) found a single prolonged spawn- 
ing period along the year, with spawning 
activity every month. In the Gulf of 
Mexico, three spawning periods with one 
extended and two discrete events in 
spring or summer were described by 
Hicks, TUNNELL AND MCMAHON (2001) 
and Hicks, MCMAHON AND INGRAO 
(2001). In view of this variability in repro- 
ductive cycles, it was useful to study 
gametogenesis in local populations of P. 
perna. 

As P. perna lives along the southern 
Atlantic coasts of Morocco, a research 
programme was carried out to deter- 
mine the spawning episodes of these 
local mussels and to analyse the charac- 
teristics of their gametogenesis. A first 
histological study (ID HALLA, BOUHAIMI, 


ZEKHNINL, NARBONNE, MATHIEU AND 
MOUKRIM, 1997) demonstrated a single 
spawning period along the year, with a 
major event in spring, and this work 
was completed by ultrastructural 
studies on spermatogenesis in the same 
Moroccan population of P. perna 
(BENOMAR, BELHSEN, GOUX, MATHIEU 
AND MOUKRIM, 2007). The main aim of 
the present paper is to describe the 
ultrastructural stages of female gamete 
formation and of oocyte degeneration in 
P. perna. The follicular cells and their 
function are also examined. 


MATERIALS AND METHODS 


Samples of five mussels each (3-4 cm 
long) were collected in December 1999, 
January, March, June and July 2000 from 
the mid-tide level at Cap Ghir (50 km 
north of Agadir town). Small portions of 
female gonad (1-3 mm? each) were fixed 
for 60 min in 2% glutaraldehyde (0.4 M 
sodium cacodylate buffer, pH 7.2) at 
4*C. The tissue was then washed in 0.4 
M cacodylate buffer (3 x 10 min) and 
postfixed for 90 min in 1% osmium 
tetroxyde (in 0.4 M cacodylate buffer) at 
4%C. After dehydration through a 
gradual ethanol series, the tissue was 
directly embedded in Epon resin at 37 
“C for 60 min and was subsequently 


BENOMAR £7 AL.: Ultrastructural study of oogenesis in the African mussel, Perna perna 


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Figure 1. A. Primary oogonia (Og) and follicular cells (Ec) of Perna perna, along the inner side of 
the acinar wall. Some vesicular cells (Vc) were located along the outer side. da, dense aggregates; 
G, Golgi apparatus; gl, glycogen; Li, lipid droplets; m, mitochondria; N, nucleus, RER, rough 
endoplasmic reticulum; pc, cytoplasmic projections of follicular cells; v, vacuoles. B. Interacinar 
vesicular cells showing two types of granules (g1, and g2). Scale bars, 2 um. 

Figura 1. A. Oogonia primaria (Og) y células foliculares (Fc) de Perna perna, sobre el lado interno de 
la pared acinar. Algunas células vesiculares (Vc) se ubicaron en el lado externo. da, agregados densos; G, 
Aparato de Golgi; gl, glucógeno; Li, gotitas de lípidos; m, mitocondria; N, núcleo, RER, retículo endo- 
plasmático rugoso; pc, proyecciones citoplasmáticas de células foliculares; v, vacuolas. B. Células vesicu- 
lares interacinares mostrando dos tipos de gránulos (g1, and g2). Escalas, 2 ym. 


placed at room temperature during the were collected on copper grids and were 
following 12 hours. Semi-thin sections stained for 20 min with urany]l acetate, 
of each gonad portion were stained at followed by lead citrate for 5 min. Sec- 
room temperature with 0.5% toluidine tions were examined using a Siemens 


blue in 2.5% Na2CO». Ultrathin sections 102 electron microscope. 


Iberus, 28 (2), 2010 


E 


Figure 


2. Oogoniae of Perna perna. A. Zonula occludens (arrows) and zonula adherens (arrow 


heads) between an oogonia (Og), a follicular cell, and a spindle-shaped cell. B-C. Nuclei of two 
oogoniae in metaphasis (first meiotic division). Scale bars, 2 um. 

Figura 2. Oogonias de Perna perna. A. Zonula occludens (flechas) y zonula adherens (puntas de flecha) 
entre una oogonia (Og), una célula folicular, y una célula fusiforme. B-C. Núcleos de dos oogonias en 


metáfase (primera división meiótica). Escalas, 2 ym. 


Different measurements for each cell 
stage of ovogenesis (a least of 10 cells 
per stage) were also performed. Indivi- 
dual values recorded for each measure- 
ment and each cell stage were averaged. 


RESULTS 


Numerous acini, surrounded by con- 
nective tissue and an external ciliated 
epithelium, constituted each gonad (Fig. 
1A). Along the inner side of each acinar 
wall, the different developmental stages 
of oogenesis, from oogoniae up to the 
first stages of the first meiotic division, 
could be easily observed. All of them 
were surrounded by follicular cells and 
spindle-shaped cells containing glycogen 


inclusions (Fig. 1A). In the interacinar 
connective tissue, other vesicular cells, 
each containing two types of granules, 
were found (Fig.1B). Along the outer side 
of the acinus, several muscular fibers 
were sometimes observed (Fig. 3A). 
Oogoniae: Groups of primary oogo- 
niae (primordial cells), each constituted 
by 2 or 4 cells, were found along the 
inner side of each acinar wall (Fig. 1A, 
B). Follicular cells characterized by their 
cytoplasmic extensions were close to 
these cell groups. The primary oogoniae 
were round or elongated, measured 5 to 
8 mun in size, and were characterized by 
a high nucleo-cytoplasmic ratio. They 
were interconnected by zonulae occlu- 
dens and zonulae adherens (Fig. 24). 
Their nuclei (3-6 um) contained small 


BENOMAR E7 AL.: Ultrastructural study of oogenesis in the African mussel, Perna perna 


a Pz, 
¿ ie A IE 


e 


Figure 3. Previtellogenic oocyte of Perna perna. A. Early previtellogenic oocyte. The nucleus (N) 


showed a wide perinuclear cisterna (*) and is positioned near the basal lamina (bl) of the acinus. 
G, Golgi apparatus; Li, lipid droplets; m, elongated mitochondria; mf, muscular fibers; r, ribo- 
somes. B. Late previtellogenic oocyte. Zonula occludens (double arrows) and zonula adherens 
(arrow heads) between one oocyte and two follicular cells. Scale bars, A: 1 qm; B: 0.2 qm. 

Figura 3. Ovocitos previtelogénicos de Perna perna. A. Ovocito previtelogénico temprano. El núcleo (N) 
presentaba una cisterna perinuclear (*) ancha y se sitúa cerca de la lámina basal (b1) del acino. G, 
Aparato de Golgi; Li, gotitas de lípidos; m, mitocóndria alargada; mf, fibras musculares; r, ribosomas. 
B. Ovocito previtelogénico tardío. Zonula occludens (flechas dobles) y zonula adherens (puntas de flecha) 
entre un ovocito y dos células foliculares. Escalas, A: 1 um; B: 0,2 um. 


patches of chromatin. In several cells, 
the chromatin formed a thin border 
along the inner side of the nuclear enve- 
lope, a single or two nucleoli could be 
observed, and the nuclear envelope was 
indented. Numerous ribosomes, spheri- 
cal mitochondria, some cisternae of 
endoplasmic reticulum, several lipid 
droplets with no membrane, and mem- 
brane-bounded dense vesicles were 
found in the cytoplasm (Figs. 1-2). 

The secondary oogoniae measured 5 
to 6 um in size, and the diplotene stages 
of the first meiotic division (Fig. 2B, C) 
were easily recognizable in their volu- 
minous nuclei (4.5-5.5 um). However, 
synaptonemal complexes were not 
found in these nuclei. These cells con- 
tained the same organelles as described 
for primary oogoniae. 

Previtellogenic oocytes: The early pre- 
vitellogenic oocytes (Fig. 3A) were con- 
nected together and to follicular cells via 
zonulae  occludens and  zonulae 
adherens (Fig. 3B). They were round or 
elongated, with irregular outlines, and 


their size ranged from 9 to 15 um. Their 
cytoplasm was more voluminous than 
that of oogoniae and was increased by 
the addition of ribosomes, of often elon- 
gated mitochondria, and of smooth or 
rough endoplasmic reticulum (ER). 
Several vacuoles around the Golgi appa- 
ratus could be observed. Their nucleus 
became elongated, with scattered chro- 
matin and a thicker nuclear envelope, 
showing a clearly wider perinuclear cis- 
terna and the presence of ribosomes 
along the outer side of the outer nuclear 
membrane. 

At a later stage (Fig. 4A-E), the 
oocytes were elongated and their size 
reached up to 18 um. Their nuclei 
became irregular, multilobulated, and 
were eccentrically positioned. Numer- 
ous pores through both membranes of 
the nuclear envelope could be seen and 
the nucleolus was often in an eccentric 
position. At the same time, dense aggre- 
gates with no membrane appeared in 
the cytoplasm, near the nucleus. The 
rough ER increased in size and some 


Iberus, 28 (2), 2010 


Figure 4 A-E. Late previtellogenic oocytes of Perna perna. Al, annulate lamellae; da, dense aggre- 
gates; G, Golgi apparatus with a vesicle (gv): g, granule; Li, lipid droplets; m, mitochondria; N, 
nucleus; RER, rough endoplasmic reticulum; arrow, yolk granule. Scale bars, 1 pm. 

Figura 4 A-E. Ovocitos previtelogénicos tardíos de Perna perna. Al, laminillas anulares; da, agregados 
densos; G, Aparato de Golgi con una vesícula (gu): g, gránulo; Li, gotitas de lipidos; m, mitocóndria; N, 
núcleo; RER, retículo endoplasmático rugoso; flecha, gránulo de vitelo. Escalas, 1 ym. 


10 


BENOMAR ET AL.: Ultrastructural study of oogenesis in the African mussel, Perna perna 


q 


VE 


Figure 5. Pinocytosis vesicle (large arrow) with a lipid droplet and glycogen particles (small 
arrows). *, glycogen deposit in a spindle-shaped cell. Zonula occludens (double arrows) and zonula 
adherens (arrow heads) between one oocyte and two follicular cells. Scale bar, 1 qm. 

Figure 5. Vesícula de pinocitosis (flecha grande) con una gotita de lípidos y partículas de glucógeno 
(flechas pequeñas). *, depósito de glucógeno en una célula fusiforme. Zonula occludens (flechas dobles) y 
zonula adherens (puntas de flecha) entre un ovocito y dos células foliculares. Escala, 1 ym. 


lipid vesicles became bigger in its vicin- 
ity. In close proximity, elongated gran- 
ules containing a dense granular mater- 
ial were sometimes observed. Lamellar 
structures of ER and the first yolk gran- 
ules appeared near the Golgi apparatus 
(Fig 4C). In the basal part of the 
ooplasm and particularly in the zones of 
contact between the oocyte and its sur- 
rounding cells (follicular cells, and 
spindle-shaped cells), some pinocytosis 
vacuoles, each containing lipid droplets 
and several deposits probably of glyco- 
gen, could be seen (Fig. 5). 

Vitellogenic oocytes: Their develop- 
ment within the gonad can be arbitrarily 
divided into four stages, from A the 
youngest, to D the oldest. Each early 
vitellogenic oocyte (A stage) kept 
contact with the acinar wall by cytoplas- 
mic projections (Fig. 6A, B) and was sur- 
rounded by follicular cells which are 
apically connected by desmosome-like 
junctions. Measuring 30 yum in size, this 
oocyte contained an elongated nucleus 
(9 um) with a very dense nucleolus. In 
the zone of attachment of the oocyte 
with the acinar wall, and particularly in 
cytoplasmic projections, some cisternae 


of rough ER could be observed. Several 
microvilli appeared on the outer surface 
of the apical oolemma (Fig. 6A). At a 
later stage in vitellogenesis (B stage), the 
nucleus became spherical and increased 
in size (Fig. 6C). The nucleolus was in 
an eccentric position and was sur- 
rounded by an electron-lucent material, 
while the nucleoplasm was constituted 
by moderately dense patches of hete- 
rochromatin and scattering euchro- 
matin. A previtelline space was forming 
between the apex of the oocyte and the 
surrounding follicular cells, and an 
amorphous electron-lucent material, 
originating from the oocyte, settled into 
this space between the numerous apical 
microvilli formed by the oolemma. Pro- 
gressively, this previtelline space 
extended along the lateral sides of the 
oocyte and the microvilli became coated 
by the amorphous material. Concur- 
rently, there was an increase in the 
number of saccules forming the Golgi 
apparatus and in the number of mito- 
chondria (mainly in the attachment zone 
of the oocyte) (Fig. 64). The rough ER 
greatly increased in volume and showed 
circular lamellae, surrounded by numer- 


11 


Iberus, 28 (2), 2010 


Figure 6. Vitellogenic oocytes of Perna perna. A. A-stage oocytes, with cytoplasmic projections (cp) 
and microvilli (mv) developing at the apex of cells. Flattened rough ER cisternae (RER) were posi- 
tioned near the nucleus or in cytoplasmic projections (cp). A ring of rough ER surrounded a yolk 
granule (Y). Li, lipid droplets; arrow head, a Balbiani's body. B. cortical granules (cg) near several 
cisternae of rough ER (arrows). C. B-stage oocytes. Their nucleolus (nu) was in an eccentric posi- 
tion and was surrounded by an electron-lucent nuclear material (arrows). A myelin-like figure (mf) 
in a follicular cell can be observed. Flattened cisternae of rough ER were located in the vicinity of 
the nucleus. Scale bars, A, C: 2 qm; B: 1 um. 

Figura 6. Ovocitos vitelogénicos de Perna perna. A. Ovocitos de estadio A, con proyecciones citoplasmáticas 
(cp) y microvellosidades (mv) desarrollandose en el ápice de las células. Cisternas aplanadas del RE rugoso 
(RER) eran situadas cerca del núcleo o en proyecciones citoplasmáticas (cp). Un anillo de RE rugoso rodeaba 
un gránulo de vitelo (Y). Li, gottas de lípidos; punta de flecha, un cuerpo de Balbiani. B. gránulos corticales 
(cg) cerca de varias cisternas de RE rugoso (flechas). C. Ovocitos de estadio B. Su nucleolo se situaba en posi- 
ción excéntrica y era rodeado por un material electrón lúcido del núcleo (flechas). Se puede observar una 
figura parecida con mielina (mf) en una célula folicular. Algunas cisternas aplanadas de RE rugoso se situ- 
aban en las inmediaciones del núcleo. Escalas, A, C: 2 ym; B: 1 um. 


12 


BENOMAR £7 AL.: Ultrastructural study of oogenesis in the African mussel, Perna perna 


Figure 7. Vitellogenic oocytes of Perna perna. A-E. C-stage oocytes showing cytoplasmic indentations (oi, 
or arrows) in their nuclei. g, granules located in the ooplasm; Li, lipid droplets; *, a dense granule with no 
membrane. See also an electron-lucent patch (ng) in the nucleus (in A), a yolk granule (Y) surrounded by 
a ring of rough ER (in B), the presence of numerous pores (arrow heads) through the nuclear envelope 
(in C), that of annulate lamellae (Al) surrounding a yolk granule (in D), and Balbianis vitelline bodies 
(arrows) near cisternae of rough ER (double arrow) (in E). Scale bars, A, B: 5 um; C-E: 1 qm. 

Figura 7. Ovocitos vitelogénicos de Perna perna. A-E. Ovocitos de estadio Cmostrando indentaciones cito- 
plasmáticas (oi, o flechas) en sus nucleos. g, gránulos situados en el ooplasma; Li, gotitas de lipidos; *, a 
gránulo densosin membrana. Nótese también una mancha electron lúcida (ng) en el núcleo (en A), un 
gránulo de vitelo (Y) rodeado por un anillo de RE rugoso (en B), la presencia de numerosos poros (puntas de 
flecha) atravesando la membrana nuclear (en C), la de laminillas anulares (Al) rodeando un gránulo de 
vitelo (en D), así como los cuerpos vitelinos de Balbiani (flechas) cerca de cisternas del RE rugoso (flechas 
dobles) (en E). Escalas, A, B: 5 ym; C-E: 1 yum. 


13 


Iberus, 28 (2), 2010 


Figure 8. Vitellogenic oocytes (D stage) of Perna perna. A-B. Different shapes of the rough ER, 
with a proliferation of lipid droplets near mitochondria (m) and rough ER cisternae (arrow heads). 
da, dense granules; arrows, glycogen particles linked to lipid droplets; *, cortical granules (in B). 
Scale bars, 1 um. 

Figura 8. Ovocitos vitelogénicos (estadio D) de Perna perna. A-B. Distintas formas del RE rugoso, con 
proliferación de gotitas de lípidos cerca de una mitocóndria (m) y cisternas del RE rugoso (puntas de 
flecha). da, gránulos densos; flechas, partículas de glucógeno ligadas a gotitas de lípido; * gránulos corti- 


cales (en B). Escalas, 1 um. 


ous lipid vesicles. Some cortical gran- 
ules (Fig. 6B) limited by an undulating 
membrane, each containing a thin gran- 
ular material (probably of glycopro- 
teinic origin), were also observed near 
the rough ER and the Golgi apparatus. 
They are formed by an autosynthetic 
way from these organelles. They became 
progressively denser and were localized 
at the periphery of the oocyte. 

The mature oocytes (C stage) 
reached 70 mm in size (Fig. 7A-E), while 
their shape became pedunculate. They 
were connected to follicular cells by 
zonulae  occludens and  zonulae 
adherens. In their nuclei (30 mm), the 
ring- or crescent-shaped nucleolus was 
dense and in an eccentric position, while 
numerous pores through the nuclear 
envelope could be observed. Deep cyto- 
plasmic indentations (Fig. 7A-C) could 


14 


be seen in the nucleus, each containing 
membrane-bounded lipid droplets and 
other dense granules with no mem- 
brane. The microvilli and the amor- 
phous material constituted a vitelline 
membrane, which became detached 
later from the oocyte and thus created a 
dense perivitelline space between the 
oocyte and the vitelline membrane. The 
lamellae of rough ER (Figs. 6-8) were of 
variable forms (annulate, circular, ovoid, 
flattened, or crescent-shaped) and were 
often positioned near the nucleus. The € 
oocytes contained the same organelles 
as described for B cells. However, four 
other structures, often positioned in the 
vicinity of the rough ER and mitochon- 
dria, could be noted. First, membrane- 
bound lipid vesicles (0.8 mm) became 
larger via their fusion (Fig. 9A). Sec- 
ondly, other lipid inclusions (0.5 mm) 


BENOMAR ET AL.: Ultrastructural study of oogenesis in the African mussel, Perna perna 


Figure 9 A-E. Vitellogenic oocytes (D stage) of Perna perna, different types of granules. See the single mem- 
brane-bounded lipid (Li) droplets (in A), several lipid inclusions with no membrane, near mitochondria 
(m) and cisternae of rough ER (in B), some membrane-bounded inclusions containing a granular mate- 
rial (*) and resulting from pinocytosis at the periphery of the ooplasm (arrow) (in A and C), several elec- 
tron-lucent granules (IV), each showing two zones of different densities (d1, and d2) (in D), lipid 
droplets, each linked to a particle of glycogen (arrows) (in E, see also Fig. 8B). Scale bars, 1 jm. 
Figura 9 A-E.. Ovocitos vitelogénicos (estadio D) de Perna perna, distintos tipos de gránulos. Nótense las 
gotitas de lípidos delimitadas por una membrana sencilla (Li) (en A), varias inclusiones de lípidos sin 
membrana, cerca de mitocondrias (m) y cisternas de RE rugoso (en B), algunas inclusiones delimitadas por 
membranas, conteniendo material granular (*%) y resultando de pinocitosis a la periferia del ooplasma 
(flecha) (en A y C), varios gránulos electrón lucidos (IV), cada uno con dos zonas de densidad diferente 
(41 y d2) (in D), gotitas de lípidos, cada uno ligado a una partícula de glucógeno (flechas) (en E, véase 
también Fig. 8B). Escalas, 1 ym. 


15 


Iberus, 28 (2), 2010 


e 


Figure 10. Follicular cells of Perna perna. A-E Irregularly-shaped nuclei. g, granules; gl, particles of glycogen; 
Li, lipid droplets; Ly, lysosomes; m, mitochondria; mf, myelin-like figures; N, nucleus; nu, nucleolus; RER, 
cisternae of rough ER; SER, smooth ER. See also lipid droplets in the folicular cells (in D), the presence of 
granules with two type of electron-lucent materials (in E), and a single inclusion containing a lipid droplet 
and several glycogen particles forming a rosette (in F). Scale bars, A, B, E: 1 pm; C, D: 5 qm. 

Figura 10. Células foliculares de Perna perna. A-E Núcleos con forma irregular. g, gránulos; gl, partículas de 
glucógeno; Li, gotitas de lípidos; Ly, lisosomas; m, mitocóndria; mf. figuras con aspecto de mielina; N, núcleo; 
nu, nucleolo; RER, cisternas del RE rugoso; SER, RE liso. Nótense también gotitas de lípidos en las células folic- 
ulares (en D), la presencia de gránulos con dos clases de materiales electrón lucidos (en E), y una inclusión ais- 
lada conteniendo una gotita de lípido y varias partículas de glucógeno formando una roseta (en E). Escalas, 
ABE CAD ID 


16 


BENOMAR £7 AL.: Ultrastructural study of oogenesis in the African mussel, Perna perna 


E 


Ed $ SS , 
S EF 
ESPA e 


E 


Figure 11. Septate desmosome (arrows) between two follicular cells of Perna perna. Scale bar, 1 um. 
Figura 11. Desmosoma septado (flechas) entre dos células foliculares de Perna perna. Escala, 1 um. 


with no membrane and showing a 
denser peripheral condensation (Fig. 9B) 
could be observed. Thirdly, membrane- 
bound inclusions (Fig. 9C), containing a 
granular material generally denser at 
their periphery, resulted from pinocyto- 
sis in the periphery of the ooplasm at 
the end of vitellogenesis. These last 
granules, which resulted from material 
extracellular endocytosis contained thin 
glycogen particles and were different 
from the cortical granules, described in 
the late previtellogenic oocytes. Lastly, 
some granules contained two granular 
materials, the first being more electron- 
lucent than the other (Fig. 9D). All these 
types of inclusions acted in the forma- 
tion of yolk. Ringed lamellae, probably 
deriving from the rough ER, sur- 
rounded a big yolk granule or several 
smaller vesicles sometimes with several 
mitochondria, and developed in the 
vicinity of lipid droplets (Fig. 7E). They 
corresponded to Balbiani's vitelline 
bodies (PIPE, 1987). 

In spite of their polyhedral shape, 
the D oocytes (postvitellogenic oocytes) 
were deformed by the presence of other 
oocytes in the lumen of the acinus. The 
nuclear envelope showed numerous 
pores and was very indented, while the 
ring- or crescent-shaped nucleolus was 
in an eccentric position. Numerous cor- 
tical granules, numerous regular yolk 


vesicles of variable density, and some 
elycogen deposits linked to lipid 
droplets (Fig. 9E) were found. In several 
zones of the cytoplasm, are stacks of 
rough ER parallel lamellae. 

Follicular cells: During previtellogen- 
esis, each oocyte was completely sur- 
rounded by small and irregularly- 
shaped follicular cells (Fig. 10A-D). Each 
of these last cells contained a big 
nucleus, of variable form, and projected 
cytoplasmic extensions into the lateral 
part of the oocyte. The follicular cells 
were interconnected together by septate 
desmosomes (Fig. 11). Their cytoplasm 
also contained numerous stacks of 
rough ER cisternae (or annulate lamel- 
lae), a smaller zone of smooth ER, 
numerous mitochondria, well-devel- 
oped lysosomes, multivesicular bodies, 
myelin-like figures, and small single or 
clumped granules of glycogen, some- 
times linked to lipid droplets (Fig. 10P). 
Some other inclusions, each containing a 
dense material and another more lucent 
(Fig. 10E), were also present near lipid 
droplets. The peripheral part of their 
cytoplasm also contained some pinocy- 
tosis vesicles, each containing a lipid 
droplet and/or a glycogen granule (Fig. 
5). When the oocyte was at the B stage 
of vitellogenesis, the follicular cells 
became detached from the apex of the 
oocyte, which bulged freely into the 


Iberus, 28 (2), 2010 


acinar lumen and became pedunculate, 
as it was still attached to the inner wall 
of the acinus. At the C stage of vitelloge- 
nesis, the follicular cells were reduced to 
their basal part and, finally, became 
completely detached from the oocyte. 

In the spaces between the follicular 
cells and the spindle-shaped cells, free 
particles of glycogen were frequently 
observed, particularly during previtello- 
genesis (Fig. 5). 


DISCUSSION 


The female gamete formation in P. 
perna was similar to that described in 
other bivalvia, such as Brachidontes vig- 
iliae (BERNARD, DAVIES AND HODGSON, 
1988), Crassostrea virginica (ECKELBARGER 
AND Davis, 1996), Mytilus edulis (ALBER- 
TINI, 1985; PIPE 1987), Pecten maximus 
(DORANGE AND LE PENNEC, 1989), Pinc- 
tada margaritifera (lHIELLEY, 1993), or 
Pinna nobilis (GAULEJAC, HENRY AND 
VICENTE, 1995). The accumulation of 
ribosomes and the presence of numer- 
ous nuclear pores in previtellogenic 
oocytes indicated a great synthesis of 
proteins and an increased transport of 
material. The perinuclear dense aggre- 
gates present in the cytoplasm of these 
oocytes might correspond to extruded 
nucleolar material (especially ribonucle- 
oproteins), as described by several 
authors (ALBERTINI, 1985; DORANGE AND 
LE PENNEC, 1989; THIELLEY, 1993; 
GAULEJAC ET AL., 1995). The presence of 
mitochondria in the stalk of the previtel- 
logenic oocyte suggested a transfer of 
material. However, the present study 
did not allow us to observe micro- 
tubules in this stalk, such as described 
in the previtellogenic oocyte of Pinna 
nobilis (GAULEJAC ET AL., 1995), or in that 
of Anodonta (BEAMS AND SEKHON, 1966). 

In the vitellogenic oocytes, the 
growth of cytoplasm in volume was 
mostly due to the accumulation of inclu- 
sions. Among them, the cortical gran- 
ules were found during all stages of 
vitellogenesis and this finding agreed 
with the report by GAULEJAC ET AL. 
(1995) in Pinna nobilis. The nature of 


18 


their contents can only be speculated 
upon based on morphological investiga- 
tions. However, evidence for a glycopro- 
teinic content is recognized in oocytes of 
most bivalve species (e.g. ALBERTINI, 
1985; GAULEJAC ET AL., 1995). According 
to PIPE (1987), the number of these corti- 
cal granules might increase via their 
division. The formation of these cortical 
granules involved the synthesis of yolk 
materials by the  proteosynthetic 
organelles of the oocyte. This autosyn- 
thetic-type formation has been 
described in some molluscan species (DE 
JONG-BRINK, BOER AND JOOSSE, 1983; 
MEDINA, GARCIA, MORENO AND LOPEZ- 
CAMPOS, 1986). The production of yolk 
appears to involve the collaboration of 
Golgi complexes and RER, as these 
organelles were observed in close asso- 
ciation with yolk bodies, as observed in 
several mollusc species (ECKELBARGER 
AND Davis, 1996; ECKELBARGER AND 
YOUNG, 1997). The other types of inclu- 
sions found in mature oocytes warrant 
special comment. First, the two types of 
lipid droplets, i.e. small inclusions with 
no membrane and bigger, membrane- 
bound inclusions, recorded in P. perna 
might correspond to two successive 
stages in the formation of these lipids 
via the action of smooth ER, of Golgi 
apparatus, of mitochondria, and of Bal- 
biani's bodies (DE JONG-BRINK ET AL., 
1983). Thus, in the caenogastropod Colus 
stimpsoni, West (1983) reported that 
lipid formation derived from the 
autosynthetic activities of the oocyte via 
the endoplasmic reticulum. Secondly, 
the granules, observed at the C stage of 
vitellogenesis and containing two types 
of electron-lucent materials, have not 
been reported in the literature on 
bivalvia. To explain this last finding, the 
most likely hypothesis was to admit that 
their contents would be of lipoproteinic 
origin via the fusion of lipid droplets 
and of proteins originating from rough 
ER. Lastly, the inclusions resulting from 
pinocytosis at the end of vitellogenesis 
were also reported by several authors in 
other species of molluscs (WesrT, 1981, 
1983; DE JONG-BRINK ET AL., 1983; ECKEL- 
BARGER AND  BLADES-ECKELBARGER, 


BENOMAR ET AL.: Ultrastructural study of oogenesis in the African mussel, Perna perna 


1989; ECKELBARGER AND YOUNG, 1997). 
Heterosynthetic process involved the 
Golgi complex and RER on the one 
hand, and endocytosis of extracellular 
material on the other hand, so that lipid 
reserves would be produced by the 
oocyte and glycogen would be endocy- 
tosed (West, 1981,1983). In fact, in P. 
perna, the inclusions might originate, 
either from spindle-shaped cells, which 
contained glycogen deposits, or from 
interacinar vesicular cells, which had 
two types of granules. As free glycogen 
particles could be seen in the intercellu- 
lar spaces between the oocyte, the follic- 
ular, and the spindle-shaped cells, it 
might be hypothesized that the free par- 
ticles would be pinocytosed by the cyto- 
plasm of the vitellogenic oocyte, or by 
that of follicular cells. Yolk synthesis in 
P. perna was similar to that described in 
other molluscs species (WEsT, 1983; Eck- 
ELBARGER AND YOUNG, 1997). However, 
this massive intake of exogenous sub- 
stances in the ooplasm was probably not 
the single way for the formation of yolk. 
Indeed, the presence of glycogen parti- 
cles, linked to lipid droplets, in the 
ooplasm suggested that they might 
form lipid-carbohydrate complexes, as 
those described in the oocytes of Mytilus 
edulis (ALBERTINI, 1985). Another way in 
P. perna might be the direct transforma- 
tion of mitochondria into yolk granules, 
as reported in the oocytes of many 
bivalve species (GAULEJAC ET AL., 1995). 
The formation of the vitelline mem- 
brane occurred during the B and C 
stages of vitellogenesis, and the material 
constituting it originated from the 
oocyte (e.g. PIPE, 1987), the follicular 
cells (DORANGE AND LE PENNEC, 1989), 
or both. A part of this material might be 
composed of the contents of cortical 
granules, as ALBERTINI (1985) noted the 
release of mucus from these granules in 
the vitelline membrane surrounding the 
oocyte of Mytilus edulis. The results 
noted in the present study indicated the 
material constituting the vitelline mem- 
brane was produced by the oocyte. Our 
results agreed with those of Wourms 
(1987) on the fact that electron 
microscopy reveals both microvilli and 


an extracellular coat in the overwhelm- 
ing majority of invertebrate oocytes. 

The ultrastructural changes noted in 
the atretic oocytes of P. perna were 
similar to those noted in other species of 
bivalvia, even though the outcome of 
reserves in P. perna was different. The 
accumulation of yolk granules consti- 
tuted a dense mass, which is released in 
the lumen of the acinus, and this 
oosorption allowed the turn over of 
nutrients to assure the energetic needs 
for the bivalve basal metabolism (Hout- 
TEVILLE, 1974). Another way for re-using, 
oocyte remnants in P. perna was consti- 
tuted by the epithelial cells bordering 
the gonoducts, as that reported by PIPE 
(1987) in Mytilus edulis, and also by the 
follicular cells (see the review by DE 
JONG-BRINK ET AL., 1983). 

Some reports were already per- 
formed on the relationship between the 
follicular cells and the maturation of 
oocytes in bivalvia. The presence of fol- 
licular cells completely surrounding the 
young oocyte and their detachment from 
it in later stages were already reported in 
many species (e.g. BERNARD ET AL., 1988; 
GAULEJAC ET AL., 1995) and this finding 
underlined the important role of these 
cells in the growth of oocytes, and their 
nutrition (DE JONG-BRINK ET AL., 1983). 
Owing to their organelles, the follicular 
cells had the capacity to participate in 
the synthesis of many substances and to 
phagocytose materials originating from 
atretic or degenerating oocytes. The ex- 
ogenous substances, which are intaken 
by the oocyte of P. perna during vitello- 
genesis via pinocytosis, might originate 
from the secretions of these follicular 
cells. The capacity of these last cells to 
phagocytose the remnants of oocytes in 
P. perna is reflected by the presence of 
their well-developed lysosomes and of 
deposits of lipid and glycogen deposits 
in their cytoplasm. It is possible that 
products originating from atretic oocytes 
permitted a transfer of precursors neces- 
sary for the vitellogenesis of other devel- 
oping oocytes, and the synthesis and 
storage of lipid and glycogen inclusions 
which can be used later for vitellogene- 
sis, as suggested by GAULEJAC ET AL. 


19 


Iberus, 28 (2), 2010 


(1995) in Pinna nobilis. According to 
WesT (1983), the follicular cells synthe- 
siz-ed glycogen and the oocytes se- 
questered it through an endocytotic 
process. 

The studies made by ID HALLA ET AL. 
(1997) on the gonad of P. perna demon- 
strated the presence of a single type of 
reserve cells. However, in the present 
work, lipid droplets and inclusions of 
glycogen were present in reserve cells, 
so that further studies are necessary to 
determine if different types of reserve 
cells exist in the gonad of P. perna or if 
the cell found in the present study 
shows different cellular stages which 
succeed in relation to the reproductive 
cycle of P. perna. 

In conclusion, the formation of 
female gametes in P. perna was greatly 
similar to the oogenesis described in 
other species of bivalvia and particu- 
larly in Mytilidae. The only difference 


BIBLIOGRAPHY 


ALBERTINI L. 1985. Recherches cytologiques et ex- 
périmentales sur l'ovogénese chez la moule, 
(Mytilus edulis L., mollusque bivalve). Doc- 
toral thesis, University of Caen, 147 p. 

BEAMS H.W. AND SEKHON 5.5.1966. Electron 
microscope studies on the oocyte of the 
fresh-water mussel (Anodonta), with spe- 
cial reference to the stalk and mechanism 
of yolk deposition. Journal of Morphology, 
119: 477-502. 

BENOMAR S., BELHSEN O.K., GOUX D., MATHIEU 
M. AND MOUKRIM A. 2007. Ultrastructural 
studies on the spermatogenesis of the African 
mussel Perna perna (Bivalvia: Mytilidae). 
Iberus, 26(2): 1-14. 

BERNARD R.T.F., DAVIES B.R. AND HODGSON 
A.N. 1988. Reproduction in a brackish-water 
mytilid: gametogenesis and embryonic de- 
velopment. The Veliger, 30: 278-290. 

BERRY P.F. 1978. Reproduction, growth, and 
production in the mussel Perna perna, on the 
east coast of South Africa. Investigational Re- 
ports of the Oceanography Research Institute, 
Durban, 48: 1-28. 

DE JONG-BRINK M., BOER H.H. AND JOOSSE J. 
1983. Mollusca. Chapter 14. In: Adiyodi KG, 
Adiyodi RG, (eds.). Reproductive biology of 
invertebrates. I. Oogenesis, oviposition, and 
oosorption, John Wiley and Sons, New York, 
pp. 297-355. 


20 


concerned the composition of the yolk 
in the mature oocyte, as it was consti- 
tuted of several endogenous substances 
(lipoproteins mainly) and also of exoge- 
nous materials intaken by the oocyte via 
pinocytosis. In P. perna, vitellogenesis 
combined both autosynthetic and het- 
erosynthetic processes, and involved the 
Golgi complex and RER on the one 
hand, and endocytosis of extracellular 
material on the other hand. Further 
studies are necessary to investigate the 
composition of vitelline granules in the 
oocytes using cytochemical methods 
and transmission electron microscopy. 


ACKNOWLEDGEMENTS 


The authors are grateful to Ms. A.M. 
Renou and Mr. D. Goux for the realiza- 
tion of ultrathin sections and Mr. J.L. 
Herblot for photographic services. 


DORANGE G. AND LE PENNEC M. 1989. Ultra- 
structural study of oogenesis and oocytic de- 
generation in Pecten maximus from the Bay of 
St. Brieux. Marine Biology, 103: 339-348. 

ECKELBARGER K.J. AND BLADES-ECKELBARGER 
P.I. 1989. Structure of the ovotestis and evi- 
dence for hetero synthetic incorporation of 
yolk precursors in the oocytes of the nudi- 
branch Mollusc, Spurilla neapolitana. Journal 
of Morphology, 201: 105-118. 

ECKELBARGER K.J. AND Davis C.V. 1996. 
Ultrastructure of the gonad and gametogenesis 
in the eastern oyster, Crassostrea virginica. L. 
Ovary and oogenesis. Marine Biology, 127: 79-87. 

ECKELBARGER K.] AND YOUNG C.M. 1997. Ul- 
trastructure of the ovary and oogenesis in the 
dmethane-seep mollusc Bathynerita naticoidea 
(Gastropoda: Neritidae) from the Louisiana 
slope. Invertebrate Biology, 116 (4): 299-312. 

GAULEJAC B. DE, HENRY M. AND VICENTE N. 
1995. An ultrastructural study of gametoge- 
nesis of the marine bivalve Pinna nobilis (Lin- 
naeus 1758). I. Oogenesis. Journal of Mollus- 
can Studies, 61: 375-392. 

Hicks D.W., TUNNELL J.W. JR AND MCMAHON 
R.F. 2001a. Population dynamics of the non- 
indigenous brown mussel, Perna perna (Lin- 
naeus 1758), in the Gulf of Mexico compared 
to other world-wide populations. Marine 
Ecology Progress Series, 211: 181-192. 


BENOMAR ET AL.: Ultrastructural study of oogenesis in the African mussel, Perna perna 


Hicks D.W., MCMAHON R.F. AND INGRAO D.A. 
2001b. Two invasive mussels in the genus 
Perna in the Gulf of Mexico. In: Virtual Pro- 
ceedings for the State of the Bay Symposium 
V,31 January-2 February 2001. Texas Natural 
Resource Conservation Commission, Austin, 
Texas, pp. 159-170. 

HOUTTEVILLE P. 1974. Contribution a l'étude 
cytologique et expérimentale du cycle du 
tissu de réserve du manteau de Mytilus edulis. 
Doctoral thesis, University of Caen. 

ID HALLA M., BOUHAIMI A., ZEKHNINI A., NAR- 
BONNE J.F., MATHIEU M. AND MOUKRIM A. 
1997. Etude du cycle de reproduction de 
deux especes de moules Perna perna (Linné, 
1758) et Mytilus galloprovincialis Lamarck, 
1819 dans la baie d'Agadir (Sud du Maroc). 
Haliotis, 26: 51-62. 

LasraK T. 1986. The reproductive cycles of the 
intertidal bivalves Crassostrea cucullata (Born, 
1778) and Perna perna (Linnaeus, 1758) from 
Transkei coast, Southern Africa. The Veliger, 
29: 226-230. 

MEDINA A., GARCIA J.C., MORENO F.J. AND 
LOPEZ-CAMPOSJ.P. 1986. Comparative stud- 
ies on the histology of the ovotestis in 
Hypselodoris tricolor and Godiva banyulensis 
(Gastropoda, Opistothobranchia), with spe- 
cial reference to yolk formation. Journal of 
Morphology, 186: 105-118. 


PrPE R.K. 1987. Oogenesis in the marine mus- 
sel Mytilus edulis: an ultrastructural study. 
Marine Biology, 95: 405-414. 

SCHURINK C.E. AND GRIFFITHS C.L. 1991. A com- 
parison of reproductive cycle s and repro- 
ductive output in four Southern African mus- 
sel species. Marine Ecology Progress Series, 76: 
123-134. 

THIELLEY M. 1993. Etude cytologique de la 
gamétogenese, de la sex-ratio et du cycle de 
reproduction chez l'huítre perliere Pinctada 
margaritifera (L) var. cumingí (Jameson), (Mol- 
lusque, bivalves). Comparaison avec le cycle 
de Pinctada maculata (Gould). Doctoral the- 
sis, Université Francaise du Pacifique, 233p.. 

WEsT D.L. 1981. Reproductive biology of Colus 
stimpsoni (Prosobranchia: Buccinidae). IV. 
Oogenesis. The Veliger, 24: 28-38. 

WesT D.L. 1983. Reproductive biology of Colus 
stimpson1 (Prosobranchia: Buccinidae). V. Nu- 
tritive egg formation. The Veliger, 25: 299- 
306. 

WOURMS J.P. 1987. Oogenesis. In Giese A.C,, 
Pearse J.S. and Pearse V.B. (Eds.): Reproduc- 
tion of marine Invertebrates, Vol. IX, General 
aspects: seeking unity in diversity: 50-157. 
Blackwell Scientific Publications and Box- 
wood Press, Pacific Grove, California. 


2 


O Sociedad Española de Malacología  —_—_———  lberus, 28 (2): 23-38, 2010 


New species of Mollusca Solenogastres from the 
Bell" -"shausen Sea and the Antarctic Peninsula (Bentart- 
2006 Expedition) 


Nuevas especies de Moluscos Solenogastres del Mar de 
Bellingshausen y Península Antártica (Expedición Bentart-2006) 


Oscar GARCÍA-ÁLVAREZ*, María ZAMARRO** and Victoriano 
URGORRI** 


Recibido el 27-IV-2010. Aceptado el 31-X-2010 


ABSTRACT 


The Bentart Projects aim to study the benthos of West Antarctica (South Shetland 
Islands, Antarctic Peninsula and Bellingshausen Sea). In this paper, one genus and two 
species new to science (Neomenia expleta sp. nov. and Plicaherpia papillata gen. and 
sp. nov.), as well as an incomplete juvenile of Phyllomenia sp. and an anterior body of 
Amboherpia sp. are studied, all collected during the expedition Bentart-2006 to the 
Antarctic Peninsula and Bellingshausen Sea and belonging to the families: Neomeni- 
idae lhering, 1876, Phyllomeniidae Salvini-Plawen, 1978 and Acanthomeniidae 
Salvini-Plawen, 1978. 


RESUMEN 


Los proyectos Bentart tienen como objetivo el estudio del bentos en la Antártida del Oeste 
(Islas Shetland del Sur, Península Antártica y Mar de Bellingshausen). En este artículo se 
estudian un género y dos especies nuevas para la ciencia (Neomenia expleta sp. nov. y 
Plicaherpia papillata gen. y sp. nov.), así como un juvenil incompleto de Phyllomenia sp. y 
la parte anterior de Amboherpia sp., recogidas durante la expedición Bentart-2006 en la 
Península Antártica y Mar de Bellingshausen, y pertenecientes a las familias: Neomenii- 
dae lhering, 1876, Phyllomeniidae Salvini-Plawen, 1978, and Acanthomeniidae Salvini- 
Plawen, 1978. 


INTRODUCTION 

During the Spanish expedition the area of the Antarctic Peninsula and 
Bentart-2006 for the study of Antarctic its adjacent islands some research had 
benthos to the Antarctic Peninsula and already been done regarding these Mol- 
Bellingshausen Sea, a small collection of lusca, with the result of about 30 
Mollusca Solenogastres was made. In described species, whereas the know- 


* Estación de Bioloxía Mariña da Graña. Universidade de Santiago de Compostela. A Graña. 15590 Ferrol. 
Spain. ogarcia.alvarezGedu.xunta.es 

** Departamento de Zooloxía e Antropoloxía Física. Facultade de Bioloxía. Universidade de Santiago de 
Compostela. Campus Sur s/n. 15782 Santiago de Compostela. Spain. 


23 


Iberus, 28 (2), 2010 


ledge of the Solenogastres fauna in the 
Bellingshausen Sea is very poor, with 
only four known species (GARCÍA- 
ÁLVAREZ AND |URGORRL, 2003a, b; 
GARCÍA-ÁLVAREZ, ZAMARRO AND 
URGORRI, 2009; SALVINI-PLAWEN, 1978). 

In this study four specimens from 
Bentart-2006 are studied, one collected 
in the Antarctic Peninsula and three in 
the Bellingshausen Sea, belonging to 
the families Neomeniidae lhering, 
1876, Phyllomeniidae Salvini-Plawen, 
1978, and Acanthomeniidae Salvini- 
Plawen, 1978 and among which we 
recognize one genus and two species 
new to science. We also provide data 
about an incomplete juvenile of Phyl- 
lomenia sp. and an anterior body of 
Amboherpia sp. 

The biodiversity of the Mollusca 
Solenogastres is still very little known; 
about 260 species have been described 
so far, which represent about 40% of the 
total of the species estimated to exist 
this group of Mollusca. In particular, the 
Antarctic Ocean is the best known geo- 
graphical area concerning the 
Solenogastres fauna, as approximately 
45% of the described species are Antarc- 
tic or Subantarctic. So far, the research of 
the Bentart collection from the four 
expeditions carried out to the South 
Shetland Islands, Antarctic Peninsula 
and Bellingshausen Sea, has resulted in 
the study of 14 species of Solenogastres 
(including those described here), 9 of 
which were new species to science, 2 
were innominate species and only 3 
were already known species (GARCÍA- 
ÁLVAREZ AND URGORRI, 2003a; GARCÍA- 
ÁLVAREZ ET AL., 2009). These facts 
clearly support the idea that the biodi- 
versity of these Mollusca is still poorly 
known. 

One of the greatest difficulties 
which hampers the study of Solenogas- 
tres is the scarcity of research material, 
as can be observed in most publications 
about this topic and as also happens in 
the study of the Bentart collection. It is 
very common to have only very few 
specimens available or even a single 
specimen for the characterization of the 
new species. This is due to the fact that 


24 


a high percentage of the samples of 
Solenogastres come from studies 
directed to the knowledge of the 
general marine benthos, for which the 
sampling methods used are not the 
most appropriate for the collection of 
small-sized fauna. When the capture 
systems used while sampling are more 
appropriate for the study of meiofauna, 
as for example epibenthic sleds, the 
results are different. These sampling 
systems are being systematically used, 
both in the expeditions for the study of 
the Atlantic abyssal basin and in the 
expeditions we are carrying out off the 
northwest coast of Spain. In both pro- 
jects, in which a priority objective is the 
knowledge of the deep-sea biodiversity, 
the results obtained in the collection of 
small fauna are very satisfactory and a 
large number of specimens of 
Solenogastres were collected (GIL-MAN- 
SILLA, GARCÍA-ÁLVAREZ AND URGORRI, 
2008, 2009). 


MATERIAL AND METHODS 


The four specimens studied were 
collected during the expedition Bentart- 
2006, which was carried out on board 
the BIO Hespérides with an Agassiz 
trawl in January-February 2006 in the 
Bellingshausen Sea and the Antarctic 
Peninsula, with five stations sampled. 
Specimens were fixed and preserved in 
70% ethanol. For the study of sclerites, 
small pieces of cuticle of the dorsal 
middle part of the body and of the 
ventral groove were separated. These 
pieces were treated with sodium 
hypochlorite at 5% for 12 hours to 
isolate the sclerites. They were then 
washed in water, dried in a drying 
chamber at 40"C and mounted on 
Canada balsam. For their anatomical 
study, specimens were decalcified in an 
EDTA solution for 12 hours, included in 
paraffin, cut in transverse series of 5 and 
10 um in section, and stained in Azan 
and Mallory trichromic. Reconstruction 
of the internal anatomy of their anterior 
and posterior body was performed 
manually. 


GARCÍA-ÁLVAREZ ET AL.: Solenogastres from the Bentart-2006 Expedition 


SYSTEMATICS (see GARCÍA-ÁLVAREZ AND SALVINI-PLAWEN, 2007) 


Order NEOMENIAMORPHA Salvini-Plawen, 1978 
Family NEOMENIIDAE lhering, 1876 
Genus Neomenia Tullberg, 1875 


Neomenia expleta sp. nov. (Figs. 1, 2) 


Type material: Holotype 1 specimen, cut in serial sections. Antarctic Peninsula. (Bentart-2006, 
Station PA 42; coordinates: 65.16654” S; 68.9364” W). Water depth: 1272 m. The holotype (cut in 
serial sections) is deposited in the Museo Nacional de Ciencias Naturales of Madrid, namber MNCN 


02/26: 


Etymology: Latin, expletus = completed, finished, achieved. Regarding its complete anatomical 


organization. 


Diagnosis. Specimen 3.4 cm x 0.95 cm 
with 5-7 longitudinal keels. Cuticle 270 
um thick. Three types of sclerites with 
radial insertion: groove-shaped sclerites 
without spear-shaped distal end; solid, 
slightly bent needles; and solid blade- 
shaped scales. Pedal groove with 25-35 cil- 
iated folds that do not get into the pallial 
cavity. Separate cerebral connectives. 
Pharynx with three regions. Posterior 
midgut with a pair of short ventrolateral 
caeca that surround the copulatory stylets. 
Seminal vesicles and seminal receptacle 
present. Opening of the paired spawning 
duct into a genital pouch. Spawning duct 
gland and stylet gland present. Supra- 
pallial glands present. Two pairs of cop- 
ulatory stylets. Prepallial spicules absent. 
Number of respiratory folds 40-45. A dor- 
soterminal sense organ prsent. 

Description. Habitus. The specimen 
is 3.4 cm long, 0.95 cm thick in its 
middle area, narrower in its anterior 
and posterior parts, with 5-7 dorsal and 
lateral longitudinal keels, little marked 
(Fig. 1A). The pedal groove and the atri- 
obuccal and pallial cavities are well- 
marked. Colour in alcohol is brown. 

Mantle. The cuticle is thick, up to 270 
mm with globular to club-shaped epi- 
dermal papillae. the matrix is up to 575 
mm thick. There are three types of scle- 
rites with radial insertion (Fig. 1B): 
groove-shaped sclerites without spear- 
shaped distal end (up to 290 mm long x 
20 mm wide) dorsally more abundant; 
solid, slightly bent needles (up to 260 
mm long) and solid blade-shaped scales 
(up to 125 mm long). 


Pedal groove and pallial cavity. The 
pedal groove bears 25-35 ciliated folds 
that do not reach into the pallial cavity. 
The pallial cavity occupies a subtermi- 
nal position and has 40-45 ciliated respi- 
ratory folds, long and narrow and radi- 
ally arranged (Figs. 1D, 2A) in the dor- 
soposterior region; many folds have 
abundant yellow dyed secretory gran- 
ules in their distal region. The pallial 
cavity has dorsally circular musculature 
and suprapallial glands (Fig. 1D). The 
pallial cavity forms in its ventroanterior 
region a genital pouch (Figs. 1D, 2B), 
into which both spawning ducts come 
out independently in its dorsolateral 
region (Fig. 2B). Both pairs of copula- 
tory stylets also come out ventrolater- 
ally into this pouch (Fig. 1D). 

Digestive system (Fig. 1C). The mouth 
opens into the atrium, which is pro- 
vided with papillae arranged in 
bundles. Three regions can be distin- 
guished lengthwise in the pharynx. The 
first one bears 2 to 4 dorsal folds with 
folded walls; one of these folds sepa- 
rates mediofrontally the mouth from the 
anterior part of the atrium and a ventral 
fold is separated from the ventral wall 
of the digestive tract by a cleft. The folds 
make this first region of the pharynx lat- 
erally narrow and form small lateral 
caeca. The pharynx is internally covered 
in this region with a thin cuticle that 
continues in the second pharyngeal 
region; all walls are folded and it lacks 
glands. It has no defined buccal tube 
and lacks a buccal sphincter. In the 
second region, the pharynx narrows 


23 


Iberus, 28 (2), 2010 


dorsoventrally and  lacks  glands, 
frontally two caeca come out from it: a 
dorsoanterior one and a ventral one. In 
this area it has a dorsal wall without 
folds and with a very soft musculature, 
its ventral wall is folded and provided 
with a thick muscular layer; this ventral 
muscular layer becomes very weak pos- 
teriorly. The third region is character- 
ized by presenting a strong sphincter 
and a folded interior wall, it opens 
frontally into the intestine, which it 
clearly penetrates. It lacks a radular 
system and ventral foregut glandular 
organs. The intestine has a short dor- 
soanterior caecum and a strong serial 
fold due to the dorsoventral muscula- 
ture. Posteriorly, a pair of short ventro- 
lateral caeca come out from the intes- 
tine; each of them surround a pair of 
copulatory stylets (Figs. 2E, F). The 
rectum is short and tubular; its interior 
wall is folded and ciliated. The anus 
opens dorsally on the frontal wall of the 
pallial cavity, flanked by respiratory 
folds. 

Nervous system and sense organs (Figs. 
1C, D). The cerebral ganglion is flat and 
relatively large (750 mm long, 360 mm 
high), several pairs of nerves come out 
from it to the atrial region and two pairs 
of independent connectives come out 
from its posterior region. The pair of 
ventral ganglia is located ventrolaterally 
in the anterior part of the pharyngeal 
region of the sphincter and is joined to 
the digestive tract by a ventral commis- 
sure. Both buccal ganglia are very small 
and located ventrolaterally to the pha- 
ryngeal region of the sphincter. The pos- 
terior part of the body possesses two 
pairs of thick ganglia, a ventral one and 
a lateral one. The supra-rectal commis- 
sure is wide and long and located dor- 
sally to the pallial cavity, immediately 
posterior to the position of the anus. The 
atrial sense organ is provided with 
dense bundles of thin, basally joined 
papillae, mostly in dorsal and lateral 
positions. A dorsoterminal sense organ 
is located in the posterior end of the 
body. 

Reproductive system (Fig. 1D) A pair 
of tubular and narrow gonads is located 


26 


on both sides of the dorsal blood sinus, 
no oocytes are observed, but they 
present spermatozoids in the posterior 
part close to the pericardium, in this 
region the gonads are provided with 
some small ventrolateral pouches, in 
which spermatozoids can be observed 
and which can be interpreted as seminal 
vesicles. The pericardium is voluminous 
and contains a very large heart joined to 
the pericardium wall only through its 
anterior and posterior ends; the heart is 
divided into two parts, a ventricle and 
an auricle. The pericardioducts come 
out ventrolaterally from the posterior 
part of the pericardium, they consist of a 
pair of narrow and internally folded 
ducts that run ventrolaterally to the 
pericardium and the digestive duct and 
present a pair of small seminal vesicles 
almost spherical (Fig. 2C) with sperma- 
tozoids, before joining the spawning 
ducts. The pair of spawning ducts is 
narrow and sinuous, not very glandular 
and come out dorsolaterally and sepa- 
rately into a ventral genital pouch of the 
pallial cavity (Fig. 2B). Each spawning 
duct has a narrow and sinuous seminal 
receptacle, located in the opening area 
of the pericardioducts into the spawn- 
ing ducts as well as a spawning duct 
gland with glandular epithelium that 
comes out into the medial area of the 
spawning duc:t. 

The copulatory system is paired, 
each part is made up of two stylets 
(Figs. 2E, F): a groove-shaped stylet and 
a Spine within a common sheath; they 
are surrounded by the ventrolateral 
caeca of the intestine. Each pair of copu- 
latory stylets is connected to a gland 
(stylet gland) that opens in the distal 
end of the stylets and from which a 
narrow duct comes out to the distal end 
of the spawning duct (Fig. 2D). No pre- 
pallial spines are present. 

Discussion. Eighteen species of the 
genus Neomenia are described at present. 
Among the combination of characteris- 
tics that distinguish Neomenia expleta sp 
nov. from the other species in this 
genus, we should highlight the follow- 
ing for their specific nature: presence of 
keels (having keels: N. carinata Tullberg, 


GARCÍA-ÁLVAREZ ET AL.: Solenogastres from the Bentart-2006 Expedition 


Figure 1. Neomenia expleta sp. nov. A: habitus; B: mantle sclerites; C: schematic organization of the 
anterior body; D: schematic organization of the posterior body. Abbreviations, At: atrial sense organ; 
Bg: buccal ganglion; Bm: posterior caecum of midgut; Cg: cerebral ganglion; Cs: copulatory stylet; 
De: dorsal caecum; Dso: dorsoterminal sense organ; Gp: genital pouch; Ht: heart; Mg: midgut; Mu: 
musculature; Pc: pallial cavity; Pd: pericardioduct; Ph: pharynx; Pr: pericardium; Rf: respiratory folds; 
Sc: suprarectal commissure; Sd: spawning duct; Sdg: spawning duct gland; Sg: copulatory stylet gland; 
Sp: sphincter; Spg: suprapallial gland; Sr: seminal receptacle; Sv: seminal vesicle; Vg: ventral ganglion. 
Figura 1. Neomenia expleta sp. nov. A: habitus; B: escleritos del manto; C: esquema de la organización 
de la parte anterior del cuerpo; D: esquema de la organización de la parte posterior del cuerpo. Abre- 
viaturas, Át: órgano sensitivo atrial; Bg: ganglio bucal; Bm: ciego posterior del intestino; Cg: ganglio 
cerebral; Cs: estilete copulador; Dc: ciego dorsal; Dso: órgano sensitivo dorsoterminal; Gp: bolsa genital; 
Ht: corazón; Mg: intestino; Mu: musculatura; Pc: cavidad paleal; Pd: pericardioducto; Ph: faringe; Pr: 
pericardio; Rf: pliegues respiratorios; Sc: comisura suprarrectal; Sd: conducto de desove; Sdg: glándula 
del conducto de desove; Sg: glándula del estilete copulador; Sp: esfínter; Spg: glándula suprapaleal; Sr: 


receptáculo seminal; Sv: vesícula seminal; Vg: ganglio ventral. 


2 


Iberus, 28 (2), 2010 


Figure 2. Neomenia expleta sp. nov. A: respiratory folds; B: spawning duct opening; C: seminal 
vesicle; D: copulatory stylet gland; E: copulatory stylet; EF: posterior caecum of midgut. Abbrevia- 
tions, Bm: posterior caecum of midgut; Cs: copulatory stylet; Gp: genital pouch; Mg: midgut; Pc: 
pallial cavity; Pd: pericardioduct; Rf: respiratory folds; Sd: spawning duct; Sdg: spawning duct 
gland; Sg: copulatory stylet gland; Sr: seminal receptacle; Sv: seminal vesicle. Scale bars, 200 um. 
Figure 2. Neomenia expleta sp. nov. A: pliegues respiratorios; B: abertura del conducto de desove; C: vesícula 
seminal; D: glándula del estilete copulador; E: estilete copulador; E- ciego posterior del intestino. Abreviaturas, 
Bm: ciego posterior del intestino; Cs: estilete copulador; Gp: bolsa genital: Mg: intestino; Pc: cavidad paleal; 
Pd: pericardioducto; Rf. pliegues respiratorios; Sd: conducto de desove; Sdg: glándula del conducto de desove; 
Sg: glándula del estilete copulador; Sr: receptáculo seminal; Sv: vesícula seminal. Escalas, 200 ym. 


1875, N. labrosa Salvini-Plawen, 1978, N. Gausch, 2004 and N. trivialis Salvini- 
trapeziformis Salvini-Plawen, 1978, N. Plawen and Paar-Gausch, 2004); lacking 
megatrapezata Salvini-Plawen and Paar- distally spear-shaped sclerites (having 


28 


GARCÍA-ÁLVAREZ ET AL.: Solenogastres from the Bentart-2006 Expedition 


Table I. Differences between genera of the family Phyllomeniidae (+ present; - absent). 
Table 1. Diferencias entre los géneros de la familia Phyllomentidae (+ presente; - ausente). 


Phyllomenia  Harpagoherpia  Lituiherpia  Ocheyohempia  Plicaherpia gen.nov. 
Aciculare  * + + + 4 ze 
Paddle-shupeu >uerites + - de a 4 
Hook-shaped sclerites + 4 + 
Groove-shaped scales - - , si 
Atrio-buccal oppening Separated Separated Common Common Common 
Midgut constrictions + - + ji se 
Gonoducts + - z S E 
Genital opening Paired Unpaired Unpaired Unpaired Unpaired 
Genital papilla - - - y + 
Copulatory stylets + p A 
Copulatory stylets gland + 
Cammera pallial cavity One One One One Two 
Respitary organs - - : Z + 
Dorsoterminal sense organ - + z 


these sclerites: N. carinata, N. trapezi- 
formis Salvini-Plawen, 1978 and N. 
naevata Salvini-Plawen and  Paar- 
Gausch, 2004); presence of anterior cleft 
in ventral pharynx (having ventral cleft: 
N. carinata and N. crenagulata, Salvini- 
Plawen, 1978); presence of separate cere- 
bral connectives (present in: N. oscari 
Salvini-Plawen, 2006, N. simplex Salvini- 
Plawen, 2006 and partly N. herwig1 
Kaiser, 1976); presence of suprapallial 
glands (present in: N. verrilli Heath, 
1918 and N. naevata); presence of midgut 
caecum (present in: N. microsolen Wirén, 
1892); presence of  latero-terminal 
mideut saes (present Um. UN. 
megatrapezata, N. trivialis and N. oscari). 
(GARCÍA-ÁLVAREZ AND URGORRI, 2003a; 
SALVINI-PLAWEN, 1978, 2006; SALVINI- 
PLAWEN, AND PARA-GAUSCH, 2004 see 
Table 1) 

Five species of the genus Neomenia 
can be found in the same biogeographi- 
cal range (García-Álvarez and Urgorri, 
2003a; Salvini-Plawen, 1978, 2006), 
Antarctic Peninsula and  adjacent 
islands, together with Neomenia expleta 
sp nov.. These are N. labrosa Salvini- 
Plawen, 1978; N. laminata Salvini- 
Plawen, 1978; N. monolabrosa Salvini- 
Plawen, 2006; N. megatrapezata; and N. 


trivialis. Each of these species has a 
series of characteristics that clearly dis- 
tinguish them from Neomenia expleta sp 
nov. 

N. labrosa (South Shetland Islands, 
220-240 m. deep), has an anterior cleft in 
the lateral pharynx; it lacks a terminal 
foregut sphincter, a midgut caecum and 
lateroterminal midgut sacs in the diges- 
tive duct; the opening of the spawning 
duct is unpaired; it possesses prepallial 
spines and a subvaginal epithelial gland 
and lacks both seminal vesicles and 
suprapallial glands (SaLvINI-PLAWEN, 
1978, 2006). 

In N. laminata (South Orkney 
Islands, 298-302 m. deep) some anatomi- 
cal data about its posterior part remain 
unknown; it has no pharyngeal lip for- 
mation, is provided with 4 foregut 
regions and lacks a terminal foregut 
sphincter and lateroterminal midgut 
sacs (SALVINI-PLAWEN, 1978, 2006). 

Of N. monolabrosa (South Shetland 
Islands, 80 m deep) only the anterior 
part is known; it has a 30-50 mm 
cuticle, has an anterior cleft in the 
lateral pharynx, 4 foregut regions and 
no midgut caecum (GARCÍA-ÁLVAREZ 
AND URGORRI, 2003a; SALVINI-PLAWEN, 
2006). 


29 


Iberus, 28 (2), 2010 


N. megatrapezata (South Shetland 
Islands, 640-670 m deep) can reach 18 
cm long and has 4 well-marked longitu- 
dinal ridges; it possesses an anterior 
cleft in the lateral pharynx, 4 foregut 
regions and lacks a midgut caecum, the 
opening of the spawning duct is 
unpaired; it has prepallial spines and 
genital papilla and has neither seminal 


vesicles nor  suprapallial 
(SALVINI-PLAWEN, 2006). 

N. trivialis (South Shetland Islands, 
640-670 m deep) has no pharyngeal lip 
formation, lacks a midgut caecum, 
seminal vesicles and suprapallial glands; 
the opening of the spawning duct is 
unpaired and it possesses prepallial 
spines (SALVINI-PLAWEN, 2006). 


glands 


Order STERROFUSTIA Salvini-Plawen, 1978 
Family PHYLLOMENIIDAE Salvini-Plawen, 1978 
Genus Phyllomenia Thiele, 1913 


Phyllomenia sp. (Figs. 3, 4) 


Material examined: 1 specimen, cut in serial sections. Bellingshausen Sea. (Bentart-2006, Station 
MB 34; coordinates: 70.12258” S; 84.8682” W). Water depth: 603 m. Only the posterior part of the 
body could be studied and reconstructed, as the anterior part was histologically strongly damaged. 


Description. Habitus. The specimen 
is elongated, 8 mm long by 0.85 mm 
thick in its middle area, of circular 
section, with an anterior end 0.6 mm 
thick and a posterior one 0.58 mm thick. 
There are no longitudinal keels or 
swellings, the anterior end is truncated 
and the posterior end slightly acumi- 
nated (Fig. 3A). There are well-marked 
atriobuccal cavity, pedal groove and 
pallial cavity. The interwoven sclerites 
do not protrude from the cuticle. Colour 
in alcohol is white. A dorsoterminal 
sense organ is present at the posterior 
end of the body. 

Mantle. The cuticle is thin, 30-40 mm 
thick in the lateral areas of the body and 
50-60 mm thick in the dorsal area, with 
several layers of interwoven tangential 
sclerites. Five types of sclerites can be 
seen (Fig. 3B): solid acicular with 
pointed distal end and rounded proxi- 
mal end slightly sigmoid (200 mm long 
x 15 mm wide); narrow solid paddle- 
shaped sclerites (230 mm long x 15 mm 
wide); wide solid paddle-shaped scle- 
rites (200 mm long x 32 mm wide). Both 
types of solid paddle-shaped sclerites 
are abundant in the ventral area on both 
sides of the pedal groove; elongated and 
wide scales with a very pointed distal 
end and a straight proximal one (100 
mm long x 30 mm wide) located above 


30 


all in its anterior end, atriobuccal cavity 
and pedal groove; blade-shaped scales 
(75 mm long x 13 mm wide). 

Pedal groove and pallial cavity. The 
pedal groove bears a ciliated fold that 
gets into the pallial cavity. The pallial 
cavity is in subterminal position, it is 
small, without cilia on its epithelium 
and has no respiratory folds (Fig. 3C). 
The rectum is short and opens through a 
wide and circular anus located in the 
dorsoanterior wall of the pallial cavity. 
Both spawning ducts come out sepa- 
rately through a pair of narrow grooves 
located on the ventral wall of the pallial 
cavity. It has three pairs of copulatory 
stylets and abdominal spicules in a pair 
of small pouches located on both lateral 
walls of the pallial cavity. 

Reproductive system (Fig. 3C). It has a 
pair of gonads, from which two long 
and narrow real gonoducts (350 mm 
long x 10-25 mm wide) come up, they 
run laterally to the pericardium until 
they join the proximal end of the peri- 
cardioducts (Fig. 4). The pericardium is 
short and wide; anteriorly it is blind, 
posteriorly two blind extensions come 
up from it and it extends dorsolaterally 
on both sides of the rectum, it lacks cilia 
on its interior epithelium and the dorsal 
blood sinus comes out into the dorsoan- 
terior part of the pericardium. The heart 


GARCÍA-ÁLVAREZ ET AL.: Solenogastres from the Bentart-2006 Expedition 


Figure 3. Phyllomenta sp. A: habitus; B: mantle sclerites; C: schematic organization of the posterior 
body. Abbreviations, Asp: Abdominal spicules pouch; Cs: copulatory stylet; Dso: dorsoterminal 
sense organ; Gd: gonoduct; Go: gonad; Ht: heart; Mg: midgut; Pc: pallial cavity; Pd: pericardiod- 
uct; Pr: pericardium; Sd: spawning duct. 

Figure 3. Phyllomenia sp. A: habitus; B: escleritos del manto; C: esquema de la organización de la 
parte posterio del cuerpo. Abreviaturas, Asp: bolsa de espículas abdominales; Cs: estilete copulador; Dso: 
órgano sensitivo dorsoterminal; Gd: gonoducto; Go: gónada; Ht: corazón; Mg: intestino; Pc: cavidad 
paleal; Pd: pericardioducto; Pr: pericardio; Sd: conducto de desove. 


Figure 4.- Phyllomenia sp. Union of gonoduct and pericardioduct. Gd — gonoduct; Pd — pericar- 
dioduct; Pr - pericardium; Re — rectum. 


Figure 4.- Phyllomenia sp. Unión del gonoducto y del pericardioducto. Gd — gonoducto; Pd — pericar- 
dioducto; Pr - pericardio; Re — recto. 


3] 


Iberus, 28 (2), 2010 


hangs from the dorsal wall of the peri- 
cardium; anteriorly it is narrow and 
little lobulate (ventricle), posteriorly it is 
wide and extensively bilobulate (auri- 
cles). The pair of pericardioducts come 
up on both sides of the medial area of 
the pericardium, where they join the 
gonoducts (Fig. 4), they are short and of 
circular section and extend ventrally 
until they come out laterally into the 
spawning ducts, near the opening of 
these ducts into the pallial cavity. Both 
spawning ducts are tubular and nar- 
rower in their anterior half, posteriorly 
they get wider and come out separately 
through a short and narrow duct onto 
the ventral wall of the pallial cavity. It 
has three pairs of copulatory stylets 
located ventrolaterally to the spawning 
ducts, each group bears musculature 
and the three stylets; each is located dor- 
sally to the other; they are short, flat and 
wide (80-100 mm long x 10-12 mm high 
x 40 mm wide). 

Remarks. The organization of the 
anterior part of the body is unknown. 
However, its size (8 mm long, 0.85 mm 
thick) and the combination of sclerites 


and caracters of the posterior part of the 
body identify the specimen hereby 
studied as a juvenile individual of genus 
Phyllomenia. Two species are described 
in the genus Phyllomenia Thiele, 1913 
(GARCÍA-ÁLVAREZ AND SALVINI-PLAWEN, 
2007): Phyllomenia austrina present in 
different locations of the Antarctic 
Ocean (South Sandwich Island, Brans- 
field Strait, Ross Sea and Davis Sea), 
148-465 m deep; and Phyllomenia cornu- 
adentata Salvini-Plawen, 1978, present in 
Tierra del Fuego (South America), 384- 
903 m deep. Both species have a 
complex and very similar organization 
of the pallial cavity and a gonopericar- 
dial system that changes as specimens 
become mature. In young specimens, 
the organization of the posterior part of 
the body is very similar to the rest of 
species of Solenogastres (SALVINI- 
PLAWEN, 1978), while adults develop 
from the pallial cavity and the spawning 
duct, a set of pouches, of which the 
function and relationship to the struc- 
tures they originate from, are only par- 
tially known (SALVINI-PLAWEN, 1970, 
1978; THIELE, 1913). 


Genus Plicaherpia gen. nov. 


Diagnosis. Solid acicular sclerites com- 
bined with hook-shaped solid sclerites and 
elongated groove-shaped scalesina layer. 
With common atrio-buccal opening. 
Midgut with constrictions. Unpaired gen- 
ital opening with genital papilla. Pallial 


cavity with two chambers. Coputatory 
stylets absent. Respiratory organs pre- 
sent. Dorsoterminal sense organ absent. 

Etymology. Latin, plicare = to fold. 
Greek, herpeton = to slither. Concerning 
the fact that it has respiratory folds. 


Plicaherpia papillata sp. nov. (Fig. 5) 


Type material: Holotype: 1 specimen, cut in serial sections. Bellingshausen Sea. (Bentart-2006, 
Station MB 34-2; coordinates: 70.11620* S; 84.8604” W). Water depth: 603 m. Holotype (cut in 
serial sections) deposited in the Museo Nacional de Ciencias Naturales of Madrid, number 


MNCN 15.02/27. 


Etymology: Latin, papilla = papilla; -atus = provided with. Concerning the fact that it has genital 


papilla. 


Diagnosis. Specimen 2 mm x 0.77 
mm, without longitudinal keels or 
swellings. Cuticle 10-20 mm thick. Four 
types of solid sclerites: slightly curved 
acicular; hook-shaped; long and narrow 


32 


groove-shaped scales, with a very 
pointed distal end, and long and wide 
groove-shaped scales. Pedal groove 
with a fold that does not reach into the 
pallial cavity. Radula with 2 medial den- 


GARCÍA-ÁLVAREZ ET ALz.: Solenogastres from the Bentart-2006 Expedition 


S0 um 


Figure 5. Plicaherpia papillosa gen. and sp. nov. A: habitus; B: mantle sclerites; C: radular tooth; D: 
schematic organization of the anterior body; E: schematic organization of the posterior body. 
Abbreviations, At: atrial sense organ; Cg: cerebral ganglion; Gp: genital papilla; Mg: midgut; Oe: 
oesophagus; Pc: pallial cavity; Pd: pericardioduct; Ph: pharynx; Pr: pericardium; Ra: radula; Re: 
rectum; Rf: respiratory folds; Rs: radular sac; Sc: suprarectal commissure; Sd: spawning duct; Sdp: 
spawning duct pouch; Sp: sphincter; Sr: seminal receptacle; Sv: seminal vesicle; Vfg: ventrolateral 
foregut glandular organs. 

Figure 5. Plicaherpia papillosa gen. y sp. nov. A: habitus; B: escleritos del manto; C: diente radular; D: 
esquema de la organización de la parte anterior del cuerpo; E: esquema de la organización de la parte 
posterior del cuerpo. Abreviaturas, At: órgano sensitive atrial; Cg: ganglio cerebral; Gp: papila genital; 
Mg: intestino; Oe: esófago; Pc: cavidad paleal; Pd: pericardioducto; Ph: faringe; Pr: pericardio; Ra: 
rádula; Re: recto; Rf: pliegues respiratorios folds; Rs: saco radular; Sc: comisura suprarrectal; Sd: con- 
ducto de desove; Sdp: bolsa del conducto de desove; Sp: esfínter; Sr: receptáculo seminal; Sv: vesícula 
seminal; Vfe: órgano glandular ventrolateral del intestino. 


39 


Iberus, 28 (2), 2010 


ticles. Oesophagus with sphincter. 
Ventral foregut glandular organs with 
short ducts. Without dorsoanterior 
caecum of the intestine. Vesicles and 
seminal receptacles present. Opening of 
the unpaired spawning duct through a 
genital papilla. Number of respiratory 
folds 15. Pallial cavity with two cham- 
bers: a dorsal one with respiratory folds 
and anus and a ventral one with the 
opening of the spawning duct. Dor- 
soterminal sense organ absent. 

Description. Habitus. The specimen 
is 2 mm long by 0.77 mm thick in its 
medial area, with an anterior end 0.45 
mm thick and a posterior one 0.6 mm 
thick. Without longitudinal keels or 
swellings. There is a well-marked pedal 
groove, the anterior end of the body 
appears truncated due to the anterior 
position of the opening of the atriobuc- 
cal cavity; in the posterior end, the sub- 
terminal opening of the pallial cavity is 
clearly observed (Fig 5A). Sclerites pro- 
trude clearly from the cuticle and point 
posteriorly. Colour in alcohol is white. 

Mantle. The cuticle is thin, 10-20 mm 
thick, with a layer of sclerites in oblique 
arrangement, an inclination of 70” point- 
ing towards the posterior part of the 
body. Four types of sclerites are 
observed (Fig. 5B): solid acicular with a 
pointed distal end and a rounded proxi- 
mal one slightly curved in its distal half 
(200 mm long x 5 mm wide); solid and 
hook-shaped, slightly protruding from 
the cuticle, similar to Ocheyoherpia hook- 
shaped  sclerites  (fragment, only 
observed at stereomicroscope in the 
animal); grooved scales elongated and 
narrow with thick margins, a pointed 
distal end and a rounded proximal one 
(210 mm long x 15 mm wide); and 
grooved scales elongated and wide with 
thick margins, a pointed distal end and 
a rounded proximal one (100 mm long x 
11 mm wide). 

Pedal groove and pallial cavity. The 
pedal groove bears a ciliated fold that 
does not get into the pallial cavity. The 
pallial cavity has a subterminal position 
and bears two chambers divided by a 
septum (Fig. 5E): a dorsal chamber that 
extends anterodorsally, where there are 


34 


15 short and wide respiratory folds with 
a radial arrangement and into which the 
rectum opens; the second chamber is 
ventral, it extends anteroventrally and 
the unpaired spawning duct opens into 
it through a genital papilla. It lacks cop- 
ulatory stylets and abdominal spicles. 
Digestive system (Fig. 5D). The 
common  atriobuccal cavity opens 
frontally in the anterior end of the body, 
with the mouth located in the atrium 
bottom. There is a long pharynx, inter- 
nally folded and encircled by a layer of 
soft circular musculature and a glandular 
layer which is thicker dorsally. It has a 
pair of ventrolateral foregut glandular 
organs tubular, narrow and short, encir- 
cled by subepithelial glands along its 
entire length, of type A (SALVINI-PLAWEN 
1978; HAND AND TopDrT, 2005) they open 
in the radular anterior area. The distic- 
hous radula is made up of pairs of hook- 
shaped teeth (25 mm long) with 2 medial 
denticles (Fig. 5C). It has a ventral 
radular sac wide and short, in which 
several radular teeth can be observed. 
The oesophagus bears a soft sphincter 
and opens frontally into the intestine. The 
intestine lacks a dorsoanterior caecum 
and is provided with serial constrictions 
made up of the dorsoventral muscula- 
ture. The rectum is tubular and narrow, 
located dorsally to the spawning duct. 
The anus opens in the anterior wall of the 
dorsal chamber of the pallial cavity. 
Nervous system and sense organs. 
Anteriorly, only the cerebral ganglion, 
located dorsally to the pharynx and the 
pair of small buccal ganglia located on 
both sides of the pharynx in the area of 
the ventrolateral foregut glandular 
organs, were observed. There is a long 
supra-rectal commissure on the rectum. 
The atrial sense organ is large with 
simple and thick sense papillae. There is 
no dorsoterminal sense organ. 
Reproductive system (Fig. 5E). There is 
a tubular and narrow pair of gonads. In 
the posterior area, the gonads present a 
pair of large seminal vesicles full of sper- 
matozoids. A short pair of gonopericar- 
dioducts of circular section open anteri- 
orly into the pericardium. The tubular 
heart is joined to the dorsal wall of the 


GARCÍA-ÁLVAREZ ET AL.: Solenogastres from the Bentart-2006 Expedition 


anterior part of the pericardium only 
through its anterior and posterior ends. 
The pericardioducts come out from the 
ventroposterior part of the pericardium 
and open dorsally into the seminal recep- 
tacles. The pair of seminal receptacles are 
located laterally in the anterior half of the 
spawning duct and comprise two 
pouches (a smaller anterior one and a 
posterior one) divided by a constriction. 
The posterior pouch of seminal recepta- 
cle opens laterally into the spawning 
duct. The spawning duct is unpaired 
along its entire extension; in its anterior 
half, its section cut is higher that wide 
due to the fact that a ventral pouch comes 
out from it, posteriorly it gets narrower 
till becoming tubular and opens unpaired 
in the centre of the genital papilla. The 
genital papilla is encircled by soft muscu- 
lature and extends ventroposteriorly till 
it opens into the ventral chamber of the 
pallial cavity. Two pouches open directly 
into the genital papilla. Their section cut 
is high and narrow and extends anteri- 
orly on both sides of the spawning duct 
to the medial area of the seminal recepta- 
cles. 

Discussion. The set of features 
present in Plicaherpia gen. nov. defines it 
clearly as a new genus within the family 
Phyllomeniidae Salvini-Plawen, 1978, of 
the order Sterrofustia Salvini-Plawen, 
1978, which is characterized by the com- 


bination of solid sclerites (with hook- 
shaped elements), distichous radula and 
ventrolateral foregut glandular organs 
with subepithelially arranged gland cell 
bodies (type A). (GARCÍA-ÁLVAREZ AND 
SALVINI-PLAWEN, 2007; SALVINI-PLAWEN, 
1978). This family is hitherto composed 
of four genera: Phyllomenia Thiele, 1913, 
Harpagoherpia Salvini-Plawen, 1978, 
Lituiherpia Salvini-Plawen, 1978 and 
Ocheyoherpia Salvini-Plawen, 1978. 

Of the features that define Plicaherpia 
gen. nov., the following are especially 
significant and make it differ from the 
rest of the genera of the family (Table [): 
the groove-shaped scales that can be 
observed on the mantle and the struc- 
ture of the pallial cavity that is divided 
into two chambers, with respiratory 
folds and with the opening of the 
spawning duct through a genital 
papilla. Besides, Phyllomenia is provided 
with a mouth separated from the 
atrium, it has gonads with true gonod- 
ucts, a paired opening of the spawning 
duct and copulatory stylets. Harpagoher- 
pia is provided with a mouth separated 
from the atrium. Lituiherpia has mantle 
sclerites arranged in several layers. 
Ocheyoherpia is provided with copula- 
tory stylets with gland (GARCÍA- 
ÁLVAREZ AND SALVINI-PLAWEN, 2007; 
GARCÍA-ÁLVAREZ AND URGORRI, 2003b; 
SALVINI-PLAWEN, 1978; THIELE, 1913). 


Family ACANTHOMENIIDAE Salvini-Plawen, 1978 
Genus Amboherpia Handl and Salvini-Plawen, 2002 


Amboherpia sp. (Fig. 6) 


Material examined: 1 specimen, cut in serial sections. Bellingshausen Sea. (Bentart-2006, Station 
MB 33; coordinates: 70.28911” S; 84.2841” W). Water depth: 430 m. Only the anterior part of the body 
and a part of the posterior could be studied and reconstructed, as the posterior part of the body 


was histologically damaged. 


Description. Habitus. The specimen 
is 3 mm long by 0.50 mm thick in its 
anterior part, 0.40 mm in its medial part 
and 0.35 mm in its posterior part. There 
are no swellings or longitudinal keels 
(Fig. 6A). The pedal groove is well- 
marked. Sclerites protrude radially from 
the cuticle. Colour in alcohol is white. 


Mantle. The cuticle is thin, 15-20 mm 
thick, without epidermal papillae. The 
sclerite layer has radial insertion, and 
there are three types of sclerites (Fig. 
6B): hollow acicular with both ends 
pointed and slightly curved (235 mm 
long x 10 mm wide); narrow groove- 
shaped scales with a very pointed distal 


35 


Iberus, 28 (2), 2010 


At 


10 um 


100 um 


50 um 


Figure 6. Amboherpia sp. A: habitus; B: mantle sclerites; C: radular teeth; D: schematic organiza- 
tion of the anterior body. Abbreviations, At: atrial sense organ; Bg: buccal ganglion; Cg: cerebral 
ganglion; Mg: midgut; Oe: oesophagus; Ph: pharynx; Ra: radula; Rs: radular sac; Vfg: ventrolateral 
foregut glandular organs. 

Figure 6. Amboherpia sp. A: habitus; B: escleritos del manto; C: dientes radulares; D: esquema de la 
organización de la parte anterior del cuerpo. Abreviaturas, At: órgano sensitive atrial; Bg: ganglio bucal; 


Cg: ganglio cerebral; Mg: intestino; Oe: esófago; Ph: faringe; Ra: rádula; Rs: saco radular; Vfe: órgano 


glandular ventrolateral del intestino. 


end and a slightly rounded one (125 mm 
long x 12.5 mm wide); and wide groove- 
shaped scales with a pointed distal end 
and a rounded proximal one (100 mm 
long x 13 mm wide). 

Pedal groove and pallial cavity. The 
pedal groove bears a ciliated fold that 
does not get into the pallial cavity. The 
pallial cavity is very small, has a subter- 


36 


minal position and lacks both respira- 
tory folds and copulatory stylets. The 
unpaired opening of the spawning duct 
into the pallial cavity could be observed. 

Digestive system (Fig. 6D). There is a 
common atriobuccal cavity with mouth 
located in the posterior part of the 
atrium. The pharynx is encircled by a cir- 
cular layer of musculature and a thicker 


GARCÍA-ÁLVAREZ ET AL.: Solenogastres from the Bentart-2006 Expedition 


glandular layer. The radular sac is wide; 
several broken teeth of a monoserial 
radula were observed in it, possibly with 
two hollow denticles (Fig. 6C). Among 
the broken parts of the radula, several 
boomerang-shaped bases of the teeth 
could be recognized (up to 32 mm long x 
8 mm wide) corresponding to a monoser- 
ial radula; a hollow circular part belong- 
ing possibly to the intermediate part of 
the tooth; and a pair of sharp denticles 
possibly located in the distal end of the 
radular tooth. The ventral foregut glan- 
dular organs consist of two short ducts 
encircled by musculature and opening 
into the pharynx at the level of the 
radula. In the posterior region of each 
duct there are bundles of glandular cells 
(type A according to SALVINI-PLAWEN, 
1978; or type Acanthomenia according to 
HANDL AND TobrT, 2005). It possesses a 
narrow oesophagus that opens dorsally 
into the intestine. The intestine lacks a 
dorsoanterior caecum. 

Nervous system and sense organs. Only 
the cerebral ganglion located dorsally to 
the pharynx could be observed. The 
atrial sense organ is large with numer- 
ous sense papillae simple and very long. 
Despite some reservations, it seems to 
present a dorsoterminal sense organ 
located in the posterior part of the body. 

Remarks. At present three genera of 
the family Acanthomeniidae Salvini- 
Plawen, 1978 are known: Acanthomenia 
Thiele, 1913; Amboherpia Handl and 
Salvini-Plawen, 2002 and Veromenia Gil- 
Mansilla, García-Álvarez and Urgorri, 


ACKNOWLEDGEMENTS 


This paper is part of the research 
projects: Bentart (MEC-Spanish Govern- 
ment REN2003-01881/ANT) and Diva- 


BIBLIOGRAPHY 


GARCÍA-ÁLVAREZ O. AND SALVINI-PLAWEN L.V. 
2007. Species and diagnosis of the Families 
and Genera of Solenogastres (Mollusca). 
Iberus, 25 (2):73-143. 


2008. The three genera differ in the com- 
bination of several characteristics: orga- 
nization of the atrio-buccal cavity; pres- 
ence/absence of radula, presence/ab- 
sence of respiratory folds and pres- 
ence/absence of a dorsoterminal sense 
organ (GIL-MANSILLA ET AL., 2008 see 
table 2; HANDL AND SALVINI-PLAWEN, 
2002; SALVIN-PLAWEN, 1978). The speci- 
men is to be clearly included within the 
genus Amboherpia as it has a common 
atriobuccal cavity, monoserial radula, it 
lacks respiratory folds and has, despite 
some reservations, a dorsoterminal sense 
organ. Two species are described in the 
genus Amboherpia: A. heterotecta Handl 
and Salvini-Plawen, 2002, Bergen (Nor- 
way), 610 m deep and A. dolicopharyngea- 
ta Gil-Mansilla, García-Álvarez and Ur- 
gorri, 2008, Angola Basin, 5415 m deep. 
Although the structure of its reproduc- 
tive system is still unknown Amboherpia 
sp., has a combination of characteristics 
that enable us to distinguish it from the 
known species of the genus (GIL-MAN- 
SILLA ET AL., 2008 HANDL AND SALVINI- 
PLAWEN, 2002). Amboherpia sp. differs 
from A. heterotecta in the fact that: it lacks 
a preradular sphincter, it is provided 
with an oesophagus, and the ventrolater- 
al foregut glandular organs have short 
ducts and a glandular association re- 
stricted to the terminal part. It differs 
from A. dolicopharyngeata in: having only 
groove-shaped scales, the pharynx is 
shorter and not divided in two regions, 
the oesophagus is shorter and lacks an 
intestinal dorso-anterior caecum. 


Artabria II (MEC-Spanish Government 
CTM-2004-00740; Xunta de Galicia 
PGIDITO7PXIB000120PR). 


GARCÍA-ÁLVAREZ O. AND URGORRI V. 2003a. 
Solenogastres molluscs from the BENTART 
Collection (South Shetland Islands, Antarc- 
tica), with a descripción of a new species. 
Iberus, 21 (1):43-56. 


37 


Iberus, 28 (2), 2010 


GARCÍA-ÁLVAREZ O. AND URGORRI V. 2003b. A 
new species of Phyllomeniidae (Mollusca 
Solenogastes: Sterrofustia) from the South 
Shetland Islands, Antarctica. Iberus, 21 (2): 99- 
107. 

GARCÍA-ÁLVAREZ O., ZAMARRO M. AND UR- 
GORRI V. 2009. Proneomeniidae (Solenogas- 
tres, Cavibelonia) from the Bentart-2006 Ex- 
pedition, with description of a new species. 
Iberus, 27 (1): 67-78. 

GiL-MAnNsILLA E., GARCÍA-ÁLVAREZ O. AND UR- 
GORRI V. 2008. New Acanthomeniidae 
(Solenogastres, Cavibelonia) from the abyssal 
Angola Basin. In: Martínez Arbizu, P. and 
Brix, S. (Eds). Bringing Light into Deep-sea 
Biodiversity. Zootaxa, 1866: 175-186. 

GIL-MANSILLA E., GARCÍA- ALVAREZ O. AND UR- 
GORRI V. 2009. A new genus and two new 
species of Simrothiellidae (Solenogastres, 
Cavibelonia) from the Abyssal Angola Basin. 
Journal of the Marine Biological Association of 
the United Kingdom, 89 (7): 1507-1515. 

HANDL C. AND ToDrT C. 2005. Foregut Glands 
of Solenogastres (Mollusca): Anatomy and 
Revised Terminology. Journal of Morphology, 
265: 28-42. 


38 


HANDL C. AND SALVINFPLAWEN L.v. 2002. New 
records of Solenogastres-Cavibelonia (Mol- 
lusca) from Norwegian fjords and shelf wa- 
ters including three new species. Sarsia, 87: 
423-450. 

SALVINI-PLAWEN L.v. 1970. Phyllomenia aust- 
rina ein Phylogenetisch bedeutsamer 
Solenogaster (Mollusca, Aculifera). Zeitschrift 
fir Zoologie, Systematik u. Evolutionsforschung, 
8: 297-309. 

SALVINI-PLAWEN L.v. 1978. Antarktische und 
subantarktische Solenogastres-Eine Mono- 
graphie: 1898-1974. Zoologica, Stuttgart, 128: 
1-315. 

SALVIN-PLAWEN L.v. 2006. Five new Iberian 
Neomeniamorpha (Mollusca, Solenogastres). 
Iberus, 24 (2): 1-26. 

SALVINI-PLAWEN L.V. AND PAAR-GAUSCH l. 
2004. Three new species of Neomenia (Mol- 
lusca, Solenogastres) from the Southern 
Hemisphere. New Zealand Journal of Marine 
and Freshwater Research, 38: 137-162. 

THIELE J. 1913. Antarktische Solenogastren. 
Deutsche Súdpolar Expedition, 14, Zoologie, 
6 (1): 35-65. 


O Sociedad Española de Malacología 


Iberus, 28 (2): 39-50, 2010 


Fauna Malacológica da cidade de Coimbra (Beira Litoral). 
Mo! :<cos “urbanos” de Portugal. 1 


Malacological Fauna from Coimbra (Beira Litoral). Portuguese 


“urban” Molluscs. 1 


Álvaro DE OLIVEIRA* 


Recibido el 21-VIL-2010. Aceptado el 31-X-2010 


RESUMO 


É listada a fauna malacológica da cidade de Coimbra (Beira Litoral, Portugal), compa- 
rando resultados da pesquisa bibliográfica com registos próprios decorrentes de trabalho 
de campo efectuado no decorrer dos últimos dois anos e meio. 197 espécies nominais 
foram citadas, das quais 100 sáo aqui consideradas válidas. Sessenta e uma espécies 
sáo reportadas como ocorrendo actualmente na cidade, uma diversidade considerável, 
tendo em conta o confinamento a que estáo votadas, e a constante e ininterrupta transfor- 
macáo do seu espaco vital. 


ABSTRACT 


The malacological fauna from Coimbra (Beira Litoral, Portugal) is listed, the results of 
bibliographic research compared with own records of field work done in the last two and 
a half years. 197 nominal species were cited, 100 of which here considered as valid 
ones. Sixty-one species are reported as presently living in the town, a notable diversity 
having in mind their confinement, and the constant and uninterrupted transformation of 


their own vital space. 


INTRODUCAO 


Nas margens do Rio Mondego e a 
cerca de 45 km da sua foz, a cidade de 
Coimbra, capital da província de Beira 
Litoral, está situada no limite oriental da 
Bacia Lusitánica, caracterizada pela natu- 
reza sedimentar, calcária, dos seus terre- 
nos. Á sua localizacáo estabelece a transi- 
cáo entre o curso superior do Rio, alcanti- 
lado e serrano, e o início do extenso e 
amplo vale, seu remoto estuário. 

A fauna malacológica terrestre e 
aquática é reportada, confrontados 
registos clássicos com resultados de tra- 


balho de campo efectuado pelo autor 
nos anos de 2008, 2009 e 1” semestre de 
2010. 

“Arredores de...” ou “environs 
de...”, expressóes  frequentemente 
usadas na bibliografia antiga, apontam 
sítios actualmente absorvidos pelo cres- 
cimento da cidade: as espécies estáo 
hoje em dia contidas em pequenos 
“ghettos” urbanos, jardins e parques 
públicos, dominados pelo exotismo da 
sua Flora. Em alguma medida este é um 
relato de extincó0es. 


* Av. Lagos, 219-Y, 4405-658 Gulpilhares, Portugal. deoliveira1004Csapo. pt 


39 


Iberus, 28 (2), 2010 


MATERIAL E MÉTODOS 


O trabalho de campo foi efectuado 
no decorrer dos últimos dois anos e 
meio (2008, 2009, 1” semestre de 2010), 
em oito sítios caracterizados na Tabela 1, 
na qual é mostrada a distribuicáo das 
espécies na cidade. Trés desses sítios 
(Jardim Botánico, Jardim de Santa Cruz 
e Quinta das Lágrimas, Figs. 1-2), pela 
diversidade faunística ¡inicialmente 
detectada, foram objecto de um maior 
esforco de amostragem: durante o ano 
de 2009, para além da amostragem 
directa, foram  colhidas  amostras 
mensais de manta morta e camada 
superficial do solo, sujeitas a imediata 
triagem. 

Todas as amostras (excepc0es apon- 
tadas) estáo depositadas na coleccáo do 
autor. A sua determinacáo específica foi 
baseada em caracteres conquiliológicos, 
e, no caso das lesmas, na sua morfología 
externa. Particularmente quanto a estas 
últimas, a sua classificacáo, náo baseada 
em estudo anatómico, foi necessaria- 
mente conservadora (exemplos: o con- 
ceito de ambas Deroceras laeve (Miller O. 
F., 1774) e Arion nobreí Pollonera, 1889 
poderá acolher mais do que um taxon). 

Foi também estudado material 
museológico, quase em exclusivo prove- 
niente das coleccóes depositadas no 
Museu de História Natural da Universi- 
dade de Coimbra, com vista á confirma- 
cáo de alguns registos. 

Paralelamente foi realizado um 
extenso trabalho de compilacáo biblio- 
gráfica, tendo como base, essencial- 
mente, cinco obras de referéncia: 
MORELET (1845), HIDALGO (1875), 
LOCARD (1899), NOBRE (1885, 1930). 
Estes registos sá0 apontados na Tabela II 
(colunas 1 a 4). Na mesma Tabela sáo 
reportados os registos próprios (coluna 
5) e Os primeiros registos para Coimbra 
(coluna 6) quando posteriores a NOBRE 
(1930). 

Na Tabela III é dada a lista de sinoní- 
mias, particularmente pertinente para 
compreender os registos de LOCARD 
(1899), que para Coimbra cita 113 espé- 
cies, das quais apenas 58 sáo aqui consi- 
deradas válidas. 


40 


RESULTADOS E DISCUSSAO 


Moluscos de Coimbra 

(1) Sáo 197 as espécies nominais de 
moluscos terrestres e aquáticos citadas 
para Coimbra (Tabelas II e II), desde 
meados do séc. XIX até a actualidade. 

(11) 100 espécies sáo consideradas 
válidas (Tabela IM), das quais 61 (61%) 
foram detectadas pelo autor (Tabela I). 

(111) Do conjunto dos registos clássi- 
cos (colunas 1-4, Tabela Il) sáo reconhe- 
cidas 76 espécies, 41 (54%) encontradas 
no presente estudo. Áquelas acrescem 
24 registos posteriores, nove dos quais 
aqui apontados como novos para 
Coimbra: Ferrissia fragilis, Carychium 
minimum, Spermodea lamellata, Vertigo 
pygmaea, Zonitoides nitidus, Lehmannia 
valentiana, Microxeromagna lowei, Otala 
lactea, Corbicula fluminea. 

(iv) O grupo no qual é mais patente 
o número de espécies entretanto desapa- 
recidas é sem dúvida os Bivalvia. Das 
oito espécies apontadas apenas uma 
(13%) foi encontrada pelo autor: a 
espécie exótica Corbicula  fluminea 
(conchas e valvas roladas). 

(v) Entre os Gastropoda o grupo 
mais afectado é o dos caracóis aquáticos: 
apenas 10 (42%) das 24 espécies foram 
detectadas. Na Superfamilia Planorboi- 
dea o cenário é ainda mais preocupante: 
das 13 espécies citadas apenas quatro 
(31%) foram recentemente observadas, 
das quais trés exóticas: Planorbella duryi, 
Ferrissia fragilis, Physa acuta. 

(vi) Quanto aos Gastropoda terres- 
tres é de notar o aparente desapareci- 
mento dos dois representantes da 
Familia  Chondrinidae,  Granopupa 
granum e Chondrina lusitanica, particular- 
mente esta última, endemismo da Bacia 
Lusitánica, em franca regressáo em toda 
a área de distribuicáo (registos próprios 
náo publicados). 

(vii) Das 61 espécies detectadas no 
presente estudo (Tabela 1), 10 (16%) 
foram observadas num único sítio: Theo- 
doxus cf. fluviatilis, Belgrandia lusitanica, 
Planorbella duryi, Vallonia cf. enniensis, 
Spermodea  lamellata, Balea  heydeni, 
Limacus flavus, Microxeromagna loweli, 
Otala lactea, Corbicula fluminea. 


DE OLIVEIRA: Fauna Malacológica de Coimbra. Moluscos “urbanos” de Portugal. 1 


CA 
ts BOO 
st TO AON 
RE a rre 
1 


Figuras 1, 2. Coimbra. 1: Margem direita do Rio Mondego e Mata do Jardim Botánico, 21-IV- 
2009. 2: Rio Mondego e Quinta das Lágrimas (margem esquerda), 10-XI1-2009. 

Figures 1, 2. Coimbra. 1: Right bank of Mondego River and Wood of the Botanical Garden, 21-IV- 
2009. 2: Mondego River and Quinta das Lágrimas (left bank), 10-X11-2009. 


41 


Iberus, 28 (2), 2010 


Tabela I. Lista das espécies observadas/colhidas em Coimbra (registos próprios). X. Presenca. (X). 
Apenas conchas. Sítios [UTM 1km]: 1. Jardim Botánico [NE4950]. 2. Jardim de Santa Cruz 
[NE4951]. 3. Penedo da Saudade [NE4950]. 4. Vale da Ribeira de Coselhas [NE4752-4852]. 5. 
Margens do Mondego [NE4849-4949-4850-4950-4851-5148]. 6. Instituto Geofísico [NE5051]. 
7. Penedo da Meditacáo [NE5052]. 8. Quinta das Lágrimas [NE4849]. 

Table I. List of species noticed/collected in Coimbra (own records). X. Presence. (X). Shells only. Sites 
[UTM 1km]: 1. Botanical Garden [NE4950]. 2. Santa Cruz Garden [NE4951]. 3. Penedo da 
Saudade [NE4950]. 4. Ribeira de Coselhas Valley [NE4752-4852]. 5. Mondego River banks 
[NE4849-4949-4850-4950-4851-5148]. 6. Instituto Geofísico [NES051]. 7. Penedo da Meditacáo 
[NE5052]. 8. Quinta das Lágrimas [NE4849). 


Espécies 1 2 3 4 5 6 / 


Theodoxus cf. fluviatilis (Linnaeus, 1758) 

Pomatias elegans (Miller O. E, 1774) X X 

Potamopyrgus antipodarum (Gray J. E., 1843). X X X (1) 

Mercuria tachoensis (Fravenfeld, 1865) X 

Belgrandia lusitanica (Palodilhe, 1867) 

Galba truncatula (Miller O. E, 1774) X 

Radix balthica (Linnaeus, 1758). 0) 
X 
X 


> >>> [o -] 


o 
=_— 


Planorbella duryi (Wetherby, 1879) 

Ferrissia fragilis (Tryon, 1863) 

Ancylus fluviatilis (Miller O. E, 1774) X X 
Physa acuta Drapamaud, 1805 X X X 

Carychium ibazoricum Bank e Gittenberger, 1985  X X X X 
Carychium minimum Múller O. E, 1774 X 
Oxyloma elegans (Risso, 1826) 

Cochlicopa lubrica (Miller 0. E, 1774) 

Cochlicopa lubricella (Rossmássler, 1834) X 
Lauria cylindracea (da Costa, 1778) X 
Leiostyla anglica (Férussac, 1821) X 
Vallonia costata (Miller O. E, 1774) 

Vallonia cf. enniensis (Gredler, 1856) 

Vallonia pulchella (Múller O. E, 1774) 

Plagyrona placida (Shuttleworth, 1852) X 
Acanthinula aculeata (Múller O. E, 1774) X 
Spermodea lamellata (Jeffreys, 1830) X 
Truncatellina cylindrica (Férussac A., 1807) X X 
Vertigo pygmaea (Draparnaud, 1801) X 

Clousilia bidentata (Stróm, 1765) X X X 0) 

Balea heydeni Maltzan, 1881 ( 
Rumina decollata (Linnaeus, 1758) 
Testacella maugei (de Férussac, 1819) 
Punctum pygmaeum (Draparnaud, 1801) 
Paralaoma servilis (Shuttleworth, 1852) 
Discus rotundatus (Miller O. E, 1774) 
Vitrea contracta (Westerlund, 1871) 
Euconulus fulvus (Miller O. F., 1774) 
Zonitoides nitidus (Múller O. F., 1774) X X 
Oxychilus cellarius (Miller O. E, 1774) X X (X 
Oxychilus draparnaudi (Beck, 1837) X X (0) 0) 09) X X 
Aegopinella nitidula (Draparmaud, 1805) X X X X X X 


>< 
>< 
>< 
>< 
>< 
>< >< >>> >< 


>< >> >> > 
>< 
>< 


>< >>> 


> >>> > 
>< >>>> 
>< 
>< 
== 
== 


42 


DE OLIVEIRA: Fauna Malacológica de Coimbra. Moluscos “urbanos” de Portugal. 1 


Tabela 1. Continuacáo. 
Table I. Continuation. 


Espécies 1 


Milax gar=*es (Draparnaud, 1801) 
Limax maxitnus ¡Linnaeus, 1758) 


Limacus flavus (Linnaeus, 1758) X 
Lehmannia valentiana (Férussac, 1822) 

Deroceras laeve (Miller O. E, 1774) X 
Deroceras reticulatum (Miller O. E, 1774) X 
Arion ater (Linnaeus, 1758) 

Arion intermedivs Normand, 1852 X 
Arion nobrei Pollonera 1889 X 
Cochlicella barbara (Linnaeus, 1758) 

Oestophora barbula (Rossmússler, 1838) X 


Oestophora lusitanica (Pfeiffer, 1841) 

Ponentina subvirescens (Bellamy, 1839) 

Candidula olisippensis (Servain, 1880) X 
Cernuella virgata (da Costa, 1778) 
Microxeromagna lowei (Potiez e Michaud, 1838) 
Theba pisana pisana (Múller O. E, 1774) 

Cepaea nemoralis (Linnaeus, 1758) X 
Otala lactea (Miller O. F, 1771) 

Portugala inchoata (Morelet, 1845) 


>< 


Cornu aspersum (Miller O. E, 1774) X 
Corbicula fluminea (Miller O. F., 1774) 
61 spp. 35 


(viii) De quatro das espécies apenas 
foram detectados vestígios (conchas 
roladas): Radix balthica, Balea heydeni, 
Otala lactea, Corbicula fluminea. 


Comentários sobre algumas espécies 

Assiminea eliae: É surpreendente o 
registo de LOCARD (1899) desta espécie 
de águas salobras. No entanto, uma 
outra espécie que compartilha o mesmo 
habitat, Myosotella myosotis, é citada 
para Coimbra por CARVALHO (1945) [ver 
“Material museológico” abaixo]. Recen- 
temente o autor encontrou M. myosotis 
associada a Assiminea cf. grayana 
FLEMING, 1828 na Ria de Aveiro, a cerca 
de 20 km do mar (registos próprios náo 
publicados). 

Mercuria tachoensis: Foi detectada 
uma colónia inédita deste endemismo 
portugués na Fonte da Sereia (Jardim de 
Santa Cruz). A sua identidade foi confir- 


>< >>> 


34 


X X 
X X X 
X X X 
X X X X X 
X 
X X X X X 
X X 
X X X 
X X X X X 
X 00 
X X X 
X X X X 
X X 
A 
X X 00 X X 
X X 00 X X 
09) 
X X X 
00 X X X X X 


22 24 18 2| 33 


mada por estudo anatómico (seis espéci- 
mes na coll. Emilio Rolán). 

Ferrissia fragilis: Registo novo para 
Coimbra desta espécie exótica recente- 
mente referenciada para Portugal 
(HOLYOAK, 2009). Foram entretanto 
detectadas mais duas colónias da 
espécie: No Vale do Baixo Mondego, 
numa nascente em Vila Nova da Barca, 
Beira Litoral; e num jardim público de 
Faro, Algarve (registos próprios náo 
publicados). 

Vallonia cf. enniensis: Esta forma foi 
detectada num único local da cidade, 
Jardim de Santa Cruz, associada a Vallo- 
nia costata e Vallonia pulchella. Apenas 
foram encontrados dois exemplares de 
V. cf. enniensis em oito colheitas com 
representantes do genus (58 exemplares 
de V. costata, sete de V. pulchella); náo 
foram detectados exemplares intermé- 
dios. 


A3 


Iberus, 28 (2), 2010 


Tabela II. Lista das espécies citadas para Coimbra. X. Presenga. (X). Apenas conchas. (+). Registos 
novos. * Espécies comentadas em “Resultados”. Registos: 1. MORELET (1845). 2. HIDALGO 
(1875). 3. LOCARD (1899). 4. NOBRE (1885, 1930). 5. Registos próprios (espécies inscritas a 
negrito). 6. Primeiros registos posteriores a NOBRE (1930): (a) BOETERS (1988). (b) DE OLIVEIRA 
(2009a). (c) CARVALHO (1945). (d) DE OLIVEIRA (2009b). (e) MARTÍNEZ-ORTÍ (2006). (f) Cas- 
TILLEJO (1990b). (g) CARVALHO (1944). (h) CASTILLEJO (19904). 

Table II. List of species noticed/collected in Coimbra. X. Presence. (X). Shells only. (+). New records. (*) 
Species commented under “Resultados”. Records: 1. MORELET (1845). 2. HIDALGO (1875). 3. LOCARD 
(1899). 4. NOBRE (1885, 1930). 5. Own records (species inscribed in bold). 6. First records after 
NOBRE (1930): (4) BOETERS (1988). (b) DE OLIVEIRA (2009a). (c) CARVALHO (1945). (4d) DE OLr- 
VEIRA (2009b). (e) MARTÍNEZ-ORTÍ (2006). (f) CASTILLEJO (1990b). (g) CARVALHO (1944). (h) 
CASTILLEJO (19904). 


Espécies | 2 


Theodoxus cf. fluviatilis (Linnaeus, 1758) X 

Pomatias elegans (Miller O. F., 1774) X X 
Bithynia tentaculata (Linnaeus, 1758) 

Assiminea eliae Paladilhe, 1875 * 

Potamopyrgus antipodarum (Gray J. E., 1843) X X (0) 
Mercuria tachoensis (Frauenfeld, 1865) * 

Belgrandia lusitanica (Polodilhe, 1867) X 
Valvata piscinalis (Miller O. E, 1774) X 
Galba truncatula (Miller O. F,, 1774) 

Radix balthica (Linnaeus, 1758) 

Radix auricularia (Linnaeus, 1758) 

Bulinus truncatus contortus (Michaud, 1829) X 
Planorbarius metidiensis (Forbes, 1838) * 

Planorbella duryi (Wetherby, 1879) X X (b) 
(+) Ferrissia fragilis (Tryon, 1863)* X 

Planorbis planorbis (Linnaeus, 1758) 

Planorbis carinatus (Múller O. E, 1774) X 
Anisus spirorbis (Linnaeus, 1758) X 
Gyraulus albus (Múller O. E, 1774) 

Gyraulus loevis (Alder, 1838) 

Gyraulus crista (Linnaeus, 1758) 

Hippeutis complanatus (Linnaeus, 1758) 

Ancylus fluviatilis (Miller O. F, 1774) 

Physa acuta Draparnaud, 1805 

Myosotella myosotis (Draparaud, 1801) * X (c) 
Carychium ibazoricum Bonk e Gittenberger, 1985 X X 
(+) Carychium minimum Miiller O. F, 1774 

Oxyloma elegans (Risso, 1826) 

Cochlicopa lubrica (Miller O. F., 1774) 

Cochlicopa lubricella (Rossmissler, 1834) 

Lauria cylindracea (da Costa, 1778) X X X 
Leiostyla anglica (Férussac, 1821) X 

Vallonia costata (Miller O. F., 1774) 

Valonia cf. enniensis (Gredler, 1856) * 

Vallonia pulchella (Miller O. F, 1774) 

Plagyrona placida (Shuttleworth, 1852) * X 
Acanthinula aculeata (Miller O. F., 1774) X X 


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44 


DE OLIVEIRA: Fauna Malacológica de Coimbra. Moluscos “urbanos” de Portugal. 1 


Tabela ll. Continuacáo. 
Table 1. Continuation. 


Espécies 


(+) Spermodea lamellata (Jeffreys, 1830) * 
Pupilla muscorum (Linnaeus, 1758) 
Pyramidula rupestris (Draparnaud, 1801) * 
Granopupa granum (Draparnaud, 1801) 
Chondrina lusitanica (Pfeiffer, 1848) * 
Truncatellina cylindrica (Férussac A., 1807) 
(+) Vertigo pygmaea (Drapamaud, 1801) * 
Merdigera obscura (Múller O. E, 1774) 
Clousilta bidentuta (Stróm, 1765) 

Balea heydeni Maltzan, 1881 

Balea perversa (Linnaeus, 1758) * 
Ferussacia folliculus (Gmelin, 1791) 
Cecilioides acicula (Miller O. F, 1774) 
Rumina decollata (Linnaeus, 1758) 
Testacella maugei (de Férussac, 1819) 
Punctum pygmaeum (Draparnaud, 1801) 
Paralaoma servilis (Shuttleworth, 1852) 
Discus rotundatus (Miller 0. F, 1774) 
Vitrea contracta (Westerlund, 1871) 
Vitrea crystallina (Miller O. E, 1774) 

Euconulus fulvus (Múller O. F, 1774) 

(+) Zonitoides nitidus (Miller O. F, 1774) * 
Oxychilus cellarius (Miller O. F, 1774) 

Oxychilus drapamaudi (Beck, 1837) 

Aegopinella nitens (Michaud, 1831) 

Aegopinella nitidula (Draparnaud, 1805) 

Parmacella valenciennii (Webb e Van Beneden, 1836 * 
Milax gagates (Draparmaud, 1801) 

Limax maximus (Linnaeus, 1758) 

Limacus flavus (Linnaeus, 1758) 

(+) Lehmannia valentiana (Férussac, 1822) 
Deroceras agreste (Pollonera, 1891) 

Deroceras loeve (Miller O. F., 1774) 

Deroceras lombricoides (Morelet, 1845) 

Deroceras reticulatum (Miller O. F., 1774) 

Arion ater (Linnaeus, 1758) 

Arion nobrei Pollonera 1889 

Arion intermedivs Normand, 1852 

Cochlicella acuta (Miller O. F., 1774) 

Cochlicella barbara (Linnaeus, 1758) 

Oestophora barbula (Rossmissler, 1838) 

Oestophora lusitanica (Pfeiffer, 1841) 

Ponentina subvirescens (Bellamy, 1839) 

Xerotricha apicina (Lamarck, 1822) 

Xerotricha conspurcata (Draparnaud, 1801) 

Candidula belemensis (Servain, 1880) 

Candidula intersecta (Poiret, 1801) 


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45 


Iberus, 28 (2), 2010 


Tabela Il. Continuacáo. 
Table IT. Continuation. 


Espécies 1 


Candidula olisippensis (Servain, 1880) 

Cemuella virgata (da Costa, 1778) 

(+) Microxeromagna lowei (Potiez e Michaud, 1838) 
Theba pisana pisana (Miller O. F, 1774) 

Cepaea nemoralis (Linnaeus, 1758) 

(+) Otala lactea (Miller O. E, 1771) 

Portugala inchoata (Morelet, 1845) 

Cornu aspersum (Miller O. E, 1774) 

Potomida littoralis (Cuvier, 1798) * 


Unio delphinus Spengler, 1793 * X 
Anodonta anatina (Linnaeus, 1758) * X 


(+) Corbicula fuminea (Miller O. F., 1774) 
Sphaerium corneum (Linnaeus, 1758) * 
Musculium lacustre (Múller O. F., 1774) 
Pisidium amnicum (Miller O. F, 1774) * 
Pisidium casertanum (Poli, 1791) 


100 spp. 12 spp. 


V. cf. enniensis foi também colhida 
num jardim da cidade do Porto, Douro 
Litoral, associada a V. pulchella [registos 
próprios náo publicados]. Aqui, numa 
amostra de nove exemplares, trés sáo 
atribuíveis a V. cf. enniensis. Dos restan- 
tes seis exemplares, atribuíveis a V. pul- 
chella, pelo menos um apresenta caracte- 
res intermédios: as primeiras voltas da 
espira com estriacáo axial típica de V. 
pulchella, que se vai gradualmente espa- 
cando, até que no último quarto de volta 
é patente já a estriacáo axial típica de V. 
enniensis. 

GIUSTI E MANGANELLI (1992) póem 
em causa a validade da maior parte das 
espécies europeias de Vallonia (entre as 
quais V. enniensis) considerando: (1) 
estáo baseadas apenas em caracteres 
conquiliológicos; (2) as populacóes de 
quase todas as espécies sáo completa- 
mente desprovidas de complexo penial; 
(3) o facto de várias “espécies” ocorre- 
rem associadas no mesmo local [p. ex. 
BECKMANN (2007) reporta V. pulchella, V. 
enniensis e V. costata associadas em dois 
locais da ilha de Mallorca, Baleares], 


46 


2 3 4 5 6 
X X 
X X 
X 
X X X X 
X X X 
0) 
X X X 
X X X 
X (9) 
X 
X X 
0) 
X X 
X 
X (9) 
X X 
l3spp.  58spp.  64spp. | 6lspp. 15 spp. 
76 spp. 


apresentada por vários autores como 
prova suficiente da sua validade taxonó- 
mica, pode náo bastar: poderá tratar-se 
de populacóes de diferentes estirpes 
genéticas ou clones da mesma espécie, 
perpetuadas por autofecundacáo ou 
partenogénese. 

Plagyrona placida: Uma das espécies 
terrestres mais raras na cidade [rara- 
mente obtido mais que um exemplar em 
qualquer das amostragens realizadas 
nos trés sítios onde foi encontrada 
(Tabela 1)]. 

Spermodea lamellata: Registo novo 
para Coimbra desta espécie rara da 
fauna portuguesa, da qual apenas sáo 
conhecidas mais trés populacóes, conti- 
das na Bacia Lusitánica, províncias de 
Beira Litoral e Estreemadura (GITTENBER- 
GER, 1989; DE OLIVEIRA, 2007). 

Vertigo pygmaea: Registo novo para 
Coimbra e para a província de Beira 
Litoral. Conhecida apenas das províncias 
de Minho e Douro Litoral nos registos clás- 
sicos, foi recentemente encontrada em Bra- 
ganca, província de Trás-os-Montes (regis- 
tos próprios náo publicados). 


DE OLIVEIRA: Fauna Malacológica de Coimbra. Moluscos “urbanos” de Portugal. 1 


Zonitoides nitidus: Registo novo para 
Coimbra e para a província de Beira 
Litoral. Desta espécie, até agora apenas 
conhecida da província de Douro Litoral 
(DE OLIVEIRA, 2008), foram entretanto 
detectadas populacóes em Valenca 
(prov. Minho) e Braganca (prov. Trás-os- 
Montes) [registos próprios náo publica- 
dos]. 


Material museológico 

Em Fevereiro de 2010 foram estuda- 
dos vários lotes de Gastropoda e Bival- 
via náo-marinhos provenientes de 
Coimbra depositados no Museu de His- 
tória Natural da Universidade de 
Coimbra (MHNC) reportados no Catá- 
logo da Coleccáo de Invertebrados de 
Portugal (CARVALHO, 1944, 1945). 
Nenhuma das amostras possui data de 
colheita e apenas quatro apontam o 
nome do autor: 

Planorbarius metidjensis: Dois lotes, um 
dos quais (131b) identificado como “Pla- 
norbis corneus (L.)”, o outro (132) como 
“Planorbis corneus (L.) var. metidjensis 
Forbes”, compostos por, respectivamente, 
cinco e duas conchas. Biometria (em mm; 
conchas maiores de cada uma das amos- 
tras): altura — 6,6-7,3; diámetro — 14,0-16,3; 
n” de voltas — 3,8-4,4. A ocorréncia em Por- 
tugal continental de Planorbarius corneus 
(LINNAEUS, 1758) náo foi confirmada. O 
estudo da populacáo reportada do Jardim 
Botánico da Universidade de Coimbra 
revelou tratar-se de Planorbella duryi (WEr- 
HERBY, 1879), espécie exótica de origem 
neárctica (DE OLIVEIRA, 2009a). 

Myosotella myosotis: Este é o único 
vestígio da ocorréncia em Coimbra 
desta espécie de águas salobras. O lote 
estudado (117a) é composto por trés 
conchas com os restos secos do animal. 
Biometria: altura — 6,9-7,8; diámetro — 
3,2-3,6; n* de voltas — 7,0. 

Pyramidula rupestris: O único lote (49b) 
procedente de Coimbra é composto por 
sete conchas, cinco das quais em muito 
mau estado de conservacáo. O estudo das 
duas conchas em melhor estado revelou 
tratar-se de Plagyrona placida [ver também 
“Registos duvidosos ou errados” abaixo]. 
Biometria (concha maior): altura — 1,6; diá- 
metro — 2,2; n” de voltas — 3,1. 


Chondrina lusitanica: O lote estudado 
(92a), identificado como “Abida secale 
(Drap.)” é composto por duas conchas. 
Biometria: altura — 7,6-8,1; diámetro — 
2,8-3,0; n* de voltas — 7,0-7,5. 

Balea perversa: A amostra (98c) é com- 
posta por sete conchas pertencentes a 
juvenis de Clausilia bidentata! Biometria 
(concha maior): altura — 4,8; diámetro — 
2,0; n* de voltas — 8,0. A ocorréncia de 
Balea perversa em Portugal continental é 
apenas suportada por material conqui- 
liológico depositado no Museo de Cien- 
cias Naturales de Madrid [uma amostra 
(pars) procedente de Coimbra na coll. 
Azpeitia (MARTÍNEZ-ORTÍ, 2006)] e no 
Museu de História Natural da Universi- 
dade do Porto [uma amostra prove- 
niente da Serra de Montesinho, prov. de 
Trás-os-Montes, na coll. Nobre (registos 
próprios náo publicados)]. 

Parmacella valencienni: A amostra 
(17a) náo foi encontrada. A espécie é 
classicamente reportada em Portugal do 
Vale do Tejo para sul (MORELET, 1845; 
NOBRE, 1930). Mais recentemente, a sua 
área de distribuicáo é ampliada para 
norte, até ao Vale do Mondego (RODRÍ- 
GUEZ, HERMIDA E OUTEIRO, 1993). No 
entanto, a sua ocorréncia actual em 
Coimbra náo foi confirmada. 

Potomida littoralis: O lote (431) é com- 
posto por trés conchas. Biometria 
(concha maior): altura - 37,8; compri- 
mento — 55,6; espessura — 21,8. Este é o 
único registo conhecido para Coimbra. 

Unio delphinus: A amostra (430) iden- 
tificada como “Unio pictorum (L.)” é 
constituído por cinco conchas. Biometria 
(concha maior): altura — 36,3; compri- 
mento — 72,0; espessura — 25,7. 

Anodonta anatina: A amostra (432) 
identificada como “Anodonta cygnea 
(L.)” é constituído por uma concha. Bio- 
metria: altura — 51,0; comprimento -— 
94,3; espessura — 28,5. 

Pisidium amnicum: A amostra (485) 
identificada como “Pisidium amnicum 
(Múller)” é constituída por trés conchas, 
as duas menores pertencentes a Sphae- 
rium corneum. Biometria: P. amnicum: 


altura — 8,9; comprimento - 11,0; espes- 


sura — 6,2; S. corneum: altura - 6,7-7,1; 
comprimento — 7,7-8,5; espessura — 4,8- 


47 


Iberus, 28 (2), 2010 


Tabela III. Lista de sinonímias: (1). MORELET (1845). (2). HIDALGO (1875). (3). LOCARD 


(1899). (4). NOBRE (1885, 1930). 


Table II. Sinonymy list: (1). MORELET (1845). (2). HIDALGO (1875). (3). LOCARD (1899). (4). 


NOBRE (1885, 1930). 
Espécies 


Theodoxus cf. fluviatilis (Linnaeus, 1758) 
Pomatias elegans (Múller O. E, 1774) 
Bithynia tentaculata (Linnaeus, 1758) 
Assiminea eliae Paladilhe, 1875 

Mercuria tachoensis (Frauenfeld, 1865) 
Belgrandia lusitanica (Paladilhe, 1867) 
Valvata piscinalis (Miller O. E, 1774) 
Galba truncatula (Múller O. E, 1774) 
Radix balthica (Linnaeus, 1758) 


Radix auricularia (Linnaeus, 1758) 
Bulinus truncatus contortus (Michaud, 1829) 
Planorbarius metidiensis (Forbes, 1838) 


Planorbis planorbis (Linnaeus, 1758) 
Planorbis carinatus (Múller 0. E, 1774) 
Anisus spirorbis (Linnaeus, 1758) 
Gyraulus albus (Múller O. E, 1774) 
Gyraulus laevis (Alder, 1838) 

Gyraulus crista (Linnaeus, 1758) 
Hippeutis complanatus (Linnaeus, 1758) 
Ancylus fluviatilis (Miller O. E, 1774) 
Physa acuta Draparmaud, 1805 


Carychium ibazoricum Bank e Gittenberger, 1985 


Oxyloma elegans (Risso, 1826) 
Cochlicopa lubrica (Miller O. E, 1774) 
Lauria cylindracea (da Costa, 1778) 
Leiostyla anglica (Férussac, 1821) 
Plagyrona placida (Shuttleworth, 1852) 
Acanthinula aculeata (Múller O. F., 1774) 
Pyramidula rupestris (Draparnaud, 1801) 
Granopupa granum (Draparnaud, 1801) 
Chondrina lusitanica (Pfeiffer, 1848) 
Truncatellina cylindrica (Férussac A., 1807) 
Merdigera obscura (Múller O. E, 1774) 
Clousilia bidentata (Stróm, 1765) 

Balea perversa (Linnaeus, 1758) 
Ferussacia folliculus (Gmelin, 1791) 
Cecilioides acicula (Miller O. E, 1774) 
Rumina decollata (Linnaeus, 1758) 
Punctum pygmaeum (Draparaud, 1801) 
Paralaoma servilis (Shuttleworth, 1852) 
Vitrea contracta (Westerlund, 1871) 
Oxychilus cellarius (Miller O. E, 1774) 
Oxychilus draparaudi (Beck, 1837) 


48 


Sinónimos 


Neritina violacea (1, 4), Theodoxia elongatula (3), Neritina fluviatilis (4) 
Cyclostoma elegans (1, 2, 3, 4) 

Bythinia tentaculata (3, 4), B. decipiens, B. matritensis (3) 

Assimina eliae (3) 

Amnicola lanceolata, A. castroiana (3), A. similis, Hydrobia similis (4) 
Paludina gibba (1), Belgrandia gibba, Hydrobia gibba (4) 

Valvata tolosana, V. eximia (3) 

Limnaea truncatula (3, 4) 

Limnaea acutalis, L. acronica, L. limosa, L. angustana, L. paulinoi, L. rubella, 
L lusitanica (3), L. peregra, L ovata (4) 

Limnaea auricularia (3, 4), L. intermedia (3) 

Physa contorta (1), P brondeli, P. paulinoi, P castroi (3), Bulinus contortus (4) 
Planorbis metidjensis, P. algericus, P aclopus, P lusitanus, P rosai (3), P dufouri 
(3, 4), P coreus var. metidiensis (4) 

anorbis umbilicatus, P intermedivs (3) 

anorbis complanatus var. carinata (4) 

anorbis leucostoma (1, 4), P rotundatus (3), P spirorbis (4) 

anorbis albus (3, 4) 

Planorbis glaber (3) 

Planorbis imbricatus (3), P crista, P nautileus (4) 

Planorbis fontanus (3, 4), P. complanatus (4) 

Ancylus strictus, A. gibbosus (3), A. striatus (4) 

Physa subopaca (3) 

Auricula gracilis (1), Carychium gracile (2, 3, 4) 

Succinea longiscata (3), S. elegans, S. pfeifferi (4) 

Zua subcylindrica (3), Cionella subcylindrica (4) 

Pupa umbilicata (2, 4), Pupilla umbilicata (3) 

Pupilla paulinoi (3) 
Helix debauxiana (3) 
Helix aculeata (3, 4) 
Helix rupestris (3, 4) 
Pupa graniformis (3), P. granum (4) 

Pupa lusitanica (3), P. avenacea var. lusitanica (4) 

Vertigo muscorum (4) 

Bulimus obscurus (3), Buliminus obscurus (4) 

Clousilia rugosa (1, 2, 4), C. moniziana (3) 

Balia perversa (3) 

Ferussacia vescoi, Eamblya (3) 

Caecilianella acicula (4) 

Bulimus decollatus (2) 

Helix pygmaea (4) 

Helix poupillieri (3) 

Hyalinia vitreola (3) 

Helix cellaria (2), Hyalinia chersa, H. lusitanica, H. blidahensis (3), H. cellaria (4) 
Hyalinia raterana, H. kraliki (3) 


= >= 


P 
P 
P 


—T TS 


DE OLIVEIRA: Fauna Malacológica de Commbra. Moluscos “urbanos” de Portugal. 1 


Tabela MIT. Continuagáo. 
Table HI. Continuation. 


Espécies 


Aegopinella nitens (Michaud, 1831) 
Aegopinella nitidula (Draparnaud, 1805) 
Milox gagates (Draparaud, 1801) 
Deroceras agreste (Pollonera, 1891) 
Deroceras loeve (Miller O. E, 1774) 
Arion intermedius Normand, 1852 
Cochlicella acuta (Miller O. F, 1774) 
Cochlicella barbara (Linnaeus, 1758) 
Oestophora barbula (Rossmússler, 1838) 
Oestophora lusitanica (Pfeiffer, 1841) 
Ponentina subvirescens (Bellamy, 1839) 


Xerotricha apicina (Lamarck, 1822) 
Xerotricha conspurcata (Draparaud, 1801) 
Candidula belemensis (Servain, 1880) 
Candidula intersecta (Poiret, 1801) 
Candidula olisippensis (Servain, 1880) 
Cernuella virgata (da Costa, 1778) 

Theba pisana pisana (Miller 0. E, 1774) 


Cepaea nemoralis (Linnaeus, 1758) 
Portugala inchoata (Morelet, 1845) 
Cornu aspersum (Miller O. E, 1774) 
Unio delphinus Spengler, 1793 
Anodonta anatina (Linnaeus, 1758) 


Sphaerium corneum (Linnaeus, 1758) 
Musculium lacustre (Miller O. F., 1774) 


Sinónimos 


Hyalinia nitens, H. castroi (3), Zonites nitens (4) 

Hyalinia nitidula (4) 

Amalia gagates (4) 

Agriolimax agrestis (4) 

Agriolimax laevis (4) 

Arion hessei (4) 

Cochlicella barbara (3), Helix barbara (4) 

Helix acuta, H. ventrosa (4) 

Helix barbula (3, 4), H. barbella, H. despicta (3) 

Helix lusitanica (3, 4) 

Helix occidentalis (2, 3, 4), H. villula, H. atachypora, H. conimbricensis, 
H. rosai (3) 

Helix apicina (2, 3, 4) 

Helix conspurcata (2, 3), H. moricola (3) 

Helix absidiata (3) 

Helix caperata (2), H. intersecta (3, 4), H. herbarum (3) 

Helix defectiva, H.unifasciata, H. olisippensis (3) 

Helix virgata, H. variabilis (4) 

Helix pisana (2, 3, 4), H. pisanella, H. pisanopsis, H. carpiensis, 

H. dierbanica (3) 

Helix nemoralis (3, 4) 

Helix inchoata (3, 4), H. paulinoi, H. nobrei, H. goltzi, H. pochi (3) 
Helix aspersa (3, 4) 
Unio pictorum (1, 4), U. oeschrus, U. amblyus (3) 


Anodonta macilenta (1, 3, 4), A. acyrta, A. silvae, A. embiella, A. bocageana, 


A. rosai (3) 
Sphaerium lusitanicum, S. nucleatum (3) 
Sphaerium lacustris (4) 


5,6. Este é o único registo conhecido de 
P. amnicum para Coimbra. 


Registos duvidosos ou errados 

Recentemente DE OLIVEIRA (2009b) 
cita para Coimbra Pyramidula pusilla 
(GITTENBERGER E BANK, 1996). Este 
registo, após análise mais detalhada, é 
errado e corresponde a Plagyrona placida. 


CONCLUSAÁO 


Pelo menos 61 espécies de moluscos 
continentais sobrevivem ainda na cidade 
de Coimbra. Uma diversidade ainda 
assim notável dados (i) o isolamento e a 


exiguidade dos habitats actualmente dis- 
poníveis e (ii) a continua manutencáo e 
artificializacio desses mesmos espacos 
(limpeza, recolha de manta morta, obras 
de “beneficiacáo”, eventual uso de 
moluscicidas...), que periódica e sistema- 
ticamente váo sendo responsáveis por 
fenómenos de mortalidade. 


AGRADECIMENTOS 


Material bibliográfico e malacoló- 
gico foi disponibilizado para consulta e 
estudo na Biblioteca de Zoologia e no 
Museu de História Natural da Universi- 
dade de Coimbra. No Museu de Histó- 


49 


Iberus, 28 (2), 2010 


ria Natural da Universidade do Porto 
foram estudadas as amostras da Familia 
Clausiliidae depositadas na coll. Nobre. 


BIBLIOGRAFIA 


BECKMANN K.-H., 2007. Die Land- und Súis- 
swassermollusken der Balearischen Inseln. 
Conchbooks, Wiesbaden. 255 p. 

BOETERS H. D. 1988. Moitessieriidae und Hy- 
drobiidae in Spanien und Portugal (Gastro- 
poda: Prosobranchia). Archiv fúr Mollusken- 
kunde, 118 (4-6): 181-261. 

CARVALHO R. N. 1944. Catálogo da Coleccáo de 
Invertebrados de Portugal existentes no Mu- 
seu Zoológico da Universidade de Coimbra. 
Mollusca. 2* Parte: A. Classe Pelecypoda 
Goldfuss (Acephala Cuv. Lamellibranchiata 
Blv.). Memórias e Estudos do Museu Zoológico 
da Universidade de Coimbra, 162: 1-22. 

CARVALHO R. N. 1945. Catálogo da Coleccáo de 
Invertebrados de Portugal existentes no Mu- 
seu Zoológico da Universidade de Coimbra. 
Mollusca. 2* Parte: B. Classe Gastropoda Cu- 
vier. Memórias e Estudos do Museu Zoológico 
da Universidade de Coimbra, 167: 1-50. 

CASTILLEJO J., 1990a. Babosas de la Peninsula 
Iberica. 1. Los Arionidos. Catalogo critico y 
mapas de distribucion (Gastropoda, Pulmo- 
nata, Arionidae). Comunicaciones del VIII 
Congreso Nacional de Malacologia, Valencia, 
Octubre de 1990, Iberus, 9 (1-2): 331-345. 

CASTILLEJO J. 1990b. Babosas de la Peninsula Ibe- 
rica. II. Los Agriolimacidos. Catalogo critico 
y mapas de distribucion (Gastropoda, Pul- 
monata, Agriolimacidae). Comunicaciones 
del VIII Congreso Nacional de Malacologia, 
Valencia, Octubre de 1990, Iberus, 9 (1-2): 
347-358. 

DE OLIVEIRA A., 2007. Spermodea lamellata (Jef- 
freys, 1830) (Pulmonata: Valloniidae, Acant- 
hinulinae): novos dados sobre a sua distri- 
buicáo em Portugal. Noticiario de la Sociedad 
Española de Malacologia, 47: 29-31. 

DE OLIVEIRA A., 2008. Materiais para o estudo 
da Malacofauna náo-marinha de Portugal. 1. 
Seis gastrópodes (Pulmonata, Stylomma- 
tophora) náo citados na obra de Augusto 
Nobre. 2. Monacha cartusiana (Muller O. F., 
1774) e Helicigona lapicida (Linnaeus, 1758) 
(Pulmonata, Helicoidea). Noticiario de la So- 
ciedad Española de Malacologia, 49: 40-45. 51: 
48 [errata]. 

DE OLIVEIRA A. 2009a. Materiais para o estudo 
da Malacofauna náo-marinha de Portugal. 4. 
Revisáo das espécies aquáticas introduzidas. 
Noticiario de la Sociedad Española de Malacolo- 
gia, 52: 31-37. 


50 


Emilio Rolán estudou anatomica- 
mente a colónia de Mercuria da Fonte da 
Sereia confirmando a sua identidade. 


DE OLIVEIRA A. 2009b. Materiais para o estudo 
da Malacofauna náo-marinha de Portugal. 5. 
Cochlicopa lubricella (Rossmássler, 1834) (Pul- 
monata, Cochlicopidae). Noticiario de la So- 
ciedad Española de Malacología, 51: 55-58. 

GITTENBERGER E., 1989. Additional data con- 
cerning the systematics and the remarkable 
ranges of three species of landsnails, known 
from Sintra. Publicacóes Ocasionais da Sociedade 
Portuguesa de Malacologia, 13: 13-16, fig. 1-5. 

GiusTI F. E MANGANELLIG., 1992. The problem 
of the species in Malacology after clear evi- 
dence of the limits of morphological syste- 
matics. In Gittenberger E. e Goud J. (Eds.): 
Proceedings of the Ninth International Malaco- 
gical Congress, Edinburgh, 1986. Unitas Mala- 
cologica, Leiden: 153-172. 

HIDALGO J. G. 1875. Catálogo iconográfico y des- 
criptivo de los moluscos terrestres de España, 
Portugal y las Baleares. Madrid. pp. 223, lam. 
A, 1-44, pp. 46. 

HOLYOAK G. A. 2009. Ferrissia fragilis (Gastro- 
poda: Planorbidae) in Portugal. Noticiario de 
la Sociedad Española de Malacología, 52: 41-42. 

LOCARD A. 1899. Conchyliologie Portugaise. Les 
coquilles terrestres des eaux douces et sauma- 
tres. Archives du Muséum d'Histoire Naturelle 
de Lyon, 7 (1): i-iv, 1-303. 

MARTÍNEZ-ORTÍ A. 2006. Balea heydeni Von Malt- 
zan, 1881 (Gastropoda, Clausilidae) en Es- 
paña: características conquiológicas y distri- 
bucción. Noticiario de la Sociedad Española de 
Malacologia, 45: 30-37. 

MORELET A. 1845. Description des Mollusques 
terrestres et fluviatiles du Portugal. J.-B. Bailliere, 
Paris. 115 pp., 14 pl. 

NOBRE A., 1885. Catalogue des Mollusques des 
environs de Coimbre (Portugal). Annales de 
la Societé Royale Malacologique de Belgique, 20: 
45-62. 

NOBRE A. 1930. Moluscos terrestres, fluviais e das 
águas salobras de Portugal. Companhia Editora 
do Minho, Barcelos. 259 pp., 18 pl. 

RODRÍGUEZ T., HERMIDA J. E OUTEIRO A. 1993. 
La Superfamilia Zonitoidea Mórch, 1864 en 
Portugal. Revista da Academia Galega de Cien- 
cias, 12: 57-66. 


O Sociedad Española de Malacología Iberus, 28 (Oy 51-62, 2010 


Notes on Mediterranean Dizoniopsis (Gastropoda: 
Cerithiopsidae), with the description of two new species 


Apuntes sobre los Dizoniopsis (Gastropoda: Cerithiopsidae) del 
Mediterráneo, con la descripción de dos especies nuevas 


Philippe BOUCHET*, Serge GOFAS** and Anders WARÉN*** 


Recibido el 8-XI-2010. Aceptado el 10-XI-2010 


ABSTRACT 


Species-level taxonomy is reviewed for the Mediterranean Cerithiopsidae with a sculpture 
formed by two rows of beads. The two previously known species are assigned respectively 
to Dizoniopsis coppolae [Aradas, 1870) and D. concatenata (Conti, 1864) and are held 
as distinct from the fossil type species Dizoniopsis bilineata (Hórnes, 1848). Two addi- 
tional species, which are found sympatrically with these in the Strait of Gibraltar, are des- 
cribed as new. Dizoniopsis micalii Cecalupo and Villari 1997 is reassigned to the genus 
Cerithiopsis. 


RESUMEN 


Se revisa la taxonomía de las especies mediterráneas de Cerithiopsidae con escultura for- 
mada por dos filas de gránulos. Las dos especies previamente conocidas se asignan res- 
pectivamente a Dizoniopsis coppolae (Aradas, 1870) y D. concatenata (Conti, 1864) y 
se consideran distintas de la especie tipo fósil Dizoniopsis bilineata (Hórnes, 1848). Dos 
especies adicionales, que se encuentran en el estrecho de Gibraltar en simpatría con las 
anteriores, se describen como nuevas. Dizoniopsis micaliiCecalupo y Villari 1997 se reu- 
bica en el género Cerithiopsis. 


INTRODUCTION 


The Cerithiopsidae are a family of 
small gastropods, distributed world- 
wide in tropical and temperate shelf 
environments. There are many genera 
and species, difficult to recognize 
because there is very little differentia- 
tion in shell morphology. All may be 
characterized by small (3 to 10 mm) size, 
brown to dark colour, high spire and 


usually a spiral sculpture of beaded 
cords. Cerithiopsidae are specialized for 
feeding on sponges and this feature is 
shared with the Triphoridae, another 
family in which differentiation at 
generic level is not reflected in shell 
characters. The question of the relation- 
ship between the two families remains 
beyond the scope of this paper. 


* Muséum National d'Histoire Naturelle, Département Systématique et Evolution USM 603/UMR 7138 * 
Systématique, Adaptation, Evolution”, Equipe “Exploration de la Biodiversité”, Case Postale 51, 55, Rue 
Buffon, F-75231 Paris Cedex 05, France. 

** Departamento de Biología Animal, Facultad de Ciencias, Universidad de Málaga, Campus de Teatinos - E- 
29071 Málaga, Spain. 

** Department of Invertebrate Zoology, Swedish Museum of Natural History, Box 50007, SE-10405 
Stockholm, Sweden. 


51 


Iberus, 28 (2), 2010 


The genus-level taxonomy of the 
Cerithiopsidae remains to be worked 
out in most areas of the world, inclu- 
ding Europe. The most relevant contri- 
bution is that of MARSHALL (1983) based 
on New Zealand species. The Medite- 
rranean species have been described in 
numerous papers, scattered in the litera- 
ture and usually dealing with one or 
two species at a time. lt was our inten- 
tion, many years ago, to provide a revi- 
sion of all European species known to 
date. For this we accumulated substan- 
tial data, but the revision was not com- 
pleted and, meanwhile, most of the 
species then undescribed were named 
by other authors. 

Among European cerithiopsids, a 
noteworthy and easily recognized mor- 
phological group is composed by 
species in which the teleoconch sculp- 
ture is composed of only two rows of 
beads until at least the penultimate 
whorl. In most of the other Mediter- 
ranean species of the family, there are 
two (rarely three) cords on the first 
teleoconch whorl and a third one is 
added very early, adapically along the 
suture. 

SACCO (1895) introduced Dizoniopsis 
as a subgenus of Cerithiopsis, for 


Neogene fossils which were said to 
differ by having two rows of granules 
per whorl only. This character is shared 
by some Recent Mediterranean repre- 
sentatives, for which the name of the 
Miocene type species has been repeat- 
edly used. There are two clearly differ- 
ent Recent species, long recognized by 
Mediterranean authors, but much con- 
fusion has arisen regarding which name 
should be used for one or another of 
them. 

This report is mostly based on mate- 
rial collected around Ceuta, in the west- 
ernmost part of the Mediterranean, 
during a workshop organized by 
Philippe Bouchet (Muséum National 
d'Histoire Naturelle, Paris) and José 
Carlos Garcia Gómez (Universidad de 
Sevilla), in May 1986. We here summa- 
rize published records on the two previ- 
ously known species, and describe two 
additional species from the Strait of 
Gibraltar. 


Abbreviations: 


MNHN Muséum National d'Histoire 
Naturelle, Paris 

sh. shell(s) 

spm live taken specimen(s) 


Genus Dizontopsis Sacco, 1895 


Type species: Cerithium bilineatum Hórnes, 1848, by original designation (Steinebrunn, Middle 


Miocene, Vienna Basin) 


The type species of Dizoniopsis has 
been illustrated by LANDAU, LA PERNA 
AND MARQUET (2006) who designated a 
lectotype. The illustrated protoconch is 
incompletely preserved but conserves 
nearly two whorls, and therefore is pre- 
sumably multispiral with nearly three 
whorls if complete. The sculpture of this 
protoconch consists of two moderate 
spiral keels on the penultimate whorl, 
the abapical one becoming concealed by 
the suture on the last whorl, and of flex- 
uose axial riblets forming a reticulate 
pattern with the keels. 

This protoconch morphology differs 
radically from that of the two Recent 


iZ 


Mediterranean species, adequately illus- 
trated and discussed by PALAZZI AND 
VILARI (2001). Conversely, it is quite 
similar to the protoconch of the Recent 
Dizoniopsis apexclarus Rolán, 2007, 
described from the islands of the Gulf of 
Guinea, which differs in having a single 
keel on the penultimate whorl and 
hardly more than two whorls in total. 
The outcome is that the name Dizo- 
niopsis bilineata cannot be used for a 
Recent Mediterranean species. Other 
available names which have been used 
for this group of species, viz. Cerithiopsis 
clarkii Forbes and Hanley, 1851, 
Cerithium concatenatum Conti, 1864 and 


BOUCHET ET AL.: Two new Mediterranean species of the genus Dizoniopsis 


Cerithiopsis coppolae Aradas, 1870, are 
discussed hereafter. 

An additional Mediterranean species 
was originally described as Dizoniopsis 
micalii by CECALUPO AND VILLARI (1997) 
and is still currently held in this binomen. 
It differs from the species discussed herein 
in both the protoconch and teleoconch 


characters, and we consider it better 
assigned to Cerithiopsis s. 1. Its protoconch 
has axial ribs resembling Cerithiopsis scalaris 
Locard, 1892 whereas the teleoconch starts 
with two rows on granules on the early 
whorls as most Cerithiopsis species, but 
later completes three rows like in the type 
species Cerithiopsis tubercularis. 


Dizoniopsis coppolae (Aradas, 1870) (Figs. 1-4) 


Cerithiopsis coppolae Aradas, 1870: Atti Accad. Gioenia Sc. Nat., (3) 4: 263-268 [Ognina near Catania, 
Sicilia] — Pallary, 1920, Expl. Scient. Maroc: 45 . 

Cerithiopsis bilineata var. ventricosa Brusina, 1871, Bull. Malac. Ital., 4: 5-7 [Croatia] 

Cerithiopsis bilineata [non (Hórnes, 1848)] — Monterosato, 1877, J. Conchy]l., 25: 41; Monterosato, 
1878, Giorn. Sc. Nat. Econ. Palermo, 99; Kobelt, 1908, Iconogr. vol. 4: 118, pl. 120, fig. 14-15. 

Cerithiopsis (Dizoniopsis) bilineata [non (Hórnes, 1848)] - Nordsieck, 1968, Europ. Meeres-Gehius- 
eschnecken: 70, pl. 11 fig. 43.10. 

Dizoniopsis euxinica haifensis Nordsieck, 1972, Arch. Molluskenk., 102: 234 [Shiqmona, Israel]. 

Cerithiopsis (Dizontopsis) bilineata [non (Hórnes, 1848)] and var. concatenata [non (Conti, 1864)] — 
Fekih and Gougerot, 1974, Bull. Inst. Océanogr. Péche Salammbó: 184-185, 207-208. 

Cerithiopsis (Dizoniopsis) coppolae Aradas, 1870 — Nordsieck, 1976, La Conchiglia 87-88: 7 — Palazzi 
and Villari, 2001, La Conchiglia, 297, suppl.: 15-18, 38-40. 

Dizoniopsis coppolae (Aradas, 1870) — Oliver Baldoví, 2007, Iberus 25(2): 32. 


Type material: C. coppolae, whereabouts unknown. C. bilineata var. ventricosa: 61 syntypes from 
mixed Dalmatian localities (Otok, Lapad, Lokrum) in Croatian Natural History Museum, Zagreb, 
reg. n” 1295, and 13 syntypes in Jeffreys collection ex Brusina, USNM 187810. D. euxinica haifensis: 
7 syntypes SMF 239381 /1 (the shell figured by Nordsieck) and 239382 /6. 

Material examined: Portugal. - Sagres, Ponta da Baleeira, 37” 00.37 N — 08” 55.5” W, 17-23 m, 2 sh. 
(4.0 x 1.5 to 4.8 x 1.8 mm without protoconch), Mission Algarve 1988 (MNHN). Morocco.- Asilah, 
from beach drift, 4 sh. (up to 3.5 x 1.3 mm without protoconch). Strait of Gibraltar. — Ceuta, Punta 
Almina, 35” 54.1” N-05* 16.5” W, submarine cliff 25-40 m, 2 sh. (4.4 x 1.7 mm, 5.1 x 1.8 mm, MNHN). 
Ceuta, Punta del Desnarigado, 35” 53.6” N — 05” 16.8” W, 16-20 m, 1 old sh. (4.8 x 1.8 mm, MNHN). 
Italy, Sicily. - Acitrezza, 1 sh. (3.9 x 1.3 mm), Mission Sicile 1990 (MNHN). Brucoli, 1 sh. (4.7 x 1.4 
mm), Mission Sicile 1990 (MNHN). Tunisia. — Djerba, 78 sh. (3.5 x 1.2 to 4.7 x 1.4 mm), col. Bouchet 


and Warén, 1982 (UNHN). 


Description: Shell up to a little more 
than 5 mm, high conical, solid, with 
about 2 * /4 protoconch whorls and 10-12 
teleoconch whorls. Protoconch narrow 
and styliform, easily broken off, with 
convex whorls and a sculpture of flexu- 
ous axial riblets which start from the 
adapical suture and gradually fade out; 
these are irregularly distributed on the 
last protoconch whorls and vary in 
strength among individuals. Sculpture 
of teleoconch composed by two spiral 
cords, approximately as broad as the 
intervening space, crossed by axial ribs 
which form very distinct beads at their 
intersection with the cords. On the first 


teleoconch whorl, the abapical cord is 
more prominent than the other one and 
the axial ribs are not very conspicuous. 
On the following whorls, the adapical 
cord progressively takes over as the 
thickest one. On the last whorl, the 
beads of the adapical cord become 
slightly more elongated but do not split. 
The abapical part of the body whorl 
bears one cord, slightly ragged but not 
beaded, in prolongation of the suture, 
and two more similar cords between 
this and the siphonal canal; these cords 
are markedly narrower than the beaded 
cords on the ribs and the spaces 
between them are crossed by raised 


53 


Iberus, 28 (2), 2010 


Figures 1-4. Dizoniopsis coppolae (Aradas, 1870). 1: shell from Djerba, Tunisia (actual size 4.4 
mm); 2: protoconch of another specimen from the same locality (scanning electron micrograph; 3, 
4: shell from Ceuta, Strait of Gibraltar, Punta Almina 25-40 m (actual size 5.1 mm). 

Figuras 1-4. Dizoniopsis coppolae (Aradas, 1870). 1: concha de Djerba, Túnez (tamaño real 4,4 
mm); 2: protoconcha de otro ejemplar de la misma localidad (micrografía electrónica de barrido); 3, 4: 
concha de Ceuta, estrecho de Gibraltar, Punta Almina 25-40 m (tamaño real 5,1 mm). 


threads parallel to the growth lines. 
Aperture oval, with a broad and short 
siphonal canal and a smaller channel at 
the opposite end. Adult shells have a 
rounded, but not thickened edge to the 
outer lip. Colour brown, grading to 
paler or even white towards the apical 
whorls; the nodes are paler with a 
greyish hue contrasting with the dark 
brown on spaces between nodes on the 
cords, and the abapical cords are also 
somewhat darker than the intervening 
spaces. Animal unknown. 

Remarks: It is strange that ARADAS 
(1870) dedicated a special paper to the 
description of this species whereas a few 
years later ARADAS AND BENOIT (1876: 
239) do not list it as valid, mentioning 
instead that some authors hold it as a 
variety of Cerithium tuberculare, and do 
not even retain Cerithiopsis as distinct 
from Cerithium. Most Mediterranean 
authors subsequently used the name 


54 


Cerithiopsis bilineata for this species, fol- 
lowing the lead by MONTEROSATO (1878). 

This species has been adequately 
figured and discussed by PALAZZI AND 
VILLARI (2001) who noted the variability 
in the protoconch sculpture. The distrib- 
ution of this species seems to be 
restricted. It is widespread in the 
Eastern and Central Mediterranean, but 
there are hardly any records from the 
coasts of France and Spain in the 
Western basin, and it is represented in 
our material from the Strait of Gibraltar 
by only three old shells. It is neverthe- 
less cited by OLIVER BALDOVÍ (2007) in 
shell grit surrounding Posidonia grounds 
in the bay of Valencia, and by PALLARY 
(1920) from the Atlantic coast of 
Morocco. The specimens collected in the 
Strait of Gibraltar and in the Algarve are 
somewhat larger and broader than those 
from Tunisia and Sicily, and all lack a 
protoconch. 


BOUCHET ET AL.: Two new Mediterranean species of the genus Dizoniopsis 


Dizontopsis concatenata (Conti, 1864) (Figs. 5-12) 


Cerithium concatenatus Conti, 1864, Il Monte Mario..: 51. — Landau, La Perna and Marquet, 2006, 
Palaeontos 10: 11, 15-16. 

Cerithi == clarkii [non Forbes and Hanley, 1851] — Monterosato, 1877, J. Conchyl., 25: 41; Mon- 
te. 378, Giorn. Sc. Nat. Econ. Palermo, 99; Kobelt, 1908, Iconogr. vol. 4: 126-127, pl. 120, 
fig. 11-12. 

Cerithiposis bilineata var. concatenata (Conti, 1864) - Cerulli-Irelli, 1912, Palaeontogr. Ital., 18: 149, pl. 
23 fig. 50-51. 

Cerithiopsis bilineata var. ventricosa [non Brusina, 1871] — Bucquoy, Dautzenberg and Dollfus, 
1884, Moll. Roussillon, vol. 1: 205-206, pl. 27 fig. 10-12. 

Cerithiopsis (Cerithiopsida) clarki [non Forbes and Hanley, 1851] - Nordsieck, 1968, Europ. Meeres- 
Geháuseschnecken: 70, pl. 11 fig. 43.31. 

Cerithiopsis (Dizoniopsis) clarki [non Forbes and Hanley, 1851] — Fekih and Gougerot, 1974, Bull. 
Inst. Océanogr. Péche Salammbó 3: 184-185, 207-208. 

Cerithiopsis (Dizoniopsis) bilineata [non (Hórnes, 1848)] - Nordsieck, 1976, La Conchiglia 87-88: 7, 18 
(en parte) — Ros and Altimira 1977: 53 “de color pardo oscuro en los individuos recolectados 
vivos” 

Cerithiopsis bilineata [non (Hórnes, 1848)] - Giannuzzi Savelli et al., 1999: 44-45, fig. 76 a,b. — 
Oliver Baldoví, 2007, Iberus, 25 (2): 49, fig. 34-35. 

Cerithiopsis (Dizoniopsis) concatenata (Conti, 1864) — Palazzi and Villari, 2001, La Conchiglia, 297, 
suppl.: 15-18, 38-40. 


Type material: Lectotype (Landau, La Perna and Marquet, 2006) in coll. Cerulli-Irelli, Museo di 
Paleontología dell'Universita di Roma “La Sapienza” (see remarks below). 

Material examined: Portugal. - Sagres, Bay of Baleeira, 37” 00.7” N — 08* 55.0” W, 3-15 m, 1 spm. 
drawing AL 132. (3.6 x 1.4 mm without protoconch), Mission Algarve 1988 (MNHN). Strait of 
Gibraltar. - Ceuta, Punta del Saudiño, 35” 54.1'N — 05” 18.0"W, submarine cliff 17-35 m; drawing CE 
38, 2 spm. (4.1 x 1.4, 4.0 x 1.4 mm); Punta del Desnarigado, 35” 53.6" N — 05” 16.8” W, 16-20 m, 1 old 
sh. (UNHN). Mediterranean France. - Le Dramont 22-30 m, 3 sh. (up to 3.7 x 1.2 mm), leg. Pelorce 
(MNHN). Illes d'Hyeres, Grande Passe 112-113 m, 2 sh. (3.8 x 1.2, 5.2 x 1.7 mm) leg. Picard 1956 
(MNHN). Cap Morgiou, Calanque de la Triperie 22 m, 1 sh. juv., leg. Zibrowius 1996 (MNHN). 
Unknown origin, possibly Roussillon, 2 spm. figured in Bucquoy, Dautzenberg and Dollfus (1884), 
pl. 27 fig. 10-11 as C. bilineata var. ventricosa (3.9 x 1. 3 mm). Corsica. - Calvi, 1 spm. (4.5 x 1.5 mm, 
MNHN). Italy. - Camogli, Genova, 42 m, 1 sh. (3.0 x 1.2 mm). Capraia, Le Formiche, 3 sh. (3.1 x 
1.1 to 4 x 1.2 mm), leg. Palazzi (MNHN); Livorno 7-17 m, 2 spm. (4.2 x 1.2), leg. Palazzi (MNHN). 
Sicily, Acitrezza 36 m, 4 sh. (full grown 3.2 x 1.1 to 4.0 x 1.4 mm), leg. Spada (SMNH). Acitrezza, 
1 sh. (4.3 x 1.3 mm), Mission Sicile 1990 (MNHN). 


Description: Shell up to a little more 
than 5 mm, high conical, solid, with 
about 2 */4 protoconch whorls and 7 to 8 
1/2 teleoconch whorls. Protoconch 
narrow and styliform, easily broken off, 
the first whorl rounded with a frosted 
surface, the following with two well- 
defined keels appearing progressively 
and ruming rather close together along 
the middle part of the whorl, more or 
less connecting to the start of the beaded 
cords on the first teleoconch whorl. 
Limit protoconch-teleoconch not very 
distinct. Sculpture of teleoconch com- 
posed by two spiral cords, approxi- 
mately as broad as the intervening 


space, crossed by axial ribs which form 
very distinct beads at their intersection 
with the cords. On the first teleoconch 
whorl, the abapical cord is markedly 
more prominent than the other one and 
overhangs the suture of the following 
whorl; the axial ribs are there compara- 
tively more apparent and more crowded 
than on the second teleoconch whorl. 
On the following whorls, the adapical 
cord progressively takes over as the 
thickest one. On the last whorl, the 
beads of the adapical cord become elon- 
gated in the axial direction and, on the 
section preceding the aperture of adult 
specimens, this cord tends to split and 


33 


Iberus, 28 (2), 2010 


form two contiguous rows of beads, 
which become still more narrow and 
elongate. The abapical part of the body 
whorl bears one thick beaded cord in 
prolongation of the suture and another 
one, flatter and not distinctly beaded, in 
the intervening space between this and 
the siphonal canal. Aperture oval, with a 
broad and short siphonal canal and a 
smaller channel at the opposite end. 
Adult shells have a rounded, but not 
thickened edge to the outer lip. Colour 
brown, grading to paler or even white 
towards the apical whorls. 

Animal with an elongated foot, with 
propodium truncated in front and bor- 
dered anteriorly by a mentum, the 
median part rather narrowing, and the 
metapodium broadly ovate. Opening of 
the pedal gland conspicuous at the ante- 
rior end of metapodium, prolonged 
towards the posterior end by an axial 
groove on the sole. Head devoid of 
snout, provided with two elongate 
cephalic tentacles which are not sepa- 
rated at their base by any rim or 
swelling, and form a V-shape when 
extended. Eyes black, rounded, situated 
within the base of the tentacles without 
forming a distinct swelling, not sur- 
rounded by any opaque granular mater- 
ial. Opercular lobe rather thick, not con- 
taining any opaque granular material. 
Mantle with a smooth edge. 

Remarks: This species is clearly sepa- 
rated from Dizoniopsis coppolae and D. 
bilineata by both protoconch and teleo- 
conch characters. The protoconch lacks 
axial sculpture, whereas on D. coppolae it 
is covered with flexuous axial ribs and 
lacks spiral keels (see PALAZZI AND 
VILLARI, 2001: figs 65-69); its two periph- 
eral keels remain on the median part of 
the whorl contrary to the fossil D. bilin- 
eata in which the abapical keel is con- 
cealed by the suture on the last proto- 
conch whorl. The teleoconch differs 
from D. coppolae in several clearcut char- 
acters of the body whorl. The subsutural 
cord definitely splits before reaching the 
outer lip of the aperture, and there 
accounts for more than one-third of the 
height of the whorl, whereas in D. coppo- 
lae this cord does not split and does not 


So) 


exceed one-quarter of the height of the 
whorl. The most obvious distinguishing 
feature is that there are only two abapi- 
cal cords on the base (one in prolonga- 
tion of the suture, and one more) 
whereas there are three (one in the pro- 
longation of the suture and two more) in 
D. coppolae. The colour pattern is also 
different. Although grading from lighter 
to darker from apex to later whorls, it is 
uniformly brown on one particular 
whorl whereas in D. coppolae the nodes 
have a greyish hue contrasting with the 
dark brown on spaces between nodes on 
the cord, and with an intermediate light 
brown in the intervening spaces 
between cords; it is clear from this that 
Ros AND ALTIMIRA (1977) observed this 
species, and not D. coppolae, on the 
Catalan coast. 

The areas behind the eyes and 
beneath the operculum are translucent 
and similar in texture to the remaining 
surface of the body, contrary to most 
cerithiopsids we have observed and 
particularly to Cerithiopsis tubercularis, 
where these areas are densely furnished 
with whitish or yellowish granules. The 
animal was observed alive in two differ- 
ent localities (Ceuta and Algarve) which 
rules out that this difference is acciden- 
tal. 

MONTEROSATO (1877, 1878) was the 
first to distinguish two species among 
Mediterranean cerithiopsids having two 
rows of granules, and used for them 
respectively the names Cerithiopsis bilin- 
eata (considering C. coppolae as a 
synonym) and C. clarkii. Cerithiopsis 
clarkii Forbes and Hanley, 1851 (vol. 3 p. 
368, vol. 4 pl. 103 fig. 6) was introduced 
conditionally, based on a specimen col- 
lected at Exmouth, in the English 
Channel. All the species of Cerithiopsi- 
dae found in Britain, including the type 
species Cerithiopsis tubercularis 
(Montagu, 1803), have normally three 
rows of granules on the whorls but may 
occasionally display two or four rows, 
usually after an accident in shell 
growth. Therefore JEFFREYS (1867: 267) 
was definitely correct in his interpreta- 
tion of the specimen as a teratological 
“Monstr. Clarkii” of Cerithiopsis tubercu- 


BOUCHET ET 4L.: Two new Mediterranean species of the genus Dizoniopsis 


Figures 5-11. Dizoniopsis concatenata (Conti, 1864). 5, 6: specimen from Ceuta, Strait of Gibral- 
tar, Punta del Saudiño 17-35 m (actual size 4.1 mm, same specimen as fig. 12); 7: another speci- 
men from the same locality (scanning electron micrograph, actual size 4.1 mm); 8: protoconch, 
same specimen; 9, 10: shell from Le Dramont, Mediterranean coast of France, 22-30 m (actual 
size 3.6 mm); 11: shell from Aci Trezza near Catania, Sicily (actual size 4.1 mm). 

Figuras 5-11. Dizoniopsis concatenata (Conti, 1864). 5, 6: ejemplar de Ceuta, estrecho de Gibraltar, 
Punta del Saudiño 17-35 m (tamaño real 4,1 mm, mismo ejemplar que fig. 12); 7: otro ejemplar de la 
misma localidad (micrografía electrónica de barrido, tamaño real 4,1 mm); 8: protoconcha, mismo 
ejemplar; 9, 10: concha de Le Dramont, costa mediterránea de Francia, 22-30 m (tamaño real 3,6 
mm); 11: concha de Aci Trezza cerca de Catania, Sicilia (tamaño real 4,1 mm). 


/ 


Iberus, 28 (2), 2010 


Figure 12. Dizoniopsis concatenata (Conti, 1864), living animal from Ceuta, Strait of Gibraltar, 
Punta del Saudiño 17-35 m (same specimen as fig. 5-6). Figure 13. Dizoniopsis aspicienda spec. 
nov., living animal of the holotype from Ceuta, Strait of Gibraltar, Punta Almina 25-40 m (same 


specimen as fig. 14-15). 


Figura 12. Dizoniopsis concatenata (Conti, 1864), animal vivo de Ceuta, estrecho de Gibraltar, 
Punta del Saudiño 17-35 m (mismo ejemplar que fig. 5-6). Figura 13. Dizoniopsis aspicienda spec. 
nov., animal vivo del holotipo de Ceuta, estrecho de Gibraltar, Punta Almina 25-40 m (mismo ejem- 


plar que fig. 14-15). 


laris, noting that the early whorls have 
the usual three rows, and this was also 
the view of MARSHALL (1911). NORD- 
SIECK (1968) first used the name 
Cerithiopsis (Cerithiopsida) clarki (written 
with one “i”) for this species, but later 
(Nordsieck, 1976) assigned it to C.bilin- 
eata and considered C. clarki (based on 
the figure of Forbes and Hanley) to 
occur only in the Atlantic. His figure of 
C. bilineata on p. 18 is nevertheless C. 
coppolae. 

CERULLFIRELLI (1912) used the name 
C. bilineata var. concatenata (Conti, 1864) 
and illustrated a specimen from the type 
locality, the lower Pleistocene strata of 
Monte Mario near Rome, which is defi- 
nitely the species considered here. The 
same shell is again figured by LANDAU, 


58 


LA PERNA AND MARQUET (2006) and 
designated as lectotype of Cerithium con- 
catenatus Conti, 1864. Cerulli-Irelli stated 
to have identified his specimens by 
comparison with Conti's type material. 
This interpretation was also that of 
PALAZZI AND VILLARI (2001) but contra- 
dicts MONTEROSATO (1884: 134; 1890: 
163) and MARSHALL (1895: 38) who con- 
sidered that Cerithium concatenatus was 
the same as Cerithiopsis pulchella Jeffreys, 
1858 (= C. jeffreysi Watson, 1885). 
However, both are somewhat at odds 
with Conti's original description (repro- 
duced in VAN AARTSEN, MENKHORST 
AND GITTENBERGER, 1984: 29), which 
mentions three rows of granules on the 
whorls and four on the body whorl of 
which there is one smaller next to the 


BOUCHET ET AL.: Two new Mediterranean species of the genus Dizoniopsis 


200 um 


Figures 14-17. Dizoniopsis aspicienda spec. nov. 14, 15: holotype, specimen from Ceuta, Strait of 
Gibraltar, Punta Almina 25-40 m (actual size 4.9 mm); 16: paratype, specimen from Ceuta, 
Benzú, 24 m (scanning electron micrograph, actual size 4.2 mm); 17: protoconch, same specimen. 
Figuras 14-17. Dizoniopsis aspicienda spec. nov. 14, 15: holotipo, ejemplar de Ceuta, estrecho de 
Gibraltar, Punta Almina 25-40 m (tamaño real 4,9 mm); 16: paratipo, ejemplar de Ceuta, Benzú, 24 


m, (micrografía electrónica de barrido, tamaño real 4,2 mm); 17: protoconcha, mismo ejemplar. 


suture and the two on the middle of the 
whorl merging into a single one (hence 
the name). 

Our understanding of the taxon is 
here based on the lectotype illustrated 
by LANDAU, LA PERNA AND MARQUET 
(2006). This specimen may not be eligi- 
ble as such because of not forming part 
of Conti's original material, the where- 
abouts of which are unknown. It is not 


desirable as a neotype either, because it 
lacks a protoconch. Therefore, if it is 
demonstrated that this is not Conti's 
species, it may have to be named as a 
new species but we nevertheless con- 
sider it more parsimonious, for the sta- 
bility of nomenclature, to continue 
usage of this name instead of leaving 
Cerithium concatenatus as a nomen 
dubium. 


Dizoniopsis aspicienda spec. nov. (Figs. 13-17) 


Type material: Holotype, live-collected specimen, Ceuta, Punta Almina, 35” 54.1” N- 05” 16.5 W, 
38-40 m, drawing CE 18 (4.9 x 1.8 mm), MNHN 23283. Paratype, Ceuta, Benz, 35” 55.0” N, 05” 22.5" 


W, 24-26 m (4.2 x 1.6 mm), MNHN 23284. 


Etymology: meaning “worth looking at”, alluding to the handsome aspect of the species. 


99 


Iberus, 28 (2), 2010 


Figures 18, 19. Dizoniopsis abylensis spec. nov. 18: holotype, shell from Ceuta, Strait of Gibraltar, 


200 um 


Punta del Saudiño 17-35 m (actual size 3.8 mm); 19: protoconch of a juvenile shell from Ceuta, 


Punta Bermeja, 27-32 m. 


Figuras 18, 19. Dizoniopsis abylensis spec. nov. 18: holotipo, concha de Ceuta, estrecho de Gibraltar, 
Punta del Saudiño 17-35 m (tamaño real 3,8 mm); 19: protoconcha de una concha juvenil de Ceuta, 


Punta Bermeja, 27-32 m. 


Description: Shell up to nearly 5 mm, 
definitely cyrtoconoid, solid, with about 
2 protoconch whorls and 7 to 7 ?/2 teleo- 
conch whorls. Protoconch narrow and 
styliform, easily broken off, the first 
whorl with a frosted surface, the follow- 
ing with one well-defined keel appearing 
progressively and running at about two- 
thirds of the whorl, closer to the adapical 
suture, and a less pronounced one situ- 
ated more abapically; both more or less 
connecting to the start of the beaded 
cords on the first teleoconch whorl. Limit 
protoconch-teleoconch not very distinct. 
Sculpture of teleoconch composed of two 
spiral cords, approximately as broad as 
the intervening space, crossed by axial 
ribs which form very distinct beads at 
their intersection with the cords. On the 
first teleoconch whorl, the abapical cord 
is markedly more prominent than the 
other one and overhangs the suture of the 
following whorl. On the following 
whorls, the adapical cord progressively 
takes over as slightly thicker. On the last 
whorl, the beads of the adapical cord 
become elongated in the axial direction 
and, on the section preceding the aper- 


60 


ture of adult specimens, this cord tends 
to split and form two contiguous rows of 
beads. The abapical part of the body 
whorl bears one thick beaded cord in 
prolongation of the suture and another 
one, flatter and not distinctly beaded, in 
the intervening space between this and 
the siphonal canal. Aperture oval, with a 
broad and short siphonal canal and a 
smaller channel at the opposite end. 
Adult shells have a rounded, but not 
thickened edge to the outer lip. Colour 
white to very pale tan. 

Animal essentially like in D. concate- 
nata, but markedly smaller in relation to 
the shell and with cephalic tentacles 
broader and flatter. 

Remarks: This species is only repre- 
sented by these two specimens, but is 
clearly separable from sympatric D. con- 
catenata by having a pale, nearly white 
shell, by the cyrtoconoid, rather pupoid 
teleoconch which is markedly broader 
at the same height, and by lacking the 
paired keels on the later larval whorls. 
Like D. concatenata, it has been observed 
to lack the granular masses behind the 
eyes and beneath the operculum. 


BOUCHET ET AL.: Two new Mediterranean species of the genus Dizoniopsis 


Dizoniopsis abylensis spec. nov. (Figs. 18-19) 


Type material: Holotype, shell, Ceuta, Strait of Gibraltar, Punta del Saudiño, 35” 54.1” N — 05” 18.0' 
W, submarine cliff 17-35 m; m (3.8 x 1.6 mm), MNHN 23285. Paratype, juvenile shell from Ceuta, 
Punta Bermeja, 35” 54.6' N — 05” 20.3” W, 27-32 m (1.8 mm), MNHN 23286. 

Etymology: from Abyla, the name of Ceuta in classical Antiquity. 


Description: Shell up to nearly 4 mm, 
slightly cyrtoconoid, solid, with about 2 
protoconch whorls and 7 teleoconch 
whorls. Protoconch narrow and styli- 
form, easily broken off, the first whorl 
rounded with a frosted surface, the fol- 
lowing with two well-defined keels 
appearing progressively and running 
along the middle part of the whorl, 
more or less connecting to the start of 
the beaded cords on the first teleoconch 
whorl. Limit protoconch-teleoconch not 
very distinct. Sculpture of teleoconch 
composed by two spiral cords, approxi- 
mately as broad as the intervening 
space, crossed by axial ribs which form 
very distinct beads at their intersection 
with the cords. On the first teleoconch 
whorl, the abapical cord is markedly 
more prominent than the other one and 
overhangs the suture of the following 
whorl. On the last two whorls, the 
adapical cord progressively takes over 
as slightly thicker. On the last whorl, 
this cord tends to split and form two 
contiguous rows of beads. The abapical 
part of the body whorl bears one well 
defined, smooth cord in prolongation of 
the suture and the intervening space 
between this and the siphonal canal is 
smoothish and excavated, with a weak 
spiral swell which is too poorly defined 
to be taken as a cord. Aperture oval, 


BIBLIOGRAPHY 


ARADAS A. 1870. Descrizione di una specie 
malacologica nuova. Atti dell" Accademia Gioe- 
nia di Scienze Naturali, (3) 4: 263-268. 

ARADAS A. AND BENOIT L. 1872-1876. Conchigli- 
ologia vivente marina della Sicilia. Atti del- 
l'Accademia Gioenia di Scienze Naturali, (3) 6: 
1-112 + pl. 1-2 [1872]; 113-226 + pl. 3-4 [1874); 
227-324 + pl. 5[1876] (dates of publication ac- 
cording to Bouchet P. 1982, Bollettino Mala- 
cologico 18: 177-180). 


with a broad and short siphonal canal 
and a smaller channel at the opposite 
end. Adult shells have a rounded, but 
not thickened edge to the outer lip. 
Colour pale brown, with the beads on 
the spiral cords lighter, the intervening 
spaces between beads on the cords and 
the smooth cord on the base darker. 
Animal unknown. 

Remarks: Although represented by 
only one adult specimen and a juvenile, 
this species is so strikingly different 
from the other two sympatric species of 
Dizoniopsis that we venture its descrip- 
tion as new. The shell is stouter than D. 
concatenata and D. aspicienda, but the 
most noteworthy character is the config- 
uration of the base with one smooth 
cord in prolongation of the suture (dis- 
tinctly beaded in the other two species) 
and an excavated surface between this 
and the suture (furnished with a distinct 
cord in the other two species); this char- 
acter is held as significant since the 
holotype is a completely adult individ- 
ual with a well-formed aperture. The 
colour pattern is also different, and rem- 
iniscent of D. coppolae with lighter beads 
contrasting on cords which are darker 
than the intervening spaces; however 
the latter differs in having three cords 
on the base and a very different proto- 
conch, with axial ribs. 


CECALUPO A. AND VILLARI A. 1997. Dizoniopsis 
micalíí. Una nuova specie per il Mediterraneo 
(Mesogastropoda: Cerithiopsidae). Bollettino 
Malacologico, 32 (1-4): 41-44. 

CERULLFIRELLI S. 1912. Fauna Malacologica 
Mariana. Parte 6. Gastropoda: Cerithiidae 
Cerithiopsidae, Triforidae, Diastomidae, Ver- 
metidae, Turritellidae, Mathildidae, Caecidae. 
Palaeontographia Italica, 18: 141-169. pl. 23-25. 


61 


Iberus, 28 (2), 2010 


CONTI A. 1864. 1! Monte Mario ed i suoi fossili sub- 
appennini raccolti e descritti dallo scultore e pa- 
leontologo. Roma, G. Cesaretti, 57 pp. 

FEKIH M. AND GOUGEROT L. 1974. Liste com- 
mentée des gastéropodes testacés marins re- 
cueillis dans les dépots littoraux actuels du 
Golfe de Tunis. Bulletin de l'Institut National 
Scientifique et Technique d'Océanographie et de 
Péche de Salammbo, 3 (1-4): 185-232 

FORBES E. AND HANLEY S.C. 1848-1853. A his- 
tory of British Mollusca and thetr shells. London, 
van Voorst. Vol. 1: 1-486 (1848). Vol. 2: 1-480 
(1 dec. 1849); 481-557 (1850). Vol. 3: 1-320 
(1850) 321-616 (1851). Vol. 4: 1-300 (1852). In- 
troduction, plates -LXXX [1853]. 

GIANNUZZESAVELLIR., PUSATERI F., PALMERI A. 
AND EBREO C. 1999. Atlante delle conchiglie 
marine del Mediterraneo, vol. 3 : Caenogastropoda 
(parte 2: Ptenoglossa). Evolver, Roma. 127 p. 

JEFFREYS J. G. 1862-1869. British Conchology. Lon- 
don, van Voorst. Vol. 1: pp. CXIV + 341 [1862]. 
Vol. 2: pp. 479 [1864]. Vol. 3: pp. 394 [1865]. 
Vol. 4: pp. 487 [1867]. Vol. 5: pp. 259 [1869]. 

KOBELT W. 1906-1908. Iconographie der schalen- 
tragende europiáischen Meeresconchylien. Part 4: 
1-80, pl. 99-114 [1906]; 81-172, pl. 115-126 
[1908]. Wiesbaden, C.W. Kreidel. 

LANDAU B., La PERNA R. AND MARQUET R. 
2006. The Early Pliocene Gastropoda (Mol- 
lusca) of Estepona, southern Spain, Part 6: 
Triphoroidea, Epitonioidea, Eulimoidea. 
Palaeontos, 10: 1-96, pl. 1-22. 

MARSHALL B.A. 1978. Cerithiopsidae (Mollusca: 
Gastropoda) of New Zealand, and a provi- 
sional classification of the family. New Zealand 
Journal of Zoology, 5: 47-120. 

MARSHALL J.T. 1895. Alterations in “British Con- 
chology”. [Part !1.]. Journal of Conchology 8: 24- 
41 [January 1895]. 

MARSHALL J.T. 1911-1912. Additions to British 
Conchology. Part VII. Journal of Conchology 13: 
179-190 [April 1911]; 192-209 [July 1911]; 223- 
231 [October 1911]; 294-306 [April 1912], 324- 
338 [July 1912]. 


62 


MONTEROSATO T. A. DI 1877. Note sur quelques 
coquilles provenant des cótes d'Algérie. 
(Trad. di H. Crosse). Journal de Conchyliolo- 
ge, 25 (1): 24-49, pl. 2, 3. 

MONTEROSATO T. A. DI 1878. Enumerazione e 
sinonimia delle Conchiglie mediterranee. 
Giornale di Scienze Naturali ed Economiche di 
Palermo, 13: 61-115. 

NORDSIECK F. 1976. Il genere Cerithiopsis Forbes 
€ Hanley, 1849 nei mari d'Europa. La 
Conchiglia, 87-88: 3-7. 

OLIVER BALDOVÍ J.D. 2007. Catálogo de los 
Gasterópodos testáceos marinos de la parte 
Sur del Golfo de Valencia (España). Iberus, 25 
(2): 29-61. 

PALAZZI S. AND VILLARI A. 2001. Molluschi e 
Brachiopodi delle grotte sottomarine del 
Taorminense. La Conchiglia, 297, suppl., 56 pp. 

PALLARY P. 1920. Exploration scientifique du Maroc 
organisée par la Société de Géographie de Paris 
et continuée par la Société des Sciences Naturelles 
du Maroc. Deuxieme fascicule. Malacologie 
(1912). 108 p., 1 pl., 1 map. Rabat and Paris, 
Larose. 

ROLÁN E. 2007. A new species of Dizoniopsis 
(Prosobranchia, Cerithiopsidae) from the 
Gulf of Guinea Islands. Iberus, 25 (1): 33-36. 

RosJ.D. AND ALTIMIRA C. 1977. Comunidades 
bentónicas de sustrato duro del litoral NE es- 
pañol. V. Sistemática de moluscos. Miscel-la- 
nia Zoologica, 4 (1): 43-55. 

SACCO F. 1895. I Molluschi dei terreni terziarii del 
Piemonte e della Liguria. Parte XVII (Cerithiidae, 
Triforidae, Cerithiopsidae e Diastomidae). 83 p, 
3 pl. Torino, Carlo Clausen. 

VAN AARTSEN J.J., MENKHORST H.P.M.G. AND 
GITTENBERGER E. 1984. The marine Mollusca 
of the Bay of Algeciras, Spain, with general 
notes on Mitrella, Marginellidae and Turri- 
dae. Basteria, supplement 2: 1-135. 


O Sociedad Española de Malacología —_—_—_———  Iberus, 28 (2): 63-72, 2010 


Spawn and early development of NE Atlantic species of 
Hypselodoris (Gastropoda: Opisthobranchia) 


Puesta y desarrollo de especies del género Hypselodoris del Atlántico 
nororiental (Gastropoda: Opisthobranchia) 


Rita COELHO* ** and Goncalo CALADO*** 


Recibido el 23-X-2010. Aceptado el 24-X[1-2010 


SUMMARY 


Despite an existing review of the Atlantic species of the family Chromodorididae (Mol. 
lusca: Nudibranchia], little is known about early development patterns of these species 
due to the difficulty of collecting data from living animals. Six species of the genus 
Hypselodoris inhabit the Portuguese continental coasts: H. bilineata, H. cantabrica, H. 
fontandravi, H. picta, H. tricolor and H. villafranca. This paper is based on new data and 
extensive studies made under laboratory conditions and aims to describe several aspects 
of their reproduction, namely egg mass type, egg size and colour, duration of embryonic 
development and development patterns. The data here obtained and existing developmen- 
tal data for Atlantic Hypselodoris from the literature are also compared. 


RESUMEN 


A pesar de que existe una revisión de las especies atlánticas de la familia Chromodoridi- 
dae (Mollusca: Nudibranchia), muy poco se conoce acerca de la biología y patrones de 
desarrollo de estas especies debido a la dificultad de obtener datos a partir de ejempla- 
res vivos. Seis especies del género Hypselodoris habitan en las costas continentales por- 
tuguesas: H. bilineata, H. cantabrica, H. fontandravi, H. picta, H. tricolor y H. villafranca. 
En el presente trabajo de aportan nuevos datos obtenidos de estudios en condiciones de 
laboratorio con el objetivo de describir algunos aspectos de la biología reproductora de 
estas especies, como el tipo de puesta, tamaño del huevo y color, duración del desarrollo 
embrionario y patrón de desarrollo. Los datos obtenidos en el presente trabajo se (sinteti- 
zan y) comparan con los existentes en la bibliografía. 


INTRODUCTION 

The Family Chromodorididae (Mol- AND GARCÍA-GÓMEZ, 1996) of the At- 
lusca: Nudibranchia) comprises one of lantic species of the family Chromodori- 
the most fantastic coloured Nudibranch didae Bergh, 1891 (Mollusca: Nudi- 
groups of the Iberian Peninsula East branchia), studies specifically devoted 
coast. Despite a review (ORTEA, VALDÉS to their reproductive biology, from egg 


* Instituto Portugués de Malacologia, Zoomarine, E.N. 125, Km 65 Guia; 8201-864 Albufeira; Portugal. E- 
mail: ipmalacOgmail.com 

** Center of Marine Science-CCMAR, University of Algarve, Faro, Portugal. 

*** Universidade Lusófona de Humanidades e Tecnologias, Av. do Campo Grande, 376 
1749 - 024 Lisboa, Portugal. 


63 


Iberus, 28 (2), 2010 


to metamorphosis are absent. General 
data on egg mass shapes, egg size or 
egg-to-juvenile periods are scarce, scat- 
tered or absent. Quite often available 
data come from one single specimen. 
Furthermore, dispersion measures or 
confidence intervals are mostly absent. 

Like with most aspects of nudi- 
branch biology, there is no general rule 
concerning their life history and life 
span. Most species seem to live for 
about one year, although the tropical 
Sea hare Dolabella auricularia has been 
reported to live for six years in an 
aquarium (HADFIELD AND SWITZER- 
DUNLAP, 1984) and the cephalaspidean 
Philine aperta can live for up to four 
years in nature (LANCASTER, 1983). On 
the other hand, there are small nudi- 
branchs, especially those that live and 
feed on short-lived cnidarian colonies 
which can complete a life cycle in a few 
weeks. For example, both the European 
aeolid Tenellia pallida (Alder and 
Hancock, 1842) [= T. adspersa (Nord- 
mann, 1845)] and the coral-eating tropi- 
cal aeolid Cuthona poritophages Rudman, 
1979 can mature in three weeks (RAs- 
MUSSEN, 1944; RUDMAN, 1979) and prob- 
ably die within two or three months. In 
the case of these very short-lived 
species, they need to complete their life- 
cycle before the colony they live and 
feed on dies. If they are too slow then 
they will themselves die from starvation 
before they can breed. 

Six species of the nudibranch genus 
Hypselodoris inhabit the Atlantic coast of 
Portugal (CERVERA, CALADO, GAVAIA, 
MALAQUIAS, TEMPLADO, BALLESTEROS, 
GARCÍA-GÓMEZ AND MEGINA, 2006): 
Hypselodoris billineata (Pruvot-Fol 1953), 
Hypselodoris cantabrica (Bouchet and 
Ortea, 1980), Hypselodoris fontandraui 
(Pruvot-Fol, 1951), Hypselodoris picta 
webbi (D'Orbigny, 1839), Hypselodoris tri- 
color (Cantraine, 1835), and Hypselodoris 
villafranca (Risso, 1818). 

In this paper we report for the first 
time data on the spawn and early devel- 
opment of six sympatric Hypselodoris 


species from Portuguese continental 


coasts maintained under similar labora- 
tory conditions. Several aspects were 


64 


considered, namely egg mass type, egg 
size and colour, duration of embryonic 
development and development type. 


MATERIAL AND METHODS 


Specimens studied were collected on 
subtidal surveys using scuba diving in 
Arrábida (West Coast — 38” 30' 18” N, 8? 
59 18” W) and the Algarve (South coast- 
37” 00 08” N, 7” 49” 20” W), Portugal, 
from April 2004 to June 2005. After col- 
lection, the animals were brought to the 
laboratory and placed in closed-circuit 
40 L aquaria, where water quality was 
monitored daily for temperature, pH 
and salinity, and weekly for the pres- 
ence of nitrites, nitrates and phosphates. 
Individuals were kept at a constant tem- 
perature (18+1*C), as similar as possible 
to that of their natural environment, 
because egg development timing is 
known to be strongly affected by tem- 
perature; this parameter was kept con- 
stant in order to make developmental 
comparisons. Adults of the same species 
were kept together and fed with one of 
their natural prey items, the corneous 
demosponge Dysidea fragilis (Montagu, 
1818), which is very common along the 
Portuguese coasts. 

Egg masses were generally laid on 
the glass walls of the aquaria. The 
oviposition was considered complete 
when the adult abandoned the egg 
mass. Then, the egg masses were care- 
fully removed from the aquaria, incu- 
bated individually in 500 L beakers at a 
constant temperature (18"C+1) and 
checked periodically until larval hatch- 
ing. Extensive observations during 
embryonic development were made 
with an optical microscope at regular 
periods (4-5 times a day) in the centre of 
the egg mass, in order to register the 
main development stages before hatch- 
ing. Daily inspection of all aquaria was 
necessary to ensure the measurement of 
zygote diameters before first cleavage 
since in these planktotrophic species 
initial development is very fast. 

The classification followed for 
Hypselodoris egg masses is the one pro- 


COELHO AND CALADO: Spawn and early development of NE Atlantic Hypselodoris 


Figure 1. Egg ribbons of A. bilineata (A), H. picta webbi (B), and H. villafranca (C). Scale bars, 2 cm. 


Figura 1. Puesta de H. bilineata (A), H. picta webbi (B) y H. villafranca (C). Escalas, 2 cm. 


posed by WILSON (2002), which deals 
exclusively with the family Chromodor- 
ididae. According to this author, 
Hypselodoris egg masses are grouped in 
the following types: A) flat egg masses 
attached to the substratum by the broad 
side of the ribbon, B) egg masses with a 
free edge, shorter or equal than the 
attached edge causing the ribbon to 
slope toward the centre or stand upright 
respectively; C) egg masses with a free 
edge, slightly longer or much longer 
than the attached edge, causing the 
ribbon to slope away from the centre of 
the spiral or causing ondulations/waves, 
with an outward slope. 

The criteria used to classify the 
species larval development were those 
outlined by THomPSON (1967, 1976): 
Type 1) planktotrophic development, 
with a free veliger stage that can be 
pelagic for an extended period of time 
and that feeds obligatorily on plankton; 
Type 2) development with lecitotrophic 
larvae, with a short-life pelagic veliger 
that can dispense with plankton feeding 
due to their large yolk reserves; type 3) 
direct development, without a free 
pelagic larval phase. 


RESULTS 


Egg masses produced in the labora- 
tory by these nudibranchs are identical 
to those collected in the field. All 


Hypselodoris spp. egg masses are shaped 
like a spiral ribbon attached to the sub- 
stratum along one edge and consisting 
of embryos embedded in a gelatinous 
matrix (Fig. 1). 

Hypselodoris egg masses found are 
grouped in two types: H. billineata, H. 
cantabrica, H. fontandraui, H. picta, H. tri- 
color have egg masses corresponding to 
type C of WILSON (2002), whereas H. vil- 
lafranca have type A ones. Developmen- 
tal characteristics of the studied species 
are summarized in Table 1. 

H. villafranca is the only species stud- 
ied with direct development. All ob- 
served egg masses from this species had 
embryos passing through a suppressed 
veliger stage before hatching as benthic 
juveniles. This form of direct develop- 
ment has been categorized by BONAR 
(1978) as ametamorphic direct develop- 
ment, which means that this species 
does not fully develop into a veliger be- 
fore undergoing metamorphosis. 

All other five Hypselodoris species 
studied present a free living plank- 
totrophic veliger, which have similar 
developmental characteristics. Despite 
the differences in egg dimensions (Table 
D), hatching times were also very similar 
(Fig. 2). 

Right after complete oviposition we 
could often observe two to three differ- 
ent stages of development in the same 
spawn. The first part to be released 
could show second cleavage (4 cells), by 


65 


Iberus, 28 (2), 2010 


Table I. Comparative table of developmental characteristics of species of the genus Aypselodoris of the 
Portuguese Coast, at 18+1%C. When appropriate, mean measures are given + standard deviation. 
Tabla I. Tabla comparativa de las características del desarrollo en especies del género Hypselodoris en 
las costas portuguesas, a 18+1"C. 


3 

=> => EA 

El 3 2 3 = E 

> E E 2 E Eg $ 

S 3 E a E Ss B $ 

2 É SS E = e 

5 3 E. a E 5 5d 

3 3 A E ES 

H. bilineata Eastern Atlantic 73.07+7.19  123.16+9.89 1  Plonktotrophic 7,75+0,25 139,5+4,4 10 
H. contabrico  Eosterm Atlantic 95.73+8.41 126.960.228 1  Planktotrophic 8.25+0.35 161.5+6.0 10 


H. fontandravi — Eostem Atlantic 86.41+4.10  135.61+22.15 1  Planktotrophic 7.50+0.14 156.9+5.8 10 


Caribbean and 


H. pict ¡ : 
picto webói Eastern Atlantic 


171.16+10.52 314.73+41.73 1-2 Planktotrophic 8.04+0.13 219.2+8.3 10 


H. tricolor Eastern Atlantic. 85.65+6.97 147.93+25.74 1  Planktotrophic 7.71+0.17 143.2=2.8 10 


Eostern Atlantic, 
except Canary 
Islands. Madeira 
and Azores 


H. villafranca 243.69 + 46.06 389.60+45.32 1 Direct 28.00+0.82 515+23 10 


DUAPIUL LY YIVUP - VAalabIe. HIS Halnctiig 


Time hatching 
_ E _ _— N N 
oO Y co [Ce] o — 
o o o o o Oo 


— 
a 
Oo 


o Mean 
[] Mean + SE 
| Mean + SD 


140 


HB HT HF HC HP 
Species 


Figure 2- Time hatching duration for all planktotrophic Hypselodoris species studied (HB= A. 
bilineata, HT= H. tricolor, HF= H. fontandraui, HC= H. cantabrica, HP= H. picta). 

Figura 2- Tiempo hasta la eclosión de las especies plAde Hypselodoris estudiadas (HB= H. bilineata, 
HT= H. tricolor, HF= H. fontandraui, AC= H. cantabrica, AP= Hypselodoris picta). 


66 


COELHO AND CALADO: Spawn and early development of NE Atlantic Hypselodoris 


Table H. Overview and timing of embryonic development in species of the genus Hypselodoris of 


the Portuguese Coast, at 18+1%C. 


Tabla II. Resumen de la cronología del desarrollo embrionario desarrollo en especies del género 


Hypselodoris en las costas portuguesas, a 18+1"C. 


E E 

2 A S = 
H. bilineata 2h 4h 7h 12 
H. cantabrico Zh 4h 8h 13 
H. fontandravi 2h 5h 7h 14h 
H. picto 2h 3h 30' 8h 13h 
H. tricolor 2h 5h 7h 13h 
H. villafranca 2h 24h 47h 72h 


the time the later eggs were emerging 
from the oviduct, still undivided. The 
stages from morula to blastula were 
observed during day 0 for all species 
except H. villafranca. 

Gastrulation was seen during day 1 
except in H. villafranca, where it was only 
observed by day 9. Only after day 5/6 
could we designate larvae as true 
veligers because a shell and a bilobed 
velum could be clearly seen. In this stage 
veligers were very active inside the cap- 
sules. Duration of the embryonic period 
from egg to gastrula of all species is pre- 
sented in Figure 3. The hatching stage 
occurred between days 7 and 8 after 
oviposition for planktotrophic species. 

The general pattern of cleavage, 
gastrulation and early embryogenesis of 
all species examined is typical of that 
described for other Opisthobranch gas- 
tropods (GOHAR AND  SOLIMAN, 
1967a,b,c). The main embryonic devel- 
opment stages and timings are pre- 
sented in Table II. Most planktotrophic 
species of Hypselodoris studied present a 
veliger with a size within the range of 
139.5 um to 161.5 qm. Exception is made 
for Hypselodoris picta webbi veliger which 
reaches 219.2 um in length at hatching 


= E a 

z 3 2 pa 
A 
E a TE 
= Ss E 3 = 
o a 
= E E 3 3 
17h 1d:14h  2d:19h  5d:20h  7d:18h 
20h 1d:18h 30:13h  6d:4h  8d:1h 
A A 
25h 1d: 21h 3d;18h 6d; 6h 8d; 6h 
20h 1d:14h 24:19 6d:lh 74:17h 
168h 9d 11d; 6h —— 28d 


time. Juveniles of Hypselodoris villafranca 
are dorid-like in shape. They measure 
515+23 um in length at hatching and 
present a translucent mantle with bright 
yellow random spots. A structural spic- 
ular-like network can be seen in the 
mantle tissue. This network is main- 
tained in adults of many dorids but is 
lost in most chromodorids. No eyes or 
rhinophores buds are visible (Fig. 4). 
During the first 10-12h post-hatching 
juveniles crawl on top of the gelatinous 
matrix of the spawn, sometimes 
seeming to graze on it. 

In Table !II we summarise the avail- 
able information on the developmental 
Characteristics of the studied species. 


DISCUSSION 


In this study several aspects of the 
spawn and development of six species 
of the genus Hypselodoris Stimpson, 1855 
are described. 

The type of egg mass presented by 
H. villafranca is quite remarkable, since it 
is to our knowledge the first observation 
of direct development among chro- 
modorids. 


67 


Iberus, 28 (2), 2010 


0 100 200 


[_]  1st Cleavage 


[Al 2nd Cleavage 


a Morula 


300 400 500 600 
Hours 
ES 3rd Cleavage Blastula 
ES] Gastrula 


Figure 3. Embryonic period duration from egg to gastrula in all species studied of Hypselodoris. 
Figura 3. Duración del desarrollo embrionario hasta el estado de gástrula. 


Concerning egg size, the mean 
diameter in H. bilineata, H. cantabrica, H. 
fontandraui and H. tricolor is below 100 
um, within the range reported for most 
Nudibranchia species (HADFIELD AND 
MILLER, 1987) and consistent with the 
planktotrophy exhibited. Egg and 
capsule sizes are classically considered 
to be good predictors of development 
type (HADFIELD AND SWITZER-DUNLAP, 
1984) and used for comparative pur- 
poses (HADFIELD AND MILLER, 1987). In 
the case of the Atlantic Hypselodoris 
picta, however, this extrapolation 
clearly predicts direct development, as 
in H. villafranca, but fails to predict 
pelagic-lecithotrophy which should be 
the case of H. picta webbi if one follows 
HADFIELD AND  SWITZER-DUNLAP's 
(1984) predictions. The same situation 
was reported by THOMPSON (1967) for 
Archidoris pseudoargus, a planktotrophic 
developer whose eggs are 170 um in 
diameter. A free-swimming, plank- 
totrophic veliger can emerge from eggs 
of very different sizes, from 73 um in H. 
bilineata (average adult length 30mm), 


68 


to 171 ym in H. picta webbi (average 
adult length 100mm). The latter is the 
largest planktotrophic egg size reported 
so far among nudibranchs (see TODD, 
LAMBERT AND bDAvIEs, 2001 for a 
review). Clearly the amount of energy 
packed into each ovum is very differ- 
ent. Nevertheless, survivorship and 
time to competence in these larvae 
remains unknown. The difference in 
hatching time between species with 
planktotrophic larvae and the one with 
direct development is certainly due to 
developmental constraints. Even so, the 
total egg-to-juvenile period is generally 
longer and variable in the plank- 
totrophic strategy since it is dependent 
of an external source of energy to 
undergo metamorphosis (ToDD, 1983; 
HAVENHAND, 1993). 

One of the reasons for supporting 
egg size with capsule size data, in order 
to have a more accurate development 
extrapolation (CLARK AND JENSEN, 1981; 
HADFIELD AND MILLER, 1987) is the fact 
that there are species with lecithotrophic 
or even direct development although 


COELHO AND CALADO: Spawn and early development of NE Atlantic Hypselodoris 


Figure 4. Newly hatched juvenile of H. villafranca. 


Figura 4. Juvenil recién eclosionado de H. villafranca. 


having small eggs. This is common 
among Sacoglossa species as stated by 
JENSEN (2001). The situation can be 
explained based on the presence of 
albumen, an intracapsular substance 
rich in proteins and very common in 
Opisthobranchia spawns (CLARK AND 
JENSEN, 1981) and the existence of extra- 
capsular yolk in some species, which 
gives additional nutritional support for 
embryos, larvae and juveniles (MARIN 
AND Ros, 1993). 

All studied species show a holoblas- 
tic and spiral cleavage, typical of the 
Gastropoda group (BIGGELAAR AND 
HASZPRUNAR, 1996). With the exception 
of H. villafranca with its slower direct 
development, all species reach morula 
stage 12-14h after oviposition and gas- 
trula stage before the 48h mark. Both 
planktotrophic and direct development 
species gastrulate by emboly, as 
described for Dendrodoris and Chro- 
modoris by GOHAR AND  SOLIMAN 
(1967a,b) and for Platydoris by SOLIMAN, 
(1978). 


The development from gastrula 
stage to veliger, with all typical struc- 
tures present lasts between 102-107h. 

It is obvious that Opisthobranch 
life-cycle and developmental strategies 
are inextricably linked to adult food 
specificity. Species that feed on tran- 
sient food organisms are usually small 
and present a short life cycle, with a 
very rapid embryonic development. On 
the other hand, species, like the 
Hypseloris here presented, that depend 
on large long-living colonies such as 
sponges, are often bigger in size and 
have longer life cycles with slower 
embryonic development (RUDMAN AND 
WILLAN, 1998). 

Although some  developmental 
timetables have been reported for a 
number of opisthobranch species, it is 
often difficult to use them to make com- 
parisons due to uncontrolled husbandry 
conditions or severe differences in water 
culture temperature, which is known to 
be one of the key factors affecting 
embryonic development. 


69 


Iberus, 28 (2), 2010 


Table III. Summary of developmental characteristics of Atlantic species of the genus Aypselodoris 


from available literature and present study. 


SPECIES Type of egg mass Egg mass color 
H. bilineata 1 whorl spiral ribbon White 
Spiral ribbon White 
3 whorls ribbon Red-orange 
2 Whorls spiral ribbon White 
2-2,5 Whorls Spiral ribbon White 
2 Whorls spiral ribbon White 
H. cantabrica 4 whorls spiral ribbon White 
Spiral ribbon White 
Spiral ribbon White 
4 whorls spiral ribbon White 
H. fontandravi Spiral ribbon White 
Spiral ribbon White 
Spiral ribbon White 
2 Whorls spiral ribbon White 
H. picta webbi Waved spiral ribbon Red-orange 
Smoothly waved spiral ribbon Orange 
5 Whorls waved spiral ribbon 
4 Whorls waved spiral ribbon  Pink-orange 
H. tricolor 3 whorls spiral ribbon White 
Spiral ribbon White 
2.5 Whorls Spiral ribbon White 
2 Whorls spiral ribbon White 
H. villafranca 2 Whorls spiral ribbon Orange 
Spiral ribbon Orange 
1 whorl spiral ribbon Orange 


The studies on development pre- 
sented by MARTÍNEZ-PITA, SÁNCHEZ- 
ESPANÑA AND GARCÍA (2006) on species 
of Polycera, conducted at 19C, report 
similar embryonic stage timings to those 
presented for planktotrophic species of 
Hypselodoris in the current study. 
YONOW (1996) cultured Acteon tornatilis 
at 12"C water temperature and noted 
that the 4-cell stage was reached about 
24h after oviposition and the gastrula 
stage within 4.9-6d. Although there are 
no significant differences in develop- 
ment pattern between compared 
species, these last data report a much 
slower embryonic development than 
our study, which is probably due to the 
6”C difference in water temperature. 

All Hypselodoris planktotrophic species 
exhibit a similar embryonic growth 


7O 


Egg mass width (mm) No. of specimens Type of capsule 


3 
5 l 
4 
3 / Oval 
8 Spherical 
l Oval 
] 
10 Oval 
| 
- 10 
6 
10 1 
2 y . 
- 3 Spherical 
3 5 - 
2.5mm - - 
- 2 Oval or spherical 


Spherical 


pattern and despite differences in egg size 
no differences are registered for hatching 
times. Hypselodoris picta webbi presents the 
biggest veliger (219.2+8.3 yum) but this size 
discrepancy with other planktotrophic 
species seems to be more related to adult 
size (approximately 100mm length) than 
to any developmental pattern. Neverthe- 
less, survivorship and time to competence 
in these larvae remains unknown. 

When  hatching,  morphological 
resemblance of Hypselodoris villafranca 
crawling juveniles to adult specimens is 
very limited and no rhinophores are 
visible. The crawling behaviour that the 
juvenile exhibits for 10-12h on top of the 
gelatinous matrix of the spawn, seeming 
to graze on it, leads us to think that 
matrix nutrients are not only important 
during embryonic development but 


COELHO AND CALADO: Spawn and early development of NE Atlantic Aypselodoris 


Tabla III. Resumen de las principales aspectos del desarrollo de las especies atlánticas del género Hypse- 
lodoris obtenidas a partir de la bibliografía y del presente estudio. 


Capsule diameter (1m) No. ofeggs per copsule Egg diameter (t1m)  Embryonic period in days SOURCE 
100-120 - 79-109 Ortea et al (1996) 
110-120 l - Garcia Gómez (2002) 
E 175-208 - Bouchet and Ortea (1980) 
120 l 11 (18-20%) Gantes (1962) 
158(=+3,5)| l 85-100 9 (21%) Sánchez-Tocino ef al (2007) 
100-133 l 67-100 1.1520:25 Current Study 
. - 95-125 1 (22%) Ortea ef al (1996) 
90-140 l - - Garcia Goméz (2002) 
95-125 l 80-85 Sánchez-Tocino ef al (2007) 
117-144 l 75-120 8.25+0.35 Current Study 
150-180 - 110-120 - Ortea ef al. (1996) 
150-180 1 - - Garcia-Gómez (2002) 
145-180 1 110 13 (18%) Sánchez-Tocino ef al. (2007) 
95-233 1 80-100 7.50+0.14 Current Study 
- - 175-208 - Ortea et al (1996) 
160-190 l - Garcia-Gómez (2002) 
170x200 12 130-135 - Sánchez-Tocino ef al. (2007) 
240-453 1-2 141-210 8.04+0.13 Current Study 
90-120 - 85-100 13 (20-22%) Ortea et al. (1996) 
90-120 l - - Garcia-Gómez (2002) 
170x200 1-2 130-135 - Sánchez-Tocino ef al. (2007) 
85-181 l 71-90 IAEA Current Study 
300-400 - Bit smaller than capsules - Ortea ef al. (1996) 
320-360 l - - Garcia-Gómez (2002) 
300-599 | 173-430 28.00+0.82 Current Study 
serve as first food for juveniles after ACKNOWLEDGMENTS 


hatching (GIBSON AND CHIA, 1991). 
Although some work is still required 
to understand some of the differences 
here recognized in development pat- 
terns between these species, the data 
presented in this work with a systema- 
tised character and controlled hus- 
bandry conditions of specimens, will 
certainly be useful regarding potential 
taxonomic or phylogenetic discussions. 


BIBLIOGRAPHY 


BIGGELAAR J. M. AND HASZPRUNAR G. 1996. 
Cleavage patterns and mesentoblast forma- 
tion in the Gastropoda: An evolutionary per- 
spective. Evolution, 50 (4): 1520-1540 


This work was carried out under a 
research project funded by Prémio 
Milénio Sagres/Expresso 2002. R.C. 
holds a grant from the Fundacáo para a 
Ciéncia e Tecnologia, Portugal (BDE 
15577/2005). António Monteiro, and 
Manuel Malaquias, read an early 
version of the manuscript and helped to 
improve it. 


BONAR D. B. 1978. Morphogenesis at meta- 
morphosis in opisthobranch molluscs. In 
Chia F.S. and Rice M.E. (Eds.): Settlement and 
metamorphosis of marine invertebrate larvae. El- 
sevier North-Holland, New York: 177-196. 


71 


Iberus, 28 (2), 2010 


CERVERA J.L., CALADO G., GAVAIA C., 
MALAQUIAS M.A.E., TEMPLADO J., BALLES- 
TEROS M., GARCÍA-GÓMEZ J.C. AND MEGINA 
C. 2006. An annotated and updated check- 
list of the opisthobranchs (Mollusca: Gas- 
tropoda) from Spain and Portugal (including 
islands and archipelagos). Boletin del Instituto 
Español de Oceanografía, 20 (1-4): 5-111. 

CLARK K.B. AND JENSEN K.R. 1981. A compar- 
ison of egg size, capsule size, and develop- 
ment patterns in the order Ascoglossa 
(Sacoglossa) (Mollusca: Opisthobranchia). 
International Journal of Invertebrate Reproduc- 
tion 3: 57-68 

GIBSON G.D. AND CHIA F. 1991. Contrasting 
reproductive modes in two sympatric 
species of Haminaea (Opisthobranchia: 
Cephalaspidea). Journal of Molluscan Stud- 
1es, 57: 49-60. 

GOHAR H.A.F. AND SOLIMAN G.N. 1967a. The 
biology and development of Dendrodoris (Do- 
riopsis) fumata (Rúpp. and Leuck.) (Gas- 
tropoda, Nudibranchia). Publications of the 
Marine Biological Station, Al-Ghardaga, Egypt, 
14: 31-54. 

GOHAR H.A.F. AND SOLIMAN G.N. 1967b. The 
biology and development of Chromodoris in- 
ornata Pease (Gastropoda, Nudibranchia). 
Publications of the Marine Biological Station, 
Al-Ghardaga, Egypt, 14: 78-95. 

GOHAR H.A.F. AND SOLIMAN G.N. 1967c. The 
biology and development of Chromodoris tinc- 
toria (Rúpp. and Leuck.) (with reference to the 
taxonomic value of spawning characters). 
Publications of the Marine Biological Station, 
Al-Ghardaga, Egypt, 14: 95-108. 

HADFIELD M.G. AND MILLER S.E. 1987. On de- 
velopmental patterns of Opisthobranchs. 
American Malacological Bulletin, 5 (2): 197-214. 

HADFIELD M. AND SWITZER-DUNLAP M. 1984. 
Opisthobranchs. In Wilbur K. (Ed.): The Mo- 
llusca, 7: Reproduction. Academic Press, New 
York: 209-350. 

HAVENHAND J. 1993. Egg to juvenile period, 
generation time, and the evolution of larval 
type in marine invertebrates. Marine Ecology 
Progress Series, 97: 247-260. 

JENSEN K.R. 2001. Review of reproduction in the 
Sacoglossa (Mollusca, Opisthobranchia). Bol- 
lettino Malacologico, 37, 81-98 

LANCASTER S.M. 1983. The biology and repro- 
ductive ecology of Philine aperta (Opistho- 
branchia: Bullomorpha) in Oxwich Bay. 
Journal of Molluscan Studies, Suppl 12A: 82- 
88 

MARIN A. AND ROS J.D. 1993. Ultrastructural 
ecological aspects of the development of 
chloroplast retention in the sacoglossan Elysia 
timida. Journal of Molluscan Studies, 59: 95- 
104. 


72 


MARTÍNEZ-PITA I.; SÁNCHEZ-ESPANÑA A.Í. AND 
GARCÍA FJ. 2006. Some aspects of the repro- 
ductive biology of two Atlantic species of 
Polycera (Mollusca: Opisthobranchia) Jour- 
nal of the Marine Biological Association of the 
U.K. 86 (2): 391-399. 

MCDONALD, G. AND NYBAKKEN J. 1997. A List 
of the Worldwide Food Habits of Nudi- 
branchs. http: / /people.ucsc.edu/=mecduck/ 
nudifood.htm (viewed October 2005) 

ORTEA J., VALDÉS A. AND GARCÍA-GÓMEZ J.C. 
1996. Revisión de las especies atlánticas de 
la familia Chromodorididae (Mollusca: Nudi- 
branchia) de grupo cromático azul. Avicen- 
nia, Suppl. 1: 1-165. 

RASMUSSEN E. 1944. Faunistic and biológical 
notes on marine invertebrates. Videnskabelige 
Meddelelserfra Dansk Naturhistorisk Forening, 
107: 207-33. 

RUDMAN, W.B. 1979. The ecology and anatomy 
of a new species of aeolid opisthobranch 
mollusc: a predator of the reef-forming coral 
Porites. Zoological Journal of the Linnean Soci- 
ety, 65: 339-359. 

RUDMAN W.B. AND WILLAN R.C. 1998. Opist- 
hobranchia. In Beesley P.L., Ross G.J.B. and 
Wells, A. (Eds.): Mollusca: The Southern syn- 
thesis. Fauna of Australia. CSIRO, Melbourne, 
pp. 915-1035. 

SOLIMAN G.N. 1978. The redescription, repro- 
duction and development of the dorid nudi- 
branch Platydoris Scabra (Cuvier) from the 
Northwestern Red Sea, Journal of Molluscan 
Studies, 44 (2): 151-165. 

STRATHMANN R.R. 1978. The evolution and loss 
of feeding larval stages of marine inverte- 
brates. Evolution, 32: 894-906. 

THOMPSON T.E. 1967. Direct development in 
the nudibranch Cadlina laevis, with a discus- 
sion of developmental processes in Opistho- 
branchia. Journal of Marine Biological Associ- 
ation of the United Kingdom, 47: 1-22. 

THOMPSON T.E. 1976. Biology of Opistobranch 
molluscs 1. London: The Ray Society 

Topp C.D. 1983. Reproductive and trophic ecol- 
Ogy of nudibranch molluscs. In Russel- 
Hunter W.D. (Ed.): The Mollusca. 6: Ecology. 
Academic Press, New York, 6: 225-259. 

Topp C.D., LAMBERT W.J. AND DAVIES J. 2001. 
Some perspectives on the biology and ecol- 
ogy of nudibranch molluscs: generalisations 
and variations on the theme that prove the 
rule. Bollettino Malacologico, 37: 105-120. 

YONOW N. 1996. Gametogenesis, egg produc- 
tion and development'in Acteon tornatilis 
(Opisthobranchia: Cephalaspidea). Malaco- 
logical Review, Supplement 6: 31-52. 

WILSON N. 2002. Egg masses of chromodorid 
nudibranchs (Mollusca: Gastropoda: Opistho- 
branchia). Malacología, 44 (2): 289-305. 


ERRATA en Rolán y Fernández-Garcés (Iberus, 28 (1) 
p. 84-86) sobre R. cancellina spec. nov. 


Sustituir el primer párrafo de "Remarks" por el siguiente: 


Remarks: The holotype is a shell in good conditions and good protoconch (Fig. 3A) 
with 5.7 mm; one paratype is a shell with 8.5 mm (ZMB 115039) and labelled as Ris- 
soina cancellata Phil. v. pulchra C. B. Adams, Jamaica, coll. Paetel (Fig. 3B). 


pe 
1 


O 1 
A es 


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e Los manuscritos, así como toda la correspondencia relacionada con los mismos, deberán ser remitidos a: 
Serge Gofas, Editor de Publicaciones, Departamento de Biología Animal, Universidad de Málaga, Campus de 
Teatinos, s/n, 29071, Málaga, España y/o al correo electrónico <sgofasQuma.es>. 

e El texto del trabajo podrá estar redactado en español, inglés, italiano, francés o portugués. 

e Los artículos, notas y monografías deberán presentarse sobre DIN A-4, por una sola cara a doble espacio 
(incluyendo referencias, pies de figura y tablas), con al menos 3 centímetros de margen por cada lado. Los tra- 
bajos se entregarán por triplicado (original y dos copias) y se incluirá una versión en un CD, o bien remitida 
por correo electrónico, utilizando procesadores de texto en sus versiones de corrientes de Windows (pero no 
en el formato .docx de Word 2007, el habitual de Windows Vista) o Macintosh. En caso de autoría compartida, 
uno de los autores deberá hacerse responsable de toda la correspondencia. 

e Junto con el trabajo debe incluirse una lista de al menos 4 posibles revisores del mismo, sin perjuicio de los 
que el propio Editor considere oportunos. 

+ Los manuscritos se presentarán de acuerdo al siguiente esquema: 

Primera página. Deberá incluir un título conciso, pero sugerente del contenido del trabajo, así como una traduc- 
ción al inglés del mismo (si el artículo no está escrito en inglés). Cuando sea preciso, deberá incluir referencia a 
familia o taxones superiores. A continuación figurarán, por este orden, el nombre y apellidos completos del 
autor o autores, las direcciones completas de los mismos, y un resumen del trabajo y su traducción al inglés. 
Dicho resumen deberá sintetizar, en conjunción con el título, tanto los resultados como las conclusiones del 
artículo; se sugiere una extensión de 100 a 200 palabras. 

Páginas siguientes. Incluirán el resto del artículo, que debe dividirse en secciones precedidas por breves encabe- 
zamientos. Siempre que sea posible, se recomienda seguir el siguiente esquema: Introducción, Material y 
métodos, Resultados, Discusión, Conclusiones, Agradecimientos y Bibliografía. Si se emplean abreviaturas no 
habituales en el texto, deberán indicarse tras el apartado de Material y Métodos. 

e Las notas breves deberán presentarse de la misma forma, pero sin resumen. 

e Deberán evitarse notas a pie de página y referencias cruzadas. Deberán respetarse estrictamente los Códigos 
Internacionales de Nomenclatura Zoológica y Botánica (últimas ediciones). Cuando un taxón aparezca por 
primera vez deberá citarse su autor y fecha de su descripción. En el caso de artículos sistemáticos, cuando se 
den las sinonimias de los taxones, éstas deberán citarse COMPLETAS, incluyendo en forma abreviada la 
publicación donde fueron descritas, y la localidad tipo si es conocida entre corchetes, según el siguiente 
esquema (préstese especial cuidado a la puntuación): 

Dendrodoris limbata (Cuvier, 1804) 

Sinonimias 

Doris limbata Cuvier, 1804, Ann. Mus. Hist. Nat. Paris, 4 (24): 468-469 [Localidad tipo: Marsella]. 

Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop.-Car., 10: 275. 

Dichas referencias no deberán incluirse en la lista de Bibliografía si es la única vez que se nombran en el texto. 
Si se incluyen una lista completa de referencias de un taxón inmediatamente tras éste, deberá seguirse el 
mismo esquema (sin incluir en Bibliografía las referencias que no se mencionen en otro lugar del texto). 

e Sólo los nombres en latín y los de taxones genéricos y específicos deberán llevar subrayado sencillo o prefe- 
rentemente ir en cursiva. En ningún caso deberá escribirse una palabra totalmente en letras mayúsculas, ni 
siquiera el Título. Las unidades a utilizar deberán pertenecer al Sistema Métrico Decimal, junto con sus correc- 
tas abreviaturas. En artículos escritos en castellano, en los números decimales sepárese la parte entera de la 
decimal por una coma inferior (,), NUNCA por un punto (.) o coma superior (*). 

e Las referencias bibliográficas irán en el texto con minúsculas o versalitas: Fretter y Graham (1962) o FRETTER 
Y GRAHAM (1962). Si son más de dos autores se deberán citar todos la primera vez que aparecen en el texto 
[Smith, Jones y Brown (1970)] empleándose et al. las siguientes veces [Smith et al. (1970)]. Si un autor ha publi- 
cado más de un trabajo en un año se citarán con letras: (Davis, 1989a; Davis, 1989b). No deberá emplearse op. 
cit. La lista de referencias deberá incluir todas las citas del texto y sólo éstas, ordenadas alfabéticamente. Se 
citarán los nombres de todos los autores de cada referencia, sea cual sea su número. Los nombres de los 
autores deberán escribirse, en letras minúsculas o Versalitas. No deberán incluirse referencias a autores 
cuando éstos aparezcan en el texto exclusivamente como autoridades de un taxón. Los nombres de las publica- 


ciones periódicas deberán aparecer COMPLETOS, no abreviados. Cuando se citen libros, dése el título, editor, 
lugar de publicación, n* de edición si no es la primera y número total de páginas. Deberán evitarse referencias 
a Tesis Doctorales u otros documentos inéditos de difícil consulta. Síganse los siguientes ejemplos (préstese 
atención a la puntuación): 

Fretter V. y Graham A. 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp. 

Ponder W.F. 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. En Ponder, W.F. 
(Ed.): Prosobranch Phylogeny. Malacological Review, supp!. 4: 129-166. 

Ros J. 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Mis- 
celánea Zoológica, 3 (5): 21-51. 

e Las gráficas e ilustraciones deberán ser originales y presentarse preferentemente en formato electrónico al 
formato de caja de la revista o proporcional a éste. Este formato es de 57 mm (una columna) o 121 mm (dos) de 
anchura y hasta 196 mm de altura, si bien se recomienda utilizar el formato a dos columnas. Es importante que 
todas las figuras sean remitidas en su formato original (por ejemplo, las fotografías en jpg de alta calidad o .tif, 
las gráficas en hojas de cálculo Excel o documentos de CorelDraw), puesto que las ilustraciones insertadas en 
el manuscrito WORD son inservibles en la fase de imprenta. Las imágenes digitales deben ser formateadas en 
su tamaño de impresión con una resolución mínima de 300 ppp para imágenes en color o escala de grises y de 
600 ppp para las de blanco y negro. Considérese la reducción que será necesaria a la hora de decidir el tamaño 
de las escalas o letras en las figuras, que no deberán bajar de los 2 mm. En figuras compuestas, cada parte 
deberá etiquetarse con letras mayúsculas, el resto de las letras deberán ser minúsculas. Las escalas de dibujos y 
fotografías deberán ser gráficas, utilizando unidades del sistema métrico decimal; no deberán hacerse referen- 
cias a los aumentos de una determinada ilustración, ya que éstos cambian con la reducción. En su caso, se 
recomienda la utilización de mapas con proyección UTM. Cada figura, gráfica o ilustración deberá presentarse 
en hojas separadas y con numeración arábiga (1, 2, 3,...), sin separar “Figuras” y “Láminas”. Los pies de 
figura, en una hoja aparte, deberán acompañarse de su traducción al inglés (o español si el inglés es la lengua 
del trabajo). Utilícese el esquema siguiente: 

Figura 1. Neodoris carvi. A: animal desplazándose; B: detalle de un rinóforo; C: branquia. 

Las abreviaturas empleadas en las ilustraciones deberán incluirse en los pies de figura. 

Los autores interesados en incluir láminas en color deberán consultar con el editor y sufragar el sobrecoste con 
una contribución de 30 € por página. Por lo demás, deberán ajustarse a los mismos requisitos indicados para 
las figuras. 

Si se pretende enviar gráficas o ilustraciones en impresión de papel es imprescindible presentar originales de 
buena calidad. Las imágenes en semitonos deben estar bien contrastadas y ajustarse al tamaño definitivo de 
impresión; al componer fotografías sobre una hoja, procúrese que los espacios entre ellas sean regulares y que 
estén debidamente alineadas. Téngase en cuenta que incluir fotografías de distinto contraste en una misma 
página conlleva una pobre reproducción final. Las gráficas de ordenador deberán imprimirse con impresora 
láser sobre papel de buena calidad. 

e Las Tablas se presentarán en hojas separadas, siempre con numeración romana ([, II, 1IL....). Las leyendas se 
incluirán en una hoja aparte acompañándose de una traducción al inglés. Deberán evitarse las tablas particu- 
larmente complejas. Se recomienda reducir el número y extensión de ilustraciones, láminas o tablas al mínimo 
necesario. 

e Los artículos que no se ajusten a las normas de publicación serán devueltos al autor con las indicaciones de 
los cambios necesarios. 

e El Comité Editorial comunicará al autor responsable del trabajo la fecha de recepción del trabajo y la fecha 
de envío a revisión. Cada original recibido será sometido a revisión por al menos dos investigadores. El 
Comité Editorial, a la vista de los informes de los revisores decidirá sobre la aceptación o no de cada manus- 
crito. El autor recibirá en cada caso copia de los comentarios de los revisores sobre su artículo. En caso de 
aceptación, el mismo Comité Editorial, si lo considera conveniente, podrá solicitar a los autores otras modifica- 
ciones que considere oportunas. Si el trabajo es aceptado, el autor deberá enviar una copia impresa del mismo 
corregida, acompañada por una versión en un CD, utilizando procesadores de texto en sus versiones corrien- 
tes de Windows (pero no el formato .docx generado por Word 2007, el habitual de Windows Vista) o Macin- 
tosh. La fecha de aceptación figurará en el artículo publicado. ; : 

e Las pruebas de imprenta serán enviadas por correo electrónico al autor responsable, exclusivamente para la 
corrección de erratas, y deberán ser devueltas en un plazo máximo de una semana. Se recomienda prestar 
especial atención en la corrección de las pruebas. 

e De cada trabajo se entregarán gratuitamente 50 separatas, además de una versión electrónica en formato 
pdf. Aquellos autores que deseen un número mayor, deberán hacerlo constar al devolver las pruebas de 
imprenta, y nunca posteriormente. El coste de las separatas adicionales será cargado al autor. 


INSTRUCTIONS TO AUTHORS 


* Iberus publishes research papers, notes and monographs devoted to the various aspects of Malacology. 
Papers are manuscripts of more than 5 typed pages, including figures and tables. Notes are shorter papers. 
Monographs should exceed 50 pages of the final periodical, and may be published as Supplements. Authors 
wishing to publish monographs should contact the Editor. Manuscripts are considered on the understanding 
that their contents have not been published or simultaneously submitted for publication elsewhere. 

e Manuscripts and correspondence regarding editorial matters must be sent to: Serge Gofas, Editor de Publica- 
ciones, Departamento de Biología Animal, Universidad de Málaga, Campus de Teatinos, s /n, 29071, Málaga, 
Spain and /or to the e-mail <sgofasQuma.es>. 

+ Manuscripts may be written in Spanish, English, Italian, French or Portuguese. 

e Manuscripts must be typed double spaced (including the references, figure captions and tables) on one side 
on A-4 (297x210 mm) with margins of at least 3 cm. An original and two copies must be submitted, together 
with a CD or e-mail remittance containing the article written with a current Windows (but not .docx format 
generated by Word 2007, mainly used with Windows Vista) or Macintosh word processor. When a paper has 
joint authorship, one author must accept responsibility for all correspondence. 

e The authors must include a list of at least 4 possible referees; the Editor can choose any others if appropriate. 

e Papers should conform the following layout: 

First page. This must include a concise but informative title, with mention of family of higher taxon when 
appropriate, and its Spanish translation. It will be followed by all authors” names and surnames, their full 
address(es), an abstract (and its Spanish translation) not exceeding 200 words which summarizes not only con- 
tents but results and conclusions. 

Following pages. These should content the rest of the paper, divided into sections under short headings. When- 
ever possible the text should be arranged as follows: Introduction, Material and methods, Results, Discussion, 
Conclusions, Acknowledgements and References. Unusual abbreviations used in the text must be grouped in 
one alphabetic sequence after the Material and methods section. 

e Notes should follow the same layout, without the abstract. 

e Footnotes and cross-references must be avoided. The International Codes of Zoological and Botanical 
Nomenclature must be strictly followed. The first mention in the text of any taxon must be followed by its 
authority including the year. In systematic papers, when synonyms of a taxon are given, they must be cited IN 
FULL, including the periodical, in an abbreviate form, where they were described, and the type localities in 
square brackets when known. Follow this example (please note the punctuation): 

Dendrodoris limbata (Cuvier, 1804) 

Synonyms 

Doris limbata Cuvier, 1804, Ann. Mus. Hist. Nat. 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. If a 
full list of references of the taxon is to be given immediately below it, the same layout should be followed (also 
excluding from the Bibliography list those which are not cited elsewhere). 

Only Latin words and names of genera and species should be underlined once or be given in italics. No word 
must be written in UPPER CASE LETTERS. SI units are to be used, together with their appropriate symbols. In 
Spanish manuscripts, decimal numbers must be separated with a comma (,), NEVER with a point (.) or upper 
comma (*). 

e References in the text should be written in small letters or Small capitals: Fretter and Graham (1962) or 
FRETTER AND GRAHAM (1962). The first mention in the text of a paper with more than two authors must 
include all of them [Smith, Jones and Brown (1970)], thereafter use et al. [Smith et al. (1970)]. If an author has 
published more than one paper per year, refer to them with letters: (Davis, 1989a; Davis, 1989b). Avoid op. cit. 

The references in the reference list should be in alphabetical order and 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 tax- 
onomic name. Titles of periodicals must be given IN FULL, not abbreviated. For books, give the title, name of 
publisher, place of publication, indication of edition if not the first and total number of pages. Keep references 
to doctoral theses or any other unpublished documents to an absolute minimum. See the following examples 
(please note the punctuation): 

Fretter V. and Graham A. 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp. 

Ponder W.F. 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. In Ponder W.F. 
(Ed.): Prosobranch Phylogeny. Malacological Review, suppl. 4: 129-166. 

Ros J. 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Mis- 
celánea Zoológica, 3 (5): 21-51. 


e Figures must be original and provided preferably in electronic format and adjusted to page format and 
column size. These should be one column (57 mm) or two columns (121 mm) wide and up 196 mm high, or be 
proportional to these sizes. Two columns format is recommended. It is essential that all figures be supplied in 
their original format (e.g. photographs as high-grade .jpg or as .tif files, graphs as Excel spreadsheets or Corel- 
Draw files), as the files inserted into WORD documents cannot be used for printing. Digital images must be 
given their final printing size with a resolution at least 300 dpi for colour and halftones, and at least 600 dpi for 
black /white. 

Take into account possible reduction in lettering drawings; final lettering must be at least 2 mm high. In com- 
posite drawings, each figure should be given a capital letter; additional lettering should be in lower-case 
letters. A scale line, labelled with SI units, must be used to indicate size; magnification ratio must be avoided 
as it may be changed during printing. UTM maps are to be used if necessary. Figures must be submitted on 
separate sheets, and numbered with consecutive Arabic numbers (1, 2, 3,...), without separating “Plates” and 
“Figures”. Legends for Figures must be typed in numerical order on a separate sheet, and a Spanish translation 
must be included. Follow this example (please note the punctuation): 

Figure 1. Neodoris carvi. A: animal crawling; B: rhinophore; C: gills. 

If abbreviations are to be used in illustrations, they should be included in the figure captions. 

Authors wishing to publish illustrations in colour should consult with the editor and will be charged a contri- 
bution of 30 € per page. They should otherwise follow the same standards as black and white prints. 

If the authors want to send Figures in printed format, it is essential to supply good quality originals. Half-tone 
images must be of good contrast, and should be submitted in the final printing size. When mounting pho- 
tographs in a block, ensure spacers are of uniform width. Remember that grouping photographs of varied con- 
trast results in poor reproduction. Computer graphics must be printed on high quality white paper with a 
laser printer. 

e Tables must be numbered with Roman numbexs (1, IL, HL...) and each typed on a separate sheet. Headings 
should be typed on a separate sheet, together with their English translation. Complex tables should be 
avoided. As a general rule, keep the number and extension of illustrations and tables as reduced as possible. 

e Manuscripts that do not conform to these instructions will be returned for correction before reviewing. 

e Authors submitting manuscripts will receive an acknowledgement of receipt, including receipt date, and the 
date the manuscript was sent for reviewing. Each manuscript will be critically evaluated by at least two refer- 
ees. Based of these evaluations, the Editorial Board will decide on acceptance or rejection. Anyway, authors 
will receive a copy of the referees” comments. If a manuscript is accepted, the Editorial Board may indicate 
additional changes if desirable. Acceptable manuscripts will be returned to the author for consideration of 
comments and criticism; a finalized manuscript must then be returned to the Editor, together with a CD con- 
taining the article written with current Windows (but not a .docx format generated by Word 2007, mainly used 
under Windows Vista) or Macintosh word processor. Dates of reception and acceptance of the manuscript will 
appear in all published articles. 

e Proofs will be e-mailed to the author for correcting errors and must be returned corrected within one week. 
At this stage no stylistic changes will be accepted. Pay special attention to references and their dates.in the text 
and the Bibliography section, and also to numbers of Figures and Tables appearing in the text. 

e Fifty reprints per article and a .pdf file will be supplied free of charge. Additional reprints must be ordered 
when the page proofs are returned, and will be charged at cost. NO LATER orders will be accepted. 


La SOCIEDAD ESPAÑOLA DE MALACOLOGÍA 


Junta Directiva desde el 11 de octubre de 2005 


Presidente José Templado González 
Vicepresidente Emilio Rolán Mosquera 
Secretaria María Carmen Salas Casanovas 
Tesorero Luis Murillo Guillén 
“Editor de Publicaciones Serge Gofas 
Bibliotecario Rafael Araujo Armero 
Vocales Ramon M. Álvarez Halcon 
Benjamín Gómez Moliner 
Alberto Martínez Ortí 


Diego Moreno Lampreave 
José Ramón Arrébola Burgos 


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 2010: 


Socio numerario (en España): 40 euros 
(en Europa) 40 euros 
(fuera de Europa): 48 euros 
Socio estudiante (en España): 23 euros 
(en el extranjero): 29 euros 
Socio Familiar: (sin recepcion de revista) 4euros 
Socio Protector: (mínimo) 48 euros 
Socio Corporativo (en Europa): 48 euros 
(fuera de Europa): 54 euros 


INSCRIPCIÓN: 6 euros, además de la cuota correspondiente. 


-Alos socios residentes en España se les aconseja domiciliar su cuota. Todos los abonos deberán enviarse al 
Tesorero (dirección reseñada anteriormente) el 1 de enero de cada año. Los abonos se harán sin recargos para la 
sociedad y en favor de la Sociedad Española de Malacología y no de ninguna persona de la junta directiva. Aque- 
llos socios que no abonen su cuota anual dejarán de recibir las publicaciones de la Sociedad. Los bonos de ins- 
cripción se enviarán junto con el abono de una cuota anual al Tesorero. 

A los residentes en el extranjero se les ruega que abonen su cuota mediante giro postal en euros (internatio- 
nal postal money orders in euros sent to the Treasurer). Members living in foreing countries can deduce 6 euros 


if paid before 15 April. 


Cada socio tiene derecho a recibir anualmente los números de /berus, Reseñas Malacológicas y Noticiarios que 
se publiquen. 


N LIBRARIES 


| 


| 


WICNOLON O 


9088 01570 


ÍNDICE 
Iberus 28 (2) 2010 


HORRO J., GORI S. AND ROLÁN E. Haedropleura ryalli, a new species from Sáo Tomé Island (Gas- 
tropoda, Turridae) 
Haedropleura ryalli, nueva especie de la isla de Sáo Tomé (Gastropoda, Turridae) ..... 1-4 
BENOMAR S., BELHSEN O.K., MATHIEU M. AND MOUKRIM A. Ultrastructural study of oogenesis 
in the African mussel, Perna perna (Bivalvia: Mytilidae) 
Estudio ultraestructural de la ovogénesis en el mejillón africano, Perna perna (Bivalvia: 
ANT AO o Hp ON E os EEE UE A ed RARA EN ONIS 5-21 
GARCÍA-ÁLVAREZ O., ZAMARRO M2? AND URGORRI V. New species of Mollusca Solenogastres 
from the Bellingshausen Sea and the Antarctic Peninsula (Bentart-2006 Expedition) 
Nuevas especies de Moluscos Solenogastres del Mar de Bellingshausen y Península Antártica 


(PAE O cto 2 DUO) LN e o ALI Ae E ae O. 23-38 
DE OLIVEIRA A. Fauna Malacológica da cidade de Coimbra (Beira Litoral). Moluscos “urbanos” de 
Portugal. 1 


Malacological Fauna from Coimbra (Beira Litoral). Portuguese “urban” Molluscs. 1... 39-50 
BOUCHET P., GOFAS S. AND WARÉN A. Notes on Mediterranean Dizoniopsis (Gastropoda: Cerit- 

hiopsidae), with the description of two new species 

Apuntes sobre los Dizoniopsis (Gastropoda: Cerithiopsidae) del Mediterráneo, con la descrip- 

INE NOS ES DECIS MUBUIS Aa DAR aa LAO SON ed A 51-62 
COELHO R. AND CALADO G. Spawn and early development of NE Atlantic species of Hypselodoris 

(Gastropoda: Opisthobranchia) 

Puesta y desarrollo de especies del género Hypselodoris del Atlántico nororiental (Gastropoda: 

TEO EN aca Ue AN a ADO laa ala CON 63-72 


ISSN 0212-3010 


ll 


' 


DL. 
4D! 
1 


“Iberus 


Vol. 29 (1) 


REVISTA DE LA 
SOCIEDAD ESPAÑOLA 
DE MALACOLOGÍA 


Oviedo, junio 2011 


Iberus 
Revista de la 
SOCIEDAD ESPAÑOLA DE MALACOLOGÍA 


Comité DE REDACCIÓN (BOARD OF EDITORS) 
EDITOR DE PUBLICACIONES (EDITOR-IN=-CHIEF) 
Serge Gofas | Universidad de Málaga, España 


DIRECTOR DE REDACCIÓN (EXECUTIVE EDITOR) 
Gonzalo Rodríguez Casero Mieres del Camino, Asturias, España 


EDITORA EJECUTIVA (MANAGING EDITOR) 
Eugenia M* Martínez Cueto-Felgueroso Mieres del Camino, Asturias, España 


EDITORES ADJUNTOS (ASSOCIATE EDITORS) 


Francisco Javier Conde de Saro Embajada de España, Japón 

Benjamín Gómez Moliner Universidad del País Vasco, Vitoria, España 

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

Emilio Rolán Mosquera Universidad de Vigo, Vigo, España 

José Templado González Museo Nacional de Ciencias Naturales, CSIC, Madrid, España 
Jesús S. Troncoso Universidad de Vigo, Vigo, España 


Comité EDITORIAL (BOARD OF REVIEWERS) 


Kepa Altonaga Sustacha Universidad del País Vasco, Bilbao, España 

Eduardo Angulo Pinedo Universidad del Poís Vasco, Bilbao, España 

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

Thierry Bockeljau Institut Royal des Sciences Naturelles de Belgique, Bruselas, Bélgica 
Ridiger Bieler The Field Museum, Chicago, Estados Unidos 

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

Jose Castillejo Murillo Universidad de Santiago de Compostela, Santiago de Compostela, España 
Karl Edlinger Noturhistorisches Museum Wien, Viena, Austria 

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

José Carlos García Gómez Universidad de Sevilla, Sevilla, España 

Gonzalo Giribet de Sebastián Harvard University, EE.UU. 

Edmund Gittenberger Notional Notuurhistorisch Museum, Leiden, Holanda 

Ángel Guerra Sierro Instituto de Investigaciones Marinas, CSIC, Vigo, España 

Gerhard Hoszprunar -Zoologische Staatssammlung Múnchen, Múnchen, Alemania 

Yuri |. Kantor AN. Severtzov Institute of Ecology and Evolution, Moscú, Rusia 
María Yolanda Manga González Estación Agrícola Experimental, CSIC, León, España 

Jordi Martinell Callico Universidad de Barcelona, Barcelona, España 

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

Tokashi Okutani Nihon University, Fujisawa City, Japón 

Marco Oliverio Universitá di Roma “La Sapienza”, Roma, !talia 

Pablo E. Penchaszadeh Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina 
Winston F. Ponder Australian Museum, Sydney, Australia 

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

Me de los Ángeles Romos Sánchez Museo Nacional de Ciencias Naturales, ESIC, Madrid, España 
Francisco Javier Rocha Valdés Instituto de Investigaciones Marinas, CSIC, Vigo, España 

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

Joandoménec Ros ¡ Aragones Universidad de Barcelona, Barcelona, España 

María Carmen Solas Casanovas Universidad de Málaga, Málaga, España 

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

Victoriano Urgorri Carrasco Universidad de Santiago de Compostela, Santiago de Compostela, España 
Anders Warén Swedish Museum of Natural History, Estocolmo, Suecia 


PORTADA DE Jlberus 
Iberus gualtieranus (Linnaeus, 1758), una especie emblemática de la península Ibérica, que da 
nombre a la revista. Dibujo realizado por José Luis González Rebollar “Toza”. 


NT HSON/g py 


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REVISTA DE LA 
SOCIEDAD ESPAÑOLA 
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Vol. 29 (1) Oviedo, junio 201 1 


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. 


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

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


Dep. Leg. B-43072-81 
ISSN 0212-3010 
Diseño y maquetación: Gonzalo Rodríguez 


Impresión: LOREDO, S. L. - Gijón 


O Sociedad Española de Malacología ——__—_—_———  Iberus, 29 (1): 1-7, 2011 


Cima apicisbelli Rolán, 2003 (Gastropoda: Cimidae): nuovo 


immigrante in Mediterraneo o specie autoctona? 


Cima apicisbelli Rolán, 2003 (Gastropoda: Cimidae): ¿nuevo inmi- 
grante en el Mediterraneo o especie autoctona? 


Danilo SCUDERI* e Francesco CRISCIONE** 


Recibido el 11-11-2010. Aceptado el 21-1-2011 


ABSTRACT 


The recovery of several living specimens of Cima apicisbelli Rolán, 2003 ¡is reported in 
the coast of Acitrezza (lonian Sea, E Sicily) within a marine protected area. This is the first 
citation in Italian waters, and the second ever for the Mediterranean, but of primary impor- 
tance as based on abundant material found alive. This finding opened a taxonomic issue 
regarding its actual validity as a distinct species. The resemblance of their shells with that 
of the autochthonous C. cylindrica [Jeffreys, 1856), re-proposes an old question regarding 
the alien species entering the Mediterranean via Gibraltar strait: are they really aliens or 
are they ¡just rare and/or overlooked native species to be better understood taxonomi- 
cally? Based on observations on fresh material of C. apicisbelli and comparison with 
shells of the native congener we here support the validity of both taxa, therefore consider- 
ing our record a real case of new finding of a species of Atlantic origin. 


RIASSUNTO 


Si segnala il ritrovamento di 26 esemplari viventi della specie alloctona Cima apicisbelli 
Rolán, 2003 nelle acque di Acitrezza (Mar lonio, Sicilia Orientale), all'interno di un'area 
marina protetta: si tratta del primo rinvenimento per le acque italiane, e del secondo in 
assoluto per il Mediterraneo, ma di primaria importanza poiché quello presente si basa su 
abbondante materiale rinvenuto vivente. Insieme ai dettagli del ritrovamento, viene di 
seguito riportata una discussione circa la reale consistenza tassonomica di questa specie: 
la congenere autoctona C. cylindrica [Jeffreys, 1856) risulta avere una conchiglia morfolo- 
gicamente molto simile. Viene esplorata la possibilitá che le due specie possano essere 
solo sinonimi: dal confronto tra individui di C. cylindrica, rispondenti all'attuale concetto 
della specie, e C. apicisbelli, rinvenute ad Acitrezza, sono emerse differenze a supporto 
della distinzione delle due specie che conducono a riferire ¡l presente rinvenimento come 
una nuovo reperimento in Mediterraneo di una specie aliena di provenienza atlantica. 


RESUMEN 


Se señala la recolección de 26 ejemplares vivos de la especie alóctona Cima apicisbelli 
Rolán, 2003 en aguas de Acitrezza (mar Jónico, este de Sicilia), dentro de un área 
marina protegida. Este es | primera cita de la especie para aguas italianas , y la segunda 
para el Mediterráneo, pero de suma importancia por tratarse de un abundante material 


* Dipartimento di Biologia Animale, Laboratorio di Biologia Marina, Universitá di Catania, Via Androne, 81 - 
95124 Catania, Italy. E-mail: danscuCtin.it 

** Australian Museum, 6 College Street Sydney NSW - 2010 Sydney, Australia. E-mail: cicciocriscio- 
neOgmail.com 


Iberus, 29 (1), 2011 


recolectado vivo. En esta ocasión comparamos esta especie con el congénere mediterrá- 
neo C. cylindrica (Jeffreys, 1856), que tiene una concha muy similar morfológicamente y 
podría ser sinónimo. Comparando ejemplares de C. cylindrica correspondiendo al con- 
cepto actual de la especie, y los C. apicisbelli de Acitrezza, las diferencias respaldan la 
distinción de las dos especies, lo cuál nos lleva a considerar este descubrimiento como 
una nueva ocurrencia en el Mediterráneo de una especie exótica de origen atlántico. 


INTRODUZIONE 


Durante questi ultimi dieci anni, 
l'ingresso in Mediterraneo di specie di 
molluschi non autoctone e apparso 
essere un fenomeno particolarmente fre- 
quente ed in crescita. Ció sia perché tale 
fenomeno viene piú spesso rimarcato ed 
enfatizzato in quanto ricollegabile al 
fenomeno piú generale del cambia- 
mento del clima su scala globale e della 
tropicalizzazione piú in particolare, sia 
anche perché la bibliografia e gli studi 
prodotti relativamente alla malacofauna 
mediterranea sono ormai estremamente 
approfonditi, cosicché la individuazione 
di nuove entita all'interno di questo 
bacino risulta maggiormente facilitata. 

Secondo GALIL (2008), la consistenza 
del fenomeno di invasione del Mediter- 
raneo puo essere spiegata come un 
effetto della sinergia tra fattori antropici 
(inquinamento, eutrofizzazione, de- 
grado degli habitats, sovrapesca, mari- 
coltura, cambiamento climatico e traf- 
fico marittimo) e la peculiare storia geo- 
fisica e climatica dell'area. Una delle 
modalita con cui le specie aliene si 
diffondono sembra essere la normale 
dispersione larvale e ció e particolar- 
mente vero per le specie cosiddette “les- 
sepsiane”, che transitano attraverso il 
canale di Suez per poi attestarsi lungo le 
coste prossime ad esso, quelle levantine 
o egiziane (OLIVERIO, 1995). In questo 
caso, la diffusione segue una succes- 
sione temporale e direzionale (“step- 
ping stones”), che si rispecchia nelle 
segnalazioni scientifiche. Ovviamente, 
nulla esclude che meccanismi di media- 
zione antropica possano essere in alcuni 
casi prime responsabili del trasporto di 
queste specie attraverso il canale o di 
una loro secondaria diffusione in Medi- 
terraneo. Rimorchiatori o chiatte ormeg- 


glate per lungo tempo ad una estremita 
per poi passare all'altro lato del canale 
possono aver costituito un ottimo sub- 
strato per specie aliene (Fox, 1926). Navi 
che fanno la spola tra il bacino di 
Levante e di Ponente possono essere un 
veicolo di diffusione per specie sessili, 
come nel caso di Brachidontes pharaonis 
(Fischer, 1870) (DI GERONIMO, 1971) e 
Pinctada radiata (Leach, 1814) (in 
ZIBROWIUS, 1992). Per quest'ultima, anzi, 
il veicolo di trasporto puo dipendere 
anche da substrati naturali, come ad 
esempio il carapace di rettili marini 
(OLIVERIO, GEROSA E Cocco, 1993). Il 
trasporto mediato da mezzi di naviga- 
zione marittima e probabilmente la 
maggiore causa di importazione di 
specie aliene sensu stricto, cioe specie 
introdottesi in Mediterraneo diretta- 
mente dal luogo d'origine (OLIVERIO, 
1995). Questo e particolarmente vero per 
quei mezzi con ridotta velocita o per 
quelle strutture che operano in condi- 
zioni di frequente ancoraggio, come le 
piattaforme di esplorazione dei fondali. 
Dodici specie di Molluschi originari del- 
l'Australia vennero per la prima volta 
segnalate nel nostro mare in prossimitá 
della piattaforma “Southern Cross” anco- 
rata presso le coste israeliane (MIENIS, 
2004). L'altra importante modalita con la 
quale si realizza 1'”invasione aliena” e 
certamente legata alla maricoltura, atti- 
vita che e cresciuta esponenzialmente 
nelle ultime due decadi. Da non trascu- 
rare e, inoltre, l'introduzione che si rea- 
lizza tramite le cosiddette ballast water, o 
acque di zavorra, riversate dalle navi da 
carico nelle localita di destinazione 
assieme al loro contenuto in larve 
(STREFTARIS, ZENETOS E PAPATHANAS- 
SsIOU, 2005). L'introduzione intenzionale 


SCUDERI E CRISCIONE: Czia apicisbellí, immigrante in Mediterraneo o specie autoctona? 


dell'ostrica del Pacifico, Crassostrea gigas 
(Thunberg, 1793), e della vongola filip- 
pina Ruditapes philippinarum (Adams e 
Reeve, 1850), oltre ad aver messo in 
pericolo le popolazioni delle specie 
autoctone  corrispondenti  (BODOY, 
MAITRE-ALLAIN E RIVA, 1981; MADHIOUB 
E ZAOUALI, 1988), fu seguita da quella 
delle specie aliene ad esse associate 
(RIBERA E BOUDOURESQUE, 1995). 

Se da una parte la fauna proveniente 
dall'area indopacifica risulta piuttosto 
semplice da distinguere da quella autoc- 
tona, quella, invece, di provenienza 
atlantica offre maggiori problemi per 
levidente somiglianza morfologica 
delle sue specie con quelle del Mediter- 
raneo. Inoltre, poiché la malacofauna 
del Mediterraneo risulta composta pre- 
valentemente da specie provenienti dal- 
l'area lusitanica, risulta spesso difficile 
stabilire se una specie sia entrata nel 
nostro mare solo recentemente oppure 
sia stata sempre presente e male inter- 
pretata o non ben identificata o ancora 
geograficamente distribuita con popola- 
zioni circoscritte nello spazio o nel 
tempo. Un'ulteriore complicazione 
deriva, infatti, da quelle specie lusitani- 
che che, tuttora presenti in Mediterra- 
neo, potrebbero averlo colonizzato a piú 
riprese nel tempo, come ad esempio 
Panopea glycymeris (Born, 1778), per le 
quali un'indagine approfondita deve 
partire dallo studio delle conchiglie 
fossili. Molte specie atlantiche, per 
questo, sono state escluse dalla lista 
delle specie alloctone della CIESM 
(ZENETOS, GOFAS, RUSSO E TEMPLADO, 
2004). 

In questi ultimi anni in certe aree 
della Sicilia orientale si sono susseguiti 
numerosi rinvenimenti di specie aliene: 
in particolare ad Acitrezza, piccolo vil- 
laggio poco a nord di Catania, che vive 
di pesca e della commercializzazione 
del pesce, ospitando uno dei maggiori 
mercati ittici siciliani, il fenomeno della 
presenza di specie non autoctone si e 
recentemente amplificato, portando a 
peculiari segnalazioni (SCUDERI E RUSSO, 
2003; 2005) e giustificando la realizza- 
zione di specifici lavori di ricerca, solo 
in parte conclusi (PANE, 2008). 


Il recente rinvenimento nella stessa 
localita di numerosi individui viventi 
determinati come Cima apicisbelli Rolán, 
2003 (Cimidae), ha offerto la possibilitá 
di investigare meglio la tassonomia di 
questa specie ed i rapporti tra malaco- 
fauna atlantica e quella mediterranea. 
Questa specie, descritta per le coste del 
Senegal, e stata rinvenuta per la prima 
volta in Mediterraneo nel 2007 lungo le 
coste di Denia, Valencia, Spagna meri- 
dionale, dove un'unica conchiglia vuota 
e stata ritrovata in un detrito raccolto a 
bassa profondita, in un fondale misto di 
sabbia e roccia, con presenza della fane- 
rogama Posidonia oceanica (L.) Delile, 
1813 e di alghe fotofile (OLIVER 
BALDOVI, 2007). 

ll rinvenimento qui descritto costi- 
tuisce il primo per le acque italiane e il 
secondo in assoluto per il Mediterraneo. 
Non e” da trascurare anche il fatto che 
sia avvenuto all'interno dei confini di 
un'area protetta. 

La disponibilita” di abbondante 
materiale vivente ha consentito un 
esame approfondito, dal quale e emersa 
una grande somiglianza di C. apicisbelli 
con la congenerica autoctona C. cylin- 
drica (Jeffreys, 1856). Tale rimarchevole 
somiglianza non era stata discussa 
all'atto della istituzione di C. apicisbell:. 

Il presente lavoro affronta una breve 
disamina tassonomica introduttiva sulle 
due specie di Cima per arrivare ad una 
soluzione logica circa il ritenere, e 
quindi segnalare, la specie di recente 
descrizione come alloctona. 


MATERIALI E METODI 


C. apicisbelli. Acitrezza (Catania, NE 
Sicilia), Marzo 2006, profonditá -0,20/1 
m su un substrato costituito in preva- 
lenza da Caulerpa racemosa, 21 individui. 
Altri 5 individui a nord di Acitrezza, 
estate 2006, profondita -1/2 m, fotofilo 
roccioso. 

C. cylindrica. Ganzirri (Messina, NE 
Sicilia), 1995, profonditá -0,20/1 m su 
un substrato costituito in prevalenza da 
Caulerpa taxifolia, 3 individui. Is Linosa 
(Agrigento, Canale di Sicilia), 1995, 


Iberus, 29 (1), 2011 


profondita -25 m in detrito, 2 conchiglie. 
Vendicari (Siracusa, SE Sicilia), 1994, 
detrito spiaggiato, 2 conchiglie. 


DISCUSSIONE 


C. apicisbelli e caratterizzata da una 
conchiglia minuta, fragile, con scultura 
delicata (Figs. 1, 2). La caratteristica che 
immediatamente la fa distinguere dalle 
altre congeneri mediterranee, C. minima, 
C. cylindrica, e la scultura della conchi- 
glia, che non e liscia, ma presenta sottili 
costicine assiali intersecate a linee spirali 
a formare un reticolo; tale trama puo 
ricordare la scultura delle varie specie di 
Graphis presenti in Mediterraneo, dalle 
quali si distingue agevolmente per l'apice 
non liscio. Come gia evidenziato in lette- 
ratura (vAN AARTSEN, 1981), la conchigjlia 
di C. cylindrica risulta liscia, solcata da 
deboli strie spirali solo negli ultimi giri: 
anche se non riportato esplicitamente, 
Vapice risulta pressoché liscio. 

Sono stati posti a confronto gli indi- 
vidui di C. apicisbelli rinvenuti nell'area 
jonica sopra indicata e alcuni individui 
di C. cylindrica conservati nella colle- 
zione di uno degli autori (D.S.), che 
confrontano perfettamente con la 
descrizione di VAN AARTSEN (1981): 
sono stati confrontati tra loro individui 
di pari dimensioni. Dal confronto degli 
individui delle due specie, riportati in 
materiali e metodi e di cui si da 
un'immagine in Figs. 1 e 4, si eviden- 
ziano differenze morfologiche, sia a 
carico della teleoconca che della proto- 
conca. Infatti la conchiglia adulta di C. 
apicisbelli, a paritá di dimensioni (circa 1 


mm), rispetto a C. cylindrica, appare piú 
inflata, con meno giri e piu arrotondati 
con sutura piu netta; la scultura appare 
molto marcata su tutta l'altezza di ogni 
giro, compresi quelli apicali; inoltre la 
bocca appare piú aperta ed ampia (si 
veda il disegno schematico di Fig. 9). 
Circa la protoconca, invece, in C. apici- 
sbelli risulta ornata da scultura consi- 
stente di linee assiali rade e spirali piú 
marcate, che sul nucleo, piuttosto 
grosso, divengono ortogonali a causa 
dell'iperstrofia protoconcale (Figs. 3 e 
7). In C. cylindrica, invece, la protoconca 
appare liscia (Figs. 5 e 6), con solo 
qualche linea d'accrescimento appena 
rintracciabile: il nucleo e piú piccolo 
(0,057 mm contro 0,071 mm di C. apici- 
sbelli). Sul nucleo embrionale di 
entrambe e visibile un cingolo apicale 
(Figs. 6 e 7) che va poi a fondersi con la 
sutura gia del primo giro di protoconca. 

Le differenze morfologiche rilevate, 
quindi, se da un lato potrebbero essere 
viste come lievi e rientrare nell'ambito 
della variabilita intraspecifica di C. 
cylindrica, che del resto risulta tuttora 
specie scarsamente rinvenuta ed affatto 
investigata tassonomicamente, come gia 
denunciato da VAN AARTSEN (1981), dal- 
l'altro sono sufficienti a discriminare le 
due entitá tassonomiche: esistono, 
infatti, gruppi di specie (vedi nel genere 
Turbonilla o Chrysallida ad esempio) che 
sono divergenti in maniera piu lieve e 
differiscono per un piú esiguo numero 
di caratteri. 

Per tale motivo C. apicisbelli e C. 
cylindrica vengono qui considerate 
specie diverse, sulla base di tali diffe- 
renze morfologiche. 


(Pagina destra) Figuras 1-3, 7, 9. Cima apicisbelli, Acitrezza (Catania, Sicilia); 1, 2: interi individui 
(0,95 mm e 0,70 mm); 3: protoconca; 7: disegno della protoconca; 9: disegno schematico del 
profilo conchiliare. Figuras 4, 5, 6, 8. Cima cylindrica, Ganzirri (Messina, Sicily); 4: intero indivi- 
duo (1,05 mm); 5: protoconca; 6: disegno della protoconca; 8: disegno schematico del profilo con- 


chiliare. 


(Right page) Figuras 1-3, 7, 9. Cima apicisbelli, Acitrezza (Catania, Sicilia); 1, 2: individuos enteros 
(0,95 mm y 0,70 mm); 3: protoconcha; 7: dibujo de la protoconcha; 9: dibujo esquemático del perfil 
de la concha. Figuras 4, 5, 6, 8. Cima cylindrica, Ganzirri (Messina, Sicily); 4: individuo entero 
(1,05 mm); 5: protoconcha; 6: dibujo de la protoconcha; 8: dibujo esquemático del perfil de la 


concha. 


SCUDERI E CRISCIONE: C2xa apicisbelle, immigrante in Mediterraneo o specie autoctona? 


Iberus, 29 (1), 2011 


Il dubbio che si possa trattare di 
forme estreme della stessa specie, 
comunque, permane, non prefiggendosi 
il presente lavoro la loro investigazione 
puramente tassonomica, ed apre una 
finestra sulle polemiche che descrizioni 
troppo affrettate, senza operare le 
dovute comparazioni con materiale 
tipico di specie molto vicine portano a 
rilevare: un monito, quindi, va necessa- 
riamente profuso affinche non si prose- 
gua con superficialita alla descrizione di 
nuovi taxa che, una volta istituiti, e 
lavoro duro e improbo, poi, andare a 
confrontare e revisionare. 

Essendo quindi maggiormente pro- 
pensi alla validita di C. apicisbelli, pos- 
siamo qui dire che quello presente 
risulta essere il primo rinvenimento per 
le acque italiane e per il Mediterraneo 
orientale, mentre risulta essere il 
secondo in ordine cronologico per 
l'intero Mediterraneo dopo quello di 
Valencia (OLIVER BALDOV1, 2007). 
Quest'ultimo, pero, e stato fondato su 
un unico individuo rinvenuto morto, da 
cui l'importanza assunta dalla presente 
segnalazione di numerosi individui 
viventi che accertano con sicurezza la 
presenza in Mediterraneo di popola- 
zioni di questa specie. 

Rimangono ancora da stabilire le 
modalita di  ingresso nel Mare 
Nostrum. 

Rispetto a quello di Valencia, infatti, 
il rinvenimento siciliano non sembra 
rappresentare una tappa di colonizza- 
zione naturale del nostro mare, sia per i 
tempi con cui si e attuata (contempora- 
nea rispetto al rinvenimento spagnolo), 
sia anche per il numero di individui, 
svariati e tutti viventi. Va inoltre osser- 
vato che, come giá precedentemente 
notato per altre specie alloctone, rinve- 
nute nelle  medesime  condizioni 
(Scuderi é£ Russo, 2003; 2005), risulta 
piuttosto anomalo rinvenire in tempi 
brevi ed al centro del Mediterraneo 
popolazioni gia ben acclimatate altrove 
assenti: ció risulta contrario alle dinami- 
che di invasione naturale conosciute 
(OLIVERIO, 1995) e sembra piuttosto con- 
forme ad un modello d'invasione 
mediato dall'uomo. 


Inoltre, 1 diversi precedenti rinveni- 
menti di specie aliene (SCUDERI E 
Russo, 2003; 2005) nella stessa area, 
nonché la presenza a ridosso di 
quest'ultima di uno dei piú fiorenti 
mercati ittici siciliani, ha suggerito 
Videa che tali specie possano arrivare 
all'interno delle cassette di pesce, non a 
caso proveniente spesso dalle aree geo- 
grafiche di origine delle stesse. In parti- 
colare la presente specie e stata des- 
critta solo molto recentemente, per il 
Senegal: la richiesta di pescato “non 
autoctono”,  particolarmente del 
Senegal, e cresciuta enormemente 
durante questi ultimi anni (osserv. 
pers.). Questo perviene ai nostri mercati 
sempre meno in condizioni di congela- 
mento e sempre piú in condizioni di 
ottima freschezza, grazie anche ai 
moderni ed efficienti canali di approv- 
vigionamento (spedizioni aeree). Per 
questo motivo, esiste a nostro parere la 
possibilitá che individui di specie aliene 
arrivino in buono stato di conserva- 
zione e riescano poi a sopravvivere e 
adattarsi, una volta accidentalmente re- 
immesse in mare. Questo meccanismo 
potrebbe spiegare anche altri fenomeni 
di colonizzazione da parte delle specie 
alloctone avvenuto in altre aree del 
Mediterraneo. Ulteriori studi di compa- 
razione tra il materiale biologico che 
accompagna il pescato importato e 
quello rinvenuto nelle indagini sul 
campo, potrebbero avvalorare tale 
ipotesi. 


RINGRAZIAMENTI 


Gli autori desiderano qui ringraziare 
la prof.ssa Grazia Cantone del Dip. di 
Biologia Animale dell'Universita di 
Catania per l'appoggio e gli utili pareri 
al riguardo. I ringraziamenti vanno 
anche al Direttore (Dott. Emanuele 
Mollica) e allo staff delll AMP “Isole 
Ciclopi” per avere consentito e facilitato 
il campionamento. Un ringraziamento 
particolare va al prof. S. Gofas (UMA) 
per le utili critiche al manoscritto e le 
sempre opportune, puntuali e precise 
disquisizioni scaturite. 


SCUDERI E CRISCIONE: Czma apicisbells, imimigrante in Mediterraneo o specie autoctona? 


BIBLIOGRAFIA 


BODOY A., MAITRE-ALLAIN T. E RIVA A. 1981. 
Croissance comparée de la palourde euro- 
péenne Ruditapes decussatus et de la palourde 
japonaise Ruditapes philippinarum dans un 
écosysteme artificiel méditerranéen. Vie Ma- 
rine, 2: 39-51. 

DI GERONIMO 1. 1971. Prima segnalazione su- 
lle coste italiane di Brachiodontes variabilis 
Krauss. Bollettino delle Sedute dell" Accademia 
Gioenia di Scienze Naturali in Catania, 10: 
847-852. 

Fox H.M. 1926. Zoological results of the Cam- 
bridge expedition to the Suez Canal, 1924. 1. 
General part. Transactions of the Zoological So- 
ciety of London, 22: 1-64. 

GaLIL B.S. 2008. Alien species in the Mediter- 
ranean Sea - which, when, where, why? Hy- 
drobiologia, 606: 105-116. 

MADHIOUB M.N. E ZAOUALIJ. 1988. Captage de 
l'huítre Crassostrea gigas dans le lac Ichkeul. 
Bulletin de l'Institut National Scientifique et 
Technique d'Océanographie et de Péche de Sa- 
lammbó, 15: 47-60. 

MIENIS H.K. 2004. New data concerning the 
presence of Lessepsian and other Indo-Pacific 
migrants among the molluscs in the Mediter- 
ranean Sea with emphasize on the situation 
in Israel. In B. Oztúirk €: A. Salman (Eds.): Pro- 
ceedings 1st National Malacology Congress, 
1-3 September 2004, Izmir. Turkish Journal of 
Aquatic Life, 2 (2): 117-131. 

OLIVER BALDOVIJ.D. 2007. Catalogo de los Gas- 
terópodos testaceos marinos de la parte Sur 
del Golfo de Valencia (España). Iberus, 25 (2): 
29-61. 

OLIVERIO M. 1995. The status of the living Me- 
diterranean Strombus, or: what is a lessepsian 
migrant. Notiziario CISMa, 16[1994]: 35-40. 

OLIVERIO M., GEROSA G. E COCCO M. 1993. First 
record of Pinctada radiata (Bivalvia, Pteriidae) 
epibiont on the loggerhead sea turtle Caretta 
caretta (Chelonia, Cheloniidae). Bollettino 
Malacologico, 28 (5-12): 149-152. 


PANE F. 2008. Malacofauna associata ad alghe fo- 
tofile in un sito dell AMP “Is. Ciclopi” Acitrezza, 
con particolare riferimento alle specie aliene. Tesi 
di laurea, Universita di Catania, 54 pp. 

RIBERA M.A. E BOUDOURESQUE C.F. 1995. In- 
troduced marine plants with special reference 
to macroalgae: mechanisms and impacts. 
Progress in Phycological Research, 11: 187-268. 

ROLÁN E. 2003. A new species of the genus 
Cima (Gastropoda, Cimidae) from Senegal. 
Novapex, 4 (1): 21-23. 

SCUDERI D. E RUSSO G.F. 2003. Due nuovi Gas- 
teropodi per le acque italiane: Melibe fim- 
briata Alder €£ Hancock, 1864 e Tricolia tin- 
gitana Gofas, 1982 (Mollusca: Gastropoda). 
Atti XXXV Congr. SIBM, Biologia Marina Me- 
diterranea, 12 (1): 618-621. 

SCUDERI D. E RUSSO G.F. 2005. Prima segnala- 
zione di Aplysia dactylomela Rang, 1828 e pro- 
babile presenza di Syphonota geographica 
(Adams e Reeve, 1850) (Gastropoda: Opist- 
hobranchia: Anaspidea) per le acque del Me- 
diterraneo. Atti XXXV Congr. SIBM, Biologia 
Marina Mediterranea, 12 (1): 338-341. 

STREFTARIS N., ZENETOS A. E PAPATHANASSIOU 
E. 2005. Globalisation in marine ecosystems: 
the story of non-indigenous marine species 
across european seas. Oceanography and Ma- 
rine Biology: An Annual Review. 43, 419-453. 

VAN AARTSEN J.J. 1981. European marine mol- 
lusca: notes on less well-known species II. The 
genus Cima Chaster, 1896. Basteria, 45: 117- 
TO; 

ZENETOS A., GOFAS S., RUSSO G. E TEMPLADO J. 
2004. CIESM Atlas of Exotic Species in the 
Mediterranean, Vol.3. Molluscs, 376 p. 
(http: / /www.ciesm.org / atlas / molluscsintro 
html). 

ZIBROWIUS H. 1992. Ongoing modifications of 
the Mediterranean marine fauna and flora by 
the establishment of exotic species. Mesogee, 
51: 83-107. 


A 
TES 
2 


A 


En 


8 


O Sociedad Española de Malacología 


Iberus, 29 (1): 9-33, 2011 


The family Tornidae (Gastropoda, Rissooidea) in the East 


Atlantic, 2. Circulinae 


La familia Tornidae (Gastropoda, Rissooidea) en el Atlántico orien- 


tal, 2. Circulinae 


Joan Daniel OLIVER* and Emilio ROLÁN** 


Recibido el 23-1-2011. Aceptado el 17-11-2011 


ABSTRACT 


The species of the subfamily Circulininae from the East Atlantic belonging to the genus Cir- 
culus are studied. There is a total of 8 species, of which 2 are previously undescribed. The 
shell morphology is ilustrated for all the species with scanning electron micrographs which 
show the protoconch and, in some cases, the microsculpture. 


RESUMEN 


Se estudian las especies del género Circulus de la subfamilia Circulininae del Atlántico 
occidental. En total son 8 especies, de las cuales 2 son nuevas para la ciencia. De todas 
ellas se ilustra la morfología de la concha con microscopía electrónica de barrido, inclu- 
yendo la protoconcha y, en algunos casos, la microescultura. 


INTRODUCTION 


In their 1969 paper, ADAM AND 
KNUDSEN (1969) studied the small 
discoid species then scarcely known 
from the West African coast. Subse- 
quently, new species were described 
(ROLÁN AND RUBIO, 1991, 1996; RUBIO 
AND ROLÁN, 1991; ROLÁN, RUBIO AND 
RYALL, 2000; ROLÁN AND RYALL, 2002). 
ROLÁN AND RUBIO (2002) published a 
paper on the species of the family 
Tornidae Sacco 1986, from West Africa 
reviewing and illustrating 39 species of 
which 23 were new to science. 

More recently, BOUCHET AND ROCROI 
(2005) recorded the Gastropod taxa of 
family level and listed as accepted the 
family Tornidae with the following sub- 
families: Torninae Sacco, 1884; Circuli- 
nae Fretter and Graham, 1962; Teinos- 


* c/Alcorisa 83, 12-C 28043, Madrid. 


tomatinae Cossmanmn, 1917; Vitrinellinae 
Bush, 1897. 

In the above mentioned work on the 
Tornidae from West Africa (ROLÁN AND 
Rubro, 2002), the family had been 
studied somewhat incompletely because 
some of these subfamilies were not 
included in Tornidae at that time but in 
other taxonomic groups. The genus Cir- 
culus Jeffreys, 1865, was treated by 
FISCHER (1887: 824) as a subgenus of 
Gibbula Risso, 1826; later DALL (1927) 
placed it in synonymy with Lydiphnis 
Melville, 1906 and THIELE (1929) as a 
genus within Cyclostrematidae. Most of 
them were placed in Archaeogas- 
tropoda, and this remained so until 
FRETTER (1956) researched the anatomy 
of Circulus striatus (Philippi, 1836), type 


** Museo de Historia Natural, Campus Universitario sur, 15782 Santiago de Compostela. 


Iberus, 29 (1), 2011 


species of Circulus, and noticed that the 
radula of this species was taenioglos- 
sate, not rhipidoglossate. 

For this reason we have resumed the 
study of the family Tornidae with the 
intention of reviewing the subfamilies 
not included in the previous study. In 
the present work we begin with subfam- 
ily Circulinae. 

In the present work, as well as in the 
previous ones, we intend to show a 
complete iconography of the species 
pointing out the differential morpholog- 
ical characters which allow a clear 
placement of each taxon. 


MATERIAL AND METHODS 


The material studied comes from 
several collections made by the authors, 
mainly in the Mediterranean, Morocco, 
Mauritania, Senegal, Sáo Tomé and 
Principe, Annobón and Angola. Other 
material collected by other malacolo- 
gists such as Francisco Fernandes, 
Anselmo Peñas, Peter Ryall and José 
María Hernández was also examined. 
There is also some material from the 
Muséum National d' Histoire Naturelle 
of Paris (MNHN). 

The material was collected mainly 
from sediments from the intertidal level 
to 8-10 m, while diving with snorkel 
and SCUBA diving, and also by dredg- 
ing from a boat. The material from the 
MNHN was collected in several expedi- 
tions to Guinea Conakry, Ivory Coast 


RESULTS 


and Congo with R/V “André Nizery” 
and “Antea Benchaci I”, and in Angola 
(coll. Serge Gofas) through manual 
dredging. 

The sediments were sieved and 
examined under a binocular microscope 
either by the collectors or the authors. 

The numbering of the protoconch 
whorls was made following the method 
of VERDUIN (1976) in which the whorls 
are counted following an initial nucleus. 


Abbreviations 


AMNH American Museum of Natural 
History, New York 

MNCN Museo Nacional de Ciencias 
Naturales, Madrid 

MNHN Muséum national d'Histoire 
naturelle, Paris 

MHNS Museo de Historia Natural, Uni- 
versidad, Santiago de Compostela 

NHMUK The Natural History Museum, 
London 

RBINS Royal Belgian Institute of 
Natural Sciences, Bruxelles 

ZMB Zoologisch Museum fir Natur- 
kunde, Berlin 

CAP collection of Anselmo Peñas, 
Vilanova i la Geltrú 

CDO collection of Daniel Oliver, Madrid 

CPR collection of Peter Ryall, Maria 
Rain 

CJH collection of José María Hernández, 
Gáldar 

sp specimen with soft parts 

s shell empty 

f fragment 


Family TORNIDAE 
Subfamily CIRCULINAE Fretter and Graham, 1962 


Genus Circulus Jeffreys, 1865 


Circulus Jeffreys, 1865. Brit. Conch., vol 3: 315 [established as a subgenus of Trochus; type species: 
Delphinula duminyi Requien, 1848 = Circulus striatus]. 


General characters of the shell: Proto- 
conch: multispiral without any sculpture. 
The transition to the teleoconch is diffi- 
cult to observe, because the separation is 


10 


scarcely apparent. The beginning of the 
teleoconch may be observed by looking 
at the beginning of the spiral cords, 
although sometimes these are eroded. 


OLIVER AND ROLÁN: The family Tornidae in the East Atlantic, 2. Circulinae 


Teleoconch: circular, flat, with a wide 
deep umbilicus. Spiral ornamentation 
formed by spiral cords. Some of them are 
prominent, forming keels. They lack any 
axial sculpture except for growth lines. 

In order to describe more easily the 
sculpture of the shell in view of its taxo- 
nomic importance, we can represent the 
apertural peristome as a hexagon (Fig. 
1) (even though its shape is frequently 
irregular, tending sometimes to be rec- 
tangular or circular): 

1: Adapical insertion of the external 
lip with the shell 

1-2: Subsutural cords 

2: Cord /upper keel 

2-3: Lateral upper cords 

3: Cord /peripheral keel 

3-4: Lateral lower cords 


Dichotomous key of Circulus: 


4: Cord /basal keel 

4-5: Basal cords 

5: periumbilical cord (change of 
direction) 

5-6: Umbilical cords 

6: Abapical insertion of the external 
lip in the shell 

6-1: Callus: the callus is a simple pro- 
longation of the inner part on the col- 
umella. 

It is important to mention that the 
border of the external lip touches the 
teleoconch a little above the peripheral 
keel (3) while the internal border does it 
a little above the basal keel (4). 

Some species may lack some of these 
cords-keels, or additional cord-keels 
may appear. Also some intraspecific 
variability can be observed. 


1. - Shell with smooth spire. There are only umbilical cords ........ C. senegalensis 


- Shell with spiral cords /or keels .... 


DES hellwithoutkeels.. o 
Snellmtkeels. 


3 - Shell with evident spiral and basal cords 


A a a as > C. congoensis 


- Shell with lateral and basal areas without spiral cords or cords only appreciable 


A o C. striatus 
4 - Peripheral keel clearly more developed than the basal keel ............... 5 
- Shell with the basal keel as developed or more than the peripheral keel ...... 6 


5 - Shell with vitreous aspect, without basal keel and without subsutural 


COMAS O UA o a 


O Ea O C. pseudopraecedens 


- Shell of white color with basal cord /keel less developed that the peripheral one 


and with subsutural clear cords .... 


A Rae: C. ryalli 


6 - Shell lacking lower lateral cords with a basal keel more developed than the peri- 


O e A A 
- Lower lateral cords present ........ 


E e E C. stephani 


7 - Peristome tending to be rectangular and having microsculpture under magnifi- 


CALOR e ESO de 


O O C. microsculpturatus 


- Peristome subcircular, lower lateral cords, tending to form subperipheral 


ES e o e Rd E IS EE 


A o E C. smithi 


Circulus striatus (Philippi, 1836) (Figs. 2A-G, 3A-G, 4A-G) 


Valvata striata Philippi, 1836, p. 147, pl. IX, fig. 3A-C. [Type locality: Cefalú near Catania, Sicily, 


Pleistocene] 


Adeorbis tricarinatus Wood, 1848. Ann. Mag. Nat. Hist. 9: 530. 
Delphinula duminyi Requien, 1848. Cat. Cog. Corse: 64 [Type locality: Ajaccio, Corsica]. . 


11 


Iberus, 29 (1), 2011 


Delphinula triangulata Rayneval and Ponzi, 1854. Cat. Monte Mario: 18 [Type locality: Monte 
Mario near Rome, Italy, Pleistocene] 

Skenea striatula Weinkauff, 1862. J. Conch. 10: 343 [incorrect subsequent spelling of striata Philippi, 
1836]. 

Circulus costulatus Locard, 1889. Bull. Soc. Mal. France, 6: 297. [Type locality: France: Brest, Morbi- 
han and Vendée]. 

Circulus carinulatus Locard, 1889. Bull. Soc. Mal. France, 6: 300. [Regions armoricaine and aqui- 
taine; Provence]. 

Circulus striatus bicarinatus Altimira, 1977: 25. [Type locality: Sant Pol de Mar, Barcelona]. 

Delphinula costata Danilo and Sandri, 1856. 


Type material: Probably in ZMB. 

Other material examined: Usual form: Spain: 1 s, Santander (MHNS); 2 s, Ria de Vigo, 10 m (MHNS); 
2s, Tarragona (CAP); 5 s, Denia (beached) (MHNS); 133 s, 11 f, Cullera (beached) (CDO); 16 s, Denia 
(beached) (CDO”; 18 s, Oliva (beached) (CDO)”; 6 s, Jávea (beached) (CDO”; 2 s, Ibiza (CDO). Morocco: 
3 s, Agadir (MHNS). Carinate form: Spain: 7 s, Cullera (beached) (CDO). Tricarinate form: Spain: 


3 s, Cullera (beached) (CDO). 


Description: Shell (Figs. 2A-E, 3A-E): 
the best description can be seen in FRETTER 
AND GRAHAM (1978). Shell flat with about 
4 whorls (including protoconch) reaching 
1.6 mm in diameter and 0.8 mm in height; 
shells with five whorls reaching 2.75 mm 
diameter and 1.25 mm height respectively 
(GRAHAM, 1988). 

The protoconch (Figs. 2F, 3F, 4F-G) is 
multispiral, has a little less than 2.25 
whorls, and is about 390 um in width. It 
is smooth without any ornamentation 
and its end is barely marked with a light 
line coincident with the beginning of the 
spiral cords of the teleoconch. 

The shell with typical morphology has 
spiral cords not forming keels. Apically 6- 
7 spiral cords can be observed whose 
width is approximately half that of the cor- 
responding interspaces. The subsutural 
adapical one is weak, a little deep and 
limited on both sides by granules. Below, 
there are three more similar cords; the 
lowest one situated on the upper keel. 
Below these, there are four more cords 
(upper laterals) the fourth being in the 
emplacement of the peripheral keel. 

Examined from the umbilical side, 
the shell has very weak low lateral 
cords, which in some shells give the 
impression that there are no cords, up to 
an isolated cord, situated in the position 
of the basal keel and which may some- 
times be double. Apparently, there are 
no basal cords between the basal keel 
and the umbilical cords, the latter being 
always present and evident under mag- 


12 


nification of the umbilicus. The lack of 
basal cords may be caused by erosion or 
they may decrease as the shell grows. 

There are four or five umbilical 
cords, separated by sulci and crossed by 
an axial irregular and granulose sculp- 
ture. The aperture is circular. 

The operculum is corneous and 
rounded. 

The animal of C. striatus has been fig- 
ured in FRETTER AND GRAHAM (1962: 284) 

Distribution: The species is known 
from Ireland and the British Isles 
(FRETTER AND GRAHAM, 1978; GRAHAM, 
1982, 1988), Atlantic Iberian Peninsula 
(ROLÁN, 1983); Portugal (NOBRE, 1940); 
Mediterranean (HIDALGO, 1917; PoPPE 
AND GOTO, 1991). The record from Sáo 
Tomé and Príncipe (FERNANDES AND 
ROLÁN, 1993), refers to another species 
with which it was confused at the time. 

Remarks: Adeorbis tricarinatus Wood, 
1848 was described as a species distinct 
from C. striatus. The description of this 
species was based on fossil shells and 
authors like JEFFREYS (1865) considered it, 
as well as C. supranitidus (Wood, 1848), a 
fossil variety of C. striatus. In spite of this 
opinion, even recently some authors, such 
as TERRENI (1981) consider it valid. 
AARTSEN, MENHORST AND GITTENBERGER 
(1984, fig. 57) indicate that this taxon as 
dubious, but admit that it could be valid 
and compare it to other similar fossil 
species. COPPINI, CUNEO, MARGELLI AND 
CAMPANI (2005, fig. 2c) present a similar 
shell and consider that itis a valid species 


OLIVER AND ROLÁN: The family Tornidae in the Fast Atlantic, 2. Circulinae 


umbilical 
cords 


peripheral keel 


subperipheral 
keel 


periumbilical cord 
"ECETIA CI 


e 


peripheral á 
keel l 
upper keel 


subsutural cords 


ELCIEINTTD 


y cords A 


ENTE A 
A y 


A 
kN 


basal cords 


ENTE 


upper 
- cords 


ve ] El 
keel 


subsutural upper 
cords keel 


protoconch-teleoconch 
scar 


second scar 


Figure 1. Schematic images showing the different sculpture and the terms employed for these char- 


acters. 


Figura 1. Imágenes esquemáticas mostrando los distintos tipos de escultura y la terminología empleada 


para estos caracteres. 


which is called Circulus tricarinatus. This 
is reflected in the CLEMAM web page 
where this name is also mentioned as valid. 

In Cullera (Valencia, East Spain), a 
great quantity of material of Circulus 


striatus has been collected. In this mater- 
ial, there were many shells which do not 
represent exactly the typical form of the 
species, but have some peculiar charac- 
ters instead: a form, which is carinate 


13 


Iberus, 29 (1) 


AS 


200 um 


Figure 2. Circulus striatus (Philippi, 1836). A-E: shells, 1.6, 1.6, 1.5, 1.2, 1.2 mm, Cullera, Valen- 
cia; E: protoconch; G: detail of the umbilicus. 
Figura 2. Circulus striatus (Philippi, 1836). A-E: conchas, 1,6; 1,6; 1,5; 1,2; 1,2 mm, Cullera, 
Valencia; F: protoconcha; G: detalle del ombligo. 


14 


OLIVER AND ROLÁN: The family Tornidae in the East Atlantic, 2. Circulinae 


ATA 
a ss 


o: 


Figure 3. Circulus striatus (Philippi, 1836). A-C: shells, 1.68, 1.7, 1.9 mm, Agadir, Morocco 
(MHNS); D, E: shells, 1.9, 2.1 mm, from Tarragona (CAP); F: protoconch, Agadir; G: detail of 
the microsculpture, Tarragona. 

Figura 3. Circulus striatus (Philippi, 1836). A-C: conchas, 1,68; 1,7; 1,9 mm, Agadir, Marruecos 
(MANS); D, E: conchas, 1,9; 2,1 mm, de Tarragona (CAP); F: protoconcha, Agadir; G: detalle de la 


microescultura, Tarragona. 


Iberus, 29 (1), 2011 


le o Mr 
y s e 


200 um 200 um 


Figure 4. Circulus striatus (Philippi, 1836). A, B: shells of carinate form 1.24, 1.4 mm (CDO); C- 
E: shells of tricarinate form, 1.9, 1.5, 1.7 mm, Cullera, Valencia (CDO); E G: protoconchs. 

Figura 4. Circulus striatus (Philippi, 1836). A, B: conchas de la forma con quilla, 1,24; 1,4 mm 
(CDO); C-E: conchas de la forma con tres quillas, 1,9; 1,5; 1,7 mm, Cullera, Valencia (CDO); E G: 


protoconchas. 


1ó 


OLIVER AND ROLÁN: The family Tornidae in the East Atlantic, 2. Circulinae 


(Fig. 4A-B), has a more developed 
peripheral cord / keel; another form, is 
tricarinate (Figs. 4C-E) presenting three 
cords more developed in the shape of a 
keel. These cords are placed in the 
upper position (2), peripheral position 


(3) and basal position (4). The presence 
of these different forms in only a small 
area suggests to us that they are simply 
extreme forms or variants of one 
species, Circulus striatus with variable 
development of the keels. 


Circulus smithi Bush, 1897 (Figs. 5A-E, 6A-E 7A-D) 


Cyclostrema tricarinata Smith, E. A., 1871. p. 737, p1.75, fig. 26. [Type locality: Whydah, Dahomey]. 
Circulus smithi Bush, 1897. New name for Cyclostrema tricarinatus Smith 1871, non Adeorbis tricari- 


natus Wood 1842. p. 126. 


Type material: Probably in BMNH. Not examined. 


Other material examined: Western Sahara: 4 s, 3 j, Dakhla, 50-60 m, (MHNS). Mauritania: 8 s, 
Nouakchott, 80-100 m (CJH). Ivory Coast: 1 j, off Grand Bassam, R/V “Antea Benchaci 1”, 5* 11,3 
N, 3 46' W, (MNHN ); 3 s, off Grand Bassam R/V “Antea Benchaci I” L, 30 m, 5 09,2 N, 39 47,1” W 
(MNAHN). Ghana: 30 s, 13 j, 8 £, Miamia, 38-40 m (MHNS); 8 j, 8 £, Miamia, 45-55 m (MHNS). Angola: 
1 s, Luanda, 50 m (MHNS); 22 s, 1 f, Luanda, 20-100 m (MHNS). 


Description: Shell (Figs. 5A-C, 6A-D, 
7A-D) flat reaching 1.8 mm in diameter, 
with 4.1 whorls (protoconch included) 
and height of about 0.86 mm. In lateral 
view, the first whorls barely extend 
beyond the level of the last whorl, 
although this character is somewhat 
variable. 

The protoconch (Figs. 5D, 6E-F) is 
smooth with 2 Y whorls, and about 560 
um in diameter. Well preserved proto- 
conchs may show very fine slightly 
sinusoidal growth lines. The teleoconch 
begins with a not very noticeable line, 
slightly sinusoidal, and with the onset 
of the spiral cords. In some shells the 
protoconch seems to have more whorls 
due to the fact that the first whorl of the 
teleoconch is practically smooth. In such 
shells, the transition to the protoconch is 
not clear, but can be seen as the subsu- 
tural cord continues parallel to the 
suture and the other cords appear pro- 
gressively below, sometimes after a new 
interruption as a scar, where they are 
more evident. 

As in other Circulus, the shell sculp- 
ture consists of spiral cords which are 
narrower than their interspaces (Fig. 
5E). Growth lines can be seen in these 
interspaces. The umbilicus is wide and 
may present 2-3 umbilical cords in the 
interior, but sometimes these may be 


transformed into 8-9 very fine and 
closely set cords. 

Distribution: From Western Sahara 
south to Angola. 

Remarks: This species presents some 
variability, but most of the shells have 
three cords which are more developed, 
forming keels: the upper one in position 
2 (upper keel), 3 (peripheral keel) and 4 
(basal keel) (see Fig. 1 for these posi- 
tions). The external lip is inserted on the 
previous whorls a little above the 
peripheral keel, while the internal lip 
begins at the level of the lower keel. 
Between the suture and the upper keel 
there are 4-5 cords, the first one of 
which is very close to the suture. 
Between the upper keel and the periph- 
eral one there are two cords; and 
between this latter and the lower one 
there are three or four. Occasionally, one 
of these cords can be more developed 
forming a keel above the basal one. In 
basal view, five basal cords can be seen. 

The separation with those C. striatus 
which present more prominent periph- 
eral keels is based mainly on the umbili- 
cal axial striation present between the 
umbilical cords, while C. smithi lacks 
this sculpture completely, only showing 
spiral cords. Furthermore, the cords of 
C. striatus have more volume than those 
of C. smithi. 


17 


Iberus, 29 (1), 2011 


Figure 5. Circulus smithi Bush, 1897. A-C: shells, 1.7, 1.89, 1.92 mm, Miamia, Ghana (MHNS); 


D: protoconch; E: details of the microsculpture. 


Figura 5. Circulus smithi Bush, 1897. A-C: conchas, 1,7; 1,89; 1,92 mm, Miamia, Ghana (MHNS); 
D: protoconcha; E: detalles de la microescultura. 


18 


OLIVER AND ROLÁN: The family Tornidae in the East Atlantic, 2. Circulinae 


500 um 


Figure 6. Circulus smithi Bush, 1897. A-D: shells, 2.9, 3.2, 2.3, 2.6 mm, Congo (MNHN); E, F: 
protoconchs. 
Figure 6. Circulus smithi Bush, 1897. A-D: conchas, 2,9; 3,2; 2,3; 2,6 mm, Congo (MNHN); E, F: 


protoconchas. 


Iberus, 29 (1), 2011 


Circulus congoensis (Thiele, 1925) (Figs. 8A-D, 9A-I) 


Vitrinella congoensis Thiele, 1925. Gast. Deut. Tiefsen Expedition: 147, pl.9, fig. 3A-C. [Type locality: 
Congo mouth]. 
Circulus striatus in ADAM AND KNUDSEN (1969): 10, fig. 5. 


Type material: Holotype (Figs. 8A-C) in ZMB. 

Other material studied: Western Sahara: 1 c, Dakhla (50-60 m) (MHNS); Senegal: 1 s, Casamance 
12” 20,7 N, 16” 53,1” W (MNHN). Ghana: 14 s, Miamia, 38-40 m (MHNS). Guinea Conaktry: 1 s, 1 j, 
W Ile Quito, R/V “André Nizery” Sedigui II, 10 00' N,15” 46' W, 28 m (MNHN); 3 s, 1 j, W Ile Quito, 
R/V “André Nizery” Sedigui IL, 10%00'N, 15” 58” W, 34 m (MNHN); 3 s, W Ile Quito, 10 00' N, 15% 
43 W, 26 m (MNHN); 1 s, Río Yomponi 10? 24” N, 14” 50 W, 22 m (MHNS); 3 s, Río Nuñez 10% 35,5' 
N, 15” 26" W 9 m (MN HN). Guinea Bissau: 1 s, Bissau 11 1045 N, 15” 44,5” W, 25 m (MHNS); Angola: 


3 s, Luanda, 20-100 m (MHNS). 


Description: Shell (Figs. 8A-C, 9A-D) 
with 4.3 whorls (protoconch included) 
reaching 1.7 mm in diameter and 0.9 
mm in height. 

The protoconch (Figs. 8D, 9F-G) is 
smooth and reaches nearly three whorls 
and about 660-710 um in diameter. The 
spiral cords begin after a small scar. 

The teleoconch is ornamented with 
spiral cords which are clearly narrower 
than the corresponding interspaces. All 
cords are similar in size; none of them is 
more developed and so they do not 
form any keel. Only the subsutural cord 
(Fig. 9H-I) is clearly narrower than the 
others just when it appears at the end of 
the protoconch, and is only visible 
under strong magnification. After this 
first whorl, this cord increases in size, its 
width subsequently approaching that of 
the other cords. 

The shell has a little more than 20 
spiral cords: 7-8 from the suture to the 
place where the basal keel (which does 
not exist) would be; about ten in the 
basal area and 5-6 more inside the 
umbilicus. The cords are a little more 
flat at the base than at the dorsal area. 

The larger shells, like the holotype, 
may have fewer cords or they may be 
less apparent. 

Distribution: This species is known 
from the Western Sahara south to Angola. 

Remarks: When ADAM AND KNUDSEN 
(1969) reviewed the taxon Circulus stria- 
tus from West Africa, they mixed up 
more than one species. They studied 21 
shells from 7 localities, 14 of which from 
Cotonou (Dahomey). They described 


20 


and figured as C. striatus the only shell 
from Illes de Los, West of Crawford 
Banc, which in our opinion does not 
belong to this species but to another that 
we will describe below. They compared 
the shells from Cotonou, which were 
also illustrated, commenting that they 
could correspond to Vitrinella congoensis 
Thiele, 1925, but considering it as a 
variety of C. striatus. After examining 
the photographs of the holotype of Vit- 
rinella congoensis Thiele in ZMB, we 
confirm that the shell of V. congoensis is 
really very similar to the shells repre- 
sented by Adam and Knudsen as C. 
striatus from Cotonou, but in our 
opinion it is different from the true C. 
striatus. For this reason we consider that 
V. congoensis Thiele is a valid species of 
Circulus, different from C. striatus. 

Probably, the confusion of Adam 
and Knudsen could be due to the study 
of a large lot of Circulus (approximately 
150 shells) from Arcachon in the 
Dautzenberg collection. The shells in 
this lot (not examined by us) were 
apparently recent and showed great 
variability: some of them had a smooth 
base, which is usual in the typical Euro- 
pean C. striatus, whereas others had a 
striated base and even intermediate 
grades appeared. Therefore, the decision 
of Adam and Knudsen to consider all of 
them (those from Arcachon and those 
from West Africa) as C. striatus is under- 
standable in view of the low number of 
shells examined from some areas, their 
similarity and small size, and the lack of 
electronic microscopy. 


OLIVER AND ROLÁN: The family Tornidae in the East Atlantic, 2. Circulinae 


Figure 7. Circulus smithi Bush, 1897. A, B: shells, 2.7, 2.7 mm, Nouakchott, Mauritania (CJH); 
CD: shells2.2,2.9'mm, Dakar, Senegal. 
Figure 7. Circulus smithi Bush, 1897. A, B: conchas, 2,7; 2,7 mm, Nouakchott, Mauritania (CJH); 
C, D: conchas, 2,2; 2,9 mm, Dakar, Senegal. 


The use of electronic microscopy 
allowed us to evaluate that the variabil- 
ity in C. striatus could occasionally lead 
to consider different species (like C. tri- 
carinatus). The study of many shells of 
C. striatus from the Mediterranean 
(Cullera, Valencia) and the Atlantic 
(Galicia, Morocco) showed us that those 
typical characters are constant and dif- 
ferent from other species. 


The most important differences 
between C. striatus and C. congoensis are: 

1- The presence on the entire shell of 
a light subsutural sculptured cord in C. 
striatus. In C. congoensis this cordlet is 
barely sculptured, and it is only found 
at the beginning in the teleoconch, later 
changing into a normal cord. 

2- The cords in the lateral low area 
and basal area in C. congoensis are 


21 


Iberus, 29 (1), 2011 


Figure 8. Circulus congoensis (Thiele, 1925). A-C: holotype, 1.93 mm, Congo mounth (ZMB); D: 
protoconch. 
Figura 8. Circulus congoensis (Thiele, 1925). A-C: holotipo, 1,93 mm, desembocadura del Congo 
(ZMB); D: protoconcha. 


evident and have the same size as the 
dorsal ones, but are a little flatter on the 
base. In C. striatus the base area appar- 
ently lacks any cord and has a very light 
and almost smooth aspect, which can be 
seen under strong magnification. It is 
possible that in the material from Arca- 
chon studied by Adam and Knudsen 
some shells of C. striatus with basal 
cords were present, due to the great 


22 


variability of the species. These shells 
never have stronger marked cords as 
does C. congoensis. 

3- The umbilicus of C. striatus is 
more open than that of C. congoensis. 

4- C. striatus is proportionally flatter 
than C. congoensis. 

5- The umbilical cords in C. striatus 
are more axially sculptured than those 
of C. congoensis. 


OLIVER AND ROLÁN: The family Tornidae in the Fast Atlantic, 2. Circulinae 


200 um. 50 um 


50 um 


Figure 9. Circulus congoensis (Thiele, 1925). A-D: shells: 1.7, 1.7, 1.6, 1.6 mm, Miamia, Ghana 
(MHNS); E: detail of the umbilicus; E G: protoconchs; H, l: microsculpture. 

Figura 9. Circulus congoensis (Thiele, 1925). A-D: conchas: 1,7; 1,7; 1,6; 1,6 mm, Miamia, Ghana 
(MHNS), E: detalle del ombligo; E G: protoconchas; H, 1: microescultura. 


Iberus, 29 (1), 2011 


Circulus senegalensis Adam and Knudsen, 1969 (Figs. 10A-F) 
Circulus senegalensis Adam and Knudsen, 1969: 13, fig. 6. [Type locality: Senegal, 60 m]. 


Type material: Holotype and 29 paratypes in RBINS. 

Other material examined: Western Sahara: Dakhla: 1 s, beach sediments (MHNS). Mauritania: 1 
s, Nouakchott, fishermen dredgings, 80-100 m (CJH). Ivory Coast: 11 s, Centre Oceanografique, 
Abdijan, stn 13, (MNHN); 1 s, Radiale Grand Bassam R/V “Antea Benchaci I”, 5 05' N, 3” 46.6' W, 
55 m (MNHN): 1 s, Radiale Gd. Bassam R/V “Antea Benchaci I”, 5" 06' N, 3 46.6! W, 50 m (MNHN). 
Ghana: 2 s, Miamia, 50 m (MHNS); 31 s, Miamia, 45-50 m (MHNS); 4 s, 38-40 m (MHNS); 12 s, Cap 
Three points, 35-65 m (MHNS). Congo: 1 s, Sta. 964, R/V André Nizery, 5” 25' S, 12” 01 E, 70 m 
(MNAHN ) 1 s, Sta. 949, R/V André Nizery, 5” 23' S, 11” 48' E, 40 m (MNHN); 5 s, Sta. 916, R/V André 
Nizery, 9405 101804 E 00m (MNHN ); 1 s, Sta. 1031, R/V André Niza Slots OZ 2 ESO 
m (MNHN). Angola: 2 s, Cabinda, W Landana 5” 07' S, 12” 01' E, 9 m (MNHN); 32 s, 5 j, Mussulo, 
Luanda, 20-100 m (MNHN); 25 s, 50 j, Luanda, 50-70 m (MNHN); 4 s, Luanda, 100 m (MHNS). 


Description: See ADAM AND KNUDSEN 


(1969). Shell (Figs. 10A-D) flat, with 4.5. 


whorls, solid, whitish, reaching up to 2.8 
mm in width and 1.7 mm in height. 

Protoconch (Fig. 10E) multispiral with 
almost two whorls and a diameter of about 
540 um. The transition to the teleoconch 
is difficult to see because of the lack of 
sculpture on this part of the surface. 

The teleoconch is smooth except for 
the presence of three (sometimes four) 
umbilical cords separated by sulci, the 
innermost one being more developed. 
Axial ornamentation formed by little 


sinusoidal growth lines, which when 
crossing the umbilical cords give them 
an undulating aspect only visible under 
strong magnification. 

Aperture circular. At those points 
where keels appear in other species, a 
sinusoidal profile may be observed. 

Distribution: This species is found all 
along the African coast from Western 
Sahara south to Angola. 

Remarks: This species is very distinct 
because it is smooth on the dorsum and 
thus quite different from other species 
occurring in the area studied. 


Circulus pseudopraecedens Adam and Knudsen, 1969 (Figure 11A-D) 


Circulus pseudopraecedens Adam and Knudsen, 1969: 14, fig. 7. [Type locality: Grand Cess, 
Liberia]. 


Type material: Holotype and 7 paratypes in RBINS. Holotype figured in ADAM AND KNUDSEN 
(1969). 

Other material examined: Senegal: 2 j, Dakar, 20 m (MHNS); 2 s, Casamance, 12” 20,7" N, 16" 53,1" 
W, 15 m (MNAHN); 1 s, Le Tacoma, 15 m (CJP). Ghana: 22 s, 50 j, 3 f, Miamia, 35-40 m (MHNS); 8 s, 
1 j, Miamia, 40-55 m (MHNS). Equatorial Guinea: 2 s, Río Núñez, 1? 35' N, 15” 26" W, 9 m (MHNS); 
1 s, W Ile Quito, R/V “André Nizery” Sedigui II, 10% 00' N, 15” 46' W, 28 m (UNHN). Angola: 7 s, 


Luanda, 20-100 m (MHNS). 


Description: Shell (Figs. 11A-C): see 
ADAM AND KNUDSEN (1969). 

Shell circular, depressed, with 4.8 
whorls (protoconch included) which 
reaches 3.3 mm in diameter and 1.3 mm 
in height. Protoconch (Fig. 11D), with a 
little less than 3 whorls and about 800 um 
in width. The transition to the teleoconch 
is difficult to see, but appears at the begin- 
ning of the upper cord. The shell has a vit- 


24 


reous, slightly transparent aspect. It has a 
clear peripheral keel and a spiral cord like 
an upper keel. In most of the shells there 
is no basal keel but ADAM AND KNUDSEN 
(1969) mention that they found a shell from 
Rufinesque Bay with this keel. 

The subsutural area is flat and 
smooth without subsutural cords. The 
upper lateral area can present some 
scarcely prominent narrow cords. A 


OLIVER AND ROLÁN: The family Tornidae in the East Atlantic, 2. Circulinae 


P 
1 
4 
. 


Figure 10. Circulus senegalensis Adam and Knudsen, 1969. A, B: shells, 1.9, 2.0 mm, Abidjan, 
Ivory Coast (MHNS); C, D: shells, 2.1, 1.3 mm, Luanda, Angola (MNHN); E: protoconch; E: 


detail of the umbilical sculpture. 
Figura 10. Circulus senegalensis Adam y Knudsen, 1969. A, B: conchas, 1,9; 2,0 mm, Abidjan, Costa 


de Marfil (MHNS); C, D: conchas, 2,1; 1,3 mm, Luanda, Angola (MNHN); E: protoconcha; F: 
detalle de la escultura umbilical. 


25 


Iberus, 29 (1), 2011 


Figure 11. Circulus pseudopraecedens Adam and Knudsen, 1969. A-C: shells, 3.2, 2.8, 3.1 mm, 


Miamia, Ghana (MHNS); D: protoconch. 


Figura 11. Circulus pseudopraecedens Adam y Knudsen, 1969. A-C: conchas, 3,2; 2,8; 3,1 mm, 


Miamia, Ghana (MHNS); D: protoconcha. 


cord, placed at */3 between the suture 
and the peripheral cord appears at the 
beginning of the teleoconch and repre- 
sents the upper keel. Below, other small 
cords may appear but are barely notice- 
able. 


26 


The peripheral keel is the most out- 
standing feature of the sculpture. The 
lateral low cords, the basal keel and the 
basal cords are absent or are very slight 
in juvenile shells. On the contrary, in 
adult shells, after some scars they can 


OLIVER AND ROLÁN: The family Tornidae in the East Atlantic, 2. Circulinae 


appear more evidently. From a basal 
view the shell shows a wide and deep 
umbilicus. In its interior there are 5-6 
umbilical cords. 

When it comes to the axial 
microsculpture, only sinusoidal growth 
lines can be found. 

The aperture is circular and proso- 
cline, and from a basal view the upper 


part of the external lip clearly extends 
beyond the lower part. 

Distribution: Known from Senegal to 
Angola. 

Remarks: The characteristic shape 
with constant and prominent keels and 
lacking other intermediate cords sepa- 
rates this species from other congeners 
in the studied area. 


Circulus stephani Rolán and Ryall, 2002 (Figure 12A-F) 


Circulus stephani Rolán and Ryall, 2002. Iberus, 20 (1): 95, figs. 1-6. [Type locality: Mimia, Ghana, 
38-40 ml). 


Type material: Holotype (Fig. 12A) in MNCN and paratypes in several museums (MNHN, MHNS, 
etc.) mentioned in the original description. 

Other material examined: Ivory Coast: 1 s, Abdijan Grand Bassan (col. Leboeuf Ortson) (MNHN). 
Ghana: 8 s, 38-40 m; (MHNS); 4 s, Cap Three Points, 35-65 m (MHNS). Angola: Cabinda: 1 s, W. 
Luanda 10? 05' S, 119 59 E, 25 m; R/V André Nizery (col. MNHN) (also that mentioned in the orig- 


inal description). 


Description: Shell (Figs. 12A-E): see 
ROLÁN AND RYALL (2002). Shell circular, 
depressed, with about 4.5 whorls reach- 
ing 2.1 mm in diameter. Protoconch (Fig. 
12F) with almost 2.75 whorls and 740 um 
in width. Circulus stephani presents three 
evident keels: upper, peripheral and basal. 
The upper one is the least developed, while 
the basal one is the strongest, reaching the 
maximum width of the shell. 

The subsutural cords appear at the 
beginning of the teleoconch. They are 


clearly narrower than the corresponding, 
interspaces. The first one to appear will 
be the upper keel. Later, on the subsu- 
tural area the other four cords appear. 
As the shell grows in size, more cords 
appear, while the largest shells can have 
up to 10 subsutural cords. 

Distribution: Known from Ivory 
Coast to Angola. 

Remarks: See ROLÁN AND RYALL 
(2002) for differentiation from other 
species of the genus in West Africa. 


Circulus microsculpturatus spec. nov. (Figures 13A-F) 


Type material: Holotype in MNHN (23687) (Figs. 13A-B). Paratype; 1 s, Dakhla, 50 m, Western 


Sahara in MNCN (15.05 /55052). 


Type locality: Sediments trawled at 50 m, Guinea Conakry. 
Etymology: The specific name alludes to the microsculpture characteristic of the present species. 


Description: Shell (Fig. 13A-B) circu- 
lar, depressed, solid, with almost 4.5 
whorls, reaches 3.5 mm in diameter and 
1.5 mm in height. Protoconch (Fig. 13C) 
with 2 whorls and 510 um in diameter. 
The main sculpture is formed by spiral 
cords, narrower than the corresponding, 
interspaces. In these  interspaces 
appears a granular microsculpture 
which is characteristic of this species 


(Figs. 13D-F). Three of the cords are 
more developed forming keels: upper, 
peripheral and basal. The basal one is 
particularly developed, giving a rectan- 
gular appearance to the peristome. 
Between the suture and the upper keel 
there are 5-6 spiral cords. Between the 
upper and the peripheral keels there are 
23 cords, of which the upper one is 
nearly as large as the keel itself. 


27 


Iberus, 29 (1), 2011 


Figure 12. Circulus stephani Rolán and Ryall, 2002. A: holotype, 2.1 mm (MNCN); B: paratype, 
1.7 mm (MNHN); C: paratype, 2.0 mm (AMNH); D: paratype, 1.6 mm (NHMUK); E: 
paratype, 2.1 mm (MHNS); F: protoconch of the holotype. 

Figura 12. Circulus stephani Rolán y Ryall, 2002. A: holotipo, 2,1 mm (MNCN); B: paratipo, 1,7 
mm (MNHN); C: paratipo, 2,0 mm (AMNH); D: paratipo, 1,6 mm (NHAMUK); E: paratipo, 2,1 
mm (MHNS); F: protoconcha del holotipo. 


Between the peripheral and the basal 
keels there are four cords. There is not a 
clear basal cord, the base being occu- 
pied by about 14 cordlets with a width 
similar to that of the interspaces. They 
go into the umbilicus. 


28 


Distribution: Only known from the 
type material (From Sahara to Guinea 
Conakry). 

Remarks: The microsculpture of the 
present species differentiates it from the 
other congenericspeciesin the studied area. 


OLIVER AND ROLÁN: The family Tornidae in the East Atlantic, 2. Circulinae 


200 um 


Figure 13. Circulus microsculpturatus spec. nov. A, B: holotype, 3.4 mm, Guinea Conakry 
(MNHN); C: protoconch of the holotype; D-F: microsculpture. 


Figura 13. Circulus microsculpturatus spec. nov. A, B: holotipo, 3,4 mm, Guinea Conakry (MNAN); 
C: protoconcha del holotipo; D-F: microescultura. 


Iberus, 29 (1), 2011 


Circulus ryalli spec. nov. (Figure 14A-D) 


Circulus striatus in Adam and Knudsen (1969): 10, fig. 4. 


Type material: Holotype (Fig. 14A) in MNCN (15.05 /55053). Paratypes in the following: MNHN 
(23688) (1 s, Fig. 14B), MHNS (5 s, 1 j, Fig. 14C), CPR (1 s). 

Type locality: Miamia, Ghana, in sediments dredged in 38-45 m. 

Etymology: The species is named after Peter Ryall, a malacologist who lived for many years in 
Ghana and helped us during the collection of the type material. 


Description: Shell (Figs. 14A-C) circu- 
lar, depressed, with 4.6 whorls (proto- 
conch included), reaching 1.9 mm in 
diameter and 0.9 mm in height. Proto- 
conch (Fig. 14D) smooth, with 2 Y 
whorls, and about 510 um in diameter. 
Teleoconch with slightly prosocline spiral 
cords and axial growth lines which cross 
the interspaces, giving them a striated 
appearance. The shell has a clear periph- 
eral keel, and in the places which would 
correspond to the other keels there are 
somewhat developed cords; as the shell 
increases in size the basal cord progres- 
sively becomes keel-shaped. The subsu- 
tural area is flat with six or seven cords 
which appear after the protoconch scar. 
These cords are wider than their inter- 
spaces at the beginning, but subsequently 
the interspaces increase in size while the 
cords continue the same, so that at the 
end of the spire the cords are clearly nar- 
rower than their interspaces. Among the 
subsutural cords, the adapical one is a 
little wider and placed in the position of 
the upper keel. The upper lateral area is 
flat and so the transition has an angled 
aspect (about 100). Between the upper 
cord /keel and the peripheral keel, there 
are 6-7 cords and, with the shell in lateral 
view, they seem to be flatter and wider 
than the corresponding interspaces. In 
the upper lip one or two cords are 
inserted on the peripheral keel. The 
lateral lower area does not have cords 
and is slightly concave. The basal area is 
flat with 4-5 cords, two of which are more 
developed, in the position of the perium- 


REMARKS AND FINAL DISCUSSION 


Eight species of the genus Circulus 
from the East Atlantic area have been 


30 


bilical cord /keel and 2-3 lighter, in the 
transition with the umbilical area. 
Between these two groups of cords there 
is a wide space. The umbilicus is open 
and in its interior there are about 7-8 
spiral cords as wide as the interspaces, 
crossed by growth lines which give them 
a striated aspect. 

Dimensions: the holotype is 1.97 mm. 

Distribution: Only known from the 
type material. 

Remarks: This species appears to be 
that which ADAM AND KNUDSEN (1969) 
considered as C. striatus from Iles de 
Los, to the W of Crawford Bank. 

Circulus striatus lacks true cords at the 
base which is smooth; in the umbilical 
cords, there is a typical microsculpture 
while in C. ryalli there only appear fine 
axial growth lines; the protoconch of C. 
striatus has between 2 and a little more 
whorls, with 390 um in diameter, while C. 
ryallihhas a protoconch with 2 Y whorls and 
about 510 um. The beginning of the teleo- 
conch in C. ryalli shows evident cords, all 
similar, while those in C. striatus are gen- 
erally more attenuate and the subsutural 
one has constantly visible microsculpture. 

Circulus senegalensis has a smooth 
shell lacking keels and cords. 

Circulus pseudopraecedens has keels 
but not spiral cords between them. 

Circulus smithi has fewer cords on 
the dorsal part, the space between the 
keels being more occupied by cords. 

Circulus microsculpturatus spec. nov. 
has a more evident microsculpture over 
the entire shell. 


studied in the present work. Most of the 
species have a tropical distribution, 


OLIVER AND ROLÁN: The family Tornidae in the East Atlantic, 2. Circulinae 


200 um 


Figure 14. Circulus ryalli spec. nov. A: holotype, 1.97 mm, Miamia, Ghana (MNC); B: paratype, 
1.9 mm (MNAHN); C: paratype, 1.9 mm (MHNS); D: protoconch of the holotype. 

Figura 14. Circulus ryalli spec. nov. A: holotipo, 1,97 mm, Miamia, Ghana (MNC); B: paratipo, 1,9 
mm (MNAN); C: paratipo, 1,9 mm (MHNS); D: protoconcha del holotipo. 


from West Sahara south to Angola, have been collected throughout the 


which seems to be an area in common to entire area. 
all of them although not all (mainly the Of those, the oldest and type species 
ones of which less material is available) of the genus is C. striatus. This species 


31 


Iberus, 29 (1), 2011 


extends from the cold waters of north- 
ern Europe south to the temperate 
waters of the Mediterranean and north- 
ern and western Morocco. It is not 
present in the rest of the African west 
coast but has been erroneously confused 
with other species on several occasions. 

As is usual in species with a wide 
dispersal area, all of them have multi- 
spiral protoconchs (between 2 and 3 
spiral whorls). 

Some European taxa which have 
been included in Circulus and do not 
belong there are the following; 

Adeorbis subcarinatus Montagu, 1803, 
now considered as a species of the 
genus Tornus. 

Circulus jeffreysii Monterosato, 1872: 
according to WARÉN (1992) it is a 
Skeneid species, not a Circulus. 

Circulus formossisimus Brugnone, 
1873. Synonymized with Circulus je- 


ffreyst. 


BIBLIOGRAPHY 


AARTSEN J.J. VAN, MENKHORST H.P.M.G. AND 
GITTENBERGER E. 1984. The marine Mollusca 
of the Bay of Algeciras, Spain, with general 
notes on Mitrella, Marginellidae and Turri- 
dae. Basteria. Suppl. 2: 1-135. 

ADAM W. AND KNUDSEN J. 1969. Quelques 
genres de Mollusques prosobranches ma- 
rins inconnus ou peu connus de l'Afrique 
occidentale. Bulletin Institut Royal des Sciences 
Naturelles de Belgique, 44 (27): 1-69. 

BOUCHET P. AND ROCROI J.P. (Ed.); Eryda J., 
Hausdorf B., Ponder W., Valdés Á. and Wa- 
rén A. 2005. Classification and nomenclator 
of gastropod families. Malacología: Interna- 
tional Journal of Malacology, 47 (1-2): 397 pp. 
ConchBooks, Hackenheim, Germany. 

COPPINI M., CUNEO F., MARGELLI A. AND CAM- 
PANI E. 2005. Gastropoda e Scaphopoda del 
porto de Livorno. Bollettino Malacologico, 41 
(5-8): 1-8. 

DaLL W.H. 1927. Small shells from dredgings 
off the southeast coast of the United States 
by the United States Fisheries steamer Alba- 
tross in 1885 and 1886. Proceedings of the United 
States National Museum, 70 (2667): 1-134. 

FERNANDES F. AND ROLÁN E. 1993. Moluscos 
marinos de Sáo Tomé y Principe: actualiza- 
ción bibliográfica y nuevas aportaciones. 
Iberus, 11 (1): 31-47. 


32 


ACKNOWLEDGEMENTS 


The authors want to express their 
thanks to Philippe Bouchet of the 
MNHN for the loan of numerous 
samples from this museum. Jesús 
Méndez and Inés Pazos of the Centro de 
Apoyo Científico y Tecnológico a la 
Investigación (CACTI) of the University 
of Vigo and to Ramiro Barreiro Pérez of 
the Centro de Apoyo Científico y Tec- 
nológico of the University of Santiago 
de Compostela (CACTUS) for the 
microphotographs of the material 
studied in this work. Other SEM pho- 
tographs were made by Marta Furió, 
Laura Tormo and Alberto Jorge in the 
MNCN. Also to José María Hernández 
(+) from Gran Canaria, Anselmo Peñas 
from Vilanova i la Geltrú and Peter 
Ryall, from Maria Rain, for the loan of 
material. António A. Monteiro of Lisbon 
made the corrections to the language. 


FISCHER P. 1887. Manuel de conchyliologie et 
de paléontologie conchyliologique : ou, His- 
toire naturelle des mollusques vivants et fos- 
siles. Paris, Savy. xx1v + 1369 p., 23 pl. 

FRETTER V. AND GRAHAM A. 1962. British proso- 
branch molluscs: their functional anatomy and 
ecology. Ray Society, London, xvi + 755 pp. 

FRETTER V. AND GRAHAM A. 1978. The proso- 
branch molluscs of Britain and Denmark. 
Part 4 - Marine Rissoacea. Journal of Mollus- 
can Studies, supplement 6: 153-241. 

GRAHAM A. 1982. Tornus subcarinatus (Proso- 
branchia, Rissoacea) anatomy and relation- 
ships. Journal of Molluscan Studies, 48: 144-147. 

GRAHAM A. 1988. Molluscs: Prosobranch and 
Pyramidellid Gastropods. Synopses of the 
British Fauna, 2. Linnean Society of London, 
662 pp. 

HIDALGO J.G. 1917. Fauna malacológica de Es- 
paña, Portugal y las Baleares. Trabajos del Mu- 
seo Nacional de Ciencias Naturales, 30, Ma- 
drid: 792 pp: 

JEFFREYS J.G. 1865. British conchology, or an ac- 
count of the Mollusca which now inhabit the 
British Isles and the surrounding seas. Vol. 111. 
Marine shells, comprising the remaining 
Conchifera, the Solenoconchia, and Gasteropoda 
as far as Littorina. ]. van Noorst, London, 393 
(+ 1) p., 8 pls. 


OLIVER AND ROLÁN: The family Tornidae in the East Atlantic, 2. Circulinae 


LOCARD A. 1889. Matériaux pour servir l'histoire 
de la malacologie francaise. VII. Note sur les 
especes francaises appartenant au genre Cir- 
culus. Bulletin de la Société Malacologique de 
France, 6: 283-307. 

NOBRE A. 1938-40. Fauna malacológica de Portu- 
gal. Moluscos marinhos e das aguas salobras. 
Companhia Editora do Minho, Barcelos. 807 
pp, 87 láms. 

ROBBEYG+I""AND.GOLO Y. 1991 “European 
Seashells. vol 1. Christa Hemmen, Darmstadt. 
352 pp. 

ROLÁN E. 1983. Moluscos de la Ría de Vigo, 1 
Gasterópodos. Thalassas, 1 (1), supl. 1: 1-383 

ÓN E. AND RUBIO F. 1991 “1990”. Aporta- 
ciones a los conocimientos sobre los micro- 
moluscos de África Occidental. 1. Tornidae 
de Sáo Tomé y Principe. Iberus, 9 (1-2): 181- 
186, 1 lám. 

ROLÁN E. AND RUBIO F. 1996. Un nuevo vitri- 
nélido (Mollusca, Gastropoda, Vitrinellidae) 
de la isla de Sáo Tomé (África occidental). 
Iberus, 14 (2): 143-146. 

ROLÁN E. AND RUBIO F. 2002. The family 
Tornidae (Gastropoda, Rissooidae) in the 
east Atlantic. Reseñas Malacologicas, supp!l.: 1- 
98. 


ROLÁN E., RUBIO F. and RYALL P. 2000. A new 
species of Teinostoma (Mollusca, Gastropoda, 
Vitrinellidae) from Ghana (W. Africa). Ar- 
gonauta, 13 (2): 81-82. 

ROLÁN E. AND RYALL P. 2002. A new species of 
the genus Circulus (Gastropoda, Adeorbidae) 
from West Africa. Iberus, 20 (1): 95-98. 

RUBIO F. AND ROLÁN E. 1991 “1990”. Aporta- 
ciones a los conocimientos sobre los micro- 
moluscos de África Occidental. 2 Archae- 
gastropoda de Sáo Tomé y Principe. Iberus, 
9 (1-2): 209-219, 5 lám. 

TERRENI G. 1981. Molluschi conchiferi del mare an- 
tistante la costa Toscana. 1-102 pp. Livorno. 
THIELE J. 1929-1935. Handbuch der Systematischen 

Weichtierkunde. 2 vols. 1154 p., 584 figs. 

VERDUIN A. 1976. On the systematic of recent 
Rissoa of the subgenus Turboella Gray, 1847, 
from the Mediterranean and European At- 
lantic coasts. Basteria, 40: 21-73. 

WARÉN A. 1992. New and little known 
“Skeneimorph” gastropods from the Mediter- 
ranean Sea and the adjacent Atlantic Ocean. 
Bollettino Malacologico, 27 (10-12): 149-248. 

WOOD S.V. 1848. A Monograph of the Crag Mol- 
lusca. 1. Univalves. Paleontological Society, 
London. 208 pp, 21 pls. 


33 


O Sociedad Española de Malacología 


Iberus, 29 (19:35-38, 2011 


First Mediterranean record of Diaphana marshalli (Sykes, 
1904) (Gastropoda, Diaphanidae) - a rare bathyal species 


Primera cita para el Mediterráneo de Diaphana marshalli (Sykes, 


1904) (Gastropoda, Diaphanidae) - una especie rara del batial 


Enzo CAMPANT*, Cesare BOGI** and Bella S. GALIL*** 


Recibido el 4-11-2011. Aceptado el 20-111-201 1 


ABSTRACT 


The rare bathyal gastropod Diaphana marshalli (Sykes, 1904), previously known only for 
the Atlantic, is reported for the first time in the Mediterranean. The specimens, collected off 
Hadera, Israel, are photographed and compared with its Mediterranean congener, D. jon- 


ica Di Geronimo, 1974 


RESUMEN 


Se cita por primera vez en el Mediterráneo el gasterópodo Diaphana marshalli (Sykes, 
1904), una especie rara del batial, previamente conocida solamente en el Atlántico. La 
cita se basa en ejemplares recogidos frente a Hadera, Israel, que se ilustran y se compa- 
ran con su congénere del Mediterráneo, D. jonica Di Geronimo, 1974 


INTRODUCTION 


The bathyal molluscs of the Levan- 
tine Sea are scarcely known. Nearly a 
century passed from STUARNY's (1896) 
publication of the specimens collected 
during the voyages of the “POLA” 
(1890-1893), JANSSEN's (1989) report of 
those collected by “METEOR” [cruise 5] 
(1987) and Bogi's account of the few 
specimens collected by “POSEIDON” 
[cruise 201 /2] (1994) from Eratosthenes 
Seamount (GALIL AND  ZIBROWIUS, 
1998). A total 42 species were identified 
from material collected during a series 
of cruises conducted between 1994 and 
1999 as part of pollution monitoring 
surveys by the Israel Oceanographic 
and Limnological Research (IOLR) off 


the northern coast of Israel (BOGI AND 
GALIL, 2004). Of these, five species con- 
stituted new records for the eastern 
Mediterranean, and 8 were newly 
recorded from the Levantine Sea. Two 
specimens, earlier considered juveniles 
of an unidentified gastropod, have been 
recently re-examined and recognized as 
Dipahana marshalli (Sykes, 1904). 


MATERIALS AND METHODS 


The area investigated is located off 
the coast of Israel, at depths between 
1227 and 1454 m. The material was col- 
lected during monitoring surveys of a 


* Corso G. Mazzini 299, 57126 Livorno, Italy, E-mail enzo.campaniCfastwebnet.it 
** Via delle Viole 7, 57124 Livorno, Italy, E-mail bogicesarePtiscali.it 
*** Israel Oceanographic and Limnological Research, National Institute of Oceanography, P.O.B. 8030, Haifa 


31080, E-mail bellaGocean.org, il 


35 


Iberus, 29 (1), 2011 


deepwater coal fly ash disposal site off 
Hadera (between 32* 38” N 34? 02 E and 
32 36” N 34" 16' E). The samples were 
collected aboard the R/V Shikmona, 
using a 45 ft Marinovich type deep 
water trawl (codend mesh 6 mm) with a 
0.5 mm plankton net secured atop. The 
samples were preserved in 10% buffered 
formalin aboard ship. In the laboratory, 


TAXONOMY 


the samples were washed and sieved 
through a 250 um mesh, preserved in 
70% alcohol, stained in Rose Bengal and 
sorted. Two specimens (with soft parts) 
were collected at a depth of 1400 m in 
September 1997 [1,5 x 0,91 mm (fig. 2); 
1,3 x 0,8 mm (figs. 3 — 5)]. The smaller 
specimen was damaged, the larger one 
is preserved in the Bogi collection. 


OPISTHOBRANCHIA Milne-Edwards, 1848 
CEPHALASPIDEA P. Fischer, 1883 
DIAPHANIDAE Odhner, 1914 
Diaphana Brown, 1827 


Diaphana marshalli (Sykes, 1904) 


Discussion: Diaphana marshall is an 
exceedingly rare species. Aside from the 
type series collected off Portugal, only 
two additional specimens are known: a 
shell collected by the “THALASSA” 
Expedition (station Z 447, 48” 47”, 11 
12, 1450 — 1500 m) and cited as Retusa 
(2) marshalli by BOUCHET (1975), and a 
2.1 mm high specimen with soft parts, 
collected by the “INCAL” Expedition 
1976 (station CP08, 50” 15” N, 13” 14 W, 
2644 m) (SCHIOTTE, 1999). 

Retusa marshalli was described and 
illustrated (SYKEs, 1904: 31, figs. 5, 6, 6a, 
b) from specimens collected by the 
“PORCUPINE” Expedition of 1870, off 
Portugal, at station 17, at 600-1095 
fathoms. Only a single specimen was 
measured (“Alt. 2, diam. max. 1.1 mm”, 
there, p. 32), though the author refers to 
another, larger specimen (“one specimen, 
broken, is larger than the type”, there, p. 
32). SYKES'” illustration (1904, fig. 6) is 
reproduced (fig. 1). 

BOUCHET (1975: 334) expressed 
doubt on the species” generic placement: 
“Des caracteres tels que la présence d'un 
ombilic ou la conformation du sommet sem- 
blent tres originaux pour une Retusa et il 
est vraisemblable que lorsque l'animal sera 
connu, une séparation générique deviendra 
nécessaire”. BOUCHET AND WARÉN (1979) 
examined Sykes” material, illustrated 


36 


the shell and referred to it as “holotype” 
(there, fig. 18 1), though that shell, at 2.45 
mm, was rather larger that the specimen 
measured by Sykes. These authors 
declared *Retusa* marshalli “probably a 
Diaphana” (there, p. 237). Schiotte (1999) 
reported that the type material consisted 
of a “holotype” 2.45 mm high 
(BMNH1922122) and “pieces of 4 or 5 
additional specimens, labelled as syntypes, 
all dry shells” (there, p. 87). 

Few species of Diaphana Brown, 
1827 have been recorded from the 
Mediterranean Sea, but the only one 
resembling D. marhalli is D. jonica Di 
Geronimo, 1974. The latter has been 
considered a junior synonym of D. lactea 
(Jeffreys, 1877) by BOUCHET AND WARÉN 
(1979) who examined the type. 
SCHIOTTE (1999: 124) who compared 
illustrations of both species declared ”... 
there is a very good resemblance between the 
shells of D. lactea and D. jonica.”. 

BOUCHET AND WARÉN (1979: 237) 
separated “Retusa' marshalli from its con- 
gener D. lactea as the former has ”... a 
whitish unsculptured shell with an umbili- 
cal chink, but the larval shell is white, not 
brown as in lactea, and the broadest part of 
the aperture is at the lower third of the shell, 
instead of the middle of the shell. In “R'. 
marshalli the larval shell is completely 
visible even in adult specimens when seen 


CAMPANI £T AL.: First Mediterranean record of Diaphana marshalli, a rare bathyal species 


Figures 1-5. Diaphana marshalli (Sykes, 1904). 1: Sykes” illustration, reproduced, height 2.45 mm; 
2: Scanning electron micrograph of the larger Levantine specimen, height 1.5 mm; 3: adapertural 
view of smaller Levantine specimen, height 1.3 mm; 4: side view of the same specimen; 5: detail of 
the apex of the same specimen. 

Figuras 1-5. Diaphana marshalli (Sykes, 1904). 1: Reproducción de la ilustración de Sykes, altura 2,45 
mm; 2: Micrografía electrónica de barrido del mayor ejemplar levantino, altura 1,5 mm, 3: Vista ada- 


pertural del menor ejemplar levantino, altura 1,3 mm, 4: vista lateral del mismo ejemplar; 5: detalle 
del ápice del mismo ejemplar. 


3/ 


Iberus, 29 (1), 2011 


from the adapertural side, while in lactea 
only the topmost part of the larval shell pro- 
trudes above the top of the body whorl. “R”. 
marshalli is smaller, being only 2.45 mm 
when adult”. 

On comparing DI GERONIMO's (1974) 
description and SEM image of the holo- 
type of D. jonica with the Levantine 
specimens of D. marshalli it was appar- 
ent that the shell of the former is more 
cylindrical and slender than the shell of 
the latter species, its height-to-width 
ratio being 1,84 compared with 1,6-1,65 
in marshalli; the columella differently set 
and the umbilicus narrower; the apex of 
D. jonica is described as ”piúu o meno 
sporgente”, but is similar to that of D. 
lactea as figured in BOUCHET AND 
WARÉN (1979) with only the topmost 
part of the larval shell protruding above 
the body whorl. 

The reproductive biology of D. mar- 
shalli is unknown, but the size of its larval 
shell indicates planktotrophic larval 
development (BOUCHET AND WARÉN, 


BIBLIOGRAPHY 


BOGI C. AND GALIL B., 2004. The bathybenthic 
and pelagic molluscan fauna off the Levan- 
tine coast, eastern Mediterranean. Bollettino 
Malacologico, 39: 79-91 

BOUCHET P., 1975. Opisthobranches de pro- 
fondeur de l'Ocean Atlantique. 1 Cephalas- 
pidea. Cahiers de Biologie Marine, 16: 317-365 

BOUCHET P. AND WARÉN A,, 1979. The abyssal 
molluscan fauna of the Norvegian Sea and 
its relation to other faunas. Sarsia, 64: 211-243 

DI GERONIMO L, 1974. Molluschi bentonici in 
sedimenti recenti batiali e abissali dello Jo- 
nio. Conchiglie, 10 (7-8): 133-172. 

GALIL B.S. 2004. The limit of the sea: the bathyal 
fauna of the Levantine Sea. Scientia Marina, 
68 (Suppl.3): 63-72. 

GALIL B.S. and ZIBROWIUS H. 1998. First benthos 
samples from Eratosthenes seamount, East- 
ern Mediterranean. Senckenbergiana maritima, 
28(416): 111- 121. 

JANSSEN R., 1989. Benthos-Mollusken aus dem 
Tiefwasser des óstlichen Mittelmeeres, 
gesammelt wáhrend der “METEOR”- Fahrt 
5 (1987). Senckenbergiana maritima, 20 (5/6): 
265-276. 


38 


1979: 231). SCHIOTTE (1999: 133) agrees 
“... Interestingly, deep-water species usually 
seem to have planktotrophic development” . 
This may explain the occurrence of a rare 
Atlantic bathyal diaphanid gastropod in 
the Levantine Sea. The common mollusks 
at depths greater than 1000 m off the 
Israeli coast are eurybathic Atlanto- 
Mediterranean and Boreal species with 
an upper bathymetric range enabling 
them to overcome the barriers posed by 
the shallow Gibraltar Straits and the 
Siculo-Tunisian sill (< 400 m), or steno- 
bathic species with epipelagic larvae. 
Indeed, the present record of D. marshalli 
agrees with the distribution patterns of 
the Levantine deep water benthos 
(GALIL, 2004). 


-_ ACKNOWLEDGEMENTS 


We thank Dr. A. Warén, for the SEM 
images and Mr. S. Bartolini for the 
digital photographs. 


SCHIOTTE T. 1999. A taxonomic revision of the 
genus Diaphana Brown, 1827, including a dis- 
cussion of the phylogeny and zoogeography 
of the genus (Mollusca: Opisthobranchia) 
Steenstrupia, 24: 77-140 

STURANY R. 1896. Berichte der Commission fúr 
Tiefsee-Forschungen XVIII. Zoologische 
Ergebnisse VII. Mollusken I (Prosobranchier 
und Opisthobranchier; Scaphopoden; Lamel- 
libranchier) gesammelt von S.M. Schiff “Pola” 
1890-1894. Denkschriften der Kaiserlichen Aka- 
demie der Wissenschaften, Mathematische-Na- 
turwissenschaftlischen Classe, 63: 1-36, pl.1-2 

SYKES E.R. 1904. On the Mollusca procured dur- 
ing the “Porcupine” Expeditions 1869-1870. 
Supplemental notes, part I. Proceedings of the 
Malacological Society of London, 6: 23-40, pl. 


O Sociedad Española de Malacología Iberus, 29 (1):39-46, 2011 


Otala punctata (O.F. Miiller, 1774) (Stylommatophora: 
Helicidae) in Italy 


Otala punctata (O.E Miller, 1774) (Stylommatophora: Helicidae) 
en Italia 


Willy DE MATTIA* and Francesco MASCIA**! 


Recibido el 2-X-2010. Aceptado el 24-111-2011 


ABSTRACT 


A living population of Otala punctata [O.F. Muller, 1774) has been found for the first time 
in Sardinia, Italy. In the island the species was previously recorded by several authors, but 
only finding badly preserved shells, often considered of doubtful origin. An anatomical 
and conchological description is given, and a comparison with other populations of the 
species and O. lactea (Muller, 1774) from the W-Mediterranean region is carried out. The 
species distribution is limited to a coastal sector of NW-Sardinia characterized by Mediter- 
ranean maquis and rural environments. Conservation status of the population is discussed, 
and its presumable origin from the Iberian peninsula through active human transportation 
is proposed. 


RESUMEN 


Una población viva de Otala punctata [(O.F. Múller, 1774) se ha encontrado por primera 
vez en Cerdeña, Italia. Existen citas anteriores por parte de varios autores, pero sólo 
basadas en conchas mal conservadas y a menudo de procedencia dudosa. Se hace una 
descripción anatómica y conquiliológica, y se compara con otras poblaciones de la espe- 
cie y con O. lactea (Muller, 1774) de la región W-Mediterráneo. La distribución de la 
especie se limita a un sector costero del noroeste de Cerdeña, un medio rural caracteri- 
zado por matorral mediterráneo. Se examina el estado de conservación de la población 
y se sugiere su presumible origen desde de la península Ibérica a través de transporte 
antrópico. 


INTRODUCTION 


In Italy O. punctata was recorded for 
the first time (as Archelix apalopena 
(Bourguignat, 1867)) by MALATESTA AND 
SETTEPASSI (1954) from Alghero (N-W 
Sardinia). Its presence in the same area 
has been confirmed later by CARRADA, 
PARISI AND SACCHI (1967). Significantly, 
both papers reported finding badly pre- 


served shells only and no living speci- 
men was ever found. Paulucci (1886) 
reported O. lactea (Muller, 1774) from 
the Tuscan Archipelago (Argentarola 
island). This record, based on a single, 
badly preserved shell, could be referred 
to O. punctata, but Eobania vermiculata 
(O.F. Múller, 1774) cannot be excluded 


* Via di Monte San Giovanni 8, 1 34147 Muggia, Italy; wdemattiagmail.com 
** Via Oreste Salomone 32, 1 09030 Elmas, Italy; fr.maxiaCgmail.com 


' Corresponding author 


37 


Iberus, 29 (1), 2011 


(GiustTI, 1976; BODON, FAVILLI, GIUSTI 
AND MANGANELLI, 1995). 

O. punctata has a Western-Mediter- 
ranean distribution. It is found in north- 
west Algeria, eastern Spain and the 
Baleares, south-western France and 
Malta (FALKNER, 1990; CLANZIG AND 
BERTRAND, 2001; FALKNER, RIPKEN, AND 
FALKNER, 2002; BARBARA AND SCHEMBRI, 
2008). The species is widely raised in 
heliciculture and thus present also in 
North and South America, and South 
Africa (HERBERT AND SIRGEL, 2001; 
COwIE, DILLON, ROBINSON AND SMITH, 


RESULTS 


2009). The aim of this work is to clarify 
the Status of Otala punctata in Sardinia. 


MATERIAL AND METHODS 


The snails were killed in water and 
then fixed in 75% ethanol. Bodies were 
isolated from the shell and then dissected 
under an optical microscope. Anatomical 
details were drawn using a camera 
lucida. Empty shells were measured 
(n=20 for each population). The material 
is kept in the collections of the Authors. 


Otala punctata (O.F. Múller, 1774) 


Material examined [dsp, dissected specimen(s)]. — Italy: Sardinia, Alghero, Sorgente Lu Cantar, 
407 32 18” N 8? 19 30” E, 30 m alt., 07. ii. 2008, 3 dsp. F. Mascia leg.; Sardinia, Alghero, Cala Bona, 
20 m alt., 40? 32 *40” N 8919 20” E, 07. ii. 2008, 2 dsp. F. Mascia leg.; Malta: Mosta, 80 m alt., 35 54” 
34” N 14? 25' 05” E, 17. i. 2010, 2 dsp, N. Barbara leg. Spain, Málaga, El Tarajal, 30 m alt., 36? 70' 


204 NAS S0023 E, 12:x1.2008, 2 dsp, Je: Torres leg; 


Shell (Figs 9-10): The protoconch is 
dark violet-brown to dark brown. The 
teleoconch is thick and robust, globose 
to subglobose, conical above and convex 
below. There are up to 5 main, dark 
brown to light brown, variably wide 
bands, which are fused with a superim- 
posed pattern of whitish reticulation. 


Sometimes the main bands abruptly 
fade becoming barely visible and the 
shell becomes almost uniformly creamy 
in colour. The polished external surface 
shows irregular, fine growth lines and 
very fine, dense spiral lines. The spire is 
conical and not much raised with 4-5 
regularly growing whorls. The last 


(Right page) Figures 1-3. Otala punctata (O.F. Múller, 1774). Italy: Sardinia, Alghero, Sorgente Lu 
Cantar. 1, 3: genitalia (ovotestis excluded); 2: inner walls of atrium and penis. Abbreviations, A: 
atrium; AG: albumen gland; BC: bursa copulatrix; BCD: diverticulum of the bursa copulatrix; 
BW: body wall; CLS: crest-like structure; DBC: duct of the bursa copulatrix; DG: digitiform 
elands; DP: distal penis; DS: dart sac; dsp: dissected specimen[s]; DV: distal vagina; E: epiphallus; 
E: flagellum; FHD: first hermaphrodite duct; FO: free oviduct; G: glans or penial papilla; PP: 
proximal penis; PPP: pseudo-penial papilla; PR: penial retractor muscle; PSP: pilaster-shaped 
pleat; PV: proximal vagina; PW: penial wall; TILS tongue-like structure; UOS: uterine ovisper- 
miduct; VD: vas deferens. 

(Página derecha) Figuras 1-3. Otala punctata (O.E Múller, 1774). Italia: Cerdeña, Alghero, Sorgente 
Lu Cantar. 1, 3: órganos genitals (ovotestis excludo); 2: paredes internas del atrio y del pene. Abreviatu- 
ras, A: atrio; AG: glándula del albumen ; BC: bursa copulatrix; BCD: divertículo de la bursa copula- 
trix; BW. pared del cuerpo; CLS: estructura en forma de cresta; DBC: conduto de la bursa copulatrix; 
DG: glándulas digitiformes ; DP: pene distal ; DS: saco del dardo; DV: vagina distal ; E: epifalo; F: 
flagelo; FHD: primer conducto hermafrodita; FO: oviducto libre; G: glans o papila penial ; PP: pene 
proximal ; PPP: papilla pseudo-penial; PR: músculo retractor penial; PSP: pliegue en forma de pilas- 
tra; PV: vagina proximal ; PW: pared del pene; TLS estructura linguiforme; UOS: oviespermiducto 
uterino; VD: vas deferens. 


40 


De MArria AND Mascla: Otala punctata (Stylommatophora: Helicidae) in Italy 


4] 


Iberus, 29 (1), 2011 


Figures 4-6. Otala punctata (O.E Miller, 1774). 4: Spain, Málaga, El Tarajal, genitalia (ovotestis 
excluded); 5, 6: Malta, Mosta; 5: genitalia (ovotestis excluded); 6: inner walls of atrium and penis. 
Figures 4-6. Otala punctata (O.E Múller, 1774). 4: España, Málaga, El Tarajal, genitalia (ovotestis 
excluído); 5, 6: Malta, Mosta; 5: genitalia (ovotestis excluído); 6: paredes internas del atrio y del pene. 


42 


DE MArTIA AND MAscIa: Otala punctata (Stylommatophora: Helicidae) in Italy 


Figure 7. Otala lactea (Miller, 1774). Spain, Menorca, Mahon, genitalia (ovotestis excluded). 
Figura 7. Otala lactea (Múller, 1774). España, Menorca, Mahón, órganos genitales (ovotestis excluido). 


whorl is very large (70% of shell height), 
well rounded and markedly descending 
where approaching the aperture. The 
sutures are moderately deep and the 
umbilicus is closed. The aperture is 
markedly oblique and oval. The peris- 
toma is interrupted, thick and reflected 
and is whitish in colour. A columellar 
tooth, more or less evident, is often 
present. The columellar part of aperture, 
palatum and parietum are always dark 
brown in colour. 

Diagnostic characters of genitalia (Figs 
1-6): Proximal part of the vagina with 
two digitiform glands. Each gland con- 
sists of a short base which bifurcates 
into three or four main branches. Each 
main branch gives rise to numerous 
long digitiform appendices. A single, 
very large dart-sac enters about midway 
along the vagina. The dart-sac contains 
a single, straight long dart with four 
lateral wings and a spear-like head; it 
opens into the vagina with a tongue-like 
structure. Along the inner wall of vagina 
there is a huge pilaster-shaped pleat 
which reaches the large crest-like struc- 
ture in the atrium. A moderately short 


copulatory (5-7 mm) duct divides into a 
diverticulum of bursa copulatrix and a 
duct of bursa copulatrix. Diverticulum 
and duct have almost the same length. 
The bursa copulatrix is usually oval or 
round. The vagina enters the atrium 
side by side with the penis. A moder- 
ately short (8-10 mm) penial flagellum 
arises where the vas deferens enters the 
penial complex. A rather short epiphal- 
lus (6-8-mm) enters the penis (8-10 mm) 
where the penial retractor meets the 
penial wall. The penis has a proximal 
widened portion (almost twice as long 
as the distal) and enveloped in a thin 
penial sheath. The short penial papilla is 
surrounded by a solid and long pseudo- 
penial papilla with a grooved surface. 

Dimensions of Sardinian specimens: 
Shell diameter: 29 + 1.7 mm (range: 27 - 
32 mm) Shell height: 18 + 1.1 mm 
(range: 16-20 mm) (average value + 
standard deviation). 

Anatomical and conchological inves- 
tigation allowed to easily distinguish 
Otala punctata from O. lactea (Miller, 
1774) and Eobania vermiculata. Otala 
lactea has a smaller shell (average diam- 


A3 


Iberus, 29 (1), 2011 


Alghero 


Figure 8. Distribution of Otala punctata (Miller, 1774) in Sardinias Italy. 


y 


Figura 8. Distribución de Otala punctata (Múller, 1774) en Cerdeña, Italia. 


eter of 20 mm) and the inner part of the 
aperture (peristome, parietum and 
palatum) is uniformly dark brown in 
colour (Fig. 11). E.vermiculata has a spire 
somewhat more elongated and the inner 
part of the aperture is uniformly creamy 
white in colour (BODON ET AL., 1995). 
Eobanía vermiculata and O. lactea can be 
easily distinguished from O. punctata by 
the anatomy of the genitalia. Eobania 
vermiculata has a much longer flagellum 
and vagina, while the free oviduct is 
markedly shorter. The diverticulum of 
the bursa copulatrix is 6-10 times longer 
than the duct of bursa copulatrix. On 
the distal penis, a slender false penial 
papilla is present, which originates from 
the penial wall and does not surround 
the penial papilla. Otala lactea can be 
distinguished mainly in virtue of the 
much longer epiphallus and penial fla- 
gellum (Fig. 7). 

The present investigation allows us 
to confirm the presence of O. punctata in 
Italy. The species occurs exclusively in 
the southern surroundings of the town 
of Alghero (Sassari, Sardinia), along the 


44 


southern rocky shore area of the town 
(Fig. 8), from sea level to 40 meters in 
altitude. Population cover an area of 
approximately 0.4 km?. In the same area, 
the population density of Eobania ver- 
miculata (O.F. Múller, 1774) and Cornu 
aspersum (O.F. Múller, 1774) is markedly 
higher than that of O. punctata. This area 
belongs to the Mediterranean upper 
thermomediterranean thermotype (BAc- 
CHETTA, BAGELLA, BIONDI, FARRIS AND 
FILIGHEDDU, 2009). Geologically the area 
is characterized by marine and conti- 
nental Quaternary deposits (BARCA, 
CARMIGNANI, OGGIANO, PERTUSAT, SAL- 
VADORI, CONTI, ELTRUDIS, FUNEDDA AND 
Pasci, 1996) and is mainly covered by 
Mediterranean coastal maquis, belong- 
ing to the western-Sardinian, calcicole, 
thermomediterranean Chamaerops 
humilis-Juniperetum  turbinatae (Bac- 
CHETTA ET AL., 2009; BIONDI, FILIGHEDDU 
AND FARRIS, 2001; BLASI, ANGIUS AND 
BACCHETTA, 2009). Mediterranean 
coastal maquis alternates with rural 
landscape principally represented by 
grazing fields and cultivated lands. 


DE MATTIA AND MascIa: Otala punctata (Stylommatophora: Helicidae) in Italy 


l0 mm 


Figures. 9-11. Shells of Otala spp. 9-10, Otala punctata (O.F. Miller, 1774). 9: Sardinia, Alghero, 
Sorgente Lu Cantar; 10: Spain, Málaga, El Tarajal; 11: Otala lactea (Miller, 1774), Spain, 


Menorca, Mahon. 


Figures. 9-11. Shells of Otala spp. 9-10, Otala punctata (O.F Miller, 1774). 9: Cerdeña, Alghero, 
Sorgente Lu Cantar; 10: España, Málaga, El Tarajal; 11: Otala lactea (Miller, 1774), España, 


Menorca, Mabón. 


DISCUSSION 


CARRADA ET AL. (1967) stressed the 
presence of fossil shells of O. punctata in 
travertines from Alghero but this has not 
been confirmed. Preliminary field research 
revealed that in the surroundings of 
Alghero, quaternary deposits do not show 
the presence of O. punctata. BALDINO, 
CARENTI, GRASSI, ORGOLESU, SECCHI AND 
WILKENS (2008) and WILKENS (2004) did 
not cite the species from the archaeologi- 
cal sites of north-western Sardinia. Also 
PAULUCCI (1882) did no mentions for O. 
punctata in Sardinia. 


Most probably O. punctata is not 
autochthonous to Sardinia. The intro- 
duction of O. punctata could be referred 
to the Aragonese occupation during the 
14th. The traditional local denomination 
of this species, which is “Caragol 
español” (Spanish snail), could support 
the hypothesis. Nowadays snail farming, 
is a common practice in Sardinia, but 
not in the surroundings of Alghero. In 
the island O. punctata and O. lactea were 
commonly used for heliciculture in the 
past, but less frequently at present. 

The population of O. punctata shows 
a very low density of specimens which 


A5 


Iberus, 29 (1), 2011 


could be referred to its alien origin. Prob- 
ably a feeding competition occurs with the 
native syntopic Helicidae species, such as 
Cantareus apertus (Born, 1778), Cornu asper- 
sum (O.F. Miller, 1774) and Eobania ver- 
miculata (O.F. Múller, 1774). Moreover, 
because of its culinary interest, the species 
is frequently collected by people. The very 
limited distribution of O. punctatais a clear 
risk factor. Destruction of habitat by city 
expansion, mostly related to tourism facil- 
ity development and collecting as a food 
item represent the main threats. Further 


BIBLIOGRAPHY 


BACCHETTA G., BAGELLA S., BIONDI E., FARRIS 
E., FILIGHEDDU R.S. AND MOSSA L. 2009. Ve- 
getazione forestale e serie di vegetazione de- 
lla Sardegna (con rappresentazione carto- 
grafica alla scala 1:350.000). Fitosociologia, 46, 
suppl. 1(1): 3-82. 

BALDINO B., CARENTI G., GRASSI E., ORGOLESU T., 
SECCHI F. AND WILKENS B. 2008. L'economia 
animale dal Medioevo all'Etá moderna nella 
Sardegna nord occidentale. Sardinia, Corsica 
et Baleares Antiquae, 6: 103-155. 

BARBARA N. AND SCHEMBRI P.J. 2008. The sta- 
tus of Otala punctata (Múller, 1774), a recently 
established terrestrial gastropod in Malta. 
Bollettino Malacologico, 44(5-8): 101-107. 

BARCA S., CARMIGNANI L., OGGIANO G., PER- 
TUSAT I P.C., SALVADORI l., CONTI P., ELTRU- 
DIS A., FUNEDDA A. AND PASCIS. 1996. Carta 
geologica della Sardegna (scala 1:200.000). Ser- 
vizio Geologico d'Italia, Firenze. 

BIONDI E., FILIGHEDDU R. AND FARRIS E. 2001. 
Il paesaggio vegetale della Nurra. Fitosocio- 
logia, 382), suppl. 2: 3-105. 

BLASI C., ANGIUS R. AND BACCHETTA G. 20009. 
Clima e Bioclima. /n: Bacchetta G., Bagella S., 
Biondi E., Farris E., Filigheddu R., Mossa L. 
2009. Vegetazione forestale e serie di vege- 
tazione della Sardegna (con rappresenta- 
zione cartografica alla scala 1:350.000). Fito- 
sociología, 46(1), suppl. 1: 3-82. 

BODON M,, FAVILLI L., GIUSTI F. AND MANGA- 
NELLI G. 1995. Gastropoda pulmonata. In: 
Minelli A. Ruffo S. €: La Posta S. (Ed.), Chec- 
klist delle specie della fauna d'Italia, 16: 60 pp. 

CARRADA G., PARISI V. AND SACCHI C.F. 1967. 
Dati per una biogeografia dei molluschi con- 
tinentali in Sardegna. Atti Societa Italiana 
Scienze Naturali Museo Civico Storia Naturale, 
Milano, 105: 377-388. 

CLANZIG S. AND BERTRAND A. 2001. Otala pun- 
ctata (O.F. Miller, 1774) en France. Docu- 
ments Malacologiques, 2: 47-48. 


46 


investigations are actually in progress in 
order to clarify the origin of the Sardinian 
population of O. punctata. 


AKNOWLEDGEMENTS 


The Authors are grateful to N. 
Barbara, P. Schembri (Zabbar, Malta) 
and J.S. Torres (Málaga, Spain) for pro- 
viding alcohol-preserved material and 
E. Zallot (Monster, Holland) for advice 
and critical comments. 


CowIE R.H., DILLON R.T. JR, ROBINSON D.G. 
AND SMITH J.W. 2009. Alien non-marine snails 
and slugs of priority quarantine importance 
in the United States: A preliminary risk as- 
sessment. American Malacological Bulletin, 
27(1-2):113-132. 

FALKNER G. 1990. Binnenmollusken. In: R. 
Fechter €: G. Falkner, Weichtiere, 112-280. 
Munchen. 

FALKNER G., RIPKEN T.E.J. AND FALKNER M. 
2002. Mollusques continentaux de France. Liste 
de référence annotée et bibliographie. Muséum 
national d'Histoire naturelle, Patrimoines 
naturels 52, Paris, 350 pp. 

Giusti F. 1976. 1 Molluschi terrestri, salmastri 
e di acqua dolce dell'Elba, Giannutri e sco- 
gli minori dell'Arcipelago toscano. Lavori, 
Societá Italiana di Biogeografía, Siena, 5 (1974): 
99-355. 

MALATESTA A. AND SETTEPASSI F. 1954. Risul- 
tati del rilevamento del foglio 192 (Alghero- 
Isola di Sardegna). III. Fossili delle formazioni 
continentali quaternarie. Bollettino Servizio 
Geologico Italaliano, 76: 33-39. 

HERBERT D.G. AND SIRGEL W.F. 2001. The recent 
introduction of two potentially pestiferous 
alien snails into South Africa and the out- 
comes of different pest management prac- 
tices: an eradication and a colonization re- 
search in action. South African Journal of 
Science, 97: 301-304. 

PAULUCCI M. 1882. Note malacologiche sulla 
fauna terrestre e fluviale dell'isola di Sar- 
degna. Bullettino della Societa Malacologica Ita- 
liana, 8: 139-381. 

PAULUCCI M. 1886. Conchiglie terrestri e d'acqua 
dolce del M. Argentario e delle isole circos- 
tanti. Bullettino della Societa malacologica Ita- 
liana, 12: 5-64, pl. 1, 2. 

WILKENS B. 2004. La fauna sarda durante 
l'Olocene: le conoscenze attuali. Sardinia, Cor- 
sica et Baleares Antiquae, 1 (2003): 181-197. 


O Sociedad Española de Malacología —_—_—_—_———  Iberus, 29 (1): 47-57, 2011 


Reproduction of Donax trunculus in the littoral of Huelva 
(southern Atlantic Spain): is there any difference with the 
Mediterranean population from the Andalusian coast? 


Reproducción de Donax trunculus en el litoral de Huelva (suroeste 
Atlántico de España): ¿hay diferencias con la población mediterránea 
de la costa andaluza? 


Cristina TIRADO, José Luis RUEDA and Carmen SALAS* 


Recibido el 13-IV-2011. Aceptado el 11-V-2011 


ABSTRACT 


The reproductive cycle of D. trunculus L., 1758 was studied using histology and changes 
in flesh dry weight, in the littoral of Huelva (southern Atlantic Spain) from June 1990 to 
May 1991. The spawning period is essentially synchronic, and extends from February to 
August, with different individual intensity. Three peaks of spawning have been recorded, 
April, June and July. The resting period ranges from September to December, with the 
whole population in cytolized stage in October. The favourable environmental variables, 
such as high levels of chlorophyll a and relatively mild seawater temperatures allow this 
extensive reproductive period. This cycle shows a diphase of the spawning period with the 
Mediterranean population from Málaga, which should imply a different close season for 
each population. 


RESUMEN 


Se estudia el ciclo reproductor de D. trunculus L., 1758 mediante histología y cambios en 
la biomasa, en el litoral de Huelva (sur de España Atlántico) entre junio de 1990 y mayo 
de 1991. El periodo de puesta es básicamente sincrónico y se extiende desde febrero 
hasta agosto, con intensidad variable según los individuos. Se han registrado tres picos 
de puesta: abril, junio y julio. El periodo de reposo ocurre entre septiembre y diciembre, 
con toda la población en la fase citolítica en octubre. Las variables ambientales favora- 
bles, tales como los altos niveles de clorofila a y las temperaturas relativamente templadas 
del agua permiten un periodo reproductor largo. Este ciclo muestra un desfase del perí- 
odo de puesta con relación a la población mediterránea de Málaga, por lo que se reco- 
mienda un periodo de veda diferente para cada población. 


INTRODUCTION 

The overexploitation, for many the fisheries. Until 2003, the law in 
years, of some marketable shellfish Andalusia (autonomous region inclu- 
species in southern Spain has mainly ding eight southern provinces of Spain) 
been due to an incorrect management of ruled an extensive close season (over six 


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


47 


Iberus, 29 (1), 2011 


months) for most of the marketable 
bivalves of this area. Because of the 
incompatibility of these extensive 
periods with a profitable economic acti- 
vity, it was not enforced. The study of 
the reproduction of the populations of 
Donacidae in the Mediterranean littoral 
of Málaga (TIRADO AND SALAS, 1998, 
1999) pointed out the possibility of esta- 
blishing an effective rule with a reduced 
close season that would still be compati- 
ble with profitable fishery activities. 
These results led regional fishery autho- 
rities to promote a research project for 
the study of reproductive cycles of the 
most important shellfish, aimed at 
adjusting the close season to the biology 
of each species and consequently to 
improve the management of these fishe- 
ries (TIRADO AND RODRIGUEZ DE LA RUA, 
2000). 

One of the studied species was 
Donax trunculus L. 1785 (“coquina”) 
with a very important consumer 
market, mostly in the province of 
Huelva (southwest coast). The repro- 
duction of this species was studied in 
the Mediterranean littoral of Andalusia 
(TIRADO AND SALAS, 1998), but it is well 
known that reproduction in bivalves is 
strongly related to environmental varia- 
bles, such as temperature, availability of 
food or type of beach, among others 
(BAYNE 1976; BROWN AND MCLAHLAN, 
2006). The environmental characteristics 
of the Atlantic shallow littoral are very 
different from those of the Mediterra- 
nean. The fishing methods are also diffe- 
rent, in the Mediterranean littoral the 
fishery is made by authorized boats, 
whereas in the Atlantic littoral the fis- 
hermen use individual dredges drawn 
backwards in shallow water. For these 
reasons, the study of the Donax truncu- 
lus population from the Atlantic littoral 
of Huelva was undertaken. 

Donax trunculus is a littoral species 
widely distributed from Brittany 
(French coast) (Lucas, 1965; ANSELL 
AND LAGARDERE, 1980; GUILLOU AND LE 
MOAL, 1980) to Southern Morocco 
(PASTEUR HUMBERT, 1962) and in the 
Mediterranean (SABELLI, GIANUZZI- 
SAVELLI AND BEDULLI, 1990). As a com- 


48 


mercial species, D. trunculus has been 
intensively studied for fisheries mana- 
gement (BALDACCINI AND BIANUCCI, 
1984, FISCHER, BAUCHOT AND SCHENEI- 
DER, 1987), parasitism and predation 
(RAMÓN, GRACENEA AND GONZÁLEZ- 
MORENO, 1999, SALAS, TIRADO AND 
MANJÓN-CABEZA, 2001), population 
dynamics (MOUÉEZA AND CHESSEL, 1976; 
GUILLOU AND LE MOAL, 1980; BAYED 
AND GUILLOU 1985; MAZÉ AND LABORDA 
1988; NEUBERGER-CYWIAK, ACHITUV 
AND MIZRAHI, 1990; ZEICHEN, ÁGNESI, 
MARIANI, MACCARONI AND ARDIZZONE, 
2002) and growth and reproduction at 
different places of the distribution area 
(Lucas, 1965; BADINO AND MARCHIONI 
1972; MOUEZA AND FRENKIEL-RENAULT, 
1973; BAYED 1990; NEUBERGER-CYWIAK 
ET AL. 1990; TIRADO AND SALAS 1998; 
GASPAR, FERREIRA AND MONTEIRO, 1999; 
DevaL 2009). All this research has revea- 
led a biological and physiological varia- 
bility of the populations in relation to 
environmental factors. Therefore, the 
aim of this paper is to study the repro- 
ductive cycle of D. trunculus from 
Huelva, and to point out the differences 
between the Atlantic and the Mediterra- 
nean populations in order to request the 
authorities to take into account the 
biology of the species in a particular 
area for a better management of the 
resources. 


MATERIAL AND METHODS 


The sampling area is located in the 
littoral of Doñana (Southwest Spain) 
(367 52 N - 6” 26'W) (Fig. 1), in an exten- 
sive beach, more than 20 km long and 
100-200 m wide, with a tidal range 
which can reach 3.6 m in spring tides. 
The beach is of fine grain and gentle 
slope that can be considered as a dissi- 
pative beach (SCHLACHER, SCHOEMAN, 
DUGAN, LASTRA AND JONES, 2008). The 
samples were collected, on a sandy 
bottom at 0.4 m depth, from June 1999 
to May 2000, with monthly frequency 
from October to February and twice a 
month during the spring and summer 
months. The specimens were captured 


TIRADO £7 AL.: Reproduction of Donax trunculus in the littoral of Huelva 


W06%50'  06%0' 0620" 


d 36030 
06%10' 2 


1 37%N 


13650" 


3640' 


Figure 1. Sampling area. Figure 2. Gear used for commercial collecting of Donax trunculus. 
Figura 1. Área de muestreo. Figura 2. Rastro usado para la pesca de Donax trunculus. 


with a dredge of 45 cm width, with 9 cm 
long teeth. The mesh was 1.75 cm, usual 
among the fishermen of the area (Fig 2). 

A total of 4333 specimens of D. trun- 
culus, between 13 and 44 mm in length, 
were examined; of these 3798 were used 
for the analysis of the flesh dry weight 
variation (about 200 individuals /sample) 
and 535 specimens for histological study 
(usually 30 per month). 

The length (L) of every specimen 
was measured to the nearest millimetre, 
and the soft parts were dried in an oven 
at 100 *C for 24 h, and weighed to the 
nearest milligram (flesh dry weight, 
FDW). Two different indices of condi- 
tion were applied, FDW/L? variation, 
and that proposed by Crosby and Gale 
(1990) as FDW*1000/volume of the 
internal cavity of the shell (considering 
the millilitres of water as milligrams) 
which is referred to as CI. The regres- 
sion of flesh dry weight on the length 
was calculated for each sample to esti- 
mate the variation in biomass of a stan- 
dard individual, based on the logarith- 
mic transformation of Ricker”s function 
W= aL? (RICKER, 1975), where W is the 
weight, L is the length, a is the ordinate 
at origin, and b is the slope. To minimize 
the bias introduced by the somatic 
growth of individuals during the cycle 


and by the variation of the size of the 
specimens between successive samples, 
the variation of flesh dry weight was 
estimated for an individual of 29 mm 
length (mean size of the population 
studied). For that we took into account 
the regression lines for every sample. 

For histological processing, speci- 
mens were anaesthetized with MgClz 
fixed in 10% formaldehyde, embedded 
in paraffin, sectioned at 10 um and 
stained with haematoxylin of Carazzi 
and eosin, and a trichromic staining 
(VOF according to GUTIÉRREZ (1967)) of 
haematoxylin of Carazzi, light green, 
orange G and acid fuchsine. The stages 
of development of the gonad were 
scored according to the scale proposed 
by DE VILLIERS (1975) for D. serra 
Róding, 1798 in South Africa, who pro- 
posed five stages: 

Cytolized. The alveoli are very small 
and wide apart. Some clams can be 
sexed when a few gametes are present. 

Preactive. The alveoli have clearly 
defined alveolar walls. They are inter- 
sected by broad, continuous transverse 
fascicles. Most clams can be sexed. 

Active. The alveoli are large and 
usually adjacent. The alveolar walls are 
always complete. Germ cells in various 
stages of development fill the alveoli 


49 


Iberus, 29 (1), 2011 


and are both actively increasing and 
enlarging. 

Spawning. The alveolar pattern is 
disturbed and the alveolar walls are 
often broken. The alveoli are often flatte- 
ned and show an orientation towards 
the centre of the gonad. 

Postactive. The amount of germ cells 
varies, depending on the intensity of 
spawning and the time that has elapsed 
since spawning took place. Phagocitic 
cells are common. 

The species showed a sexual differen- 
tiation during the time of sexual activity. 
During the active period, the ovaries of 
D. trunculus are dark blue whereas the 
testes have a viscous aspects and a 
whitish-orange colour, which makes pos- 
sible to identify the sex macroscopically 
of most of the specimens. 

To evaluate the possible influence of 
environmental factors on the cycle, the 
temperature of sea water at the surface 
was measured simultaneously with the 
collection of the individuals. Samples of 
water (11) were taken for determination 
of chlorophyll a. Pigment analyses were 
carried out by filtering the water 
through Whatman GF/C glass filters. 
The pigments of the retained cells were 
then extracted with acetone for 12 h in 
cool, dark conditions, following the 
recommendations of LORENZEN AND 
JEFFREY (1980). Concentrations of chlo- 
rophyll 4 were calculated using the 
trichromatic equations of JEFFREY AND 
HUMPHREY (1975). 

The test of Kolmogorov-Smirnov and 
Kendal and Pearson's rank correlations 
included in the program SPSS 8.0, were 
used to check the distribution of the data. 
Cross correlation between both condition 
indexes and percentage of spawning 
with temperature and chlorophyll a 
levels were calculated to asses their 
influence on the reproductive cycle. 


RESULTS 


Environmental factors 

The monthly temperature data 
showed important fluctuations (Fig. 3), 
with the maximum value in the second 


50 


half of September (23 *C) and the 
minimum (12.5 “C) in December. The 
most important decrease was recorded 
from October to November (9.5 *C), and 
conversely, the highest increase (4.5 *C) 
was observed from February to the first 
half of March. 

The chlorophyll a levels were higher 
than 2 ug/l throughout the cycle, 
showing an irregular pattern, with two 
important peaks, in September (24.3 
ug /1) and March (16.1 ug/)). 


Sex ratio 

The sex ratio was determined on 
specimens with shell length ranging 
from 13 to 44 mm. In the samples from 
December to the end of July it is possi- 
ble to identify the sex of the whole 
population according to the colour and 
external aspect of the gonad. A total of 
2564 specimens were examined for sex- 
ratio study (all the specimens from his- 
tology and those of biomass study from 
December to July), of which 1268 were 
males (49.45%) and 1296 females 
(50.55%) (Fig. 4). The sex ratio for all 
them can be considered as 1:1 (x2= 0.7, 
P>0.95). The smallest specimen exami- 
ned with gonad differentiated was a 
female of 13.8 mm. 


Sexual cycle 

Flesh dry weight: The variation of 
flesh dry weight —length ratio (FDW /L?) 
during the annual cycle is shown in 
Figure 5. The mean monthly values of 
both variables, flesh dry weight (FDW) 
and size (L?) were considered in 3798 
specimens. The standard deviations 
range between 10 and 21%, being larger 
than the standard deviations observed 
in the monthly mean size (between 6 
and 15%) (Fig. 6) There is also a broad 
weight range in most of the samples 
(Fig. 7), with standard deviation 
between 23 and 45%. 

According to flesh dry 
weight-length ratio (FDW/L?), the 
population showed low values during, 
summer and autumn, with an increase 
from December to February. From 
February, the flesh dry weight- length 
ratio decreases until summer, with a 


TIRADO ET AL.: Reproduction of Donax trunculus in the littoral of Huelva 


Ta AL AD: SE 520, N  D. JJ]. FE MrlMr.Apl Ap2 Myl My2 


Figure 3. Sea water temperature and concentration of chlorophyll 4 in sea water at the sampling 
site through the year. 

Figura 3. Temperatura del agua de mar y concentración de clorofila a en el agua del mar en el sitio de 
muestreo durante todo el año. 


a %EF O %M 


A A Ll D J FE — Mrl Mr2 Apl Ap2 Myl My2 


Figure 4. Relative frequency (%) of sexes during the year of study. F: females. M: males. 
Figura 4. Frecuencia relativa (%) de los sexos durante el año de estudio. F: hembras. M: machos. 


FDW/T? 
E 


Jaija2 JJ AL c0A22 81 520 N.D. E MriMr2Apl Ap2 My1My2 


Figure 5. Flesh dry weight (FDW)/ Length (L?) ratio during the year of study. Bars show standard 
deviation. 
Figure 5. Relación de peso seco de biomasa (EDW)/ Longitud (L?) durante el año de estudio. Las barras 


muestran la desviación estándar. 


dl 


Iberus, 29 (1), 2011 


Lm (mm) 


A ME E 0 ASI SO AN O E Mrl Mr2 Apl Ap2 Myl My2 


Figure 6. Monthly mean length of shells (L) during the year of study. Bars show standard devia- 
tion. 

Figura 6. Media mensual de la longitud de las conchas (L) durante el año de estudio. Las barras mues- 
tran la desviación estándar. 


Jnl Jn2 Jl J2 Al A2 S1 S2 O N D Jl F Mrl Mz Apl Ap2 Myl My2 


Figure 7. Monthly mean flesh dry weight (DW) during the year of study. Bars show standard 
deviation. 

Figura 7. Media mensual del peso seco de biomasa (DW) durante el año de estudio. Las barras muestran 
la desviación estándar. 


Cl 


Jnl Jn2 Jl J2 Al A2 SI S2 O N D J  F MrlMr2Apl Ap2 Myl My2 


Figure 8. Index of condition of Crosby and Gale (CI): flesh dry weight x 1,000 / volume of the 
internal cavity of the shell, during the year of study. 
Figura $. Índice de condición de Crosby y Gale (CD): Peso seco de la biomasa x 1000 / volumen de la 


cavidad interna de la concha, durante el año de estudio. 


DL 


TIRADO ET ALz.: Reproduction of Donax trunculus in the littoral of Huelva 


Table I. Linear regression calculated for each sample. Lm: mean length; flesh dry weight for a stan- 
dard individual; R?: coefficient of determination; R: coefficient of correlation; N: number of speci- 
mens; W: monthly flesh dry weight for a standard individual of 29 mm. 

Tabla [. Regressión linear calculada para cada muestra. Lm: longitud media; R': coeficiente de deter- 
minación; L: coeficiente de correlación; N: número de individuos; W: peso seco de biomasa mensual para 


un individuo estándar de 29 mm. 


lm Regression lines 

Jn] 3127 y=2.3423 x-1.1468 
Jn2 31 y= 2.2581x- 1.0263 
1 28.15 y=2.3057 x-1.201 

12 21.31 y=2.0405 x-0.82724 
Al 26.21 y =2.3454x -1.2257 
A2 31:19 y=2.2426x-1.0357 
Sl 34.57 y=2.5491 -1.5168 

92 MS y=2.3769 x-1.2753 
0 27.81 y=2.6152x-1.6298 

N 250] y=2./123 x-1.6903 
D 25.40 y=2.701 x- 1.7035 
| 21.58 y = 2.8042 - 1.7363 
F 28.09 y =2.8/14x- 1.7802 
Mr 29.49 y=2:605X- 1.3919 
Mr2 28.62 Y =2.0323%- 11993 
Apl 2452 y =2.785x 1.7727 

Ap2 29.43 y =2.3754x-1.2251 
My] 31.09 y =2.4862x-1.3195 
My2 29.54 y = 2.4818x- 1.3567 


R R N W(L=29 mm) 
0.7171 0.8468 200 66.78 
0.8105 0.2003 199 64.46 
0.8062 0.8979 198 67.15 
0.7073 0.8410 200 58.35 
0.8452 0.9193 199 66.79 
0.7991 0.8939 197 64.00 
0.4281 0.6543 202 12.41 
0.2006 0.9490 201 67.65 
0.8392 0.9161 199 14.21 
0.8741 0.9349 200 16.97 
0.8329 0.9126 200 16.62 
0.9229 0.9607 200 19.58 
0.9071 0.9524 200 81.49 
0.8814 0.9388 199 14.15 
0.8816 0.9389 200 80.35 
0.862 0.9284 199 18,79 
0.1727 0.8790 200 67.66 
0.7618 0.8728 202 10.78 
0.8038 0.8965 199 10.62 


small peak in May. The CI index 
showed a similar pattern (Fig. 8). The 
most important decrease was recorded 
from March to April. 

The monthly regression lines for 
weight-length relationship are shown in 
Table L, and the monthly variation of 
flesh dry weight for a standard indivi- 
dual (W) is represented in Figure 9. 
According to these data the population 
from Huelva had a relatively synchro- 
nous reproductive pattern, with 18 
samples (from a total of 19) explaining 
70% of the variations of weight by the 
length. The pattern of the standard indi- 
vidual was similar to that of Cl index. 

According to the test of Kolmogorov 
Smirnov, only the temperature and 
FDW /L* data showed a normal distri- 
bution. The coefficients of correlation of 
Pearson and Kendall have been calcula- 
ted. According to these data, the tempe- 
rature is inversely correlated with 


FDW /L? (r=0.541p<0.05) and with the 
FDW of the standard individual 
(tau=0.438, p<0.01). No other correla- 
tions were significant. 


Gametogenic cycle 

The data from the histological study 
are presented in Figure 10. Activation of 
gonads started in January while the 
regression began in the second half of 
August with more than 66% of the 
population in postactive and cytolized 
stages. In October, the whole population 
was in cytolized stage. 

During the active period, the histolo- 
gical data showed continuous spaw- 
ning, with percentages higher than 55% 
from February to July with three peaks: 
first half of April (96.67%), June (100%) 
and second half of July (96.67%), while 
only 13% of the individuals at the end of 
August were spawning. Different stages 
in the same gonad and new activation 


IÍ 


Iberus, 29 (1), 2011 


85 
80 
o 
5h 70 
E 6 
= 60 
A ss 
50 
45 
40 


Ja A AO SS IO NADIA 


F- Mrl Mr2 Ap1 Ap2 My1My2 


Figure 9. Variation in flesh dry weight in a standard Donax trunculus animal 29 mm long. 
Figura 9. Variación del peso seco biomasa en un animal estándar de Donax trunculus de 29 mm de 


longitud. 


A A E E e Y 
AI A 
A A AS 
A E A es 
A O 
A E 
A A 


A O 
SU 


% 


AS SAO NO E 


2 ha hd 


E Mrl Mr2 Apl Aps Myl My2 


Figure 10. Monthly cumulative frequency of different stages of development of the gonads in D. 
trunculus. C: cycolized; Pr: preactive; EA: early active; A: active; S: spawning; Ps: postactive. 

Figura 10. Frecuencia mensual acumulada de los diferentes estados de desarrollo de las gónadas en D. 
trunculus. C: cicolizado; Pr: preactivo; EA: activo temprano; A: activo; S: en puesta; Ps: postactivo. 


without a total regression of the gonad 
have been observed in many indivi- 
duals. 

The resting period started in Septem- 
ber and finished in December. At the begin- 
ning of this period, it was impossible to 
identify the sex of 36% of the individuals, 
and in October all the individuals were in 
cytolized stage and it was impossible to 
identify the sex of 50% of the sample. 


DISCUSSION 


Sex ratio 

The sex-ratio of the population of 
Donax trunculus from Huelva agrees 
with observations of other authors 
regarding this species (Lucas, 1965; 


54 


MOUEZA AND FRENKIEL-RENAULT 1973, 
TIRADO AND SALAS, 1998, DEVAL, 2009, 
La VALLE, 2006), but disagrees with the 
results obtained in the neighbour popu- 
lation of southern Portugal, where 
GASPAR ET AL. (1999) found a higher pro- 
portion of males in all size classes. 

According to the macroscopic identi- 
fication of the gonads, the active period is 
more extensive in the Atlantic population, 
in which it is possible to identify the sex 
of the whole population from February to 
the end of July (Fig 3), while in the Medi- 
terranean population, the gonads are 
coloured in the whole sample only in May 
and June. Percentages of specimens with 
non coloured gonads higher than 70% 
were observed from October to February 
(TIRADO AND SALAS, 1998) 


TIRADO ET AL.: Reproduction of Donax trunculus in the littoral of Huelva 


Sexual cycle 

Condition index: The wide range of 
the standard deviation was partially 
related to the presence of different 
stages of development of the gonads. 
The data for the different condition 
indexes of this Atlantic population 
showed a long reproductive period, 
from December to September, very 
similar to those found by BAYED (1990) 
in the Atlantic coast of Morocco and by 
GASPAR ET AL. 1999 in the South of Por- 
tugal, although in the latter there was a 
continuous decrease of the index of con- 
dition from February to August. 

However, there are differences with 
the Mediterranean population of 
Málaga, in which the most important 
and continuous decrease of the index of 
condition was observed from April to 
June, with an important increase in July. 
In the population from Huelva, the 
major decrease was recorded from 
March to April, with a slight increase in 
May, but with continuous decrease 
during the summer. 

In Turkey, the population studied by 
DeEvaL (2000) had a shorter reproductive 
period; the spawning occurred from 
April to July with a peak between May 
and June. This reproductive cycle is 
coincident with that of the population of 
Málaga. The duration of the reproduc- 
tive period of the Turkish and Málaga 
populations was similar to that found in 
other Mediterranean populations 
(ANSELL AND BODOY 1979 in Camargue, 
French Mediterranean coast; MÓUEZA 
AND  [FRENKIEL-RENAULT 1973 in 
Algeria), although these were not coinci- 
dent in time, due to differences in the 
environmental variables. 

Gametogenic cycle: The data obtained 
point to continuous spawning for 
almost the whole population from April 
to August, coincident with the decrease 
of the index of condition during this 
period. The coexistence of different 
stages in the same gonad, together with 
a direct transformation from a postac- 
tive stage to an active one without the 
intermediate step of a cytolized stage, 
have been reported for other donacids 
(DE VILLIERs, 1975, for D. serra; TIRADO 


AND SALAS 1998 for D. trunculus in 
Málaga; GASPAR ET AL. 1999 in D. truncu- 
lus in Faro (Portugal) Tirado and Salas, 
1999 for D. venustus and Donax semistria- 
tus), or other species, such as Tapes rhom- 
boides (see MORVAN AND ANSELL, 1988), 
Callista chione (see TIRADO, SALAS AND 
LóPEz, 2002) and Venus verrucosa (see 
TIRADO, SALAS AND MÁRQUEZ, 2003). 
This renewed activation of the gonad 
seems to be at the origin of the fluctua- 
tions of the indexes of condition during 
this period. After the most important 
release of gametes at the beginning of 
spring, there was new but less intense 
spawning by the same individuals in 
the same cycle. The occurrence of suc- 
cessive individual spawnings has also 
been described in the population of the 
Adriatic Sea (ZEICHEN ET AL. 2002). 

The most important drop of the 
indexes of condition occurred between 
March and April, with 100% of the 
population in spawning, and after an 
important increase of chlorophyll a in 
March. The strong regression of the 
gonad from September to October 
(100% of the population in postactive or 
cytolized stage) could be related to the 
strong decrease of the temperature 
(9.5%C). An inverse correlation has been 
obtained between temperature and flesh 
dry weight (FDW/L and FDW of a stan- 
dard individual). The availability of 
food in September at the same time as a 
maximum of clorophyll 4 could be 
advantageous for the storage of reser- 
ves, after an extensive spawning period. 
This result is also described in the Alge- 
rian population (MOUEZA AND FREN- 
KIEL-RENAULT, 1973). On the other hand, 
the increase of chlorophyll a in March 
could be advantageous for the larva 
after the highest peak of spawning of 
the population from March to April. 

According to the histological data, 
the Moroccan population of D. trunculus 
have three months of resting period 
(BAYED, 1990), whereas in the popula- 
tion of Huelva there is only one month 
(October). Moreover, the percentages of 
spawning in the population of Huelva 
are around 100%, whereas in the popu- 
lation of Morocco the percentages never 


SS 


Iberus, 29 (1), 2011 


reach 50%. The data of the population 
from Faro (Portugal) are more similar to 
those found in the population of 
Huelva, although the whole population 
is never in spawning (maximum 80% in 
August, GASPAR ET AL., 1999). 
According to SCHLACHER ET AL. 
(2008) and BROWN AND MCLACHLAN 
(2006) the populations from dissipative 
beaches have a more extensive repro- 
ductive cycle than those from reflective 
ones. However, our data on the repro- 
duction of D. trunculus from the littoral 
of Málaga, collected in  reflective 
beaches (TIRADO AND SALaAs, 1998) 


BIBLIOGRAPHY 


ANSELL A.D. AND BODOY A. 1979. Comparison 
of events in the seasonal cycle for Donax vit- 
tatus and Donax trunculus. In Naylor, E. and 
R. G. Hartnoll (Ed.): Cyclic phenomena in ma- 
rine plants and animals. Proceedings of the 
13th European Marine Biology Symposium. 
Pergamon Press, Oxford and New York: 191- 
198. 

ANSELL A.D. AND LAGARDERE F. 1980. Obser- 
vations on the biology of Donax trunculus 
and Donax vittatus at lle d'Oléron (French 
Atlantic coast). Marine Biology, 57: 287-300. 

BADINO G. AND MARCHIONNI V. 1972. Neu- 
rosecretion and gonad maturation in a pop- 
ulation of Donax trunculus L. from Leghorn 
(Italy). Bollettino di Zoologia, 39: 321-326. 

BALDACCINI G. AND BIANUCCI P. 1984. Contri- 
buto alla conoscenza di Donax trunculus (Bi- 
valvia) del litorale versiliese: aspetti tecno- 
logici, commerciali e statistici della raccolta. 
Nova Thalassia, 6: 441-449. 

BAYED A. 1990. Reproduction de Donax trun- 
culus sur la cóte Atlantique marocaine. Cahiers 
de Biologie Marine, 31: 159-169. 

BAYED A. AND GUILLOU J. 1985. Contribution á 
l'étude des populations du genre Donax: la 
population de D. trunculus L. (Mollusca, Bi- 
valvia) de Mehdia (Maroc). Annales de 
l'Institut Océanographique, Paris, 61 (2): 139- 
147. 

BAYNE B.L. 1976. Marine mussels, their ecology and 
physiology. International Biological Pro- 
gramme. Ed. Cambridge University Press, 
441 pp. 

BROWN A.C. AND MCLAHLAN A. 2006. Ecology 
of sandy beaches (2nd). Elsevier, Amsterdam: 
SIP 


5Ó 


showed a similar extension of the repro- 
ductive cycle to that of the population of 
Huelva, which lives in a dissipative 
beach, but with an offset of three 
months between the beginning of the 
spawning period in each population. 
These results justify the necessity of 
taking into account the biology of the 
species in a particular area for a better 
management of the resources by ruling 
different close seasons for the Atlantic 
and the Mediterranean populations, 
from March to April in the Atlantic 
populations and from May to June in 
the Mediterranean ones. 


CROSBY M.P. AND GALE L. 1990. A review 
and evaluation of bivalve condition index 
methodologies with a suggested standard 
method. Journal of Shellfish Research, 9 (1): 
233-237. 

DE VILLIERS G. 1975. Reproduction of the sand 
mussel Donax serra Róding. Investigational 
Reports of the Sea Fisheries Branch of South 
Africa, 109: 1-31: 

DevaL M.C. 2009. Growth and reproduction 
of the wedge clam Donax trunculus in the sea 
of Marmara, Turkey. Journal of Applied Ichth- 
yology, 25 (5): 551-558. 

FISCHER W., BAUCHOT M.L. AND SCHENEIDER M. 
1987. Fiches FAO d'identification des especes 
pour les besoins de la péche. (Révision 1). Mé- 
diterranée et mer Noire. Zone de péche 37. 
Vol. IVegetaux et Invertébrés, 760 pp. 

GASPAR M.B., FERREIRA R. AND MONTEIRO C.C. 
1999. Growth and reproductive cycle of Do- 
nax trunculus L., (Mollusca:Bivalvia) off Faro, 
southern Portugal. Fisheries Research, 41: 309- 
316. 

GUILLOU J. AND Le MOAL Y. 1980. Aspects de 
la dynamique des populations de Donax vit- 
tatus et D. trunculus en baie de Douarnenez. 
Annales de l'Institut Océanographique, Paris, 
56 (1): 55-64. 

GUTIÉRREZ M. 1967. Coloración histológica para 
el ovario de peces, crustáceos y moluscos. In- 
vestigación Pesquera, 31 (2): 265-271. 

JEFFREY S.W. AND HUMPHREY G.T. 1975. New 
spectrophotometric equation for determi- 
ning chlorophylls a, b, c! and c? in higher 
plants, algae and natural phytoplankton. Bio- 
chemie und Physiologie der Pflanzen, 167: 191- 
194. 


TIRADO ET ALz.: Reproduction of Donax trunculus in the littoral of Huelva 


LA VALLE P. 2006. Donax trunculus (Bivalvia: Do- 
nacidae) quale indicatore biologico degli 
equilibri costieri e del bilancio sedimenta- 
rio. Doctoral thesis, Universita degli studi 
di Roma “La Sapienza”. pp. 1-148, i-xv and 
appendix, 1-44. 

LORENZEN C.J. AND JEFFREY S.W. 1980. Deter- 
mination of chlorophyll in seawater. 
UNESCO Technical Papers in Marine Science, 
35: 1-20. 

LUCcas A. 1965. Recherches sur la sexualité des 
Mollusques Bivalves. Bulletin Biologique de la 
Prance et de la Belgique, 99: 115-247 

MaZÉ R.A. AND LABORDA A.J. 1988. Aspectos 
de la dinámica de población de Donax trun- 
culus (Linnaeus, 1758) (Bivalvia: Donacidae) 
en la ría de El Barquero (Lugo, NO España). 
Investigación Pesquera, 52 (3): 299-312. 

MORVAN C. AND ANSELL A.D. 1988. Stereolo- 
gical methods applied to reproductive cycle 
of Tapes rhomboides. Marine Biology, 97: 355- 
364. 

MOUEZA M. AND FRENKIEL-RENAULT L. 1973. 
Contribution a l'étude de la biologie de Do- 
nax trunculus L. (Mollusques Lamelli- 
branches) dans l'Algérois: la reproduction. 
Cahiers de Biologie Marine, 14 (3): 261-283. 

MOUEZA M. AND CHESSEL D. 1976. Contribution 
a l'étude de la biologie de Donax trunculus L. 
(Mollusque, Lamellibranche) dans 1 Algerois: 
analyse statistique de la dispersion le long 
d'une plage en Baie du Bou-Ismail. Journal of 
Experimental Marine Biology and Ecology, 21: 
211-221. 

NEUBERGER-CYWIAK L., ACHITUV Y. AND MIZ- 
RAHI L. 1990. The ecology of Donax truncu- 
lus Linnaeus and Donax semistriatus Poli from 
the Mediterranean coast of Israel. Journal of 
Experimental Marine Biology and Ecology, 134: 
203-220. 

PASTEUR-HUMBERT C. 1962. Les Mollusques 
marins testacés du Maroc. Vol. II. Les La- 
mellibranches et les Scaphopodes. Travaux de 
l'Institut Scientifique Chérifien, Série Zoolo- 
gie, 28, 188 pp. 

RAMÓN M., GRACENEA M. AND GONZÁLEZ-MO- 
RENO O. 1999. Bacciger bacciger (Trematoda, 
Fellocistomidae) infection in commercial 
clams Donax trunculus (Bivalvia, Donacidae) 
from the sandy beaches of the Western Me- 
diterrranean. Diseases of Aquatic Organisms, 
35: 37-46. 


RICKER W. 1975. Computation and interpreta- 
tion of biological statistics of fish popula- 
tion. Bulletin of the Fisheries Research Board of 
Canada, 191: 1-382. 

SABELLI B, GIANUZZI-SAVELLI R. AND BEDULLI 
D. 1990. Catalogo annotato dei Molluschi marini 
del Mediterraneo. Societa Italiana dei Mala- 
cologia, vol 1, 348 pp. 

SALAS C., TIRADO C. AND MANJÓN-CABEZA M.E. 
2001. Sublethal foot-predation on Donaci- 
dae (Mollusca, Bivalvia). Journal of Sea Re- 
search, 46: 43-56. 

SCHLACHER T.A., SCHOEMAN D.S., DUGAN J., 
LASTRA M. AND JONES A. 2008. Sandy beach 
ecosystems: key features, sampling issues, 
management challenges and climate change 
impacts . Marine Ecology, 29 (Suppl. 1) 70-90 

TIRADO C. AND RODRÍGUEZ DE LA RÚA A. 2000. 
Estudio del ciclo reproductor de los moluscos bi- 
valvos y gasterópodos del litoral andaluz. In- 
forme Técnico. Consejería de Agricultura y 
Pesca, Junta de Andalucía, 357 pp. 

TIRADO C. AND SALAS C. 1998. Reproduction 
and fecundity of Donax trunculus L., 1758 
(Bivalvia: Donacidae) in the littoral of Málaga 
(Southern Spain). Journal of Shellfish Research, 
17 (1): 169-176. 

TIRADO C. ANDSALASC. 1999. Reproduction of 
Donax venustus Poli 1795, Donax semistriatus 
Poli 1795 and intermediate morphotypes (Bi- 
valvia: Donacidae) in the littoral of Málaga 
(Southern Spain). P.S.Z.N.: Marine Ecology, 20 
(2): 111-130. 

TIRADO C., SALAS C. AND LÓPEZ J.I. 2002. Re- 
production of Callista chione (L., 1758) (Bi- 
valvia: Veneridae) in the littoral of Málaga 
(Southern Spain). Journal of Shellfish Research, 
21 (2): 643-648. 

TIRADO C., SALAS C. AND MÁRQUEZ Il. 2003. Re- 
production of Venus verrucosa L., 1758 (Bi- 
valvia: Veneridae) in the littoral of Málaga 
(southern Spain). Fisheries Research, 63: 437- 
445. 

ZEICHEN M.M., AGNESI S., MARIANI A., MAC- 
CARONI A AND ARDIZZONE D. 2002. Biology 
and Population Dynamics of Donax truncu- 
lus L., (Bivalvia: Donacidae) in the South 
Adriatic Coast (Italy). Estuarine; Coastal and 
Shelf Science, 54: 971-982. 


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O Sociedad Española de Malacología 


Iberus, 29 (1):59-74, 2011 


Bittium nanum (Gastropoda, Cerithiidae), una especie 


válida de las islas Azores 


Bittium nanum (Gastropoda, Cerithiidae), a valid species from the 


Azores Islands 
Diego MORENO* 
Recibido el 20-IV-2011. Aceptado el 19-V-2011 
RESUMEN 


Se hace una redescripción de la especie del género Bittium Gray, 1847 que constituye 
uno de los elementos más abundantes del zoobentos de la zona litoral rocosa en las islas 
Azores. La especie ha sido confundida por casi todos los autores con las especies euro- 
peas B. reticulatum (da Costa, 1778) o B. latreillii (Payraudeau, 1826) pero se diferencia 
suficientemente para considerarse un endemismo del archipiélago. Se propone usar para 
ella el binomen Bittium nanum (Mayer, 1864), un nombre olvidado pero nomenclatural- 
mente válido y basado en fósiles de una terraza de edad Pleistoceno en Prainha, isla de 
Santa María. Este (introducido originalmente en el binomen Cerithiopsis nana Mayer, 
1864) es un homónimo primario de Cerithiopsis tubercularis var. nana Jeffreys, 1867, por 
lo cual esta última especie debería de ser renombrada o redescrita. 


ABSTRACT 


A redescription is given for the species of the genus Bittium Gray, 1847 which is one of 
the most abundant components of the zoobenthos on rocky shores of the Azores. The 
species has been confused by almost all authors with the European species B. reticulatum 
(da Costa, 1778) or B. latreillii (Payraudeau, 1826) but differs enough to be considered a 
species endemic to the archipelago. lt is proposed to use for this species the binomen Bit 
tium nanum (Mayer, 1864), a forgotten but nomenclaturally valid name based on fossils of 
a terrace of Pleistocene age in Prainha, Island of Santa Maria. This name (first introduced 
in the binomen Cerithiopsis nana Mayer, 1864) is a primary homonym of Cerithiopsis 
tubercularis var. nana Jeffreys, 1867, which should be renamed or better described anew. 


INTRODUCCIÓN 


En las islas Azores vive una especie 
del género Bittium Gray, 1847 que cons- 
tituye uno de los elementos más abun- 
dantes del zoobentos de la zona litoral 
rocosa. Sin embargo, no existe un con- 
senso acerca del estatus taxonómico de 
esta especie, la cual ha sido general- 
mente confundida con otras similares 
que viven en el litoral de Europa conti- 
nental. 


El Bittium de las Azores ha recibido 
generalmente el nombre de Bittium reti- 
culatum (Da Costa, 1778), la especie tipo 
del género y el representante del mismo 
más ampliamente distribuido y más 
común en las costas continentales euro- 
peas, por parte de numerosos autores, 
siguiendo en ello a Mac ANDREW (1857: 
124 y 150). Este nombre se siguió utili- 
zando durante muchos años por otros 


* c/ Araña, apartamentos Las Dunas 2, 04150 — Cabo de Gata, Almería (España) 


39 


Iberus, 29 (1), 2011 


autores, como JEFFREYS (1885: 57), DAUT- 
ZENBERG (1889: 41), NOBRE (1924: 79; 
1930: 09) MORTON AO 
BULLOCK, TURNER Y FRALICK (1990: 43), 
AZEVEDO (1991: 23), HOUBRICK (1993: 
274), MORTON, BRITTON Y MARTINS 
(1998: 64 y 97). Posteriormente, se citó 
provisionalmente como Bittium cf. reti- 
culatum por ÁVILA Y AZEVEDO (1997: 
326). También ha sido utilizado el 
nombre B. reticulatum para los fósiles del 
Cuaternario de la isla de Santa María 
por ZBYSZEWSKY Y FERREIRA (1962: 215), 
GARCÍA-TALAVERA (1990: 440) y CALLA- 
PEZ Y SOARES (2000: 314). También, ha 
sido considerada como Bittium depaupe- 
ratum (Watson, 1897) por NORDSIECK 
(1976: 7) y NORDSIECK Y GARCÍA-TALA- 
VERA (1979: 90). 

El nombre Bittium latreillii (Payrau- 
deau, 1826), ha sido sugerido para la es- 
pecie de Azores por VERDUIN (1976: 135) 
y, más recientemente, por ÁVILA, AMEN, 
AZEVEDO, CACHAO Y GARCÍA-TALAVERA 
(2002: 349), ÁvILA, CARDIGOS Y SANTOS 
(2004: 77; 2007: 46-47) y ÁVILA, SANTOS, 
PENTEADO, RODRIGUES, QUINTINO Y MA- 
CHADO (2005: 73). Por último, MARTINS, 
BORGES, ÁVILA, COsTa, MADEIRA Y MOR- 
TON (2009: 58) mencionan la especie co- 
mo Bittium cf. latreilli1, y citan también a 
B. latreilliií como muy raro frente a Vila 
Franca do Campo en la isla de Sáo Mi- 
guel a 180 m de profundidad. La dudas 
en la asignación de un nombre a la espe- 
cie han hecho que se haya quedado co- 
mo Bittium sp. por algunos autores, co- 
mo ÁvILA (1998: 483; 2000: 102), ÁvILA, 
AZEVEDO, GONCALVES, FONTES Y CARDI- 
GOS (1998: 493; 2000: 144) y CosTA Y Ávr- 
LA (2001: 120, 124). 

El objeto de este trabajo es aportar 
una descripción detallada del Bittium de 
las Azores y de mostrar que se trata de 
una especie distinta a las que viven en 
las costas europeas. En un principio se 
había contemplado su descripción como 
especie nueva (MORENO, 1998, no publi- 
cado en el sentido de la CINZ). En una 
posterior revisión bibliográfica realizada 
en el Muséum National d' Histoire Natu- 
relle de París (MNHN) se pudo hallar 
para la misma un nombre disponible, 
aunque caído en el olvido, basado en un 


60 


fósil del Cuaternario de la isla de Santa 
María y originalmente publicado en el 
binomen Cerithiopsis nana Mayer, 1864. 
La descripción de Mayer es explícita y la 
figura es suficiente para reconocer la 
especie, que ÁVILA, AMEN, AZEVEDO, 
CACHAO Y GARCÍA-TALAVERA (2002) han 
vuelto a encontrar en su localidad tipo 
aunque identificada como Bittium latrei- 
Ilii. El nombre ha sido ignorado por 
autores posteriores, con la única excep- 
ción de GARCÍA-TALAVERA (1990) quien 
se limita a repetir la cita original de 
Mayer sin relacionarla con el Bittium de 
su propio material (identificado p. 440 
como Bittium reticulatum). 

Pese a ello, se considera que la falta 
de uso para este nombre se debe a la 
falta de atención para la especie o a su 
confusión con otras especies, no al uso 
acostumbrado de un sinónimo frente al 
nombre nomenclaturalmente válido. 
Tampoco ha sido rechazado como 
“nomen oblitum” durante el periodo de 
vigencia de dicho concepto (1961-1973, 
artículo 23b de la edición de 1961 del 
CINZ). Por lo tanto, se propone la nueva 
combinación Bittium nanum (Mayer, 
1864) como el nombre válido para el 
Bittium de las Azores. 

Cerithiopsis nana Mayer, 1864 es un 
homónimo primario con respecto a 
Cerithiopsis tubercularis var. nana Jeffreys, 
1867, un nombre en uso para una 
especie taxonómicamente válida del 
género Cerithiopsis (familia Cerithiopsi- 
dae). Recientemente, CECALUPO Y 
ROBBA (2010), designaron Cerithiopsis 
nana Jetfreys, 1867 como especie tipo de 
un género nuevo, Nanopsis Cecalupo y 
Robba. Los mismos autores anuncian su 
intención de solicitar a la CINZ que 
suprima el nombre más antiguo de 
Mayer (1864). Esta acción no parece 
deseable al tratarse del nombre poten- 
cialmente válido de otra especie y, como 
solución alternativa para el problema 
nomenclatural, se debería contemplar el 
renombrar o redescribir Cerithiopsis nana 
Jeffreys, 1867. 

El nombre Cerithiopsis nana Mayer, 
1864 es homónimo secundario de Cerit- 
hium tuberculare var. nanum Wood, 1848, 
actualmente considerado como una 


MORENO: Bittium nanum, una especie válida de las islas Azores 


especie del género Cerithiopsis (Marquet, 
1997). Sin embargo, al no haberse reem- 
plazado y al no considerarse especies 
congenéricas en la actualidad, se aplica 
el artículo 59.2 del CINZ según el cual 
bajo estas circunstancias “el nombre 
más moderno no debe rechazarse, 
incluso si uno de los nombres de nivel 
especie se propuso originalmente en el 
género actual del otro”. 

Resulta sorprendente que un 
nombre específico tan trivial como 


nanum no resulte preocupado en combi- 
nación con el nombre genérico Bittium, 
pero este parece ser el caso. En la misma 
familia hallamos a Cerithium nanum C.B. 
Adams, 1850, actualmente considerado 
como perteneciente en la familia Tripho- 
ridae y sin uso como nombre válido, por 
lo tanto ni homónimo primario ni 
secundario; este nombre preocupa a 
Cerithium nanum Pallary, 1912 (un Ceri- 
thium del grupo de C. vulgatum Bru- 
guiere, 1792). 


SISTEMÁTICA 
Bittium nanum (Mayer, 1864) comb. nov. (Figs. 1-32) 


Cerithiopsis nana Mayer, 1864. Syst. Verz. der foss.: 66-67, lám.6, fig. 46 [citado por error fig. 49 en 
el texto]. 


Material estudiado: se han estudiado un total de 6.358 ej., todos procedentes de las islas Azores: 
Banco Joáo do Castro: St. 1, 1971, 7 ej., máx. 5,0 mm, MNHN, mission Biacores, 40-50 m, Faial: 
Castelo Branco, jul-79, 5 ej., máx. 3,5 mm, MNHN, col. Zibrowius, 28 m; Castelo Branco, St. P-11, 
1971, 15 ej. máx. 3,0 mm, MNHN, mission Biacores, 5-7 m; Horta, St. 29, 1971, 34 ej., máx. 2,5 mm, 
MNHN, mission Biacores, 2-3 m; Horta, St.L-9, 1971, 5 ej., máx. 2,0 mm, MNHN, mission Biacores, 
3 m; Horta, St. P-13, 1971, 10 ej., máx. 5,0 mm, MNHN, mission Biacores, 17 m; Monte da Guia, jul- 
79, 6 ej., máx. 4,5 mm, MNHN, Zibrowius, 47-60 m; Monte da Guia, St.P-19, 1971, 17 ej., máx. 4,5 
mm, MNHN, mission Biacores, 4 my St. L- 8, 1971, 1 ej., máx. 5,8 mm, MNHN, mission Biacores, 8 
m; St. L-11, 1971, 12 ej., máx. 8,0 mm, MNHN, mission Biacores, 2-4 m; St. P-13, 1971, 4 ej., máx. 2,0 
mm, MNHN, mission Biacores, 3-4 m;, Flores: I. Muda, St. P-24, 1971, 8 ej., máx. 4,0 mm, MNHN, 
mission Biacores, 24 m, 5. Cruz de Flores, 1989, 300 ej., máx. 4,0 mm, MNHN, Exp. Flores, col. Gofas, 
20 m; S. Cruz de Flores, 1989, 1 ej., máx. 7,0 mm, MNHN, Exp. Flores, col. Gofas, 1 m; S. Cruz de 
Flores, 1989, 1000 ej., máx. 6,0 mm, MNHN, Exp. Flores, col. Gofas, 20 m; S. Cruz de Flores, 1989, 
1000 ej., máx. 7,0 mm, MNHN, Exp. Flores, col. Gofas, 20 m; Formigas: (E) St. P-44, 1971, 3 ej., máx. 
4,0 mm, MNHN, mission Biacores, 35-45 my (E) St. P-44, 1971, 13 ej., máx. 4,5 mm, MNHN, mission 
Biacores, 43 m; (W) St. P-43, 1971, 4 ej., máx. 5,5 mm, MNHN, mission Biagores, 15 m; Pico: Lajes 
do Pico, 04/07/1995, 6 ej., máx. 9,5 mm, col. D. Moreno, 1 my Lajes do Pico, 04/07/1995, 18 ej., máx. 
8,5 mm, col. D. Moreno, 1 m; Lajes do Pico, 04/07/1995, 7 ej., col. D. Moreno, 1 m, Lajes do Pico, 
04/07/1995, 10 ej., máx. 9,5 mm, col. D. Moreno, 1 m;, Lajes do Pico, 04/07/1995, 41 ej., máx. 9,0 
mm, col. D. Moreno, 1 m; S. Roque, 02/07/1995, 15 ej., col. D. Moreno, 6 m; S. Maria: P. Malbusca, 
O1/10/19712 ej., máx. 5,5 mm, MNHN, mission Biacores, 12-25 m; P. Malbusca, St.P-3, 1971, 5 ej., 
máx. 3,5 mm, MNHN, mission Biacores, 30 m; Ponta Marváo, jun-90, 41 ej., máx. 7,5 mm, MNHN, 
col. S. y C. Gofas, 0-1 m; Praia Formosa, jun-90, 300 ej., máx. 7,0 mm, MNHN, col. S. y C. Gofas, 0 
m; Vila do Porto, jun-90, 3 ej., máx. 6,5 mm, MNHN, col. S. y C. Gofas, 0-1 m; Vila do Porto, jun- 
90, 52 ej., máx. 6,0 mm, MNHN, col. Gofas y Azevedo, 9 m;, Vila do Porto, jun-90, 300 ej., máx. 9,0 
mm, MNHN, col. Gofas y Azevedo, 6 m; Sáo Miguel: Agua d'Alto, jul-88, 9 ej., MNHN, col. Gofas, 
50 m; Caloura, B. Areia, jul-88, 111 ej., máx. 7,0 mm, MNHN, col. Gofas, 15 m; Caloura, Pta. Galera, 
jul-79, 3 ej., máx. 4,0 mm, MNHN, col. Zibrowius, 15 m; Caloura, Pta. Galera, 10/07/1983, 8 ej., 
máx. 4,0 mm, MNHN, col. Bouchet et al., 20 m; Caloura, Pta. Galera, 10/07/1983, 300 ej., máx. 6,3 
mm, MNHN, col. Bouchet et al., 20 m; Caloura, Pta. Galera, jul-88, 500 ej., máx. 7,0 mm, MNHN, 
Gofas, 13-18 m; Caloura, Pta. Galera, 29/06/1995, 58 ej., máx. 5,0 mm, col. D. Moreno, 20 m; Capelas, 
St. P-5, 1971, 8 ej., máx. 3,0 mm, MNHN, mission Biacores, 12 m; Capelas, M. St.P-41, 1971, 40 ej, 
máx. 5,5 mm, MNHN, mission Biacores, 29 m, Feteiras, jul-88, 500 ej., máx. 6,0 mm, MNHN, col. 
Gofas, 15-24 m;, Lagoa, jul-88, 500 ej., máx. 6,5 mm, MNHN, col. Gofas, Infralitoral; Ponta Delgada, 
09/07/1983, 300 ej., máx. 6,0 mm, MNHN, col. Bouchet, et al., 10-20 m; Ponta Delgada, jul-88, 53 


61 


Iberus, 29 (1), 2011 


ej., máx. 8,0 mm, MNHN, col. Gofas, 0 m; Ponta Delgada, St.P-29, 1971, 3 ej., máx. 3,5 mm, MNHN, 
mission Biacores, 12 m; Ponta Piramide, jul-88, 300 ej., máx. 5,0 mm, MNHN, col. Gofas, 13 my St. 
P-32, 1971, 12 ej., máx. 4,3 mm, MNHN, mission Biacores, 7 m; Vila Franca, jul-88, 100 ej., máx. 4,0 
mm, MNHN, col. Gofas, 10 m; Vila Franca, jul-88, 57 ej., máx. 8,0 mm, MNHN, col. Gofas, 0-5 m; 
Vila Franca, Ilheu, jul-88, 113 ej., máx. 7,5 mm, MNHN, col. Gofas, 0-1 m; Vila Franca, Ilheu, 
29/06/1995, 88 ej., máx. 4,5 mm, col. D. Moreno, 15 m;, Vila Franca, St.P-36, 1971, 21 ej., máx. 6,0 
mm, MNHN, mission Biacores, 24 m; Terceira: P. Diego, St. P-7, 1971, 22 ej., máx. 5,0 mm, MNHN, 


mission Biacores, 40 m;, St. P-7, 1971, 7 ej., máx. 4,3 mm, MNHN, mission Biacores, 39-40. 


Redescripción: La concha (figs. 1-7) es 
pequeña, de unos 6 a 9 mm en los 
adultos, 9,5 mm es el máximo medido 
en el material estudiado (6.358 ejempla- 
res). Sin embargo, existen ejemplares 
aislados que superan los 10 mm ($. 
ÁVILA, com. pers.). Los adultos tienen 
unas 10 vueltas de espira. La superficie 
está surcada por numerosas costillas 
espirales y axiales que, al encontrarse, 
forman un pequeño tubérculo. El 
número de costillas espirales es de 5 Ó 6. 
Los adultos más pequeños tienen 5, pero 
los más grandes tienen 6 en la vuelta del 
cuerpo o en la anterior. 

La formación de las costillas espira- 
les es muy similar a la conocida para B. 
latreillii (VERDUIN, 1976) pero algo más 
rápida, por ello tienen una 6* costilla. El 
origen de las distintas costillas espirales 
es el siguiente. La 3* costilla (en el 
tiempo, no por su posición) se forma en 
posición subsutural, aproximadamente 
en la primera o segunda vuelta de la 
teleoconcha, y rápidamente alcanza en 
tamaño a las anteriores. Posteriormente, 


la 4* costilla comienza a formarse por 
debajo de la 3* costilla o subsutural, 
aproximadamente en la tercera vuelta 
de espira (Figs. 5 y 6), antes que en B. 
latreillí1. La 5* costilla nace por debajo de 
la 3% o subsutural, aproximadamente en 
la sexta vuelta de espira de la teleocon- 
cha (Fig. 6). Por último, la 6* costilla 
espiral nace igual que las anteriores, por 
debajo de la 3? o subsutural, aproxima- 
damente en la novena o décima vuelta 
de espira. La microescultura de la 
superficie de la concha es similar a la 
conocida para B. reticulatum y B. latreilli1, 
con microtubérculos en los espacios 
intercostales (espirales) y pequeñas cavi- 
dades en las costillas (Fig. 7). 

El número de costillas axiales, que 
también es más numeroso que en B. 
latreillit, es de 24 a 28 en la vuelta del 
cuerpo o en la anterior. Las costillas son 
finas, ortoclinas en las primeras vueltas 
de la teleoconcha y prosoclinas u opisto- 
cirtas en las últimas. Los tubérculos que 
se forman al cruzarse las costillas axiales 
con las espirales son brillantes, redonde- 


(Página derecha) Figuras 1-11. Bittium nanum. Conchas, microescultura al MEB y opérculo. 1, 2: 
conchas de Lajes do Pico, Is. Pico, Azores (4/V11/1995) (8,1 y 7,4 mm de longitud); 3, 4: conchas 
de Marváo, Is. Santa María, Azores (V1/1990) (8 y 7 mm); 5: formación de la 42 costilla espiral 
(flecha) y microescultura, S. Roque do Pico, Is. Pico (2/VI1/1995); 6: concha juvenil mostrando la 
formación de la 42 y 52 costillas espirales (flechas), S. Roque do Pico, ls. Pico (2/V11/1995); 7: 
detalle de la microescultura de la concha, S. Roque do Pico, Is. Pico (2/V11/1995); 8, 9: conchas 
con pérdida de la escultura, Lajes do Pico, Is. Pico (4/V11/1995) (6 y 5,7 mm); 10: concha juvenil 
mostrando la pérdida de escultura, S. Roque do Pico, ls. Pico (2/V11/1995); 11: opérculo, Lajes do 
Pico, Is. Pico (4/V11/1995). 

(Right page) Figures 1-11. Bittium nanum. Shells, SEM of microsculpture and operculum. 1, 2: shells 
from Lajes do Pico, Pico Is., Azores (4/VI1/1995) (8.1 and 7.4 mm in length); 3, 4: shells from 
Marváo, Santa Maria ls., (VI/1990) (8 and 7 mm); 5: formation of the 4' spiral rib (arrow) and 
microsculpture, S. Roque do Pico, Pico Is. (2/V1/1995); 6: juvenile shell showing the formation of the 
4” and 5” ribs spirals (arrows), S. Roque do Pico, Pico ls. (2/VI/1995); 7: detail of shell microsculp- 
ture, S. Roque do Pico, Pico ls. (2/VH/1995); 8, 9: shells with loss of sculpture, Lajes do Pico, Pico l., 
(4/VI1/1995) (6 and 5.7 mm); 10: juvenile shell showing loss of sculpture, S. Roque do Pico, Pico l. 
(2/VI/1995); 11: operculum, Lajes do Pico, Pico Is. (4/VH/1995). 


62 


álida de las islas Azores 


1ttium nanum, una especie v 


B 


MORENO 


63 


Iberus, 29 (1), 2011 


ados, se encuentran muy próximos unos 
a otros y son muy uniformes. Como en 
las demás especies del género, las costi- 
llas axiales no continúan en la base. 

Las costillas espirales de la base son 
lisas, unas 6 ó 7 en los adultos. Las dos 
adapicales suelen ser más fuertes, segui- 
das de una 3* más débil que la 4”. Las 
últimas van haciéndose más débiles 
hasta la columela. 

Con respecto a la escultura de la 
concha, una característica que hay que 
destacar, por ser muy peculiar, es la 
existencia de numerosos ejemplares que 
la pierden totalmente en alguna fase de 
su crecimiento o en la mayor parte de la 
teleoconcha (Figs. 8, 9, 10, 24, 29, 30 y 
32). Estas espiras suelen desviarse con 
frecuencia del eje de la columela, con lo 
que el perfil de la concha se hace irregu- 
lar. Dicho crecimiento deforme, que no 
parece afectar al normal desarrollo del 
animal, no se encuentra aislado en 
determinadas poblaciones, puesto que 
se ha encontrado en numerosas localida- 
des de distintas islas y podría estar pre- 
sente en todo el archipiélago (es muy 
frecuente, al menos, en las islas de Pico, 
S. Miguel y S. María). HOUBRICK (1993) 
no hace referencia en ningún momento 
a esta alteración en el crecimiento, que 
parece ser propio de B. nanum, y no se 
ha observado o no se conoce en otras 
especies del género. 

Cuando la escultura se pierde, las 
conchas aparecen más o menos lisas, sin 
costillas espirales, costillas axiales ni 
microescultura, y con aspecto deforme, 
muy tosco (Figs. 8, 9 y 29). En primer 
lugar podría pensarse que se trata de 
conchas rodadas, pero no es así al 
haberse observado en individuos vivos 
y en conchas frescas. También se podría 
pensar que se trata de una degeneración 
o de alguna malformación, debida 
quizás al aislamiento genético que sufre 
la especie en su conjunto O cada una de 
sus distintas poblaciones insulares, pero 
no parece ser así, puesto que muchos 
individuos que sufren esta pérdida de 
escultura vuelven a recuperarla en fases 
sucesivas. Así, encontramos ejemplares 
con concha normal, seguida por unas 
vueltas de espira (1-3 vueltas) sin escul- 


64 


tura, y, de nuevo, vueltas con escultura 
normal (Figs. 24 y 30). Probablemente se 
trate de un morfo más, que se presenta 
en un número elevado de individuos y 
que es favorable en ciertas circunstan- 
cias para el camuflaje del animal. Estos 
ejemplares deformes presentan un 
ombligo incipiente (Figs. 8, 9 y 10), que 
no se encuentra en las conchas de creci- 
miento normal. 

El perfil general de las vueltas de 
espira en las conchas de crecimiento 
normal con escultura es bastante recto, 
pero no tan marcado como en B. latreilliz. 
El perfil general de la concha no es tan 
triangular como en B. latreillii, sino lige- 
ramente pupoide, con las primeras 
vueltas de bordes divergentes y las 
últimas con bordes cada vez más parale- 
los. La periferia de la concha en los 
ejemplares que han perdido la escultura 
espiral, como ya se ha comentado, es 
totalmente irregular. 

La abertura es ligeramente ovalada, 
con el labio externo apenas engrosado 
en los adultos, aunque aumenta en 
dimensiones y vuelo, pero de forma 
menos marcada que en B. latreillii. La 
especie posee varices, sobre todo los 
ejemplares de mayor tamaño en la 
vuelta del cuerpo 

El color de fondo de la concha es 
uniforme, pardo claro o castaño (Figs. 
26-30 y 32). Los tubérculos suelen distin- 
guirse muy bien, pues son de color 
claro, ya sea su tono natural o acen- 
tuado por una ligera erosión de los 
puntos más sobresalientes de la concha. 
Es frecuente que, como en B. latreillii, 
algunos tubérculos estén pigmentados 
de blanco, principalmente a lo largo del 
cordón subsutural, aunque también en 
todos los tubérculos que constituyen 
una costilla axial determinada o en 
tubérculos sueltos en cualquier posición. 
Las costillas espirales de la base suelen 
estar pigmentadas de blanco y castaño, 
intermitente, muy marcado en las dos 
costillas adapicales. 

La protoconcha de B. nanum (Figs. 
12-16) es muy similar a la B. latreillii, lo 
que confirma el estrecho parentesco 
entre ambas especies, ya observado al 
estudiar la teleoconcha. Tanto las 


MORENO: Bittium nanum, una especie válida de las islas Azores 


Figuras 12-19. Bittium nanum. Protoconcha y rádula al MEB. 12-16: S. Roque do Pico, ls. Pico, 
Azores (2/VI1/95); 17-19: Lajes do Pico, Is. Pico (4/V11/1995). 12: protoconcha con borde sinusí- 
gero roto; 13, 14: protoconcha completa; 15, 16: microescultura y detalle; 17: rádula completa; 
18, 19: varias filas de la rádula con los dientes marginales cerrados y abiertos. 

Figures 12-19. Bittium nanum. SEM of protoconch and radula. 12-16: S. Roque do Pico, Pico l»., 
Azores (2/VI1/95); 17-19: Lajes do Pico, Pico ls. (4/VH/1995). 12: protoconch with broken sinusigera 
rim; 13, 14: complete protoconch; 15, 16: microsculpture and detail; 17: complete radula; 18, 19: 
several rows of'the radula with closed and open marginal teeth. 


65 


Iberus, 29 (1), 2011 


dimensiones de la protoconcha en su 
conjunto (385 um: n= 4), como la presen- 
cia de dos costillas espirales en la última 
vuelta de la protoconcha II y una micro- 
escultura con tubérculos patentes, es 
prácticamente igual a la observada de B. 
latreilli1. Sin embargo, existen algunas 
ligeras diferencias. La anchura de la pro- 
toconcha es algo mayor en esta especie 
(media de 289 um: n= 4) que en B. latrei- 
1lii (media de 260 um; n= 7). Los tubér- 
culos son de base ovalada, de entre 7 y 9 
um de diámetro mayor (Figs. 15 y 16), 
algo más grandes que los de B. latreilliz. 
Entre estos tubérculos hay otros diminu- 
tos, de unos 0,2 um (Fig. 16), que están 
también presentes en B. latreillii. 

El animal de B. nanum (Figs. 20-24, 
26, 28 y 32) es similar al de otras espe- 
cies del género, tanto en forma como en 
pigmentación. Los tentáculos cefálicos 
son más largos que el morro y el borde 
del manto posee papilas. Las hembras 
en la época de reproducción presentan 
un ovopositor desarrollado (Fig. 20: 
ovp). El color de fondo es pardo claro, 
sobre el que se disponen manchas de 
color pardo oscuro, principalmente en la 
cabeza y dorso, y puntos de color blanco 
Opaco, dispersos por toda la superficie 
(Figs. 26, 28, 32). Los tentáculos tienen 
manchitas oscuras y blancas en los 
bordes y centro, con frecuencia, de color 
rojizo, como en B. latreilli1. El extremo de 
los tentáculos tiene manchas blancas. 
Existe una mancha amarillenta detrás 
del ojo, oculta por manchas oscuras más 
superficiales. En la base del morro, que 
es la zona más oscura, se distingue con 
dificultad, por transparencia, el bulbo 
bucal, que es anaranjado. Los labios son 
de color claro con puntos blancos. El 
propodio está poco pigmentado de 
oscuro en el dorso y posee pequeñas 
manchas en el centro. Los bordes latera- 
les del propodio tienen pequeñas 
papilas redondeadas pigmentadas de 
blanco. El surco ciliado derecho es más 
pálido que las áreas próximas. El borde 
del manto tiene manchitas de color 
pardo entre las papilas, que no poseen 
manchas blancas. Más internamente, el 
manto tiene una banda amarilla paralela 
al borde. Los lóbulos del opérculo, a 


66 


ambos lados del animal, poseen una 
gran mancha de color blanco, como en 
B. latreillii. El pie es blanquecino con 
puntos blancos y la suela es de color 
crema con dos bandas internas amari- 
llentas paralelas junto a los bordes y 
numerosos puntos blancos por toda su 
superficie. 

La rádula de B. nanum (Figs. 17-19) 
es muy similar a la de B. reticulatum y a 
la de B. latreillii. Su longitud, en ejem- 
plares adultos de entre 8 y 9 mm de lon- 
gitud de concha, es de poco más de 900 
um y tiene unas 40 filas de dientes. El 
diente raquídeo tiene entre 2 y 3 dentí- 
culos a cada lado del diente central. El 
diente lateral tiene entre 1 y 2 cúspides 
en la cara interna y entre 3 y 7 en la 
externa. El diente marginal externo tiene 
entre 3 y 5 dentículos en la cara interna 
y entre 3 y 7 en la externa. Por último, el 
diente marginal externo tiene entre 3 y 9 
cúspides en el lado interno, mientras 
que el externo es liso. . 

Comentarios: Bittium latreillii es una 
especie de amplia distribución en las 
costas europeas, desde el Golfo de 
Vizcaya hasta el Sahara Occidental, 
incluyendo Madeira y Canarias, y todo 
el Mediterráneo, que vive en fondos 
infra- y circalitorales. Es similar a B. 
latreillii, pero posee una concha más 
pequeña (hasta unos 10 mm), con un 
perfil más redondeado, a veces cirtoco- 
noide. La protoconcha de B. nanum es 
más ancha y con tubérculos más 
grandes que la de B. latreillii. No es de 
extrañar que en las islas Azores, además 
de la especie endémica y de amplia dis- 
tribución en el archipiélago, B. nanum se 
pueda encontrar alguna población 
aislada de B. latreillii, como parecen 
haber demostrado MARTINS ef al. (2009) 
que podría provenir de Madeira o de la 
costa europea continental, gracias a las 
larvas planctotróficas que pueden reco- 
rrer grandes distancias en el mar. 

Otros autores, como NORDSIECK (1976) 
y NORDSIECK Y GARCÍA-TALAVERA (1979), 
han utilizado para esta especie el nombre 
Bittium depauperatum (Watson, 1897). Sin 
embargo, B. depauperatum fue descrito por 
WATSON (1897) para Madeira. Se han con- 
sultado 3 sintipos de Bittium depauperatum 


MORENO: Bittium nanum, una especie válida de las islas Azores 


Figuras 20-24. Bittium nanum. Anatomía externa. 20-22. Lajes do Pico, Is. Pico, Azores 
(4/V11/1995). 20: hembra mostrando el ovopositor (ovp); 21: animal visto desde el lado izquierdo; 
22: pigmentación de la cabeza, de los tentáculos cefálicos y del propodio; 23: pigmentación del 
animal, vista ventralmente. S. Cruz, Ís. Flores, Azores (1989, MNHN) (Dibujo de S. Gofas); 24: 
ejemplar con concha que ha perdido la escultura y que ha vuelto a recuperarla. Ponta Piramide, ls. 
S. Miguel, Azores (V11/88, MNHN) (dibujo de S. Gofas con algunas modificaciones). 

Figures 20-24. Bittium nanum. External anatomy. 20-22. Lajes do Pico, Pico ls., Azores 
(4/VI1/1995). 20: female showing the ovipositor (ovp); 21: animal viewed from the left side; 22: pig- 
mentation of the head, cephalic tentacles and propodium; 23: pigmentation of the animal, ventral view. 
S. Cruz, Flores Ís., Azores (1989, MNHN) (Drawing by S. Gofas); 24: specimen with shell sculpture 
interrupted and later recovered. Ponta Piramide, S. Miguel ls., Azores (VIL/88, MNHN) (drawing by 
S. Gofas with some modifications). 


67 


Iberus, 29 (1), 2011 


(Watson, 1897) del Natural History 
Museum de Londres, y se trata de una 
especie con protoconcha pauciespiral, no 
multiespiral como la especie de Azores. 

Datos anteriores: La única descripción 
previa conocida de la concha es la de 
HOUBRICK (1993: 274, fig. 3, A, B y C), 
pero al considerar todas las especies de 
Bittium del Atlántico oriental y Medite- 
rráneo como una sola (B. reticulatum), 
parte de la descripción y algunas de las 
ilustraciones se refieren a la especie de 
Azores, mientras que otros datos e ilus- 
traciones están basados en otras pobla- 
ciones del continente. Así, sus figuras 3 
A, B y €, corresponden a la especie de 
Azores (S. Miguel), mientras que las 
figuras 3 D, E y H, corresponden a B. 
reticulatum, las dos primeras fotografías 
a un ejemplar procedente de Túnez, 
mientras que la última se trata de un 
juvenil del que no da la localidad. En los 
datos de su descripción, la talla que 
señala corresponde, probablemente a 
datos bibliográficos, puesto que da 15 
mm de longitud máxima, cuando la 
especie de Azores no suele superar los 
10. Sin embargo, los datos sobre las cos- 
tillas espirales sí coinciden con la 
especie de Azores (“5 costillas espirales 
principales en la vuelta del cuerpo”). 

El material que utilizó HOUBRICK 
(1993: 281) en su revisión de la subfamilia 
Bittiinae para estudiar la anatomía del 
género Bittium era de las islas Azores, y 
por tanto corresponde con la especie que 
aquí nos ocupa. Este autor, aunque 
conocía la variabilidad de Bittium reticula- 


tum, y que algunos autores consideraban 
a las poblaciones de las islas Azores como 
distinta a B. reticulatum, y que por tanto 
en aguas europeas existe un complejo de 
especies, consideró a todas ellas en 
sentido amplio (sensu lato) como una sola: 
B. reticulatum. HOUBRICK (1993: 282) 
utilizó como argumento para considerar 
el complexo de especies de B. reticulatum 
como una única especie el hecho de que 
la anatomía conocida hasta ahora de 
otras poblaciones, como la de JOHANSSON 
(1947) o los bocetos realizados por 
Ponder de ejemplares de Suecia (PONDER, 
in litt.) correspondían básicamente con 
las observaciones realizadas sobre su 
material de Azores. 

Las conchas que seleccionó Hou- 
BRICK (1993) para ilustrar a B. reticulatum 
tenían distintas procedencias y corres- 
pondían a diferentes especies. En con- 
creto, las figuras 3 A, B y C correspon- 
den a Bittium nanum y proceden de la 
isla de Sáo Miguel (Azores), mientras 
que el resto son de Bittium reticulatum y 
son de Túnez (fig. 3 D y E), o no se 
indica procedencia (fig. 3 H). 

La protoconcha que describe e 
ilustra HOUBRICK (1993: 274, Fig. 3 G) 
para B. reticulatum corresponde real- 
mente a esa especie. Aunque dicho 
autor trabajó material de Azores donde 
no vive B. reticulatum, el ejemplar que 
utilizó para describir e ilustrar la proto- 
concha (sin detallar localidad), procede 
sin duda del continente, pues posee una 
superficie lisa. Además, la protoconcha 
ilustrada por reste autor no se observa 


(Página derecha) Figuras 25-32. Bittium nanum. Hábitat y animales vivos. 25-30. Lajes do Pico, ls. 
Pico, Azores (4/V11/1995). 25: laguna costera en Lajes do Pico; 26-28: animales vivos; 29: ejem- 
plar con concha de escultura normal (abajo) y ejemplar con concha sin escultura (arriba); 30: 
ejemplar con concha que ha perdido la escultura normal y la ha vuelto a recuperar posteriormente; 
31: hábitat en arena gruesa junto a Ervilia castanea, -15 m, lheu de Vila Franca, Is. Sáio Miguel 
(29/V1/1995); 32: ejemplar con concha que ha perdido la escultura, Ilheu de Vila Franca, Is. Sáo 
Miguel (29/V1/1995). 

(Right page) Figures 25-32. Bittium nanum. Living animals and habitats. 25-30. Lajes do Pico, Pico 
ls., Azores (4/V11/1995). 25: coastal lagoon in Lajes do Pico; 26-28: living animals; 29: specimen with 
normal shell sculpture (below) and other with no shell sculpture (above); 30: specimen in which normal 
shell sculpture has been lost and. later recovered; 31: habitat in coarse sand with Ervilia castanea, -15 
m, Ilheu de Vila Franca, Sáo Miguel ls. (29/V1/1995); 32: Specimen in which normal shell sculpture 
has been lost, lheu de Vila Franca, Sáo Miguel ls. (29/V1/1995). 


68 


MORENO: Bttium nanum, una especie válida de las islas Azores 


Iberus, 29 (1), 2011 


en su totalidad, debido a que la abertura 
de la misma queda oculta por la concha. 

El animal de B. nanum (Figs. 20-24, 
26, 28 y 32) fue descrito e ilustrado con 
mucho detalle por HOUBRICK (1993: 276, 
fig. 4). El opérculo ilustrado por Hou- 
BRICK (1993: 276, fig. 3 f) para B. reticula- 
tum, podría corresponder a B. nanum, 
aunque de nuevo este autor no detalla la 
localidad. En base a material procedente 
de las islas Azores recogido durante el 
presente estudio, el opérculo (Fig. 11) es 
muy similar al conocido para otras espe- 
cies del género, con forma ligeramente 
ovalada, núcleo excéntrico, superficie 
lisa y última vuelta más estrecha que el 
resto del opérculo. La impresión muscu- 
lar es algo más larga que la mitad de la 
longitud del opérculo. 

La anatomía descrita en detalle por 
HOUBRICK (1993) para B. reticulatum 
corresponde realmente a B. nanum. Esta 
fundamental aportación de dicho autor 
se basa en disecciones y cortes histológi- 
cos de material recogido vivo por él 
durante el “Workshop” celebrado en 
Sáo Miguel (Azores) en julio de 1988. 
Las observaciones realizadas para esta 
tesis coinciden plenamente con las reali- 
zadas por dicho autor. De la descripción 
de HOUBRICK (1993) destaca el detallado 
estudio del sistema reproductor. El ovi- 
ducto paleal está compuesto por dos 
láminas gruesas. En la lámina media se 
encuentran la bursa y el receptáculo 
seminal, y en la lámina lateral se 
encuentra el surco de lamelas ciliadas y 
el receptáculo del espermatóforo (que él 
llama bursa). Todo es muy similar a lo 
descrito para B. reticulatum por JOHANS- 
SON (1947). La completa descripción de 
la especie realizada por HOUBRICK 
(1993) coincide en casi todo con las 
observaciones efectuadas en el presente 
trabajo. Dicho autor ilustra (Fig. 4 d) los 
ganglios bucales en posición dorsal res- 
pecto al bulbo bucal, lo que debe tra- 
tarse de un error ya que su posición real 
es ventral. 

HOUBrICK (1993: 276-278, Tabla. 2 y 
Fig. 5) ofrece datos contradictorios sobre 
la rádula; no coincide lo que describe con 
lo que se ve en las fotografías que presenta, 
aunque los valores indicados entran dentro 


7O 


de la variabilidad del material propio, pro- 
cedente de la isla Pico. 

Las puestas fueron descritas por 
HOUBRICK (1993), se supone que basán- 
dose en el material obtenido por él en 
julio de 1988 en Azores. Nosotros no las 
hemos observado. HOUBRICK (1993: 279) 
dice: “Puesta constituida por un cordón 
gelatinoso fino (de unos 25 mm de lon- 
gitud estirado) en ajustada espiral en el 
sentido de la agujas del reloj o doblado 
sobre sí mismo de forma irregular y 
fijada al sustrato. Cordón gelatinoso que 
contiene muchos huevos opacos peque- 
ños (de 0.65 um de diámetro) [sic, por 65 
um] cada uno dentro de una fina, trans- 
parente y hialina cápsula (de 110 um de 
diámetro). Toda la puesta contiene unos 
800 huevos”. 

Hábitat y distribución: El hábitat de B. 
nanum es preferentemente infralitoral. 
La especie es muy abundante en todo el 
archipiélago de Azores donde vive 
desde fondos muy someros hasta 50 m 
de profundidad (ejemplares recogidos 
vivos). Algunas conchas estudiadas pro- 
ceden de hasta 60 m, pero no correspon- 
den con material fresco. En la bibliogra- 
fía se indica que se ha encontrado vivo 
hasta una profundidad de 38 m y 
conchas hasta 360 m (MARTINS ET AL., 
2009). Ocupa todo tipo de fondos 
rocosos infralitorales, siendo poco fre- 
cuente en el cinturón de algas del meso- 
litoral y en las pequeñas lagunas coste- 
ras de las islas como la de Lajes do Pico, 
en Pico (Fig. 25). Sin embargo, es una de 
las especies más abundantes en Azores 
en las algas fotófilas infralitorales 
(AZEVEDO, 1991: 23, 30-33); COSTA Y 
ÁvILA (1998) la consideran dominante 
sobre Halopteris. También se encuentra 
en enclaves de arena gruesa, constitui- 
dos por un detrito de origen volcánico, 
al menos en zonas situadas entre 
grandes bloques o en la base de acanti- 
lados rocosos. Este hábitat marginal lo 
comparte con el bivalvo Ervilia castanea, 
también abundantísimo en las islas (Fig. 
31). Este tipo de hábitat es diferente al 
observado en B. reticulatum y B. latretlli1, 
que aunque abundan en la arena rete- 
nida por las algas que cubren piedras o 
bloques rocosos, nunca se encuentran 


MORENO: Bittium nanum, una especie válida de las islas Azores 


vivos en los detritos que quedan entre 
ellos, que tienen cúmulos de conchas y 
el sedimento está suelto. Este último 
tipo de hábitat es más parecido al de B. 
submamillatum. Se desconoce si la 
especie vive también en fondos detríti- 
cos o de cascajo, aunque es muy posible 
dado el amplio rango batimétrico que 
posee. MORTON (1990: 17, como B. reticu- 
latum) cita a la especie en una plata- 
forma rocosa cubierta de arena en el 
interior del islote de Vila Franca (S. 
Miguel, Azores). En la misma isla 
BULLOCK, TURNER Y FRALICK (1990: 47 y 
54, como B. reticulatum) la citan sobre 
Codium adhaerens y sobre algas feofíceas. 

La distribución de B. nanum se 
encuentra restringida exclusivamente a 
las islas Azores. Existen citas de todas 
las islas del archipiélago; ÁvILA (1998: 
483) aporta una extensa recopilación de 
citas de las distintas islas. Para este 
trabajo se han estudiado muestras de 
casi todas ellas, en concreto de S. María, 
S. Miguel, Terceira, Pico, Faial y Flores, e 
incluso de los pequeños islotes llamados 
Formigas, situados entre S. Miguel y S. 
María, y en el Banco Joáo de Castro que 
se encuentra entre las islas orientales (5. 
María y S. Miguel) y las centrales (Ter- 
ceira, Graciosa, S. Jorge, Faial y Pico), 
donde ya había sido citada por ÁVILA Y 
AZEVEDO (1997) y por ÁMILA ET AE. 
(2004), respectivamente. Se creía que era 
la único representante del género 
Bittium presente en las islas Azores, pero 
recientemente MARTINS ef al. (2009) ilus- 
tran un material profundo que parece 
corresponder al auténtico B. latreillii, 
especie que consideran muy rara, proce- 
dente de 180 m de profundidad frente a 
Vila Franca do Campo en la isla de Sáo 
Miguel. 

Lo más probable es que B. nanum sea 
una especie formada a partir de una 
antigua población de B. latreillii que 
quedara aislada en Azores. El origen 
europeo de la especie se ve apoyado por 
el estudio de GOFAS (1990: 123) sobre las 
familias Rissoidae y Anabathridae en 
Azores, en el que demuestra que la afi- 
nidad de la fauna de las islas es casi 
completa con la de las costas europeas, 
mientras no existen apenas elementos 


pertenecientes a la fauna norteameri- 
cana. Lo mismo ocurre con el resto de la 
flora y fauna del archipiélago según 
observaciones personales. Las islas 
Azores deben considerarse parte de la 
región biogeográfica Lusitana (BRIGGS, 
1974: 208). 

B. latreillii debió colonizar en épocas 
pasadas las islas Azores, gracias, proba- 
blemente a una circulación de corrientes 
superficiales diferente de la actual, 
quizá en esa época las islas se encontra- 
ran más cerca de Europa de lo que lo 
están ahora. Esta población, muy alejada 
del continente, debió sufrir un prolon- 
gado aislamiento reproductivo, acen- 
tuado en la actualidad por unas corrien- 
tes superficiales que impiden el contacto 
entre las poblaciones del continente y 
las de las islas Azores, en ambos senti- 
dos. Este aislamiento ha producido la 
divergencia específica. 

Según GOFAS (1990b: 123, fig. 13), los 
movimientos actuales de aguas superfi- 
ciales de la zona están dominados por la 
corriente del Golfo, con dirección oeste- 
este que llega a las Azores y continúa 
hacia el norte de Europa. Las corrientes 
procedentes de las costas europeas 
tienen dirección sur hasta Madeira y 
Canarias, donde giran en dirección oeste 
pasando muy al sur de las Azores. El 
archipiélago queda en una zona central 
rodeado por corrientes que giran a su 
alrededor en el sentido de las agujas del 
reloj (anticiclón). Este movimiento del 
agua superficial es similar al atmosfé- 
rico. Es muy conocido el anticiclón de 
las Azores, centrado sobre las islas, que 
con frecuencia se traduce en calmas de 
varios días, a pesar de estar en medio 
del océano. 


Nota nomenclatural acerca de Cerit- 
hiopsis tubercularis var. nana Jeffreys, 
1867 

Como se ha mencionado anterior- 
mente, Cerithiopsis nana Mayer, 1864 es 
homónimo primario de Cerithiopsis 
tubercularis var. nana Jeffreys, 1867 y, 
pese a ello, este último nombre está 
actualmente en uso para una especie 
taxonómicamente válida de la familia 
Cerithiopsidae. CECALUPO Y ROBBA 


YN 


Iberus, 29 (1), 2011 


(2010) citan un buen número de referen- 
cias (entre ellas, VAN AARTSEN MENK- 
HORST Y GITTENBERGER, 1984: 29, fig. 
(SI PALAZZO ISO CAGADAS 
MIFSUD Y SAMMUT, 1996: p. 132, pl. 
14,fig. 1; GIRIBET Y PEÑAS, 1997: 50, fig. 
25, 29) y es previsible que se reúnan las 
condiciones requeridas en el artículo 
23.9 del CINZ sobre “inversión de pre- 
cedencia”, O sea al menos 25 citas del 
nombre por parte de al menos 10 
autores en un espacio de tiempo no 
menor de 10 años en los últimos 50 
años. 

No obstante, aunque se contemple la 
inversión de precedencia, el nombre es a 
su vez homónimo secundario de Ceri- 


AGRADECIMIENTOS 


Mi más sincero agradecimiento a 
todas las personas que han colaborado de 
una forma u otra en el presente trabajo, 
como a José Templado, director de Tesis 
que me animó y ayudó en todo momento 
en el estudio del género Bittium; a Serge 
Gofas que siempre me apoyó y permitió 
realizar mis estancias en París, a Philippe 
Bouchet, Bernard Métivier, Pierre 
Lozouet y Virginia Héros del Muséum 
National d'Histoire Naturelle de París 
que facilitaron en todo momento los 
trabajo en el Laboratorio de Malacologie; 
a Sergio P. Ávila por su amistad y el 


BIBLIOGRAFÍA 


ÁvILA S.P. 1998. Zonacáo intertidal de uma co- 
munidade malacológica na “Poca de Barra”, 
uma lagoa localizada na plataforma costeira 
da vila das Lajes do Pico, Acores. Agoreana, 
8 (4): 457-485. 

ÁviLA S.P. 2000. Shallow-water marine mol- 
luscs of the Azores: biogeographical rala- 
tionships. Arquipélago. Life and Marine Sci- 
ences, Suppl. 2 (Part A): 99-131. 

ÁviLA S.P.. AMEN R. AZEVEDO J.M.N., 
CACHAO M. Y GARCÍA-TALAVERA F. 2002. 
Checklist of the Pleistocene marine mol- 
lusks of Praínha and Lagoínhas (Santa 
Maria Island, Azores). Acoreana, 9 (4): 343- 
370. 


AZ 


thium tuberculare var. nanum Wood, 1848 
(Crag Mollusca, 1: 70, pl. 8, fig.5c), como 
ya ha sido notado por parte de varios 
autores anteriores (CECALUPO Y BUZZU- 
RRO, 2005: 134; LANDAU, LA PERNA Y 
MARQUET, 2006: 13). El nombre de Wood 
está en uso (SACCO, 1895: 66; REGTEREN 
ALTENA, BLOKLANDER Y POUDEROYEN, 
1955: 30, pl. 6, fig. 64; MARQUET, 1997: 
82; CLEVERINGA, MEJER, VAN LEEUWEN, 
DE WOLF POUWER, LISSENBERG Y 
BURGER, 2000: 204-205). Una solución 
más satisfactoria al problema nomencla- 
tural, pero que se sale del ámbito del 
pesente trabajo, sería que Cerithiopsis 
nana Jeffreys fuera renombrada o, mejor, 
descrita como nuevo taxón. 


envío de numerosa bibliografía e infor- 
mación sobre las islas Azores; a M”* 
Ángeles Ramos y Josefo Bedoya (triste- 
mente fallecido) por su colaboración y 
obtención de las fotografías de Scanning 
en el Museo Nacional de Ciencias Natu- 
rales de Madrid; y a Nuria Martín 
Herrero por todo su apoyo y ayuda en 
los muestreos, incluido Lajes do Pico, en 
un viaje de bodas malacológico inolvida- 
ble. También se agradece a dos revisores 
anónimos las sugerencias y correcciones 
realizadas que han mejorado sustancial- 
mente el trabajo. 


ÁvILA S.P. Y AZEVEDO J. M. N. 1997. Shallow- 
Water molluscs from the Formigas islets, 
Azores, collected during the “Santa Maria e 
Formigas 1990" scientific expedition. Acore- 
ana, 8 (3): 323-330. 

ÁvILA S.P., AZEVEDO J.M.N., GONGALVES J.M., 
FONTES]. Y CARDIGOS F. 1998. Checklist of the 
shallow-water marine molluscs of the Azores: 
1- Pico, Faial, Flores and Corvo. Acoreana, 8 
(4): 487-523. 

ÁVILA S.P., AZEVEDOJ.M.N., GONCALVES J.M., 
FONTES J. Y CARDIGOS F. 2000. Checklist of the 
shallow-water marine molluscs of the Azores: 
2- Sáo Miguel Island. Acoreana, 9 (2): 139- 
WS: 


MORENO: Bittium nanum, una especie válida de las islas Azores 


ÁvILA S.P., CARDIGOS F. Y SANTOS R.S. 2004. D. 
Joáo de Castro Bank, a shallow water hy- 
drotermal-vent in the Azores: checklist of 
the marine molluscs. Arquipélago. Life and 
Marine Sciences, 21 (A): 75-80. 

ÁviLA S.P., CARDIGOS F. Y SANTOS R.S. 2007. 
Comparison of the community structure of 
the marine mollusks of the “Banco D. Joáo de 
Castro” seamount (Azores, Portugal) with that 
of typical inshore habitats on the Azores ar - 

_ chipelago. Helgoland Marine Research, 61: 43-53. 

ÁVILA S.P., SANTOS A.C., PENTEADO A.M., Ro- 
DRIGUES A.M., QUINTINO I. Y MACHADO M.I. 
2005. The molluscs of the intertidal algal turf 
in the Azores. Iberus, 23 (1): 67-76. 

AZEVEDO J.N. 1991. Estudo das comunidades ma- 
lacológicas fitais do litoral em Sáo Miguel, Aco- 
res. Universidade dos Acores, 75 pp. 

BRIGGSJ.C. 1974. Marine zoogeography. McGraw 
Hill, Inc. New York, 475 pp. 

BULLOCK R.C., TURNER R.D. Y FRALICK R.A. 
1990. Species richness and diversity of algal- 
associated micromolluscan communities 
from Sáo Miguel, Acores. En: Martins A.M.F. 
(Ed.). The marine fauna and flora of the Azores. 
Acoreana Suplemento. (Proceedings of the First 
International Workshop of Malacology, Sáo 
Miguel, Azores, 1988): 39-58. 

CACHIA C., MIFSUD C. Y SAMMUT P.M. 1996. 
The marine Mollusca of the Maltese Islands (Part 
Two: Neotaenioglossa). Leiden, Backhuys Pu- 
blishers, 228 pp. 

CALLAPEZ P. Y SOARES A.F. 2000. Late Quater- 
nary warm marine mollusks from Santa 
Maria (Azores) paleoecologic and paleobio- 
geographic considerations. Ciencias da Terra, 
14: 313-322. 

CECALUPO A. Y BUZZURRO G. 2005. Cerithiopsis 
annae: una nuova specie di Cerithiopsidae per 
le coste tunisine. Bollettino Malacologico, 40 (9- 
213285 

CECALUPO A. Y ROBBA E. 2010. The identity of 
Murex tubercularis Montagu, 1803 and des- 
cription of one new genus and two new spe- 
cies Of the Cerithiopsidae (Gastropoda: Tri- 
phoroidea). Bollettino Malacologico, 46 (2): 45-64. 

CLEVERINGA P., MEJER T., VAN LEEUWEN 
R.J.W., DE WOLF H., POUWER R., LISSENBERG 
T. Y BURGER A.W. 2000. The Eemian strato- 
type locality at Amersfoort in the central 
Netherlands: a reevaluation of old and new 
data. Geologie en Mijnbourw, 79 (2/3): 197-216. 

CosTA A.C. Y AVILA S.P. 2001. Macrobenthic 
mollusc fauna inhabiting Halopteris spp. sub- 
tidal fronds in Sáo Miguel island, Azores. 
Scientia Marina, 65 (2): 117-126. 

DAUTZENBERG P. 1889. Contribution a la faune 
malacologique des Iles Acores. Résultats des 
dragages effectués par le yacht l'Hirondelle. 
Résultats des Campagnes Scientifiques accomplies 
sur son yatch par Albert I de Monaco, Fascicule 
A 


GARCÍA-TALAVERA F. 1990. Fauna tropical en el 
Neotirreniense de Santa Maria (1. Azores). La- 
vori S.I.M., 23: 439-443. 

GIRIBET G. Y PEÑAS A. 1997. Fauna malacológica 
del litoral del Garraf (NE de la Península 
Ibérica). Iberus, 15 (1): 41-93. 

GOFAS 5. 1990. The littoral Rissoidae and An- 
abathridae of Sáo Miguel, Azores. En Martins 
A.M.F. (Ed.): The marine fauna and flora of the 
Azores. Acoreana Suplemento. (Proceedings of 
the First International Workshop of Mala- 
cology, Sáo Miguel, Azores, 1988): 97-134. 

HOUBRICK R.S. 1993. Phylogenetic relationships 
and generic review of the Bittiinae (Proso- 
branchia: Cerithioidea). Malacología, 35: 261- 
315 

JEFFREYS J.G. 1885. On the mollusca procured 
during the Lightning and Porcupine expedi- 
tions. Proceedings of the Zoological Society of Lon- 
don: 27-63. 

JOHANSSON J. 1947. Úber den offenen Uterus bei 
einigen Monotocardiern ohne Kopulation- 
sorgan. Zoologiska Bidrag fran Uppsala, 25 102- 
110. 

LANDAU B., La PERNA R. Y MARQUET RR. 2006. 
The Early Pliocene Gastropoda (Mollusca) of 
Estepona, southern Spain, Part 6: 
Triphoroidea, Epitonioidea, Eulimoidea. 
Palaeontos, 10: 1-96, pl. 1-22. 

MAC ANDREW R. 1857. Report on the marine tes- 
taceous Mollusca of the North-east Atlantic 
and neighbouring Seas, and the physical con- 
ditions affecting their development. Report of 
the British Association for the Advancement of 
Science, (1856): 101-158. 

MARQUET R. 1997. Pliocene gastropod faunas 
from Kallo (oost-Vlaanderen, Belgium) - Part 
3. Caenogastropoda: Aporrhaidae to Muri- 
cidae, and Part 4. Buccinidae to Helicidae. 
Contributions to Tertiary and Quaternary Ge- 
ology, 34: 69-149. 

MARTINS A.M.F., BORGES]J.P., ÁVILA S.P., COSTA 
A.C., MADEIRA P. Y MORTON B. 2009. Illus- 
trated checklist of the infralittoral mollusks 
off Vila Franca do Campo. Acoreana, Suple- 
mento 6 (Setembro): 15-103. 

MAYER K. 1864. Systematisches Verzeichniss der 
fossilen Reste von Madeira, Porto Santo und 
Santa Maria, nebst Beschreibung der neuen Arten. 
Zúrich (Reedition Brill /Backhuys, Leiden, 
1988), I-VI, 109 pp. 

MORENO D. 1998. Sistemática y biología del género 
Bittium (Gastropoda, Prosobranchia) en la Pe- 
nínsula Ibérica. Tesis Doctoral, Universidad 
Complutense de Madrid, inédita, 473 pp. 

MORTON B. 1990. The intertidal ecology of 
Ilheu de Vila Franca - a drowned volcanic 
crater in the Azores. En Martins A.M.F. 
(Ed.): The marine fauna and flora of the Azores. 
Agoreana Suplemento. (Proceedings of the 
First International Workshop of Mala- 
cology, Sáo Miguel, Azores, 1988): 3-20. 


as 


Iberus, 29 (1), 2011 


MORTON B., BRITTON J.C. Y MARTINS A.M.F. 
1998. Ecologia Costeira dos Acores. Sociedade 
Afonso Chaves, Ponta Delgada, x + 249 pp. 

NOBRE A. 1924. Contribucoes para a fauna dos 
Acores. Anais do Instituto de Zoología da Uni- 
versidade do Porto, 1: 41-90. 

NORDSIECK F. 1976. Il genere Bittium Leach, 
1847 nei mari d'Europa. La Conchiglia, 8 (93- 
94): 6-9. 

NORDSIECK F. Y GARCÍA-TALAVERA F. 1979. Mo- 
luscos marinos de Canarias y Madera (Gastro- 
poda). Aula de Cultura Tenerife, 208 pp. 

PALAZZIS. 1994. Cerithiopsis nana (Jeffreys, 1867) 
vivente su Suberites. Bollettino Malacologico, 30: 
79-80. 

REGTEREN ALTENA C.O. VAN, BLOKLANDER A. 
Y POUDEROYEN L.P. 1955. De fossile schelpen 
van de Nederlandse stranden en zeegaten, 2. 
Basteria, 19: 2736. 

SACCO F. 1895. I Molluschi dei terreni terziaril del 
Piemonte e della Liguria. Parte XVII (Cerithi1dae, 
Triforidae, Cerithiopsidae e Diastomidae). To- 
rino, Carlo Clausen, 83 pp, 3 pls. 


74 


VAN AARTSEN J.J.; MENKHORST H.P.M.G. Y GIT- 
TENBERGER E. 1984. The marine Mollusca of 
the Bay of Algeciras, Spain, with general 
notes on Mitrella, Marginellidae and Turri- 
dae. Basteria, supplement 2: 1-135. 

VERDUIN A. 1976. On characters, variability, 
and distribution of the Europaean marine 
gastropods Bittium latreillii (Payraudeau) and 
Bittium lacteum (Philippi). Basteria, 40: 133-142. 

WATSON R.B. 1897. On the Marine Mollusca of 
Madeira. Linnean Journal of Zoology, 26: 233- 
329, 2:p1S: 

WOOD S.V. 1848. A monograph of the Crag Mol- 
lusca, with descriptions of shells from the upper 
Tertiaries of the British Isles. The Palaeonto- 
graphical Society Monographs, London, 1-208 
Pp 2115: 

ZBYSZEWSKY G. Y FERREIRA O.V. 1962. Étude 
géologique de l'ile de Santa Maria (Acores). 
Comunicacoes dos Servicos Geológicos de Por- 
tugal, 46, 209-245. 


NORMAS DE PUBLICACIÓN 


e La revista Iberus publica artículos de fondo, notas y monografías que versen sobre cualquiera de los aspectos 
relacionados con la Malacología. Se entiende por artículo un trabajo de investigación de más de 5 páginas de 
texto, incluidas láminas, gráficos y tablas. Las notas son trabajos de menor extensión. Las monografías son tra- 
bajos sobre un tema único, de extensión superior a las 50 páginas de la revista y que serán publicadas, si 
procede, como un suplemento de Iberus. Los autores interesados en publicar monografías deberán ponerse 
previamente en contacto con el Editor de Publicaciones. Se entiende que el contenido de los manuscritos no ha 
sido publicado, ni enviado simultáneamente a otra revista para su consideración. 

e Los manuscritos, así como toda la correspondencia relacionada con los mismos, deberán ser remitidos a: 
Serge Gofas, Editor de Publicaciones, Departamento de Biología Animal, Universidad de Málaga, Campus de 
Teatinos, s/n, 29071, Málaga, España y / o al correo electrónico <sgofasuma.es>. 

e El texto del trabajo podrá estar redactado en español, inglés, italiano, francés o portugués. 

e Los artículos, notas y monografías deberán presentarse sobre DIN A-4, por una sola cara a doble espacio 
(incluyendo referencias, pies de figura y tablas), con al menos 3 centímetros de margen por cada lado. Los tra- 
bajos se entregarán por triplicado (original y dos copias) y se incluirá una versión en un CD, o bien remitida 
por correo electrónico, utilizando procesadores de texto en sus versiones de corrientes de Windows (pero no 
en el formato .docx de Word 2007, el habitual de Windows Vista) o Macintosh. En caso de autoría compartida, 
uno de los autores deberá hacerse responsable de toda la correspondencia. 

e Junto con el trabajo debe incluirse una lista de al menos 4 posibles revisores del mismo, sin perjuicio de los 
que el propio Editor considere oportunos. 

e Los manuscritos se presentarán de acuerdo al siguiente esquema: 

Primera página. Deberá incluir un título conciso, pero sugerente del contenido del trabajo, así como una traduc- 
ción al inglés del mismo (si el artículo no está escrito en inglés). Cuando sea preciso, deberá incluir referencia a 
familia o taxones superiores. A continuación figurarán, por este orden, el nombre y apellidos completos del 
autor o autores, las direcciones completas de los mismos, y un resumen del trabajo y su traducción al inglés. 
Dicho resumen deberá sintetizar, en conjunción con el título, tanto los resultados como las conclusiones del 
artículo; se sugiere una extensión de 100 a 200 palabras. 

Páginas siguientes. Incluirán el resto del artículo, que debe dividirse en secciones precedidas por breves encabe- 
zamientos. Siempre que sea posible, se recomienda seguir el siguiente esquema: Introducción, Material y 
métodos, Resultados, Discusión, Conclusiones, Agradecimientos y Bibliografía. Si se emplean abreviaturas no 
habituales en el texto, deberán indicarse tras el apartado de Material y Métodos. 

e Las notas breves deberán presentarse de la misma forma, pero sin resumen. 

e Deberán evitarse notas a pie de página y referencias cruzadas. Deberán respetarse estrictamente los Códigos 
Internacionales de Nomenclatura Zoológica y Botánica (últimas ediciones). Cuando un taxón aparezca por 
primera vez deberá citarse su autor y fecha de su descripción. En el caso de artículos sistemáticos, cuando se 
den las sinonimias de los taxones, éstas deberán citarse COMPLETAS, incluyendo en forma abreviada la 
publicación donde fueron descritas, y la localidad tipo si es conocida entre corchetes, según el siguiente 
esquema (préstese especial cuidado a la puntuación): 

Dendrodoris limbata (Cuvier, 1804) 

Sinonimias 

Doris limbata Cuvier, 1804, Ann. Mus. Hist. Nat. Paris, 4 (24): 468-469 [Localidad tipo: Marsella]. 

Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop.-Car., 10: 275. 

Dichas referencias no deberán incluirse en la lista de Bibliografía si es la única vez que se nombran en el texto. 
Si se incluyen una lista completa de referencias de un taxón inmediatamente tras éste, deberá seguirse el 
mismo esquema (sin incluir en Bibliografía las referencias que no se mencionen en otro lugar del texto). 

e Sólo los nombres en latín y los de taxones genéricos y específicos deberán llevar subrayado sencillo o prefe- 
rentemente ir en cursiva. En ningún caso deberá escribirse una palabra totalmente en letras mayúsculas, ni 
siquiera el Título. Las unidades a utilizar deberán pertenecer al Sistema Métrico Decimal, junto con sus correc- 
tas abreviaturas. En artículos escritos en castellano, en los números decimales sepárese la parte entera de la 
decimal por una coma inferior (,), NUNCA por un punto (.) o coma superior ('). 

e Las referencias bibliográficas irán en el texto con minúsculas o versalitas: Fretter y Graham (1962) o FRETTER 
Y GRAHAM (1962). Si son más de dos autores se deberán citar todos la primera vez que aparecen en el texto 
[Smith, Jones y Brown (1970)] empleándose et al. las siguientes veces [Smith et al. (1970)]. Si un autor ha publi- 
cado más de un trabajo en un año se citarán con letras: (Davis, 1989a; Davis, 1989b). No deberá emplearse op. 
cit. La lista de referencias deberá incluir todas las citas del texto y sólo éstas, ordenadas alfabéticamente. Se 
citarán los nombres de todos los autores de cada referencia, sea cual sea su número. Los nombres de los 
autores deberán escribirse, en letras minúsculas o Versalitas. No deberán incluirse referencias a autores 
cuando éstos aparezcan en el texto exclusivamente como autoridades de un taxón. Los nombres de las publica- 


ciones periódicas deberán aparecer COMPLETOS, no abreviados. Cuando se citen libros, dése el título, editor, 
lugar de publicación, n” de edición si no es la primera y número total de páginas. Deberán evitarse referencias 
a Tesis Doctorales u otros documentos inéditos de difícil consulta. Síganse los siguientes ejemplos (préstese 
atención a la puntuación): 

Fretter V. y Graham A. 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp. 

Ponder W.F. 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. En Ponder, W.F. 
(Ed.): Prosobranch Phylogeny. Malacological Review, suppl. 4: 129-166. 

Ros J. 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Mis- 
celánea Zoológica, 3 (5): 21-51. 

e Las gráficas e ilustraciones deberán ser originales y presentarse preferentemente en formato electrónico al 
formato de caja de la revista o proporcional a éste. Este formato es de 57 mm (una columna) o 121 mm (dos) de 
anchura y hasta 196 mm de altura, si bien se recomienda utilizar el formato a dos columnas. Es importante que 
todas las figuras sean remitidas en su formato original (por ejemplo, las fotografías en .j¡pg de alta calidad o .tif, 
las gráficas en hojas de cálculo Excel o documentos de CorelDraw), puesto que las ilustraciones insertadas en 
el manuscrito WORD son inservibles en la fase de imprenta. Las imágenes digitales deben ser formateadas en 
su tamaño de impresión con una resolución mínima de 300 ppp para imágenes en color o escala de grises y de 
600 ppp para las de blanco y negro. Considérese la reducción que será necesaria a la hora de decidir el tamaño 
de las escalas o letras en las figuras, que no deberán bajar de los 2 mm. En figuras compuestas, cada parte 
deberá etiquetarse con letras mayúsculas, el resto de las letras deberán ser minúsculas. Las escalas de dibujos y 
fotografías deberán ser gráficas, utilizando unidades del sistema métrico decimal; no deberán hacerse referen- 
cias a los aumentos de una determinada ilustración, ya que éstos cambian con la reducción. En su caso, se 
recomienda la utilización de mapas con proyección UTM. Cada figura, gráfica o ilustración deberá presentarse 
en hojas separadas y con numeración arábiga (1, 2, 3,...), sin separar “Figuras” y “Láminas”. Los pies de 
figura, en una hoja aparte, deberán acompañarse de su traducción al inglés (o español si el inglés es la lengua 
del trabajo). Utilicese el esquema siguiente: 

Figura 1. Neodoris carvi. A: animal desplazándose; B: detalle de un rinóforo; C: branquia. 

Las abreviaturas empleadas en las ilustraciones deberán incluirse en los pies de figura. 

Los autores interesados en incluir láminas en color deberán consultar con el editor y sufragar el sobrecoste con 
una contribución de 30 € por página. Por lo demás, deberán ajustarse a los mismos requisitos indicados para 
las figuras. 

Si se pretende enviar gráficas o ilustraciones en impresión de papel es imprescindible presentar originales de 
buena calidad. Las imágenes en semitonos deben estar bien contrastadas y ajustarse al tamaño definitivo de 
impresión; al componer fotografías sobre una hoja, procúrese que los espacios entre ellas sean regulares y que 
estén debidamente alineadas. Téngase en cuenta que incluir fotografías de distinto contraste en una misma 
página conlleva una pobre reproducción final. Las gráficas de ordenador deberán imprimirse con impresora 
láser sobre papel de buena calidad. 

e Las Tablas se presentarán en hojas separadas, siempre con numeración romana (1, II, II1...). Las leyendas se 
incluirán en una hoja aparte acompañándose de una traducción al inglés. Deberán evitarse las tablas particu- 
larmente complejas. Se recomienda reducir el número y extensión de ilustraciones, láminas o tablas al mínimo 
necesario. 

e Los artículos que no se ajusten a las normas de publicación serán devueltos al autor con las indicaciones de 
los cambios necesarios. 

e El Comité Editorial comunicará al autor responsable del trabajo la fecha de recepción del trabajo y la fecha 
de envío a revisión. Cada original recibido será sometido a revisión por al menos dos investigadores. El 
Comité Editorial, a la vista de los informes de los revisores decidirá sobre la aceptación o no de cada manus- 
crito. El autor recibirá en cada caso copia de los comentarios de los revisores sobre su artículo. En caso de 
aceptación, el mismo Comité Editorial, si lo considera conveniente, podrá solicitar a los autores otras modifica- 
ciones que considere oportunas. Si el trabajo es aceptado, el autor deberá enviar una copia impresa del mismo 
corregida, acompañada por una versión en un CD, utilizando procesadores de texto en sus versiones corrien- 
tes de Windows (pero no el formato .docx generado por Word 2007, el habitual de Windows Vista) o Macin- 
tosh. La fecha de aceptación figurará en el artículo publicado. 

e Las pruebas de imprenta serán enviadas por correo electrónico al autor responsable, exclusivamente para la 
corrección de erratas, y deberán ser devueltas en un plazo máximo de una semana. Se recomienda prestar 
especial atención en la corrección de las pruebas. 

e De cada trabajo se entregarán gratuitamente 50 separatas, además de una versión electrónica en formato 
.pdf. Aquellos autores que deseen un número mayor, deberán hacerlo constar al devolver las pruebas de 
imprenta, y nunca posteriormente. El coste de las separatas adicionales será cargado al autor. 


INSTRUCTIONS TO AUTHORS 


e Iberus publishes research papers, notes and monographs devoted to the various aspects of Malacology. 
Papers are manuscripts of more than 5 typed pages, including figures and tables. Notes are shorter papers. 
Monographs should exceed 50 pages of the final periodical, and may be published as Supplements. Authors 
wishing to publish monographs should contact the Editor. Manuscripts are considered on the understanding 
that their contents have not been published or simultaneously submitted for publication elsewhere. 

+ Manuscripts and correspondence regarding editorial matters must be sent to: Serge Gofas, Editor de Publica- 
ciones, Departamento de Biología Animal, Universidad de Málaga, Campus de Teatinos, s/n, 29071, Málaga, 
Spain and / or to the e-mail <sgofasQuma.es>. 

e Manuscripts may be written in Spanish, English, Italian, French or Portuguese. 

e Manuscripts must be typed double spaced (including the references, figure captions and tables) on one side 
on A-4 (297x210 mm) with margins of at least 3 cm. An original and two copies must be submitted, together 
with a CD or e-mail remittance containing the article written with a current Windows (but not .docx format 
generated by Word 2007, mainly used with Windows Vista) or Macintosh word processor. When a paper has 
joint authorship, one author must accept responsibility for all correspondence. 

e The authors must include a list of at least 4 possible referees; the Editor can choose any others if appropriate. 

e Papers should conform the following layout: 

First page. This must include a concise but informative title, with mention of family of higher taxon when 
appropriate, and its Spanish translation. It will be followed by all authors” names and surnames, their full 
address(es), an abstract (and its Spanish translation) not exceeding 200 words which summarizes not only con- 
tents but results and conclusions. 

Following pages. These should content the rest of the paper, divided into sections under short headings. When- 
ever possible the text should be arranged as follows: Introduction, Material and methods, Results, Discussion, 
Conclusions, Acknowledgements and References. Unusual abbreviations used in the text must be grouped in 
one alphabetic sequence after the Material and methods section. 

e Notes should follow the same layout, without the abstract. 

e Footnotes and cross-references must be avoided. The International Codes of Zoological and Botanical 
Nomenclature must be strictly followed. The first mention in the text of any taxon must be followed by its 
authority including the year. In systematic papers, when synonyms of a taxon are given, they must be cited IN 
FULL, including the periodical, in an abbreviate form, where they were described, and the type localities in 
square brackets when known. Follow this example (please note the punctuation): 

Dendrodoris limbata (Cuvier, 1804) 

Synonyms 

Doris limbata Cuvier, 1804, Ann. Mus. Hist. Nat. 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. If a 
full list of references of the taxon is to be given immediately below it, the same layout should be followed (also 
excluding from the Bibliography list those which are not cited elsewhere). 

Only Latin words and names of genera and species should be underlined once or be given in italics. No word 
must be written in UPPER CASE LETTERS. SI units are to be used, together with their appropriate symbols. In 
Spanish manuscripts, decimal numbers must be separated with a comma (,), NEVER with a point (.) or upper 
comma ('). 

e References in the text should be written in small letters or Small capitals: Fretter and Graham (1962) or 
FRETTER AND GRAHAM (1962). The first mention in the text of a paper with more than two authors must 
include all of them [Smith, Jones and Brown (1970)], thereafter use et al. [Smith et al. (1970)]. If an author has 
published more than one paper per year, refer to them with letters: (Davis, 1989a; Davis, 1989b). Avoid op. cit. 

The references in the reference list should be in alphabetical order and 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 tax- 
onomic name. Titles of periodicals must be given IN FULL, not abbreviated. For books, give the title, name of 
publisher, place of publication, indication of edition if not the first and total number of pages. Keep references 
to doctoral theses or any other unpublished documents to an absolute minimum. See the following examples 
(please note the punctuation): 

Fretter V. and Graham A. 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp. 

Ponder W.F. 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. In Ponder W.F. 
(Ed.): Prosobranch Phylogeny. Malacological Review, suppl. 4: 129-166. 

Ros J. 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Mis- 
celánea Zoológica, 3 (5): 21-51. 


e Figures must be original and provided preferably in electronic format and adjusted to page format and 
column size. These should be one column (57 mm) or two columns (121 mm) wide and up 196 mm high, or be 
proportional to these sizes. Two columns format is recommended. It is essential that all figures be supplied in 
their original format (e.g. photographs as high-grade .jpg or as .tif files, graphs as Excel spreadsheets or Corel- 
Draw files), as the files inserted into WORD documents cannot be used for printing. Digital images must be 
given their final printing size with a resolution at least 300 dpi for colour and halftones, and at least 600 dpi for 
black / white. 

Take into account possible reduction in lettering drawings; final lettering must be at least 2 mm high. In com- 
posite drawings, each figure should be given a capital letter; additional lettering should be in lower-case 
letters. A scale line, labelled with SI units, must be used to indicate size; magnification ratio must be avoided 
as it may be changed during printing. UTM maps are to be used if necessary. Figures must be submitted on 
separate sheets, and numbered with consecutive Arabic numbers (1, 2, 3,...), without separating “Plates” and 
“Figures”. Legends for Figures must be typed in numerical order on a separate sheet, and a Spanish translation 
must be included. Follow this example (please note the punctuation): 

Figure 1. Neodoris carvi. A: animal crawling; B: rhinophore; C: gills. 

If abbreviations are to be used in illustrations, they should be included in the figure captions. 

Authors wishing to publish illustrations in colour should consult with the editor and will be charged a contri- 
bution of 30 € per page. They should otherwise follow the same standards as black and white prints. 

If the authors want to send Figures in printed format, it is essential to supply good quality originals. Half-tone 
images must be of good contrast, and should be submitted in the final printing size. When mounting pho- 
tographs in a block, ensure spacers are of uniform width. Remember that grouping photographs of varied con- 
trast results in poor reproduction. Computer graphics must be printed on high quality white paper with a 
laser printer. 

e Tables must be numbered with Roman numbers (1, IL, TIL...) and each typed on a separate sheet. Headings 
should be typed on a separate sheet, together with their English translation. Complex tables should be 
avoided. As a general rule, keep the number and extension of illustrations and tables as reduced as possible. 

e Manuscripts that do not conform to these instructions will be returned for correction before reviewing. 

e Authors submitting manuscripts will receive an acknowledgement of receipt, including receipt date, and the 
date the manuscript was sent for reviewing. Each manuscript will be critically evaluated by at least two refer- 
ees. Based of these evaluations, the Editorial Board will decide on acceptance or rejection. Anyway, authors 
will receive a copy of the referees” comments. If a manuscript is accepted, the Editorial Board may indicate 
additional changes if desirable. Acceptable manuscripts will be returned to the author for consideration of 
comments and criticism; a finalized manuscript must then be returned to the Editor, together with a CD con- 
taining the article written with current Windows (but not a .docx format generated by Word 2007, mainly used 
under Windows Vista) or Macintosh word processor. Dates of reception and acceptance of the manuscript will 
appear in all published articles. 

e Proofs will be e-mailed to the author for correcting errors and must be returned corrected within one week. 
At this stage no stylistic changes will be accepted. Pay special attention to references and their dates in the text 
and the Bibliography section, and also to numbers of Figures and Tables appearing in the text. 

e Fifty reprints per article and a .pdf file will be supplied free of charge. Additional reprints must be ordered 
when the page proofs are returned, and will be charged at cost. NO LATER orders will be accepted. 


La SOCIEDAD ESPAÑOLA DE MALACOLOGÍA 


Junta Directiva desde el 11 de octubre de 2005 


Presidente José Templado González 

Vicepresidente Emilio Rolán Mosquera 

Secretaria María Carmen Salas Casanovas 

Tesorero Luis Murillo Guillén 

Editor de Publicaciones Serge Gofas 

Bibliotecario Rafael Araujo Armero 

Vocales Ramon M. Álvarez Halcon 
Benjamín Gómez Moliner 
Alberto Martínez Ortí 


Diego Moreno Lampreave 
José Ramón Arrébola Burgos 


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 2011: 


Socio numerario (en España): 40 euros 
(en Europa) 40 euros 
(fuera de Europa): 48 euros 
Socio estudiante (en España): 23 euros 
(en el extranjero): 29 euros 
Socio Familiar: (sin recepcion de revista) 4euros 
Socio Protector: (mínimo) 48 euros 
Socio Corporativo (en Europa): 48 euros 
(fuera de Europa): 54 euros 


INSCRIPCIÓN: 6 euros, además de la cuota correspondiente. 


A los socios residentes en España se les aconseja domiciliar su cuota. Todos los abonos deberán enviarse al 
Tesorero (dirección reseñada anteriormente) el 1 de enero de cada año. Los abonos se harán sin recargos para la 
sociedad y en favor de la Sociedad Española de Malacología y no de ninguna persona de la junta directiva. Aque- 
llos socios que no abonen su cuota anual dejarán de recibir las publicaciones de la Sociedad. Los bonos de ins- 
cripción se enviarán junto con el abono de una cuota anual al Tesorero. 

A los residentes en el extranjero se les ruega que abonen su cuota mediante giro postal en euros (internatio- 
nal postal money orders in euros sent to the Treasurer). Members living in foreing countries can deduce 6 euros 


if paid before 15 April. 


Cada socio tiene derecho a recibir anualmente los números de /berus, Reseñas Malacológicas y Noticiarios que 
se publiquen. 


1 


| 


| 


wii 


ÍNDICE 
Iberus 29 (1) 2011 


SCUDERI D. E CRISCIONE E. Cima apicisbelli Rolán, 2003 (Gastropoda: Cimidae): nuovo immi- 
grante in Mediterraneo o specie autoctona? 
Cima apicisbelli Rolán, 2003 (Gastropoda: Cimidae): ¿nuevo inmigrante en el Mediterraneo 0 
ESPECIE AULOCIORAS A A A A EN 1-7 
OLIVER J.D. AND ROLÁN E. The family Tornidae (Gastropoda, Rissooidea) in the East Atlantic, 
2. Circulinae 
La familia Tornidae (Gastropoda, Rissovidea) en el Atlántico oriental, 2. Circulinae .. 9-33 
CAMPANI E., BOGI C. AND GALIL B.S. First Mediterranean record of Diaphana marshall (Sykes, 
1904) (Gastropoda, Diaphanidae) - a rare bathyal species 
Primera cita para el Mediterráneo de Diaphana marshalli (Sykes, 1904) (Gastropoda, Diap- 


PEMAAE) UNA ESPECIE TIT A 35-38 
DE MArTTIA W. AND MAscIiaA E Otala punctata (O.E Miller, 1774) (Stylommatophora: Helicidae) 

in Italy 

Otala punctata (O.E Múlter, 1774) (Stylommatophora: Helicidae) en Italia ....... 39-46 


TIRADO C., RUEDA J.L. AND CARMEN SALAS C. Reproduction of Donax trunculus in the littoral 
of Huelva (southern Atlantic Spain): is there any difference with the Mediterranean popu- 
lation from the Andalusian coast? 

Reproducción de Donax trunculus en el litoral de Huelva (suroeste Atlántico de España): ¿hay 
diferencias con la población mediterránea de la costa andaluza? ico 47-57 

MORENO D. Bittium nanum (Gastropoda, Cerithiidae), una especie válida de las islas Azores 

Bittium nanum (Gastropoda, Cerithiidae), a valid species from the Azores Islands .... 59-74 


ISSN 0212-3010 


UTION LIBRARIES 


UN 


619 4623 


] berus 


mor 
aos 


INVE REVISTA DE LA 
SOCIEDAD ESPAÑOLA 
DE MALACOLOGIA 


Vol. 29 (2) 


| Vigo, diciembre 201 1 


Iberus 
Revista de la 


SOCIEDAD ESPAÑOLA DE MALACOLOGÍA 


Comité DE ReDAcciÓN (BOARD OF EDITORS) 
EDITOR DE PUBLICACIONES (EDITOR=IN=CHIEF) 


Serge Gofas 


Universidad de Málaga, España 


DIRECTOR DE REDACCIÓN (EXECUTIVE EDITOR) 


Gonzalo Rodríguez Casero 


Mieres del Camino, Asturias, España 


EDITORA EJECUTIVA (MANAGING EDITOR) 


Eugenia M* Martínez Cueto-Felgueroso 


Mieres del Camino, Asturias, España 


EDITORES ADJUNTOS (ASSOCIATE EDITORS) 


Francisco Javier Conde de Saro 
Benjamín Gómez Moliner 
Angel Antonio Luque del Villar 
Emilio Rolán Mosquera 

José Templado González 

Jesús S. Troncoso 


Embajada de España, Japón 

Universidad del País Vasco, Vitoria, España 

Universidad Autónoma de Madrid, Madrid, España 
Universidad de Vigo, Vigo, España 

Museo Nacional de Ciencias Naturales, CSIC, Madrid, España 
Universidad de Vigo, Vigo, España 


Comité EDITORIAL (BOARD OF REVIEWERS) 


Kepa Altonaga Sustacha 
Eduardo Angulo Pinedo 
Rafael Araujo Armero 

Thierry Bockeljau 

Ridiger Bieler 

Sigurd v. Boletzky 

Jose Castillejo Murillo 

Karl Edlinger 

Antonio M. de Frias Martins 
José Carlos García Gómez 
Gonzalo Giribet de Sebastián 
Edmund Gittenberger 

Angel Guerra Sierra 

Gerhard Haszprunar 

Yuri |. Kantor 

María Yolanda Manga González 
Jordi Martinell Callico 

Ron K. 0'Dor 

Takashi Okutani 

Marco Oliverio 

Pablo E. Penchaszadeh 
Winston E. Ponder 

Carlos Enrique Prieto Sierra 
MY de los Angeles Ramos Sánchez 
Francisco Javier Rocha Valdés 
Paul 6. Rodhouse 
Joandoménec Ros ¡ Aragones 
María Carmen Salas Casanovas 
Gerhard Steiner 

Victoriano Urgorri Carrasco 
Anders Warén 


PORTADA DE Jberus 


Universidad del País 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 

Harvard University, EE.UU. 

Notional Natuurhistorisch Museum, Leiden, Holanda 

Instituto de Investigaciones Marinos, CSIC, Vigo, España 

Zoologische Staatssammlung Múnchen, Múnchen, Alemania 

AN. Severtzov Institute of Ecology and Evolution, Moscú, Rusia 
Estación Agrícola Experimental, CSIC, León, España 

Universidad de Barcelona, Barcelona, España 

Dalhousie University, Halifax, Canada 

Nihon University, Fujisawa City, Japón 

Universitá di Roma “La Sapienza”, Roma, Italia 

Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina 
Australian Museum, Sydney, Australia 

Universidad del Poís Vasco, Bilbao, España 

Museo Nacional de Ciencias Naturales, CSIC, Madrid, España 
Instituto de Investigaciones Marinas, CSIC, Vigo, España 

British Antarctic Survey, Cambridge, Reino Unido 

Universidad de Barcelona, Barcelona, España 

Universidad de Málaga, Málaga, España 

Institut fúr Zoologie der Universitút Wien, Viena, Austria 

Universidad de Santiago de Compostela, Santiago de Compostela, España 
Swedish Museum of Natural History, Estocolmo, Suecia 


Iberus gualtieranus (Linnaeus, 1758), una especie emblemática de la península Ibérica, que da 
nombre a la revista. Dibujo realizado por José Luis González Rebollar “Toza”. 


Iberus 


REVISTA DE LA 
SOCIEDAD ESPAÑOLA 
DE MALACOLOGÍA 


Vol. 29 (2) Vigo, diciembre 201 1 


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: Serge Gofas, Editor de Publicaciones, Departamento de Bio- 
logía Animal, Universidad de Málaga, Campus de Teatinos, s/n, 29071, Málaga, España. 

Los trabajos se entregarán por triplicado (original y dos copias). Se recomienda a los autores leer 
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, P.O. Box 321, 
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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 


THE FAMILY TORNIDAE (GASTROPODA, 
RISSOOIDEA) IN THE CARIBBEAN AND 
NEIGHBORING AREAS 


LA FAMILIA TORNIDAE (GASTROPODA, RISSOOIDEA) 


EN EL CARIBE Y ÁREAS VECINAS 


Federico RUBIO 
Pintor Ribera, 4-16* 
46930 Quart de Poblet 
(Valencia), Spain 
e-mail: federubioono.com 


Raúl FERNÁNDEZ-GARCÉS 
Centro de Estudios Ambientales de Cienfuegos (CEAC), 
División de Gestión Ambiental (DGA), 
calle 17, esquina Ave. 46, 
Cienfuegos, Cuba 


$ 


Emilio ROLÁN 
Museo de Historia Natural, 
Campus Universitario Sur, 
15782, Santiago de Compostela, Spain 
e-mail: erolanCemiliorolan.com 


INDEX OF SPECIES: 


Iberus 29 (2) 2011 


PLATES 


BN 


FAMILY TORNIDAE 
SPECIES PAGE 
GENUSCIRECULTS 
Cireulus semaisculptus (Olsson $e MeGinty, 1958) connaccciiicinccnociconioo cdta idas 6 
SAS SS IL A A 8 
Erreulusitatus (Ad Mea once iobacaradcdaa riada 10 
Circula stexamis ((MOOre, LODO) .astio acto sincadon ia cionaadocaco rodeado icrabaciiiod 11 
GENUS TEINOSTOMA 
VEAS rra lA A A 15 
Temostoma contosyrus Pilsbry Se MeGinty, 1945 cecccooninnonianiorocanesccnsinacsonónass 16 
Ternostomolenteculare EL €. Eear Sho) cuota dioadasiooinibecrecn caia 19 
Terrostornia seclisun (Dal. LO tool iici ados id Sig icióósn 21 
MEA ERAN TAS NA 24 
Temostoma incertum. Pilsbry.£ MeGinty, TIAS vavecionscscoaninccasonodniaconccuens donó 24 
Teinostoma anastomosis spec. nov. Rubio, Rolán 8 Le€ .....ooconionicnonicnconcnnos 27 
Teinostoma panamense spec. nov. Rubio, Rolán é Lee..........cononcnonmo.*... 29 
Teinostoma biscaynense Pilsbry 6 McGinty, 1945 .oooocccccncciccnnoononnnnnnnanicnnonos 29 
Ternostomaobteciun Pilsbry (E MEGINIEy, LIA caocoraccnmiaiosiniocn serian 33 
MAA E ES AS O A AS 34 
lenostomamesación PilsbiSMecGinty, LOAD ccotcneconnorionoien caso ea pirineos 36 
Temmostoma semistriatumtd Orbleny, 1842) occiso nionicoiamaiasaciiós 38 
Termostomto miau scada SI decido noni 41 
Jemmostomalderema Pisbry e MeGiiy II tapones tancia 43 
Temostoma mbilicatunm (El: Lea, 143 e ccanosanacotaninonnitocccal cie saordanenacóan 44 
MR a AS A 46 
Memos tonta soma un (Dal in ena adan a carios 48 
MS A A A 50 
Temostonta cocolitoris Pilsbry E MeGinty, TIAS e ccccociania rdancenecininn rincon 50 
TesostontaMeliciiun pe proto incio oline 53 
TemostomamesasiomaxEb: Adams, 1850 urinarias 54 
emos toma cienjuegosense Spec MOV ose cistuiocidsatass siaancrosidoso soba edocbe sica 56 
Temostoma parvicallum Pilsbry $ MeGinty, 19D escnorrtoninciciniancacinninniónicanenas 9 
Temostomvayme saca lumispec:MO Ve ss oooicasaio colono ciencia iento cotas 99 
Temmostoma carimicallusilalsbiy E MeGinty, LIO ccotovonodiccincnandcococccaco natación 61 
Temostoma lituspalmarum Pilsbry € MeGinty, 194 vncncoiicnnionoscinnineneriianess 61 
GENUS TORNUS 
Tornusicaraboboensis (Welsbord, 1962) 0 cridncncnncconiolon cito endodancodnonacaino seas 63 
MISS A E O A ENE 64 
GENUS ANTICLIMAX 

Annolmaricrass tabs (Aguayo: Se BOLLO, LIO) macrotcntoncan cucuta 68 
cas pec nov. Rubio, Rola de Pelorce iaiciinaenciooeeiciosonines 70 
Anticlimax decorata Rolán, Fernández-Garcés éz Rubio, 199 inc... 70 


Sl) Anticiimazx provosciad ca a e 73 
38 ARECIÓMAPIS YM 75 
39 Antcrmax lock Olsson A 74 
40 Ancla aaa cias O O A 76 
41 Anticlimax hispantolensis cratera Pilsbry €: Olsson, 1950 cocooccnicnicnicnncinn... IS 
GENUS AOROTREMA 
42 Aro tema stronum DAMAS A o ÓN 80 
Aorotrema pontogenes (Schwengel €: MeGinty, 1942) ..ooocoocciciccociciciccocacannos 81 
GENUS CYCLOSTREMISCUS 
43 Cyclostremiscus beans eS 84 
44 Cyclostremiscus multiliratus spec. nov. Rubio, Rolán éz Garcia ................. 86 
45 Eyclostrenmiscus dal Bus 88 
46 Eyclostremiscuspentasonas Ci 88 
47 Eyclostrentiscus al US 91 
48 Eyelostreniscus te O 93 
409 Cyelostrentiscus DaniscaMans el O 95 
50 Eyclostremiscus jeanaae lso E MEGA 97 
Sl Euclostremiscus ss uppressus DM A 97 
52 Cyclostremiscus vanbruggent de Jong $: Coomans, 1988 ....ooconcoccccciccconooo. LOO 
d9 Cyclostremiscus diminutus spec. nov. Rubio, Rolán éz Pelorce ................ 100 
54 Cyclostremiscus cubanus (Pilsbry € Aguayo, 1933 )tc.acccconncaconoioinononcion: 103 
3) Eyclostremiscus eu oy pls cuy OS BOO MA eta aio 105 
56 Cyclostremiscus microstriatus spec. nov. Rubio, Rolán € Lee.................. 105 
GENUS COCHLIOLEPIS 
57 Cochliolepis adams NA e 0 108 
58 Cochtiolepas RAM 109 
59 Cochttolepis nautltormisiClolmes 1 ei 1012 
6061762 Cochlholepts paras O a on 110 
63 Cochliolepss striata Da neto o naaa, Ao 118 
64 Cochilolepis PU A 118 
65 Cochliolepis patriciorspec, nov. Rubio, Rolán S Lee coacciones sonetos 120 
66 Cochliolepis diferens spec. nov. Rubio, Rolán € Leen 125 
67 Coco an ace tol e DE 123 
GENUS EPISCYNIA 
68/69. Episcyia mona A e ed 126 
GENUS PARVITURBOIDES 
7071 Baroiturbordes mteruptus CEBA 129 
GENUS PLEUROMALAXIS 
Ja Plewromalaxis balest (Búisbiy MCG a a 133 
GENUS SOLARIORBIS 
NS) solariornbis antillensiside Jon: E CIMA ocaso 135 
73 SOLARIS AMIA 135 
74 Solamorbisblaken (Render 14 A O 187 
7) Solarntorbis elezans (Olsson EME 139 
705177. Solarionbisiamacarimatus (CA 139 
78 Solamorbismoorcanas (Y ON 143 


79 Solaronbistimultistaatus A. Es Vernill. LO dc ot 145 
80 Solarmiorbis incas Gabba acen 145 
81 Solaiorbiss clio Vanatta LLO) ni tasas 1147 
82 SOLOMON DIS isemipunacios Nlo0te IO DE oconreoconanoicicncioioe ciencias ileso besaacs 149 
83, 84 Solariorbisterminalis(Bilsbry Se MeGinty 1946) conoccaconi nic idiccacentestano 150 
8) AS A 153 
86 AS A A 159 
86 TI A A A A A 159 
87 Solariorbis punctostriatus spec. nov. Rubio, Rolán dz Le€...oooonconinninnnnn..... 157 
Ll ASIS EAN A A A 159 
GENUS VITRINELLA 
90 Vitimella annelñtesge de Jong ECoomans, IO iii enatoterualon sondin cen nennenisinicnes 162 
91 Witinella comeracia (Vanaltta, LI analice 164 
92 MAT Us En Dela SAR A A 165 
93 inem suelos (Dal ISI ct o AÍ  E diota 167 
94,95 Witnella relico idea Cb. Adams LSO scada iaa aiodatados 167 
96 Vitnella tlordara Pisbry E MeGinty, 1I4O ceocconasenentoóniosionincotaneocitcadadoio 71 
97 Vitanet canalla SDE MON ooo tcteces leonado ioneinio is iadd asin 173 
98 A E RO 173 
99 Virinella pseudo ais tata Spec Vulcan lectoras aaineidoo soso 176 
100 A 176 
101 E 178 
102 NE E e 181 
103 Wiimnela cuptdinensis Altena, TIO sapicacaccnaciin ion donostia in etade latas 181 
104 Vitanella fuera Risbry E MeGinty, LORO caicinicnonccincocicninoecaciocasd notado ccinaós 184 
105 VE A 186 
GENUS VITRINORBIS 
106 Vitrinorbiselecans Olsson E MeoGinty 199 ecscacicinosnconiiacacanocraconniicnsesrnecnss 186 
OTHER SPECIES 
SPECIES NOT REPRESENTED IN OUR MATERIAL 
Amtelimas senumot(Vanattas LIS lr adiaietiiiadaccidoniiacids 189 
Eochttolepis surimamensis Altena, LIO me cvcnarcinionecionandbadeccn coa sesacccacos iaa añendo 189 
Discopists.omalos (olla ASTON coito dote nta iaconrmaoncennconneedid soldier dns 189 
Pleuromalaxis paul Olsson € MeGinty, LODOS. oc icncosinonoiiccniacoiatindososascncipano 190 
solariorbis hondurasensis (Vanatta TIO coacitanncat irianan di nidriincinacndas 190 
107 Solarorbis petita (Batischer LEÍ) coiecoosennenitocencente neo caresiancns rodando sesimpaerisins 190 
Temostona avunculusLlsbiy LIDO aries said 191 
Ternostomamnonierer Joussearme 1972 conncconoonconatonsnonocconno rado roo netas csosenenenens 191 
Temostoma paroumilSamapson LS). ccunsiracaoalo cnn artesslrarasotacósonaisdias 192 
108 Vitrenellaanomala (Orsay SA) Dieron concaciononinnionretano des canaiacatin se dercadiadono 192 
Vienmeltarcarmara (diOibieny 142 arrancan ocasionaron cri ndnpesepegetonos 192 
Vine nea pt anatta Ll roo ioio rai ccndieos dino acageno sreeesosia ries 192 
036 109 Viirinela Dust (ll Bterter SO) icon necneonioo dico cnecandeneno enanos semen 193 
A O A AA EE NS A 195 


FOSSILS OR SUPPOSEDLY FOSSIL SPECIES, NOT RECENT 
Antclnaxathleenae (Pilsbry E MeGinty, TI4O) uncccincncacancann cion cocoa aer 195 


110 


pill 


112 
(US; 


114 
114 
ls 


116 
116 
17 
118 
119 
120 


v 


A0H0 Henao 197 


Eyclosteniscus 197 
EyclostremiscusioumteriMastieid OS 197 
Eyclostremiscusolss on 197 
Parorturbomides ate 197 
Solariorbis.eugenes alsbry LIS ao 197 
Termostomiacaloosaense DM 199 
Temostoma tectispua ld eso Eo 199 
SPECIES FROM NEARBY OR RELATIVELY NEARBY AREAS 

Neusas marshall (ye oie iS 199 
Pond ernnellaxacrtaba Absalao, 2000 a O 200 
Ternostomadabnormie EA Sai de 200 
SPECIES RECORDED ERRONEOUSLY FROM THE CARIBBEAN 

Adeo rise O 201 
Vitirinelare ita 201 

SUPPLEMENT 


DUBIOUS SPECIES, APPARENTLY NOT TORNIDAE 


A0rotrema erraticumblsbiy E Me A atea cia coSS 202 
Vitrinellatinecta (<bB Adams LO E dee Ed EU 202 
«Vitrimella carte Da O an Doo aa coa e asada 202 
Vitrinella cyclostomordes (eii 203 
Vitrnella ayala Ec toa 204 


VITRINELLID SPECIES TRANSFERRED TO SUBFA MILY 


SKENEINAE IN THIS WORK 
FAMILY TURBINIDAE 
Ernsonella ondas MD o ato 205 
Ensonela ceo rana DL 207 
Erxsonella murio MD O 207 
Miro certo DEA SI 208 
Ilse y aio 210 
Xy lOs encata MD 21 
VWitraneltasmassarta MD IL OD LS 2D 


SPECIES THAT WERE INCLUDED IN TORNIDAE AND 
NOW BELONG TO OTHER FAMILIES 


FAMILY TURBINIDAE 
Ganesa proxima Uryon, 1990 reseca cocoa seno Ledo aNnd iba Edo cola 215 
Ganesa vushae Dal LI esoo a eo Na OSA Zo 
Ganesa conica Da TIZT ta le rai odo a dee SUBO oooO DS 
Conesa ers a el AS 
Ganesa valvata DIM esla RETOS 215 
Ganesa diaprana AE. Verrill 1er conato leona sao es ZO 
Canesa stat BS ee 215 


EAN US E KA A LEO RO 215 


(AT es ZAS 
Canesa Ona Aa E odo 215 
CURE aaa IAE NV ii ioneaaad A ko) 
Ganesarabrss cola Bus Io enteraste cibbib tados 215 
AD A e On 216 
EI AD A A A AN 216 
Cranioyrasspunulosa Bush, 1 lili tease abetos 216 
12 Ditwornellamodesta DASS) strains coda asco nidad re decis liliana 216 
Molero pss abyssicola Bush, 1 codi oneiasicdalds 216 
IMoNeriopsis scera Da IS o decidiendo dpto 216 
AA E A US A o A A 216 
A a AS A O 216 
ED O A 216 
FAMILY LEPETELLIDAE 
E a A 217 
a A A 217 
FAMILY ODOSTOMUDAE 
Envclostremelatóntds Bush, 197 coo acotaciones 217 


vil 


OA 


O Sociedad Española de Malacología —__—— Iberus, 29 (2):1-240 , 2011 


The family Tornidae (Gastropoda, Rissooidea) in the 
Caribbean and neighboring areas 


La familia Tornidae (Gastropoda, Rissooidea) en el Caribe y áreas 
vecinas 


Federico RUBIO*, Raúl FERNÁNDEZ-GARCÉS** and Emilio ROLÁN*** 


Recibido el 28.04.2011. Aceptado el 26.09.2011 


RESUMEN 


Se estudian las especies del Caribe, incluidas en la familia Tornidae y que comprenden 
los géneros Circulus, Teinostoma, Tornus, Anticlimax, Aorotrema, Cyclostremiscus, Cochlio- 
lepis, Episcynia, Parviturboides, Pleuromalaxis, Solariorbis, Vitrinella y Vitrinorbis, sobre la 
base de unos 2700 ejemplares y conchas examinados. Los taxones estudiados de Torni- 
dae son 101 en total, siendo 4 del género Circulus, 27 de Teinostoma, 2 de Tornus, 8 de 
Anticlimax, 2 de Aorotrema, 14 de Cyclostremiscus, 9 de Cochliolepis, 1 de Episcynia, 1 
de Parviturboides, 1 de Pleuromalaxis, 16 de Solariorbis, 15 de Vitrinella y 1 de Vitrinor- 
bis. De estas especies 23 son nuevas para la ciencia y se describen aquí: 8 en Teinos- 
toma, 1 en Anticlimax, 3 en Cyclostremiscus, 3 en Cochliolepis, 4 en Solariorbis y 4 en 
Vitrinella. Se discuten las asignaciones a estos géneros y sus relaciones. Además, se apor- 
tan datos sobre 40 especies más, relacionadas en algún momento con la familia en estu- 
dio. Un nuevo nombre se propone para “Cyclostrema” thomasi Pilsbry, 1945, previa- 
mente ocupado. 

Teinostoma nessaeum y leinostoma obtectum son tratados como especies válidas, no sinó- 
nimos de leinostoma biscaynense; Cyclostremiscus trilix no es sinónimo de Cyclostremis- 
cus pentagonus; Cyclostrema thomasi no es sinónimo de Vitrinella filifera. Por el contrario, 
Teinostoma clavium es aquí considerada como sinónimo de Teinostoma semistriatum. 

Se designan lectotipos para Teinostoma reclusum, Teinostoma solidum, Parviturboides inte- 
rruptus, Solariorbis petitii, Episcynia inornata y Cochliolepis parasitica. 

Se designan neotipos para leinostoma megastoma, Teinostoma semistriatum y Circulus 
orbignyi. 

Algunos tipos de las colecciones Dall y KJ Bush, todos en USNM, se fotografiaron por pri- 
mera vez por SEM. 

Del examen de los tipos de Dalt (1927), llegamos a la conclusión de que un número de 
especies descritas en el mismo no son tornidos y sí skeneidos incluidos en los géneros Cir- 
sonella, Micro y Xyloskenea. 


ABSTRACT 


The Caribbean species included within the family Tornidae in the genera Circulus, Teinos- 
toma, Tornus, Anticlimax, Aorotrema, Cyclostremiscus, Cochliolepis, Episcynia, Parvitur- 


* Pintor Ribera, 4-16, 46930 Quart de Poblet (Valencia), Spain e-mail: federubioGono.com 

** Centro de Estudios Ambientales de Cienfuegos (CEAC), División de Gestión Ambiental (DGA), calle 17, 
esquina Ave. 46, Cienfuegos, Cuba 

*** Museo de Historia Natural, Campus Universitario Sur, 15782, Santiago de Compostela, Spain e-mail: 
erolanCemiliorolan.com 


Iberus, 29 (2), 2011 


boides, Pleuromalaxis, Solariorbis, Vitrinella and Vitrinorbis are studied, based on about 
2700 specimens and shells examined. There are a total of 101 species, including 4 in the 
genus Circulus, 27 in Teinostoma, 2 in Tornus, 8 in Anticlimax, 2 in Aorotrema, 14 in 
Cyclostremiscus, 9 in Cochliolepis, 1 in Episcynia, 1 in Parviturboides, 1 in Pleuromalaxis, 
16 in Solariorbis, 15 in Vitrinella, and 1 in Vitrinorbis. Of all these species 23 are new 
and are described here: 8 in Teinostoma, 1 in Anticlimax, 3 in Cyclostremiscus, 3 in 
Cochliolepis, 4 in Solariorbis, and 4 in Vitrinella. The assignation of these species to these 
genera and their relationships are discussed. Furthermore, some information is given on 
40 species more which have been related at times to the family studied. A new name is 
proposed for “Cyclostrema” thomasi Pilsbry, 1945, which is preoccupied. 

Teinostoma nessaeum and Teinostoma obtectum are treated as valid species, not synonyms 
of Teinostoma biscaynense; Cyclostremiscus trilix is not a synonym of Cyclostremiscus pen- 
tagonus; Cyclostrema thomasi is not a synonym of Vitrinella filifera. Conversely, Teinos- 
toma clavium is herein considered a synonym of Teinostoma semistriatum. 

Lectotypes are here designated for Teinostoma reclusum, Teinostoma solidum, Parvitur- 
boides interruptus, Solariorbis petitii, Episcynia inornata and Cochliolepis parasitica. 
Neotypes are designated for Teinostoma megastoma, Teinostoma semistriatum, and Circu- 
lus orbignyi. 

Some types from the Dall and K.J. Bush collections, all in USNM, are photographed for 
the first time using SEM. 

From the examination of Dall's (1927) types, we concluded that a number of species 
described therein are not tornids but skeneids included in the genera Cirsonella, Micro 


and Xyloskenea. 


INTRODUCTION 


The  “vitrinellids”  (Vitrinellidae 
Bush, 1897), here considered synony- 
mous with Tornidae Sacco, 1896, are 
prosobranch molluscs generally living 
in shallow waters of tropical seas. In the 
Atlantic only Tornus subcarinatus 
(Montagu, 1803) and Circulus striatus 
(Philippi, 1836) live in the marine waters 
of Western Europe but there are many 
species known from the shores of West 
Africa (ADAM é KNUDSEN, 1969; ROLÁN 
éz RUBIO, 2002; OLIVER 6 ROLÁN, 2011). 
There is no known case of anfiatlantism 
in species of the Tornidae family. 

STIMPSON (1858) described the first 
vitrinellid for the malacological fauna of 
the SE United States: Cochliolepis para- 
sitica, which was found alive under the 
scales of the giant scaly worm Poly- 
dontes lupina. GABB (1873, 1881) 
described several fossil species from the 
Miocene of Santo Domingo and the 
Pliocene of Costa Rica. Subsequently 
some of these species have been found 
living. Circulus liratus (A.E. Verrill, 
1882) was described from north of Cape 
Hatteras, North Carolina 


MOORE (1964) stated that no vit- 
rinellids have been found south of 
Trinidad. Yet Rios (1975, 1994) and Díaz 
MERLANO é PUYANA HEGEDUS (1994) 
mention several species from Brazil and 
Colombia respectively. 

The supposed “vitrinellid” recorded 
alive in deeper water off the Azores by 
CLARKE (1962): Teinostoma azorica 
Dautzenberg « Fischer, 1896, was 
recently shown to be a turbinid. lts 
radula is rhipidoglossan, and its general 
characters are coincident with those of 
the Skeneinae species of the genera 
Skenea and Dikoleps (RUBIO éz ROLÁN, 
2009). Other vitrinellids collected in 
deep water such as Circulus dalli Bush 
(618 m), found very close to the north 
coast of Little Bahama Bank, have been 
considered to be transported downs- 
lope by the currents. Teinostoma 
solidum (Dall, 1889) was found below 
the continental shelf. MookrE (1964) 
mentions seeing specimens with animal 
and operculum in natural position, but 
in his opinion they could not be vit- 
rinellids. 


RUBIO ET 41£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Most of the records of living vit- 
rinellids are between one and several 
metres deep. Among the living species 
collected by PiLSBRY 2 MCGINTY (1945a 
and 1946b) are: Vitrinella helicoidea C.B. 
Adams, Teinostoma lerema Pilsbry € 
McGinty, Teinostoma  parvicallum 
Pilsbry £ McGinty, Teinostoma carini- 
callus Pilsbry £ McGinty and Pleuroma- 
laxis balesi Pilsbry € McGinty. Other 
species collected alive: Cyclostremiscus 
pentagonus (Gabb) in sand-muddy 
bottom in Mississippi Sound; 
Cyclostremiscus suppressus (Dall) on 
Thalassia off Matheson Hammock, Bis- 
cayne Bay, less than 1 meter; Solariorbis 
mooreana (Vanatta), Solariobis infracari- 
nata (Gabb), Cyclostremiscus cubanus 
(Pilsbry € Aguayo) and Cyclostremis- 
cus pentagonus (Gabb) in sandy bottom 
and/or muddy-sandy bottom. Circulus 
striatus (Philippi), type species of the 
genus, lives on sandy bottoms off the 
Atlantic coasts of southern France and 
the Iberian Peninsula. The vitrinellids 
supposedly feed on detritus and 
diatoms. 

We have scanty information on the 
soft parts of most of the species of the 
family Tornidae Sacco, 1896; it is a het- 
erogeneous assortment of species funda- 
mentally grouped together on the basis 
of some similarity of the shells. The 
ignorance of the soft anatomy and the 
radula of most of the species described 
up to now makes a correct generic and 
subgeneric placement difficult. The only 
anatomical studies published on these 
groups correspond to the type species of 
the genera Circulus Jeffreys, 1865 [C. 
striatus (Philippi, 1836) in FRETTER 
(1956)], Cochliolepis Stimpson, 1858 [C. 
parasitica Stimpson, in MOORE, 1972], 
Tornus Turton € Kingston, 1830 [T. sub- 
carinatus (Montagu, 1803), in GRAHAM, 
1982]. Also Cyclostremiscus beaui (P. 
Fischer, 1857) and Circulus texanus 
(Moore, 1965) in BIELER éz MIKKELSEN 
(1988). We consider this an insufficient 
basis for understanding the family. 

After a detailed anatomical examina- 
tion of living samples of Cochliolepis 
parasitica, type species of the genus, 
MOORE (1972) placed Cochliolepis in 


Vitrinellidae, due their anatomical simi- 
larity. He went on to remark that, in 
spite of the fact that some of the species 
are more conchologically similar to 
those of Tornus, the relationship of 
Cochliolepis with Tornus and Tornidae 
is not close. 

PONDER (1988) in his study on the 
phylogeny of Truncatelloidea (now 
known as Rissooidea), incorporated the 
available studies  (FRETTER, 1956; 
MOORE, 1972; GRAHAM, 1982) into his 
own, apparently not published, and 
inferred that there was a complex of 
tornids-vitrinellids-adeorbids (= cir- 
culids), as a single family, which equates 
to Tornidae, but warned that two or 
more families could be separated from 
this group. This author affirmed that the 
Tornidae (included with vitrinellids for 
this analysis) have some characters in 
common, but in their overall anatomy 
approach the hydrobiids-truncatellids 
and more closely the lIravadiidae and 
Elachisinidae. 

BIELER € MIKKELSEN (1988) studied 
several populations of two western 
Atlantic vitrinellids: Cyclostremiscus 
beauii (P. Fischer, 1897) and Circulus 
texanus (Moore, 1965) giving enough 
anatomical and radular data for generic 
differentiation. 

PONDER (1994) described the exter- 
nal and internal morphology of three 
vitrinelliform species from Hong Kong. 
Two of these, Sigaretornus plana and 
Circulus mortoni, are placed in the Vit- 
rinellidae, the other, a new genus and 
species, Lantauta taylori, is included in 
the Iravadiidae. The variation in organ 
systems in vitrinelliform species and the 
generic relationships are discussed, 
using the most comprehensive anatomi- 
cal studies then available, provided by 
BIELER é MIKKELSEN (1988). They 
confirm that the anatomy of Sigaretor- 
nus plana and that of Circulus mortoni 
are very similar to that of Cyclostremis- 
cus beauti, described in detail by BIELER 
éz MIKKELSEN (1988). This shows the 
necessity of establishing a profile with 
the most important characters in vit- 
rinellids and clarifying the relation 
between the type species of Vitrinella 


Iberus, 29 (2), 2011 


and Tornus, as both are the type genera 
of the families Vitrinellidae and 
Tornidae, due to the controversy on the 
relationships of both families. 

Authors such as GRAHAM (1982) 
have suggested that few differences 
were found between both families and 
so, they could be grouped together, 
while other authors, like MOORE (1965) 
and BIELER éz MIKKELSEN (1988) have 
suggested that both could be kept sepa- 
rate. 

RUBIO éz ROLÁN (1998) reported on 
the radulae of Pachystremiscus ornatus 
(Olsson € McGinty, 1958) and P. pul- 
chellus (Olsson €£ McGinty, 1958) and 
placed both species in the genus Lod- 
derena (Archaeogastropoda, Skenei- 
dae), considering Pachystremiscus a 
synonym of Lodderena. 

The species of tornids and vitrinel- 
lids are very similar in soft anatomy 
and radula as shown in ROLÁN éz RUBIO 
(2002) in their report on the Tornidae of 
the east Atlantic. 

BOUCHET éz ROCROI (2005) summa- 
rized the state-of-the-art for taxonomy 
of the Gastropoda, casting new light on 
the historical evolution of the phylum 
Mollusca. Malacologists currently con- 
sider this classification a “hybrid” of 
the pre-existing traditional taxonomy 
based on morphological characters and 
the more recent far-reaching revisions, 
based on the molecular characteristics 
of DNA and RNA. Also there are opin- 
ions about the classification of families 
into subfamilies which are often not 
well resolved, and should be regarded 
as the best possible hypotheses. In our 
opinion, the new classification of gas- 
tropods drastically changes existing 
systematics and is an important step 
forward the gastropod nomenclature. 
In this work, the family Tornidae Sacco, 
1896 is divided into four subfamilies: 
Torninae Sacco, 1896; Circulinae Fretter 
éz Graham, 1962; Teinostomatinae Coss- 
mann, 1917; and Vitrinellinae Bush, 
1897. The subfamily Torninae includes 
the genera previously placed in Tornus; 
Circulinae only includes the genus Cir- 
culus; the subfamily Teinostomatinae 
only has the genus Teinostoma and Vit- 


rinellinae groups together the other 
genera included in the old family Vit- 
rinellidae Bush, 1897, herein considered 
a subfamily. This new classification will 
be employed in the present work. 

As most of the species included in 
the group under consideration here 
have a typical shell form (orbicular to 
lenticular), we must point out that 
some species with similar shells actu- 
ally belong to other families: for 
example, the genus Cyclostrema 
Marryat, 1818 is in Liotiidae Gray, 1850; 
the genera Ganesa Jeffreys, 1883, 
Lydiphnis Melvill, 1906, Dillwynella 
Dall, 1889, Molleriopsis Bush, 1897 and 
Leptogyra Bush, 1897 are in Turbinidae 
Rafinesque 1815, subfamily Skeneinae 
Clark, 1851; the genus Choristella Bush, 
1897 is in Lepetellidae Dall, 1881 and 
the genus Cyclostremella Bush, 1897 is 
in Pyramidelloidea, Odostomiidae 
Pelseneer, 1928. Some of these generic 
names have  erroneously been 
employed for species included in the 
group studied here. 


MATERIALS AND METHODS 


Most of the material studied in the 
present work was acquired from sedi- 
ments obtained by diving or from 
dredgings and later separated with a 
binocular lens under magnification. 
Consequently most of the material is 
composed of empty shells in shell grit; 
occasionally a shell with soft parts 
could be obtained. An important part of 
the material studied is from Cuba, to 
which area we initially planned to limit 
our work; this material was mainly 
obtained from the collections of the 
second author in Cienfuegos Bay and 
later deposited mostly at the MHNS. 
For this reason, at the beginning of this 
study we only examined Cuban shells. 
Subsequently we included new mater- 
ial obtained by the third author on 
several trips to Yucatan (Mexico), 
Guatemala, Nicaragua, and south and 
west Cuba. Finally, we added other 
materials collected by Colin Redfern in 
the Bahamas, by Jacques Pelorce in 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


several islands of the Caribbean, by 
some collectors from Itaparica, Brazil, 
and an important quantity from the col- 
lection of Harry G. Lee, which consid- 
erably amplified the study geographi- 
cally and in terms of biodiversity. Some 
shells were lent by Marlo Krisberg, and 
we studied types and other material in 
other private collections and several 
Museums. 


Abbreviations: 


AMNH American Museum of Natural 
History, New York 

ANSP Academy of Natural Sciences of 
Philadelphia 

FLMNH Florida Museum Natural 
History, Gainesville 

IES Instituto de Ecología y Sistemática, 
Havana 

MCZ Museum of 
Zoology, Philadelphia 

MHNS Museo de Historia Natural 
“Luis Iglesias”, University of San- 
tiago de Compostela (coll E. Rolán) 

MNCN Museo Nacional de Ciencias 
Naturales, Madrid 


Comparative 


SYSTEMATIC PART 


MNHN Muséum National d' Histoire 
Naturelle, Paris 

MPH Museo Poey, Havana 

NHMUK National History Museum 
United Kingdom, London 

PRI Paleontological Research Institu- 
tion, New York 

RNHL Rijksmuseum van Natuurlijke 
Historie, Leiden 

USNM National Museum of Natural 
History, Washington 

YPM Peabody Museum of Natural 
History, Yale University, New 
Haven, Connecticut 

CCR collection of Colin Redfern, Boca 
Raton 

CEG collection of E.F. Garcia, Louisiana 

CFG collection of R. Fernández-Garcés, 
Cienfuegos 

CFR collection of F. Rubio, Valencia 

CHE collection of Harry G. Lee, Florida 

CJP collection of J. Pelorce, Le Grau de 
Roi 

CMK collection of M. Krisberg, Florida 

sp specimen with soft parts 

s empty shell 

] juvenile 

f fragment 


Superfamily RISSOOIDEA Gray, 1847 
Family TTORNIDAE Sacco, 1896 
Subfamily CIRCULINAE Fretter €: Graham, 1962 


Genus Circulus Jeffreys, 1865 


Lydiphnis Dall, 1927, pp. 123 


Circulus Jeffreys, 1865: 315. British Conchology. Vol. VII. Marine shells comprising the remain- 
ing Conchifera, the Solenoconchia and Gasteropoda as far as Littorina. J. Van Voorst. Plenum 
Press, New York and London, 393(+1) pp., 8 pls + frontispiece. 

Type species (by monotypy): Trochus duminyi Requien, 1848 (= Valvata striata Philippi, 1836). 


Diagnosis: “Shell relatively strong, 
depressed; base smooth, rounded, umbili- 
cus opened and deep spiral sculpture 
presenton the dorsum and on the periph- 
exnye: 

Remarks: Circulus has been habitually 
treated as a section or subgenus of a 
better known genus. FISCHER (1887) 
places it as a subgenus of Gibbula Risso 
and BusH (1897) as a section of Vitrinella. 


DaLL (1927) reassigned to Lydiphnis 
Melville, 1906 some of the species placed 
in Circulus by Bush, but without giving 
an explanation. 

THIELE (1929: 63) considered Circulus 
as a genus within Cyclostrematidae and 
mentions some details in the text about 
the radula. 

FRETTER (1956) researched the 
anatomy of Circulus striatus (Philippi), 


Iberus, 29 (2), 2011 


type species of Circulus, and found 
several discrepancies in Thiele in refer- 
ence to this genus. The radula is tae- 
nioglossate, not rhipidoglossate, and the 
animal is similar in appearance to that of 
the Rissoidae. The animal has a pair of 
pallial tentacles on the right margin of the 
mantle, and epipodial tentacles are 
wanting. The sexes are separate, and the 
male has a penis. 

KEEN (1960) places the genus in 
Cyclostrematidae, Order Archaeogas- 
tropoda. But the detailed anatomical 
work of FRETTER (1956) has shown that 
the animal belongs in Mesogastropoda 
close to the family Rissoidae. 

Species have been placed in the genus 
Circulus in the Caribbean, West Africa 
and Australia. Circulus striatus (Philippi) 
has been found on the southeast coast of 
Europe and in the Mediterranean. Circu- 
lus liratus (A.E. Verrill, 1882) was found 
from the east coast of Florida up to New 
England in the United States and Circulus 
semisculptus (Olsson £ McGinty, 1958) 
from both coasts of Florida and Panama. 
Several species have been described from 
the latter region, but apparently they 
must be reassigned to other genera. 

The shells are very similar in their 
general aspect to those of Solariorbis, but 
they lack the strong rib close to the 
umbilicus. 

According to MOORE (1964) Adeorbis 
orbignyi P. Fischer, 1857, described from 
Cuba, may be included in this genus, but 
the type in the Laboratoire de Malacolo- 
gie of MNAN is lost and so he considered 
the taxon a nomen dubium. 

BIELER 6 MIKKELSEN (1988) give a 
complete account of the anatomy and 
reproductive biology of two species 
grouping them in  Vitrinellidae: 
Cyclostremiscus beaui (P. Fischer, 1857) 
and Circulus texanus (Moore, 1965), from 


populations of both species found in 
Florida, in burrows in fine sand of the 
stomatopod  crustacean  Lysiosquilla 
scabricauda (Lamarck, 1818). The great 
size of Cyclostremiscus beaui (6-8 mm 
maximum diameter) simplified the 
detailed study of its anatomy and mor- 
phology and at the same time allowed the 
authors to confirm FRETTER'S (1956) 
hypothesis that some characters found in 
Circulus striatus, are in proportion to its 
overall size. 

PONDER (1994: 258) says: “species of 
Circulus can be defined as vitrinellids 
having a spirally ridged shell with a 
simple non varicose aperture, a simple 
penis and a posteriorly notched foot 
lacking a metapodial tentacle. This latter 
character is absent in Vitrinella texana 
Moore, a species included in Circulus by 
BIELER 62 MIKKELSEN (1988)”. 

There are seven recent species of Cir- 
culus described from the east coast of 
USA and the Caribbean: Circulus liratus 
(A.E. Verrill, 1882), Circulus texanus 
(Moore, 1965), Circulus semisculp tus 
(Olsson € McGinty, 1958), Circulus 
orbignyi (P. Fischer, 1957), Circulus hen- 
dersoni (Dall, 1927), Circulus margariti- 
formis (Dall, 1927) and Circulus translu- 
cens (Dall, 1927). In the present work all 
these taxa have been studied and figured. 
A neotype is designated for C. orbignyi, a 
taxon considered by some authors as a 
nomen dubium, the type material having 
been lost. DALL (1927) described C. hen- 
dersoni, C. margaritiformis and C. 
translucens, in the genus Lydiphnis, all of 
them from deep water dredgings in 
Georgia and Fernandina. These taxa will 
be discussed in the section following the 
main revision inasmuch as we have deter- 
mined that they are not tornids but 
species in Skeneinae as will be demon- 
strated below. 


Circulus semisculptus (Olsson £ McGinty, 1958) (Figures 1A-D) 


Vitrinella semisculpta Olsson €: McGinty, 1958. Bulletins of American Paleontology, 39: 30-31, pl. 
3, figs. 2-2b. [Type locality: Bocas Island, Panama]. 


Type material: Holotype represented in MOORE (1964: 220, fig. 7). Deposited in the ANSP (211888). 


Not examined. 


RUBIO £7 AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Rancho Luna Beach, Cuba; D: protoconch. 


Figures 1A-D. Circulus semisculptus (Olsson 82 McGinty, 1958 


200 yn 


). A-C: shells, 1.9, 1.6, 1.6 mm, 


Figuras 1A-D. Circulus semisculptus (Olsson e McGinty, 1958). A-C: conchas 1,9 1,6, 1,6 mm, 


Playa Rancho Luna, Cuba; D: protoconcha. 


Other material examined: Cuba: 2 s, Rancho Luna Beach, Cienfuegos, 10-20 m (MHNS); 3 s, Rancho 
Luna Beach, 20-54 m (MHNS). Antigua and Barbuda: Antigua: 1 s, north St. John, 5-6 m (CJP). Mar- 
tinique: 1 s, Pointe Borgnesse, 12 m, on sand-muddy on the base of the reef (CJP). Florida, USA: 1 
s, SE Raccoon Key, Monroe Co., 0.5 m subtidal with Halimeda sps (CHL). Bahamas: 1 s, Olympus 
Reef, Grand Bahama Island, 36 m coralline algae bottom (CHL). 


Description: Shell (Figs. 1A-C) 
rather strong, whorls rounded with 
spiral sculpture on the dorsum and 
base smooth. Protoconch (Fig. 1D) rel- 
atively large, about 310 um maximum 


diameter, with 1 Y spiral whorls 
where two stages of development are 
easily observable. Its surface is appat- 
ently smooth although some rough 
parts are present in the terminal 


Iberus, 29 (2), 2011 


portion. A strong varix marks the sep- 
aration from the teleoconch. Last 
whorl with 5 spiral cords distributed 
on the dorsal half of the whorl 
between the suture and the periphery. 
A fine axial striation covers the inter- 
spaces between the spiral cords. Base 
rounded, totally smooth except 
growth lines slightly oblique, peris- 
tome not continuous. 

Habitat: MOORE (1964: 73) suggests 
that the species seems to prefer 
shallow water, and all the shells in the 
type series were in beach sediments. 
However, several others came from 
between 10 and 54 m deep. 

Distribution: lt has been recorded 
from Colón island and Bocas island, 


Panama (OLssoN € MCGINTY, 1958); 
from South Plonida and Panama 
(MOORE, 1964); from South of Florida 
and Central America (HOUBRICK, 
1968): from Portete, Costa Rica 
(HOUBRICK, 1968; ROBINSON € 
MONTOYA, 1987); from south Florida 
and the western Caribbean (ABBOrTT, 
1974); from Aruba (DE JONG éz 
COOMANS, 1988); from Colombia (DÍAZ 
MERLANO € PUYANA HEGEDUS, 1994); 
from Abaco, Bahamas (REDFERN, 2001). 
In the present work Cuba, Antigua 
and Martinique are added. 

Remarks: Our material matches the 
figure in MOORE (1964: 220, fig. 7) and 
the shell photographed by REDFERN 
(2001, fig. 184) from Abaco, Bahamas. 


Circulus orbignyi (P. Fischer, 1857) (Figures 2A-D) 


Adeorbis orbignyi Fischer, P. 1857. Journal de Conchyliologie, 6: 173. [Type locality: Cuba]. 
Circulus sp.: In REDFERN, 2001: 43, pl. 21, figs. 185. 


Type material: The holotype is lost. Neotype (Fig. 2A), here designated, in MNHN (24227). 

Other material examined: Cuba: 1 s, Jibacoa, 3-6 m (MHNS); 1 s, Baracoa, on beach (MHNS); 5 s, 
Cienfuegos Bay, 20 m (MHNS); 1 s, Cienfuegos Bay, 30 m (MHNS); 1 s, Rancho Luna Beach, 12 m 
(MHNS); 23 s, Rancho Luna Beach, 10-20 m (MHNS); 11 s, Rancho Luna Beach, 45 m (MHNS); 1 s, 
Punta Tamarindo, 15 m (MHNS). Grenadines: 1 s, Mayreau, 8 m, coralline sand with coral, gor- 
gonies and sponges (CJP). Trinidad and Tobago: Tobago, 1 s, Horse Shoe reef, Tobago Cays, 15 m, 
in sediments (CJP). Nicaragua: 1 s, Cayo Witties, 15 m (MHNS). Panama: 1 s, Portobello, drift (CHL). 


Florida, USA: 1 s, Hallandale Beach, Broward Co., Florida, drift (CHL). Bahamas: 1 s, Grand Bahama 


Island, 0.5 m on sand (CHL). 


Type locality: Cuba; the neotype is from Rancho Luna Beach, Cienfuegos, Cuba. 


Description: Shell (Figs. 2A-C) 
strong, depressed, spire slightly ele- 
vated, composed of 4 Y whorls with 
spiral sculpture on its dorsal half and 
fine growth striae on the base. 
Umbilicus wide and deep. Protoconch 
(Fig. 2D) of almost 1 % whorls, and 
about 302 um in diameter, with two 
stages of development well sepa- 
rated; initially smooth, continuing 
with a slightly rough surface and 2-3 
threads on its terminal portion. A 
strong varix marks the beginning of 
the teleoconch, which is composed of 
2 Y rounded whorls, smooth on the 
base except for growth lines, and 14 
to 18 spiral cords on the dorsal 
middle on the last whorl. Spaces 
between cords covered by fine irregu- 


lar reticulation. Aperture rounded, a 
little oblique. Columella slightly 
reflected. Under high magnification, 
faint spiral cordlets can be seen inside 
the umbilicus. 

Dimensions: The neotype is 2.5 
mm in diameter and 1.7 mm in height. 

Animal and radula unknown. 

Habitat: In Cuba it seems to be a 
widely distributed species, most shells 
were found between 10 and 20 m 
deep. 

Distribution: The species is known 
from Cuba, Bahamas, the Grenadines 
and Tobago. It is mentioned from Sáo 
Sebastiáo, Sáo Paulo, Brazil (Rios, 
2008). It has been recorded from 
Abaco, Bahamas (REDFERN, 2001) as 
Circulus sp. 


RUBIO ET 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 2A-D. Circulus orbignyi (P. Fischer, 1857). A: neotype, 2.5 mm (MNHN); B: shell, 2.7 
mm, shell, Rancho Luna Beach, Cuba (CFG); C: 2.4 mm, Cayo Witties, Nicaragua (MHNS); D: 
protoconch. 
Figuras 2A-D. Circulus orbignyi (P Fischer, 1857). A: neotipo, 2,5 mm (MNHN); B: concha, 2,7 
mm, concha, Playa Rancho Luna, Cuba (CFG); C: 2,4 mm, Cayo Witties, Nicaragua (MHNS); D: 


protoconcha. 


Discussion: FISCHER (1857) in the 
original description of Adeorbis 
orbignyi said: “It is distinguished 
from congeneric species by the regu- 
larity of the ornamentation consti- 
tuted by a dozen of transversal ribs 
placed at equal distance”. MOORE 


(1964: 70-71) treated it as a nomen 
dubium only because the type was 
not found. This taxon cannot be con- 
sidered as nomen nudum with a 
good description like the one avail- 
able. The type, supposedly in the 
MNHN, is considered lost (Virginie 


Iberus, 29 (2), 2011 


Héros pers. comm.). In our opinion 
there is no doubt that the description 
of this species corresponds to the 
shells we have from Cuba (type local- 
ity), where it is relatively common. 
For this reason and also because the 
taxon has been accepted by other tax- 
onomists (e.g., www.malacolog.org) 


we have designated a neotype in 
order to maintain nomenclatural 
stability. 

This species can be  distin- 
guished from C. semisculptus by 
the great number of spiral cords 
and the dense  microsculpture 
between them. 


Circulus liratus (A.E. Verrill, 1882) (Figures 3A-E) 


Cyclostremiscus pentagonus auct. non Gabb, 1873. 


Omalaxis lirata A.E. Verrill, 1882. Transactions of the Connecticut Academy of Arts and Sciences 
5: 529. In Bush, 1893, pl. 1, fig. 11-12]. [Type locality: USFC sta. 770, off Newport, Rhode 


Island, 16 ml). 


Type material: Holotype in USNM (406741). Not examined. 
Other material examined: Florida, USA: 1 s, Atlantic Beach, Duval Co. (CHL); 4 s, 30 m, 35 mi E St. 
Augustine, St. Johns Co. (CHL); 2 s, 53 m, 75 mi E St. Augustine, St. Johns Co. (CHL); 4 s, 16th Ave. 


S, Jacksonville Beach, Duval Co. (CHL). 


Description: ratter MIO OKE 
(196 Sell dep mes sed, 
rather hice + wltorls oro undieds 


inner half of base smooth. Spiral 
seulpture o abomuteronito ter 
spiral= rides separated by 
esrooVes  oniyo ss iiontly iden, 
Unmnbilicustuidetand deep ros 
toconch projecting with nearly 2 
3% smooth whorls, about 530 um 
in maximum diameter. Teleo- 
conch with only 1 % whorls, 
ornamented with 10 spiral cords 
distributed between the dorsum 
and the outer base. The inter- 
spaces are wider, without axial 
sculpture except fine growth 
lines. Base flat, without sculp- 
ture. Umbilicus wide, the previ- 
ous whorls being visible in its 
interior, randidelimited" bye 
strong cord and 2-3 more on its 
inner wall. 

Holotype measures 2.1 mm. 
The figured shells are 2.3-2.7 mm 
in diameter and about 1.01 mm in 
height. 

Habitat: This species seems to 
be found living in depths of a few 
meters (MOORE, 1964). The 
bathimetry reported in the litera- 
ture is between 7 and 165 m. 


10 


Distribution E astCOasS Ok 
Florida to Rhode Island. Recorded 
from Newport, Rhode Island 
(VERRILLE, 13882) tromol Cape 
Hatteras, North Carolina (Bush, 
1897); from Rockaway, New York 
and off Lantana, Florida (MOORE, 
1964); from St. Lucie Co., Florida 
(LYONS,. 19ISO) from Atlantic 
Beach, Jacksonville Beach, Batten 
Island and St. Augustine, Florida 
(LEE, 2009). 

Remarks: C. liratus is the only 
species of Circulus from the West 
Indies similar in morphological 
characters to  Circulmsserialtis 
(Philippi, 1836) type species of the 
genus. C. liratus has been placed 
in the genera Lydiphnis and 
Skenea. Circulus liratus differs 
from C. semisculptus by its pro- 
jecting protoconch while C. semi- 
sculptus is almost planispiral. 

Solariorbis mooreana (Vanatta, 
OO) AS Aim lar batas ita 
narrow umbilicus and lacks the 
thickening of the umbilical wall, 
which is typical of the Solariorbis; 
also this latter species lacks orna- 
mentation in the  interspaces 
between cords, and these cords do 
not extend to the base. 


RUBIO £7 AZ.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 3A-E. Circulus liratus (A.E. Verrill, 1882). A-C: shells, 2.18, 2.14, 2.33 mm, Jacksonville 
Beach, Florida (CHL); D: detail of the umbilicus; E: protoconch. 

Figuras 3A-E. Circulus liratus (4.E. Verrill, 1882). A-C: conchas, 2,18, 2,14, 2,33 mm, Jacksonville 
Beach, Florida (CHL); D: detalle del ombligo; E: protoconcha. 


Circulus texanus (Moore, 1965) (Figures 4A-F) 


Vitrinella texana Moore, 1965. The Nautilus, 78: 76-77, pl. 7 figs. 4-6. [Type locality: Mustang 
Island, near Port Aransas, Texas]. 
Vitridomus texanus (Moore, 1965), auct. 


Type material: Holotype from Mustang Island, near Port Aransas, Texas, diameter, 1.72 mm, height, 
0.78 mm. Deposited in the Division of Mollusks, USNM (636311). Not examined. 


11 


Iberus, 29 (2), 2011 


Other material examined: Brazil: 1 s, Itaparica (MHNS). Florida, USA: 1 s, Delray Beach, Palm Beach 


Co. (CHL); 1 s, Longboat Key, Sarasota Co. (CHL). 


Description: Original descrip- 
tion (MOORE, 1965): “The shell is 
depressed, and has a flattened 
apex. The umbilicus is narrow but 
deep, and is almost flat sided. Sides 
os the shell curve out and idilio wn 
gently so that the periphery forms 
an angle with the base of the shell. 
The aperture is oblique. 

“The protoconchiconsists o ess 
glassy whorls. The teleoconch con- 
sists of about 1 Y whorls, and is 
sculptured on the upper side with 
espiral 10 ves ando re 
lower side with numerous short 
sadiatino rnibletso Iihese ribletstare 
eros sed by a eel s penal 
ero oves mando theretamenas ev exa 
stronger spiñal  Sroo Ves mine 
umbilicus. The ventral side is flat- 
tened, sand, mn tre ho lotype Deals 
about 30. radiatinio nibletso he 
riblets become indistinct on the last 
half of the whorl, and become diffi- 
cult to count”. 

“The aperture is oblique, and is 
broadly ovate. The peristome is 
deeply notched at the upper inner 
angle. The parietal wal is rates 
thiek, and. is extended. a Little 
forward ojthe tapertunes e 
umbilicus is narrow and almost flat 
sided, but there is no angle with the 
base of the shell. The shell itself is 
quite thin and fragile, and only the 
holotype and one  immature 
paratyupe. cane unbroken. Oe 
paratype is actually only half of the 
body whorl of a broken shell”. 

After BIELER 6  MIKKELSEN 
(1988): “Shell small (1.7-1.8 mm in 
diameter, 0.55-0.65 mm in height). 
With about 1 Y teleoconch whorls; 
almost planispiral, sculptured dor- 
sally and ventrally with about 18 
fine spiral ribs; transparent when 
aloe, opaque aujter deati. eibis 
slightly stronger, more widely 
spaced. qustbelowitsuture om 
dorsum and at periphery, where 
about ia os porro undeda Meeel 


12 


below lateral midline. Suture 
impressed. Ventral surface below 
keel less convex. Often with 30-40 
widely-spaced, low axial ribs which 
are primarily evident from inside of 
body whorl. Umbilicus wide. Outer 
ip very sliehtly re lected some 
specimens with one former varix. 
Aperture at oblique angle to dorso- 
ventral axis. Sutural sinus shallow. 
Periostracum thin, transparent, 
with spiral grooves more numerous 
than on shell surface. Protoconch 
smooth, 0.5 mm diameter, about 2 
whorls. No sculptural demarcation 
separating protoconch I and proto- 
conch Il. 

Maximum 

We can add to this description 
the following: The protoconch 
(Figs. 4D-E) is about 340 um in 
diameter, and it has a little more 
than 1 Y whorls. Apparently it is 
smooth, but its nuclear portion is 
covered by very small tubercles of 
various sizes. This kind of 
microsculpture is also present on 
the dorsum of the teleoconch (Fig. 
4F). 

Habitat: Living specimens were 
taken from stomatopod burrows in 
shallow-water sand flats in the 
Indian River lagoon, St. Lucia 
County, eastern Florida (BIELER éz 
MIKKELSEN, 1988). It prefers a 
rather narrow range of temperature 
and salinity, and is probably most 
abundant in shallow coastal waters. 
Its bathymetric range is between 0 
and 44 m in depth, and it was 
found alive between 0 and 0.5 m. 

Distribution: USA: Florida: East 
Florida; Texas. Reported from Port 
Aransas, Texas (MOORE, 1964); from 
Texas (ODÉ, 1987b); from the north- 
western Gulf of Mexico (BIELER € 
MIKKELSEN, 1988). 

Remarks: This species was 
described in the genus Vitrinella 
(MOORE, 1965). BIELER €: MIKKELSEN 


ANIOS SÍAES Lol 


RUBIO £7 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 4A-F. Circulus texanus (Moore, 1965). A-C: shell, 1.78 mm, Sarasota Co., Florida (CHL); 
D: protoconch; E: detail of the protoconch; F: detail of the microsculpture of the teleoconch. 
Figuras 4A-E Circulus texanus (Moore, 1965). A-C: concha, 1,78 mm, Sarasota Co., Florida (CHL); 
D: protoconcha; E: detalle de la protoconcha; E: detalle de la microescultura de la concha. 


(1988) placed it in the genus Circu- the eastern Atlantic. It differs from 
lus sensu lato because it agrees in all other western Atlantic Circuli- 
shell shape and sculpture with the nae principally in having radiating 
type species, Circulus striatus from riblets on the ventral side. 


13 


Iberus, 29 (2), 2011 


Subfamily TEINOSTOMATINAE Cossmamn, 1917 
Genus Teinostoma H. € A. Adams, 1853 


Teinostoma H. £ A. Adams, 1853. Genera of Recent Mollusca 1: 122. 
Type species: Teinostoma politum H. 6 A. Adams, 1853 (by monotypy). Proc. Zool. Soc. Lond. 


pl. 10, figs. 1-3. 


Diagnosis: Shell minute, depresed- 
turbiniform, thin, glassy, smooth, 
umbilicate. Protoconch of about 1.3 
convex whorls. Teleoconch whorls 
convex, base and umbilical rim angu- 
late. Multispiral operculum. Rádula: 
Central tooth cutting area broadly 
“V” shaped, serrated, 2 basal denti- 
cles, ventral process “U”. shaped. 
Lateral tooth cutting area at inner 
third, broadly angulated and ser- 
rated. Inner marginal cutting area 
very broad, comb-like. Outer mar- 
ginal cutting area short. 

MOORE (1964) defines the species 
as: “shell small to minutecompact, 
depressed, low spired, few flattened 
whorls which are rounded or carl- 
nate; smooth or sculptured by fine 
striations, suture not impressed, 
umbilicus covered partly or entirely 


Identification key 


by a heaoy calls pad Wi tehos 
extended over the parietal wall, col- 
umella concave, thick. Animal very 
similar to Vitrinella but with a longer 
oO e 

Remarks: According to PILSBRY 
(1953) the species included in the 
genus Teinostoma are known from 
the Upper Cretaceous to Recent, 
being very common in many Tertiary 
deposits. The genus Teinostoma has 
been subdividided by some authors 
into several subgenera (Annulicallus, 
Pseudorotella, Idioraphe); unfortu- 
natly, the types of the type species of 
two of these subgenera are lost or in 
such poor condition that identifica- 
tion is uncertain. As the objective of 
this work is not supraspecific classifi- 
cation, we will group all the studied 
species in the genus Teinostoma. 


In order to make schematic the separation of the species in this group with so 
many species we present the following identification key for the genus pointing out 
the most important characters for each species: 


1 - Shell With Protocol 2 
- Shell with protoconch fully or partially covered by athin coat ............ 3 
2 Snell with tounded mictopits spraly An - 
- onell with spral nas e 5 
= Shell completely smooth... 6 
3 Shell withispiral orde di iodo 7 
- Shell completelyssmooth o. o e 8 
4 ohell elobosesandifracilen a lo. le la Ad T. ciskae 


- Shell globose with low spire ....... 
- Shell with strong peripheral keel .... 


- Shell angular at the periphery 


nel suba 
5 - Shell obtusely subangular ....... 


O T. baldingeri 
A A T. goniogyrus 
o e EI e A T. lenticulare 
O A e A T. reclusum 


E NN RS A UR y RO T. incertum 


- Shell with spiral irregular microcordlets fused between them in the first 


A A ON 
- Shell with spire slightly elevated and striated callus 


14 


RRE E UN ADN ES T. anastomosis 


E e T. panamense 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


GAtumbilicus completely covered PycallUs A E. 9 
Un bllicus partyico vereda e A 10 
Shell totallyscovered by ispital COLAS: 2 T. semistriatum 


=snellidorsallyicovered by tine spiralicordlets: T. nesaeum 
- Shell with dorsum and umbilicus surrounded by a strong spiral carina T. carinicallus 
- Shell with dorsum and umbilicus surrounded by a strong spiral carina and weak spiral 


A e e A T. lituspalmarum 
8 - Shell strongly depressed, transversely dilated” ...........T.... T. obtectum 
ohclttiexpanded apertute e a as T. expansum 
- Shell minute, flattened above and below... ..::2.....0.... T. minusculum 
- Shell more elongated by extension (outwards from the outer lip ...... T. lerema 
HS ES A A T. megastoma 

- Shell with periphery very rounded and strong umbilical callus . .T. umbilicatum 
9 - broadly ovate aperture, rather strongly oblique.............. T. biscaynense 
- protoconch placed below the next whorl, rounded aperture, peristome almost con- 
A A E A O T. cienfuegoensis 
¡penistomeexternally reflected toward Dack.................- T. helicinum 

So hellpyrttorm, umbilical callus very large... ono co T. megacallum 

- a fine groove separates the umbilical callus from the columella . . .T. parvicallum 
Wa tmanetlar callus atend of the columella..... oo... ....-. T. solidum 
- no groove of separation between columella and callus ......... le cocolitorts 
- spire moderately elevated, callus with half moon shape .......... T. lunense 


- a groove separating the umbilical callus from the columella 


Teinostoma ciskae Faber, 1995 (Figures 5A-C) 


Teinostoma millepunctata Nowell-Usticke, 1969 non T. millepunctatum Pilsbry £ Olsson, 1945. 
A Supplementary Listing of New Shells, to be Added to the Check List of the Marine Shells of 
St «Croix: 10,pl. 2, fe, 307. 

Teinostoma millepunctata Nowell-Usticke, 1971. A Supplementary Listing of New Shells, to be 
Added to the Check List of the Marine Shells of St. Croix, revised edition: 6. [Type locality: 
Antigua, Secret Harbor, 40 ft]. 

Teinostoma ciskae Faber, 1995. De Kreukel, 31: 62 [replacement name for T. millepunctata 
Nowell-Usticke, 1969]. 


Type material: Represented in NOwELL-UsTICKE (1969, pl. 2, fig. 307). The lectotype of T. 
millepunctata was deposited in AMNH (n* 195413) (BoYko € CORDEIRO, 2001). Not exam- 
ined. 

Other material examined: Cuba: 5 s, Bahía de Cienfuegos, 20-30 m (MHNS); 20 s, Rancho 
Luna Beach, 20-54 m (MHNS). Trinidad and Tobago: Tobago, 1 c, Horseshoe reef, 15 m, from 
shell grit (CJP). Virgin Islands: 1 s, Peter Island, 18 m, shell grit (CHL). Bahamas: 1 s, Olympus 
Reef, NNW West End, Grand Bahama Island, 36 m, from coralline algal fragments (CHL). 
Florida, USA: 2 s, APAC Pit, Sarasota Co. Plio-Pleistocene (CHL). 


Description: Shell (Figs. 5A-B) 
globose, fragile, whitish and with its 


quent part, about 166 um in diameter, 
and with two strong varices separating 


surface totally covered by punctiform 
incisions aligned spirally. 

Protoconch (Fig. 5C) of about one 
whorl, with a finely granular surface at 
its beginning and smooth in the subse- 


the two stages. Teleoconch of about 2 Y 
globose whorls, totally covered by the 
microsculpture mentioned above (Fig. 
5C). Aperture rounded, external lip 
fine, inner lip and columella thickened. 


iS 


Iberus, 29 (2), 2011 


fuegos, Cuba; C: protoconch. 


50 um 


Figures SA-C. Teinostoma ciskae Faber, 1995. A-B: shells, 1.8, 1.2 mm, Rancho Luna Beach, Cien- 


Figuras SA-C. Teinostoma ciskae Faber, 1995. A-B: conchas, 1,8, 1,2 mm, Playa Rancho Luna, 


Cienfuegos, Cuba; C: protoconcha. 


Umbilicus totally closed by an extension 
from the columella. 

Dimensions: Holotype 1.59 mm in 
diameter. We have shells with about 2.5 
mm in maximum dimension. Maximum 
reported size: 2.6 mm 

Habitat: The species is distributed in 
the deep infralittoral, found between 15 
and 54 m deep, on coralline bottoms. 

Distribution: Known from Antigua, 
its type locality (NOwELL-USTICKE, 1969 
and 1971; FABER, 1995); from Aruba (DE 
JONG € COOMANS, 1988); from Abaco, 
Bahamas (REDFERN, 2001) and from 
Virgin Islands, Bahamas, Tobago, and 
Cienfuegos, Cuba, in the present work. 


Remarks: NOWwELL-UsSTICKE 
(1969) described Teinostoma 
millepunctata. This name was preoc- 
cupied by T. millepunctatum Pilsbry 
€ Olsson, 1945, from Ecuador, for 
this reason FABER (1995) proposed 
the replacement name Teinostoma 
ciskae, for Nowell-Usticke”s species. 
T. ciskae may be distinguished from 
the other known species of Teinos- 
toma by the more globose and 
fragile shell, by its peculiar proto- 
conch, and mainly by its typical 
microsculpture of punctiform inci- 
sions. No similar species exists in 
the Caribbean region. 


Teinostoma gontogyrus Pilsbry £ McGinty, 1945 (Figures 6A-G) 


Teinostoma goniogyrus Pilsbry £ McGinty, 1945a. The Nautilus, 59: 3, pl. 1, figs. 8. [Type local- 


ity: Off Destin, west Florida]. 


Rotella carinata H. C. Lea, 1846. Trans. Amer. Philos. Soc., 9: 263, pl. 36, fig. 78. (non d'Orbigny, 
1842) [Type locality: Petersburg, Virginia, Neogene fossil]. 


16 


RUBIO ET 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Type material: Represented in PILSsBRY ££ MCGINTY (1945a). Not examined. 

Other material examined: Cuba: 1 s, Guajimico, 15 m (MHNS); 1 s, Cienfuegos Bay, stn. 12a, 22%07'N 
— 8026'W, 4 m (MHNS); 12 s, Cienfuegos Bay, 10 m (CFG); 5 s, Rancho Luna Beach, 10-54 m (CFG). 
Florida, USA: 1 s, 65 mi. E St. Augustine, St. Johns Co., Fl., 53 m, dredged (CHL); 3 s, 32 mi. E St. 
Augustine, St. Johns Co., Fl., 30 m, dredged (CHL); 1 s, 23 mi. ENE Mayport, Duval Co., Fl., 26 m. 
(CHL); 1 s, Caloosahatchee Formation, La Belle, Hendry Co., Plio-Pleistocene (CHL). 


Description: Shell (Figs. 6A-D) 
subconical, depressed, solid, 
whitish, and with a strong keel at 
the periphery. Protoconch (Figs. 6ÉE- 
G) of about 2 whorls and about 360 
um in diameter, with a smooth 
surface at its beginning and fine 
lateral granulation and 5-6 spiral 
lines of small perforations on the 
subsequent part, varix scarcelly 
marked. Teleoconch of about 1 l4 
whorl, rapidly expanding, dorsally 
convex and ventrally concave in the 
umbilical area; surface totally 
covered by very fine clearly sepa- 
rated perforations, spirally aligned 
(Figs. 6G-H) and with a prominent 
cord-like keel at the periphery. A 
fine callus completely covers the 
umbilicus; a fine groove runs 
between the columella and the 
callus. Aperture ovoid, a little 
depressed, with the upper part of 
the external lip sharp and advanced. 

Dimensions: Holotype measures 
1.95 mm in diameter. We have shells 
about 1.3 mm in maximum diame- 
ter. 

Habitat: Marl Bottom, in 18-20 
fms (32-56 m) (Pilsbry € McGinty, 
1945a). It has been found alive 
between 42 and 59 m, but shells 
have been found in sediments col- 
lected between 10 and 100 m. 

Distribution lt. has > been 
recorded from off Destin, west 
Florida (PILsBRY ér MCGINTY, 1945a); 
from Bocas island, Panama (OLsson 
éz MCGINTY, 1958); from off north- 
west Florida, southern Haiti and 
Panama (MOORE, 1964); from north- 
west Gulf of Mexico (ODÉ, 1987); 
from Florida to Caribbean Panama 
(Lyons, 1989; LEE, 2009); from 
Colombia (Díaz MERLANO  éz 
PUYANA HEGEDUS, 1994); also from 
Cuba. 


Remarks: PILSBRY é€ MCGINTY 
(1945a) mention that Teinostoma 
goniogyrus resembles Rotella cari- 
nata (d'Orbigny) from St. Thomas in 
shape, but instead of the small 
umbilical callus of that species, has 
a remarkable, extremely thick callus, 
exceeding that of any other Teinos- 
toma except T. pilsbryi. Under high 
power some faint traces of close 
spiral striation can be seen in a few 
places on the unique type. MOORE 
(1964) commented that this species 
is also similar to T. incertum in the 
spiral punctiform lines and in the 
shape of the umbilical callus. T. 
incertum is more depressed and has 
a strong peripheral keel. The strati- 
graphic distribution of this species 
is from the Upper Miocene to 
Recent. There is a considerable vari- 
ation of size between the fossil 
shells from the Miocene and Plio- 
Pleistocene and the recent ones. 

PILSBRY dé MCGINTY (1945a) 
described T. goniogyrus on the basis 
of a single shell, citing the wider 
umbilical callus as the specific dif- 
ference from R. carinata. PILSBRY 
(1953) figured fossil shells of T. 
goniogyrus from Smithfield, Vir- 
ginia and St. Petersburg, Plio-Pleis- 
tocene of southern Florida and kept 
the size of the umbilical callus as the 
only difference between species. We 
have examined shells from Florida 
and Cuba and found very little dif- 
ference between them. We have also 
examined shells from the Pliocene, 
Caloosahatchee Formation, from La 
Belle, Florida, and we have not 
observed important differences in 
the size of the umbilical callus. K.J. 
Bush (1897) identified two speci- 
mens from station 2278, off Cape 
Hatteras, in 16 fathoms (29 m), as 
the R. carinata of d'Orbigny. 


1 


Iberus, 29 (2), 2011 


Figures GA-G. Teinostoma goniogyrus Pilsbry 82 McGinty, 1945. A-D: shells, 1.5, 1.5, 1.4, 1.5 mm, 
Cienfuegos Bay, Cuba; E: protoconch; F-G: microsculpture. 

Figuras GA-G. Teinostoma goniogyrus Pilsbry + McGinty, 1945. A-D: conchas, 1,5, 1,5, 1,4, 1,5 
mm, Bahía de Cienfuegos, Cuba; E: protoconcha; F-G: microescultura. 


In our opinion, R. carinata and T. 
goniogyrus may be the same 
species, and the different size of the 
umbilical callus is not enough for a 
specific separation. The problem is 
that the shells identified by K.J. 
Bush as R. carinata in USNM were 


18 


not found. So, lacking comparative 
material, we keep both species- 
level taxa waiting until more mate- 
rial from the type locality is 
obtained in the future in order to 
decide if there is any specific dif- 
ference. 


RUBIO £7 412.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Teinostoma lenticulare (H.C. Lea, 1846) (Figures 7A-K) 


Rotella lenticularis H.C. Lea, 1846. Trans. Amer. Philos. Soc., 9: 264, pl. 36, fig. 79. [Type locality: 


Petersburg, Virginia, Neogene fossil!]. 


Type material: Type material in ANSP. Not examined. 

Other material examined: Cuba: 12 s, Cienfuegos Bay, 22%07'N 80%27'W, 9 m (MHNS); 5 s, Cien- 
fuegos Bay, sta. 12a, 22"07'N 80%26'W, 4 m, 19 s, Cienfuegos Bay, 10 m (MHNS); 1 s, Cienfuegos 
Bay, 20-30 m (MHNS); 15 c, Cienfuegos Bay, 10 m; 6 c, Cienfuegos Bay, 12 m (MHNS). 


Description: This is the short original 
description: “Shell lenticular, depressed, 
thin, smooth, polished, spire very short, 
sub-ovate; obtuse; sutures small, linear; 
whorls four, convex; last whorl angulate; 
base smooth; callus small; mouth sub- 
rotund; columella broad, curved”. At 
same time H.C. Lea comments: “The 
angle of the last whorl is very variable. It 
sometimes amounts almost to a carina. 
The mouth is nearly round. The callus is 
slightly depressed below the surrounding 
surface. This shell is, in part, allied to 
both the preceding species [Rotella cari- 
nata], but differs in the number of whorls, 
shape of the columella and spire, and the 
angle on the last whorl. They also differ 
much in thickness”. 

The shell (Figs. 7A-H) has the shape 
of a small trochoid, relatively solid, with a 
shagreen appearance due to minute 
punctae. Protoconch (Figs. 71-J) of about 1 
34 whorls and with about 310 um in diam- 
eter, ornamented with very small and dis- 
persed tubercles and 4-5 very fine spiral 
threads. Teleoconch of about 1 Y whorls, 
rapidly expanding; the whorls are totally 
covered by very small punctiform pits, 
clearly separated from each other, spirally 
aligned and very dense. The middle of 
the last whorl is angular, and this angle is 
almost at the periphery, fading progres- 
sively and almost disappearing near the 
aperture. In adult specimens, from the 
last Y. whorl a thickening of the inner lip 
is present extending and projecting over 
the umbilicus nearly totally covering it 
and forming the characteristic callus of 
this species (Figs. 7 E-F). This callus can 
be observed in several degrees of devel- 
opment (Fig. 7K). The different forms of 
umbilical occlusion are related to the age 
and development of the individual. 

Dimensions: Holotype 1.6 mm in 


diameter by 0.95 mm in height. Our 
largest shells measure 1.3 mm in diameter 
and 0.80 mm in height. 

Habitat: The shells studied were col- 
lected in sediments between 4 and 30 m in 
depth, on a coralline sand bottom. 

Distribution: Only known as recent 
species from Cienfuegos, Cuba. 

Remarks: T. lenticulare is a fossil 
species described from the Miocene of 
Smithfield, Virginia, Yorktown Formation. 

BusprY (1953: pl. .50, figs. 3-3d), 
figured shells of T. lenticulare with sizes 
between 1.65 mm and 1.80 mm, and com- 
pared them to the type of Lea, with the 
intention of making a comparison with 
shells of T. goniogyrus. He did not 
mention any similarity to T. incertum, 
commenting that it is less depressed than 
T. goniogyrus, which in the first half of the 
last whorl is clearly angled, rather than 
keeled, and the angulation becomes 
obscure near the labrum. The columella is 
not clearly marked towards the external 
part from the umbilical callus. 

MOORE (1964) also did not mention T. 
lenticulare. Further, he made no reference 
to T. lenticulare in his discussion of T. 
incertum only making a comparison to T. 
parvicallum. 

In this species as well as others the 
callus form is very variable, and this is 
due to different developmental stages of 
the shell. 

Teinostoma lenticulare as well as T. 
goniogyrus, T. reclususm, T. ciskae and T. 
baldingeri spec. nov. (see below), have a 
microsculpture formed by very small 
punctiform pits, clearly separated from 
each other, and spirally aligned. In con- 
trast T. incertum, has punctiform pits at 
the beginning, of the teleoconch, but they 
immediatly become incised spiral lines or 
sulci. 


lberus 23220 


ES 
y 
22999 
AA E 
JS 


Figures 7A-K. Teinostoma lenticulare (H.C. Lea, 1846). A-H: shells in several positions, 1.1-1.3 
mm, all from Cienfuegos Bay. l: protoconch; J: detail of the protoconch; K: detail of the umbilicus 
and microsculpture. 

Figuras 7A-K. Teinostoma lenticulare (H.C. Lea, 1846). A-H: conchas en diferentes posiciones, 1,1- 
1,3 mm, todas de la Bahía de Cienfuegos. I: protoconcha; J: detalle de la protoconcha; K: detalle del 


ombligo y microescultura. 


20 


RUBIO £7 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Teinostoma reclusum (Dall, 1889) (Figures 8A-G, 9A-F) 


Ethalia reclusa Dall, 1889. Bull. Mus. Comp. Zoology, 18: 361, pl. 28, figs. 7. [Type locality: 
Yucatan Strait, 640 fms (1157 m); North Carolina, 12-63 fms (22-113 m)]. 


Type material: Syntype in MCZ (007552), from off Yucatan Strait, in 640 fms (1157 m) (Figs. 8A-G). 


This shell is here designated as the lectotype. 


Other material examined: Florida, USA: 11 s, 32 mi. E St. Augustine, St. Johns Co., dredged 30 m 
(CHD); 1 s, 65 mi. ESt. Augustine, St. Johns Co., dredged 53 m (CHL); 4 s, 29 mi. E Mayport, Duval 


Cop23 mi (GEL) 


Description: This is the original 
description in DaALL (1889a): “Shell 
small, when fresh, vitreous transparent 
white, of three visible whorls, the last 
much the largest, smooth and polished 
above, or with only faint incremental 
lines below; periphery rounded, spire 
and base moderately rounded; margin 
of last whorl appressed at the suture so 
that the thin edge runs up over the pre- 
ceding whorl and the real suture is 
almost invisible in fresh specimens; the 
outline of the preceding whorl being 
visible through the shell, the appear- 
ance of a suture is presented much 
nearer the periphery than the suture 
really is. Aperture nearly circular, 
obligue; the columella thick, appressed; 
umbilical callus sparse, not polished, in 
adolescent specimens not quite com- 
pieles 

We add: The shell (Figs. 8A-D, 9A-C) 
has 3 9% whorls, 2 corresponding to the 
protoconch and 1 % to the teleoconch. 
The protoconch (Fig. 8FE, 9D) is relatively 
large, about 260 um in diameter, appar- 
ently smooth and two phases can be 
observed separated by a varix. The teleo- 
conch is totally covered by rounded 
micropits clearly separated from each 
other, spirally aligned (Fig. 8G, 9E-F). 
The periphery of the last whorl is slightly 
angled near the base. Aperture quadran- 
gular and peristome thick. Parietal callus 
wide. Columella and external lip wide 
and reflected outward. Base slightly 
convex, with a wide callus covering all 
the umbilicus. 

Dimensions: The figured lectotype 
measures 1.7 mm in maximum diameter 
and 0.9 mm in height (ratio H/D= 0.52). 

Habitat: This species is considered 
as being from deep water, having been 


described from shells collected in the 
Yucatan Strait in 1152 m depth (640 
fathoms). In North Carolina (DALL, 
1889) it was collected between 12 and 63 
fms (22-113 m), on sandy and gravelly 
bottom in the warmer area. ODÉ (1987) 
recorded it at 22 m from North Carolina. 
LEE (2009) recorded it at 65 miles east of 
St. Augustine, St. Johns Co., Florida, 
dredged at 53 m. 

Distribution: USA: North Carolina 
(JOHNSON, 1934; ODÉ, 1987a); Florida 
(LEE, 2009); Gulf of Mexico, 640 fms 
1057 m; Yucatan Strait, Gulf of Mexico, 
640 fms (1057 m) (DaLL, 1889a). 

Remarks: DALL (1889) reported the 
following: “This species is nearest to 
Ethalia diaphana d'Orbigny, so far as 
the base is concerned, but resembles E. 
anomala d'Orbigny in its upper surface, 
and was inadvertently referred to that 
species in my Preliminary Report (Bull, 
IX, p. 52). It has, however, a more ele- 
vated shell and a proportionately larger 
last whorl, while E. anomala has no 
basal callus over the umbilicus”. 

The figure in PiLsBRY (1953, pl. 56, 
fig. 5) of the holotype of T. subconicum 
(H.C. Lea) is very similar to that of the 
holotype of Ethalia reclusa Dall, differ- 
ing in the size of the callus, which does 
not totally cover the umbilicus and also 
because it lacks any microsculpture on 
the shell. 

One of the distinguishing characters 
of T. reclusum are the micropits aligned 
spirally, which completely cover the 
shell. This character was not mentioned 
by DaLL (1889) in the original descrip- 
tion but was mentioned by LEE (2009, 
fig. 328) despite the companion shell 
figure appearing a little different from 
the lectotype. 


A] 


Iberus, 29 (2), 2011 


Figures 8A-G. Teinostoma reclusum (Dall, 1889), lectotype from Yucatan Strait, 1.7 x 0.9 mm 
(MCZ 007552). A-C: optical photographs; D-E: SEM micrographs; F: protoconch; G: detail of 
the microsculpture. 

Figuras 8A-G. “Teinostoma reclusum (Dall, 1889), lectotipo del Estrecho de Yucatán, 1,7 x 0,9 mm 
(MCZ 007552). A-C: fotografías ópticas; D-E: microfotografías MEB; F: protoconcha; G: detalle de la 


microescultura. 


2 


RUBIO £7 4L.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 9A-E Teinostoma reclusum (Dall, 1889). A-C: shells, St. Augustine, St. Johns Co., Florida 
(CHL); D: protoconch; E-F: microsculpture. 
Figuras 9A-E Teinostoma reclusum (Dall, 1889). A-C: conchas, St. Augustine, St. Johns Co., Florida 
(CHL); D: protoconcha; E-F: microescultura. 


T. reclusum could be grouped with 
T. ciskae-T. goniogyrus-T. lenticulare, all 
of which have their surface covered by 
pits. 

T. ciskae is more globose and has 
fewer, larger micropits. 


From T. goniogyrus and T. lenticulare 
it differs in having a smooth protoconch, 
the lack of spiral lines of micropits and 
the peripheral keel. 

LEE (2009: 69; no. 328) figured this 
species (SEM). 


IS 


Iberus, 29 (2), 2011 


Teinostoma baldingeri spec. nov. (Figures 10A-I) 


Type material: Holotype (Figs. 10A-G) in MCZ (243769). 
Type locality: At 3-4 miles S of Fort de France, St. Louis, Martinique, in 25-29 m. 
Etymology: The specific name honors Adam J. Baldinger, Molluscs Collections Manager at the MCZ 


for his help in this paper. 


Description: Shell (Figs. 10A-C) of 
very small size, whitish in color, almost 
transparent, shining, with a low spire, 
rounded periphery and globose appear- 
ance. The minute protoconch (Fig. 101) 
has about 1 whorl, is apparently 
smooth, and measures 180 um in diame- 
ter. The teleoconch has about 2 whorls, 
the suture is distinct, the periphery 
rounded, and is totally covered by 
micropits spirally aligned. The last 
whorl covers approximately 2/3 of the 
penultimate. Aperture oblique, subcir- 
cular; columella arched. Umbilical area 
concave, umbilicus completely covered 
by a thick callus that extends from the 
columella and which is characteristic of 
the species. 

Dimensions: Holotype is 1.0 mm in 
maximum diameter. 

Habitat: Dredged in 25-29 m. 

Distribution: Only known from St. 
Louis, Martinique, the type locality. 

Remarks: Despite its small size, we 
believe that the shell studied corre- 


sponds to an adult, if we consider the 
formation of the outer lip and col- 
umella, as well as the development of 
the umbilical callus. 

Teinostoma baldingeri spec. nov. 
could be confused with other species of 
the genus Teinostoma such as T. ciskae, 
T. gontogyrus, T. lenticulare , T. anasto- 
mosis and T. reclusum, which have the 
same ornamentation, formed by micro- 
pits spirally aligned. 

T. ciskae is more globose and its 
micropits are larger. 

T. goniogyrus has a peripheral keel. 

T. lenticulare has a peripheral keel 
and a protoconch with sculpture. 

T. reclusum, is more depressed 
(ratio LM D=-0 02 as aciterent 
umbilical callus, and the  spiral 
microsculpture is formed by aligned 
micropits. 

T. anastomosis spec. nov. (see 
below) has its first whorl totally 
covered by spiral irregular interdigitat- 
ing microcordlets. 


Teinostoma incertum Pilsbry £2 McGinty, 1945 (Figures 11A-E) 


Teinostoma (Idioraphe) incertum Pilsbry £e McGinty, 1945a. The Nautilus, 59: 7, pl. 1, fig. 7. 


[Type locality: Off Destin, northwest Florida]. 


Type material: Holotype of T. incertum in ANSP (181118). Not examined. 

Other material examined: Florida, USA: 2 s, 1 m, Shoals reef, Shoals, Key West, Monroe Co. (CHL); 
1 sp and 7 s, 32 mi E. St. Augustine, St. Johns Co., 30 m. dredged (CHL); 3 s, 23 mi ENE Mayport, 
Duval Co., 28 m (CHL); 2 s, 29 mi. ESE Mayport, Duval Co., 29 m, sand shell bottom (CHL). 


Description: This is the original 
description: “The shell is depressed 
but with a low-conic spire with dis- 
tinct suture, a bluntly subangular 
periphery, microscopic spiral stria- 
tion and very little umbilical callus. 
Wiorls 3, convex, wuthimpressed 
linear suture, the peripieryloj last 
whorl very obtusely subangular. The 
base is moderately convex, concave 


24 


around the center. The oblique, circu- 
lar aperture 1s somewhat angular 
above. Peristome blunt but rather 
thin outwardly; the columella very 
thick, passing into a moderate pari- 
etal callus. Behind the columellar 
thickening an umbilical callus closes 
the umbilicus, its edge 1ll-defined 
except towards the front of the shell, 
where it terminates in a rather deep 


RUBIO ET 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 10A-I. Teznostoma baldingeri spec. nov. A-C: holotype, 1 mm, optical photographs (MCZ 
243769); D-G: holotype, SEM micrographs; H: microsculpture; l: protoconch. 
Figuras 10A-1. Teimostoma baldingeri spec. nov. A-C: holotipo, 1 mm, fotografías ópticas (MCZ 
2437069; D-G: holotipo, microfotografías MEB; H: microescultura; Í: protoconcha. 


Iberus, 29 (2), 2011 


A A A 


10 um 


Figures 11A-E. Teinostoma incertum Pilsbry 8 McGinty, 1945. A-B: shell, 1.44 mm, Pelican 
Shoals, Florida (CHL); C: protoconch; D-E: microscupture. 

Figuras 11A-E. Teinostoma incertum Pilsbry 4% McGinty 1945. A-B: concha, 1,44 mm, Pelican 
Shoals, Florida (CHL); C: protoconcha; D-E: microescultura. 


26 


RUBIO £7 4L.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


eres ema deter ico mimi event 
0.95 mm”. 

There is a better and more com- 
plete description for T. incertum in 
MOORE (1964: 88-89). 

The shell (Figs. 11A-B) is small, 
trochoid, relatively solid, with a 
shagreen appearance due to 
minute punctae. Protoconch (Fig. 
11C) of about 2 whorls and about 
380 um in diameter, ornamented 
with randomly distributed tuber- 
cles and a line of tubercles close to 
the suture, the varix at the transi- 
tion to the teleoconch is not thick- 
ened. Teleoconch of about 1 » 
whorls, increasing rapidly; whorls 
totally covered by pits in spiral 
lines connected by  shallow 
grooves which transform them into 
incised lines. Periphery subangu- 
lar, not tangulated or keeled. 
Umbilicus  totaly covered by 
numerous layers of callus originat- 
ing behind the columella. 

Dimensions: Holotype 1.6 mm 
in diameter by 0.95 mm in height. 
Our largest shells measure 1.44 
mm in diameter. 

Habitat: Marl bottom, in 32-36 
m (PILSBRY € MCGINTY, 1945a). 

Depth: 11 to 55 m. The shells 
studied were collected in sedi- 
ments obtained at 1 m near the 
base of the reef. MOORE (1964) con- 
sidered it as “a shallow shelf 
species along the Florida coasts”. 

Distribution: Known from the 
USA: East Florida, West Florida, 
Texas (PILSBRY € MCGINTY, 1945a; 
MOORE, 1964; Lyons, 1989; LEE, 
2009) Elomda and. the east of 
Brazil (Rios, 1994). 

Remarks: PILSBRY € MCGINTY 
(1945a) mention, based on the 
incomplete callus and the final 


suture, that the name “incertum” 
does not refer to the validity of the 
species but to its systematic place- 
ment. They also comment that the 
minute spiral striation is too small 
to be shown in the figure of the 
holotype, suggesting that it is not 
present in beached shells. 

It is curious to see that PILSBRY 
(1953 tin OESSON ET “AL5 11953) 
fisured shells of. T. lenticulare in 
comparison with T. goniogyrus but 
did not mention the existence of T. 
incertum, a species described by 
himself (PiLsBRY é  MCGINTY, 
1945a: 7) which has a significant 
similarity in shell shape. MOORE 
(1964) also did not mention T. 
lenticulare. Further, he considers T. 
incertumo close to 1. parvicallum, 
from which it is differentiated by 
the spiral sculpture and the deeper 
suture. The umbilical callus, which 
is projected onto the lower part of 
the peristome, can also help in the 
identification. 

We think that the characteristic 
callus of T. incertum is simply due 
to the consideration of less than 
fully-developed specimens. 

T. incertum is a species charac- 
terized by the microsculpture of 
the teleoconch, beginning with 
connected, vs. isolated, pits which 
promptly transform themselves 
into spiral lines completely cover- 
ing the shell. T. ciskae, T. gonio- 
ouas, de tenmbiculare, 0 re clususnt, 
and 1 buldingert- are different 
because the  microculpture is 
formed by discrete punctiform pits 
spirally aligned but distinctly iso- 
lated from their neighbors. 

LEE (2009: 68; no. 324) provided 
a SEM of a specimen missing some 
of its outer lip. 


Teinostoma anastomosis spec. nov. Rubio, Rolán éz Lee (Figures 12A-E) 


Type material: Holotype (Figs. 12A-C) deposited in FLMNH (448607). 
Type locality: Channel east of Seahorse Key, Cedar Keys, Levy Co., Florida, dredged 4.5-7 m. 
Etymology: The specific name is in reference to the interdigitating sculpture on the early postnu- 


clear whorls. 


2 


Iberus, 29 (2), 2011 


Figures 12A-E. Teinostoma anastomosis spec. nov. Rubio, Rolán 82 Lee. A-C. holotype, 1.79 mm, 
Channel east of Seahorse Island, Cedar Keys, Levy Co., Florida (FLMNH); D: microsculpture; E: 
microsculpture and protoconch. 

Figures 12A-E. Teinostoma anastomosis spec. nov. Rubio, Rolán (Y Lee. A-C. holotipo, 1,79 mm, 
Canal este de Seahorse Island, Cedar Keys, Levy Co., Florida (FLMNA); D: microescultura; E: micro- 
escultura y protoconcha. 


28 


RUBIO ET 4L.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Description: Shell (Figs. 12A-C) 
solid, with trochoid aspect, a little 
wider than high (H/D= 0.75), and spire 
formed by 4 whorls. Protoconch (Fig. 
12E) apparently smooth, measuring 
about 370 um in diameter, with 1 % 
whorls and with two stages, each 
delimited by a thick varix. The teleo- 
conch has 2 Y4 whorls, the suture is dis- 
tinct, the periphery rounded and 
totally covered by spiral irregular 
microcordlets (Figs. 12D-E) tend to fuse 
on the first whorl, producing micropits 
in their interspaces. Aperture rounded, 
slightly prosocline; columella thick- 
ened behind, without any canal, and 
with a callus which extends parallel to 
and behind it, partially closing the 
umbilicus. 


Dimensions: Holotype is 1.79 mm in 
maximum diameter and 1.34 mm in 
height (ratio H/D= 0.75). 

Habitat: Dredged between 4.5 to 7 m. 

Distribution: Only known from the 
type locality. 

Remarks: Teinostoma anastomosis 
spec. nov. may be distinguished from T. 
ciskae, T. goniogyrus, T. lenticulare, T. 
baldingeri and T. reclusum, because all 
these have a microsculpture formed by 
rounded micropits spirally aligned. 

T. incertum and T. panamense have 
the same ornamentation formed by 
incised spiral lines, but T. incertum has 
a subangular periphery, and T. pana- 
mense is ornamented by widely-spaced 
punctiform incisions and has a striated 
umbilical callus. 


Teinostoma panamense spec. nov. Rubio, Rolán €: Lee (Figures 13A-D) 


Type material: Holotype (Figs. 13A-B) deposited in F.MNH (448606). 


Type locality: Portobello, Panama. 


Etymology: The specific name alludes to the country where the species was collected. 


Description: Shell (Figs. 13A-B) 
solid, with trochoid aspect and spire 
slightly elevated; formed by 4 whorls. 
Protoconch (Fig. 13C) a little uncleaned 
in the sutural area, without tubercles or 
spiral sculpture, measuring about 350 
um in diameter, with 2 whorls, delim- 
ited by a weak varix. The teleoconch has 
2 whorls, is covered entirely by micro- 
pits aligned spirally, which initially are 
rounded and are closer, becoming some- 
what more punctiform incisions (Fig. 
13D). Suture covered by a thin horny 
layer uncemented. Periphery rounded, 
not keeled, angular, or subangular. 
Aperture rounded, slightly prosocline. 
Columella not thickened, separated 
from the callus by a shallow groove at 
its outer edge. Base slightly concave. A 
thick striated callus completely occludes 
the umbilicus. 


Dimensions: Holotype is 1.40 mm in 
maximum diameter. 

Habitat: Unknown. Material studied 
from drift sample. 

Distribution: Only known from the 
type locality. 

Remarks: Teinostoma panamense 
spec. nov. can be distinguished from T. 
ciskae, T. gontogyrus, T. lenticulare, T. 
baldingeri and T. reclusum because all 
of these have a microscupture formed 
by rounded micropits spirally aligned. 

T. incertum and T. anastomosis have 
the same ornamentation formed by 
incised spiral lines. But T. incertum has 
a subangular periphery and T. anasto- 
mosis is ornamented by spiral irregular 
microcordlets which are fused occasion- 
ally between them on the first whorl, 
presenting micropits in their inter- 
spaces. 


Teinostoma biscaynense Pilsbry £ McGinty, 1945 (Fig. 14A-D) 


Teinostoma (Idioraphe) biscaynense Pilsbry £ McGinty, 1945a. The Nautilus, 60: 5, pl. 1, fig. 4. 
[Type locality: Biscayne Bay at Coconut Grove, Florida]. 


Za 


Iberus, 29 (2), 2011 


Figures 13A-D. Teinostoma panamense spec. nov. Rubio, Rolán 82 Lee. A-B: holotype, 1.4 mm, 
Portobello, Panama (FLMNH); C: protoconch; D: microsculpture. 

Figuras 13A-D. Teinostoma panamense spec. nov. Rubio, Rolán € Lee. A-B: holotipo, 1,4 mm, Por- 
tobello, Panamá (ELMNAH); C: protoconcha; D: microescultura. 


30 


RUBIO £7 AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Type material: Holotype in ANSP (181104). Not examined. 

Other material examined: Florida, USA: 1 s, 50-60 mi. E Ponte Vedra, St. Johns Co., 45 m (CHL); 1 
s, Pelican Shoals, Key West, Monroe Co., 1 m, edge of reef (CHL); 1 s, 32 mi. E St. Augustine, St. 
Johns Co., 30 m (CHL); 1 £, Anclote Key, Pasco Co., sand bar (CHL). ABC: 3 s, off Palm Beach, Aruba, 
5 m (CHL). Cayman Islands: 1 s, 100 m off Seven Mile Beach, 30 m, base of coral, Grand Cayman 
(CHL). Virgin Islands: 2 s, Dead Man Reef, 18 m (CHL). Panama: 1 s, Colín Is., Bocas Islands (CEG). 
Bahamas: 3 s, South Riding Rocks, Cay Sal Bank, 28 m, base of live coral reef (CHL). Cuba: 7 s, Gua- 


jimico (MHNS). 


Description: Original description in 
PILSBRY é MCGINTY (1945a): “The 
strongly depressed shell is glossy and 
smooth except for fine weak growth- 
lines; about equally convex above and 
below, with rounded periphery and 
small umbilical callus. There are about 3 
le whorls, the first projecting, the next 
rather narrow and flat, the last whorl 
increasing very rapidly. The suture is 
distinct, visibly impressed, not obscured 
by overlaid callus. The broadly ovate 
aperture is rather strongly oblique, 
angular above. The upper margin is 
thin, arching rather strongly forward. 
The columella is rather thick, rounded, 
reflected in a broad callus covering the 
umbilicus and passing into a rather thin 
parietal callus, which is thickened in the 
posterior angle of the aperture. Diame- 
ter 1.8 mm, height 0.9 mm”. 

In our material it is possible to see 
that the largest shell has most of the col- 
umellar callus while there is a fine 
coating covering the suture. 

Habitat: It lives in shell sand in Bis- 
cayne Bay at Coconut Grove and near 
Baker”s Haulover, also on rocky sand 
bars (PILSBRY €££ MCGINTY, 1945a). Itis a 
common inshore and shallow coastal 
water species in the southeastern United 
States (MOORE, 1964). 

Distribution: Teinostoma (Idioraphe) 
biscaynense has been recorded from Bis- 
cayne Bay at Coconut Grove, Florida 
(PiLSsBRY € MCGINTY, 1945a); from East 
Florida, West Florida and Texas 
(MOORE, 1964); from Florida Peninsula 
(COOLEY, 1978); from Florida to Texas 
(EMERSON éz JACOBSON, 1976; LYONS, 
1989; LEE, 2009); from Mexico: Tabasco 
(GARCÍA-CUBAS éz REGUERO, 1990) and 
Veracruz (REGUERO ET AL., 1991); from 
Abaco, Bahamas (REDFERN, 2001). Now 
recorded from Panama and Cuba. 


Remarks: In the original descrip- 
tion, PILSBRY 6: MCGINTY (1945a) make 
reference to its similarity to T. reclusum 
in size and in the small columellar 
callus, being different because the spire 
of the latter species is more elevated, 
and the upper margin of the lip is also 
different. 

MOORE (1964: 95) remarked that he 
had examined the types of T. biscay- 
nense, T. nesaeum and T. obtectum, 
deposited in the ANSP, commenting 
that the type of de T. biscaynensis is a 
juvenile beached shell which had lost a 
great part of the dorsal callus; that of T. 
obtectum is also a beached shell but in 
better condition; finally, that of T. 
nesaeum is a specimen collected alive 
with soft parts remaining in the shell. 
After the comparison of the three types 
with hundreds of shells from Biscayne 
Bay, he commented that no differences 
between them were found except those 
related to variation in size. As for the 
spiral cordlets on the dorsum present 
in the shells of T. nesaeum, he did not 
consider them an important taxonomic 
character, making reference to them as 
“extremely evanescent”. Thus, he con- 
cluded that T. biscaynense, T. obtectum 
and T. nesaeum were the same species 
giving T. biscaynense, the first species 
published in the same work, priority. 

We do not agree with this conclu- 
sion, and, as we will show in the 
description and figures, each one has 
constant taxonomic characters sufficient 
to consider them as valid species just as 
they were described by PiLSBRY « 
MCGINTY (1945a). The shells pho- 
tographed agree perfectly with the 
material described and figured by 
PILSBRY € MCGINTY (1945a: fig. 4). 

MOORE (1964) also stated that T. bis- 
caynense is different from the other 


3] 


Iberus, 29 (2), 2011 


Figures 14A-D. Teinostoma biscaynense Pilsbry 82 McGinty, 1945. A: shell, 1.3 mm, Florida 
(CHL); B-C: shells, 1.2, 1.36 mm, Guajimico, Cuba (MHNS); D: protoconch, from Cuba. 
Figuras 14A-D. Teinostoma biscaynense Pilsbry e McGinty, 1945. A: concha, 1,3 mm, Florida 
(CHL); B-C: conchas, 1,2, 1,36 mm, Guajimico, Cuba (MHNS); D: protoconcha, de Cuba. 


species of the genus Teinostoma from 
shallow water in the West Indies 
because it has the spire totally covered 
by a fine callous coat. Also, this charac- 


32 


ter made it similar to T. cryptospira (= T. 
umbilicatum), a species from deep 
water off Cape Hatteras, North Car- 
olina. 


RUBIO ET 4L.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


We do not agree with this because 
PiLSBRY € MCGINTY (1945a) stated in 
their original description: “The suture is 
distinct, visibly impressed, not obscured 
by overlaid callus”. In relation with the 
protoconch: “There are about 3 Y 
whorls, the first projecting, the next ...” 
Based on this passage, the spire of T. bis- 
caynense cannot be totally covered by a 
callous coating, as is emphasized by 


MOORE (1964: 96, 98). This discrepancy 
may reflect an error in the identification 
of the examined types. 

T. biscaynense differs from a group 
of species formed by T. umbilicatum 
(=T. cryptospira), T. nesaeum, T. obtec- 
tum, T. lerema and T. clavium, because 
in these a fine callous coat covers the 
spire, partially or totally, the protoconch 
being hidden in some of them. 


Teinostoma obtectum Pilsbry € McGinty, 1945 (Figures 15A-B) 


Teinostoma (Idioraphe) obtectum Pilsbry €£z£ McGinty, 1945a. The Nautilus, 59: 6, pl. 1, fig. 6. 
[Type locality: “Treasure Island”, the first islet south of Singer Bridge, northern end of Lake 


Worth, Palm Beach, Florida]. 


Type material: Holotype in ANSP (181121). Not examined. 
Material examined: Florida, USA: 1 s, 29 mi. ESE Mayport, Duval Co., Fl., 23 m(CHL); 1 s, just S 
jetty, Anastasia Island, St. Augustine Beach, St. Johns Co. (CHL); 1 s, beach, Indian Pass, Port St. 


Joe, Gulf Co. (CHL). 


Description: This is the original 
description of PILSBRY € MCGINTY 
(1945a): “The moderately solid smooth 
shell is strongly depressed, trans- 
versely dilated, the spire covered with 
a translucent glaze through which the 
suture shows. About three rather 
rapid ly increasing but regularly spiral 
whorls are visible through the sub- 
transparent callous coat over the spire, 
which superficially shows no trace of 
the suture. The periphery is rounded, 
the base not very convex. Aperture is 
rounded, but angularly produced and 
slightly channelled above and with a 
flattened parietal outline. Outer margin 
of peristome thin, the concave col- 
umella rather thick, passing into the 
rather large and slightly convex umbili- 
cal callus. Parietal callus is rather thick. 
Diameter 2.2 and 1.65 mm, height 0.95 

Maximum reported size: 2.2 mm 

Habitat: Shell sand bottom (PILSBRY 
$ MCGINTY, 1945a). Bathymetric range 
0 to 500 m. 

Distribution: USA: Florida: East 
Florida (PILSBRY éz MCGINTY, 1945a; 
MOORE, 1964: 4; LEE, 2009: 68); Mexico: 
Campeche State, Yucatan State, Quin- 
tana Roo (VOKES é€ VoOKeEs, 1984); 


Venezuela: unlocalized (Princz, 1982a); 
Puerto Rico (WARMKE éz ABBOTT, 1961). 

Remarks: After the description of the 
species PILSBRY é MCGINTY (1945a) 
mention: “The elliptical outline, the 
strong depression, and the callus 
smoothly covering the spire, distinguish 
this species, which is known by a single 
shell. A small nick in the outer lip was 
restored in the figure”. 

MOORE (1964: 97) stated: “The types 
of Teinostoma biscaynensis, T. nasaeum 
and T. obtectum have been examined by 
the writer. That of T. biscaynensis is a 
worn dead shell which has lost most of 
the dorsal shelly callus. The type of T. 
obtectum is also a dead shell, but is in 
much better condition. It is near the 
maximum size of the species. The type 
of T. nasaeum was taken alive, and the 
soft parts still remain in the shell. The 
writer has compared all three types with 
each other and with several hundred 
specimens from Biscayne Bay, and can 
find no differences other than those 
resulting from wear and tear or varta- 
tion in size. As T. biscaynensis is the 
jurst aspectes listed. 1n- PIESBRY 6 
MCGINTY (19454), it is given page prece- 
dence, and the other two species are 
placed in synonomy”. 


39 


Iberus, 29 (2), 2011 


Figure 15A-B. Teinostoma obtectum Pilsbry £ McGinty, 1945. A-B: shell, 1.9 mm, Port St. Joe, 
Gulf Co, Florida (CHL). 
Figura 15A-B. Teinostoma obtectum Pilsbry e McGinty, 1945. A-B: concha, 1,9 mm, Port St. Joe, 
Gulf Co, Florida (CHL). 


We can not agree with this opinion, 
because the descriptions and the figures 
of the original papers given by PILSBRY 
éz MCGINTY (1945a) of these three 
species synonymized by MOORE (1964) 
have been enough to identify the mater- 
ial studied in them. Our only explana- 
tion is that there was mixing of the type 
material deposited in the museum. The 
shells photographed agree perfectly 
with the material described and figured 
by PILSBRY € MCGINTY (1945a: fig. 6). 


There is a real confusion between T. 
biscaynense and T. obtectum because 
the latter species was described from a 
single shell and also because neither 
was figured with its original descrip- 
tion. They were then placed in syn- 
onymy by MOORE (1964), whose judge- 
ment was accepted. T. obtectum differs 
from T. biscaynense in having the spire 
totally covered by a fine callous coating, 
and also in the shape of the umbilical 
callus. 


Teinostoma expansum spec. nov. (Figures 16A-G) 


Type material: Holotype (Fig. 16A) deposited in MNCN (15.05 /55063). Paratypes in USNM (1155031, 


1 s, Fig. 168) and MNHN (24402, 1 s, Fig. 16C). 


Other material examined: Cuba: 2 sp and 4 s, Cienfuegos Bay, 22"07'N 80%27'W, 9 m (MHNS). 


Type locality: Cienfuegos Bay, Cuba. 


Etymology: The specific name refers to the expansion of the aperture. 


34 


RUBIO £7 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


um 


e 


Figures 16A-G. Teinostoma expansum spec. nov. A: holotype, 1.1 mm (MNCN). B-C: paratypes, 1.2, 


1.3 mm, Cienfuegos, Cuba; D: protoconch; E: detail of the umbilical callus; F: operculum; G: radula. 
Figuras 16A-G. Teinostoma expansum spec. nov. A: holotipo, 1,1 mm (MNCN). B-C: paratipos, 1,2, 1,3 
mm, Cienfuegos, Cuba; D: protoconcha; E: detalle del callo umbilical; E: opérculo; G: rádula. 


Description: Shell (Figs. 16A-C) 
depressed, the spire totally covered by a 
fine callous surface which even hides 
the protoconch (Fig. 16D). By transillu- 
mination 3 Y spiral whorls can be seen, 
the last one rapidly expanding and 
smooth except for fine growth lines. 


Aperture ovoid, oblique and wide; 
external lip sharp, projected outward. 
Columella (Fig. 16E) wide, rounded, 
reflected in a large callus which cover 
the umbilicus. 

Dimensions: The largest shells 
studied were 1.3 mm in diameter. 


35 


Iberus, 29 (2), 2011 


Operculum (Fig. 16F) multispiral 
with a central nucleus. 

Radula (Fig. 16G) taenioglossate, 
with formula 2+1+R+1+2. Central tooth 
wide basally, the ventral margin without 
denticles. Cutting area formed by a 
main large and sharp cusp and 4 denti- 
cles of lesser size on each side. Lateral 
teeth similar to the central one, the bases 
are quadrangular and also without den- 
ticles; border area with a central cusp 
and 5 smaller denticles at each side, 
more elongated wich central tooth. Mar- 
ginal teeth wide and elongate; the inner 
with 28-30 slight denticles on the cutting 
edge is hook shaped; the outer marginal 
teeth are inclined outward in their upper 
third and have 14-18 denticles on the 
upper end of their internal margins. 


Habitat: Our material was collected 
in about 9 m depth. 

Distribution: Only known from Cien- 
fuegos Bay, its type locality. 

Remarks: Teinostoma expansum is 
close in shell characters to T. biscaynense, 
T. obtectum and T. lerema, all them char- 
acterized by having the spire covered 
totally or partially by a fine callous coat. 

Teinostoma biscayense lacks this fine 
coat and the spire is free, thus allowing 
the protoconch to be photographed 
easily. 

Teinostoma obtectum has a much 
more depressed shell, and the aperture is 
more deflected and oblique. 

Teinostoma lerema has the aperture 
slightly oblique, grooved in the upper 


internal angle. 


Teinostoma nesaeum Pilsbry €£ McGinty, 1945 (Figures 17A-F) 


Teinostoma (Idioraphe) nesaeum Pilsbry € McGinty, 1945a. The Nautilus, 60: 5-6, pl. 1, fig. 2. 


[Type locality: Missouri Key, Florida]. 


Type material: Holotype in ANSP (181117). Not examined. 
Material examined: Cuba: 14 s, Guajimico, 15 m (MHNS); 1 s, Bahía de Cienfuegos, 20-30 m (MHNS); 


15 s, Rancho Luna Beach, 30 m (MHNS). 


Description: The original descrip- 
tion in PILSBRY € MCGINTY (1945a: 5-6) 
is as follows: “The shell is rather 
strongly depressed, about equally 
conver above ana elote 
rounded periphery; glossy, with some 
spiral striae on the upper surface, 
none on the lower. There are appar- 
ently about 2 Y whorls, but the sutures 
are obliterated by a coat of translucent 
callus which covers the spire, with a 
shallow impression over the apex. The 
upper surface shows shallow spiral 
striae which weaken towards the 
periphery and disappear on the base. 
The aperture is oblique, rounded, but 
acute a timneuppenmanale ole 
however is filled, making the cavity 
round. Outer lip blunt. Columella 
thickened, passing into the rather 
thick parietal callus. Umbilical callus 
thick, slightly convex, a trifle rugose”. 
The shells are represented in Figs. 
17A-F) 


36 


Dimensions: Holotype 1.45 mm on 
maximum dimensions (diameter). The 
largest shell in our material is 1.4 mm. 

Habitat: Living under stones and in 
rocky sandbars (PILSBRY éz MCGINTY, 
1945a: 6). Mangrove swamps, sandy 
and rocky areas (VOKES € VOKES, 
1984). Shallow, soft bottoms (DÍAZ 
MERLANO éz PUYANA HEGEDUS, 1994). 

Distribution: The species has been 
collected from Missouri Key, Florida 
and Biscayne Bay near Baker's 
Haulover, Miami (PILSBRY € MCGINTY, 
1945a:6). From Colón and Bocas island, 
Panama (OLsson € MCGINTY, 1958). 
From South and North Carolina to the 
Caribbean Sea (HOUBRICK, 1968). From 
Campeche to Carmen and Zacatal 
cities, from Ninum Point to Campeche, 
from El Cuyo to Ninum Point, from 
Point Yalcupul to Cerritos island and 
from isla Mujeres to Isla Holbox, 
Mexico (VOKES éz VOKES, 1984). From 
Curacao, Aruba and Bonaire (DE JONG 


RUBIO ET AZz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 17A-E Teinostoma nesaeum Pilsbry 82 McGinty, 1945, shells, 1.2, 1.2, 1.2, 1.1, 1.0, 1.4 
mm, Rancho Luna Beach, Cuba (MHNS). 
Figuras 17A-E Teinostoma nesaeum Pilsbry  McGinty, 1945, conchas, 1,2, 1,2, 1,2, 1,1, 1,0, 14 
mm, Playa Rancho Luna, Cuba (MHNS). 


$ COOMANS, 1988). From North Carolina 
to Panama and. Colombia (DÍAZ 
MERLANO € PUYANA HEGEDUS, 1994). 
Also found in Cuba: Cienfuegos. 
Remarks: MOORE (1964: 95-99) placed 
the taxa T. nesaeum and T. obtectum in 
synonymy with T. biscaynense (see 
above). We consider T. nesaeum a valid 
species, easily differentiable from its con- 
geners by the following characters: 


dorsal ornamentation of fine spiral 
cordlets; base smooth; umbilical callus 
strong; suture covered by a translucent 
callus coat. These characters were also 
considered by PiLsBRY é  MCGINTY 
(1945a) as important for species separa- 
tion. 

All the shells studied have the 
cordlets on the dorsum but not on the 
base. We have not found intergradations. 


3% 


Iberus, 29 (2), 2011 


Teinostoma semistriatum (d'Orbigny, 1842) (Figures 18A-I, 19A-F) 


Trochus (Rotella) semistriata d'Orbigny, 1842. Mollusques. Histoire Physique, Politique et Natu- 
relle de lVíle de Cuba 2: 61, pl. 18, figs. 20-22. [Type locality: Cuba] 

Pseudorotella semistriata (4'Orbigny): In P. Fischer, 1957. Journal de Conchyliology 6: 52. 

Teinostoma (Idioraphe) clavium Pilsbry €: McGinty, 1945a. The Nautilus, 60: 4, pl. 1, fig. 1. [Type 
locality: Tavernier, Key Largo]. 


Type material: Two syntypes in NHMUK (in very bad condition). Neotype here designated of 
Trochus (Rotella) semistriata in MCZ (208142), from La Chorrera, Habana, Cuba. Type species of 
Teinostoma (Idioraphe) clavium in ANSP (181106). Not examined. 

Other material examined: Guadeloupe: 1 s, coralline sandy and rocky bottom, 2 m (CJP). Cuba: 5 
s, Matanzas, Varadero (N Cardenas) (MCZ 109344); 7 s, Archipielago de los Canarreos, 15 m (MHNS); 
8 s, Cayo Diego Perez, 12 m (MHNS); 19 s, Jibacoa, 3-6 m (MHNS); 1 s, Cienfuegos Bay, 20-30 m 
(MHNS); 48 s, Rancho Luna Beach, 20 m (MHNS); 2 s, Comodoro, 0 m (MHNS); 4 s, playa Girón, 
5 m (MHNS). Florida USA: 1 s, Peanut Island, Lake Worth, Palm Beach Co. (CHL); 3 s, Virginia 
Key, Dade Co. (CHL); Spoil bank, APAC Pit, Sarasota Plio-Pleistocene (CHL). Bahamas: 2 s, Matt 
Lowes Cay, Abaco, grit (CHL); 1 s, Paradise Island, New Providence, 2 m (CHL); 6 s, West End, 
Grand Bahama Island, grit (CHL). Puerto Rico: 1 s, San Juan, grit (CHL). St. Martin: 15 s, Leeward 


Island, grit (CHL). Virgin Islands: 1 s, Frederiksted, St. Croix, grit (CHL). 


Description: This is the original 
description in D'ORBIGNY (1842): “Shell 
orbicular, depressed, thin, transparent, 
white, above transversely (1.e. concen- 
trically) striae, beneath polished; umbil- 
ical callus shining; spire very short, 
obtuse, whorls four, slightly convex; 
aperture oval. Diameter 1.5 mm; height 
0.7 mm”. 

Shell (Figs. 18A-F, 19A-E) strong, 
solid, somewhat depressed, totally 
covered by spiral cords. Umbilical 
callus wide (Fig. 18C). Protoconch (Figs. 
18G-H, 19F) with about 1 Y whorls, 
smooth, about 180 um, partially 
covered by the first whorl of the teleo- 
conch so only the apex of the proto- 
conch is visible. Teleoconch with about 
2 Y4 whorls, covered with spiral cords 
lacking sculpture in their interspaces 
(Fig. 18D); periphery rounded. Umbilical 
callus strong and wide, completely cov- 
ering the umbilicus. Aperture oblique, 
with a small groove on the upper inter- 
nal angle which is somewhat extended 
giving the shell an elongate aspect. 

Dimensions: Holotype of T. clavium 
2.3 mm in maximum diameter. 

Habitat: This species lives in 
shallow water between 0 and 18 m 
depth. In Cuba it is relatively common, 
1t was collected in sediments from 
between 5 and 30 m. 


38 


Distribution: Known from USA: 
Florida: East Florida, Florida Keys 
(JOHNSON, 1934; PILSBRY é MCGINTY, 
1945a; MOORE, 1964; LEE, 2009); Mexico: 
Campeche State, Quintana Roo (ODÉ, 
1987); Colombia, Venezuela: Sucre (DÍAZ 
MERLANO é PUYANA HEGEDUS, 1994; 
Princz, 1986); Bahamas: New Provi- 
dence (MOORE, 1964), Abaco (REDFERN, 
2001); Puerto Rico (WARMKE éz ABBOTT, 
1961); Cuba (OrRBIGNY, 1842; P. FISCHER, 
1857; ESPINOSA ET AL., 1985). Now 
recorded from Guadeloupe. 

Remarks: Nomen dubium according 
to MOORE (1964: 81-82) who com- 
mented: “The types of Pseudorotella, T. 
semistriata d'Orbigny, are in the British 
Museum (Natural History), but both 
types have deteriorated badly and the 
principle characters can not be made 
out. There has been a great deal of con- 
fusion with d'Orbigny's species for over 
a century, and, without an authentic 
specimen of T. semistriata to compare 
with other species, the true characters of 
the subgenus must remain in doubt. It is 
impossible to say which of d'Orbigny's 
(1842) species belong to the genus, and, 
since the types are no longer identifi- 
able, they are here considered species 
indeterminate”. 

We have seen this type material in 
the NHMUK, and it is in very poor con- 


RUBIO ET 41£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 18A-1. Teinostoma semistriatum (d'Orbigny, 1842). A-F: shells, 1.6, 0.9, 1.6, 1.1, 2.1, 1.1 
mm, Cienfuegos, Cuba (MHNS); G: protoconch; H: detail of the protoconch; I: detail of 
microsculpture. 

Figuras 18A-1. Teinostoma semistriatum (4'Orbigny 1842). A-F: conchas, 1,6, 0,9, 1,6, 1,1, 2,1, 
1,1 mm, Cienfuegos, Cuba (MHNS); G: protoconcha; H: detalle de la protoconcha; I: detalle de la 


microescultura. 


3% 


Iberus, 29 (2), 2011 


Figures 19A-E. Teinostoma semistriatum (d'Orbigny, 1842). A-C: neotype, 2.1 mm, La Chorrera, 
La Habana (MCZ 208142); D-E: shells, 1.8, 2.2 mm, Matanzas, Cuba (MCZ); F: protoconch. 
Figuras 19A-E Teinostoma semistriatum (4'Orbigny 1842). A-C: neotipo, 2,1 mm, La Chorrera, La 
Habana (MCZ 208142); D-E: conchas, 1,8, 2,2 mm, Matanzas, Cuba (MCZ); F: protoconcha. 


dition, but in our opinion the taxon is 
not a nomen dubium. 

In the type material of the Museum 
of Comparative Zoology (MCZ) there 
were two lots labeled: “Pseudorotella 
semistriata (Orb.), 5 specimens from 


40 


Matanzas, Varadero, Cuba” deposited 
with n” 109344, which were sent by M. 
L. Jaume, a well-known Cuban malacol- 
ogist, and another “Teinostoma semistri- 
ata d'Orb., 1 specimen from La Chor- 
rera, Habana, Cuba,” with n* 208142. We 


RUBIO ET ALz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


studied them closely, and they conform 
to the description of  Trochus 
semistriata. This species seems to be rel- 
atively common along the Cuban coast, 
having being found in the beached shell 
grit. 

In SAGRA (1842: 177) Orbigny men- 
tions in relation to Rotella semistriata: 
“It lives in Cuba, together with the 
former (R. diaphana). It can be found in 
large numbers in the sands of Playa del 
Chivo y of La Chorrera; found also in 
St. Thomas”. 

For these reasons and in order to 
keep nomenclatural stability, we have 
designated as neotype the specimen 
from La Chorrera, Habana, deposited in 
MCZ with n” 208142 considering T. 
clavium Pilsbry € McGinty, 1945 a 
junior synonym. 

In the original description 
d'Orbigny also mentions: “beneath pol- 
ished”. This means the origin of the 


material employed for this description 
was beached and eroded; those shells 
have the base totally smooth and pol- 
ished due to abrasion. However, as we 
can observe in fresh shells, the base is 
totally covered by spiral cordlets as is 
the dorsum. 

Teinostoma  semistriatum is a 
species easily distinguished from its 
congeners by its robustness and the size 
of the shell, the form of the umbilical 
callus, and the ornamentation of the 
teleoconch with spiral cords which 
completely cover the dorsum. The pro- 
toconch is partially concealed by the 
first whorl of the teleoconch. As in pre- 
vious species, it can form a group with 
T. umbilicatum, T. biscaynense, T. nes- 
saeum and T. lerema, because in all of 
them the last whorl is extended cover- 
ing some or all of the previous whorls. 
MOORE (1964) mentions its similarity to 
T. biscaynense and T. lerema. 


Teinostoma minusculum (Bush, 1897) (Figures 20A-D) 


Pseudorotella minuscula Bush, 1897. Transactions of the Connecticut Academy of Arts and Sci- 
ences 10: 118-119, text-figs. 3a-c. [Type locality: USFC sta. 2283, off Cape Hatteras, North Car- 


olina]. 


Type material: Holotype in USNM (41623) is a broken shell. Examined in micrograph (Fig. 20A). 
Other material examined: USA: 1 s, off Dry Tortugas, Monroe Co., Florida, 20-50 fms (USNM); 1 s 
(more deteriorated) ENE Mayport, Duval Co., Florida, 26 m (CHL). 


Description: Original description 
in BusH (1897): “Shell thick, solid, por- 
cellanous, slightly tinted with yellow 
along the suture and on the base; flat- 
tened above and below, with the 
indented umbilical region covered 
with a thin lustrous glaze or layer of 
enamel. Surface smooth and very lus- 
trous, marked only by irregular, 
microscopie, cerotota. limes.  suture 
inconspicuous. Whorls about 2 Y%, 
coiled in the same plane, lapping well 
on to each other, rapidly enlarging, 
with a very small nuclear whorl and 
large body-whorl. Aperture very 
obligue, somewhat ovate; peritreme 
not continuous, modified into a thin, 
inconspicuous glaze on the body- 
whorl, elsewhere with rounded edge, 


with a slight callous deposit beneath 
the suture where the outer-lip extends 
obliquely well forward from the body- 
whorl, with little, 1f any, curvature 
and forms a slight sutural notch. 
Greatest diameter, about 1.5 mm; 
height, about 0.5 mm”. T. minusculum 
has the umbilical region entirely 
covered by a thin, very lustrous glaze 
or layer of enamel, not in any sense a 
thickened pad as in Teinostoma umbil- 
icatum. 

Habitat: Depth: 14 to 50 fms. 

Distribution: USA: North Carolina 
(Bush, 1897); Florida: East Florida 
(LEE, 2009) 

Remarks: This species was 
described in the subgenus 
Pseudorotella P. Fischer, 1857. BusH 


41 


Tberus 2D DONA 


Figures 20A-D. Teinostoma minusculum (Bush, 1897). A: holotype, 1.5 mm, (USNM, 41623); B- 
D: shell, 1.6 mm, Dry Tortugas, Monroe Co., Florida (CHL). 
Figuras 20A-D. Teinostoma minusculum (Bush, 1897). A: holotipo, 1,5 mm, (USNM, 41623); B- 
D: concha, 1,6 mm, Dry Tortugas, Monroe Co., Florida (CHL). 


(1897) says: “This species approaches 
Teinostoma cryptospira (A.E. Verrill) 
Dall, but it is a much smaller shell, 
with the whorls quite differently 
coiled and with the umbilical callus 


42 


represented by a thin glaze”. Really, 
the only similarity with Teinostoma 
cryptospira is that both have the spire 
covered by a fine callous covering. LEE 
(2009: 68; no. 326) figured this species. 


RUBIO £7 AZ.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Teinostoma lerema Pilsbry £ McGinty, 1945 (Figures 21A-H) 


Teinostoma (Idioraphe) lerema Pilsbry £ McGinty, 1945a. The Nautilus, 60: 6-7, pl. 2, figs. 1-1a. 
[Type locality: Missouri Key, Florida Keys]. 


Type material: Holotype in ANSP (181120). Not examined. 

Other material examined: Cuba: 3 s, Cienfuegos Bay, 22%07'N, 80%27'W, 9 m (MHNS); 7 s, Cien- 
fuegos Bay, 2207'N80"26'W, 4 m (MHNS). Martinique: 1 s, Pointe Borgnesse, 12 m, sand-muddy 
bottom, close to the reef (CJP). St. Kitts £ Nevis: 4 s, Monkey Shoals, Nevis Island, 18 m (CHL). 
Haiti: 1 s, Labaree, sand beach (CHL). Puerto Rico: 1 s, San Juan, grit (CHL). Panama: 3 s, Isla Caren- 
eros, 8-9 m (CHL); 1 s, 1 mi. Punta San Blas, San Blas Island, sand bar just inside reef (CHL). 


Description: Shell (Figs. 21A-G) very 
small, solid, smooth and shiny. Proto- 
conch (Fig. 21H) smooth, with about 2 
whorls, and a diameter of about 200 um, 
but usually partially covered by the first 
teleoconch whorl. The teleoconch has a 
little more than one whorl and is 
smooth except for fine growth lines. 
Suture covered by a fine callous coat. 
Umbilicus totally covered by a thin 
callus. Aperture oblique, rounded, 
without any sulcus on the upper inter- 
nal angle; the outer lip is extended 
outward, giving the shell a more elon- 
gated aspect. 

Dimensions: Holotype 1 mm in 
maximum diameter and 0.45 mm in 
height. The largest of our shells is 1.0 
mm in diameter. 

Animal figured by PILSBRY é 
MCGINTY (1945a, pl 2, fig. la). 

Habitat: Living under rocks (PILSBRY 
éz MCGINTY, 1945a). Collected alive 


under stones between 0.6 and 16 m in 
depth. The Cuban shells were found in 
sediments from between 4 and 9 meters. 
Records between 0 and 48 m. 

Distribution: Know from the USA: 
West Florida, Missouri Key (PILSBRY éz 
MCGINTY, 1945a); Texas (ODÉ, 1987); 
Mexico: Tabasco, Veracruz, Campeche 
State (GARCÍA-CUBAS, 1971); Colombia 
(Díaz MERLANO é PUYANA HEGEDUS, 
1994); Virgin Islands: St. John (MOORE, 
1964), Curacáo (DE JONG € COOMANS, 
1988); Cuba (SARASÚA, 1970; ESPINOSA 
ET AL., 1985). 

Remarks: It is the smallest Teinos- 
toma described up to now. Its small size, 
the suture covered by a fine callous 
layer and the elongated form of the 
aperture differentiate it from congeneric 
species. Teinostoma lerema has a certain 
similarity to T. biscaynense in general 
form, but the latter has the spire totally 
covered by a fine callus. 


Teinostoma umbilicatum (H.C. Lea, 1843) (Figures 22A-G) 


Rotella umbellicata H.C. Lea, 1843. Proc. Amer. Philos. Soc., 3: 164. (Nude name). 

Rotella umbilicata H.C. Lea, 1846. Trans. Amer. Philos. Soc. (new series) 9: 264, pl. 36, fig. 80. 
[Type locality: Miocene of Petesburg]. 

Rotella cryptospira A.E. Verrill, 1884. Transactions of the Connecticut Academy of Arts and Sci- 
ences, 6: 241-242 (not figured). [Type locality: USFC sta. 2109, off Cape Hatteras, North Car- 
olina]. 


Type material: Lectotype of Rotella cryptospira USNM 35731; it was selected and figured by JOHNSON 
(1989). MCZ Publications on Mollusks Occasional Papers on Mollusks, 5(67): 32, pl. 10, fig. 8. Not 
examined. 

Material examined: Cuba: 5 s, Rancho Luna Beach, 20-54 m (MHNS); 6 s, Cienfuegos Bay, 10 m 
(MHNS). Florida, USA: 1 s, 23 mi. ENE Mayport, Duval Co., 26 m (CHL); 1 s, Jacksonville Beach, 
Duval Co. grit (CHL); 2 s, Anclote Key, Pasco Co. sand bar (CHL); 7 s, channel E Seashore Key, 
Cedar Key, Levy Co. 4-6 m (CHL); 2 s, Louisiana: off western part, 22 m (CHL). Jamaica: 3 s, Priory, 
St. Anr's Parish, dredged shallow water (CHL). Trinidad and Tobago: Tobago: 7 s, Horse Shoe reef, 
15 m, shell sand (CJP). 


A3 


Iberus, 29 (2), 2011 


Figures 21A-H. Teinostoma lerema Pilsbry 82 McGinty, 1945; shells 1.0, 0.8, 0.7, 1.0, 0.8, 0.7, 0.9 
mm, Rancho Luna Beach, Cienfuegos, Cuba (CEG). 
Figuras 214-H. Teinostoma lerema Pilsbry 4 McGinty, 1945; conchas, 1,0, 0,8, 0,7, 1,0, 0,8, 0,7, 
0,9 mm, Playa Rancho Luna, Cienfuegos, Cuba (CFG). 


Description: Shell (Figs. 22A-C, 22E- 
G) strongly depressed, smooth, rapidly 
expanding. A fine callous layer covers a 
great part of the previous whorl, 
keeping visible the protoconch and a 


44 


small part of each whorl (Fig. 22D). 
Periphery very rounded. Aperture 
slightly oblique, rounded, grooved in 
the upper internal angle. Columella 
strong, reflected towards the external 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 22A-D. Teinostoma umbilicatum (H.C. Lea, 1843). A-C: shells, 1.9, 1.9, 1.6 mm respec- 
tively, Cienfuegos Bay, Cuba; D: protoconch; E-G: 1.8, 1.6, 1.3 mm, Tobago (CHL). 

Figuras 22A-D. Teinostoma umbilicatum (A.C. Lea, 1843). A-C: conchas, 1,9, 1,9, 1,6 mm respec- 
tivamente, Bahía de Cienfuegos, Cuba; D: protoconcha; E-G: 1,8, 1,6, 1,3 mm, Tobago (CHL). 


part forming a strong and characteristic 
callus which completely covers the 
umbilicus. 

Dimensions: The lectotype is 2.5 mm 
in maximum diameter. The largest shell 


examined was 1.93 mm in diameter and 
1.07 mm in height. 

Habitat: A species of wide bathymet- 
ric distribution, recorded between 18 
and 305 m depth. DaLL (1892) collected 


45 


Iberus, 29 (2), 2011 


it alive off the coasts of North Car- 
olina and Florida in 30 to 50 fathoms, 
U.S. Commission. Our material was 
collected on coralline sandy bottoms 
between 10 and 54 m. 

Distribution: Recorded from the 
USA: New Jersey, Campeche, North 
Carolina and Florida (A.E. VERRILL, 
1884; DaALL, 1892; LEE, 2009); from 
Mexico: Campeche State (ODÉ, 1987a); 
Venezuela (PRINCZ, 1982) and Cuba 
(ESPINOSA ET AL., 1985). 

Remarks: VERRILL (1884) says: 
“This species bears some resemblance 
to Rotella anomala D'Orbigny, but is 
peculiar in having the whorls of the 
spirecomceated o nimeanly aso 20 ele 
last whorl”. DALL (1892) mentions: 
“This little shell resembles T. umbili- 
catum Lea in having the whorls nearly 
concealed by the thinned-out edge of 
e precede ono Miel es 
appressed nearly to the apex. The 
surface is smooth and polished. The 
fossils have been identified by com- 
parison with a specimen named by 


the author, who has not yet figured 
his species”. PiLSBRY (1953: 416, in 
OLSSON ET AL., 1953) placed T. cryp- 
tospira in the subgenus Idioraphe 
Bilsbry 192241? eS eos 
characterized because the last whorl 
envelopes all of those preceeding, or 
leaves only the apical whorl exposed. 
The suture is developed only as an 
arcuate or angular line radiating from 
summit to periphery”. In this sub- 
genus are included T. umbilicatum, T. 
verrilli O. Meyer, 1885 and T. nanum 
H.C. Lea, 1833. PiLsBRY (1953) men- 
tions that in many shells observed the 
callous coat in the sutural border of 
the last whorl almost reaches the apex 
or only the apex is free. The type of T. 
umbilicatum is broken, but the apical 
area 1s preserved in good condition. 
The T. umbilicatum group of teinos- 
tomes has continued to the present 
day in species only very slightly dif- 
ferent from the ancestral form, the 
living representative being named T. 
cryptospira. 


Teinostoma altum Pilsbry, 1953 (Figures 23A-D) 


Teinostoma (Pseudorotella) altum Pilsbry, 1953 (in OLSSON ET AL., 1953). Acad. Nat. Sci. 
Philadelphia, Monographs 8: 413, pl. 49, fig. 2-2f. [Type locality: Plio-Pleistocene of North St. 


Petersburg, Florida]. 


Type material: Holotype in ANSP (18398). Not examined. 
Other material examined: Cuba: 2 s, Bahía de Cienfuegos, 20-30 m (MHNS). 


Description: The original descrip- 
tion in OLSSON ET AL. (1953) is as 
follows: “The shell is solid, depressed 
globose with conic spire, smooth 
surface, very incomplete umbilical 
callus divided from the columella by a 
groove. It is often perforate, or the 
umbilicus may be closed. There are 
fully four moderately convex whorls 
united by a distinctly impressed suture. 
The last whorl is broadly rounded at 
periphery, the base strongly convex, 
excavated around the narrow umbilical 
crevice (which is often closed). The 
aperture is nearly circular, but angular 
above. Peristome is rather long and 
evenly concave, somewhat thickened, 


46 


separated from the callus by a shallow 
groove at its outer edge. The umbilical 
callus is a rather small, convex, lunate 
lobe, which typically does not wholly 
close the umbilicus”. 

We show some images of shells 
(Figs. 23A-C) and protoconch (Fig. 
23D). We must point out these distin- 
guishing characters of the species: the 
elevated spire, the rounded periphery, 
the groove which separates the col- 
umella from the umbilical callus and 
that this callus does not always cover 
the umbilicus. 

Dimensions: Holotype 2.2 mm in 
maximum diameter and 1.6 in height. 
Our largest shell is 1.50 mm in diameter 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 23A-D. Teinostoma altum Pilsbry, 1953. A-C: shells, 1.3, 1.4, 1.3 mm, Cienfuegos Bay, 


Cuba. D: protoconch. 


100 um 


Figuras 23A-D. Teinostoma altum Pilsbry, 1953. A-C: conchas, 1,3, 1,4, 1,3 mm, Bahía de Cienfue- 


gos, Cuba. D: protoconcha. 


and 1.15 mm in height (ratio H/D: 
0.76). 

Habitat: The only two shells found 
were from shell grit between 20 and 30 m 
in depth. Elsewhere in the literature it is 
reported from 0 to 139 m. 

Distribution: Florida, USA: Fossil, 
from the Plio-Pleistocene of St. Petersburg 
(Pilsbry, 1953; Odé, 1987) Recent of: Cuba: 
Cienfuegos. USA: Georgia: 57 mi E Sapelo 
Is. 18-20 m (Lee, 2009). Florida: 28 mi E 
ESE Mayport, Duval Co. 22.5 m (LEE, 
2009). 

Remarks: Fossil species described 
from the Plio-Pleistocene of South Florida. 
The shells from Cuba seem to be recent. 

Teinostoma altum is similar to T. 
cocolitoris, but the latter is larger, the spire 
lower, has fewer whorls and lacks the 


groove separating the columella and the 
umbilical callus. 

From T. parvicallum it may be differ- 
entiated by the characters of the umbilical 
callus. Also it is a little similar to T. 
reclusum in its general form and in the 
groove in the umbilical callus, but the 
latter has a lower spire and a different 
peripheral profile. 

Another close species is Teinostoma 
subconicum (H.C. Lea), described from 
the Miocene of Smithfield, Virginia from 
only one shell. 

PrLsBRY (1953; pl. 56, fig. 5), gives a 
drawing of this species showing the dif- 
ferences with T. altum: the shell is more 
depressed, the periphery of the last whorl 
more arched, and the umbilicus is totally 
closed. 


A7 


Iberus, 29 (2), 2011 


In the web page www.jaxshell.org, 
as well as in “Marine Shells of North- 
east Florida” and “Select Images of 
Western Atlantic Gastropods” there is a 
SEM micrograph of a specimen of 
Teinostoma altum called Teinostoma sp. 


aff. altum, dredged in 30 m, 32 mi E St. 
Augustine, Florida, which is identical to 
our material from Cuba and that 
figured by PiLsBRY (1953). This figure 
also appears in (LEE, 2009: 67; species 
no. 320) 


“Teinostoma” solidum (Dall, 1889) (Figures 24A-G) 


Ethalia solida Dall, 1889. Bull. Mus. Comp. Zoology, 18: 362, pl. 28, figs. 3, 5. [Type locality: 
Station 19, Lat. 23%3'N, Lon. 83%10'W, off Bahia Honda, Cuba]. 


Type material: Syntype in MCZ (007553), from off Bahia Honda, Cuba. Range: 23.32N-83.10-W, in 
567 m. This shell is here designated the lectotype (Fig. 24). 


Description: The original description 
in DALL (1889) is as follows: “Shell 
small, solid, stout, ivory white, of three 
rounded whorls, the last much the 
largest. Sculpture of fine incremental 
lines, sometimes faintly wrinkled near 
the suture; upper surface rounded, sub- 
conic, the whorls not impressed at the 
suture, which is fairly distinct. Periph- 
ery rounded, base subconic, umbilicus 
reduced to a minute chink with a 
twisted callus above it; aperture circular, 
obligue, with a triangular callus at each 
end of the columella; the upper margin 
declining”. 

We add: The shell (Figs. 24A-E) is 
solid and compact, trochoid, spire 
formed by 3 Y rounded whorls sepa- 
rated by a distinct suture. The proto- 
conch has scarcely one whorl (which is 
not certain due to the difficulty in dis- 
cerning the separation from the teleo- 
conch). It measures about 450 um in 
diameter and the nucleus 160 um. The 
protoconch is short, bulbous, and is 
covered by small, sharp, branching 
tubercles arranged in a spiral pattern. 
Teleoconch formed by 2 Y whorls, 
totally smooth except for numerous 
growth lines. Umbilicus almost totally 
closed by a fine callous layer, which is 
the extension of the columella; within, it 
is possible to see two small folds which 
delimit several axial striae and spiral 
cordlets which cross and produce a 
reticular pattern. Aperture rounded, 
slightly angulate in its parietal part 
portion. Parietal callus, columella and 


48 


internal lip strong and wide. There is no 
sulcus between the columella and the 
callus. 

Dimensions: Lectotype 2.1 mm in 
maximum diameter and 1.6 mm in 
height. 

Habitat: This is a fairly deep-water 
species that apparently lives only on the 
continental slope (MOORE, 1964). From 
529-792 m. 

Distribution: From Georgia to West 
Florida, off Fernandina (JOHNSON, 1934). 
Recorded in Bahia Honda, Cuba (DALL, 
1889); Florida Keys (MOORE, 1964). 

Remarks: Provisionally, we keep this 
species in the genus “Teinostoma” in 
quotes, uncertainty about its correct 
placement for the reasons cited above. 
The name Teinostoma solidum is pre- 
occupied by a West African species of 
E-AS SM (ds 737 pl 75 te 20) 
which has all the typical characters of 
the genus. But the Caribbean species 
could be a Skeneid, in which case the 
name would be available because it was 
described in the genus Ethalia. At 
present, we prefer to keep it in this 
dubious status instead of creating a 
replacement name. 

It is necessary to point out the great 
similarity of “Teinostoma” solidum to 
some species of the skeneid genera 
Lisomphalia, Skenea and Trochaclis. 
Some deep water species of the genus 
Skenea living along the coast of Iceland 
and Scandinavia are characterized by 
the trochoid aspect of the shell, the short 
and bulbous protoconch, usually sculp- 


RUBIO £7 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 24A-G. * Teinostoma” solidum (Dall, 189), lectotype, 2.1 x 1.6 mm Bahía Honda, Cuba 
(MCZ 007553). A-C: optical photographs; D-E: SEM micrographs; E: protoconch; G: detail of 
the protoconch. 

Figuras 24A-G. “Teinostoma” solidum (Dall, 189), lectotipo, 2,1 x 1,6 mm Bahía Honda, Cuba 
(MCZ 007553). A-C: fotografías ópticas; D-E: microfotografías MEB; F: protoconcha; G: detalle de la 


protoconcha. 


tured, and having an umbilicus which, species of the genus Trochaclis are char- 
as in many species such as Skenea tro- acterized by a short and bulbous proto- 
choides (Friele, 1876), is very narrow conch with fine spiral cordlets. WARÉN 


and deep and has riblets within. The (1991: 179) reported that the genus 


49 


Iberus, 29 (2), 2011 


Trochaclis was originally classified in 
the Mesogastropoda and later trans- 
ferred to the Vetigastropoda by 
HICKMAN éz MCLEAN (1990) because of 
the morphology of the operculum, 
epipodium and ctenidium. 

Due to these similarities we consider 
the placement of this species in 
Tornidae, subfamily Teinostomatinae as 
dubious, but we mantain this clasifica- 
tion until anatomical, opercular and 
radular morphology allow correct sys- 


tematic placement. MOORE (1964) 
treated T. solidum and T. floridensis 
(Dall, 1889) in a similar fashion. 

“Teinostoma” solidum is more solid 
and elevated than any other species 
described from this region (DALL, 1889). 
According to MOORE (1964: 100), the 
small, bulbous and ornamented proto- 
conch, subglobose shape and narrow 
chink-like umbilicus and shelf within 
the aperture distinguish this species 
from other West Indian species. 


Teinostoma lunense spec. nov. (Figures 25A-D) 


Type material: Holotype (Figs. 25A-B) in MNCN (15.05/55066); a paratype in MHNS. 
Type locality: Rancho Luna Beach, Cienfuegos, Cuba, 20 m. 
Etymology: The specific name refers to the type locality. 


Description: Shell (Figs. 25A-C) 
rounded, a little higher than wide, spire 
moderately elevated, with four whorls, 
solid, smooth and somewhat shiny. Pro- 
toconch (Fig. 25D) of about 1 Y smooth 
whorls. Teleoconch with surface smooth 
except for fine growth lines, periphery 
rounded. Suture faintly indicated, 
visible by transparency, covered by a 
fine callous layer. Aperture rounded, 
peristome almost continuous, internal 
upper angle grooved. Columella and 
inner lip rounded, reflected towards the 
umbilicus, forming a characteristic 
callus, wide and fine, with a half moon 
crescent shaped, which partially covers 
the umbilicus. There is no groove of sep- 
aration between columella and callus. 


Dimensions: Holotype is 1.3 mm 
in maximum diameter and 0.9 mm 
of in height. 

Habitat: This species was col- 
lected in shell grit at 20 m depth. 

Distribution: Only know from 
Cienfuegos, Cuba, the type locality. 

Remarks: The figure in PILSBRY 
(1953, pl. 56, fig. 5) of the holotype 
of T. subconicum (H.C. Lea) is very 
similar to this species differing in 
the size of the callus, which, like a 
crescent moon does not completely 
cover the umbilicus. 

From T. altum it may be sepa- 
rated by the absence of the groove 
between the columella and the 
callus. 


Teinostoma cocolitoris Pilsbry £ McGinty, 1945 (Figures 26A-G) 


Teinostoma (Ellipetylus) cocolitoris Pilsbry £e McGinty, 1945a. The Nautilus, 59: 8-9, pl. 1, fig. 3. 
[Type locality: Off Lake Worth, Palm Beach County, Florida]. 


Type material: Holotype in ANSP (181122). Not examined. 

Material examined: Cuba: 1 s, Canarreos Archipelago, 5 m; 1 s, Cayo Diego Perez, 5 m (MHNS); 13 
s and 2 sp, Cienfuegos Bay, 20-30 m (MHNS); 3 s, Rancho Luna Beach, 10-20 m (MHNS); 16 s, 
Rancho Luna Beach, 10-30 m (MHNS); 10 s, Rancho Luna Beach, 45 m (MHNS); 16 s, Rancho Luna 
Beach, 20-54 m (MHNS); 3 c, Los Laberintos, Rancho Luna Beach, 35 m (MHNS); 19 s and 2 sp, Faro 
los Colorados, 56 m (MHNS); 2 s, Punta Tamarindo, 15 m (MHNS); 3 s, Havana, 30 m. USA: 31 s, 
off Louisiana, 56-65 m (CHL). Antigua and Barbuda: Antigua, 1 s, 67 m, SE Falmouth Harbour, 
dredged (CHL). Bahamas: 1 s, French Bay, San Salvador, 15 m, sand ledge (CHL); 4 s, Samphire 
Cay, NW Nassau, 15 m, base of reef (CHL). Jamaica: 3 s, Priory, St. Anr's, Parish, shallow water 
(CHL). ABC Islands: 1 s, off Klein Bonaire, Bonaire, 38 m, base of reef (CHL). 


50 


RUBIO E7 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 25A-D. Teinostoma lunense spec. nov. A-B: holotype, 1.3 mm, (MNCN); C: 1.5 mmm, 


paratype; both from Rancho Luna Beach, Cienfuegos, Cuba; D: protoconch. 
Figuras 25A-D. Teinostoma lunense spec. nov. A-B: holotipo, 1,3 mm, (MNCN); C: 1,5 mmm, para- 
tipo; ambas de Playa Rancho Luna, Cienfuegos, Cuba; D: protoconcha. 


Description: Shell (Figs. 26A-F) 
with 3 % rounded whorls, solid, 
smooth, rounded aperture, umbilicus 
partially occluded. Protoconch (Fig. 
26G) of barely one whorl and about 
230 um in diameter, with a smooth 
surface and a strong varix at its end. 
Teleoconch of about 2 % whorls. Dor- 
sally convex with very numerous, 
curved, prosocline growth lines; ven- 
trally, very slightly convex, with the 
same growth lines. Umbilicus narrow 
and deep, partially occluded by the 
callus that originates between the 
columella and the internal lip, callus 
small and semicircular. Aperture 
rounded, peristome continuous. 

Dimensions: Holotype 3.0 mm in 
diameter and 2.0 mm in height. Our 
shells reach 2.85 mm in maximum 
diameter. 

Operculum multispiral with a 
central nucleus. 

Habitat: Species with wide bathy- 
metric distribution, recorded 
between 18 and 122 m depth. In 


Cuba, it was found in sediments col- 
lected between 5 and 56 m, the living 
material examined was collected on 
coralline sandy bottom between 20 
and 56 m. In Cuba, off Rancho Luna 
Beach and Yaguanabo, living speci- 
mens were collected at 30-36 m on 
Halimeda sp. 

Distribution: Previously known 
from USA: North Carolina (PORTER, 
1974), Florida (PILSsBRY € MCGINTY, 
1945a), Louisiana (ODÉ, 1987); 
Colombia (GARCIA, 2002); Brazil: Rio 
de Janeiro (Rios, 1994); Abaco, 
Bahamian islands (REDFERN, 2001). 
Cuba: Cienfuegos Bay. 

Remarks: This is a characteristic 
species, with its surface totally 
smooth, the aperture rounded, and 
the umbilicus partially closed by the 
callus; these characters differentiate it 
from other congeneric species. Its 
callus is similar to that of  T. 
incertum, but the lack of ornamenta- 
tion in the protoconch and teleoconch 
differentiate them. 


51 


Iberus, 29 (2), 2011 


Figures 26A-G. Teinostoma cocolitoris Pilsbry 82 McGinty, 1945. A-F: shells, between 2.0 and 2.6 
mm, Cienfuegos Bay, Cuba; G: protoconch. 

Figuras 26A-G. Teinostoma cocolitoris Pilsbry € McGinty 1945. A-F: conchas, entre 2,0 y 2,6 mm, 
Bahía de Cienfuegos, Cuba; G: protoconcha. 


Teinostoma helicinum spec. nov. (Figures 27A-F) 


Type material: Holotype (Fig. 27A) in MNCN (15.05/55064) and 2 paratypes (Fig. 27B y 27D) 
(15.05/55065), from type locality. Other paratypes: AMNH (1 s), FLMNH (448613, 1 s) MNHN 
(24403, 1 s), IES (2 s), MHNS (100540, 1 s), USNM (1155032, 1 s), CFR (3 s), GHL (1 s) and CFG (6 s). 


OZ 


RUBIO ET 412.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Other material examined: Cuba: 3 s and 1 sp, Faro de los Colorados, 56 m (MHNS). 
Type locality: Rancho Luna Beach, Cienfuegos, Cuba, 40-45 m. 
Etymology: The specific name refers to the shape of the peristome, which resembles that of the land 


snail, subfamily Helicinae. 


Description: Shell (Figs. 27A-D) 
strong, solid, somewhat depressed, with 
about 4 whorls, whitish, smooth and 
shiny. Protoconch (Figs. 27E-F) of about 
1 Y smooth whorls and about 280 um in 
diameter. It is separated from the teleo- 


conch by a strong varix. Teleoconch of 
about 2 Y whorls, totally smooth. 

Convex dorsally with very numer- 
ous, curved, prosocline growth lines; 
ventrally, very slightly convex, also with 
the same growth lines. Periphery 


Figures 27A-E. Teinostoma helicinum spec. nov.; A: holotype, 2.0 mm (MNCN); B: paratype, 1.8 
mm, (MNCN); C: paratype, 1.9 mm (MNHN); D: paratype, 1.5 mm (MNCN), Rancho Luna 
Beach, Cienfuegos, Cuba; E-F: protoconch. 
Figuras 27A-E Teinostoma helicinum spec. nov.; A: holotipo, 2,0 mm (MNCN); B: paratipo, 1,8 mm, 
(MNCN); C: paratipo, 1,9 mm (MNAN); D: paratipo, 1,5 mm (MNCN), Playa Rancho Luna, Cien- 
fuegos, Cuba; E-F: protoconcha. 


39 


Iberus, 29 (2), 2011 


rounded. A well formed suture can be 
seen by transillumination, being covered 
by a fine callous coat. Umbilicus entirely 
covered by a small callus formed by a 
thickening of the columella. Aperture 
nearly ovoid without any groove in the 
upper inner angle. Peristome sharp, 
externally reflected. 

Dimensions: Holotype 2.0 mm in 
maximum diameter and 0.95 mm of in 
height. 

Habitat: Collected on sandy bottoms 
between 20 and 56 m depth. 

Distribution: Only known from Cien- 
fuegos, Cuba. 

Remarks: Temostoma helicinum spec. 
nov. may be distinguished from its con- 
geners by the fine and narrow callous 
layer which covers the suture, by the 


small umbilical callus, and particularly 
by the outwardly deflected peristome as 
seen in some species of terrestrial shells 
of the subfamily Helicinae. 

It may be distinguished from T. 
ciskae, T. goniogyrus and T. lenticulare 
because these have the teleoconch surface 
totally covered by punctiform incisions; 
from T. clavium and T. nesaeum because 
these have the teleoconch covered totally 
or partially by spiral cords, and from T. 
lerema, T. umbilicatum and T. biscay- 
nense because these have the protoconch 
totally covered by a fine callous layer. 
From T. megacallus it can be distin- 
guished by its externally reflected peris- 
tome and the smaller umbilical callus, 
and from T. megastoma by the sunken 
protoconch of the latter. 


Teinostoma megastoma (C.B. Adams, 1850) (Figures 28A-C) 


Vitrinella megastoma C.B. Adams, 1850. Monog. Vitrin.: 7. [Type locality: Port Royal, Jamaica]. 
Teinostoma biscaynense auct. non Pilsbry £e McGinty, 1945a. 


Type material: Lectotype (Figs. 28A-C) in MCZ (156269) after CLENCH éz TURNER (1950: 306, plate 
35, fig. 2). At present itis destroyed. There are 11 paralectotypes (labeled as paratypes), from Jamaica 
in MCZ (186187). A neotype is here designated from one of these shells (Figs. 28A-C) of this lot. 
Material examined: Bahamas: 1 s, Olympus Reef, 12 mi. NNW West End, Grand Bahama, 36 m, 
coralline algae fragments (CHL). USA: Florida: 1 s, Spoil, Apac Pit, Sarasota Co., fossil (CHL); 1 s, 
32 mi. E St. Augustine, St. Johns Co. 27 m ft (CHL). Louisiana: 1 s, 71-74 m (CHL). 


Description: We repeat the original 
description in C.B. ADAMS (1850): 
“Much depressed, transversely ovate: 
white, translucent: smooth and shining: 
apex very obtuse: spire convex, but little 
elevated: whorls a little more than three, 
moderately convex, with a distinct 
suture; last whorl very large, rapid ly 
increasing in the last part, well rounded: 
aperture scarcely modified by the last 
whorl: umbilical region very widely and 
deeply indented. Mean divergence 
about 130%; length of spire 0.01 inch; 
total length .033 inch; greatest bread th 
0.06 inch, least bread th 0.045 inch”. 

PiLsBRY (1946) mentioned the origi- 
nal description and figured the species 
for the first time. He complemented the 
description: “The shell is transparent, 
glassy, quite thin and globose for 
Teinostoma, with very large aperture, 


54 


and very small, sloping umbilical callus, 
which is flat or slightly concave, the 
base rising around it. The wholly super- 
icralisutuse iso nderedasby aa alse 
sutural line by transparence; sometimes 
this gives the illusion of a deeply 
impressed suture. Diameter: 2 mm, 
height 1.2 mm”. And he follows: “This 
was rather abundant. By the very small 
umbilical  callus it  agrees with 
Pseudorotella, as Miss Bush has noted 
on the label. The “paratype” figured is 
the largest in the lot. Adams's measure- 
ments were apparently from a smaller 
one. Except in size, the dozen specimens 
are all very much alike”. 

Shell (Figs. 28A-C) solid, smooth, 
whitish and shiny; spire of a little more 
than 3 rapidly-expanding whorls. Teleo- 
conch with about 2 whorls, smooth 
except for fine growth lines. Periphery 


RUBIO ET 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


(MCZ 156269). 


Figures 28A-C. Teinostoma megastoma (C.B. Adams, 1850). A-C: neotype, 1.5 mm, Jamaica 


Figuras 28A-C. Teinostoma megastoma (C.B. Adams, 1850). A-C: neotipo, 1,5 mm, Jamaica (MCZ 


156269). 


rounded. Suture seen by transillumina- 
tion to be covered by a fine coat of shell 
material which also covers the proto- 
conch and the first teleoconch whorl. 
Each whorl covers much of the previous 
one. Callus completely covers the 
umbilicus. Aperture oval, not modified. 
Base concave, umbilicus totally covered 
by a fine callus. 

Maximum reported size: 2.5 mm 

Habitat: Species with wide bathy- 
metric distribution, recorded between 0 
and 123 m depth. Collected alive 
between 0 and 42 m. It lives in man- 
groves, sand, muddy, and rocky bottom. 
Shallow soft bottom (Díaz MERLANO é 


PUYANA HEGEDUS, 1994). In Cuba, most 
of the samples were collected between 4 
and 10 m. 

Distribution: Recorded from Port 
Royal, Jamaica (C.B. ADams, 1850; 
CLENCH éx TURNER, 1950); from Colón 
and Bocas Island, Panama (OLssoN éz 
MCGINTY, 1958); from St. Croix, Virgin 
Islands (NOWELL-UsSTICKE, 1959); from 
USA: Louisiana, Texas; Mexico: 
Campeche State, Yucatan State, Quin- 
tana Roo (MOORE, 1964; ODÉ, 1987); 
from NE Florida (Lgr, 2009); from North 
Carolina and south of the Caribbean 
Sea, and Portete, Costa Rica (HOUBRICK, 
1968); from Campeche to Ciudad del 


Ss 


Iberus, 29 (2), 2011 


Carmen and Zacatal, from Ninum Point 
to Campeche, from El Cuyo to Ninum 
Point, from Yalkupul Point to Cerritos 
Islands and from Isla Mujeres to Holbox 
Island, Mexico (VOKES éz VOKES, 1984); 
from North Carolina to the western 
Caribbean (ABBOT, 1974); from Cuba 
(ESPINOSA ET AL., 1985); from Portete 
and Moín, Costa Rica (ROBINSON éz 
MONTOYA 1987); from Curacao, Aruba 
and Bonaire (DE JONG é COOMANS, 
1988); from North Carolina to Panama 
and Colombia (Díaz MERLANO € 
PUYANA HEGEDUS, 1994); from Brazil: 
Rio de Janeiro (RIOS, 1994). Martinique, 
in our material. 

Remarks: The tube with the label of 
the lectotype in MCZ (156269) had only 
a few fragments; so, from the lot of the 
paralectotypes (MCZ 186187) we have 
selected one shell which is here desig- 
nated the neotype. 

PiLsBRY (1946) accepted the descrip- 
tions and figured the species of Vit- 
rinella described from Jamaica by C.B. 
Adams, but he placed this species in the 
genus Teinostoma, subgenus 
Pseudorotella, agreeing with the 
opinion of Katherine J. Bush noted in 


the label of lectotype, MCZ 156269. The 
globose aspect, the rounded aperture, 
the umbilical callus reduced to a fine 
slightly convex callous coat, and the 
protoconch placed below the following 
whorl separate it from its congeners. 
LEE (2009, fig. 325) figured a shell 
dredged at 27 m, 32 miles East of St. 
Augustine, Florida, but in our opinion 
this is not T. megastoma but a member 
of the Cornirostridae, genus Tomura. In 
the web page www.jaxshells.org, as well 
as in “Marine Shells of Northeast 
Elorida* as: in “¿Selected Images ot 
Western Atlantic Gastropods” there is a 
SEM micrograph under the name of 
Teinostoma megastoma which, in our 
opinion, is T. umbilicatum (=T. cryp- 
tospira). The two species are similar 
because they have the spire partially 
covered by a fine callous coating. The 
differences between them are that in T. 
umbilicatum the callous cap completely 
covers the shell, including the proto- 
conch; the spire is flat, the umbilical 
callus is stronger and the aperture is 
almost circular. In T. megastoma the 
spire is more elevated, the umbilical 
callus is finer, and the aperture ovoid. 


Teinostoma cienfuegosense spec. nov. (Figures 29A-D) 


Type material: Holotype (Fig. 22A) in MNCN (15.05 /55061) and 2 paratypes (Figs. 29B-C)in MNCN 
(15.05 /55062). Other paratypes: MHNS (100547, 10 s), AMNH (2 s), NHMUK (2 s), MNHN (24393, 


2 s), (IES, 2 s), (CER, 3 s) and (CFG, 6 s). 


Other material examined: Cuba: 11 s, Cienfuegos Bay, 10 m (MHNS); 4 j, Cienfuegos Bay, 20-30 m 
(MHNS). Martinique: 1 c, Pointe Borgnesse, 12 m, sandy-muddy bottom, close the reef (CJP). 
Type locality: Cienfuegos Bay, sta. 12a, 22%07'N 80%26'W, 4 m. 

Etymology: The specific name refers to the type locality, an area extensively sampled by the second 


author. 


Description: Shell (Figs. 29A-C) a 
little depressed, rounded, trochiform, 
solid, smooth, whitish and shiny; spire 
formed by 4 rapidly-increasing whorls. 
Protoconch (Fig. 29D) bulbous, of 
about 1 % whorls, and of two clearly 
differentiated stages, the first one 
smooth and the second with small dis- 
persed granules; it measures about 280 
um and it is placed below the next 
whorl. Teleoconch with about 2 Y 
whorls, totally smooth except for fine 


3Ó 


growth lines. Periphery rounded. 
Suture barely distinguishable, seen by 
transillumination to be covered by a 
fine coat of shell material. Each whorl 
covers much of the previous one. 
Callus covers the umbilicus completely. 

Dimensions: Holotype is 1.43 mm 
of maximum diameter. 

Maximum reported size: 2.5 mm 

Habitat: ln Cuba most of the 
samples were collected between 4 and 
10 m in coralline sand bottom. 


RUBIO £7 41.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 29A-D. Teinostoma cienfuegosense spec. nov. A: holotype, 1.43 mm (MNCN); B-C: paraty- 
pes, 1.2, 1.3 mm, Cienfuegos Bay, Cuba (MNCN); D: protoconch. 

Figuras 29A-D. Teinostoma cienfuegosense spec. nov. A: holotipo, 1,43 mm (MNCN); B-C: parati- 
pos, 1,2, 1,3 mm, Babía de Cienfuegos, Cuba (MNCN); D: protoconcha. 


Distribution: Only known from 
Cienfuegos Bay, Cuba. 

Remarks: Early in the course of 
this work this species was con- 
fused with T. megastoma on the 
basis of the figure of the holotype 
provided. by. PILSBRY (1946). 
Teinostoma cienfuegosense spec. 
nov bears little similarity to” 1. 
megastoma, from which it can be 


distinguished by its uncalloused 
protoconch and the rounded, 
almost circular aperture. 

The globose aspect, the 
rounded aperture, the umbilical 
callus reduced to a fine callous 
slightly convex coat, and the pro- 
toconch placed below the follow- 
ing whorl separate it from its con- 
geners. 


Teinostoma parvicallum Pilsbry ££ McGinty, 1945 (Figures 30A-H) 


Teinostoma (Idioraphe) parvicallum Pilsbry £ McGinty, 1945a. The Nautilus, 59: 4-5, pl. 2, fig. 2. 
[Type locality: Missouri Key, Florida]. 


Type material: Holotype in ANSP (181105). Not examined. 

Material examined: Cuba: 34 s, Cienfuegos Bay, 22%07'N 80%27'W, 9 m (MHNS); 42 s, Cienfuegos 
Bay, 10 m (MHNS); 9 s, Rancho Luna Beach, 20 m (MHNS); 5 s, Rancho Luna Beach, 15-54 m 
(MHNS). Trinidad and Tobago: Tobago: 1 s, Horse Shoe Reef, 15 m, coralline sandy grit near the 
reef (CJP). 


37 


Iberus, 29 (2), 2011 


Figures 30A-H. Teinostoma parvicallum Pilsbry 8% McGinty, 1945. A-E; shells, between 1.0 and 
1.5 mm, Cienfuegos Bay Cuba; F-G: protoconch; H: radula. 
Figuras 30A-H. Teinostoma parvicallum Pilsbry + McGinty, 1945. A-E; conchas, entre 1,0 y 1,5 
mm, Bahía de Cienfuegos, Cuba; F-G: protoconcha; H: rádula. 


Description: In PiLSBRY € MCGINTY 
(1945a: 4-5) and MOORE (1964: 85-87). 

Shell (Figs. 30A-E) trochiform, with 
spire relatively high, umbilical callus 
medium in size, totally covering the 


58 


umbilicus in adult individuals. Proto- 
conch (Figs. 30F-G) of about 1 % smooth 
whorls, and about 270 um in diameter. 
Teleoconch of about 2 Y whorls, totally 
smooth except for small growth lines. 


RUBIO ET 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Periphery and base rounded. Between 
the columella and the umbilical callus 
there is a fine groove. Umbilicus gen- 
erally closed totally by the callus, 
except in young individuals, in which 
a small fissure may persist. Aperture 
slightly oblique, with a sharp border 
on the external lip and a continuous 
peristome. 

Dimensions: Holotype 2.0 mm in 
diameter by 1.5 mm in height. Our 
shells reach 1.5 mm in diameter and 
1.09 mm in height, and being smaller 
keep the same ratio (D/H: 0.75). 

Animal of holotype figured by 
PILSBRY €£ MCGINTY (1945a). 

Radula (Fig. 30H) taenioglossate, 
with formula 2+1+R+1+2. Central 
tooth wide basally, the ventral margin 
well developed, without denticles. 
Cutting surface formed by a central 
large, sharp cusp and 5 denticles of 
medium size on each side. Lateral 
teeth similar to the central, their bases 
are quadrangular and also without 
denticles; free margin with a central 
cusp and 4-5 smaller denticles on each 
side, more elongated than central 
tooth. Marginal teeth long, narrow, 
and hook-shaped; the medial aspect 
with 24-26 weak denticles on the 
upper outer margin; the outer mar- 
ginal teeth are inclined outwards in 


their distal third and possess 15-16 
denticles on the upper end of their 
medial aspects. 

Habitat: This species lives under 
stones between 10 and 50 m in depth. 
Some authors recorded it in deeper 
water (up to 90 m) based only on 
empty shells. Considered a continen- 
tal species by MOORE (1964) it is 
widely distributed among the islands 
of the Caribbean. 

Distribution: It has been recorded 
from the USA: Missouri Key, Florida 
(PILSBRY é MCGINTY, 1945a); from 
Puerto Rico (WARMKE €  ABBOTT, 
1961); from Florida Keys, Texas to 
Mexico (MOORE, 1964); from Texas 
(ANDREWS, 1977) “from. Cuba 
(ESPINOSA: SER LALO 1985) from 
Venezuela, Sucre and Isla Margarita 
(Princz, 1986); from Florida to Texas 
(Lyons, 1989); from Colombia (DíAz 
MERLANO é PUYANA HEGEDUS, 1994); 
from Abaco, Bahamas (REDFERN, 
2001). From Tobago, in the present 
work. 

Remarks: T. parvicallum is very 
similar in general aspect with the 
shell of T. incertum, from which it can 
be distinguished by the obliteration of 
the umbilicus by callus even in juve- 
nile shells and in lack of ornamenta- 
tion on the teleoconch. 


Teinostoma megacallum spec. nov. (Figures 31A-E) 


Type material: Holotype (Fig. 31A)in MNCN (15.05 /55067), and 2 paratypes (Figs. 31B-C) in MNCN 
(15.05/55068), from type locality. Other paratypes from Cienfuegos Bay, sta. 12a, 22007'N 80"26'W, 
4 m: MHNS (100548, 1 s, Fig. 31D), MNHN (24394, 1 s), FLMNH (448614, 1 s), AMNH (1 s), CFG 


(Lis) CER(2S): 


Other material examined: Cuba: 2 s, Cienfuegos Bay, sta. 12, 22%07'N 8027'W, 9 my 10 s, Cienfue- 
gos Bay, sta. 12a, 22%07'N 8026'W, 4 m; 1 s, Cienfuegos Bay, 20-30 m; 2 s, Rancho Luna Beach, 15- 


30 m. 


Type locality: Cuba, Bahía de Sagua, northeast Cuba, 3-6 m. 
Etymology: The specific name refers to the large callus, which characterizes the species. 


Description: Shell (Figs. 31A-D) tro- 
choid, pyriform, very solid, almost as 
wide as high, smooth, umbilical callus 
very large. Protoconch (Fig. 31E) of 
about 1 Y smooth whorls, and about 
230 um in diameter. Teleoconch of 
about 2 whorls, no ornamentation, with 


visible suture and rapid expansion. 
Dorsally convex with very numerous, 
curved, prosocline growth lines; ven- 
trally, very slightly convex, also with 
the same growth lines, which are more 
evident on the dorsum and periphery. 
Umbilicus totally hidden by a strong 


9% 


Iberus, 29 (2), 2011 


Figures 31A-E. Teinostoma megacallum spec. nov. A: holotype, 1.65 mm, Sagua, Cuba (MNCN); 
B: paratype, 1.5 mm, Sagua, Cuba, (MNCN); C: paratype, 1.6 mm, Cienfuegos, Cuba (MNCN); 
D: paratype 1.8 mm (MANS); E: protoconch. 

Figuras 3LA-E. Teinostoma megacallum spec. nov. A: holotipo, 1,65 mm, Sagua, Cuba (MNCN); B: 
paratipo, 1,5 mm, Sagua, Cuba (MNCN); C: paratipo, 1,6 mm; Bahía de Cienfuegos, Cuba 
(MNCN); D: paratipo 1,8 mm (MHNS); E: protoconcha. 


callus formed by the thickening of 
the columella and the internal lip. 
Aperture nearly quadrangular with 
an expansion of the peripheral 
labrum, columella wide and almost 
straight, outer lip sharp, the upper 
part advanced. 

Dimensions: Holotype 1.65 mm in 
maximum diameter and 1.12 mm of 


height. 


60 


Habitat: In Cuba this species has 
been collected in sandy grit between 
3 and 30 m deep. 

Distribution: Only know from 
Cienfuegos, Cuba. 

Remarks: Teinostoma megacallum 
may be distinguished from its con- 
geners by the robust shell and princi- 
pally by the thick columella and the 
great size of the umbilical callus. 


RUBIO ET 4Lz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Teinostoma carinicallus (Pilsbry € McGinty, 1946) (Figures 32A-D) 


Teinostoma lituspalmarum auct. non Pilsbry £: McGinty, 1945. 

Teimostoma (Annulicallus) carinicallus Pilsbry € McGinty, 1946a. The Nautilus, 60: 17-18, pl. 2, 
figs. 6-6b. [Type locality: Missouri Key, Florida]. 

Pseudorotella carinicallus Pilsbry € McGinty, 1946. 


Pseudorotella carinicallum (sic). 


Type material: Holotype in ANSP (181979). Not examined. 
Other material examined: Florida, USA: 1 s east side, Peanut Island, under rocks (CMK); 1 s, Anclote 
Key, Pasco Co. (CHL); 1 s, Pelican Shoals, Key West, Monroe Co., 5-7 m, (CHL). 


Description: The best description is 
in MOORE (1964: 101-102) “Shell 
depressed, shoulder concave, bearing a 
low spiral rib at its outer edge; umbili- 
cus surrounded by a strong spiral 
carina. Spire flattened, formed by four 
whorls, two in the protoconch and two 
in the teleoconch. Periphery rounded; 
umbilical area is bordered with a 
strong spiral carina. Aperture oblique, 
parietal callus rather thin. There is a 
small groove at the upper inner angle”. 

We add: A short channel is formed 
at the junction of the umbilical keel 
with the columella. The protoconch 
(Fig. 32D) is barely detectable due to a 
fine callous layer which covers the 
shell dorsally. In the studied material 
the dorsal cord is prominent and pro- 
duces a distinct angulation on the 
shell. 

Maximum reported size: 2.7 mm. 
Our shell (Figs. 32A-C) measures 2.15 
Aun. in. diameter and 1.25 mm in 


height. 


Habitat: Depth between 0 to 46 
m. The type specimen was taken 
alive in shallow water. However the 
species appears to be rare in 
shallow inshore waters (MOORE, 
1964). 

Distribution: USA: Florida: East 
Florida, West Florida, Florida Keys 
(PILSBRY €: MCGINTY, 1946b); Texas 
(ODÉ, 1987b); Panama (OLssoN éz 
MCGINTY, 1958). 

Remarks: This species was col- 
lected from the Plio-Pleistocene of 
St. Petersburg. PILSBRY (1953) stated: 
 amone the  Pliocene. specimens 
there are many in which the spiral 
angle of the upper surface is wholly 
absent”. Our shell presents the 
typical characters of the species. 

T. carinicallus is very similar to 
> liituspalmarum. —_Pilsbry  < 
McGinty. The main difference 
between the two species is that T. 
carinicallus is smooth and 7. litus- 
palmarum has weak spiral striae. 


Teinostoma lituspalmarum Pilsbry £: McGinty, 1945 


Teinostoma (Annulicallus) lituspalmarum Pilsbry 8: McGinty, 1945a. The Nautilus 59: 7-8, pl. 2, 


fig. 3. [Type locality: Off Palm Beach, Florida]. 


Type material: Holotype deposited in ANSP (181103). Not examined. 


Desenption: "See PIESBRY « 
MCGINTY (1945a). 

Maximum reported size: 1.7 mm. 

Habitat: Rocky reef, in 90 m 
deep (50 fms). 

Distribution: USA: Florida: East 
Florida (PiLsBRY é  MCGINTY, 


1945a). 


Remarks: See also T. lituspal- 
marum auct. = TT. carinicallus. 

Since its original description this 
species has not been recorded. 
MOORE (1964: 102-103) mentions: 
“Teinostoma carinicallus is very 
similar to T. lituspalmarum Pilsbry 
and McGinty. The main difference 


61 


Iberus, 29 (2), 2011 


Figures 32A-D. Teinostoma carinicallus Pilsbry 82 McGinty, 1946. A-C: shell, 2,15 mm, Peanut 


Island, Florida (CMK); D: protoconch. 
Figuras 32A-D. Teinostoma carinicallus Pilsbry Y McGinty, 1946. A-C: concha, 2,15 mm, lsla 


Peanut, Florida (CMK); D: protoconcha. 


betweentthe tos peciesms nat two are mierely ormsiojbessjamie 
carinicallus is smooth and T. litus- species, but it is best to keep them 
palmarum has weak spiral striae. separate until the problem can be 
More material may show that the solvedis 


62 


RUBIO ET 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Subfamily TORNINAE Sacco, 1896 


Genus Tornus Turton € Kingston, 1830 


Tornus Turton € Kingston, 1830. Testacea Britanica, pp. 438, pl. 7, fig. 9. [Type species: Helix 
subcarinata Montagu, 1803, by monotypy. Recent. Europe]. 
Adeorbis S. Wood, 1842. Ann. Mag. Nat. History, 9:530. [Type species: Adeorbis subcarinatus 


(Montagu, 1803)). 


Diagnosis: Shell of small size (2-3 
mm), solid, usually depressed, spire 
with 3-4 whorls. Protoconch smooth, 
between 1 and 2 whorls (most fre- 
quently 1 %), not elevated. Teleoconch 
with strong spiral cords crossed by 
strong axial ribs. External lip crenulated. 
Aperture  subtrigonal.  Operculum 
ovoid, paucispiral and chitinous. 

Habitat: Acording to FRETTER € 
GRAHAM (1978) and GorFas, PINTO 
AFONSO € BRANDAÓ (1985), the true 
Tornus live deeply buried in sand under 
stones, but they need clean sand 
through which the water circulates and 
allows good oxygenation. In The Straits 
of Gibraltar, Spain it lives with other 
species in areas with strong current and 


heavy waves partially buried in a sandy 
bottom among boulders and stones. 

Remarks: Numerous species of 
Tornus have been described from Euro- 
pean coasts as well as West Africa. 
ROLÁN € RUBIO (2002) revised the 
family Tornidae in the East Atlantic, 
studying 39 species of which 13 are in 
the genus Tornus. But on the other side 
of the Atlantic, no species had been 
described in the genus Tornus from 
either coast of the New World. 
However, two species previously placed 
in the genus Cyclostremiscus are, in our 
opinion, members of this genus. The 
morphology of their shells, very similar 
to some of the West African coast, is of 
interest. 


Tornus caraboboensis (Weisbord, 1962) (Figures 33A-C) 


Cyclostremiscus caraboboensis Weisbord, 1962. Bulletins of American Paleontology, 42(193): 
140-141, pl. 13, figs. 7-9. [Type locality: La Salina, west of Puerto Cabello, state of Carabobo, 


Venezuela]. Fossil record. 


Type material: Type material deposited in PRI (26094). Not examined. 
Other material examined: Guatemala: 2 s, Livingston, 2 m (MHNS). 


Description: (See WEISBORD, 1962 
and ALTENA, 1975). Shell (Figs. 33A- 
B) with a reticulated surface, pro- 
duced by the spiral cords crossing 
the axial ribs. Protoconch (Fig. 33C) 
of about 1 % smooth whorls, about 
260 um in diameter. Teleoconch 
sculpture formed by strong keel-like 
spiral cords placed one on the 
dorsum, two at the periphery, 
another one on the base, and finally 
the last one delimiting the umbilicus. 
Fine spiral threads are distributed 
between the keels. Spiral cords and 
threads are crossed by  narrow 
strongly prosocline axial ribs. Aper- 


ture rounded, outer lip with five 
prominences caused by the ends of 
the spiral keels. Columella strong, 
slightly curved. 

Dimensions: Holotype 2.3 mm in 
maximum diameter, but largest shells 
can reach 3.0 mm. Our material mea- 
sures 1.23 mm in diameter and 0.82 
mm in height. 

Habitat: In shallow water. The 
samples studied were collected in 
muddy sand bottom at 2 m in depth. 

Distribution: Species considered 
of continental distribution. Recorded 
from Venezuela (WEISBORD, 1962); 
Colombia (CoseL, 1986;  DíAz 


Ó3 


Iberus, 29 (2), 2011 


Figures 33A-B. Tornus caraboboensis (Weisbord, 1962). A-B: shells, 1.1, 1.0 mm, Livingston, 
Guatemala (MHNS); C: protoconch. 
Figuras 33A-B. 'Tornus caraboboensis (Weisbord, 1962). A-B: conchas, 1,1, 1,0 mm, Livingston, 
Guatemala (MANS); C: protoconcha. 


MERLANO € PUYANA HEGEDUS, 1994); 
Surinam (ALTENA, 1975); Curacao: 
Schottegat and Spaanse Waters (DE 
JONG é COOMANS, 1988); Brazil: 
Espirito Santo (RIOS, 1994); 
Guatemala (the present work). The 
only insular record is that of DE JONG 
$ COOMANS (1988) for Curacao, very 
close to the continent, not entirely 
inconsistent with the “continental” 
distribution paradigm. 

Remarks: This species was 
described as fossil in the Pliocene of 
Carabobo, Venezuela. The samples 
recorded from Surinam are also fossil 
but derived from the Holocene 
deposites. The material here studied 


Ss recent rom yin stone; 
Guatemala. By their development 
stage they seem to be juvenile shells. 
The morphology of the sculpture 
(strong cords and fine spiral cordlets 
crossing with axial ribs forming a 
characteristic reticule) recall some 
West African species of Tornidae, T. 
subcarinatus (Montagu, 1803); T. 
africanus Adam é€ Knudsen, 1969; T. 
aemilii Rolán €£ Rubio, 2002; T. erici 
Rolán € Rubio, 2002; and particularly 
T. umbilicorda Rolán € Rubio, 2002, 
a species with which it has consider- 
able similarity. For this reason we 
have placed Cyclostremiscus 
caraboboensis in the genus Tornus. 


Tornus schrammii (P. Fischer, 1857) (Figures 34A-E) 


Cyclostrema schrammii P. Fischer, 1857. Journal de Conchyliologie, 6: 287-288, pl. 10, fig. 11 


[Type locality: Guadeloupe]. 
Cyclostremiscus schrammii (P. Fischer, 1857) 


Type material: Type material deposited in MNHN. Lost according to HOUBRICK (1967). 
Other material examined: Guatemala: 1 s, Livingstone, 2 m (MHNS). Trinidad and Tobago: Tobago, 


4 s, Buccoo (CHL). 


64 


RUBIO £7 4L.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 34A-E. Tornus schrammii (P. Fischer, 1857). A-B: shell, 1.5, 1.55 mm, Tobago (CHL); €: 
shell, 1.15 mm, Livingstone, Guatemala (MHNS); D: protoconch, Tobago; E: microsculpture. 
Figuras 34A-E. Tornus schrammii (2 Fischer, 1857). A-B: concha, 1.5, 1,55 mm, Tobago (CHL); C: 
concha, 1,15 mm, Livingstone, Guatemala (MHNS); D: protoconcha, Tobago; E: microescultura. 


Description: Shell: see P. FISCHER 
(1857). Shell (Fig. 34A-C) very solid, 
depressed and whitish in color. Pro- 
toconch (Fig. 34D) with 1 Y whorls, 
about 230 um in diameter, with 


strong tubercles irregularly placed, 
which are smaller and more numer- 
ous near the varix which appears at 
the transition to the teleoconch. 
Teleoconch sculptured by 4 strong 


65 


Iberus, 29 (2), 2011 


nodular spiral cords (one on the 
dorsum, two on the periphery, and 
one more on the base. There are 20- 
22 strong axial ribs. At the crossing 
points there are nodules. Subsutural 
area ornamented by fine spiral 
striae. The nodules of the subsutural 
cord are more elevated than that of 
the upper peripheral cord. Aperture 
rounded, prosocline. Umbilicus 
wide and deep which allows the 
previous whorls to be seen. 

Dimensions: The holotype is 2.0 
mm in diameter. Largest shell, a 
paratype in MNHN, is 3.2 mm in 
diameter. Our shells did not exceed 
Somo: 

Habitat: Recorded from the 
external reefs (VOKES é€ VOKES, 
1984) and shallow water (ROBINSON, 


991 
Distribution: le has been 
recorded. tom. Guadeloupe (E 


FISCHER, 1857); from Colón and 
Bocas Island, Panama (OLSSsON é 
MCGINTY, 1958); from Costa Rica 
(Houbrick, 1967); from Guadeloupe 
and Central America (HOUBRICK, 


1968); from Cancún to the Belize 
border, Arcas Keys, Alacran reef and 
Cayos del Norte and Lobos from the 
Banco Chinchorro, Mexico (VOKES éz 
VOKES, 1984; ROBINSON, 1991); from 
the Caribbean Sea (ABBOTT, 1974); 
and from Colombia (Díaz MERLANO 
éz PUYANA HEGEDUS, 1994). 

Remiarias LES nO ta emy 
common species, few shells are 
known even from the type locality. 
Since the description by P. FISCHER 
(1857), no new records had been 
published until HouBrick (1967), 
who figured one of seven shells 
found in Portete, Provincia Limón, 
Costa Rica. Houbrick considered 
this — specimen of. particular 
impostance since he reported that 
the holotype in the MNHN was lost. 

Tornus schrammii is very similar 
in shell morphology to some species 
from the west coast of Africa such as 
E EVO ODE, Te ENS AO Oe, 1 
ryalli, and T. rachelae. These all 
share a depressed shell sculptured 
with strong nodulous cords or 
tubular extensions. 


Subfamily VITRINELLINAE Bush, 1897 


Marine prosobranchs that have 
small or minute depressed shells, 
which are transparent while the 
animal is alive. The aperture is 
usually oblique, rounded, and 
without a terminal varix. There are 
usually one to three postlarval 
whorls which may be smooth or 
sculptured. The operculum is circu- 
lar and multispiral, and the radula 
taenioglossate. 

The animal has small black 
eyes, a pair of cephalic tentacles, 
and a pair of pallial tentacles. Both 
pairs are provided with mobile 
cilia for producing respiratory cur- 
rents and are tipped with long stiff 
inmobile: ¡cilía, he. male 4s 
equipped with a penis located 
behind the head and which extends 
baci anto the mantle ica vity 
(MOORE, 1964). 


66 


In the Recent fauna, there are 
ten genera grouped within the 
subfamily Vitrinellinae: Anticli- 
max Pilsbry € McGinty, 1946; 
Aorotrema Schwengel € McGinty, 
1942; Cyclostremiscus Pilsbry éz 
Olsson, 1945; Cochliolepis Stimp- 
son, 1858; Episcynia Mórch, 1875; 
Parviturboides Pilsbry € McGinty, 
1950; Pleuromalaxis Pilsbry € 
McGinty, 1945; Solariorbis 
Conrad. "lsOo:. WMitrimeltlaS A 
Adams, 1850 and - Vitrinorbis 
Pilsbry é€ Olsson, 1952, and all 
them are present in the geo- 
eraphie. area studied. Acnew 
genus, Neusas Warén € Bouchet, 
2001, has been placed provision- 
ally in Vitrinellinae, although the 
authors considered it in an uncer- 
tain systematic position, probably 
not in the Tornidae 


RUBIO E7 42Z.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Genus Anticlimax Pilsbry £ McGinty, 1946 


Climacia Dall, 1903: 1633 (Preoccupied, M'Lachlan, 1869 (Neuroptera)). 

Climacina Aguayo € Borro, 1946: 11. (Preoccupied by Gemellaro, 1878 (Mollusca)). 
Anticlimax Pilsbry € McGinty, 1946. The Nautilus, 60: 12. 

Type species: Teinostoma (Climacia) calliglyp tum Dall, 1903 (by monotypy). 


Remarks: PILSBRY € OLSSON (1950) 
revised the genus and gave the follow- 
ing description: “The shell is wider than 
high, with a dome-shaped or low-conic 
spire of few (3 or 4) whorls, carinate 
periphery and more or less convex base. 
The protoconch is smooth, of scarcely 
more than one convex whorl to 1 1. 
Sculpture of close, usually punctate, 
spiral striation and radial wavelike ribs 
on the base, sometimes on the upper 
surface also. The aperture is oblique, 
quadrangular or triangular, with a 
thickened peristome, the outer lip is 
angular or often extended at the termi- 
nation of the keel. Umbilicus bordered 
by a spirally emerging callous rib, termi- 
nating at the columella or in the genus 
Subclimax it fills the umbilicus”. 

PILSBRY € OLSSON (1950) divided 
Anticlimax into two subgenera: 

Subgenus Anticlimax s. str.: charac- 
terized by having the umbilicus open, 
bordered by a spiral cord which termi- 
nates in the columella in a small trian- 
gular callus. Type species: Anticlimax 
calliglyp ta (Dall). 

Subgenus Subclimax: characterized 
by having an umbilicus, totally or par- 
tially closed by a solid column which 
terminates in a callus fused to the col- 
umella. Type species: A. hispanioliensis 
Pilsbry € Olsson, 1950. 

Nothing is known about the animal 
of Anticlimax. Its generic assignment 
has been based only on the distinguish- 
ing characters of the shell, as the form, 
the radial folds on the base, the angular 
form of the external lip, and the spiral 
grooves in zigzag, among others. 

Most of the known species are fossil 
from the Miocene and Plio-Pleistocene. 
There are few recent species described, 
and they occur in North Carolina, 
Florida and Belize. 

AGUAYO éz BORRO (1946) described 
Canimarina and placed it provisionally 


as a subgenus of Cyclostremiscus, to 
accommodate the new species 
Cyclostremiscus (Canimarina) crassi- 
labris. 

This species, in their opinion, had 
characters which could ally it to the 
genera Cyclostremiscus, Miralabrum, 
Teinostoma and Climacia, but also it 
could be considered as a new genus due 
to its own different unique characters. 

Almost at the same time, PILSBRY éz 
MCGINTY (1946a) introduced the 
replacement name Anticlimax, for the 
monotypic genus containing Teinostoma 
(Climacia) calliglyp tum Dall, 1903, since 
the genera Climacia Dall, 1903 and Cli- 
macina Aguayo € Borro, 1946, based on 
the same type species, were unavailable 
homonyms (see above). 

FABER (2007) considers Canimarina a 
valid genus, comparing it only with 
Cyclostremiscus, and places it in Vit- 
rinellidae solely on the basis of its 
lacking “a clear apertural varix”. Also 
he considers Solariorbis decipiens 
Olsson € McGinty, 1958 a junior 
synonym of Cyclostremiscus (Canima- 
rina) crassilabris. Since then databases 
treat Canimarina as a valid genus and 
place it in Vitrinellinae. In our opinion, 
Canimarina must be considered a 
synonym of Anticlimax for the follow- 
ing reasons: 

-FABER (2007) considered it a valid 
genus without comparison to the genus 
Anticlimax. 

-the description of the subgenus is 
not sufficiently detailed to allow distine- 
tion from other species placed in Anti- 
climax. 

-Cyclostremiscus (Canimarina) cras- 
silabris shares all the generic characters 
of Anticlimax. 

-The stated date of publication for 
Anticlimax is July and that of Canima- 
rina is August of the same year, giving 
the former priority. 


67 


Iberus, 29 (2), 2011 


Anticlimax crassilabris (Aguayo « Borro, 1946) (Figures 35A-G) 


Cyclostremiscus (Canimarina) crassilabris Aguayo € Borro, 1946. Revista de la Sociedad Malaco- 
lógica “Carlos de la Torre,” 4(2): 46-47, lám. 3, figs. 1-3. [Type locality: Barranco E. del rio 
Canímar. Formación “Yumurí,” Upper Miocene. Matanzas, Cubal. 

Solariorbis decipiens Olsson €: McGinty, 1958. Bulletins of American Paleontology, 39: 28-29, pl. 
3, figs. 5, 5a, 5b. [Type locality: Bocas Island, Panama]. 


Type material: Type of Cyclostremiscus (Canimarina) crassilabris deposited in Museo Poey, Uni- 
versity of Havana. Type of Solariorbis decipiens in ANSP (211914). Not examined. 

Other material examined: Cuba: 1 s, Carenas Key, Cienfuegos Bay, 3 m, under stones (MHNS); 1 
s, in the channel to Cienfuegos Bay, 30 m (MHNS); 3 c, Rancho Luna Beach, 20 m (MHNS); 7 s Cien- 
fuegos (MHNS). Martinique: 1 c, Pointe Borgnesse, 12 m, from coralline sandy grit near the reef 


(EJ)! 


Description: The description of the 
species in AGUAYO € BORRO (1946) is as 
follows: “Concha diminuta, deprimida; 
espira cónica, algo elevada sobre el 
plano de la ultima vuelta. Provista de 2 
lh vueltas, la primera lisa, las siguientes 
esculpidas radial y espiralmente. Escul- 
tura axil o radial formada por costillitas 
salientes (unas 30 en la última vuelta) 
que desaparecen al 1/8 final de la 
última vuelta. Estas costilla persisten, 
aunque menos marcadas en el cuerpo 
de la última vuelta; pero desaparecen 
en la base al llegar a la quilla espiral 
inferior. Base lisa. Escultura espiral 
formada por dos quillas salientes que 
delimitan el contorno de la última 
vuelta, quedando articuladas por las 
costillas axiles. Además hay líneas finas 
espirales entre ambas quillas, y algunas 
evanescentes en la base. Cuerpo de la 
última vuelta de contorno cuadrangu- 
lar visto de perfil. Ombligo estrecho, 
parcialmente cubierto por un callo. Al 
final de la última vuelta la quilla supe- 
rior se proyecta tangencialmente hacia 
la abertura, formando un engrosa- 
miento muy marcado en ángulo recto. 
Abertura subtriangular, engrosada por 
una callosidad formada por la citada 
prolongación de la quilla superior. 
Borde parietal de la abertura con un 
callo peculiar en su ángulo superior”. 

The protoconch (Figs. 35F-G) is 
about 230 um in diameter, and it is 
totally smooth. The teleoconch is 
totally covered by fine spiral cordlets. 
On the dorsum, strong radial ribs may 
be observed. They are slightly proso- 


68 


cline and, on the last 1/3 of the body 
whorl, become more fine and numer- 
ous and are crossed by the spiral 
cordlets to form a reticule. Basally the 
shell is very convex and expresses 
strong radial undulating ribs. The 
external lip is angled in the peripheral 
area and is projected forward forming a 
characteristic extension. Umbilicus 
small, partially occluded by the thick- 
ening and extension of the columella. 

Dimensions: The  holotype of 
Cyclostremiscus (Canimarina) crassi- 
labris 1s 1.20 mm in maximum diameter 
and 0.78 mm in height. The holotype of 
Solariorbis decipiens measures 1.25 
mm in maximum diameter and 0.50 
mm in height. 

Animal and radula unknown. 

Distribution: Panama: Bocas Island 
(OLSSON é MCGINTY, 1958; FABER, 
2007); and Portobello (FABER, 2007); 
Dominica (FABER, 2007); Cuba: Matan- 
zas, fossil (AGUAYO € BORRO, 1946); 
Rancho Luna Beach, Cienfuegos Bay 
and Martinique, in the present work. 

Habitat: In Cuba this is a uncom- 
mon species, having been found in sed- 
iments collected between 20 and 50 m 
in depth. 

Remarks: Anticlimax crassilabris 
was described as fossil from the Upper 
Miocene, umuarí Formation. 
AGUAYO é€z BORRO (1946) mention that 
the base is lacking sculpture. There are 
axial undulating folds, which do not 
appear in the original figuration. These 
axial folds on the base are observed in 
the individual examined from Cuba. 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 35A-G. Anticlimax crassilabris (Aguayo 8 Borro, 1946). A-E; shells, between 1.2 and 2.0 
mm, Rancho Luna Beach, Cuba (MHNS); F-G: protoconch. 

Figuras 35A4-G. Anticlimax crassilabris (Aguayo e Borro, 1946). A-E; conchas, entre 1,2 y 2,0 mm, 
Playa Rancho Luna, Cuba (MHNS); F-G: protoconcha. 


OLSSON € MCGINTY (1958) described 
Solariorbis decipiens on the basis of 
only one specimen. Which such limited 
material it has not been possible to 
know the morphological variability of 
the shell. Our shells conform to the 
description of Solariorbis decipiens as 
well as that of C. crassilabris and have 
a consistency in the distinguising char- 
acters. Principal among these features 
are the sculpture of the dorsum formed 
by axial ribs with spiral threads in the 


intervals and the strongly convex base 
with strong undulating radial folds. 
External lip angular and apparently not 
channeled by the keel. All specimens 
are characteristic of the genus Anticli- 
max, and for this reason we have 
placed the species in this genus and not 
in Solariorbis. At the same time the 
similarity of Anticlimax crassilabris to 
Solariorbis  decipiens Olsson € 
McGinty impels us to consider the two 
synonymous. 


69 


Iberus, 29 (2), 2011 


Anticlimax glabra spec. nov. Rubio, Rolán € Pelorce (Figures 36A-D) 


Type material: Holotype (Fig. 36A) and a paratype (Figs. 36B-C) in MNHN (24200). 
Type locality: Grenadines, Martinique, Pointe Borgnese, 12 m. 
Etymology: The specific name refers to the smooth surface of the shell in opposition to other con- 


generic species. 


Description: Shell (Figs. 36A-C) 
very small, lenticular, spire consists of 
3 rounds of rapid growth. Protoconch 
(Figs. 36D) slightly projecting, about 
ISO diameter tana toralyy 
smooth. Teleoconch totally smooth 
except for the first quarter whorl in 
which there is ornamentation formed 
by spiral cords and axial ribs that 
intersect to form a characteristic retic- 
ulate sculpture. There is a thick subsu- 
tural cord that begins at the proto- 
conch and ends in the parietal area of 
the aperture, forming a thick callus. 
Basally the shell is more convex than 
dorsally and withouth strong radial 
undulating ribs. Aperture oval and 
with a small groove at the upper inner 
angle; the external lip is angled at the 
periphery; columella straight, inner 
lip reflected toward the umbilicus. A 
thin horny layer covers the entire 
umbilical area. 

Dimensions: The holotype mea- 
sures 1.00 mm and the paratype 1.06 
mm. 


Animal and radula unknown 

Distribution: Known only from the 
type locality. 

Habitat: Collected in muddy sand 
at the base of the reef, at a depth of 12 
m. 

Remarks: Anticlimax glabra spec. 
nov. is very similar to Anticlimax 
crassilabris to the point that we 
thought that it represented eroded 
specimens of that species. But the 
characters are constant in both known 
specimens, and they are very charac- 
teristic of this new species. The proto- 
conch shape, the angular, unchan- 
neled external lip, the subsutural 
cord, and its similarity to A. crassi- 
labris militated for its placement in 
Anticlimax although it also shares 
characteristics with Teinostoma. 

Anticlimax glabra spec. nov. 
differs from A. crassilabris by its lack 
of dorsal and basal ornamentation, by 
its rounded periphery and by having 
its umbilicus completely covered by a 
thin horny layer. 


Anticlimax decorata Rolán, Fernández-Garcés éz Rubio, 1997 (Figures 37A-D) 


Anticlimax decorata Rolán, Fernándes-Garcés € Rubio, 1997. Iberus, 15(1): 31-34. figs. 1-2 [Type 


locality: Rancho Luna Beach, Cienfuegos, Cubal. 


Type material: Holotype in MNCN (15.05 /27420) (Figs. 37A-B). Paratypes in the following collec- 


tions: AMNH, NHMUK, CFG, IES, MHNS. 


Other material examined: Cuba: 1 c, Rancho Luna Beach, 10-20 m. 


Description: See ROLÁN, FERNAN- 
DEZ-GARCÉS € RUBIO (1997). Shell 
(Figs. 37A-B). Protoconch (Fig. 37D); 
microsculpture (Fig. 37C). 

Dimensions: Holotype 1.4 mm in 
maximum diameter. 

Animal and radula unknown. 

Distribution: Only known from 
Cuba. 


7O 


Habitat: Unknown. 

Remarks: The genus Anticlimax, 
as has been noted above, is divided 
in two subgenera: Anticlimax and 
Subclimax. PILSBRY € OLSSON (1950) 
mentioned that there is no species 
demonstrating intergradation 
between these subgenera. Neverthe- 
less, A. decorata presents morpho- 


RUBIO ET 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 36A-D. Anticlimax glabra spec. nov. Rubio, Rolán 82 Pelorce. A: holotype, 1.0 mm, Mar- 
tinique (MNHN); B-C: paratype, 1.06 mm, Pointe Borgnese, Martinique (MNCN); D: proto- 
conch. 
Figuras 36A-D. Anticlimax glabra spec. nov. Rubio, Rolán e Pelorce. A: holotipo, 1,0 mm, Martini- 
que (MNHN); B-C: paratipo, 1,06 mm, Pointe Borgnese, Martinique (MNCN); D: protoconcha. 


logical characters which are inter- 
mediate. The open umbilicus and 
lack of columellar callus would 
place it in Anticlimax s. st., but the 
absence of axial sculpture on the 


base and of any kind of prolonga- 
tion of the labrum, suggest Subcli- 
max. We prefer to employ only the 
full generic assignment in this 
report. 


71 


Iberus, 29 (2), 2011 


Figures 37A-D. Anticlimax decorata Rolán, Fernandez-Garcés 8 Rubio, 1997. A-B: holotype, 1.4 
mm, Rancho Luna Beach, Cuba (MNCN); C: details of sculpture; D: protoconch. 

Figures 37E-H. Anticlimax proboscidea (Aguayo, 1949). E-G: shells, 1.4, 1.4, 1.6 mm, Rancho 
Luna Beach, Cuba; H: detail of the microsculpture. 

Figuras 37A-D. Anticlimax decorata Rolán, Fernandez-Garcés 9 Rubio, 1997. A-B: holotipo, 1,4 
mm, Playa Rancho Luna, Cuba (MNCN); C: detalle de la escultura; D: protoconcha. 

Figuras 37E-H. Anticlimax proboscidea (Aguayo, 1949). E-G: conchas, 1,4, 1,4, 1,6 mm, Playa 
Rancho Luna, Cuba; H: detalle de la microescultura. 


iz 


RUBIO ET 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Anticlimax proboscidea (Aguayo, 1949) (Figures 37E-H) 


Teinostoma (Anticlimax?) proboscidea Aguayo, 1949. Revista de la Sociedad Malacológica 
“Carlos de la Torre,” 6: 93-94, pl. 4, fig. 5. [Type locality: Camarioca, Matanzas, Cuba]. 


Type material: Holotype in the Museo Poey (n” 12902), Havana, Cuba. 
Other material examined: Antigua and Barbuda: Redonda island, 1 c, among rocks, 5-15 m. Cuba: 
1 s, Rancho Luna Beach, 10-20 my 1 s, Baracoa, 15 m; 2 s, Cienfuegos Bay, 15 m. ABC Islands: 1 s, 


50 mi. off Palm Beach, Aruba, 5 m (CHL). 


Description: This is the text in the ori- 
ginal description (AGUAYO, 1949): 
“Concha diminuta, sólida, blanca, en 
forma de domo, base casi plana, periferia 
obtusamente angular; ombligo marcado 
por una línea microscópica. Provista de 3 
MY vueltas de espira, de las cuales las 
nucleares (1 Y vueltas) son lisas, y las 
restantes, con numerosas líneas espirales 
microscópicamente punticuladas (unas 
15 en la última vuelta, desde la sutura 
hasta la periferia). Base con una escultura 
similar de 12 líneas espirales punticula- 
das. Región umbilical con un callo que 
no obtura por completo el umbilicus, 
dejando una pequeña cisura muy estre- 
cha. Última vuelta prolongada tangen- 
cialmente de una manera semejante a la 
del género Miralabrum Pilsbry y Olsson, 
1945. Abertura transversalmente ovoidal, 
pero casi subcircular, prolongada en un 
estrecho canal en el ángulo basal externo. 
Un grueso callo une a la abertura con la 
región parietal y columelar de la 
concha”. 

The protoconch has a little more than 
one whorl, with a diameter of about 310 
um and under high magnification it can 
be seen that the surface is slightly rough- 
ened. 

The teleoconch (Figs. 37E-G) is 
totally covered by evenly-spaced spiral 
cords, which present small ovoid small 
ovoid cells in their interspaces. Aperture 
triangular, the external lip is elongated 
basolaterally to produce an acute angu- 
lation at its lateral margin. Aside from 
this feature, the lip lacks an internal 


channel. Peristome thickened. The col- 
umellar callus is extended towards the 
base partially closing the umbilicus. 

Dimensions: Holotype is 1.7 mm in 
maximum diameter. Our shells measure 
1.55 mm in maximum diameter and 0.74 
mm in height. 

Animal and radula unknown. 

Distribution: MOORE (1965) consid- 
ered it endemic to Cuba. DE JONG é- 
COOMAN (1988) recorded it in Bonaire 
and Aruba (5 sps). REDFERN (2001) men- 
tions numerosous shells from Abaco, 
Bahamas, collected in sediment from 23 
m in depth. One shell from Redonda 
(Antigua and Barbuda) confirms its 
presence for the Lesser Antilles. 

Habitat: It is an uncommon species. 
We have it from sediments collected 
between 10 and 20 m in depth, but the 
holotype came from 40 m. 

Remarks: AGUAYO (1949: 94) stated 
that the general aspect of T. proboscidea 
is similar to that of Teinostoma pilsbryi 
McGinty, 1945, from which it is distin- 
guished by its small size, lack of the 
globose callus in the umbilicus, and the 
prolongation at the end of the last whorl. 
The domed shape and the aforemen- 
tioned projection have similarity to some 
species of the genus Anticlimax Pilsbry 
$ McGinty, 1946. 

Teinostoma pilsbryi McGinty, 1945 is 
smaller and has a different umbilical 
callus. 

Climacia  tholus  Pilsbry and 
McGinty, 1946 is smaller and has a dif- 
ferent umbilical callus. 


Anticlimax pilsbryi (McGinty, 1945) (Figures 38A-C) 


Teinostoma pilsbryi McGinty in Pilsbry and McGinty, 1945. The Nautilus, 58: 142-143, pl. 1, fig. 
5. [Type locality: One and one half miles off Cape Florida, 70 feet]. 


73 


Iberus, 29 (2), 2011 


Climacia tholus Pilsbry ££ McGinty, 1946a. The Nautilus, 59: 79-80, pl. 1-1b, 2-2a. [Type locality: 
About four miles off Carysfort Light, Florida]. 


Type material: Holotype of Teinostoma pilsbryi in ANSP (181080). Holotype of Climacia tholus in 


ANSP (181290). Not examined. 


Other material examined: Florida, USA: 1 s, Tarpon Springs, Hillsborough County, dredged in 12 


m at SW of Anclote Key (CMK). 


Description: The shell (Figs. 38A-B) 
has a dome shaped spire, peripheral 
keel at the base, a heavy callus which 
partly or completely fills the umbilicus, 
sculpture of many low close-set zigzag 
grooves, and low radiating waves on 
the base. Protoconch (Fig. 38C) of 1 Y 
smooth whorls, size about 240 um, 
without any varix at the transition to the 
teleoconch. Sculpture formed by fine 
spiral zigzag grooves. There are about 
20 on the base and 22-23 on the dorsal 
aspect of the body whorl. A thick ridge 
borders the umbilicus, and a heavy 
callus fills it partly or completely. Aper- 
ture of triangular shape, a little oblique. 
Parietal callus well developed, col- 
umella very strong and extended 
towards the umbilicus. There is no 
sulcus at the convergence with the 
peripheral keel. 

Holotype of Teinostoma pilsbryi 3.4 
mm in diameter and 2.0 mm in height. 
The photographed shell measures 1.9 
mm in diameter. 

Habitat: This is a species of wide 
bathymetric distribution having been 
collected between 0 and 152 m in depth. 
MOORE (1986) considered it a continen- 
tal species living in shallow water on 
mud bottom. 


Distribution: USA: Florida: East 
Florida (MCGINTY, 1945; PILSBRY éz 
MCGINTY, 1946a), Texas (ANDREWS, 
1977); Mexico: Campeche State, Yucatan 
State, Ouintana Roo (ODÉ, 1987b); 
Venezuela: unlocalized (PriNcz, 1982). 
South Florida, Texas and Mexico 
(MOORE, 1964; EMERSON € JACOBSON, 
1976). 

Remarks: MOORE (1964: 167) syn- 
onymyzed Teinostoma pilsbryi McGinty, 
1945 and Climacia tholus Pilsbry é 
McGinty, 1946a, saying that the species 
has a considerable variation in size and 
extension of the umbilical callus and that 
these characters overlapped between the 
two taxa to the extent that one could not 
distinguish the two. We agree with this 
conclusion. The only difference between 
A. pilsbry and A. tholus is the size of the 
umbilical callus, which in A. pilsbry is 
large, reflected outward and occluding 
the umbilicus, which is narrow and 
deep. A. pilsbryi shows considerable 
variation in the size and extent of the 
umbilical callus. The lack of radial costu- 
lation on the base, observed in the holo- 
type of T. pilsbry, was considered a con- 
sequence of the immaturity of the speci- 
men. Moore placed both species in Anti- 
climax. 


Anticlimax locklini Pilsbry £ Olsson, 1950 (Figures 39A-D) 


Anticlimax locklini Pilsbry € Olsson, 1950. Bull. Amer. Paleont., 33: 114, n* 135, pl. 2, figs. 8, 8a, 


9. [Plio-Pleistocene of Shell Creek, Florida]. 


Type material: Holotype in ANSP (n” 18393). Not examined. 
Other material examined: Nicaragua: 1 s, Witties Reef, 20-30 m. (MHNS). 


Description: This is the original 
description in PILSBRY 6: OLSSON (1950): 
“The shell has a low-conic truncate upper 
surface, a carinate periphery and rather 
flatly convex base. The spire is small, occu- 


74 


pying about one-third of the diameter of 
the shell. There are 3 Y whorls, the first 1 Y 
glossy, convex and projecting very slightly, 
the next whorl narrower, the penultimate 
whorl having several low radiating waves 


RUBIO £T AZz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


ough Co., Florida (CMK); C: protoconch. 


Figures 38A-C. Anticlimax pilsbryi (McGinty, 1945). A-B: shell, 1.9 mm, Tarpon Spring, Hillsbor- 


Figuras 38A-C. Anticlimax pilsbryi (McGinty 1945). A-B: concha, 1,9 mm, Tarpon Spring, Hills- 


borough Co., Florida (CMK); C: protoconcha. 


which do not extend upon the last part of 
the whorl. The last whorl has nearly 
straight lateral slopes, concave above the 
peripheral keel and closely marked with 
punctuate spiral striae. The base has about 
12 somewhat protractively rad iating wave- 
like ribs and weak, partly effaced striae. 
The aperture 1s triangular. The umbilicus is 
filled by a callus, forming a raised, convex 
pad in connection with the columellar 
callus, a small umbilical notch left next to 
the parietal callus”. 

Habitat: Our shell (Figs. 39A-C) was 
collected on a coralline bottom between 20 
and 30 m in depth. 

Distribution: Only know as a fossil 
species from Plio-Pleistocene of Shell 
Creek and St. Petersburg, Florida (PILSBRY 
éz OLsson, 1950); off Yucatan, Mexico 


(TREECE, 1980) and Witties Reef, 
Nicaragua. 

Remarks: PILSBRY €: OLSSON (1950) 
reported: “This species differs from 
related Miocene forms by reduction of 
the radial waves of the upper surface 
to a few grouped near the middle of 
the penultimate whorl. The umbilical 
callus pad is quite unlike the complete 
one of A. annae, and A. locklini has a 
more pronounced peripheral keel than 
that species, the whorl being concave 
aboventat. As intother species, the 
number of radiating ribs varies ind i- 
vidually”. 

TREECE (1980) considered A. lock- 
lini a recent species recorded off 
Yucatan. Our shell confirms that A. 


locklini is a recent species. 


TS) 


Iberus, 29 (2), 2011 


Figures 39A4-D. Anticlimax locklini Pilsbry 82 Olsson, 1950. A-C: shell, 1.3 mm, Witties Reef, 
Nicaragua (MHNS); D: microsculpture. 

Figuras 39A-D. Anticlimax locklini Pilsbry e Olsson, 1950. A-C: concha, 1,3 mm, Witties Reef, 
Nicaragua (MANS); D: microescultura. 


Anticlimax annae Pilsbry € Olsson, 1950 (Figures 40A-D) 


Anticlimax annae Pilsbry € Olsson, 1950. Bull. Amer. Paleont., 33: 113. n* 135: 11, pl. 3, figs. 12 a- 
b. [Type locality: Plio-Pleistocene of St. Petersburg, Florida —fossil species-]. 


Type material: Holotype and paratype in ANSP (n” 18397). Not examined. 
Other material examined: Mexico: 1 s, Puerto Morelos, 24 m, Yucatan (MHNS). 


7Ó 


RUBIO ET 4L.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 40A-D. Anticlimax annae Pilsbry 82 Olsson, 1950. A-C: shell, 1.6 mm, Puerto Morelos, 
Yucatan (MHNS); D: microsculpture. 
Figuras 40A-D. Anticlimax annae Pilsbry € Olsson, 1950. A-C: concha, 1,6 mm, Puerto Morelos, 
Yucatán (MHNS); D: microescultura. 


Description: This is the original 
description (PILSBRY €: OLSsON, 1950): 
“The rather strong, solid shell is convex 
below, and in the form of a low truncate 
cone above the angular periphery. The 


spire is very narrow, level or slightly 
sunken. There are about three and one- 
third whorls the first 1 Y glossy, convex, 
the next narrow and depressed at first, 
but increasing rapid ly near its end. The 


DS 


Iberus, 29 (2), 2011 


last whorl is very wide, sloping to the 
periphery. Sculpture of punctuate 
spiral grooves, about twenty on the 
upper surface, the base with similar 
but weaker grooves. Radiating sculp- 
ture of rather strong waves on the 
penult whorl and about half of the last 
whorl, the base having about 8 to 15 
stronger radiating waves which fade 
out near the periphery. The aperture is 
subtriangular, a little effuse at the 
peripheral angle. The thick parietal 
callus extends well forward, and with 
the rather flattened, semicircular, col- 
umellar lobe, completely closes the 
umbilicus”. 

Holotype is 2.5 mm in diameter 
and 1.35 mm in height. Our shell (Figs. 
40A-C) is 1.6 mm in diameter. 

Habitat: Abundant in the Pliocene 
deposits of St. Petersburg. Our mater- 
ial was collected in sediments from 
coralline bottom. 


Distribution: Only known as a 
fossil from St. Petersburg and 
Orlando, Florida. Our specimen is 
clearly of recent origin. 

Remiark ss BRASS O SS ON 
(1950) stated: “This species differs 
from A. hispaniolensis and A. his- 
paniolensis cratera by the very 
strong parietal callus and the more 
flattened callus pad which wholly 
closes the umbilicus. It is rather vart- 
able in size and especially in the 
development of radial waves on the 
upper surface. In most specimens 
examined, waves are present on the 
first half of the last whorl, but occa- 
sionally they do not extend beyond 
the penultimate whorl, the whole 
last whorl lacking of them”. 

A. locklini is distinguished by the 
more pronounced peripheral keel 
and by a small thickened parietal 
callus and columella. 


Anticlimax hispaniolensis cratera Pilsbry € Olsson, 1950 (Figures 41A-E) 


Anticlimax hispaniolensis cratera Pilsbry € Olsson, 1950. Bull. Amer. Paleont., 33: 116, n* 135, pl. 
2, fig. 6, pl. 3, figs. 11-11b. [Miocene of Banana River, Costa Rica]. 


Type material: Holotype in ANSP (n” 18403). Not examined. 
Other material examined: Nicaragua: 1 s, Witties, 20-30 m (MHNS). 


Description: This is the original 
description (PILSBRY éz OLSSON, 1950): 
“The shell is similar to A. hispaniolensis 
in form, but the spire is somewhat more 
sunken. There are 3 1/ 3 whorls, the first 
two planorboid, narrow. The last whorl 
rises in the narrowly rounded ridge, 
crater-like, around the spire, then slopes 
stratghtly to the blunty carinate periph- 
ery, below which it is rather weakly 
convex. Sculpture of strongly impressed 
punctuate (or minutely z1g2a8) spiral 
lines and radiating waves on the base. 
The aperture is triangular, the upper 
margin of the peristome nearly straight, 
extending above the peripheral keel. The 
umbilicus is filled by a callus which 
expands into a strongly convex semicir- 
cular pad, united with the columellar 
callus”. 


78 


Dimensiones: Our shell (Figs. 
41A-C) has 1.0 mm in diameter. 

Habitat: Our shell was collected 
in coralline bottom at a depth of 
20-30 m. 

Distribution: Only known as a 
fossil species from Miocene of 
Banana River, Costa Rica (PILSBRY 
£z OLSSON, 1950) and Nicaragua. 

Remarks: PILSBRY € OLSSON 
(1950) stated: “The close relation- 
ship between A. hispaniolensis 
cratera and A. hispaniolensis has 
been noted under the latter species. 
Both are based upon specimens not 
jullysmiature- Wero elevo entiabktlve 
posterior grooved commissure of 
the aperture, conspicuous in these 
specimens, would be nearly or 
wholly obliterated and the peris- 


RUBIO ET 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 41A-E. Anticlimax hispaniolensis cratera Pilsbry 82 Olsson, 1950. A-C: shell, 1.0 mm, 
Witties, Nicaragua (MHNS); D: protoconch; E: details of the microscupture. 

Figuras 41A-E. Anticlimax hispaniolensis cratera Pilsbry € Olsson, 1950. A-C: concha, 1,0 mm, 
Witties, Nicaragua (MHNS); D: protoconcha; E: detalle de la microescultura. 


tome thickened in the adult stage, distinguished by the shape and 


as in A. annae”. larger extension of the columellar 
This species has some similarity callus as well as the suppression of 
to A. locklini, from which it can be the radiating waves on the base. 


79 


Iberus, 29 (2), 2011 


Genus Aorotrema Schwengel € McGinty, 1942 


Aorotrema Schwengel €: McGinty, 1942. The Nautilus, 56: 17 (as a subgenus of Cyclostrema). 
Type species: Cyclostrema (Aorotrema) pontogenes Schwengel £ McGinty, 1942 (by original 


designation). 


Diaenosis stella je 
whorls which are strongly angular 
above, at the periphery and around 
the broadly funnel shaped umbili- 
cus; te aperture  subimangular 
peristome only shortly in contact 
with preceding whorl or free”. 

Remarks: Four species from the 
western Atlantic have been 
included in the genus Aorotrema: 
A. cistronium (Dall, 1889a), A. pon- 
togenes Schwengel €  McGinty, 
(AAA ea ic un sb 
McGinty, 1945 and A. gardnerae 
Pilsbry, 1953. The first three were 
considered as recent species while 
A. gardnerae was considered a 
fossil species from the Miocene. Of 
the recent species, A. cistronium 
and A. pontogenes have been con- 


sidered valid species. MOORE (1964: 
189), after the examination of the 
holotype of A. erraticum, decides 
that it is a juvenile of Turbo cas- 
tanea Gmelin, as a result of the sim- 
ilarity of the details of the proto- 
conch, shape, and teleoconch sculp- 
ture. Also, after the examination of 
a paratype of A. pontogenes he con- 
sidered that it is a juvenile shell 
probably of an Astraea. With 
respect to A. gardnerae, the fossil 
species from the Miocene, he stated 
that by its shape it could be also a 
juvenile Turbo. Therefore the simi- 
larity between the species of 
Aorotrema with young Turbo or 
Astraea is evident, the latter differ- 
ing by their possession of a calcare- 
ous operculum. 


Aorotrema cistronium (Dall, 1889) (Figures 42A-G) 


Cyclostrema cistrontum Dall, 1889a. Bulletin of the Museum of Comparative Zoology, 18: 394, 
(DaLtL, 1889b, pl. 42, fig. 11) [Type locality: off the coast of North Carolina [USFC stas. 2595, 
2598, 2608, 2610, 2612 (DALL, 1889b)]]. 


Type material: Type in ANSP (178697). Not examined. 

Material examined: Florida, USA: 1 s, E Gulf Blvd, NE end Sand Key (CHL); 42 s, trawled 27 m, 20 
mi E St. Augustine, (CHL); 71 s, dived, sponge grunge 36 m, 45 mi E Mayport, Duval Co. (CHL); 
1 s, dredged 54 m, 20 mi SE Pensacola Co, (CHL); 4 s, Ex batfish trawled, 39-45 m, 60 mi E Ponte 
Vedra, St. Johns Co. (CHL); 1 s, dredged, 53 m, 65 mi E St. Augustine, St. Johns Co. (CHL). Fossil: 


1 s, Pliocene of La Belle, FL (CHL). 


Description: The original description 
by DaLtL (1889) is as follows: 

“Shell small, white, with a polished 
nucleus, one and half rounded and as 
many more carinated whorls; spire 
depressed; radiating sculpture of fine 
close flexuous threads, which appear 
chiefly in the interspaces of the spirals, 
giving the surface a minutely punctate 
appearance; these extend over the 
whole surface except of the nuclear 
whorls; spiral sculpture of on the 
summit seven or eight, between the 


80 


carinae six or eight, and on the base ten 
or fifteen extremely fine threads, even 
and uniform, with about equal inter- 
spaces, some a little granular from the 
radiating sculpture; beside these there 
are three very strong carinae; one forms 
the margin of the nearly flat spire, the 
second extends horizontally just below 
the periphery, the space between them 
deeply excavated; the third forms the 
edge of the funicular narrow deep 
umbilicus. The base is conical, exca- 
vated just within the peripheral carina; 


RUBIO ET 4L£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


it rises to the edge of the umbilicus, 
which is marked by a strong thread, and 
within is vertically striated. The last 
whorl descends from the general plane 
and finally becomes separated from the 
body whorl; the margin is simple, 
sharply angulated by the carinations, 
otherwise the aperture would be ovate, 
with the columellar side somewhat 
excavated ”. 

Alt. 1.6; max. diam. 2.0 mm. 

Maximum reported size: 2.5 mm. 

Habitat: Off the coast of North Car- 
olina, in 22-63 fms (40-130 m), on a sand 
and gravel bottom, in the warmer area 
(DaLL, 1989a). From sponge-associated 


debris, in 39 m, East of Mayport, Florida 
(LEE, 2009). 

Depth: 14 to 115 m (live 22 to 46 m). 

Distribution: USA: North Carolina 
(DaLL, 1989a), Texas (ODÉ, 1987b) and 
Florida (LEE, 2009). CAMPBELL (1993) 
recorded it as a fossil from the Pliocene 
of Yorktown and Chowan river forma- 
tions in Virginia. 

Remarks: The material examined 
agrees with that figured by PILSBRY 
(1953, pl. 54, figs. 5-5c) from St. Peters- 
burg Plio-Pleistocene. 

Some shells (Fig. 43E) have a separa- 
tion of the aperture from the terminal 
body whorl. 


Aorotrema pontogenes (Schwengel £: McGinty, 1942) 


Cyclostrema (Aorotrema) pontogenes Schwengel € McGinty, 1942. The Nautilus, 56: 17-18, pl. 3, 
fig. 3 [Type locality: South by West of Destin, Florida, 18 miles off shore]. 


Type material: A paratype in ANSP (178697). Not examined. 


Description: This is the original 
description in SCHWENGEL éz MCGINTY, 
(1942): “The openly umbilicate, white 
shell is strongly bicarinate with flattened 
spire, of 3 whorls, the first two planor- 
boid. The first 1 whorls are smooth, 
convex, the convexity increasing on the 
next whorl, and overhanging outward ly 
on the last whorl, forming a strong but 
blunt upper carina. There is a more 
extended and somewhat up-curved carina 
at the periphery, la deep concavity 
between the two carinae. Below the 
peripheral keel the surface slopes straight 
to the prominent ridge around the umbili- 
Cus. 

On the last whorls there is a secondary 
sculpture of fine low spiral threads, their 
intervals crossed by finer growth lines, 
giving a minutely punctate appearance in 
some places; these spirals not extending 
into the funnel-shaped umbilicus. Aper- 
ture with the columellar margin deeply 
concave in the middle. Outer lip inserted 
just below the keel of next-to-last whorl, 
gently sloping to the first carina, then 
concave to the stronger and more 
extended peripheral carina, below which 
it slants straight to the base”. 


Height 1.3 mm, diameter 1.9 mm. 

The umbilicate shell is strongly 
bicarinate. Protoconch of one whorl, 
smooth and glassy. Teleoconch con- 
sists of 1 Y whorls; the spire is 
almost flat out to the shoulder. 
Carina strongly angular and outer 
slope is concave. Peripheral carina 
strong and flat on top. Ornamenta- 
tion formed by fine punctate spirals, 
present over all the shell, even in the 
inner umbilicus. Aperture modified 
by the carina, columella large and 
concave. Umbilicus narrow and 
deep, well delimited by the umbilical 
carina. 

Habitat: Dredged in 18 to 20 
fathoms (32-36 m), off Destin, North- 
west Florida. Collected on sandy marl 
bottom. 

Distribution: USA: Florida: West 
Florida (SCHWENGEL é MCGINTY, 
1942). 

Remarks: SCHWENGEL $ 
MCGINTY (1942) mention: “This 
species is similar in size, sculpture 
and form to C. cistronium Dall, but 
the spire 18 flatter, second. carina 
mue more “extend ea ep 1e - 


8] 


Iberus, 29 (2), 2011 


Figures 42A-G. Aorotrema cistronium (Dall, 1889). A-E: shells, 2.1, 1.86, 1.8, 1.8, 1.7 mm, 
Florida, (CHL); F: microsculpture; G: protoconch. 

Figuras 42A-G. Aorotrema cistronium (Dall, 1889). A-E: conchas, 2,1, 1,86, 1,8, 1,8, 1,7 mm, 
Florida, (CHL); E: microescultura; G: protoconcha. 


crust” manner, the last whorl not MOORE (1964) thought this might be 
descending and the aperture more juvenile Astralium phoebium. 

angled and not separated from the We lack material on which to base an 
body whorl as in C. cistronium”. opinion. 


82 


RUBIO £7 4Lz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Genus Cyclostremiscus Pilsbry € Olsson, 1945 


Cyclostremiscus Pilsbry €: Olsson, 1945: 266. 


Type species: (by original designation) Vitrinella panamensis C.B. Adams, 1852. Proc. Acad. Nat. 


Sci. Phila., 97: 266. Recent, Caribbean. 


Diagnosis: The description of the 
genus (PILSBRY € OLSSON, 1945) is as 
follows: “Shell small, solid, moderately 
strong, depressed or discoidal, much 
wider than high, umbilicate, of few 
whorls, from which the first 1 Y to 2 
form a smooth glassy protoconch. Last 
whorl typically with several spiral 
angles or carinae. Their intervals 
usually with lower axial riblets or 
striae. Aperture subcircular or modi- 
fied by the angles of the shell, the peri- 
stome continuous, not thickened exter- 
nally”. 

Remarks: PILSBRY € OLSSON (1945) 
introduced Cyclostremiscus for a series 
of small or minute species, abundantly 
represented in the West American 
tropics. Some of them had been 
described in the genera Vitrinella and 
Cyclostrema. They designate Vitrinella 
panamensis C. B. Adams, 1852 as the 
type species of the genus; this species 
is characterized by “the last whorl typ- 
ically having several spiral angles or 
carinae, their intervals typically with 
lower axtal riblets or striae”. 

PILSBRY (1953) described the sub- 
genus Ponocyclus with Adeorbis 
beauii as the type species, and distin- 
guished it from Cyclostremiscus by its 
lack of axial sculpture. 

MOORE (1964) divides Cyclostrem- 
iscus species into three small groups: 
Cyclostremiscus beauti, C. dalli and C. 
pentagonus are practically identical in 
their general appearance; all have 2 or 
3 keels on the periphery and several 
grades of finer spiral sculpture. C. sup- 
pressus and C. jeannae resemble each 
other with their smaller umbilicus and 
three-keeled periphery. C. cubanus 
does not seem to have any relation to 
any of the previously mentioned 
species. 

BIELER 6: MIKKELSEN (1988) after the 
study of the anatomy of Circulus 
texanus and Cyclostremiscus beauil 


making a comparison with Circulus 
striatus, conclude saying that the differ- 
ences between both genera are mainly 
anatomical. 

PONDER (1994) mentions that the 
assignation of Cyclostremiscus beauii to 
the genus Cyclostremiscus needs a revi- 
sion. 

The genus Cyclostremiscus Pilsbry 
£ Olsson, 1945 is based on Vitrinella 
panamensis C.B. Adams, 1852. The type 
species is small, has strong axial sculp- 
ture (distinct axials lacking in Cy. 
beauii), a few strong spiral ribs which 
are rendered nodulose by the axials, and 
has an apertural varix. These shell char- 
acters are markedly discordant with 
those of “Cy”. beauii. The genus Pono- 
cyclus (Pilsbry, 1953) was erected for 
Adeorbis beauti and it can be used for 
this species. 

If we accept this establishment, we 
could differentiate this group of species 
placing them outside Cyclostremiscus 
and including them in Ponocyclus, but 
this could only be tentative, because it is 
necessary to have an anatomical study 
of each species in order to decide their 
correct final position. 

RuBIO € ROLÁN (1998) presented 
radular information on Pachystremiscus 
ornatus Olsson € McGinty, 1958 and 
Pachystremiscus pulchellus Olsson € 
McGinty, 1958 and placed both species 
in the genus Lodderena (Archaeogas- 
tropoda, Skeneidae), considering the 
genus Pachystremiscus as a synonym. 

We have considered the comments 
of BIELER é MIKKELSEN (1988) and 
PONDER (1994) about the differences 
between Circulus and Cyclostremiscus, 
which are mainly anatomical. Also there 
is not a basic model of morphological 
characters which allows us to place the 
species in one or the other genus. So, it 
is very difficult to make a generic differ- 
entiation in the known species placed in 
group 1, which could be placed also in 


83 


Iberus, 29 (2), 2011 


Circulus. Only an anatomical study of 
each of them would allow us to know 
the correct genus. For all of these 
reasons, we prefer to keep these 
species in their present status, as con- 
sidered by other authors. 

The genus Cyclostremiscus is in 
our opinion one of the most concho- 
logically heterogeneous of those 
within the subfamily Vitrinellinae. All 
the previously described recent taxa 
have been studied and figured in the 
present work, and two taxa new to 
science are also treated. 


In an attempt to better differentiate 
Cyclostremiscus species we have distin- 
guished two groups which clearly differ 
from the other congeners: 

Group 1. Cyclostremiscus beauti — C. 
multiliratus - C. dalli — C. pentagonus — 
C. trilix —C. hendersonl. 

Group 2. C. bartschi (only fossil) - C. 
jeannae - C. suppresus - C. vanbruggen1 
Es 

The species not included in these 
groups have such distinctive morpho- 
logical characters that their identifica- 
tion is relatively simple. 


Group 1 


These species are quite similar in general appearance; all have 2 or 3 
peripheral keels and several degrees of smaller spiral sculpture. 


Cyclostremiscus beauii (P. Fischer, 1857) (Figures 43A-E) 


Cyclostrema angulatum auct. non A. Adams, 1850. 

Adeorbis beauii P. Fischer, 1857. Journal de Conchyliologie, 6: 286, pl. 10, fig. 12. [Type locality: 
Guadeloupe]. 

Cyclostrema bicarinatum Guppy, 1866. Quarterly Journal of the Geological Society of London, 
22(1): 281-295, pls. 16-18. [Type locality: “Miocene” (Lower Pliocene), Jamaica]. 

Skenea sulcata Simpson, 1887: 61 [nomen nudum, see MOORE, 1964: 131]. 

Adeorbis beauii bicarinatus (Guppy, 1866): Dall, 1903: 1595. [Type locality: Oligocene of Jamaica]. 

Circulus bicarinatus (Guppy, 1866): in WOODRING, 1828: 439, pl. 37, figs. 10-12. 

Circulus stirophorus M. Smith, 1937. The Nautilus, 51: 67, pl. 6, figs. 2a, b [Type locality: Plio-Pleis- 
tocene, Florida]. 


Type material: BIELER 8 MIKKELSEN (1988) designated the only specimen extant in the MNHN, cor- 
responding to the lot deposited by Fischer, as lectotype. 

Other material examined: Florida, USA: 2 s, 39-45 m, 50-60 mi., E Ponte Vedra, St. Johns Co., Fla. 
(CHL); 3 s, Delray Beach, Palm Beach Co. Fla (CHL); 2 s, 2 j, near Mantis Shrimp Holes, St. Lucia 
Inlet, Stuart Martin Co., Fla. (CHL); 1 s, 84 m, WNW Tortugas (CHL); 2 s, 54-60 m, Cedar Keys, Fla 
(CHL). Cuba: 3 s, Cienfuegos Bay, 10 m (MHNS). Colombia: 1 s, Cayos de San Andrés, in coral bottom 
20-25 m (CHL). 


Description: It is one of the 
better studied species of the Vit- 
rinellinae, including morphology, 
radula, ecology, and anatomy, see 
BIELER 6 MIKKELSEN (1988). 

Maximum reported size: 12.5 
mm. The studied and  pho- 
tographed material (Figs. 43A-C) 
from Cuba measures 8.1 mm in 
maximum diameter and 3.0 mm 
in height. The shell from Colom- 
bia in the CHL is 10.1 mm. 


84 


Distribution: The species is 
known in the Western Atlantic, 
from North Carolina to Brazil. 
Recorded from Puerto Rico 
(WARMKE é ABBOTT, 1975). From 
Campeche to Ciudad del 
Carmen and Zacatal, from El 
Euyo ito Ninum Pont trom 
Yalkupul Point to Cerritos 
Island and from Cancún to the 
Belize border, Mexico (VOKES éx 
VOKEs, 1983). From North Car- 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 43A-E. Cyclostremiscus beauii (P. Fischer, 1857). A-C: shell, 8.1 mm, Cienfuegos Bay, 
Cuba; D: protoconch; E: detail of the microsculpture. 

Figuras 43A-E. Cyclostremiscus beauii (P Fischer, 1857). A-C: concha, 8.1 mm, Bahía de Cienfue- 
gos, Cuba; D: protoconcha; E: detalle de la microescultura. 


olina to Florida; the Antilles; 
Ceará, Pernambuco and 
Alagoas, Brazil (Rios, 1985). 
Guadaloupe; Jamaica; Florida; 
the Antilles; North Carolina; 
Dry Tortugas; Sint Maarten; 
Colombia; from North Carolina 
to Brazil (BIELER € MIKKELSEN, 
1988). "Aruba. (DE  JoNnec. € 
COOMANS, 1988). From North 
Carolina to Brazil (ROBINSON, 
1991). From North Carolina to 


the central coasts of Brazil (DíAZ 
MERLANO € PUYANA HEGEDUS, 
1994). Carolinas to West Indies 
(Morris, 1973). Laguna Indian 
River, Florida (MIKKELSEN ET AL, 
1995). Cuba (ESPINOSA ET AL., 
1985). 

Habitat: BIELER € MIKKELSEN 
(1988) reported this species 
inhabiting burrows of the stom- 
atopod Lysiosquilla scabricauda 
(Mantis Shrimp), between 0.5 


85 


Iberus, 29 (2), 2011 


and 1 m in depth. Other authors 
mention its shells in the diges- 
tive tracts of the  seastar 
Asno pe cien anciana 
1825), probably as prey, but par- 
asitism is possible. Other 
recorded habitats are between 0 
and 46 m in depth, sand, 
muddy, and rocky bottom 
(VOKES é VOKES, 1984); sand in 
shallow water (Díaz MERLANO 
é£ PUYANA HEGEDUS, 1994), but 
these are based on empty shells. 

Discussion: €. beautt is the 
largest vitrinelid in the West 
Atlantic. It may be «distin- 
guished from other known 
species by its larger size, by its 


bicarinate periphery, and by the 
numerous and strong spiral cords 
which form its dorsal sculpture. 
Two recent species (Cyclostrem- 
iscus major Olsson € Smith, 1951 
and Cyclostrema gordana 
Hertlein é Strong, 1951) from the 
Panamic region are very similar 
nsize, isculpture tan docu 
BIELER 6 MIKKELSEN (1988) con- 
sider Cyclostrema  bicarinata 
Guppy and Circulus stirophorus 
Smith synonyms of C. beauii, 
indicating that the variation in 
the basal sculpture, probably due 
to ontogenetic change, was not 
enough reason for their differen- 
tiation. 


Cyclostremiscus multiliratus spec. nov. Rubio, Rolán € Garcia (Figures 44A-F) 


Type material: Holotype (Figs. 44A-C) in USNM (1155028) (ex CEG). 
Type locality: SW Florida, 2751.79 N — 84*59.82'W, dredged in 308-323 m. 
Etymology: The name refers to the numerous cords on the dorsum. 


Description: Shell (Figs. 44A-C) 
planispiral with spire slightly elevated, 
periphery bicarinate, and a wide 
umbilicus. Protoconch (Fig. 44F) 
slightly projected, with 1 Y. whorls, and 
a maximum diameter of about 530 um, 
apparently smooth with two stages dist 
inguis-hable: the first one, embry- 
onic] with % whorls; the second part is 
almost Y whorls. Indistinct separa- 
tion from the rest of the shell. Teleo- 
conch with 2 “4  whorls; sculpture 
formed by 2 keels which define the 
periphery of the shell, one placed on 
the middle part and the other in the 
basal area. First whorl with 10-12 spiral 
cordlets, with axial growth lines in 
the interspaces. The dorsum on the last 
whorl has 18-20 spiral cords, of which 6 
are very closely-placed and near the 
suture and 3-4 near the peripheral keel; 
the interspaces between the cords are 
covered by microtubercles. Between the 
peripheral keels there are 10-12 fine 
spiral cordlets with regular 
interspaces. On the base there is a spiral 
cord limiting the umbili- 
cal infundibulum. Base and inside the 


86 


umbilicus with numerous fine spiral 
cordlets. Aperture almost quadrangu- 
lar, prosocline. Peristome modified by 
the  peripheral keels;  columellar 
area arched, parietal zone not thick- 
ened. 

The holotype measures 3.18 mm in 
maximum diameter. 

Distribution: Only known from SW 
Florida, its type locality. 

Habitat: Deep water, dredged at 
308-323 m deep. 

Remarks: C. multiliratus spec. 
nov. is very similar in its general 
appearance to C. beauii, but the 
latter is larger, its protoconch has a 
smaller diameter, and the number of 
dorsal cords is smaller (5-7). Further, it 
lacks microtubercles between the dorsal 
cords and cordlets between the two 
peripheral keels. 

Another similar species is C. dalli, 
but it lacks ornamentation between the 
cords, has fewer dorsal cords, which 
are stronger and more elevated, and 
lacks spiral cordlets between the 
peripheral keels. Both species have a 
wide bathymetric range. 


RUBIO £7 AZ.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 44A-E Cyclostremiscus multiliratus spec. nov. Rubio, Rolán 82 García. A-C: holotype, 3.18 
mm (USNM); D: dorsal microsculpture; E: peripheral microsculpture; F: protoconch. 

Figuras 44A-E Cyclostremiscus multiliratus spec. nov. Rubio, Rolán ** García. A-C: holotipo, 3,18 
mm (USNM); D: microescultura dorsal; E: microescultura periférica; E: protoconcha. 


Iberus, 29 (2), 2011 


Cyclostremiscus dalli (Bush, 1897) (Figure 45A-D) 


Circulus dalli Bush, 1897. Transactions of the Connecticut Academy of Arts and Sciences, 10: 126, 
pl. 23, figs. 3-3a, 6. [Type locality: USFC sta. 2655, 27*22'N, 78%07'30"W]. 


Lydiphnis dalli (Bush, 1897). 


Type material: Holotype in YPM (15802); paratype in USNM (44983). Not examined. 
Other material examined: Florida, USA: 1 s, 36-90 m, off Dry Tortuga, Gulf of Mexico (CHL). 


Description: The original 
description (BusH, 1897) is as 
follows: “This deep-water species is 
of more delicate texture and more 
transparent than the more northern 
shallow-water species of similar 
form. It is ornamented on the body- 
whorl with two rather inconspicu- 
ous carinae, one defining the base 
and the other on the periphery; 
above tiistle is una ce isc UanO Y 
aboutaseventa delicate mea ua 
microscopic shallow grooves or 
striae, the two uppermost being the 
most distinet; above hesenive 
surface is smooth and appears 
somewhat flattened; there are also a 
few less distinct striae below the 
peripery anda nte uno dlical 
region. Greatest oidii 0 an; 
height: 1.4 mm”. 

We add: This larger species (Figs. 
45A-C) is characterized by having 2 
strong keels, one basal and the other 
at the middle of the periphery. On 
the dorsum, between the peripheral 
keel and the suture, 4-6 strong spiral 
cords can be observed without any 
sculpture between the cords except 
for very fine growth lines. Near the 
base there are about 8 spiral cords, a 
little smaller that the dorsal ones 
and 3-4 more very fine threads 


inside the umbilicus. The proto- 
conch (Fig. 45D) is wide, and has a 
little more than one whorl, about 
540 um in diameter, apparently 
smooth but perhaps with some 
small axial cordlets mainly at its ter- 
minus. 

Maximum reported size: 4 mm. 
The shell figured here is 4,3 mm in 
diameter and 1.87 mm in height. 

Habitat: Among Foraminifera, in 
338 fms (608 m) (Bush, 1897). Bathy- 
metric range: 79 to 618 m. 

Distribution: USA: North Car- 
olina, Eloridas lord ae ys 
Bahamas: Great Bahama Bank 
(BusH, 1897 and MOORE, 1964). 

Remarks: This larger species, 
with its inconspicuous sculpture, 
seems to be a connecting link 
between the distinctly grooved ones 
and the carinated ones, so we can 
make a series of gradation in sculp- 
ture from the smooth variety of 
Cyclostremiscus supranitidus, fol- 
lowed by C. supranitidus (typical), 
a Co a Cs SUCUaLa, (Co SÉFUOS 
tus, up to the strongly grooved C. 
liratus (Bush, 1897). The latter 
species may be distinguished from 
E pentaco nus ando 
having two peripheral keels instead 
of three. 


Cyclostremiscus pentagonus (Gabb, 1873) (Figures 46A-F) 


Adeorbis supranitidus auct. non Wood, 1842. 


Cyclostrema pentagonum Gabb, 1873. Amer. Philos. Soc. Trans. New series, 15: 243. [Type local- 


ity: Miocene of the Dominican Republic]. 


Vitrinella pentagonus (Gabb, 1873): GABB, 1881: 368, pl. 47, fig. 68. 
Cyclostremiscus pentagonus (Gabb, 1873): WOODRING (1928: 73). 


Type material: Holotype in ANSP (2831) represented in MOORE (1964, fig. 23). Not examined. 
Material examined: Cuba: 5 s, Rancho Luna Beach, 40 m (MHNS); 2 s, Cienfuegos Bay, 10 m (MHNS). 


88 


RUBIO £7 AZz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 45A-D. Cyclostremiscus dalli (Bush, 1897). A-C: shell, 4.3 mm, off Dry Tortuga, Florida 
(CHL); D: protoconch. 
Figuras 45A-D. Cyclostremiscus dalli (Bush, 1897). A-C: concha, 4,3 mm, frente a Dry Tortuga, 
Florida (CHL); D: protoconcha. 


Description: Shell (Figs. 46A-C) 
rounded, depressed, whitish to yel- 
lowish brown in color, solid. Proto- 
conch (Fig. 46D), erect projecting, 
with a little more than 2 whorls, 
smooth or slightly rough, and with a 
diameter of about 420 um. Two well 
differentiated parts can be seen on it: 


the first one, embryonic, is shorter, 
with a little more that % whorl; the 
second is almost 1 Y whorls. The sep- 
aration from the rest of the shell is 
distinct. Teleoconch with 2 rapidly- 
increasing whorls; sculpture formed 
by 3 keels which define the periphery. 
There is a spiral cord which delimits 


89 


Iberus, 29 (2), 2011 


Figures 4G6A-E. Cyclostremiscus pentagonus (Gabb, 1873). A-C: shells, 2.9, 2.7, 2.3 mm, Rancho 
Luna Beach, Cienfuegos, Cuba (MHNS); D: shell, 2.1 mm, Brazil (CHL); E: microsculpture; E: 
protoconch. 
Figuras 4GA-E Cyclostremiscus pentagonus (Gabb, 1873). A-C: conchas, 2,9, 2,7, 2,3 mm, Playa 
Rancho Luna, Cienfuegos, Cuba (MHNS); D: concha, 2,1 mm, Brasil (CHL); E: microescultura; F: 


protoconcha. 


the umbilicus and fine spiral cords 
more evident in the spaces between 
the keels. Also there are fine sinuous 
axial growth lines. Umbilicus wide 
and deep exposing the previous 


90 


whorls, even the protoconch; on its 
interior up to 5 spiral cords can be 
seen. Aperture hexagonal, more dis- 
tinctly at the periphery because of the 
keels. 


RUBIO ET AZz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Dimensions: The holotype mea- 
sures 3.5 mm. The studied shells were 
up to 2.96 mm in maximum diameter. 

Habitat: The bathymetric range of 
this species is very wide, having been 
recorded between 0 and 538 m in 
depth for the three synonymized 
species; the records for living speci- 
mens are between 0 and 45 m: 
between 13 and 31 m (ABBOrTT, 1974); 
in muddy sand in the surf zone 
(ANDREWS, 1977); in shallow water 
beetween 13 and 30 m in depth (Díaz 
MERLANO éz PUYANA HEGEDUS, 1994). 

Distribution: USA, Mexico, Costa 
Rica, Colombia; Venezuela, Puerto 
Rico and Cuba. Recorded from Cape 
Hatteras, North Carolina (Bush, 1885). 
From Puerto Rico (WARMKE é€ ABBOTT, 
1961). From North Carolina, north- 
western Florida and Alabama, Texas 
and Campeche Bank, Miocene and 
Pliocene of Panama, Jamaica, Domini- 
can Republic and Florida (MOORE, 
1964). From North Carolina (PORTER, 
1974). Off North Carolina to Florida, 
Texas and the West Indies (ABBOTT, 
1974; EMERSON é€ JACOBSON, 1976). 
From the southeast of USA to the 
Antilles and Texas (ANDREWs, 1977). 
From Laguna de Termino, Campeche 
(GARCÍA-CUBAS, 1981). From Vene- 
zuela (PRINCZ, 1982). From Campeche 


to Ciudad del Carmen and Zacatal, 
and from Cancún to the Belize border, 
Mexico (VOKEÉS € VOKES, 1984). From 
Puerto Rico (ORTIZ-CORPS, 1985). From 
Moín, Costa Rica (ROBINSON  éz 
MONTOYA, 1987). From North Carolina 
to Florida, Texas and the Caribbean 
Sea (Lyons, 1989). From off North Car- 
olina to the Caribbean (ROBINSON, 
1991). North Carolina Florida, West 
Indies, South Brazil (Rios, 1994). From 
North Carolina to the South American 
coast (Díaz MERLANO é PUYANA 
HEGEDUS, 1994). From Cuba (EspI- 
NOSA, FERNÁNDEZ-GARCÉS € ROLÁN, 
1995). From Abaco, Bahamas Islands 
(REDFERN, 2001). From Cahuita to 
Gandoca, Costa Rica (ESPINOSA éz 
ORTEA, 2001). 

Remarks: Until its definitive place- 
ment in Cyclostremiscus, this species 
was placed in the genera Adeorbis, 
Circulus, Lydiphnis, Skenea and Vit- 
rinella. The prominent protoconch and 
its tricarinate periphery in the recent 
samples, give the shells of this species 
a Characteristic appearance and easily 
distinguish it from congeneric species. 
MOORE (1964) mentioned having seen 
living specimens of this species, and 
the animals showed the typical charac- 
ters of the vitrinellids, with ciliated 
tentacles and without epipodia. 


Cyclostremiscus trilix (Bush, 1885) (Figures 47A-D) 


Skenea trilix Bush, 1885. Expl. Albatross. Report U.S. Com. Fish and Fisheries for 1883: 464. 
[Type locality: Off Cape Hatteras, North Carolina, 7-17 fathoms (13-31 m) [USFC sta. 2113 


(JOHNSON, 1989)]|]. 


Circulus trilix (Bush, 1885): Bush, 1897: 127, pl. 22, figs. 6, 10, 12; pl. 41, fig. 7. 
Cyclostremiscus trilix (Bush, 1885) in MOORE (1961: 18). 


Type material: Holotype in USNM (35365). Not examined. 
Material examined: Panama: 1 s, 600-700 m, off Atlantic coast (CHL). 


Description: The original descrip- 
tion in BusH (1885: 584) is as follows: 
“This species closely resembles Adeor- 
bis supranitida Wood, in form and 
seulpture, butt has a thin, horny 
operculum and an animal like 
Skenea”. Depth range, 14 to 15 
fathoms (25-27 m). 


BIELER €  MIKKELSEN (1988) 
mention: “Based on the misconception 
that these species are archaeogas- 
tro piods; BUSsa (1997:127, 142, ple 22, 
figs. 47A-G) erroneously constructed a 
rhipidoglossate radula for Circulus 
trilix (Bush, 1885) [= Cyclostremiscus 
pentagonus (Gabb, 1837), fide MOORE, 


Sn 


Iberus, 29 (2), 2011 


Figures 47A-D. Cyclostremiscus trilix (Bush, 1885). A-B: shells, 2.3, 1.7 mm, 600-700 m, off 
Atlantic coast, Panama (CHL); C: microsculpture; D: protoconch. 

Figuras 47A-D. Cyclostremiscus trilix (Bush, 1885). A-B: conchas, 2,3, 1,7 mm, 600-700 m, frente 
a la costa Atlántica, Panamá (CHL); C: microescultura; D: protoconcha. 


1964: 138]”. Its radula is taenioglos- 
sate like any species of the Tornidae. 

We add: Protoconch (Fig. 47D) 
projecting with almost 1 % smooth 
whorls, about 440 um in maximum 


az 


diameter; under high magnification 
irregular microtubercles can be seen 
on its surface. A strong varix marks 
the transition to the teleoconch. On 
its first quarter whorl the teleoconch 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


exhibits a strong spiral cord which 
will later become a keel placed on 
the dorsal aspect of the periphery. 
There is no other dorsal spiral sculp- 
ture. On the ventral aspect, three 
spiral cords delimit the umbilicus. 
Aperture rounded; peristome contin- 
u0us. 

Maximum reported size: 3 mm 

Habitat: The bathymetric range of 
this species is quite ample, having 
been collected between 2 and 538 m 
in depth. The species is relatively 
common in Cape Hatteras between 7 
and 17 fms (19-31 m) of depth. The 
living material was collected 
between 26 and 27 m (Bush, 1897). 

Distribution: USA: Massachu- 
setts, North Carolina (Bush, 1885b; 
DaLL, 1927), Florida: West Florida 
(MOORE, 1964); Texas (PARKER, 1959; 
MOORE, 1964); Puerto Rico (WARMKE 
éz ABBOTT, 1961). 


Remarks: Cyclostremiscus trilix 
was described initially in the 
genus Skenea and later transferred 
to the genus Circulus, Lydipnis 
and Vitrinella. WOODRING (1957) 
gives a lengthy discussion on C. 
trilix and C. pentagonus, conclud- 
ing that the only difference was 
that C. trilix was larger. MOORE 
(1964) considers the two synony- 
mous. 

We believe that there are 
enough characters to distinguish 
these two taxa at the species level. 
The presence of microtubercles on 
the protoconch and the lack of 
spiral microsculpture between the 
perypheral keels allow us to differ- 
entiate Cyclostremiscus trilix from 
Cyclostremiscus pentagonus. The 
shell figured here is very similar to 
that described and figured by 
BusH (1897). 


Cyclostremiscus hendersoni (Dall, 1927) (Figures 48A-F) 


Lydiphnis hendersoni Dall, 1927. Proceedings of the United States National Museum, 70(2667): 


124-125. [Type locality: Off Georgia]. 
Circulus henderson1 Dall. 


Type material: Lectotype (Figures 48A-F) in USNM (108396). Examined by SEM. 


Other material examined: None. 


Description: This is the original 
description by DaLL (1927): “Shell 
small, white, depressed, glassy, with 
about three tricarinate whorls, the 
nucleus deeply sunken; axial sculpture 
of faint incremental lines, on the spire 
retractively oblique and on the base 
protractively arcuate; spiral sculpture 
of a prominent carina at the periphery, 
a less prominent one on the spire about 
one-third of the way from the suture to 
the periphery, and on the base another 
slightly nearer the umbilicus than to 
the periphery; these two secondary 
keels are variable, sometimes strong, 
sometimes almost obsolete, sometime 
partly one or the other on the same 
specimen; the edges of the carina are 
sometimes finely spirally striated, sug- 
gesting that they may when fresh and 


intact carry a series of epidermal 
fringes like Episcynia; the space within 
the basal carina is funicular, extending 
to a moderately narrow perforate 
umbilicus; the aperture would be 
rounded, but in those specimens in 
which the carinae are all developed the 
thin margin is modified and the aper- 
ture is angulated by them; in some 
specimens faint spiral striation is indi- 
cated on the. surface between the 
carinae; the margin of the aperture in 
the adult is continuous over the body 
of the shell. Maximum diameter, 3.5 
mm; minimum diameter, 2.9 mm; 
height, 1.3 mm”. 

We add: The protoconch (Fig. 48F) 
is of small size, has a rough aspect and 
terminates with a thickened varix. Size 
about 170 um in maximum diameter. 


73 


Iberus, 29 (2), 2011 


Figures 48A-E. Cyclostremiscus hendersoni (Dall, 1927). A-E: Lectotype in different positions, 3.5 
mm (USNM, 108396); F: protoconch. 
Figuras 48A-E Cyclostremiscus hendersoni (Dall, 1927). A-E: Lectotipo en distintas posiciones, 3,5 
mm (USNM, 108396); F: protoconcha. 


The teleoconch has a very prominent 
peripheral keel placed in the middle of 
the shell; also two less prominent cords, 
one of them on the dorsum at the mid- 
point between the suture and the outer 


94 


keel; and the other ventral, closer to the 
periphery. The dorsal cord slowly fades 
out disappearing near the middle of the 
body whorl. Under high magnification 
very fine spiral striae can be seen in the 


RUBIO ET 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


spaces between the cords and the keel. 
Another character typical of this shell 
is that the last whorl envelops the pre- 
vious one below the peripheral keel, 
which, being prominent, hides this 
suture. A thickening of the umbilical 
wall progressively covers the umbili- 
cus, which is narrow and deep. Inter- 
nal lip a little thickened and reflected 
outward, further impinging on the 
umbilicus. 

Habitat: Dredged from 805 meters 
depth, in broken coral, on a coarse 
sand and broken shell bottom, where 
numerous samples were collected. 

Distribution: Only known from the 
type locality. 

Remarks: DALL (1927) mentions the 
following: “The periostracum on Epis- 
eynia is extremely fugacious, and it is 
not prudent to assume that these and 


other small shells dredged and 
showing none are normally without 
one. Á few have a persistent perios- 
tracum, and there is no good reason to 
suppose that others never posses one”. 

The depressed spire, the tricarinate 
shell, with spiral striae between the 
keels, and the form of the umbilicus 
allow us to place this species in the 
genus Cyclostremiscus. Only the pro- 
toconch gives us some doubts, which 
can not be resolved due to the lack of 
live-collected material to permit study 
of the radula. Cyclostremiscus pen- 
tagonus, C. trilix and C. dalli are the 
species with greatest affinity, princi- 
pally because of their shells bearing 
three spiral keels. The present species 
can be distinguished from them by the 
form of the inner lip and the ornamen- 
tation of the umbilicus. 


Group 2 


This group of species clearly shows the evolution from the fossil forms with 
elaborate ornamentation (Cyclostremiscus bartschi) towards the recent 
species with more subdued sculpture (Cyclostremiscus spec. nov.). 


Cyclostremiscus bartschi (Mansfield, 1936) (Figures 49A-D) 


Cyclostrema bartschi Mansfield, 1936. Florida State Geol. Surv. Bull. N* 3: 132, pl. 20, figs. 13-15. 
[Type locality: Upper Miocene of Harveys Creek, Leon Co., Florida] Fossil. 


Material examined: Florida, USA: 1 s, Plio-Pleistocene Belle (CHL). 


Description: In PiLsBRY (1953): “Shell 
is rather solid, discoidal, the upper 
surface flattened, with a spiral ridge 
midway on the last whorl, the vertical 
peripheral zone bounded by keels above 
and below; the base with a prominent 
keel bounding a broad, conic umbilicus. 
There are 3 Y whorls, the first weakly 
convex; following half turn is narrower. 
After the second turn the whorls in 
apical view are somewhat concave and 
weakly folded radially on both sides of 
a median spiral ridge. The peripheral 
zone has oblique folds, and base is 
folded radially. Umbilicus is conic and 
has smooth, straight sides. The aperture 
is moderately oblique, more or less dis- 


tinctly hexagonal, the peristome blunt, 
thickened at the basal angle and col- 
umellar slope, continued in a rather 
thick parietal callus, which is slightly 
advanced beyond the upper angle of the 
mouth”. 

Height 0.9 mm, diameter 1.55 mm. 

Habitat: Dredged from 28 and 30 
fms (50-54 m), Sta. 2112 and 2110. 

Distribution: TREECE (1980) mentions 
this species from the northeastern shelf 
and upper slope of Yucatan. Rios (2009) 
also records it from Guatemala and East 
Brazil. 

Remarks: This species was initially 
considered exclusively fossil following 
its original description. TREECE (1980) 


SS 


Iberus, 29 (2), 2011 


Figures 49A-D. Cyclostremiscus bartschi (Mansfield, 1936). A-D: shell, 1.4 mm, Plio-Pleistocene of 


La Belle, Florida (CHL); D: protoconch. 


Figuras 49A-D. Cyclostremiscus bartschi (Mansfield, 1936). A-D: concha, 1,4 mm, Plio-Pleistoceno 


de La Belle, Florida (CHL); D: protoconcha. 


mentioned it as a recent species from 
Yucatan, Gulf of Mexico, but this record 
was considered dubious by Malacolog, 
version 4.1.1. <www.malacolog.org>. A 
Database of Western Atlantic Marine 
Mollusca. Rios (2008) recorded it again 
from Guatemala and East Brazil, and 
based on these records we have consid- 
ered Cyclostremiscus bartschi as a 


96 


recent species. Morphologically, the 
vertical periphery, limited by two 
keels, and the wide spiral cord placed 
on the dorsum and on the base of the 
shell are the main differential charac- 
ters to distinguish it from closely 
related species. The nodules on the 
cords and keels are formed by the 
intersecting axial ribs. 


RUBIO ET 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Cyclostremiscus jeannae Pilsbry ££ McGinty, 1946 (Figures 50A-D) 


Vitrinella panamensis auct. non C.B. Adams, 1852. 
Cyclostremiscus jeannae Pilsbry £ McGinty, 1946. The Nautilus, 59: 82, pl. 8, figs. 4-4a. [Type 
locality: Off Palm Beach, Florida]. 


Type material: Holotype in ANSP (181371). Figured by MOORE (1964, fig. 25). Not examined. 

Material examined: Florida, USA: 1 s, 60 m, off Sombrero Light, Key Vaca, Monroe Co. (CHL); 1 s, 
dredged, 30 m, 32 mi E St. Augustine, Dt. Johns Co. (CHL); 1 s, 26 m, Tanzler's Waters, 23 mi ENE 
Mayport, Duval Co. (CHL); 1 s, 27.75 mi ESE Mayport, Duval Co. (CHL). St. Kitts € Nevis: 1 s, 18 
m, base of the reef, Monkey Shoals (CHL). Guadeloupe: 1 s, lle d'Aves, Saint Martin Island, 8 m 


(EJE)! 


Description: Shell (Figs. 50A-B) 
depressed, spire flat, periphery flat, 
tricarinate, but central carina weak, 
umbilicus funnel shaped with strong 
delimiting carina. Protoconch (Fig. 
50C) of about 1 % -2 whorls; smooth 
and glassy, about 250 um in diameter. 
The end of the protoconch not clearly 
marked lacking a varix. Sculpture 
formed by a tricarinate periphery. 
There is a weak spiral rib on the 
middle of the dorsal side and a low 
cord at the suture. The base is smooth 
except for weak axial costulation. 
Umbilicus funnel shaped with flat 
walls, bordered by a strong descend- 
ing carina. Aperture strongly modi- 
fied by the carina and peripheral 
keels. The parietal callus is rather 
heavy and extends a little in front of 
the aperture. 

Dimensions: The holotype size is 
2.3 mm. The studied shells average 
1.52 mm in maximum dimension 
(diameter). 

Habitat: The bathymetric range of 
this species is ample, having been 
recorded between O and 128 m in 
depth; but the records for living mate- 
rial are between 42 and 91 m. 


Distribution: It has been recorded 
from USA, off Palm Beach, Florida 
(PILSBRY € MCGINTY, 1946); Colón and 
Bocas Island, Panama (OLsson é 
MCGINTY, 1958); Florida Peninsula, 
from Palm Beach to St. Petersburg 
(MOORE, 1964); North Carolina 
(PORTER, 1974); South half of Florida to 
Texas, Costa Rica and Panama (ABBOTT, 
1974); Portete, Costa Rica (HOUBRICK, 
1968; ROBINSON é MONTOYA, 1987); 
Louisiana and Texas (ODÉ, 1988); South 
Florida to Texas, Costa Rica and 
Caribbean Panama (Lyons, 1989); 
Mexico: Veracruz and Campeche State 
(REGUERO é GARCÍA-CUBAS, 1991); 
South Florida and from Texas to 
Panama and Colombia (Díaz MERLANO 
éz PUYANA HEGEDUS, 1994); northeast 
Florida (LEE, 2009). 

Remarks: The flat spire, the flat- 
tened periphery and the strong, 
descending umbilical carina are the 
important characters of the species. C. 
jeannae has a much larger umbilicus 
with a stronger bordering carina than 
C. suppressus (Dall, 1889). A similar 
species is C. bartschi (Mansfield, 1930), 
but it lacks the weak central keel on the 
periphery. 


Cyclostremiscus suppressus (Dall, 1889) (Figures 51A-H) 


Teinostoma (Ethalia) suppressa Dall, 1889a. Bulletin of the Museum of Comparative Zoology 18: 
1-492, pls. 10-40. [Type locality: Goodland Point, West Florida]. 


Circulus suppressus (Dall, 1889). 


Material examined: USA, Florida: 3 s, Anclote Key, Tarpon Springs (CHL); 1 sp, Sunset Park, Tarpon 
Springs, Pinellas Co. (CMK); 2 s, NE end Sand Key, Little Pass, Pinellas Co. (CHL); 1 s, Seaquar- 
ium Flats, Virginia Key, Dade Co. (CHL); 1 s, dredged 1.5-3 m, 5 mi SW Cedar Key, Levy Co. (CHL). 


Louisiana: 2 s, Last Island, Terra bone (CEG). 


97 


Iberus, 29 (2), 2011 


Figures SOA-D. Cyclostremiscus jeannae Pilsbry 8 McGinty, 1946. A-B; shells, 1.5 mm, San 
Martin Island (CJP); C: shell, 1.7 mm, Sombrero Light, Key Vaca, Florida (CHL); D: protoconch. 
Figuras SOA-D. Cyclostremiscus jeannae Pilsbry Y McGinty, 1946. A-B; conchas, 1,5 mm, San 
Martin Island (CJP); C: concha, 1.7 mm, Sombrero Light, Key Vaca, Florida (CHL); D: protoconcha. 


Description: Shell (Figs. 51A-D) 
depressed, spire flat, periphery strongly 
tricarinate, additional spiral carina close 
to suture, umbilicus minute, con- 
stricted, carinate. Protoconch (Fig. 51E) 
with 2 whorls, smooth and flat, and a 
diameter of about 390 um. Teleoconch 
formed by about 2 whorls. Spiral sculp- 
ture consists of 3 narrow sharp keels on 
the periphery, the middle one a little 
more prominent than the others. Near 
the suture there is another less promi- 
nent keel. Base flattened, bordered by 
the lower peripheral carina on the 
outher edge, and by another carina 
around the  umbilicus. Aperture 
oblique, parietal callus wide and with a 
sulcus on the upper inner angle. 
Umbilicus narrowed by the thickened 
inner wall. 

Dimensions: Maximum reported 
size: 2.4 mm. The figured specimens 
measure 1.5 and 1.8 mm in maximum 
diameter. 

Habitat: Depth: O to 58 m (live 0.6 
m). MOORE (1964) reported that this 


98 


species lives in shallow water strictly 
and is restricted to continental shores. 

Distribution: USA: North Carolina, 
Florida: East Florida, West Florida 
(DAEL, 18892. OANSONAA U9OSd) 
Louisiana, Texas; Mexico: Campeche 
State, Yucatan State. Recorded from 
Goodland Point; Hemphill and Baker's 
Haulover, Biscayne Bay, Florida 
(PiLSsBRY éz MCGINTY, 1946) and from 
the lower east coast and entire west 
coast of Florida and the coast of Texas 
(MOORE, 1964; EMERSON é JACOBSON 
(1976); from North Carolina (PORTER, 
1974); from Louisiana and Texas (USA) 
and Campeche State and Yucatan State, 
Mexico (ODÉ, 1988). 

Remarks: The flat spire, the three 
strong and sharp carinae, and the 
reduced umbilicus are the identifying 
characters of this species. 

The "shells+trom-*" Last Island; 
Terrabone, Louisiana, are apparently 
more depressed, present two strong 
cords on the dorsum, one of them near 
the suture, and the umbilicus is a little 


RUBIO ET ALz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 51A-H. Cyclostremiscus suppressus (Dall, 1889). A-D: shells, 1.5, 1.8 mm, Pinellas Co., 
Florida (CMK); E: protoconch. F-H: shells, 2.2, 2.1 mm, Last Island, Terrabone, Louisiana 
(CEG). 

Figuras 51A-H. Cyclostremiscus suppressus (Dall, 1889). A-D: conchas, 1,5, 1,8 mm, Pinellas Co., 
Florida (CMK); E: protoconcha. F-H: conchas, 2,2, 2,1 mm, Last Island, Terrabone, Louisiana 


(CEG) 


Iberus, 29 (2), 2011 


more closed than in shells from Florida. 
This could represent the morphological 
variability of the species. 

The closest species is Cyclostremis- 
cus jeannae, which is distinguished by 


the lack of pronounced peripheral keels 
and the presence of a wide umbilicus. 

Cyclostremiscus colombianus Pilsbry 
$ Olsson, 1845 is morphogically 
similar. 


Cyclostremiscus vanbruggeni de Jong € Coomans, 1988 (Figures 52A-E) 


Cyclostremiscus vanbruggen1 de Jong € Coomans, 1988. Studies on the Fauna of Curacao and 
other Caribbean Islands, 69: 32, pl. 2, fig. 136. [Type locality: Curacao / Arubal. 


Type material: Holotype in ZMA (3.87.063). Not examined. 
Material examined: Brazil: 2 s, Praia da Itararé, Sáo Vicente, Sáo Paulo (CHL). Trinidad and Tobago: 


Tobago, 1 s, Scarborough (CHL). 


Description: The original description 
is as follows: “Shell wider than high. 
The sculpture consists of fine spiral 
striae and low radial ribs which on the 
upper side of the last whorl are weak or 
nearly absent. On the periphery at the 
beginning of the last whorl there are 3 
smooth keels of which the middle one 
soon disappears and gradually also the 
upper one, so that near the aperture 
practically only the lower keel remains. 
On the upper side there is a ridge which 
eradually fades away. The very wide 
umbilicus is bordered by a rid ge”. 

Maximum reported size: 1.6 mm 


Habitat: Nothing is mentioned by DE 
JONG € COOMANS (1988) about the 
habitat of this species. The shell pho- 
tographed was collected in sandy sedi- 
ments. 

Distribution: ABC Islands: Aruba, 
Curacao (DE JONG é££ COOMANS, 1988). 

Remarks: The closest species concho- 
logically is Cyclostremiscus jeannae, 
from which it can be distinguished by 
the weaker dorsal ornamentation and 
the smaller umbilicus. Cyclostremiscus 
suppressus may be distinguished by its 
lack of axial sculpture except very fine 
growth lines. 


Cyclostremiscus diminutus spec. nov. Rubio, Rolán € Pelorce (Figures 53A-G) 


Type material: Holotype (Figs. 53A-D) in MNHN (24201) (ex CJP); one paratype from Pointe-á- 
Pitre, Guadeloupe (FLMNH, 448611). Other paratypes: USNM (1155030, 1 s, Fig. 53D), AMNH (1 


s, Fig. 53E), from Bocas Island, Panama (ex CHL); CHL (1 s, Puerto Rico). 


Type locality: Guadeloupe. 


Description: Shell (Figs. 53A-C) 
solid, depressed, lenticular, with a 
short spire. Protoconch (Fig. 53D) 
smooth, bulbous, about 290 um in 
diameter, with 1 Y whorls. Teleoconch 
with a little more than one whorl. 
Ornamentation produced by 5 spiral 
cords which are distributed regularly 
on the periphery and which are 
marked by axial growth striae, also 
evident near the umbilicus. At the 
beginning of the teleoconch a spiral 
cord may be seen on the dorsum but it 
promptly fades. Aperture rounded, 


100 


columella and inner lip reflected 
outward. Umbilicus wide and deep, 
spiral cords absent from its border and 
inner aspect. 

Holotype is 0.75 mm in maximum 
diameter and 0.32 mm in height. 

Habitat: No information. All the 
material was collected in drift sedi- 
ments. 

Distribution: Only known from the 
type material: Guadeloupe, Puerto Rico 
and Panama. 

Remarks: Cyclostremiscus spec. nov. 
may be distingished from the other 


RUBIO £7 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 52A-E. Cyclostremiscus vanbruggeni de Jong 82 Coomans, 1988. A-B: shells, 1.43, 1.28, 
Praia da Itararé, Sáo Vicente, Sao Paulo (CHL); C: shell, 1.7 mm, Searborough, Tobago (CHL); 
D: detail of the sculpture; E: protoconch. 

Figuras 52A-E. Cyclostremiscus vanbruggeni de Jong Y Coomans, 1988. A-B: conchas, 1,43, 1,28, 
Praia da ltararé, Sáo Vicente, Sáo Paulo (CHL); C: concha, 1,7 mm, Searborough, Tobago (CHL); D: 
detalle de la escultura; E: protoconcha. 


species of the complex bartschi-jeannae- periphery 5 spiral cords not particularly 
vanbruggeni-suppresus, by the lack of prominent, delimiting a convex periph- 
any axial sculpture, having at the ery. 


101 


Iberus, 29 (2), 2011 


Figures 53A-G. Cyclostremiscus diminutus spec. nov. Rubio, Rolán 82 Pelorce. A-C: holotype, 0.75 
mm, Guadeloupe (MNHN); D-E: paratypes, 0.95, 0.87 mm, Bocas Island, Panama (USNM and 
AMNH, ex CHL)); F: paratype, 0,92 mm, Puerto Rico (CHL); G: protoconch. 

Figuras 53A-G. Cyclostremiscus diminutus spec. nov. Rubio, Rolán «* Pelorce. A-C: holotipo, 0,75 
mm, Guadeloupe (MNAN); D-E: paratipos, 0,95, 0,87 mm, Isla Bocas, Panamá (USNM and 
AMNH, ex CHL)); F: paratipo, 0,92 mm, Puerto Rico (CHL); G: protoconcha. 


102 


RUBIO ET AZ.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Cyclostremiscus cubanus (Pilsbry £ Aguayo, 1933) (Figures 54A-G) 


Circulus cubanus Pilsbry €: Aguayo, 1933. The Nautilus, 46: 120. pl. 6 figs. 6-6b. [Type locality: 
Varadero, Matanzas, Cuba]. 

Cyclostrema cubanus (Pilsbry £ Aguayo, 1933). 

Cyclostremiscus puntagordensis Weisbord, 1962: 141-142, pl. 13, figs. 10-12. [Type locality: Playa 
Grande Formation (Maiquetía member) at W-23, north flank of Punta Gorda anticline, 
Venezuela]. Fossil record. 


Type material: Holotype in ANSP (160198). Represented in MOORE (1964: 230, fig. 26). 

Other material examined: Belize: 2 s, 18 m, Deadman's Reef, Turneffe Island (CHL). Panama: 1 s, 
1 mi N Punta San Blas, San Blas Islands (CHL). Honduras: 3 s, Calabash, Roatan Island (CHL). 
Puerto Rico: 2 s, Holiday Inn, San Juan (CHL). Cuba: 4 s, Varadero Beach, Matanzas (MCZ 109345, 
179783); 1 s, Archipelago de los Canarreos, 5 m (CFR); 1 s, Cayo Avalos, 8 m (MHNS); 1 s, Cayo 
Diego Perez, beached (MHNS); 1 s, Faro Perez Diego, 10 m (MHNS); 2 s, Jibacoa, 3-6 m (MHNS); 
5 s, Rancho Luna Beach, 10 m (MHNS); 15 s, Rancho Luna Beach, 20 m (MHNS); 4 s, Rancho Luna 
Beach, 45 m. Florida, USA: 1 s, 30 m, 32 mi E St. Augustine, St. Johns (CHL). Bahamas: 1 s, 15 m, 
French Bay, San Salvador (CHL); 4 s, 36 m, Mt Olympus Reef, 12 mi NNW Great Bahama Is (CHL); 
4 s, 15 m, Samphire Cay, near Nassau (CHL); 21 s, 18 m, South Ridring Rocks, Cay Sal Bank (CHL); 
1 s, 25 m, Marina Long Cay (CHL). Saint Kitts € Nevis: 6 s, 18 m, Monkey Shoals (CHL). Mar- 
tinique: 1 s, coralline reefs of Point Borgnesse, 12 m, sand-muddy bottom (CJP). Trinidad and 
Tobago: Tobago, 16 s, Horse Shoe Reef, 15 m, shells grit bottom (CJP). Aruba: 1 s, 15 m, off Palm 
Beach (CHL). Bonaire: 1 s, 39 m, off Klein Bonaire (CHL). Virgin Islands: 5 s, 18 m, Deadman Chest 


(CHL); 5 s, Magens Bay, N coast of St. Thomas (CHL); 1 s, Virgin Gorda (CHL). 


Description: Shell (Figs. 54A-C) dep- 
resed, whitish, robust, with spiral sculp- 
ture. Protoconch (Fig. 54D) of almost 1 Y 
whorls, about 270 um in diameter, and 
with two clearly differentiated stages; 
numerous strong irregular tubercles are 
seen on its surface. There is no varix 
evident at the transition between proto- 
conch and teleoconch. The latter has 
about 1 Y rapidly-expanding whorls. 
Ornamentation formed by 6 strong spiral 
cords: 3 on the periphery, keel-like, 
having a slightly oblique profile; dorsally 
one spiral cord extends between the 
suture and the upper peripheral cord. On 
the base there are two spiral cords: one 
around the umbilicus and the other 
between this and the basal peripheral 
cord. On the spiral cords some thickening 
can be seen at the points where the axial 
growth striae cross. The entire surface of 
the shell is covered by a microsculpture 
(Figs. 54E-G) of fine spiral anastomosing, 
threads. Aperture quadrangular, thick- 
ened; outer lip with very fine dentition on 
its inner margin. Columella and internal 
lip reflected towards the umbilicus. Pari- 
etal callus complete. Umbilicus wide and 
deep. Dimensions: The holotype is 1 mm 
in maximum diameter. 


In our material there are some larger 
shells (1.20 mm in maximum diameter). 

Animal and radula unknown. 

Habitat: The bathymetric range of this 
species is large, having been recorded 
between 0 and 140 m in depth. There is no 
record of live-collected material, so its true 
habitat is unknown. 

Distribution: This species has a wide 
Caribbean distribution. It has been 
recorded from Varadero, Cuba (PILSBRY € 
AGUAYO, 1933); from Florida, Cuba and the 
Virgin Islands (MOORE, 1964); from Florida, 
Garden Banks, Texas (ODÉ, 1988); from 
Florida, St. Augustine (LEE, 2009); from 
Abaco, Bahamas (REDFERN, 2001); and from 
Panama (OLsson é£ MCGINTY, 1958). More 
localities are added in the present work. 

Remarks: We have pointed out some 
shell features which were not mentioned in 
the original description or considered erro- 
neous in the original or subsequent descrip- 
tions. With respect to the protoconch, 
PILSBRY €: AGUAYO (1933) mentioned only 
that it is smooth and small; the dimensions 
of the holotype are erroneus because it was 
reported to be 1 mm in height and 0.45 mm 
in diameter. MOORE (1964) says that the 
protoconch has 2 Y whorls when really it 
has only 1 %; he also comments that the 


103 


Iberus, 29 (2), 2011 


Figures 54A-G. Cyclostremiscus cubanus (Pilsbry 8 Aguayo, 1933). A-C: shells, 1.1, 1.2, 1.1 mm, 
Rancho Luna Beach, Cienfuegos, Cuba (MHNS); J-K: protoconch; E-G: details of the 
microsculpture. 

Figuras 54A-G. Cyclostremiscus cubanus (Pilsbry € Aguayo, 1933). A-C: conchas, 1,1, 1,2, L,1 mm, 
Playa Rancho Luna, Cienfuegos, Cuba (MHNS); J-K: protoconcha; E-G: detalles de la microescultura. 


microsculpture covers all the shell, but it is as a fossil from the Plio-Pleistocene of 
absent from the umbilicus. Venezuela by WEISBORD (1962), but the only 

C. puntagordensis, a species placed in shell known, the hololotype, has slighty 
synonymy by MOORE (1964) was described larger dimensions (1.4 mm). 


104 


RUBIO £7 4L£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Cyclostremiscus euglyptus Aguayo € Borro, 1946 (Figures 55A-D) 


Cyclostremiscus euglyptus Aguayo € Borro, 1946. Rev. Sdad. Malac. “Carlos de la Torre,” 4(1): 
9-10, figs. 4-6. [Type locality: Matanzas, Cuba]. 


Type material: Holotype represented in AGUAYO éz BORRO (1946, figs. 4-6), in MPH (11877). Described 
as a fossil of the Upper Tertiary found in the formation “Yumurí,” Upper Miocene of Cuba. 

Other material examined: Cuba: 1 s, Cayo Avalos, 8 m (MHNS); 1 s, Cayo Perez Diego, 5 m(MHNS); 
1 s, Jibacoa, 3-6 m (MHNS); 16 s, Cienfuegos Bay, 20-30 m (CFG); 2 s, Rancho Luna Beach, 12 m 
(CFG); 47 s, Rancho Luna Beach, 20 m (MHNS); 1 s, Rancho Luna Beach, 35 m (CFG); 14 s, Rancho 
Luna Beach, 45 m (CFG); 5 s, Faro los Colorados, 56 m (CFG); 2 s, Punta Tamarindo, 25 m (CFG). 


Description: Shell (Figs. 55A-C). The 
original description of AGUAYO éz 
BORRO (1946: 9-10) is as follows: 
“Concha pequeña, discoidal, de espira 
muy poco elevada, ampliamente umbili- 
cada. Con 1 4 vueltas embrionarias lisas 
y 2 Y postnucleares fuertemente escul- 
pidas. Escultura espiral formada por 
dos quillas periféricas que limitan el 
contorno de la última vuelta y otra basal 
que bordea el ombligo. Finas líneas espi- 
rales se observan entre las expresadas 
quillas. Escultura axial formada por 
unas 40 costillas radiales en la última 
vuelta, que forman nódulos o incisiones 
al cruzar los cordones espirales. Perife- 
ria de la última vuelta con los lados casi 
paralelos y formando ángulos marcados 
con el eje de la concha. Ombligo amplio, 
dejando ver las vueltas nucleares. Pared 
del ombligo con escultura radial, pero 
sin líneas espirales. Abertura cuadran- 
gular, con el lado columelar muy arque- 
ado; el resto poligonal por la articula- 
ción de las quillas espirales”. 


Protoconch (Fig. 55D) with a diame- 
ter of about 190 um, smooth except at 
the earliest, embryonic, part where irreg- 
ular short lines can be seen. The separa- 
tion from the teleoconch is very evident. 

Dimensions: the holotype is 1.05 mm 
in diameter and 0.56 mm in height. The 
single paratype, from the same lot, mea- 
sured 1 mm in diameter and 0.60 mm in 
height. The dimensions of our material 
are similar to those of the holotype. 

Habitat: This species is relatively 
common in Cuba. Its bathymetric distri- 
bution is between 5 and 60 m; most of 
the shells of our material were collected 
between 20 and 56 m. 

Distribution: Only known from 
Cuba. 

Remarks: There is no other species in 
the genus with this shell morphology, 
thus its identification is easy. Being a 
species probably “endemic” to the 
island of Cuba, it has been overlooked 
in published works and electronic data- 
bases. 


Cyclostremiscus microstriatus spec. nov. Rubio, Rolán € Lee (Figures 56A-H) 
Vitrinorbis sp. Lee, 2009: 69, n* 333. 


Type material: Holotype (Fig. 56A) in MNCN (15.05/55055). Paratypes in the following, collections: 
MNHN (24395, 1 s, Fig. 56B), MHNS (100549, 1 s, Fig. 56C) from the type locality (all ex CFG). Other 
paratypes: FLMNH (448609, 1 s, Fig. 48D, trawled 30 m, 35 mi E Mayport, Duval Co., Fla.; 3 s, trawled 
30 m, 35 mi E St. Augustine, St. Johns Co., Fla. (Fig. 56E) (one in USNM 1155029, other in ANSP and 
a third in CHL). 

Type locality: Rancho Luna Beach, Cienfuegos, Cuba. 

Etymology: The specific name refers to the minute striation in the interspaces between cords. 


Description: Shell (Figs. 56A-E) of (Fig. 56F) with a fine granulation barely 


very small size, planispiral, robust, 
whitish, solid, not shiny. Protoconch 


observable; 1 14 whorls and about 240 
um in diameter; there is no thick rib at 


105 


Iberus, 29 (2), 2011 


———— 50 um 


Figures S5A-D. Cyclostremiscus euglyptus Aguayo 82 Borro, 1946. A-C: shells, 1.0, 0.95, 0.8 mm, 


Cienfuegos, Cuba; D: protoconch. 


Figures 55A-D. Cyclostremiscus euglyptus Aguayo d* Borro, 1946. A-C: conchas, 1,0, 0,95, 0,8 mm, 


Cienfuegos, Cuba; D: protoconcha. 


the junction with the teleoconch. 
Teleoconch with about 1 % whorls. The 
entire shell is covered by wide spiral 
cords of almost equal size. The inter- 
spaces between cords are covered by 
fine axial incisions (Figs. 56G-H). On 
the last whorl there are between 20 
and 23 cords; 7-8 on the dorsum, 8 on 
the periphery and 6-8 on the base. On 
the base, there is a space without cords 
from the middle of the last whorl to 
the umbilical border. Umbilicus 
smaller than those in congeneric 
species, the previous whorls can be 
seen on its interior. Aperture proso- 
cline, rounded, thickened, its external 
margin forms a characterístic point in 
the area where the 7th-9th cords, the 
three most prominent ones, are placed. 
The columella as well as the internal 
lip and the upper part of the external 
lip are reflected outward. The first 
peripheral cord, as well as the dorsum 
and the last peripheral cord near the 


106 


base are visible from above, the base 
being sligtly concave. 

Dimensions: The holotype is 1.3 mm 
in maximum diameter and 0.40 mm in 
height. 

Animal and radula unknown. 

Distribution: Known only from 
Cienfuegos, Cuba and Florida, USA. 

Discussion: Cyclostremiscus micros- 
triatus spec. nov., is a very characteristic 
species; its form is almost planispiral, its 
ornamentation is formed by subequal 
spiral cords and very fine axial incisions 
in the interspaces, as well as the prolon- 
gation which forms the border of the 
external lip. 

A similar species was recorded by 
Lyons (1989) as Vitrinorbis sp. It is 
much more flattened in profile and is 
markedly angulate at the level of the 
lowest peripheral keel. LEE (2009) sug- 
gested that Circulus gunteri (Mansfield, 
1930), described from the Miocene of 
Florida, was ancestral to Cyclostremis- 


RUBIO ET 4L.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 56A-H. Cyclostremiscus microstriatus spec. nov. Rubio, Rolán 82 Lee. A: holotype, 1.3 mm 
(MNCN); B: paratype, 1.1 mm (MNHN); C: paratype, 1.0 mm (MHNS), Rancho Luna Beach, 
Cienfuegos, Cuba; D: paratype, 1.3, Duval Co., Florida (FLMNH); E: paratype, 1.2 mm, E St. 
Angustin, St. Johns Co., Florida (USNM); EF: protoconch; G-H: microsculpture. 

Figures 56A-H. Cyclostremiscus microstriatus spec. nov. Rubio, Rolán e Lee. A: holotipo, 1,3 mm 
(MNCN); B: paratipo, 1,1 mm (MNAN); C: paratipo, 1,0 mm (MANS), Playa Rancho Luna, Cien- 


fuegos, Cuba; D: paratipo, 1,3, Duval Co., Florida (FLMNAH); E: paratipo, 1,2 mm, E St. Angustin, 
St. Johns Co., Florida (USNM); E: protoconcha; G-H: microescultura. 


107 


Iberus, 29 (2), 2011 


cus microstriatus spec. nov. (his Vitri- 
norbis species [no. 333]); the fossil, 
while having the same flat spire, differs 
in having the body whorl totally 
covered by cords and lacking the labral 
projection. 

Circulus quadricristatus (Aguayo, 
1949) is also similar to C. microstriatus 


spec. nov., but it only presents 4 wide 
cords on the periphery and lacks the 
labral projection. 

LeE (2009) also suggested that the 
Panamic Cyclostremiscus salvatierrensis 
Hertz, Myers € Gemmill, 1992 was 
cognate with C. microstriatus spec. nov. 
(his Vitrinorbis species [no. 333]). 


Genus Cochliolep1s Stimpson, 1858 


Cochliolepis Stimpson, 1858. Proc. Boston Soc. Nat. Hist., VI: 308. 
Type species: Cochliolepis parasiticus [sic] Stimpson, 1858, by monotypy. Recent, Caribbean. 


Diagnosis: MOORE (1964: 168) 
“Thin-shelled, strongly depressed 
and openly umbilicate. Apex flat or 
sunken, surface smooth except for 
growth lines or occasionally spiral 
striations. Aperture obligue, peris- 
tome very briefly in contact with 
preceding whorl. Adult whorls 
rapidly increase 11 size. sojt plants 
described as bright red, with two 
long pallial tentacles, and with a 
supplementary gill projecting from 
the right side of the aperture. Oper- 
culum thin and flexible”. 

C. parasitica was collected alive 
below the “elytra” of the giant worm 
Polidontes lupinus. C. parasitica is a 
commensal and not a parasite. lt 
apparently feeds on the fine particu- 
late organic material drawn into the 
tube by the worm's respiratory and 
feeding current. It possibly performs 
a cleaning function. 

Remarks: MOORE (1964) com- 
mented on the distribution of the 
genus: “the genus appears to be con- 
fined to the West Indian region. C. 
parasitica is found on the Carolina 


coast, C. nautiliformis and C. striata 
live on the Gulf of Mexico and C. 
adamsii 1s known from Guade- 
¡Oupese 

However, we have found a very 
different situation: First at all, we 
have tried to make the specific dif- 
ferentiation basing not only on the 
characters of the teleoconch but also 
the protoconch. Fortunately we were 
able to compare shells from very dif- 
ferent Caribbean localities confirm- 
ing the morphology of the proto- 
conch of Cochliolepis parasitica 
shown for the first time by REDFERN 
(2001). The sculpture is very charac- 
teristic, and it allows the distinction 
from congeneric species even as 
juveniles. 

As for Cochliolepis nautiliformis 
(Holmes, 1859), in spite of the 
opinion of some authors who con- 
sider tasas yanonymtot e: 
parasitica, we could not confirm this 
due to the difficulty of observing the 
protoconch of that species; thus we 
prefer to present it provisionally as a 
separate taxon. 


Cochliolepis adamsii (P. Fischer, 1857) (Figures 57A-D) 


Adeorbis adamsii P. Fischer, 1857. Journal de Conchyliologie, 6: 287, pl. 10, fig. 11. [Type locality: 


Guadeloupel. 


Type material: In MNHN. Not examined. 


Other material examined: Bahamas: 1 s, beach near Current Cut, Current Eleuthera (CHL); 1 s, 36 
m, Olympus Reef, 12 mi NNW West End, Grand Bahama Island (CHL). Florida, USA: 1 s, 1 m, 


Peanut Island, Palm Beach Co. (CHL). 


108 


RUBIO ET ALz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Description: This is the original 
description of P. FISCHER (1857: 287): 
“Testa translucida, corneo-alba, nitida, 
complanata, suborbiculari, umbilicata; 
supra minutissimé et concentrice striata; 
subtus laevicula; anfractibus 3 Y rapide 
accrescentes, ultimo compressiusculo, 
non carinato; aperture subovata, parum 
transversa; margine dextro  tenul, 
fragil”. This description must be suple- 
mented with some information obtained 
from the materal presently studied: 

Shell (Figs. 57A-D) depressed, 
fragile, of whitish-tan color, comprised 
of 3 Y rapidly-expanding whorls. Proto- 
conch (Fig. 57C) about 390 um in diame- 
ter with a nucleus of about 160 um. Fol- 
lowing the nucleus a prominent spiral 
sculpture appears, being formed by four 
discrete cordlets which terminate at the 
beginning of the teleoconch. The teleo- 
conch has a fine spiral striation which is 
crossed by numerous distinct growth 
lines, even into the widely-open umbili- 
cus. Aperture suboval, with a fine and 
fragile border. No columellar or parietal 
callus. 

Dimensions: Holotype 6.5 mm in 
maximum diameter and 2.0 mm in 
height. 

Our shells are 7.1 and 6.2 mm in 
diameter. 


Animal and radula unknown. 

Habitat: This species, as well as its 
congeners, prefers shallow water in 
coastal bays. No bathymetric infor- 
mation was provided in the original 
description. Our shell came from a 
sandy and rocky bottom at 2 m deep. 

Distribution: Recorded from USA: 
Florida: Florida Keys (MaAurtY, 1922; 
MOORE, 1964) and Guadeloupe 
(FISCHER, 1857). 

Remarks: The species was previ- 
ously included in the genus 
Macromphalina. 

MOORE (1964) mentioned that 
during his stay in Paris in 1963, he 
examined the type of Adeorbis 
adamsii P. Fischer, 1857, concluding 
that “it is a Cochliolepis very similar 
to C. striata Dall, 1889, but more 
inflated”. 

The four species in the West 
Indian region may be separated into 
two groups: C. parasitica and C. nau- 
tiliformis are relatively small, smooth 
and shiny; C. striata and C. adamsii 
are relatively larger, spirally striated 
and have a light tan coloration. Con- 
trary to the observations of MOORE 
(1964: 174), the base presents the 
same ornamentation as the rest of the 


shell. 


Cochliolepis holmesii (Dall, 1889) (Figures 58A-F) 


Cochliolepis parasiticus [sic] Holmes, 1859. Post-Pleiocene Fossils of South Carolina: 93-94, pl. 
14, figs. 9, 9a, 9b. [Type locality: South Carolina, Post Pliocene] non Cochliolepis parasiticus 


[sic] Stimpson, 1858. 


Vitrinella holmesi Dall, 1889a. Bulletin of the Museum of Comparative Zoology 18: 360, 392. 


Type material: Deposited in MCZ. Not examined. 


Material examined: Florida, USA: 5 s, APAC Pit, Sarasota, Plio-Pleistocene (CHL). Mexico: 2 s, 


Puerto Morelos, Yucatan, 10-16 m (MHNS). 


Description: From PiLsBRY (1953: 
433-434): “The thin shell is biconvex; 
the periphery, below the middle of the 
whorl, is quite narrowly rounded, the 
whorl being more convex above than 
below it. The spire is nearly level but 
the apex is visible in a face view. The 
suture is deeply impressed, the whorls 
convex. The umbilicus is very broad 


and open, with a deep suture, the 
umbilical side of the whorls being 
strongly convex. The weak wrinkles of 
growth become stronger as they 
approach the lip, and are often rather 
fine and close on the base of the last 
third of the body-whorl. A microscopic, 
close, spiral striation covers the last 
whorl in good specimens, being weaker 


109 


Iberus, 29 (2), 2011 


Figures 57A-D. Cochliolepis adamsii (P. Fischer, 1857). A-B; shells, 7.1, 6.2 mm, Bahamas (CHL); 
C: protoconch; D: microsculpture. 
Figuras 57A-D. Cochliolepis adamsii (P Fischer, 1857). A-B; conchas, 7,1, 6,2 mm, Bahamas 
(CHL); C: protoconcha; D: microescultura. 


at the base. The aperture is oblique. 
Peristome is thin, the columellar margin 
concave below, then in a straight line to 
the insertion just below the periphery. It 


110 


is thin throughout. The parietal callus is 
very short and thin”. 


We add: The shells (Figs. 58A-C) 
examined have the protoconch (Figs. 


RUBIO ET 4L.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 58A-E. Cochliolepis holmesii (Dall, 1889). A-C: shells, 3.66, 3.0, 2.96 mm, Plio-Pleistocene 
of Sarasota, Florida (CHL); D: microsculpture; E-F: protoconchs. 

Figuras 58A-E Cochliolepis holmesii (Dal), 1889). A-C: conchas, 3,66, 3,0, 2,96 mm, Plio-Pleisto- 
ceno de Sarasota, Florida (CHL); D: microescultura; E-F: protoconchas. 


58E-F) bulbous, about 470 um, 1 % 
whorls, and with a very marked suture. 
Two stages may be easily distin- 
guished: the embryonic shell is % of 
whorl, measuring about 170 um in 


diameter and is covered by a fine gran- 
ulation, more evident in the areas close 
to the suture. The second stage has 5-6 
fine spiral lines and minute growth 
marks. 


Dal] 


Iberus, 29 (2), 2011 


The teleoconch is completely 
covered by growth marks and spiral 
striae; on the dorsum of the shell the 
collabral growth marks are more 
robust, especially at the later stages of 
growth, while on the base they are finer 
and denser on the last third of the 
whorl. Periphery subangular. 

Maximum reported size: 2.4 mm 

Habitat: Depth: 2 to 11 m. 

Distribution: USA: Florida: East 


Florida (Lyons, 1989); Mexico: 
Cozumel (MOORE, 1973). 
Remarks: The name  Vitrinella 


holmesii was introduced by Dall 
because Cochliolepis parasiticus [sic] 
Stimpson was misidentified by HOLMES 
(1859: 93-94, pl. 14, figs. 9, 9a, 9b); it 
was, in fact, a new species. 

The type of this species appears to 
be lost; it was described and figured by 


HOLMES (1859) but no topotypic speci- 
mens have been collected. Dall's notes 
(1892) mention a specimen in the 
USNM (114368), which he identified 
with the Post-Pliocene species of 
HOLMES (1859). DALL (1889a) used the 
spellins Holmes tonic Uan 
Holniestiitcon po 392 DA is 92) 
acting as First Reviser, employed the 
latter spelling, thus establishing the 
correct original spelling. 

C. holmesíii was described as a 
fossil of the Post-Pleiocene of South 
Carolina (HoLMEs, 1859; DALL, 1889a); 
from the Miocene of Duplin County, 
North Carolina (DaALL, 1892); from the 
St. Petersburg Pliocene, the Pliocene in 
the Waccamaw formation of North Car- 
olina, and the Miocene of the Natural 
Well, Duplin Co., North Carolina 
(PILSBRY, 1953). 


Cochliolepis nautiliformis (Holmes, 1859) (Figures 59A-C) 


Adeorbis nautiliformis Holmes, 1859: 93, pl. 14, figs. 8-8b. [Type locality: Cainhoy, Wando River, 


South Carolina]. 


Type material: Unknown. 


Other material examined: Florida, USA: 1 sp, Tarpon Springs, Pinellas Co., S end of the parking lot 
Howard Park, (CHL); 1 s, Gulfport, Pinellas Co. (CHL). 


Remarks: In discussing figured speci- 
mens of Cochliolepis  nautiliformis 
(Holmes) from the St. Petersburg Plio- 
Pleistocene, PILSBRY (1953) in OLSSON éz 
HARBISON (1953) commented: “DALL 
(1889: 360) stated that A. nautiliformis 
Holmes is identical with C. parasitica 
Stimpson, and this synonymy has been 
accepted by some later authors without 
verification. There are several specific dif- 
ferences, the most conspicuous being the 


much more extensive envelopment of the 
spire in C. nautiliformis, reducing the 
visible spiral, the summit being left as a 
slight depression. The apical whorl is 
wider and fully exposed in C. parasitica”. 
We have insufficient material to for- 
mulate an opinion about this taxon, there- 
fore we present figures of what we per- 
ceive as this species. The problem can 
only be resolved with more material, 
including live-collected specimens. 


Cochliolepis parasitica Stimpson, 1858 (Figures 60A-G, 61A-H, 62A-F) 


Cochliolepis parasiticus [sic] Stimpson, 1858: 307-309, text-fig. [Type locality: Charleston Harbor, 


South Carolina]. 


Nemafera compressa Kurtz, 1960: 8. Not figured. [Type locality: Fort Johnson, South Carolina]. 


Type material: Two syntypes in USNM (95079) (corresponding to two specimens deposited with 
n” 87142, given by Stimpson to Isaac Lea). Examined in photographs (Figures 60A-G). The larger 


one is here designated the lectotype. 


112 


RUBIO ET 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 59A-C. Cochliolepis nautiliformis (Holmes, 1859). A-B: shell, 5.0 mm, Gulfport, Pinellas 
Co., Florida (CHL); C: protoconch. 

Figuras 59A-C. Cochliolepis nautiliformis (Holmes, 1859). A-B: concha, 5,0 mm, Gulfport, Pinellas 
Co., Florida (CHL); C: protoconcha. 


Other material examined: Bermuda: about 300 s, Gibbons Bay (CHL). Bahamas: 1 s, Channel, Chub 
Cay (CHL); 1 s, W end Grand Bahama (CHL); 3 s, beach, 300 m N Current Cut, Eleuthera (CHL); 
2 s, 15 m, NW Nassau (CHL); 85 s and some j, from Abaco, 10-23 m (CCR). Florida, USA: 1 s, 36- 
90 m, off Dry Tortugas (CHL); 2 s, beach, 500 m W Flamingo Visitors Center, Monroe Co (CHL). 
Mexico: 3 s, Puerto Morelos, Yucatan, 8-18 m (MHNS). Guadeloupe: 1 s, north Olive Bay, 2 m, 
among rocks (CJP). Cuba: 5 s, Maria la Gorda, 20 m (MHNS) 3 s, Playa Girón, 2 m (MHNS) 4 s, 
Cayo Carenas, Cienfuegos Bay, 5 m. Turks éz Caicos: 13 s, 18 m, French Cay (CHL). Puerto Rico: 1 
s, beach, Holiday Inn, San Juan (CHL). St. Kitts and Nevis: 1 s, 14 m, The Garden, Nevis (CHL). 
Aruba: 1 s, 15 m, off Palm Beach (CHL). Grand Cayman Island: 1 s, beach grit, 100 m S. Rum Pt., 
Cayman Kai (CHL). 


Description: Shell (Figs. 60A-D, 614, 
62A-C): See STIMPSON (1858). The best 
description is in MOORE (1972: 103): 
“The smooth glassy shell is strongly 
depressed, flattened on top, and broad ly 
umbilicate. The protoconch is slightly 
immersed in the teleoconch and consists 
of about 1 % whorls. The teleoconch 
consists of slightly more than two 


whorls in a large specimen 4.02 mm in 
diameter. The first adult whorl has a 
thin callus of shell up to, or sometimes 
covering part of, the protoconch; the 
suture is carried out toward the periph- 
ery on the last half whorl. The suture 
thus follows a nearly crescent-shaped 
curve from the protoconch to the inner 
corner of the aperture. There is a thin 


LUS 


Iberus, 29 (2), 2011 


Figures 60A-G. Cochliolepis parasitica Stimpson, 1858. A-B: lectotype, 3.4 mm (USNM 87142); 
C-D: shell of the same lot, 3.4 mm (USNM); E: detail of the operculum; F-G: protoconch of the 
both specimens. (SEMs by Yolanda Villacampa, USNM). 
Figuras G60A-G. Cochliolepis parasitica Stimpson, 1858. A-B: lectotipo, 3,4 mm (USNM 87142); C- 
D: concha del mismo lote, 3,4 mm (USNM); E: detalle del opérculo; E-G: protoconcha de ambos ejem- 
plares. (fotografías al MEB de Yolanda Villacampa, USNM). 


114 


RUBIO ET 4L.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


callus on the parietal wall, and the col- 
umella is thickened where it joins the 
precedieron ines taplerture1s 
oblique, for the upper part the peris- 
tome is extended well beyond the 
remainder of the outer lip. The aper- 
ture is wider than high, with a ratio of 
about atole Outer lp 1s EIN, 
smooth, and with no thickening to 
indicate maturity. Most adult speci- 
mens have one or more strong growth 
lines, which give a nautiloid appear- 
ance mole shell The umbilicus ts 
widely open and penetrates to the pro- 
toconch. Height-to-breadth ratio for the 
teleoconch 1s about 1.0 to 3.5. A speci- 
men 4.02 mm in diameter was 1.23 mm 
high. Maximun diameter is about 4.5 

Curiously, until the work of 
REDFERN (2001) the protoconch of 
Cochliolepis parasitica had never been 
described, probably due to the destruc- 
tion of the holotype and because puta- 
tive type material now in USNM 
(labelled “syntype”, possible 
paratypes), though juveniles, have the 
protoconch eroded or concealed in 
callus. In our opinion the description of 
the protoconch of Cochliolepis parasit- 
ica is very important, because it is nec- 
essary for the correct differentiation 
from otherwise quite similar species in 
the genus. 

According REDFERN'S (2001) 
description: “The protoconch of adult 
shells is usually abraded, showing only 
traces of the distinctive sculpture seen 
on a larval shell. The embryonic whorl 
is smooth and translucent, but the sub- 
sequent whorl is more opaque, with an 
irregular pattern of raised zigzag spiral 
imeads o Eachioj ese produces 
random  off-shoots that project 
obliquely towards the adjacent thread, 
and this sculpture tends to become 
more regular towards the periphery”. 

We were able to compare the proto- 
conchs from the Bahamas with those 
from Bermuda and Turks € Caicos, and 
we conclude that, despite small differ- 
ences among them, all fall within 
Redfern's description (Figures 60F-G, 
61B-H, 62D-F). 


Maximum reported size: 4.2 mm. 
Lectotype in USNM 3.4 mm. The other 
specimen in the lot is of a similar size. 

Size of a specimen from Playa Giron, 
Cuba: shell 3.05 mm diameter; proto- 
conch 570 um in diameter. 

Size specimen from Puerto Morelos, 
Yucatan: shell 2.63 mm diameter; proto- 
conch 463 um in diameter. 

The animal has a pair of cephalic 
tentacles, two more pallial tentacles, 
which project from the upper angle of 
the aperture, a medium-sized slipper- 
like foot, a multispiral operculum, a 
large and well developed ctenidium and 
an elongate, narrow and curved penis in 
the male (MOORE, 1972). 

Habitat: Depth: 0 to 48 m (living at 
0.3 to 0.6 m). MOORE (1972) concludes 
that C. parasitica is not a parasite of the 
annelid Acoetes lupina Stimpson, but a 
herbivore which lives as a dependent 
symbiont. Since the snails are often 
found on the worm itself, they possibly 
perform a cleaning function. 

Distribution: Cochliolepis parasitica 
Stimpson was originally found in the 
harbor of Charleston, South Carolina, 
living under scales of the annelid 
Acoetes lupina. lt was reported by 
HOLMES (1860) from the Post-Pliocene at 
Simmons Place, Wando River, South 
Carolina, but this was an erroneous 
identification; the species figured by 
him is not that of Stimpson, and it was 
renamed Vitrinella holmesii Dall. 

Its actual known range is: USA: 
North Carolina, South Carolina, Florida: 
West Florida, Florida Keys, Texas; 
Mexico: Campeche State, Yucatan State, 
Quintana Roo; Colombia; Bermuda; 
Puerto Rico. Recorded from Charleston 
Harbor, South Carolina (STIMPSON, 
1858); from (DaLL, 1892); from Cainhoy, 
Wando River, South Carolina (HOLMES, 
1859); from Fort Johson, South Carolina 
(KURTZ, 1860); from Beaufort, North 
Carolina (HARTMAN, 1945); from Grand 
Cayman (ABBOTT, 1958) but MOORE 
(1972) considers that Abbott's record 
corresponds to another different 
species); from the west coasts of Florida 
and Aransas Bay, Texas (MOORE, 1964); 
from Aransas Bay, Texas to Beaufort, 


15 


Iberus, 29 (2), 2011 


Figures 61A-H. Cochliolepis parasitica Stimpson, 1858. A: juvenile shell, 1.36 mm, Bahamas; B: 
protoconch, Bahamas; C-H: protoconchs, Bermuda (CHL). 

Figuras 61A-H. Cochliolepis parasitica Stimpson, 1858. A: concha juvenil, 1,36 mm, Bahamas; B: 
protoconcha, Bahamas; C-H: protoconchas, Bermuda (CHL). 


North Carolina (MOORE, 1972); from 
North Carolina (PORTER, 1974); from 
Campeche State, Yucatan State and 
Quintana Roo, Mexico (ODÉ, 1988); 
from Colombia (Díaz MERLANO éz 


116 


PUYANA HEGEDUS (1990); from Florida 
(Lyons, 1998); from Abaco, Bahamas 
(REDFERN, 2002) and from St. Augustine 
(Leg, 2009); from Cuba in the present 
work. 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 62A-F. Cochliolepis parasitica Simpson, 1858. A-B: juvenile shell, 1.6 mm, Guadeloupe 
(CJP); C: shell, 2.1 mm, Maria la Gorda, Cuba; D-F: protoconchs; D: from Turks 8 Caicos; E: 
Maria la Gorda; F: from Guadeloupe. 
Figuras 62A-E Cochliolepis parasitica Stimpson, 1858. A-B: concha juvenil, 1,6 mm, Guadeloupe 
(C]P); C: concha, 2,1 mm, Maria la Gorda, Cuba; D-F: protoconchas; D: de Turks * Caicos; E: 
Maria la Gorda; F: de Guadeloupe. 


C. parasitica has been recorded as 
fossil (as Adeorbis nautiliformis; a dis- 
puted synonym) from the post-Pliocene of 
South Carolina at the Wando River 


(HOLMES, 1860); from the Plio-Pleistocene 
of the Caloosahatchie beds, Florida (DALL, 
1892) and from the Plio-Pleistocene of St. 
Petersburg, Florida (PILSBRY, 1953). 


117 


Iberus, 29 (2), 2011 


Remarks: The type specimen of 
C. parasitica was lost in the 
Chicago fire of 1871, but two speci- 
mens given by Stimpson to Isaac 


Leartaretnow nte National 
Museum, n” 95079. The larger one, 
diameter 3.4 mm, is designated the 
lectotype. 


Cochliolepis striata Dall, 1889 (Figures 63A-E) 


Cochliolepis parasttica auct. non Stimpson, 1858. 


Cochliolepis striata Dall, 1889. Bull. Mus. Comp. Zool. Harvard, 18: 360. [Type locality: Egmont 


Key, Tampa Bay, Florida]. 


Type material: In USNM. Not examined. 


Other material examined: Mexico: 1 s, Puerto Morelos, Yucatan, 6-18 m (MHNS). Florida, USA: 4 
s, Sanibel, Lighthouse Beach (CHL); 1 s, dredged 4.5-7 m, Seahorse Key, Cedar Keys, (CHL); 1 s, 


APAC Pit, Sarasota Pit, Plio-Pleistocene (CHL). 


Description: Protoconch (Fig. 63D- 
E) white in color, smooth, composed of 
2 whorls (frequently not fully exposed, 
partially or totally covered by the sub- 
sequent whorls), and for this reason its 
diameter cannot be accurately mea- 
sured. The shell (Figs. 63A-C) is light 
honey in color, shows a uniform sculp- 
ture on its entire surface, including the 
base and the inner umbilical area; this 
is formed by numerous spiral cordlets 
with axial microstriae in the inter- 
spaces. There are also sinous growth 
lines. The spiral cords with microstriae 
are predominant on the dorsum of the 
shell, while on the base and in the 
umbilicus the growth lines are predom- 
inant. 

Dimensions: Holotype 6.5 mm in 
maximum diameter and 2.0 mm in 
height. 

Our largest shells measure 7.8 mm 
in diameter. 

Animal and radula are unknown. 

Distribution: This species has been 
recorded from Beaufort, North Carolina 
(Jacor, 1921); from Florida (JOHNSON, 
1934); Virgin Islands (NOwELL-USTICKE, 
1959); from North Carolina, Florida: 


East Florida, West Florida and Texas 
(MOORE, 1964, EMERSON éz JACOBSON 
(1976); from southeastern Panama 
(RADWIN, 1969); from Texas (ANDREWS, 
1977); from northwest Gulf of Mexico 
(ODÉ, 1969, 1988); from Abaco, 
Bahamas (REDFERN, 2001); also, in the 
present work, from Mexico: Yucatan 
and Cuba: Cienfuegos. 

Habitat: MOORE (1964) indicated 
that the species seems to prefer coastal 
bays with shallow waters. It is distrib- 
uted in the intertidal area between 0 
and 27 m. Various authors who have 
found it reported a depth between 0 
and 50 m, but these records were based 
only on empty shells. Nothing is 
known of the soft parts. MOORE (1964: 
41) considers C. striata an endemic 
species from the Gulf of Mexico, with 
strictly limited distribution. Our record 
indicates that the distribution is wider 
than expected. 

Remarks: The spiral cords and the 
axial microstriae in the interspaces dis- 
tinguish this species from all con- 
geners. Contrary to MOORE (1964: 174) 
the shell has the same sculpture on the 
base as on the rest of the shell. 


Cochliolepis planispiralis spec. nov. (Figures 64A-E) 


Type material: Holotype (Fig. 64A) in MNCN (15.05 /55054); paratypes: MHNS (100550, 1 s, Fig. 
64B), MNHN (24396, 1 s, Fig. 64C), IES (1 s), CFG (1 s), MCZ (1 s), USNM (1155034, 1 s), all from 


the type locality. 


Type locality: 15-20 m, Puerto Morelos, Yucatan, Mexico. 
Etymology: The specific name refers the planispiral development of the shell spire. 


118 


RUBIO £7 412.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 63A-E. Cochliolepis striata Dall, 1889. A-C: shells, 6.2, 7.8, 7.8 mm, Sanibel Light House 
Beach, Florida; D-E: protoconchs. 

Figures 63A-E. Cochliolepis striata Dall, 1889. A-C: conchas, 6,2, 7,8, 7,8 mm, Sanibel Light 
House Beach, Florida; D-E: protoconchas. 


Iberus, 29 (2), 2011 


Description: Shell (Figs. 64A-C) 
depressed, planispiral, the spire formed 
by about 2 Y whorls. The protoconch 
(Fig. 64D-E) has one whorl or a little 
more, measuring about 480 um in diam- 
eter. The embryonic whorl is smooth, 
but the subsequent whorl has an irregu- 
lar pattern of raised zigzag spiral 
threads. Each of these produces random 
off-shoots that project obliquely towards 
the adjacent thread, and this sculpture 
tends to become more regular towards 
the periphery. Varix at the transition to 
the teleoconch not very thickened. 
Teleoconch with two whorls and 
rounded periphery. Ornamentation con- 
sists of fine spiral grooves, which are 
best seen in the first whorl of the spire 
and at the periphery, and marked 
growth lines, some of them very sharp, 
spread over the dorsum and base of the 
shell. The umbilicus is widely open, and 
on its apex the protoconch can be seen. 
Aperture ovoid, parietal area not 
callous, columella straight, reflected 
outward, with slight thickening of the 


umbilical wall; no progresive ocelusion 
of the umbilicus is observed. 

Dimensions: The holotype is 2.63 
mm in maximum diameter; one 
paratype (MCZ) reachs 2.8 mm. 

Habitat: The shells were collected in 
sandy bottom. 

Distribution: Only known from the 
type locality. 

Remarks: The dorsal and basal orna- 
mentation are very constant. Cochli- 
olepis planispiralis spec. nov. shows 
great similarities to C. parasitica in pro- 
toconch ornamentation and teleoconch 
spiral grooves, but it is different because 
the periphery is regularly rounded and 
not in a low position; also in the devel- 
opment of the spire, and shape of the 
periphery, dorsal border of the aperture 
and columella (see Fig. 64). 

C. holmesti, C. patricioi and C. dif- 
ferens are different by the different 
sculpture of the protoconch. 

C. striata and C. adamsi lack the 
spiral cordlets in the protoconch, having 
a stronger striation on the teleoconch. 


Cochliolepis patricioi spec. nov. Rubio, Rolán € Lee (Figures 65A-G) 


Type material: Holotype (1 s, Figs. 65A-B) in FLMNH (448608); paratypes: USNM (1155035, 1 s), 
ANSP (1 s), CHL (1 s) from St. Augustine, St. Johns (Figs. 65C-E) (all ex CHL). 

Type locality: 60 m, off Sombrero Light, Key Vaca, Monroe Co. Florida, USA. 

Etymology: The species is named for Patricio Calviño, good friend of the first author. 


Description: Shell (Figs. 65A-E) 
depressed, almost planispiral, com- 
posed of 3 Y whorls. The protoconch 
(Fig. 65F-G) has 1 whorl measuring 
about 340 um in diameter, apparently 
smooth although very small granules 
can be seen all over, mainly on the 
areas close to the suture. Varix at the 
transition with the teleoconch not 
thickened. Teleoconch with two 
whorls; the first possessing 35-40 
somewhat robust axial ribs. At the 
beginning of the second whorl the ribs 
begin to rapidly fade and soon change 
into simple, but marked, growth lines. 
This ornamentation, although most 
easily seen dorsally, can be observed 
on the periphery as well as on the 
base. Umbilicus wide and deep. Aper- 


120 


ture ovoid, parietal area not callous, 
columella reflected outward, without 
thickening of the umbilical wall or 
progressive occlusion of the umbili- 
cus. 

Dimensions: 2.0 mm in maximum 
diameter. 

Habitat: Collected in sandy sedi- 
ment at 60 m. 

Distribution: Only known from the 
type locality and off St. Augustine, 
Florida. 

Remarks: The dorsal ornamenta- 
tion is very variable. Some shells lack 
the axial ribbing so evident on the 
first whorl of the teleoconch, showing 
only strong growth lines. 

Cochliolepis patricioi spec. nov. 
can be distingished from: 


RUBIO £7 ALz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 64A-E. Cochliolepis planispiralis spec. nov. A: holotype, 2.63 mm, Puerto Morelos, Yucatan 
(MNCN); B-C: paratypes, 2.4 mm (MNHN), 2.2 mm (MHNS); D-E: protoconchs. 

Figuras 64A-E. Cochliolepis planispiralis spec. nov. A: holotipo, 2,63 mm, Puerto Morelos, Yucatan 
(MNCN); B-C: paratipos, 2,4 mm (MNAN), 2,2 mm (MANS); D-E: protoconchas. 


C. parasitica, which has consistent C. holmesii, which also lacks this 
sculpture in the protoconch and lacks ribbing and has the periphery lower 
ribs on the first whorl of the teleo- and subangulate, rather than rounded 
conch. and at mid-whorl. 


12 


Iberus, 29 (2), 2011 


Figures 65A-G. Cochliolepis patricioi spec. nov. Rubio, Rolán 82 Lee. A-B: holotype, 2.0 mm, 
Monroe Co. Florida (FLMNH); C-E: paratypes: 1.8, 1.9, 1.7 mm, St. Augustine, Florida 
(USNM, ANSP and CHL, respectively); F-G: protoconchs: F: from the holotype; G: from the 
paratype of fig. C. 

Figuras 65A-G. Cochliolepis patricioi spec. nov. Rubio, Rolán e* Lee. A-B: holotipo, 2,0 mm, Monroe 
Co. Florida (FLMNHA); C-E: paratipos: 1,8, 1,9, 1,7 mm, St. Augustine, Florida (USNM, ANSP y 
CAL, respectivamente); F-G: protoconchas: F: del holotipo; G: del paratipo de la 


122 


RUBIO E7 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


COrsiaatatand ie adams, which 
also lack the ribs on the teleoconch 
and have  conspicuous  spiral 
striation. 


C. differens spec. nov., which lacks 
the axial ribs on the first whorl of the 
teleoconch, has a more inflexed col- 
umella, and has a smaller umbilicus. 


Cochliolepis differens spec. nov. Rubio, Rolán €: Lee (Figures 66A-D) 


Type material: Holotype (Figs. 66A-D) in FLMNH (448615) ex CHL. 
Type locality: APAC Pit, Sarasota, Florida, USA Plio-Pleistocene. 
Etymology: The specific name refers to the shell characters, which differ rather strikingly from 


those of its congeners. 


Description: Shell (Figs. 66A-C) 
depressed, fine, smooth, composed of 3 
% whorls. The protoconch (Fig. 66D) 
has 1 Y bulbous whorls, about 430 um 
in diameter, slightly raised above the 
other whorls. Two stages can be dis- 
cerned: on the first the surface is 
covered by microgranules; the second 
exhibits fine growth striae. Teleoconch 
formed by 2 Y convex and rapidly- 
expanding whorls and is totally 
smooth except for fine growth lines. 
Suture wide and strongly marked on 
all the shell except on the last quarter 
of the body whorl. Periphery nearly 
symmetrically rounded. Aperture 
strongly prosocline, angled at the 
suture. Columella arched, a little 
reflected towards the umbilicus; exter- 
nal lip sharp, angled at the periphery 
and a little sinuous basally. Parietal 
callus barely appreciable. Umbilicus 


wide and deep, with a straight and stri- 
ated wall. 

Dimensions: Holotype is 4.3 mm in 
maximum diameter and 1.76 mm in 
height. 

Habitat: Unknown. 

Distribution: Only known from the 
type material. 

Remarks: Cochiolepis differens can 
be distinguised from C. parasitica, 
because the latter has sculpture on the 
protoconch and its apex is flat. 

C. holmesii has a different proto- 
conch sculpture and a sculptured teleo- 
conch. 

Cochiolepis striata and C. adamsi 
have conspicuous spiral sculpture on 
the teleoconch. 

C. patricioi has axial ribs on the first 
whorl of the teleoconch, possesses a less 
inflexed columella, and has a larger, 
more open umbilicus. 


Cochliolepis sp. (Figures 67A-C) 


Material examined: Cuba: 2 s, Rancho Luna Beach, Cienfuegos, 20 m (MHNS). 


Description: Protoconch (Fig. 67C) 
white in color, smooth, formed by 1 Y 
whorls of spire and about 360 um of 
diameter; apparently it has two well- 
differentiated stages, the first is 
bulbous, projected from the rest of the 
shell and terminates in a varix; the 
second stage, of one whorl only has 
fine growth lines. A varix marks the 
beginning of the teleoconch and the 
onset of the spiral sculpture. The shell 
(Figs. 67A-B) is light honey in color, 
and shows a uniform sculpture on the 


entire surface, including the base and 
the imner umbilical area: this is 
formed by numerous spiral cordlets 
with axial microstriae in the spaces 
between cords. Sinuous growth lines 
are also present. 

Dimensions: Our shells measure 
2.9 and 3.0 mm in diameter and 1.63 
mm in height. 

Animal and radula are unknown. 

Remarks: Cochliolepis sp. is rather 
similar in its general appearance to C. 
adamsii, but the protoconch is pro- 


125 


Iberus, 29 (2), 2011 


Figures 6G6A-D. Cochliolepis differens spec. nov. Rubio, Rolán 82 Lee. A-C: holotype, 4.3 mm, Plio- 
Pleistocene of Sarasota, Florida (FLMNH); D: protoconch. 

Figuras 66A-D. Cochliolepis differens spec. nov. Rubio, Rolán € Lee. A-C: holotipo, 4,3 mm, Plio- 
Pleistoceno de Sarasota, Florida (ELMNH); D: protoconcha. 


jected upward, shows two different 
stages of development, and is not par- 
tially covered by the first whorl of the 
teleoconch. Since this is the main dis- 


124 


tinguishing character and we have 
such scanty material we defer naming 
this taxon until we have more mater- 
ial for study. 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 67A-C. Cochliolepis sp. A-B: shells, 2.9, 3.0 mm, Rancho Luna Beach, Cuba; C: proto- 
conch. 
Figuras 67A-C. Cochliolepis sp. A-B: conchas, 2,9, 3,0 mm, Playa Rancho Luna, Cuba; C: protocon- 
cha. 


Genus Episcynia Mórch, 1875 


Episcynia Mórch, 1875 (described as subgenus of Architectonica). Malak, Blátter, 22: 155. 
Type species: (by monotypy) Solarium inornatum d'Orbigny, 1842. 


Diagnosis: Shell trochiform, whorls 
rather convex, and with a minutely serrate 
peripheral keel, base flattened, umbilicus 
narrow and deep, flat sided and strongly 
angled with the base of the shell. Perios- 
tracum thin, produced into spiral fringes of 
filaments above and below the periphery. 

Remarks: MOORE (1964) placed Episcy- 
nia multicarinata in synonymy with Episcy- 


nia inornata comparing the types of Dall 
and d'Orbigny and reporting that there 
were no morphological differences between 
them. Species of Episcynia are known from 
the Miocene to Recent. Two valid species 
have been described from the West Atlantic: 
Episcynia inornata (a recent species) and 
Episcynia naso (a fossil species fron the 
Miocene of the Dominican Republic). 


125 


Iberus, 29 (2), 2011 


Episcynia inornata (d'Orbigny, 1842) (Figures 68A-F, 69A-C) 


Trochus (Solarium) inornatum d'Orbigny, 1842. Moll. Hist. Phys. Pol. et Nat. Líle de Cuba, 2: 67- 
68, pl. 19, figs. 25-27. [Type locality: Saint Thomas, Virgin Islands]. 

Vitrinella multicarinata Dall, 1889. Bull. Mus. Comp. Zoology, 18: 392-393. [Type locality: Off 
Cape Hatteras, North Carolina, 15 fms]. 


Type material: Lectotype (here designated) in NAMUK (Fig. 69C) and paralectotypes (Figs. 69A- 
B). 

Other material examined: Florida, USA: 1 s, beach, 16 Ave. S, Jacksonville Beach, Duval Co. (CHL); 
1 s, 39-45 m, 40-60 mi E Ponte Vedra, St Johns Co. (CHL); 1 s, 30 m, 20 mi E St. Augustine, St Johns 
Co. (CHL); 1 s, 53 m, 65 mi E St. Augustine, St Johns Co. (CHL); 1 s, 4.5-6 m, E Seahorse Key, Cedar 
Keys, Levy Co. (CHL). Cuba: 6 s, Cienfuegos Bay, 20 m (MHNS); 32 s, Rancho Luna Beach, 10-54 
m (CFG). Martinique: 1 s, Fort de France, NW of Grande Seche, 18 m (MCZ 361885); 1 s, Fort de 
France, St. Louis, 3-4 miles S. of Fort, 14-16 fms (MCZ 243767). Surinam: 1 s, 70 miles NNE Para- 
maribo, 652'N, 5446'W, 51 m (MCZ 274032). Brazil: 1 s, Praia Itararé, Sáo Vicente, Sáo Paulo (CHL); 


1 s, Itaparica, 4-5 m (MHNS); 2 s, Salvador (MHNS). 


Description: The most complete 
description is in MOORE (1964: 105-109). 

Shell (Figs. 68A-D) with 5 whorls. The 
protoconch (Figs. 68E-F) has barely one 
whorl with a slightly rough surface, about 
190 um in maximum diameter. 

Dimensions: the figured shell is 3.3 
mm in diameter and 1.73 mm in height. 

Animal and radula unknown. 

Habitat: Species with a large bathy- 
metric distribution, recorded between 15 
and 110 m in depth. According to MOORE 
(1964) the species has a wide range of dis- 
tribution, but it is uncommon; it seems 
that the species prefers shallow water and 
the deepest record for living material is 15 
fathoms, cited by DALL (1889a). 

Distribution: lt has been recorded from 
St. Thomas, Virgin Islands (D'ORBIGNY, 
1842); from off Cape Hatteras, North Car- 
olina (DALL, 1889); from Lake Worth, Palm 
Beach, Cape Florida, Miami and Destin, 
Florida (PiLSBRY € MCGINTY, 1946); from 
Pernanbuco and Alagoas, Brazil (LANGE 
DE MORRETES, 1949); from Bocas Island, 
Panama (OLsson éz MCGINTY, 1958); from 
North Carolina, South Florida, west coast 
of Florida, Texas, Panama, Puerto Rico and 
the Virgin Islands (MOORE, 1964); Puerto 
Rico (WARMKE é ABBOTT, 1961); from 
northwestern Campeche Bank, Mexico 
(RICE 62 KORNICKER, 1965); from North 
Carolina to Florida, Texas and the Greater 
Antilles (ABBOTT, 1974); from Texas shores 
(ANDREWS, 1977); from Venezuela (PRINCZ, 
1982); Cuba (ESPINOSA ET AL., 1985); from 


Z6 


Pernambuco, Brazil (MELLO éz PERRIER, 
1986); from northwest Gulf of Mexico 
(ODÉ, 1988); from Aruba island (DE JONG éz 
COOMANS, 1988); from shallow waters off 
Hutchinson Island, Florida to Texas and 
the Caribbean Sea (Lyons, 1989); from 
northeast Florida (LEE, 2009). 

Remarks: Over the years, this species 
has been included in the genera Adeorbis, 
Architectónica, Torinia, Trochus and Vit- 
rinella before its definitive placement in 
Episcynia. The fine periostracum, the 
finely serrated peripheral keel and the 
straight-sided umbilicus are the main dis- 
tinguishing characters of this species. 

DaALL (1889a) described Vitrinella 
(Episcynia?) multicarinata from Cape Hat- 
teras, North Carolina, but he did not make 
a comparison of his species with that of 
d'Orbigny, and most  malacologists 
assumed that the two were different. 
MOORE (1964) considered them synony- 
mous, indicating that he had the opportu- 
nity to review d'Orbigny's four syntypes 
in the British Museum. The types were in 
a vial labelled by d'Orbigny as Rotella car- 
inata, but they fit the description of Solar- 
ium inornatum. The specimens were com- 
pared to the American specimens, and no 
distinguishing characters could be found. 
Of the four syntypes, the smallest one (Fig. 
69C) is in good condition and is hereby 
designated the lectotype. So, Episcynia 
multicarinata may be considered simply 
as a form with extra angulations on the 


periphery of the whorls (Fig. 68D). 


RUBIO £7 4L£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 68A-E. Episcynia inornata (Y Orbigny, 1842). A-C: shells, 3.2, 2.7, 3.3 mm, Rancho Luna 
Beach, Cienfuegos, Cuba (MHNS); D: shell, form “multicarinata,” 3.6 mm, Praia Itararé, Sáo 
Vicente, Sáo Paulo (CHL); E-F: protoconchs, from Cuba and Florida. 

Figuras 68A-FE Episcynia inornata (4'Orbigny 1842). A-C: conchas, 3,2, 2,7, 3,3 mm, Playa Rancho 
Luna, Cienfuegos, Cuba (MHNS); D: concha, forma “multicarinata,” 3,6 mm, Praia Itararé, Sáo 
Vicente, Sáo Paulo (CHL); E-F: protoconchas, de Cuba y Florida. 


Iberus, 29 (2), 2011 


Figures 69A-C. Episcynia inornata (d'Orbigny, 1842). A-C: shells labeled as Rotella carinata in 
NHMUK: A-B: paralectotypes; C: lectotype. 
Figuras 69A-C. Episcynia inornata (4'Orbigny 1842). A-C: conchas etiquetadas como Rotella cari- 
nata en NAMUK: A-B: paralectotipos; C: lectotipo. 


Genus Parviturboides Pilsbry 2 McGinty, 1950 


Parviturboides Pilsbry $ McGinty, 1950. The Nautilus, 63(3): 86. 
Type species: (by original designation) Vitrinella interrupta C.B. Adams, 1850 = Cyclostrema 


sanibelense Pilsbry, 1939. Caribe, Recent. 


Diagnosis: In MOORE (1964: 155) 
be shell is small, EroCionaa, 
sculpture of spiral cords and axial 
threads, umbilicus narrow and bor- 
dered by a spiral cord. Aperture sub- 
circular but angular above, columel- 
lar margin thickened. The animal has 
two ciliated tentacles, black eyes, 
two pallial tentacles, a pair of oper- 
cular lobes amd the posterior end of 
the foot may be bilobed. The penis is 
long, straight, and glandular. The 
operculum is thin, multispiral and 
cinc ULT 


128 


Remarks: The shell is very similar 
to some of the genus Parviturbo 
Pilsbry € McGinty, 1945, from which 
it can be distinguished by its proto- 
conch, which is less than 2 whorls, 
and by the abrupt beginning of the 
sculpture on the teleoconch. PILSBRY 
(1950: 87), in describing the genus 
Parviturboides, stated that it should 
be compared with the genus 
Pseudorbis Monterosato, 1884 for the 
similarity in shell characters. MOORE 
(1964: 156) stated “but this has not 
yet been done”. This comparison is 


RUBIO ET AZ.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


no longer necessary because the 
genus Pseudorbis was placed in 
Skeneidae by RUBIO é RODRÍGUEZ 
BABÍO (1991), who noted that its 


radula is rhipidoglossate, and its 
anatomical characters are similar to 
those of the genera Skenea and 
Dikoleps. 


Parviturboides interruptus (C.B. Adams, 1850) (Figures 70A-L, 71A-E) 


Vitrinella interrupta C.B. ADAMS, 1850. Monograph of Vitrinella, a new genus of new species of 
Turbinidae: 6. [Type locality: Port Royal, Jamaica]. 

Cyclostrema zacalles Mazyck, 1913: 18. 

Cyclostrema sanibelense Pilsbry, 1939. The Nautilus, 53: 53, pl. 8, fig. 3. [Type locality: Sanibel 
Island, Florida]. 

Parviturbo zacalles (Mazyck, 1913): In PiLsBRY € MCGINTY, 1945b. The Nautilus, 59: 57, pl. 6, 
PS Zz Za O: 

“Cyclostrema” interruptum sanibelensis Pilsbry €££ McGinty, 1945b. The Nautilus, 59: 58, pl. 6, 
fig. 9. 

Cyclostrema interruptum C.B. Adams in Pilsbry, 1946. Notulae Naturae, 162: 5, pl. 1, fig. 4-5. 


Type material: CLENCH € TURNER (1950: 294, plate 35) mention two specimens in the lot of C.B. 
Adams Collection (MCZ 156270), and they represent two different shells in their plate, but they 
caption the two figures “holotype” despite the fact that there are two different shells depicted. In 
the actual lot examined by us only one shell is present and we consider it the lectotype. 

Other material examined: Cuba: 159 sp, Cañón de la Bahía de Cienfuegos, between 0-10 m (CFG); 
14 sp, Rancho Luna Beach, 10-20 m (MHNS); 28 sp, Rancho Luna Beach, 45 m (MHNS). Guade- 
loupe: 2 s, north coast, 2 m, between rocks (CJP); 3 s, Point-a-Pitre, 5 m (CHL). Florida, USA: 2 s, 
21 m, 14 mi ENE Mayport, Duval Co. (CHL); 2 s, 3-4 ft, Pelican Shoals, Key West, Monroe Co. (CHL); 
5 sp, Sebastian Inlet, Brevard County, North Jetty, under bridge (CHL); 3 s, SE Clearwater Island 
bridge Little Pass, Pinellas Co. (CHL); 1 s, 10th Avenue, Atlantic Beach, Duval Co. (CHL); 1 f, 2-3 
m, Seahorse Key, Cedar Keys, Levy Co. (CHL). Puerto Rico: 7 s, beach, Holiday Inn, San Juan, 
(CHL). Virgin Islands: 7 s, beach, Magens Bay, N coast St. Thomas (CHL). Nicaragua: 3 s, The 
Witties, 20 m (MHNS). Panama: 1 s, beach, Portobello (CHL); 5 s, Careneros Island 920.394'N, 
32713.932'W, 8-9 m (CHL); 1 s, 1 mi Punta San Blas, San Blas Island, intertidal (CHL); 1j, 09%22.027'N, 
8214.336'W (CEG). Venezuela: 12 s, near Jose Griego, N Margarita Island (CHL). Haiti: 6 s, Labaree, 


sand beach (CHL). Brazil: 1 s, Praia de Itararé, Sáo Vicente, Sáo Paulo (CHL). 


Description: The best conchologi- 
cal and anatomical description is in 
MOORE (1964: 156-161). 

Shell (Figs. 70A-C, 71A-C) a little 
depressed, globose, trochiform, spiral 
sculpture formed by strong cords and 
fine axial striae; umbilicus very 
narrow. Protoconch (Figs. 70D-F) large, 
with a diameter of about 400 um, its 
surface roughened by numerous tuber- 
cles and some spiral lines. Two differ- 
ent stages can be distinguished. Teleo- 
conch with spiral strong cords and fine 
axial ribs in the interspaces; on the last 
whorl 8-9 spiral cords can be seen. 
Base rounded, aperture oblique, almost 
circular, but a little angled at the fusion 
point of the external lip with the col- 
umella. 


Dimensions: Holotype 1.59 mm in 
diameter. Our largest shells measure 
about 2.5 mm in maximum dimension. 

Operculum (Figs. 70G-H) chitinous, 
multispiral with a central nucleus. 

Radula (Fig. 701) taenioglossate, the 
formula 2+1+R+1+2. Central tooth wide 
basally, with three denticles on the 
ventral margin. Cutting area formed by 
a large and sharp cusp and 6-7 small 
denticles of small size at each side. 
Lateral teeth similar to the central one, 
cutting area with a central cusp and 4-5 
smaller denticles at each side. Marginal 
teeth elongated, with 22-25 denticles on 
the free margin. 

Habitat: This species lives between 0 
and 48 m deep; deeper records are 
based on empty shells. In Cienfuegos 


129 


Iberus, 29 (2), 2011 


Figures 70A-I. Parviturboides interruptus (C.B. Adams, 1850). A-C: shells, 1.4, 1.4, 1.3 mm, 
Rancho Luna Beach, Cienfuegos, Cuba (CFG); D-F: protoconchs; G-H: operculum, 0.65 and 0.5 
mm, from shells of 1.5 and 1.4 mm; I: radula. 

Figuras 70A-1. Parviturboides interruptus (C.B. Adams, 1850). A-C: conchas, 1,4, 1,4, 1,3 mm, 
Playa Rancho Luna, Cienfuegos, Cuba (CFG); D-F: protoconchas; G-H: opérculos, 0,65 y 0,5 mm, 
procedentes de conchas de 1,5 y 1,4 mm, l: rádula. 


130 


RUBIO £7 ALz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 71A-E. Parviturboides interruptus (C.B. Adams, 1850). A-C: shells, 1.25, 1.1, 1.3 mm, The 
Witties, Nicaragua (MHNS); D: protoconch; E: shell, Guadeloupe (CJP). 

Figuras 71A-E. Parviturboides interruptus (C.B. Adams, 1850). A-C: conchas, 1,25, 1,1, L3 mm, 
The Witties, Nicaragua (MANS); D: protoconcha; E: concha, Guadeloupe (C]P). 


Bay, Cuba it is relatively common 
between 0 and 10 m deep. It has been 
recorded on sponges from 7 to 90 m 


deep (ABBOTT, 1974). Under rocks and 
in crevices in shallow water (ANDREWS, 


1977). On rocky coasts it has been found 
among sabellariid polychetes (WILEY ET 
AL. 1982). Also found on sandy and 
muddy bottoms (VOKES éz VOKES, 1984). 
On reefs far from the coast this species 


57 


Iberus, 29 (2), 2011 


has been seen associated with colonies 
of Oculina varicosa, algae, sponges, 
octocorals and dead coral, between 6 
and 80 m deep (REED € MIKKELSEN, 
1987). Also on algae and sand bottom 
(MIKKELSEN ET AL. 1995). It has been 
collected directly on sponge and algae 
in 1-2 feet and from colonies of Vermic- 
ularia knorri taken at 30 m (Lek, 2009). 
In Cuba it has been collected in the 
bases of Gorgonia flabellum and G. 
ventalina. 

Distribution: It is a common and 
widely distributed species in the West 
Indies. It has been recorded from Port 
Royal, Jamaica (C.B. ADaAms, 1850). 
From Sanibel Island, Florida (PILSBRY, 
1939). From Colón and Bocas Island, 
Panama (OLsson € MCGINTY, 1958). 
From South Carolina, Florida, Texas, 
Mexico, Panama, Jamaica, Haiti and 
Puerto Rico (MOORE, 1964; ANDREWS, 
1977). From South Carolina to Gulf of 
Mexico and Caribbean Sea (HOUBRICK, 
1968). From  Portete, Costa Rica 
(HOUBRICK, 1968; ROBINSON  éz 
MONTOYA, 1987). From North Carolina 
(PORTER, 1974). From punta del Morro 
to Punta Delgada, Veracruz, Mexico 
(WILEY ET AL. 1982). From Campeche to 
Ciudad del Carmen and Zacatal; from 
El Cuyo to punta Ninum, from Punta 
Yalkupul to isla Cerritos; from Isla 
Mujeres to Isla Holbox; from Cancún to 
the Belize border (VOKES € VOKES, 
1984). From North Carolina to Florida; 
Texas, the Antilles, Brazil, Uruguay 


(Rios, 1985). From Cuba (ESPINOSA ET 
AL., 1985). From Florida (REED éz MIk- 
KELSEN, 1987). Aruba; Puerto Rico (DE 
JONG € COOMANS, 1988). From Florida 
to Texas and north coast of South 
America, Colombia (Díaz MERLANO éz 
PUYANA HEGEDUS, 1994). From Indian 
River Lagoon, Florida (MIKKELSEN ET 
AL. 1995); from Abaco, Bahamas 
(REDFERN, 2001). From Cahuita to 
Gandoca, Cuba (ESPINOSA éz ORTEA, 
2001). From St. Augustine Inlet, north- 
east Florida (LEE, 2009). Venezuela, 
Haití, Panama, Nicaragua, and Brazil 
from our material. 

Remarks: Parviturboides interrup- 
tus is like no other vitrinellid from the 
West Atlantic. On the other hand, it can 
be confused with some species of the 
genus Parviturbo Pilsbry € McGinty, 
1945, distinguished from them by its 
protoconch of no more than 2 whorls, 
its sculpture commencing abruptly at 
the end of the protoconch. Another 
genus with conchologically similar 
species, Pseudorbis Monterosato, 1884, 
has been placed in Skeneidae, based on 
the rhipidoglossate radula of Pseudor- 
bis granulum (Brugnone, 1873), that 
has been recently observed (RUBIO € 
RODRÍGUEZ BABÍO, 1991). 

Some populations (shown in 
Figures 71A-C) present more numerous 
riblets in the axial sculpture (about 45 
on the first whorl versus 23) than the 
shells of other populations (Fig. 70A- 
E). 


Genus Pleuromalaxis Pilsbry £ McGinty, 1945 


Pleuromalaxis Pislbry y McGinty, 1945a. The Nautilus, 59: 1-13. 
Type species: (by monotypy) Pseudomalaxis (Pleuromalaxis) balesi Pilsbry € McGinty, 1945. 


Caribe, Recent. 


Diagnosis: Shell strongly depressed, 
with two spiral keels on the periphery. 
Space between the keels concave, axial 
riblets present on both top and bottom 
of the whorl. Umbilicus wide, shallow 
to moderately deep. 

Remarks: The taxon Pleuromalaxis 
was created as a section of Pseudoma- 


152 


laxis, as  Pilsbry € McGinty 
believed that their species was con- 
generic with Pseudomalaxis nobilis 
ARE e rn ater O LS SON es 
MCGINTY (1958) considered it a full 
genus and placed it in the Vitrinell- 
idae, removing it from the Architec- 
tonicidae. 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Pleuromalaxis balesi (Pilsbry £ McGinty, 1945) (Figures 72A-D) 


Pseudomalaxis (Pseudomalaxis) balesi Pilsbry € McGinty, 1945a. The Nautilus, 59: 10, pl. 2, fig. 
8. [Type locality: Missouri Key, Florida Keys]. 


Type material: Holotype in ANSP (181124). Represented in MOORE (1964). Not examined. 

Other material examined: Cuba: 2 s, Cayo Avalos, 8 m (MHNS); 1 s, Cayo Perez Diego, 5 m (MHNS); 
1 s, Jibacoa, 3-6 m (MHNS); 1 s, Baracoa, 0 m (MHNS); 4 s, Guajimico, 0 m (MHNS); 3 s, Cienfue- 
gos Bay, sta. 12, 2207'N 8027'W, 9 m (MHNS); 32 s, Cienfuegos Bay, 8 m (MHNS); 1 s, Cienfue- 
gos Bay, 20-30 m (MHNS); 1 s, Rancho Luna Beach, 12 m (MHNS); 1 s, Rancho Luna Beach, 35 m 
(MHNS)y; 13 s, Rancho Luna Beach, 45 m (MHNS); 2 s, Los Laberintos, Rancho Luna Beach, 35 m 
(MHNS). Granadines: 1 s, Mayreau, 8 m, coralline sand with coral blocks, gorgonians and sponges 
(CJP). Trinidad and Tobago: Tobago, 1 s, Horse shoe reef, 15 m, sandy grit (CHL). Mexico: 1 s, 


Puerto Morelos, Yucatan, 6-18 m (MHNS). 


Description: Shell (Figs. 72A-B) 
strongly depressed but not planispiral. 
Umbilicus wide. Protoconch (Fig. 72D) 
apparently smooth, measuring 260 um, 
it has two distinct parts, and terminates 
in a varix. Teleoconch completely 
covered by fine spiral cordlets. Peripher- 
ally bicarinate and concave between the 
two keels. On the dorsum there are two 
series of axial ribs: the finer ones begin 
on the suture and fade towards the 
middle of the dorsum, the stronger ones 
run from the middle of the dorsum out 
to the upper peripheral keel, forming 
strong nodules at their terminations. 
The base is divided in two by a strong 
spiral cord. Aperture oblique, peristome 
not continuous, parietal callus strong. 
Umbilicus wide and shallow which 
exposes the protoconch within. 

Dimensions: Holotype 1.59 mm in 
diameter. Our material is between 1 and 
1.5 mm, but a 1.8 mm shell is mentioned 
in the literature. 

Operculum (Fig. 72C) fine, multispi- 
ral and with a central nucleus. 

Habitat: This species has a wide 
bathymetric range, being recorded 


between 0 and 100 m in depth. It has 
been collected alive under rocks in 
shallow water (MOORE, 1964). In Cuba 
itis a relatively frequent species but 
not common, only 1-2 specimens 
appear in each station studied. In 
Cuba it was collected between 0 and 
40 m. Reported further down in other 
areas. 

Distribution: It is recorded from 
the USA: Missouri Key, Florida Keys 
(PrLsBRY é MCGINTY, 1945a); from 
Havana Province, Cuba (JAUME 
BORRO, 1946); from Bocas Island, 
Panama (OLSssON € MCGINTY, 1958); 
from Grand Cayman Island (ABBOTT, 
1958); from Puerto Rico (WARMKE «z 
ABBOTT, 1961); from Florida, Texas and 
Panama (MOORE, 1964); from South- 
east Florida, Texas and the Caribbean 
(ABBOTT, 1974); from Quintana Roo, 
Mexico (VOKES é€z VOKES, 1984). From 
Grenadines and Tobago in the present 
work. 

Remarks: The only other species 
known in this genus, Pleuromalaxis 
pauli Olsson € McGinty, 1958,is 
smaller and has a more elevated spire. 


Genus Solariorbis Conrad, 1865 


Solariorbis Conrad, 1865. Amer. Journ. Conch. 1: 30. 
Type species: (by monotypy) Delpinula depressa Lea. Eocene of Clairbonian, Alabama. 


Diagnosis: After PiLsBRY (1953): 
sotell 2 to 4 ma, white, stiong, 
depressed, with 3 or 4 whorls, either 
rounded or angular, and usually with 


some spiral striation, the grooves typi- 
cally punctate; apical whorls level and 
smooth. The umbilicus has a spiral 
ridge on a thickening of the wall, 


SS 


Iberus, 29 (2), 2011 


Figures 72A-C. Pleuromalaxis balesí (Pilsbry 8% McGinty, 1945). A-B: shells, 1.0, 1.5 mm, Rancho 
Luna Beach, Cienfuegos, Cuba; C: operculum; D : protoconch. 

Figuras 72A-C. Pleuromalaxis balesi (Pilsbry € McGinty 1945). A-B: conchas, 1,0, 1,5 mm, Playa 
Rancho Luna, Cienfuegos, Cuba; C: opérculo; D : protoconcha. 


ending in a callus lobe or ledge at the 
columellar margin. Aperture rounded, 
with a small groove at the top. Outer lip 
rather thin and evenly curved”. 
Remarks: Solariorbis was described 
as a section or subgenus of Teinostoma. 
The peculiar thickening around the 
umbilicus is one of the more important 
characteristics of the genus. Another 
character common to many species is 
the reticular punctate sculpture due to 
the interaction of spiral grooves and 
axial ribs; this character was mentioned 
by DaALL (1892) in reference to T. 
depressum as the type species of Solari- 
orbis. The genus is distributed from the 
Eocene to the Recent with numerous 
fossil species described. MOORE (1964) 
considered the following species as 
Recent: S. multistriata (A.E. Verrill, 


134 


1884), S. mooreana (Vanatta, 1904), S. 
blakei (Rehder, 1944), S. infracarinata 
(Gabb, 1881), Solariorbis schumoi 
(Vanatta, 1913) S. hondurasensis 
(Vanatta, 1913), S. terminalis (Pilsbry € 
McGinty, 1946), S. corylus Olsson € 
McGinty, 1958, S. decipiens Olsson é 
McGinty, 1958, and S. semipunctus 
Moore, 1964. To these it is necessary to 
add S. guianensis Altena, 1966 and S. 
antillensis de Jong € Coomans, 1988, 
which were described subsequently; 
also included in the present account as 
recent are S. funiculus (Dall, 1892) and 
S. contractus (Vanatta, 1913). Solariorbis 
decipiens is transferred to the genus 
Anticlimax due to the conchological 
similarity with the species of this genus. 

Some species previously placed in 
the genus Solariorbis, such as S. con- 


RUBIO ET 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


tractus, S. funiculus and S. opsitelo- 
tus present a small triangular thicken- 
ing at the junction of the columella 
and the umbilical margin, which does 
not impinge on the umbilicus; further 
more they lack the characteristic 
ornamentation of the genus Solarior- 
bis (the reticular punctate sculpture 


due to the interaction of the spiral 
grooves and axial ribs). For these 
reasons and because their spiral cord 
ornamentation, umbilical margin and 
protoconch are more similar to Vit- 
rinella anneliesae, we have deter- 
mined that they are better placed in 
Vitrinella. 


Solariorbis antillensis de Jong $: Coomans, 1988 (Figures 73A-C) 


Solariorbis antillensis de Jong € Coomans, 1988. Studies of the Fauna of Curacao and other 
Caribbean Islands, 69: 33, pl. 13, fig. 143. [Type locality: Curacao / Aruba] 


Type material: Holotype in ZMA (3.87.064). Represented in DE JONG ££ COOMANS (1988). Not exam- 


ined. 


Other material examined: Cuba: 1 s, Rancho Luna Beach, 35 m (MHNS). Puerto Rico: 1 s, E Boca 
de Cangrejos (MCZ 361886). Bahamas: 1 s, South Riding Rocks, Cay Sal Bank, base of coral reef in 
fine sediment (CHL). Antiqua and Barbuda: 1 s, [le Sister, N St. Johns, 5-6 m (CJP). Guadeloupe: 2 


s, 1 £, Port Louis, 0-2 m, brushing stones (CJP). 


Description: The original description 
given by DE JONG £: COOMANS (1988) is 
as follows: “Shell disc-shaped, wider 
than high, 1% nuclear and two postnu- 
clear whorls. The whorls are rounded, 
without carinae and covered by spiral 
striae with punctuated grooves. $. antil- 
lensis differs from S. guianensis by a 
quite different protoconch, the under 
side being less flattened, the umbilicus 
less closed, the axial wrinkles less mani- 
fest, the spirals more prominent and 
their punctuation more clear”. The ambi- 
guity of the distinguishing characters has 
given us reason to amplify the descrip- 
tion. 

We add: Shell (Figs. 73A-B) planispi- 
ral, depressed, shell of 2 % rapidly- 
expanding whorls. Protoconch (Fig. 73C) 
with 1 Y whorls, totally smooth, about 
240 um in maximum diameter. Teleo- 
conch with 1 Y whorls ornamented by 
regularly spaced axial ribs and spiral 
cords of equal size. This sculpture covers 
all the shell except a small area on the 
base close to the periphery. The intersec- 


tion of ribs and cords forms a regular 
reticulation characteristic of the genus. 
Aperture rounded, base of the columella 
and internal lip thickened and reflected 
outward forming a typical callus. 
Umbilicus narrow and deep, nearly 
closed by the thickening of the columella 
until the last Y of the body whorl, which 
then deviates laterally. 

Dimensions: Holotype 1.6 mm in dia- 
meter. The figured specimen measures 
1.29 mm in maximum dimension. 

Habitat: Nothing is mentioned by DE 
JONG £ COOMANS (1988) in the descrip- 
tion work of the species. Our material 
comes from a coralline sandy bottom 35 
m deep. 

Distribution: Only know from 
Curacao / Aruba, Puerto Rico, Bahamas, 
and Cuba. 

Remarks: S. antillensis is very close to 
S. guianensis, with which it can be con- 
fused. It is best differentiated by the reg- 
ularity of the ornamentation, which 
forms a characteristic reticulation not 
present in S. gutanensis. 


Solariorbis guianensis Altena, 1966 (Figures 73D-F) 


Solariorbis guianensis Altena, 1966. Zoologische Mededelingen, 41: 238-239, figs. 4a-c. [Type 


locality: Cupido, Maratakka River, Surinam]. 


(85 


Iberus, 29 (2), 2011 


Figures 73A-C. Solariorbis antillensis de Jong 8 Coomans, 1988. A-B: shell, 1.3 mm, Rancho 
Luna Beach (MHNS); C: protoconch. Figures 73D-FE. Solariorbis guianensis Altena, 1966. D-E: 
shell, 1.5 mm, Cienfuegos Bay (MHNS); F: protoconch. 

Figuras 73A-C. Solariorbis antillensis de Jong «* Coomans, 1988. A-B: concha, 1,3 mm, Playa 
Rancho Luna (MHNS); C: protoconcha. Figuras 73D-E Solariorbis guianensis Altena, 1966. D-E: 
concha, 1,5 mm, Bahía de Cienfuegos (MANS); E: protoconcha. 


Type material: Represented in ALTENA (1966, 1975). Not examined. 
Other material examined: Cuba: 2 s, Cienfuegos Bay, 20-30 m (MHNS). 


Description: ALTENA (1966: 238, figs. 
4a-c) figuring the holotype, presents a 
drawing without many details attempt- 
ing to depict the essential aspects of 
ornamentation: spiral pitted lines and 
axial ribs. DE JONG € COOMANS (1988: 
33) Temarked: “sculpture of the last 
whorl consisting of little pronounced 
and irregular fine radiating striae start- 
ing from the suture and the umbilicus, 
but not reaching the periphery and a 
few indistinct spirals, near the suture on 
the upper side and near the periphery 


Só 


on the under side;” pointing out the dis- 
tinguished characters of this species. 

We add: Shell (Figs. 73D-E) lenticu- 
lar, depressed, of 3 Y rapidly increasing 
whorls. Protoconch (Fig. 73F) smooth, 1 
Y whorl, measuring about 240 um; a 
strong varix is present at the transition 
to the teleoconch, on the first whorl of 
which there is a pitted reticular sculp- 
ture formed by the crossing of spiral 
cords and axial ribs. 

The principal distinguishing charac- 
ter is the ornamentation on the last 


RUBIO ET 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


whorl, formed by fine and irregular 
radial ribs which begin at the suture and 
disappear before reaching, the periphery, 
and spiral cordlets which are more 
evident near the suture and close to the 
umbilicus, keeping a central zone of the 
last whorl without sculpture. Base 
convex, without ornamentation, except 
for the growth lines. The umbilicus is 
narrow and deep, nearly obliterated by 
the thickened columella until the last Y 
of the body whorl, where it abruptly 
deviates laterally. Aperture rounded, 
external lip fine, internal lip thickened 
and reflected towards umbilicus. 

Dimensions: Holotype 0.90 mm in 
diameter. The figured shell measures 
1.47 mm in maximum diameter. 

Habitat: DE JONG € COOMANS (1988: 
33) mention that a significant number of 


specimens was found near Cabrieten- 
berg in Spaanse Water, always together 
with Teinostoma lerema. ln the area 
studied by us it is quite scarce and only 
2 specimens were found. 

Distribution: Known from Colombia 
(Díaz MERLANO é PUYANA HEGEDUS, 
1994), Aruba (DE JONG é COOMANS, 
1988), and Cuba: Cienfuegos. 

Remarks: Originally described as a 
fossil of the Holocene of Surinam. DÍAZ 
MERLANO é€ PUYANA HEGEDUS (1994) 
recorded it as live-collected in Colombia 
and DE JONG é€ COOMANS (1988) in 
Aruba. The shell presents the basic char- 
acters which distinguish this species 
from others included in the genus Solar- 
iorbis: the thickening of the umbilical 
wall and the finely pitted reticular 
sculpture. 


Solariorbis blakei (Rehder, 1944) (Figures 74A-D) 


Vitrinella blakei Rehder, 1944a. The Nautilus, 57: 97, pl. 9, figs. 1-2. [Type locality: Bed 1 (lowest 
bed) of the Talbot Formation, Wailes Bluff, near Cornfield Harbour, St. Mary's County, Vir- 


ginia]. 


Type material: Holotype in USNM (537834). Not examined. 

Material examined: Florida, USA: 4 s, E. St. Augustine, St. Johns Co., (CHL); 2 s, Cedar Key, Levy 
Co., 4.5-6 m shell /rubble bottom (CHL); 2 s, Stuart, Martin Co. (CHL); 2 s, Pinellas Co., (CHL); 1 
s, SE Mayport (CHL). Puerto Rico: 2 s, beach at Holiday Inn, San Juan (CHL). Turks € Caicos: 1 s, 


Grand Turk (CHL). 


Description: Shell (Figs. 74A-C) 
strongly depressed, small size, spire 
almost flat, very weakly sculptured, 
umbilicus nearly or entirely covered 
by the thickening of the umbilical 
wall. Protoconch (Fig. 74D) smooth 
with 2 whorls, size about 330 um in 
diameter and projecting slightly api- 
cally. Teleoconch with 1 % whorls, 
sculptured with  faint  spiral 
cords/grooves and short axial wrin- 
kles that radiate out from the suture. 
On the last half whorl the cords are 
weakened and the axial ribs almost 
disappear, persisting only in the sub- 
sutural. area. The base. 18. broad, 
smooth, and evenly rounded. Aper- 
ture oblique with a heavy parietal 
callus. Columella thickened, reflected 
towards the umbilicus, forming a 


small callus. Umbilicus reduced to a 
very narrow fisure, limited, and par- 
tially occluded by the thickening of 
the umbilical margin although the col- 
umella deviates laterally in the last Y 
whorl. On the umbilical wall strong 
growth folds can be seen. 

Dimensions: About 1.5 mm in 
maximum diameter. 

Habitat: It lives in creeks and bays 
(ANDREWs, 1977). Bathymetric range: 0 
to 26 m. 

Distribution: This species was con- 
sidered a continental species with an 
extensive distribution along the east 
coast of North America. It had never 
been reported from any of the West 
Indian islands (MOORE, 1964), but 
actually there are records from a few 
islands. USA: South Carolina, Florida: 


137 


Iberus, 29 (2), 2011 


Figures 74A-D. Solariorbis blakeí (Rehder, 1944). A-C: shell, 0.8 mm, Holiday Inn, Puerto Rico 
(CHL); D: protoconch. 
Figuras 74A-D. Solariorbis blakei (Rehder, 1944). A-C: concha, 0,8 mm, Holiday Inn, Puerto Rico 
(CHL); D: protoconcha. 


East Florida, West Florida; Mississippi; 
Texas; Mexico: unlocalized; Costa 
Rica, Panama. Reported from Colón 
and Bocas Island, Panama (OLssonN € 
MCGINTY, 1958). From Florida, Missis- 
sippi, Texas, Oregon, and Campeche 
Bank, Mexico (MOORE, 1964). From 
South Carolina to the Gulf Mexico and 
the Caribbean Sea (HOUBRICK, 1968). 
From South Carolina to Texas and the 
Caribbean Sea (ABBOTT, 1974); from 
the states of the Gulf to Mexico 
(EMERSON é JACOBSON, 1976). East 
coast of the USA; Florida; states of the 


138 


Gulf of Mexico; Mexico (ANDREWS, 
1977). Portete, Costa Rica (HOUBRICK, 
1968; ROBINSON € MONTOYA, 1987). 
From Martin, Palm Beach, Pinellas, 
and Levy Cos., Florida; Texas and 
Puerto Rico (ODÉ, 1988). From Lee Co. 
(GUNDERSEN, 1998). From SE Mayport 
and St. Augustine, Florida (LEE, 2009). 

Remarks: Described as a Pleis- 
tocene fossil. S. blakei like S. semi- 
punctus is smaller than other West 
Indian Solariorbis. S. blakei can be dis- 
tinguished from S. semipunctus by the 
rounded and smoother dorsal surface. 


RUBIO £7 AZz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Solariorbis elegans (Olsson £ McGinty, 1958) (Figures 75A-G) 


Vitrinella (Striovitrinella) elegans Olsson € McGinty, 1958. Bulletins of American Paleontology 
39: 31, pl. 3, figs. 1-1d. [Type locality: Bocas Island, Panama]. 


Type material: Holotype in ANSP (211877). Not examined. 
Material examined: Panama: 3 s, Bocas Island, drift (CHL); 2 s, Portobello, drift (CHL). 


Description: Shell (Figs. 75A-E) 
white or glassy, depressed, about 3 Y 
whorls. The protoconch (Figs. 75F-G) is 
projected over the teleoconch, has a 
little more than one whorl, smooth, 
and about 400 um in maximum diame- 
ter. In some shells two strong varices 
can be seen, which mark the two stages 
of development. Teleoconch totally 
covered by spiral cords of similar size, 
which at the crossing points with the 
axial lines form small rectangles, more 
evident and rounded on the dorsum 
and on the base of the teleoconch, and 
a little more elongate on the periphery. 
There are more than 35 spiral cordlets 
on the last whorl. Aperture rounded, 
oblique, external lip fine, columella 
strong and reflected towards the 
umbilicus, forming a small callus. 
Umbilicus deep; the peristome sharp, a 
little thickened near the base. Inner 
wall of the umbilicus with growth 
folds. 

The largest shell examined mea- 
sures 2.0 mm in maximum diameter 
and 1.1 mm in height. 

Habitat: Sand in shallow water 
(Díaz MERLANO éz PUYANA HEGEDUS, 
1994). Depth: 0 m (live 0 m). 

Distribution: Costa Rica, Panama, 
Colombia, Venezuela: unlocalized. 


Reported from Colón and Bocas 
Island, Panama (OLsson é€ MCGINTY 
1958). Central América (HOUBRICK, 
1968). Western Caribbean (ABBOTT, 
1974). Portete and Moin, Costa Rica 
(HOUBRICK, 1968; ROBINSON € 
MONTOYA 1987). From Venezuela 
(BRINEZ, 1982). From Costa Rica to 
Colombia (Díaz MERLANO é PUYANA 
HEGEDUS, 1994). 

Remarks: Described in the sub- 
genus  Striovitrinella Olsson € 
McGinty, 1950; this is characterized by 
having the entire surface of the shell 
sculptured with fine, threadlike 
spirals, operculum circular, thin, chiti- 
nous, with numerous, slowly enlarging 
spiral whorls; radula taenioglossate. 
Based on the thickening of the umbili- 
cal wall due to the enlargement and 
reflecting towards the umbilicus of the 
columella, the columellar callus, and 
the pits formed by the crossing of 
spiral cords and growth folds, we have 
decided to place this species in Solari- 
orbis. 

Solariorbis elegans is very similar 
to. Solariorbis . multistriatus, from 
which it can be distinguished by the 
shape of the umbilical callus, being 
less globose, and because the proto- 
conch is more erect. 


Solariorbis infracarinatus (Gabb, 1881) (Figures 76A-H, 77A-C) 


Adeorbis infracarinata Gabb, 1881. Journ. Acad. Nat. Sci. Philadelphia, 2 serie, 8: 365, pl. 46, 
fig. 62 [Type locality: Pliocene beds of Limon, Costa Rica]. 

Teinostoma bartschi Vanatta, 1913. Proc. Acad. Nat. Sci. Phil., 65: 26-27, pl. 2, figs. 9-11. 
[Type locality: Porto Barrios and Livingston, Guatemala]. 

Solariorbis euzonus Pilsbry € McGinty, 1950. The Nautilus, 63: 85, pl. 5, figs. 7-7a. [Type 
locality: Sebastian, Indian River County, Florida]. 


Type material: Holotype of Adeorbis infracarinata in ANSP (3380) and figured by MOORE 
(1965). Holotype of Teinostoma bartschi in ANSP (76501) figured by VANATTA (1913). Not 


examined. 


189 


Iberus, 29 (2), 2011 


Figures 75A-G. Solariorbis elegans (Olsson 8 McGinty, 1958). A-B: shell, 2.0 mm, Portobello, 
Panama (CHL); C-E: shells, 1.5, 1.4, 1.5 mm, Bocas Island, Panama (CHL); F-G: protoconchs. 
Figuras 75A-G. Solariorbis elegans (Olsson e McGinty, 1958). A-B: concha, 2,0 mm, Portobello, 
Panama (CHL); C-E: conchas, 1,5, 1,4, 1,5 mm, Isla Bocas, Panama (CHL); F-G: protoconchas. 


Other material examined: Guatemala: 2 s, Livingston, 3 m (MHNS). Brazil: 1 s, Itaparica (MHNS); 
1 s, Praia de Itararé, Sáo Vicente, Sáo Paulo (CHL). Florida, USA: 7 s, Cedar Key (CHL); 1 s, Bahia 
Honda Key, Monroe Co. (CHL); 1 s, Marco Island, Collier Co. (CHL); 1 s, SE Mayport (CHL); 2 s, 
APAC Pit, Sarasota, Plio-Pleistocene (CHL). 


140 


RUBIO ET 4L£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 76A-H. Solariorbis infracarinatus (Gabb, 1881). A-C: shell, 1.4 mm, Livingstone, 
Guatemala (MHNS); D: protoconch; E-H: form with axial ribs, 1.7, 1.8, 2.0 mm, Florida, USA 
(CHL); H: protoconch. 
Figuras 76A-H. Solariorbis infracarinatus (Gabb, 1881). A-C: concha, 1,4 mm, Livingstone, Guate- 
mala (MHNS); D: protoconcha; E-H: forma con costillas axiales, 1,7, 1,8, 2,0 mm, Florida, USA 
(CHL); H: protoconcha. 


141 


Iberus, 29 (2), 2011 


Figures 77A-C. Solariorbis infracarinatus (Gabb, 1881). A-B: shell, 1.7 mm, Itaparica, Brazil 


(MHNS); C: protoconch. 


Figuras 77A-C. Solariorbis infracarinatus (Gabb, 1881). A-B: concha, 1,7 mm, lItaparica, Brazil 


(MHNS); C: protoconcha. 


Description: Shell (Figs. 76A-C, 77A- 
B) depressed, rounded spire, carinate at 
the periphery, and narrowly umbilicate. 
Protoconch (Fig. 76D) of nearly 1 Y 
whorls and about 340 um in diameter. 
Teleoconch of about 2 whorls with a 
strong Carina near the periphery. 
Between this and the umbilicus there 
are 3 spiral cords. On the dorsum, 
between the suture and the periphery 
there are numerous axial ribs. A strong 
cord borders the umbilicus internally, 
thickening slowly towards the aperture 
and forming a callous lobe on the col- 
umellar margin typical of the genus. 

Habitat: Species common in shallow 
water in coastal bays. Amply distributed 
in continental waters. Not recorded in 
any island of the West Indies (MOORE, 
1964). 

Distribution: It has been recorded 
from Florida to Texas (USA), Campeche 


142 


Bank (Mexico), Guatemala and Panama 
(MOORE, 1964). From the USA, North 
Carolina to Gulf of Mexico and 
Caribbean Sea (HouBrick, 1968). On 
both sides of Florida; from Texas to 
Campeche Bank; Central America 
(ANDREWS, 1977). From Campeche to 
Ciudad del Carmen and Zacatal, Mexico 
(Vokes éz VOKES, 1984). From Florida 
(Lyons, 1989). South of Florida; Texas; 
Caribbean Sea (ROBINSON, 1991). Florida 
and Texas to Colombia (Díaz MERLANO 
éz PUYANA HEGEDUS, 1994). Rio Grande 
do Norte, Brazil (Rios, 1994). Our mater- 
ial is from Livingston, Guatemala. Also 
Florida Plio-Pleistocene: record from 
Chesapeake Bay, Virginia by Wass 
(1972: 125) needs confirmation. 
Remarks: MOORE (1964) consid- 
ered Teinostoma bartschi Vanatta, 
913 “and. SolariorbisteuzonWs 
Pilsbry € McGinty, 1950 synonyms 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


of Solariorbis infracarinatus not 
finding important differences. In 
spite of this, some databases treat 
T. bartschi as a valid species. The 
figured shell (Figs. 77A-C) was col- 
lected in Livingston, Guatemala, its 
type locality. 

PiLsBRY (1953) figured a shell of 
Solariorbis euzonus from the Plio- 
Blerstocene ot St. Petersburg, in 
which he distinguished 2-3 fine but 
easily-seen spiral cordlets on the 
dorsum, above the keel, as well as 
fine axial ribs on the first whorl of 
the teleoconch, which fade on the 
last whorl. Both these features can 
be found in well-preserved speci- 
mens of T. bartschi and S. infracari- 
nata. 


VANATTA (1913) and PILSBRY é 
MCGINTY (1950), in describing T. 
bartschi and 5. euzonus. respec- 
tively, did not compare these 
species ito Eos IMracanimarus 
described by GABB (1881). Based on 
the preceding analysis, and consis- 
tent with MOORE (1964), we con- 
sider lc vartschtiand 5. guzonus 
junior synonyms of S. infracarina- 
tus. 

In reference to S. infracarinatus, 
MOORE (1964) stated: “No other 
Solariobis in the western Atlantic 
has the combination of low round 
spure, rearal waves on the first 
whorl and spiral sculpture which is 
not visible from above”. We agree 
with this diagnosis. 


Solariorbis mooreanus (Vanatta, 1904) (Figures 78A-F) 


Vitrinella mooreana Vanatta, 1904. Proc. Acad. Nat. Sic. Philadelphia, 55: 757, figs. 1-3. [Type 
locality: Gulf side of Crooked Island near Panama City, Florida]. 

Solariorbis bassilisus Pilsbry, 1953. Monographs of Acad. Nat. Sci. Philadelphia, 18: 420, pl. 56, 
figs. 4-4c. [Type locality: Pliocene of St. Petersburg, Florida]. 


Type material: Five syntypes of Vitrinella mooreana in ANSP (84611). Holotype of Solariorbis bassil- 
isus deposited in ANSP (18408). Not examined. 

Other material examined: Cuba: 3 s, Cienfuegos Bay, 20-30 m (MHNS). Florida, USA: 1 sp, St. Joe 
Bay, grass flats (CHL). Brazil: 1 s, Praia de Itararé, Sáo Vicente, Sáo Paulo (CHL); 1 s, off Santos, 
Sáo Paulo State, trawled by nets at 100 m (CHL). 


Descrip tion: Shell (Figs. 78A-C, 78E- 
E) depressed, with six spiral ridges on 
dorsal of the shell. The peripheral cord 
is very prominent, like a keel. Base 
smooth, umbilicus narrow and deep. 
The protoconch (Fig. 78D) is projected 
over the teleoconch, with 2 spiral 
whorls, smooth and with about 350 um 
in maximum diameter. The teleoconch 
has two spiral whorls; the periphery is 
keeled by a peripheral cord, sharp and 
prominent, and strong cords on the 
dorsum. Ornamentation formed by 
punctiform sulcus, dorsally as well as 
ventrally. Base slightly concave. Aper- 
ture rounded, without canal on the 
upper internal angle. Columella thick- 
ened, with an expansion which from the 
internal lip is reflected outward forming 


a small triangular callus which closes 
the umbilicus gradually. 

Dimensions: Holotype of Vitrinella 
mooreana is 1.08 mm in diameter and 
1.2 mm of height. There are records up 
to 2.75 mm. Holotype of Solariorbis 
basilissus is 2.4 mm in diameter and 
1.25 mm in height. 

Animal and radula unknown. 

Habitat: The species has been found 
in a shell grit bottom at 9 m. According 
to MOORE (1964) it occurs on sandy 
bottoms in turbid waters. 

Distribution: Recorded from Gulf 
side of Crooked Island near Panama 
City, Florida (VANATTA, 1904); from Mis- 
sisippi (ANDREWS, 1977); from East and 
West Florida (Lyons, 1998); from Texas 
(ODÉ, 1988a); from Florida, Texas, north- 


143 


Iberus, 29 (2), 2011 


Figures 78A-E. Solariorbis mooreanus (Vanatta, 1904). A-C: shell, 1.9 mm, Cienfuegos Bay 
(MHNS); D: protoconch; E-F: shell, 2.6 mm, Itararé, Sao Vicente, Brazil (CHL). 
Figuras 78A-E Solariorbis mooreanus (Vanatía, 1904). A-C: concha, 1,9 mm, Bahía de Cienfuegos 
(MHNS); D: protoconcha; E-F: concha, 2,6 mm, Itararé, Sáo Vicente, Brasil (CHL). 


east Brazil, and Abrolhos Islands (Rios, 
1994). Also from Cienfuegos, Cuba. 
Remarks: This species is also 
known as a fossil. Solariorbis moore- 
anus is different from the congeneric 
species by the spiral cords which cover 
the dorsum of the shell, the sharp and 


144 


prominent peripheral cord which 
forms a keel around the shell, the 
spiral sulcus with punctiform incisions 
which cover the interspaces between 
the cords, and the triangular callus 
originating at the internal lip, which 
impinges on the umbilicus as it grows. 


RUBIO ET 4L.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Solariorbis multistriatus (A.E. Verrill, 1884) (Figs. 79A-I) 


Ethalia multistriata A.E. Verrill, 1884. Trans. Connect. Acad. Arts and Sci, 6: 242-243. [Type local- 
ity: Albatross sta. 2109, off Cape Hatteras, North Carolina]. 


Type material: Lectotype in USNM (35733). Not examined. 
Other material examined: Cuba: 5 s, Cienfuegos Bay, 20 m (MHNS); 2 s, Rancho Luna Beach, 20 m 


(MHNS). 


Description: Shell (Figs. 79A-E) 
strong, spire a little elevated, formed 
by 3 Y whorls. Protoconch (Figs. 79G- 
[) relatively large, bulbous, measuring 
about 480 um and with a little more 
than one whorl; two stages are distin- 
guised, the first one with a rough 
surface and the second with an 
evident reticulation and between one 
and seven strong varices on its dorsal 
aspect. Teleoconch with 2 Y whorls, 
totally covered by spiral cords of 
similar size, more than 50 on the body 
whorl. Between the cords there are 
spiral punctiform sulci (Fig. 79F) 
formed by the intersections with axial 
ribs. The pitting is more evident on 
the first whorl of the teleoconch, being 
barely detectable on the body whorl. 
Due to the thickening of the umbilical 
margin a characteristic triangular 
callus is formed and a gradual closing 
of the umbilicus is produced. 

Dimensions: Lectotype 1.59 mm in 
diameter. The larger shell figured is 
3.1 mm of maximum diameter. 

Animal and radula are unknown. 

Habitat: Species with a large 
bathymetric distribution, recorded 
between 5 and 312 m deep. Our shells 
were collected on coralline sandy 
bottom at 20 m. The type material was 
obtained at 142 fms (312 m). 

Distribution: Recorded off Cape 
Hatteras (A.E. VERRILL, 1884); from 
Cuba: North Havana Province 
(AGUAYO é JAUME, 1936); North Car- 


olina to West Indies (ABBOTT, 1974); 
from Bocas Island, Panama (OLssoN éz 
MCGINTY, 1958); from the Yucatan 
Peninsula, Mexico (VOKES € VOKES, 
1984); from the northeast Gulf of 
Mexico (PARKER 6 CURRAY, 1956); 
from Cape Lookout, North Carolina 
(MOORE, 1964); from Laguna de 
Termino, Campeche, Mexico (GARCÍA- 
CUBAS, 1982); from Puerto Rico and 
the Lesser Antilles (WARMKE € 
ABBOTT, 1961); from Guyana (PRINCZ, 
1977); from Venezuela (Princz, 1982). 
Also Cienfuegos, Cuba. 

Remarks: S. multistriatus was 
described by A.E. VERRILL (1884) from 
a juvenile specimen, which had not 
yet developed all the adult concholog- 
ical characters, and perhaps for this 
reason he placed the species in the 
genus Ethalia. BusH (1897) placed the 
species in Vitrinella due to the appar- 
ent similarity of its shell to V. heli- 
coidea, type species of that genus. 
MOORE (1964) moved it to Solariorbis 
after studying an adult specimen 
showing the thickening of the umbili- 
cal wall and pitted sulcus. He also 
mentioned that S. multistriatus is 
similar to S. terminalis, from which it 
can be differentiated by its smaller 
size, its higher spire and smaller 
umbilical callus. 

In our opinion the number and 
regularity of the spiral cords and the 
triangular callus are the main distin- 
guishing characters of this species. 


Solariorbis truncatus (Gabb, 1881) (Figures 80A-C) 


Vitrinella truncata Gabb, 1881. Journ. Ac. Nat. Sci.: 367, pl. 47, fig. 65. [Type locality: Limon, 


Costa Rica, Miocene]. 


Solariorbis corylus Olsson £ McGinty, 1958. Bulletins of American Paleontology, 39: 28, pl. 3, 


figs. 4-4b. [Type locality: Bocas Island, Panama]. 


145 


Iberus, 29 (2), 2011 


Figures 79A-1. Solariorbis multistriatus (A.E. Verrill, 1884). A-E: shells, 2.5, 2.6, 3.1, 2.56, 2.85 
mm, Cienfuegos Bay; F: microsculpture; G-Í: protoconchs. 

Figuras 79A-1. Solariorbis multistriatus (4.E. Verrill, 1884). A-E: conchas, 2,5, 2,6, 3,1, 2,56, 2,85 
mm, Bahía de Cienfuegos; E: microescultura; G-Í: protoconchas. 


Type material: V. truncata: Type in ANSP (3107). Holotype of S. corylus (ANSP 211909) and a 
paratype in ANSP (211910). Not examined. 
Other material examined: 1 s, Portobello, Panama (CHL). 


146 


RUBIO ET ALz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 80A-C. Solariorbis truncatus (Gabb, 1881). A-B: shell, 1.66 mm, Portobello, Panama 
(CHL); C: protoconch. 
Figuras 80A-C. Solariorbis truncatus (Gabb, 1881). A-B: concha, 1,66 mm, Portobello, Panamá 
(CHL); C: protoconcha. 


Description: Shell (Figs. 80A-B) 
small, white or glassy, with a 
strongly flattened, depressed spire 
bounded by a ridged shoulder, the 
profile of the body whorl below the 
shoulder being wider and evenly 
convex. Protoconch (Fig. 80C) with 1 


(ROBINSON é MONTOYA, 1987; 
ROBINSON, 1991). From Colón and 
Colón and Bocas Island, Panama 
(OLSSON é MCGINTY, 1958). From 
southeastern Panama  (RADWIN, 
1969). From Panama and Colombia 
(DÍAZ MERLANO $ PUYANA 


14 whorls. 
Maximum reported size: 2.4 mm 
Habitat: Sandy bottom (DÍAZ 
MERLANO 6 PUYANA HEGEDUS, 1994). 
Distribution: Recorded from the 


HEGEDUS, 1994). 

Remarks: OLSssON é MCGINTY 
(1958) stated: “Similar to S. contracta 
(Vanatta) from Monkey River, Hon- 
duras but lacks the central or periph- 


Pleistocene, Moín, Costa Rica eral keel and has a higher shell”. 


Solariorbis schumoi (Vanatta, 1913) (Figs. 81 A-D) 


Teinostoma schumoi Vanatta, 1913. Proc. Acad. Nat. Sci. Philadelphia, 65: 25-26, pl. 2, figs. 5, 10. 
[Type locality: Porto Barrios and Livingston, Guatemala]. 


Type material: Holotype in ANSP (73483) figured by VANATTA (1913). Not examined. Best repre- 
sentations in ALTENA (1966: 240-241, figs. 6a-e) and ALTENA (1975: figs. 11a-c). 

Other material examined: Guatemala: 2 s, Livingston, 3-4 m (MHNS). Brazil: 1 s, Praia Itararé, Sáo 
Vicente, Sáo Paulo, drift (CHL). 


147 


Iberus, 29 (2), 2011 


Figures 8lA-D. Solariorbis schumoi (Vanatta, 1913). A-B: shells, 0.8, 1.1 mm, Livingston, 


Guatemala (MHNS). C-D: shell, 1.8 mm, Praia Itararé, Sáo Vicente, Brazil (CHL). 
Figuras 8LA-D. Solariorbis schumoi (Vanatta, 1913). A-B: conchas, 0,8, 1,1 mm, Livingston, Guate- 
mala (MHNS). C-D: concha, 1,8 mm, Praia ltararé, Sáo Vicente, Brasil (CHL). 


Description: Shell (Figs. 81A-D) 
small, imperforate in adult individu- 
als, white, very compact, suture 
marked, rounded periphery, with 
undulant profile due to the spiral 
cords; the penultimate whorl exhibits a 
spiral sulcus near the suture. Proto- 
conch of nearly 1 % whorls and about 
340 um in diameter. Teleoconch with 2 
YM whorls, with an ornamentation 
formed by strong spiral cords and fine 
spiral threads, strong axial ribs, and 


148 


fine axial striae; the spaces between the 
cords are convex and are crossed by a 
fine axial striation. On the dorsum 
there is a strong nodulous cord pro- 
duced by the intersection with axial 
ribs, the latter progressively more 
evident as the shell grows. There are 5- 
7 spiral cords on the periphery. Near 
the periphery of the base there are two 
spiral cords. Strong growth folds 
extend from the basal cords towards 
the inner part of the umbilicus. Aper- 


RUBIO £T AZz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


ture suborbicular; external lip sharp, 
parietal area wide. Columella and 
inner lip. are reflected .outward, 
forming a wide callus which increases 
the umbilical wall and may close 
totally the umbilicus in adult shells. 
This character is fundamental in the 
placement of the related species, T. 
hondurasensis, in Solariorbis. 

The holotype measures 2.23 mm in 
diameter and 1.51 mm in height. The 
largest shell figured measures 1.2 mm 
in diameter and 0.71 mm in height, 
and it is a non-adult individual. 

Habitat: It lives in shallow water at 
about 2 meters depth. 

Distribution: Known from Guate- 
mala (WANATTA 19138) Costa Rica, 
Panama and Colombia (CoseEL, 1986); 
ABC Islands: Aruba (DE JONG é 


COOMANS, 1988); Brazil: Pernambuco, 
Sáo Paulo (MELLO € PERRIER, 1986); 
Moin, Costa Rica (ROBINSON  éz 
MONTOYA, 1987); Surinam (ALTENA, 
1966) 

Remarks: This species was originally 
included in the genus Teinostoma. 
MOORE (1964) placed them in Solari- 
orbis on the basis of the enlarge- 
ment of the umbilical wall and the 
pitted spiral sulcus in the original 
feuresiot 1. schuniol andsI. hon- 
durasensis. T. schumoi may be dif- 
ferentiated from T. solidum Smith 
by the sculpture on the dorsum; 
from S. hondurasensis Vanatta, 1913 
it may be differentiated by having 
more spiral cords and being umbili- 
cated as juvenile and imperforate, 
or nearly so, as an adult. 


Solariorbis semipunctus Moore, 1965 (Figures 82A-D) 


Solariorbis semipunctus Moore, 1965. The Nautilus, 78: 77-78, pl. 8, figs. 1-3. [Type locality: 


Northwest Campeche Bank, Mexico]. 


Type material: Holotype in USNM (636309) figured by MOORE (1965). Not examined. Best repre- 


sentation in FABER (2007). 


Other material examined: Cuba: 3 s, Rancho Luna Beach, 20 m (MHNS); 2 s, Cienfuegos Bay, 20- 


30 m (MHNS). 


Description: Shell (Figs. 82A-C) 
strongly depressed, 3 Y whorls, spiral 
sculpture punctiform with a rounded 
periphery and umbilicus sealed by 
callus. Protoconch (Fig. 82D) smooth 
with nearly 1 Y whorls and about 200 
um in diameter. Teleoconch of about 1 
% whorls; fine spiral punctiform 
cordlets cover the whole surface. In 
the middle of the dorsum and after 
the first Y whorl, the shell is keeled. 
The umbilicus of the adult shells is 
totally occluded by the columellar 
callus, which, in the form of a fine 
callous surface, also covers the 
cordlets close to the umbilicus. Base 
somewhat convex. 

Dimensions: Holotype 0.93 mm in 
diameter and 0.4 mm of height. The 
material studied and figured is 1.4 
and 1.6 mm in diameter and 0.6 mm 


in height. 


Animal and radula unknown. 

Habitat: The species lives on a 
muddy bottom at 18 m deep 
(MOORE, 1965). Our samples are 
from sediments collected on a 
coralline bottom between 25 and 50 
mA: 

Distribution: Recorded from 
Yucatan State, Mexico and Haiti 
(MOORE, 1965); from Campeche 
Bank, Mexico (18 m) and Haiti 
(ABBOTT, 1974); from East and West 
Florida (Lyons, 1998); from Florida: 
Virginia Key and  Schottegat, 
Curacao (FABER, 2007) and from 
Cuba: Cienfuegos. 

Remarks: S. semipunctus may be 
distinguished from its congeners by 
the strongly depressed spire, dorsal 
keel, and the umbilicus being totally 
occluded by the callus formed from 
the inner lip. 


149 


Iberus, 29 (2), 2011 


Figures 82A-D. Solariorbis semipunctus Moore, 1965. A-C: shells, 1.4, 1.6, 1.3 mm, Rancho Luna 
Beach (MHNS); D: protoconch. 

Figuras 82A-D. Solariorbis semipunctus Moore, 1965. A-C: conchas, 1,4, 1,6, 1,3 mm, Playa 
Rancho Luna (MANS); D: protoconcha. 


Solariorbis terminalis (Pilsbry £ McGinty, 1946) (Figures 83A-E, 84A-F) 


Vitrinella terminalis Pilsbry £ McGinty, 1946. The Nautilus, 60: 17, pl. 2, fig. 5-5a. [Type locality: 
Destin, northwestern Florida]. 


Type material: Type species in ANSP (181883). Not examined. 

Material examined: Florida, USA: 1 s, Tarpon Springs, Pinellas Co., dredged S of Anclote Key, in 
3.5-6 m (CMK); 2 s, 3018.13'N 81%06.91'W, 24 m, off Mayport, Duval Co. (CHL); 1 s, off Fort Myers 
Beach, Lee Co., trawled 20 m (CHL); 2 s, 50-60 mi E Ponte Vedra, St. Johns Co., trawled at 39-45 m 
(CHD); 1 s, 10 mi SW Seahorse Key, Cedar Keys, Levy Co., trawled at 18 m (CHL); 1 s, St. Augus- 
tine, St. Johns Co., tailings scallop processing plant (CHL); 2 s, drift, Jacksonville Beach, Duval Co. 


(CHL). 


Description: From the original 
description (PILSBRY €£ MCGINTY, 1946) 
we have copied some paragraphs which 
express important characters of the 
species: “There are 4 Y convex whorls 
joined by a rather strongly impressed 
suture. Last whorl is rounded at periph- 
ery and base. Sculpture of a few very 
weak but coarse spirals in the peripheral 
region: Mer columellaimarinis 
extremely thick, reflected over part of 
the umbilicus”. 


150 


The low conic shell (Fig. 83A-C, 84A-C) 
is relatively thick and heavy, and has a very 
small constricted umbilicus. Protoconch of 
2 smooth glassy whorls. Teleoconch has 
two whorls. The spire is elevated and 
allows all the whorls to be seen in apertural 
view. Suture impressed. The sculpture con- 
sists Of several low spiral threads on the 
peripheral area. The base is broad and 
smooth except for spiral sculpture on the 
outer part. Aperture circular; columella 
thick and heavy (MOORE, 1965: 117-118). 


RUBIO £7 AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 83A-E. Solariorbis terminalis (Pilsbry 82 McGinty, 1946). A-D: shell, 1.7 mm, Tarpon 
Spring, Pinellas Co. (CMK); D: protoconch eroded; E: detail of the spire. 

Figuras 83A-E. Solariorbis terminalis (Pilsbry 8 McGinty, 1946). A-D: concha, 1,7 mm, Tarpon 
Spring, Pinellas Co. (CMK); D: protoconcha erosionada; E: detalle de la espira. 


151 


Iberus, 29 (2), 2011 


Figures 84A-E Solariorbis terminalis (Pilsbry 8 McGinty, 1946). A: shell, 2.5 mm, off Fort Myers 
Beach, Lea Co. (CHL); B-C: shells, 2.3, 1.1 mm, off Mayport, Duval Co. Florida, 20 m (CHL); D- 
E: shells, 1.9, 1.8 mm, 50-60 mi E Ponte Vedra, St. Johns Co. (CHL); F: protoconch. 

Figuras 84A-E Solariorbis terminalis (Pilsbry e McGinty, 1946). A: concha, 2,5 mm, frente a Fort 
Myers Beach, Lea Co. (CHL); B-C: conchas, 2,3, 1,1 mm, off Mayport, Duval Co. Florida, 20 m 
(CHL); D-E: conchas, 1,9, 1,8 mm, 50-60 millas al E de Ponte Vedra, St. Johns Co. (CHL); E: protocon- 


We add the following: The proto- 
conch (Figs. 83D, 84F) has about 2 
whorls, is smooth, and measures about 
350 um in diameter, lacking any thicken- 
ing in the transition to the teleoconch. 
The teleoconch is dorsally smooth, 
having a marked suture developing into 
a sulcus which progressively enlarges 


1132 


until the middle part of the body whorl, 
thence forming a subsutural cord on the 
final quarter whorl. On the periphery of 
the last whorl 3-4 spiral cords can be 
seen. Columella wide, reflected towards 
the umbilicus, thickening the umbilical 
wall and subtotally closing the umbili- 
cus. Internal lip reflected outward. 


RUBIO £7 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Within the umbilicus there are 2-3 fine 
spiral cordlets. 

Dimensions: Maximum reported 
size: 2.4 mm 

Habitat: This species lives on sandy 
bottom in depths between 18 and 35 m. 

Distribution: USA: North Carolina 
(PORTER, 1974), West Florida (PILSBRY 
é MCGINTY, 1946b; MOORE, 1964); 
Panama (OLsson € MCGINTY, 1958). 
MOORE (1964) considered the distribu- 
tion of the species to be very limited, 
being restricted to  northwestern 
Florida, but he did not consider the 
record Of OLSSON € MCGINTY (1958) 
from Panama. ODÉ's (1973b) record 
from the northwest Gulf of Mexico, 
-must be confirmed, because he himself 
(ODÉ, 1988) did not again record this 
species. 


Remarks: PiLSBRY € MCGINTY (1946: 
17) stated: “The conic spire has more 
whorls and a deeper suture than in 
typical Vitrinellidae, and the columella 
is very heavily callused. The umbilicus 
is quite small up to the last half whorl of 
the umbilical suture, when it becomes 
rapidly much wider”. In our opinion, 
the subsutural sulcus on the last whorl, 
the peripheral cords, and the lack of 
dorsal and basal ornamentation, with 
the exception of the thickened umbilical 
margin, are the distinguishing charac- 
ters of the species. 

Vitrinella  diaphana  (d'Orbigny, 
1842) could be this species, but the type 
material is in such poor condition that 
this cannot be ascertained (see Fig. 
108G). It should be considered, there- 
fore, a nomen dubium. 


Solariorbis ruris spec. nov. (Figures 85A-J) 


Type material: Holotype (Fig. 85A) in MNCN (15.05/55058) and 1 paratype (Fig. 85B), from type 
locality, at -20 m (15.05/55059). Other paratypes: MHNS (100551, 1 s, Fig. 85C), at -20 m; MNHN 


(24397, 1 s, Figs. 85D-E), at -45 m. 


Other material examined: Martinique: 1 s, (CJP) (Figs. 85F-G). 

Type locality: Rancho Luna Beach, 20 m, Cienfuegos, Cuba. 

Etymology: The specific name is derived from the Latin third declension noun rus, ruris, meaning 
“of the farm; rural” in reference to its surface, in some areas with parallel lines, as in the surface of 


a plowed field. 


Description: Shell (Figs. 85E-G) of 
small size, depressed, solid, 4 whorls, 
with characteristic spiral threads and 
punctiform  grooves. Protoconch 
(Figs. 85H) of about 2 whorls, mea- 
suring about 290 um in diameter; its 
surface is covered by fine tubercles; a 
delicate varix is observed. Teleoconch 
of about 2 rapidly-increasing whorls; 
dorsally and ventrally convex; totally 
covered by spiral cordlets and pune- 
tiform incisions in the interspaces, 
forming the characteristic reticular 
punctiform sculpture. In the dorsal 
area of the last whorl, between the 
suture and the periphery, a central 
zone without spiral microsculpture 
can be observed; a similar area can be 
seen close to the umbilicus on the 
base of the last Y whorl. Axial orna- 
mentation consists of growth striae. 


Base slightly convex, umbilicus 
small, progresively occluded by the 
columellar callus. Aperture rounded, 
prosocline, external lip sharp, col- 
umellar margin and internal lip 
thickened, reflected towards the 
umbilicus forming a characteristic tri- 
angular callus. 

Dimensions: Holotype 1.50 mm in 
diameter and 0.57 mm in height. 

Animal unknown. 

Habitat: The species has been 
found on a coralline sandy bottom, 
sometimes with a little mud, at 45 m. 

Distribution: Only known from 
Cuba and Martinique. 

Remarks: Solariorbis ruris spec. 
nov. has a sculpture formed by spiral 
cordlets with punctiform incisions in 
the interspaces which covers all the 
shell except for a short band without 


153 


Iberus, 29 (2), 2011 


Figures 85A-J. Solariorbis ruris spec. nov. A: holotype, 1.3 mm, Rancho Luna Beach (MNCN); B: 
paratype, 1.4 mm (MNCN); C: paratype, 1.3 mm (MHNS); D-E: paratype, 1.3 mm (MNHN); 
E-G: shell, 1.5 mm, Martinique (CJP); H-I: protoconchs; H: from the shell of Fig. G; 1: from the 
holotype Fig. A; J: detail of the base (paratype Fig. B). 

Figuras 85A-J. Solariorbis ruris spec. nov. A: holotipo, 1,3 mm, Playa Rancho Luna (MNCN); Bb: 
paratipo, 1,4 mm (MNCN); C: paratipo, 1,3 mm (MANS); D-E: paratipo 1,3 mm, (MNAN); F-G: 
concha, 1,5 mm, Martinica (C]P); H-I: protoconchas; H: de la concha de la Fig. G; 1: del holotipo Fig. 
A; J: detalle de la base (paratipo Fig. B). 


154 


RUBIO E7 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


spiral microsculpture on the dorsum 
and base of the last whorl. It also 
possesses a triangular columellar 
callus. 

Solariorbis terminalis is the only 
species with somewhat similar orna- 
mentation, but the cords cover only 
the dorsum of the shell and are wider 
and less numerous. 


From Solariorbis lineopunctatus 
(see below) it may be distinguished 
by the fewer, thicker and spiral 
cordlets. 

From Solariorbis sp. (see below) it 
may be differentiated by its rounded 
convex periphery and the sulcus with 
punctiform incisions instead of quad- 
rangular hollows. 


Solariorbis lineopunctatus spec. nov. (Figures 86A-D) 


Type material: Holotype (Figs. 86A-B) in MNCN (15.05/55056) and one paratype in MNCN 


(15.05/55057). 


Type locality: Cienfuegos Bay, sta. 12, 22007'N 80%27'W, 9 m, Cuba. 
Etymology: The specific name refers to the punctiform sculpture aligned spirally which totally 


covers the shell. 


Description: Shell (Figs. 86A-B) of 
very small size, not very depressed, 
apparently not very solid; spire 
formed by three rapidly-increasing 
whorls, sligtly angled at the periph- 
ery, not keeled. Protoconch (Figs. 86C) 
of about 2 whorls, measuring about 
400 um in diameter, being placed on a 
plane slightly above the subsequent 
whorls and with its surface covered 
by small tubercles and very fine spiral 
threads. [In some places these form a 
fine reticule. Teleoconch with a little 
more than 1 rapidly-increasing whorl; 
convex dorsally as well as on the base, 
periphery angled; surface totally 
covered by punctiform incisions spi- 
rally aligned except inside the umbili- 
cus, where only numerous growth 
marks are appreciated. Aperture 
rounded, without a sulcus in the inner 
upper angle. Outer lip sharp. Col- 
umella and inner lip thickened and 
reflected outward. Umbilicus rela- 


tively wide and deep, without evident 
ornamentation except for the axial 
growth marks. 

Dimensions: Holotype 1.05 mm in 
diameter, and 0.36 mm in height. 

Animal unknown. 

Habitat: The species was found in 
depths between 10 and 60 m, on 
coralline and slightly muddy bottoms. 

Distribution: Only know from 
Cuba. 

Remarks: The shell of Solariorbis 
lineopunctatus spec. nov. is not very 
strong, with a fragile aspect, reddish 
color, and dull in luster. Its teleo- 
conch, totally covered with spirally- 
aligned punctiform incisions makes it 
easily recognizable and distinguishes 
it from its congeners. Its ornamenta- 
tion is rather similar to that present in 
some species of the genus Teinostoma 
(T. goniogyrus, T. lenticulare), but the 
wide umbilicus and the thickening of 
the umbilical border are distinctive. 


Solariorbis sp. (Figures 86E-F) 


Material studied: One shell (Figs. 86E) off Dominica, 200 m, in detritus obtained from inside sub- 


merged bottles (lost during the study). 


Description: Shell (Fig. 86E) tro- 
choid, solid, not very depressed, of dirty 
white color. Protoconch with about 2 


spiral whorls on a little higher plane 
than the later whorls. Teleoconch 
formed by about 2 whorls totally 


SS 


Iberus, 29 (2), 2011 


Figures 86A-D. Solariorbis lineopunctatus spec. nov. A-B: holotype, 1.05 mm, Cienfuegos Bay 
(MNCN); C: protoconch; D: microsculpture. Figures 86E-E. Solariorbis sp. E: shell, 1.47 mm, off 
Dominica; F: microsculpture. 

Figuras 86A-D. Solariorbis lineopunctatus spec. nov. A-B: holotipo, 1,05 mm, Bahía de Cienfuegos 
(MNCN); C: protoconcha; D: microescultura. Figuras 86É-E Solariorbis sp. E: concha, 1,47 mm, 
frente a Dominica; FE: microescultura. 


56 


RUBIO ET 4L£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


covered by spiral cords of equal size 
and axial sulci forming quadrangular 
spaces. Periphery only slighty convex, 
almost straight; a spiral cord in the 
dorsal extreme and another in the 
basal part form the limits and create 
slight angulations. Base slightly 
concave in its central part. Umbilicus 
small, almost occluded by a callus 
formed from a thickening of the col- 
umella. Aperture rounded, the supe- 
rior angle with a sulcus; parietal area 
straight and thickened, external lip 
sharp. Columella widened and 
reflected towards the external part 
forming a characteristic callus. 

Dimensions: The shell is 1.47 mm 
in diameter and 0.76 mm in height. 

Animal unknown 

Habitat: The only specimen known 
was collected in the shell grit found 
inside one of a few bottles found at 
200 m. 


Distribution: Only known from 
the island of Dominica. 

Remarks: The present shell may 
be distinguished from its congeners 
by its sculpture of spiral cordlets 
and cuadrangular hollows covering 
the shell, by the wide callus formed 
by the thickening of the columella, 
and principally by the dorsal and 
basal cords which angulate the 
periphery, giving it an almost 
straight profile. S. lineopunctatus n. 
sp. has narrower, more numerous 
spiral cords and the sulcus has punce- 
tiform incisions. From S. ruris n. sp. 
it can be separated by the latter 
having the smooth zones on the 
body whorl and its smaller, triangu- 
lar columellar. 

Unfortunately this shell was lost 
during the study, and we decided 
not to name this species until new 
material can be collected. 


Solariorbis punctostriatus spec. nov. Rubio, Rolán € Lee (Figures 87A-E) 


Type material: Holotype (Figs. 87A-B) deposited in FLMNH (448610)(ex CHL). One paratype from 
the type locality (CHL). Another paratype in USNM (1155036, 1 s, ex CHL, from Courland Bay, 


Tobago). 
Type locality: Isla Margarita, Venezuela. 


Etymology: The specific name refers to its microsculpture formed by spiral striae with punctiform 


depressions. 


Description: Shell (Figs. 87A-D) 
solid, depressed, about equally convex 
above and below, with rounded periph- 
ery and spiral striae throughout. Proto- 
conch of about 1 Y whorls, not project- 
ing upward. Teleoconch of about 2 
whorls, totally covered by spiral cords, 
in interspaces of which axial striae can 
be seen, forming characterístic small 
hollows. Rounded periphery. Aperture 
rounded, slightly prosocline, external lip 
a little thickened, columella thickened 
and reflected outward forming a small 
callus. Umbilical area concave. Umbili- 
cus almost closed by the thickening of 
the umbilical margin. 

Dimensions: Holotype 1.39 mm in 
diameter and 0.63 mm height. One 
paratype with about 1.4 mm in 
maximum dimension. 


Habitat: Unknown, the material 
was collected in sediments. 

Distribution: Only known from 
Isla Margarita, Venezuela, its type 
locality, and Courland Bay, Tobago. 

Remarks: Solariorbis punctostria- 
tus spec. nov. has a shell very similar 
in general appearance to that of V. 
cupidinensis, from which it can be 
distinguished by having the umbili- 
cus practically closed, almost reduced 
to a fissure, and by the columellar 
callus which thickens the umbilical 
wall and closes the umbilicus. Also it 
is similar to some Teinostoma species, 
from which it can be distinguished by 
the typical occlusion of the umbilicus 
by the thickening of the columella 
and by its typical Solariorbis sculp- 
ture. 


LSZ 


Iberus, 29 (2), 2011 


Figures 87A-E. Solariorbis punctostriatus spec. nov. Rubio, Rolán 8 Lee. A-B: holotype, 1.4 mm, 
Isla Margarita, Venezuela (FLMNH); C: paratype, 1.4 mm, from type locality (CHL); D: 
paratype, 1.2 mm, Tobago (USNM); E: detail of the ornamentation. 

Figuras 87A-E. Solariorbis punctostriatus spec. nov. Rubio, Rolán «* Lee. A-B: holotipo, 1,4 mm, Isla 
Margarita, Venezuela (FLMNA); C: paratipo, 1,4 mm, de la localidad tipo (CHL); D: paratipo, 1,2 
mm, Tobago (USNM); E: detalle de la ornamentación. 


158 


RUBIO ET 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


“Solariorbis” solidus spec. nov. (Figures 88A-F, 89A-F) 


Type material: Holotype (Fig. 88A) in MNCN (15.05 /55060). Paratypes: MNHN (24398, 1 s, Fig. 
88B), NHAMUK (1 s, Figs. 88E, 89A), MCZ (1 s, Figs. 89B-C), USNM (1155037, 1 s, Fig. 89E), all from 
type locality; AMNH (1 s, Fig. 88C) Cayo Diego Perez, Canarreos Archipelago, 20 m, Cuba; MHNS 
(100552, 1 s, Fig. 88D) Cienfuegos, 20-30 m. 

Other material examined: Cuba: 3 s, Canarreos (MHNS). Florida, USA: 1 s, Little Madeira Bay. Ever- 
glades N.P. Florida, Monroe Co., low tide (CHL). 

Type locality: Rancho Luna Beach, Cienfuegos, 20-45 m, Cuba. 

Etymology: The specific name refers to the solidity of the shell, more observable in the contour of 


the aperture. 


Description: Shell (Figs. 88A-E, 
89A-F) trochoid, solid, yellowish 
white, polished and shiny. Proto- 
conch (Fig. 88F) broad and flat, 
slightly projecting, with about 1 Y 
whorls, 410 um in diameter, and 
without sculpture. Teleoconch of 
about 1 Y. rapidly-increasing 


whorls; dorsally and  ventrally 
convex. The shell is generally 
smooth and without sculpture 


except for weak axial growth lines, 
which in some specimens are much 
more rough and sharp, and faint 
spiral cords on the basal periphery. 
Aperture oval, outer lip thick, col- 
umellar margin thickened and 
reflected outward. Umbilicus wide 
and deep, inside there is a thick 
cord formed by a thickening of the 
columella, which surrounds and 
limits the umbilical infundibulum. 

Dimensions: Holotype 1.3 mm in 
diameter, but there is a paratype 1.8 
mm in diameter. 

Animal unknown. 

Habitat: This species has been 
collected between 5 and 45 m, on a 
coralline sand bottom. 

Distribution: Only known from 
Cuba and Florida, USA. 


about it being a marine species. 
Therefore its generic placement in 
Solariorbis, and even in Tornidae, is 
provisional. We made a comparison 
with a paratype of Paludinella heli- 
coides “Gundlach” 1865 (MCZ), and 
our impresion is that the latter is 
not as solid. The dearth of concho- 
logical characters makes a proper 
comparison difficult. Paludinella 
helicoides is a species of fresh water 
hydrobiid considered endemic to 
Cuba, but due to its seldom being 
collected it has been recorded only 
a few times. Anyway, in our mater- 
ial there are other shells, collected 
in areas as distant as Cuba and 
Florida. We keep its provisional 
description while awaiting more 
material. 

Although the overall appearance 
of the shell, wide umbilicus and 
aperture, predisposes us to place it 
in Vitrinella, the columellar thicken- 
ing and the thickening of the umbil- 
ical wall are typical characters of 
the genus Solariorbis. 

Solariorbis solidus spec. nov. 
differs from provisional congeneric 
species by a trochoid form, its wide 
umbilicus, the lack of ornamenta- 


tion on the teleoconch, and its 
solidity in spite of its fragile 
appearance. 


Remarks: We are not totally sure 
that this species is a tornid, not 
even having complete security 


Genus Vitrinella C.B. Adams, 1850 


Vitrinella C.B. Adams, 1850. Monograph of Vitrinella, a new genus of species of Turbinidae. 10 
p. Amherst Massachusetts. 

Type species: (by original designation) Vitrinella helicoidea C.B. Adams, 1850 (by subsequent 
designation: Bush, 1897: 105. Caribbean, Recent. 


15% 


Iberus, 29 (2), 2011 


Figures 88A-F. *Solariorbis” solidus spec. nov. A: holotype, 1.3 mm, Rancho Luna Beach, Cuba 
(MNCN); B: paratype, 1.08 mm, Rancho Luna Beach (MNHN); C: paratype, 1.14 mm, 
Canarreos, Cuba (AMNH); D: paratype, 1.08 mm, Cienfuegos Bay (MHNS); E: paratype, 1.6 
mm, Rancho Luna Beach (NHMUK); FE: protoconch of the holotype. 

Figuras 88A-E “Solariorbis” solidus spec. nov. A: holotipo, 1,3 mm, Playa Rancho Luna, Cuba 
(MNCN); B: paratipo, 1,08 mm, Playa Rancho Luna (MNHN); C: paratipo, 1,14 mm, Canarreos, 
Cuba (AMNH); D: paratipo, 1,08 mm, Babía de Cienfuegos (MHNS); E: paratipo, 1,6 mm, Playa 
Rancho Luna (NHMUK); F: protoconcha del holotipo. 


160 


RUBIO ET ALz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 89A-E. “Solariorbis” solidus spec. nov. A: paratype, 1.6 mm, Rancho Luna Beach 
(NHMUK); B-C: paratype, 1.17 mm, Rancho Luna Beach (MCZ); D-E: shells, 1.2 mm, Cien- 
fuegos Bay; F: paratype, 1.8 mm, Rancho Luna Beach (USNM). 

Figuras 89A-E “Solariorbis” solidus spec. nov. A: paratipo, 1,6 mm, Playa Rancho Luna (NHMU, 
B-C: paratipo, 1,17 mm, Playa Rancho Luna (MCZ); D-E: conchas, 1,2 mm, Bahía de Cienfuegos, 
paratipo, 1,8 mm, Playa Rancho Luna (USNM).. 


Iberus, 29 (2), 2011 


Diagnosis: Shell small, fine, smooth 
or weakly sculptured, low spire and 
open, deep  umbilicus. Aperture 
rounded and only slightly modified by 
the previous whorl. Parietal callus 
usually thin. Animal provided with a 
pair of ciliated cephalic tentacles, a pair 
of epipodial tentacles, and a circular 
multispiral operculum. Radula tae- 
nioglossate. 


Remarks: C.B. ADAMS (1850) insti- 
tuted a new genus Vitrinella for five 
new species of micromolluscs found in 
beach sand from Jamaica. No type 
species was selected and none of the 
species was figured. BusH (1897) 
designed Vitrinella helicoidea as the 
type species of the genus, describing 
this species again and figuring it for first 
time. 


Vitrinella anneliesae de Jong € Coomans, 1988 (Figures 90A-I) 


Vitrinella anneliesae de Jong € Coomans, 1988. Marine gastropods from Curacao, Aruba and 
Bonaire: 31, pl. 2, fig. 131. [Type locality: Curacao]. 


Type material: Holotype in ZMA (3.87.062). Represented in de JONG £ COOMANS (1988). Notexam- 
ined. 

Other material examined: Cuba: 10 s, Canarreos Archipelago, 5 m (MHNS); 4 c, Cayo Avalos, 8 
m (MHNS); 4 c, Cayo Diego Perez, 15 m (MHNS); 2 c, Faro Diego Perez, 10 m (MHNS); 3 s, Jibacoa, 
3-6 m (MHNS); 3 c, Guajimico, 5 m (MHNS); 5 c, Cienfuegos Bay, 20-30 m (MHNS); 2 c, Cien- 
fuegos Bay, 30 m (MHNS); 14 s, Rancho Luna Beach, 12 m (MHNS); 37 s, 10-30 m (MHNS); 2 s, 
Rancho Luna Beach, 35 m (MHNS); 20 s, Rancho Luna Beach, 45 m (MHNS); 106 s, Rancho Luna 
Beach, 5-54 m (MHNS); 7 s, Los Laberintos, Rancho Luna Beach, 35 m (MHNS); 21 c, Faro los 
Colorados, 56 m (MHNS); 11 s, Tamarindo Point, 56 m (MHNS). Martinique: 8 c, Pointe Borgnesse, 
12 m, muddy sandy bottom near the reef (CJP). Grenadines: Mayreau, 1 c, west coast, 8 m, coralline 
sandy bottom with coral blocks, gorgonians, and sponges (CJP). Bahamas: 2 s, 6 m, off N Andros, 
dredgeg (CHL); 2 s, Riding Rocks, Cay Sal Bank, 18 m, base coral reef (CHL); 2 s, NW Nassau, 
50 ft. Saint Vincent: 1s (CHL). Honduras: 1 s, Roatan Island, 12 m, coralline sand. Puerto Rico: 


1 s, NW Puerto Rico (CHL). 


Description: The original description 
is as follow: “Shell wider than high. 
Whorls without a keel. Except for the 
nucleus sculptured with fine spiral 
threads, which in larger specimens 
become hardly visible, or absent at the 
periphery of the last whorl. Umbilicus 
present”. This is a very short description 
for a correct specific determination. 

Therefore we are providing a new 
description pointing out the most 
important distinguishing characters: 
Shell (Figs.  90A-F)  subconical 
depressed, solid, whitish and without 
any keel, spire formed by 3 Y convex 
whorls, each overlapping most of the 
previous whorl. Protoconch (Figs. 90H- 
[) about 300 um in diameter and about 1 
Y. whorls, with a slightly rough surface 
and a strong varix which marks the 
beginning of the teleoconch. Teleoconch 
of a little more than 2 whorls, totally 
covered by fine spiral cordlets which 


162 


reach from the suture to the umbilical 
border. On the dorsum of the body 
whorl, the cords diverge somewhat lat- 
erally to the growth axis so that new 
ones continue to appear at the suture. 
One prominent spiral cord limits and 
precipitously angulates the umbilical 
infundibulum, which is deep and not 
very wide. Aperture rounded; columella 
robust; inner lip thickened by the cross- 
ing of the umbilical cord but without 
any callus. 

Dimensions: Holotype 1.59 mm in 
diameter. We have shells reaching about 
25 mm in maximum dimension. 
Maximum reported size: 1.7 mm 

Habitat: This is one of the most 
common species in Cuba. Usually it was 
collected between 10 and 56 m, but 
sometimes shells have been found in 
shallower water. 

Distribution: ABC Islands: Curacao 
(DE JONG é COOMANS, 1988); Mar- 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 90A-I. Vitrinella anneliesae de Jong 82 Coomans, 1988. A-F: shells, 2.7, 2.8, 2.1, 2.7, 2.2, 
3.1 mm, Cienfuegos, Cuba (MHNS); G: microsculpture of the shell Fig. A; H-I: protoconchs. 
Figuras 90A-1. Vitrinella anneliesae de Jong 4 Coomans, 1988. A-F: conchas, 2,7, 2,8, 2,1, 2,7, 2,2, 
3,1 mm, Cienfuegos, Cuba (MHNS); G: microescultura de la concha de la Fig. A; H-I: protoconchas. 


163 


Iberus, 29 (2), 2011 


tinique; Mayreau and Cuba: mainly 
collected in Cienfuegos Bay, but also in 
Canarreos and other areas. 

Remarks: Since its original descrip- 
tion this species has not been recorded 
from any Caribbean locality. It is a very 
characteristic species which has only a 
little similarity to V. funiculus and V. 


contracta. From V. funiculus it may be 
distinguished by the smaller umbilicus 
and by the smaller and more numerous 
spiral cordlets. From V. contracta it 
differs in the shape of its spiral 
cordlets and because they run parallel 
to, rather than obliquely from, the 
suture. 


Vitrinella contracta (Vanatta, 1913) (Figures 91A-E) 


Omalaxis funiculus contractus Vanatta, 1913. Proc. Acad. Nat. Sci. Philadelphia, 65: 25, pl. 2, figs. 
4, 6. [Type locality: Monkey River, British Honduras]. 


Type material: Represented in VANATTA (1913) and deposited in ANSP (106.125). Not examined. 

Other material examined: Cuba: 14 s, Rancho Luna Beach, 45 m (MHNS). Trinidad and Tobago: 
Tobago, 2 c, Horse Shoe Reef, 15 m, shell grit (CJP). Bahamas: 5 s, Olympus Reef, NNW West End, 
Grand Bahama Island, 36 m, coralline algal fragments bottom (CHL); 4 s, French Bay, San Salvador, 
18 m (CHL); 6 s, Grand Bahamas, 7.5 m (CHL). St. Kitts £: Nevis: 4 s, Monkey Shoals, Nevis, 18 m 
(CHL). ABC Islands: 1 s, NW Klein Bonaire, Bonaire, 15 m (CHL). St. Vincent: 1 s (CHL). Belize: 4 
s, Dead Mans Reef, Turneffe Is., 18 m, sand (CHL). Turks € Caicos: 3 s, French Bay, 18 m (CHL). 


Florida, USA: 3 s, APAC Pit, Sarasota, Plio-Pleistocene (CHL). 


Description: Shell (Figs. 91A-C). 
Protoconch (Fig. 91D) of about 1 Y 
smooth whorls, about 200 um in diam- 
eter, set in a lower plane than the teleo- 
conch. Two different stages can be dis- 
tiguished: the embryonic, with barely 
14 whorl and the larval with % whorl 
and partially covered by the first whorl 
of the teleoconch. The entire surface of 
the teleoconch is sculptured by spiral 
cords which are placed parallel to the 
suture (Fig. 91E); there is no puncti- 
form sculpture; a more prominent 
spiral cord is placed in the middle of 
the periphery and gives a keeled 
profile to the shell. Base convex, 
umbilicus deep and narrow, delimited 
by a spiral cord and not closed by the 
columellar thickening. Aperture orbic- 
ular; columella and inner lip thickened 
and reflected outward. 

Dimensions: Holotype 1.85 mm in 
diameter. We have shells up to about 
2.2 mm in maximum dimension. 

Animal and radula unknown. 

Habitat: Not described due to the 
fact that the holotype was found on an 
anchor. Our material was collected on 
coralline bottom between 25 and 50 m 
in depth. 


164 


Distribution:  Known from 
Monkey River, British Honduras 
(VANATTA, 1913); Tobago; and Cien- 
fuegos, Cuba. 

Remarks: VANATTA (1913) 
described this taxon as a variety of 
Omalaxis funiculus Dall, from 
which it is distinguished by having 
a narrower umbilicus and because 
the spiral cords run parallel to the 
suture. S. contracta was placed in 
the genus Solariorbis due to the 
presence oia mala talas 
callus formed in the inner lip in the 
periumbilical region. We think that 
the existence of this small callus is 
insufficient indication to consider it 
a Solariorbis; instead, due to its 
similarity to several species of the 
genus Vitrinella, we consider its 
placement in this genus more accu- 
rate. Vitrinella contracta, as well V. 
funiculus and V. anneliesae, have a 
similar ornamentation, which is at 
the same time different from the 
other species included in this 
genus: smooth spiral cords, non- 
punctiform sulci, strong growth 
lines, and a strong periumbilical 
carina. 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 914-E. Vitrinella contracta (Vanatta, 1913). A-C: shells, 2.0, 2.2, 2.0 mm, Rancho Luna 
Beach, Cuba. D: protoconch, from shell of Fig. A; E: sculpture. 

Figuras 914-E. Vitrinella contracta (Vanatta, 1913). A-C: conchas, 2,0, 2,2, 2,0 mm, Playa Rancho 
Luna, Cuba. D: protoconcha, de la concha de la Fig. A; E: escultura. 


Vitrinella funiculus (Dall, 1892) (Figures 92A-E) 


Teinostoma (Solariorbis) funiculus Dall, 1892. Trans. Wagner Free Inst. Sci. Philadelphia, 3: 417, 
pl. 23, fig. 15. [Type locality: Caloosahatchee River, Florida. Described as fossil of the Plio- 
Pleistocene]. 


Type material: Holotype in USNM (112652). Represented in DaLL (1892). Not examined. 
Other material examined: Cuba: 4 s, Rancho Luna Beach, 45 m. 


Description: Shell (Figs. 92A-D). below that of the teleoconch. Two dis- 
Protoconch (Fig. 92E) heterostrophic, of tinct stages can be observed: the embry- 
about 1 Y4 smooth whorls, about 230 um onic protoconch with barely Y whorl 
in diameter, placed on a plane slightly and the larval one, which has % of 


165 


Iberus, 29 (2), 2011 


100 um 


Beach (MHNS); E: protoconch. 


Figures 92A-E. Vitrinella funiculus (Dall, 1892 


). A-D: shells, 1.6, 1.9, 1.5, 1.8 mm, Rancho Luna 


Figuras 92A-E. Vitrinella funiculus (Dall, 1892). A-D: conchas, 1,6, 1,9, 1,5, 1,8 mm, Playa 


Rancho Luna (MANS); E: protoconcha. 


whorl and is partially covered by the 
first teleoconch whorl. In adult speci- 
mens there is no peripheral angulation. 
The teleoconch has its surface totally 
covered by spiral cordlets, which run 
obliquely from the suture and are a 
little wider on the base near the peri- 
umbilical cord. There is no punctiform 
ornamentation. Umbilicus very wide, 
infundibuliform, exposing the previous 
whorls. The umbilical wall, correspond- 
ing to the columellar margin, lacks 
spiral cords. No columellar callus. 

Dimensions: Holotype 1.75 mm in 
diameter. We have shells of about 1.9 
mm in maximum dimension. 


166 


Distribution: This species has been 
recorded live-collected in Colombia (DÍAZ 
MERLANO éz PUYANA HEGEDUS, 1994). 
From Cuba, Cienfuegos in present work. 

Remarks: DALL (1892) mentions that 
V. funiculus is very similar to the juve- 
niles of Teinostoma opsitelotus, being dis- 
tinguished from this species by the 
umbilical characters. V. funiculus is very 
similar to V. anneliesae, from which it can 
be distinguished by its fewer spiral cords 
and wider umbilicus which exposes the 
previous whorls. From V. contracta it may 
be differentiated by its spiral cordlets 
running obliquely, not parallel, to the 
suture and by the larger umbilicus. 


RUBIO E7 4L.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Vitrinella opsitelotus (Dall, 1892) (Figures 93A-E) 


Teinostoma opsitelotus Dall, 1892. Trans. Wagner Free Inst. Sci., 3: 414, pl. 19, figs. 5, 5b. [Type 
locality: Plio-Pleistocene of the Caloosahatchee River and Shell Creek, Florida]. 
Solariorbis opsitelotus (Dall): In PiLsBrRY (1953). Monogr. Acad. Nat. Sci. Philad. 8: 419, pl. 53, 


figs. 4-4e. 


Type material: Holotype in USNM (113104). Not examined. Specimens figured by PiLsBRY (1953) 


deposited in ANSP (18411). 


Other material examined: Florida, USA: 7 s, APAC Pit, Sarasota Plio-Pleistocene (CHL). 


Description: Original description in 
DaLL (1892: 414). Description expanded 
in PiLsBRY (1953: 419). 

Distribution: Fossil species recorded 
from the Pliocene of the Caloosahatchee 
River and Shell Creek, Florida (DALL, 
1892); from St. Petersburg Pliocene and 
Alligator Creek at Acline, Florida 
(PILsBRY, 1953). 

Remarks: DaLL (1892) placed this 
species in the genus Teinostoma men- 
tioning in his description that the 
umbilicus was “completely filled by a 
flattish, somewhat irregular callus,” in 
shells which Dall believed to be adult of 
T. opsitelotus. PILSBRY (1953) placed it 
in Solariorbis mentioning: “That 
[imperforate] form is not represented in 
the St. Petersburg series of over forty 


specimens. Many of them are about 
equal in size, and appear to be adult. 
This condition may perhaps permit the 
suggestion that Dall's form with the 
umbilicus filled by a callus is an abnor- 
mal or gerontic individual or possibly 
another species”. 

If the columellar thickening proves 
sufficient a a reason to keep it in the 
genus Solariorbis, the form and orna- 
mentation of the protoconch, together 
with the sculpture of the teleoconch and 
the umbilicus (which in basal view, is 
not closed by the thickening of the 
umbilical wall) seem to place this 
species close to the  Vitrinella 
anneliesae, V. contracta and V. funiculus 
group. So, in our opinion it must be 
placed in the genus Vitrinella. 


Vitrinella helicoidea C.B. Adams, 1850 (Figures 94A-G, 95A-D) 


Vitrinella helicoidea C.B. Adams, 1850. Monograph of Vitrinella, a New Genus of New Species 
of Turbinidae: 9. [Type locality: Port Royal, Jamaica]. 

Vitrinella praecox Pilsbry £ McGinty, 1946b. The Nautilus, 60: 14-15, pl. 2, figs. 1-1b. [Type local- 
ity: North end of Lake Worth]. 


Type material: Lectotype in MCZ (156271), represented in CLENCH € TURNER, 1950: plate 35, fig. 1); 
1 paralectotype (labeled as paratype), also from the type locality (MCZ 186188). Vitrinella praecox 
represented in PILsBRY £ MCGINTY (1946b). Not examined. 

Other material examined: Cuba: 5 s, Sancho Pardo Shoal, 15 m (MHNS); 5 s, Guajimico, 5 m 
(MANS); 30 s, Cienfuegos Bay, 8 m (MHNS); 1 s, Cienfuegos Bay, sta. 12, 22%07'N 80%27'W, 9 m 
(MHNS); 4 s, Cienfuegos Bay, sta. 12a, 22%07'N 80%26'W, 4 m (MHNS); 3 s, Cienfuegos Bay, 20-30 
m (MHNS); 18 s, Rancho Luna Beach, 20 m (MHNS); 1 j, Rancho Luna Beach, 10-20 m (MHNS); 4 
s, Cienfuegos Bay, 8 m (MHNS); 4 s, Guajimico, figured (MHNS). Virgin Islands: 2 s, St. Thomas 
(MCZ 156276). Venezuela: 3 s, José Griego, north coast Isla Margarita, 0 m, drift (CHL). ABC 
Islands: 1 s, Newport Reef, Curacao, 20 m (CHL). Antigua: 1 s, Falmouth Harbour, 1-2 m, sand 
grass, (CHL). Turks €: Caicos: 1 s, The G Spot, French Cay, 18 m (CHL). Florida, USA: 1 s, off 
Boynton inlet, Palm Beach Co., 76-90 m (CHL). 


Description: Shell (Figs. 94A-E) tro- 
choid in shape, glossy, with a low profile, 
4 Ya whorls; each whorl overlaps the 


periphery of the previous one. Protoconch 
(Figs. 94F-G) with a finely wrinkled 
surface at its beginning and spiral irregu- 


167 


Iberus, 29 (2), 2011 


Figures 93A-E. Vitrinella opsitelotus (Dall, 1892). A-C: shells, 2.1, 2.1, 2.2 mm, Sarasota Co. 
Florida (CHL); D: detail of the sculpture; E: protoconch. 

Figuras 93A-E. Vitrinella opsitelotus (Dall, 1892). A-C: conchas, 2,1, 2,1, 2,2 mm, Sarasota Co. 
Florida (CHL); D: detalle de la escultura; E: protoconcha. 


lar cordlets in the subsequent part; about 
280 um in diameter and a little more than 
1 Ya whorls. Teleoconch formed by 2 Y 
whorls. From the varix which marks the 


168 


end of the protoconch there are two strong 
spiral cords which extend up to the end of 
the first whorl of the teleoconch where 
they fade out. The subsequent part of the 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 94A-G. Vitrinella helicoidea C.B. Adams, 1850. A-E: shells, 2.1, 1.9, 1.7, 2.5, 2.4 mm, 
Cienfuegos Bay, Cuba (MHNS); F-G: protoconchs. 

Figuras 94A-G. Vitrinella helicoidea C.B. Adams, 1850. A-E: conchas, 2,1, 1,9, 1,7, 2,5, 2,4 mm, 
Bahía de Cienfuegos, Cuba (MHNS); F-G: protoconchas. 


shells is nearly smooth, with only very Dimensions: Lectotype 2.7 mm in 
fine growth lines. A spiral cord angulates diameter and 1.6 mm in height. The 
the border of the umbilical infundibulum. largest shell in our material measures 2.95 
Umbilicus deep with smooth walls. mm in diameter and 1.7 mm in height. 


169 


Iberus, 29 (2), 2011 


Figures 95A-D. Vitrinella helicoidea C.B. Adams, 1850. A-C: shells, 1.8, 1.7, 1.4 mm, Sandro 
Pardo Shoal, Cuba (MHNS); D: protoconch. 
Figuras 95A-D. Vitrinella helicoidea C.B. Adams, 1850. A-C: conchas, 1,8, 1,7, 1,4 mm, Bajo de 
Sandro Pardo, Cuba (MHNS); D: protoconcha. 


Habitat: This species was found 
between 0 and 42 m in depth. It has been 
found under rocks close to shore 
(ANDREWs, 1977). Under rocks in shallow 


170 


waters (Díaz MERLANO é PUYANA 
HEGEDUS, 1994). Our material was col- 
lected on a coralline bottom between 5 and 


20 m. 


RUBIO ET 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Distribution: Recorded from Port 
Royal, Jamaica (ADAMS, 1850); from the 
north inlet of Lake Worth, Palm Beach 
Co., Florida (PILSBRY €: MCGINTY, 1946); 
from Colón and Bocas Island, Panama 
(OLSSON éz MCGINTY, 1958); Bermuda, off 
Cape Hatteras (JOHNSON, 1934), North 
Carolina, South Florida, Texas, Jamaica, 
Haiti, Puerto Rico and Panama (MOORE, 
1964); Virgin Islands (NOwELL-USTICKE, 
1959); SE United States; the Antilles; E 
coast of Central America (HOUBRICK, 
1968). Puerto Rico (WARMKE éz ABBOTT, 
1975). Bermuda; from southeast USA to 
the Antilles; Texas; Ouintana Roo; Costa 
Rica; Panama (ANDREWS, 1977). From 
Campeche to Ciudad del Carmen and 
Zacatal, Mexico (VOKES €: VOKES, 1984). 
Curacao (DE JONG € COOMANS, 1988). 
Bermuda; from North Carolina to Florida 
and the Caribbean Sea (RoBINSON, 1991). 
From Bermuda and North Carolina to the 
Dutch Antilles and Colombia (Díaz 


MERLANO € PUYANA HEGEDUS, 1994). 
Abaco, Bahamas (REDFERN, 2001). Our 
material extends the range to Cuba, the 
ABC Islands, Turks € Caicos, and 
Venezuela. 

Remarks: MOORE (1964: 58) considers 
V. praecox a synonym of V. helicoidea 
applying the observation that it was 
described by PILSBRY €: MCGINTY (1946) 
on the basis of juvenile specimens which 
show the spiral cords. We have found 
shells (Figs. 94C-E) with the teleoconch 
whorls more depressed, dorsally and ven- 
trally, and which exhibit three spiral cords 
instead of two on the first Y whorl of the 
teleoconch. The rest of the sculptural char- 
acters and dimensions of the shell and 
protoconch are similar (3 x 1.8 mm; proto 
270 um). Lacking radular and anatomical 
information on these slightly different 
shells, we provisionally consider them 
only morphological variations of a single 
species. 


Vitrinella floridana Pilsbry €£ McGinty, 1946 (Figures 96A-D) 


Vitrinella floridana Pilsbry y McGinty, 1946. The Nautilus, 60: 16-17, pl. 2, figs. 4-4a. [Type local- 
ity: Northern Biscayne Bay near Baker's Haulover, Miamil. 


Type material: Holotype figured by MoorE (1964, Fig. 4) deposited in ANSP (181880). Not examined. 
Other material examined: Florida, USA: 32 s, Hypoluxo Island, Lantana, Palm Beach Co., drift 
(CHL); 1 s, N Peanut Island, Lake Worth, Palm Beach Co., dredged at 2-4.5 m (CHL). 


Description: This is the original 
description in PILSBRY ££ MCGINTY (1946): 
“The minute shell is depressed, whitish, 
smooth, openly umbilicate, the umbilicus 
contained 3.25 times in the diameter. 
There are barely 3 Y% convex whorls, the 
last whorl somewhat flattened below the 
periphery, rounded at periphery, base, 
and umbilical border, the umbilicus 
rather broadly open, perspective, the 
whorls visible within it convex. The aper- 
ture is rather strongly oblique, rounded 
angular above (or in a basal view it 
appears bluntly triangular). Peristome 
thin, the upper margin only moderately 
arched forward. The columellar margin 
is thickened and runs forward above. 
Parietal callus rather thick and short. 
Diameter 1.95 mm, height 0.95 mm; 
umbilicus 0.6 mm wide”. 


We can add: Shell (Figs. 96A-C). Pro- 
toconch (Fig. 96D) of about 1 % whorls 
and about 330 um in maximum diame- 
ter, with a slightly rough surface, more 
evident on the subsutural area, which 
gives it a frosted appearance. Teleo- 
conch of about 2 whorls, smooth except 
for clear growth lines also visible within 
the umbilicus. Umbilicus wide and deep 
with rounded walls, lacking a perium- 
bilical cord. Aperture ovoid strongly 
prosocline. 

Dimensions: We have shells with 
about 1.30 mm in maximum dimension 
and 0.75 mm in height. 

Habitat: This species has a mainly 
continental distribution, living between 
0 and 46 m in depth. MOORE (1964) 
mentions that it is very common in 
some localities, having examined a lot of 


TA 


Iberus, 29 (2), 2011 


Figures 9GA-C: Vitrinella floridana Pilsbry 8% McGinty, 1946. A-C: shells, 1.3, 1.3, 1.4 mm, 
Hypoluxo Island, Lantana, Palm Beach Co., Florida (CHL); D: protoconch. 

Figuras 96A-C: Vitrinella floridana Pilsbry Y McGinty, 1946. A-C: conchas, 1,3, 1,3, 1,4 mm, 
Hypoluxo Island, Lantana, Palm Beach Co., Florida (CHL); D: protoconcha. 


924 specimens from Mud Island, Biscayne Bay near Baker”s Haulover, 
Aransas Bay, Texas. Miami; Indian River south of Sebastian, 

Distribution: The species is known Indian River Co.; North Inlet of Lake 
from the USA: Recorded from Northern Worth, Palm Beach; and Barnes Sound, 


172 


RUBIO E7 42z.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Key Largo, Florida (PILSBRY € 
MCGINTY, 1946); East and West 
Florida, Texas (Lyons, 1971; 
ANDREWS, 1977); Texas (ODÉ, 1987). 
Mexico: Quintana Roo (VOKES éz 
VOKESs, 1984); Tamaulipas, Tabasco, 
Veracruz, Campeche Bank, Quin- 
tana Roo (GARCÍA-CUBAS, 1970, 
1990). Colombia (Díaz MERLANO éz 
PUYANA HEGEDUS, 1994). Cuba: 
Cienfuegos Bay. 

Remarks: The small size, rather 
large umbilicus without a border- 
ing cord, and the feeble convexity 
of the upper margin of peristome in 


apical or basal view are its more 
conspicuous features (PILSBRY €: 
MCGINTY, 1946). Vitrinella flori- 
dana has a shell very similar in 
general appearance to that of Vit- 
rinella helicoidea from which it can 
be distinguished by the ornamenta- 
tion of the protoconch, by the lack 
of spiral cords on the first Y whorl 
of the teleoconch and the lack of a 
bordering umbilical cord. From Vit- 
rinella canaliculata it can be differ- 
entiated by the absence of the 
sutural canal and the ornamenta- 
tion of the protoconch. 


Vitrinella canalicu lata spec. nov. (Figures 97A-C) 


Type material: Holotype (Fig. 97A) in MNCN (15.05 /55070). Paratypes: MHNS (100553, 1 s); MNHN 


(24399, 1 s); CFG (1 s). 


Type locality: Rancho Luna Beach, Cienfuegos, Cuba. 
Etymology: The specific name refers to the sutural canal which is visible along the full extent of the 


teleoconch. 


Description: Shell (Figs. 97A-B) 
depressed, without sculpture and with a 
broad umbilicus. Protoconch (Fig. 97C) 
of about one whorl, with a slightly 
rough surface, and about 320 um in 
maximum diameter; immediately after 
the protoconch there commences a wide 
suture which forms a small groove. 
Teleoconch of about 2 whorls, smooth 
except for fine growth and the sutural 
canal, which is visible all the way to the 
aperture. Umbilicus wide and deep with 
rounded walls, without a periumbilical 
cord. 

Dimensions: Holotype 1.8 mm in 
diameter. 


Habitat: The studied material 
came from coralline sand bottoms 20 
m in depth. 

Distribution: Only known from 
Rancho Luna Beach, Cienfuegos, 
Cuba, its type locality. 

Remarks: V. canaliculata spec. 
nov. has a shell very similar in 
general appearance to that of V. heli- 
coidea and V. floridana, from which 
it can be distinguished by the orna- 
mentation of the protoconch, the 
lack of spiral cords on the first Y 
whorl of the teleoconch, having a 
sutural canal and the lack of perium- 
bilical cord. 


Vitrinella aristata spec. nov. (Figures 98A-I) 


Type material: Holotype (Fig. 98A) in MNCN (15.05/55069). Paratypes: ANSP (1 s); AMNH (4 s); 
FLMNH (448612, 1 s); MCZ (1 s); MHNS (100554, 24 s); MNHN (24400, 1 s); NHMUK (1 s); USNM 
(1155033, 1 s); IES (1 s); CFG (10 s), CHL (3 s) and CFR (10 s). 

Other material examined: Cuba: 3 s, Cienfuegos Bay, 20-30 m (MHNS); 2 s, Rancho Luna Beach, 12 
m (MHNS7); 1 sp, 14 s, Rancho Luna Beach, 10-20 m (MHNS); 2 s, Rancho Luna Beach, 35 m (MHNS); 
8 s, Rancho Luna Beach, 40 m (MHNS); 27 c, Rancho Luna Beach, 45 m (MHNS); 4 c, Los Laberin- 
tos, Rancho Luna Beach, 35 m (MHNS); 1 sp, 17 s Faro de los Colorados, 56 m (MHNS); 1 s, Punta 
de Tamarindo. 

Type locality: Rancho Luna Beach, Cienfuegos, Cuba. 


173 


Iberus, 29 (2), 2011 


Figures 97A-C. Vitrinella canaliculata spec. nov. A-B: holotype, 1.8 mm (MNCN); B: paratype, 
1.3 mm, Rancho Luna Beach (MHNS); C: protoconch. 
Figuras 97A-C. Vitrinella canaliculata spec. nov. A-B: holotipo, 1,8 mm (MNCN); B: paratipo, 1,3 


mm, Playa Rancho Luna (MHNS); C: protoconcha. 


Etymology: The specific name refers to the unique pattern of dorsal opisthocline and ventral proso- 
cline grooves, which meet at an acute angle at the mid-periphery creating a pattern reminscent of 


an ear, or husk, of grain, in Latin “arista”. 


Description: Shell (Figs. 98A-D) 
depressed, solid, brown, polished, and 
with characteristic oblique grooves. 
Protoconch (Figs.  98F-G)  het- 
erostrophic, about 1 % whorls, and 
about 280 um in diameter, placed on a 
plane slightly above that of the other 
whorls, with a smooth surface at its 
beginning followed by a segment with 
coarse prosocline growth lines termi- 
nating with a faint axial varix. Teleo- 
conch of about 1 Y rapidly-increasing 
whorls; dorsally and ventrally convex; 
from the suture and from the umbilical 
border, oblique sulci (opisthocline and 
prosocline respectively) converge at 
the middle of the periphery. These 
sulci are less evident on the last Y 
whorl. Aperture rounded, oblique, col- 
umellar margin not thickened, outer 
and inner lips sharp. Umbilicus wide 
and deep, exposing the previous 


174 


whorls, without ornamentation except 
for the growth lines and several faint 
spiral cords. 

Dimensions: Holotype 1.23 mm in 
diameter and 0.57 mm in height. 

Operculum corneous and multispi- 
ral with a central nucleus. 

Radula (Fig. 98H) taenioglossate, 
with formula 2+1+R+1+2. Central 
tooth wide basally, the ventral margin 
well developed, without denticles. 
Cutting area formed by a large and 
sharp cusp and 5 denticles of medium 
size at each side. Lateral teeth similar 
to the central one, but the base is nar- 
rower; free margin with a central cusp 
and 4-5 smaller denticles at each side. 
Marginal teeth narrow and elongate; 
the inner with 24-26 fine denticles on 
the upper outer margin; the outer mar- 
ginal teeth are strongly inclined 
outward in their upper third and 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 98A-1. Vitrinella aristata spec. nov. A-B: holotype, 1.23 mm, Rancho Luna Beach, Cuba 
(MNCN); C-D: paratypes, 1.0, 0.9 mm, Rancho Luna Beach (MHNS); E: microsculpture; F-G: 
protoconch; H: radula: 1: operculum, 0.5 mm in diameter. 

Figuras 98A-1. Vitrinella aristata spec. nov. A-B: holotipo, 1,23 mm, Playa Rancho Luna, Cuba 
(MNCN); C-D: paratipos, 1,0, 0,9 mm, Playa Rancho Luna (MHNS); E: microescultura; F-G: proto- 
concha; H: rádula: I: opérculo, 0.5 mm de diámetro. 


175 


Iberus, 29 (2), 2011 


possess 16-18 denticles on the upper 
end of their inner margin. 

Animal unknown. Operculum (Fig. 
981) rounded and multispiral. 

Habitat: This species has been col- 
lected between 10 and 60 m, on a 
coralline, slightly muddy bottom. 

Distribution: Only known from 
Cuba. 


Remarks: Vitrinella aristata spec. 
nov. has a glistening shell with a very 
attractive and characteristic sculpture 
which makes it unmistakable. V. 
anneliesae is the only species with a 
similar ornamentation, with slightly 
oblique cords emerging from the 
suture. The radula is similar to that of 
other tornid species. 


Vitrinella pseudoaristata spec. nov. (Figs. 99A-E) 


Type material: Holotype (Fig. 994) in MNCN (15.05/55071). Paratypes: MNHN (24401, 1 s, Fig. 


99B) and IES (1 s, Fig. 99C), both from type locality. 


Type locality: Rancho Luna Beach, Cienfuegos, Cuba. 
Etymology: The specific name refers to the similarity with the species Vitrinella aristata. 


Description: Shell (Figs. 99A-C) 
depressed, solid, polished, and with 
characteristic oblique grooves. Proto- 
conch (Fig. 99D) heterostrophic, 1 Y 
whorls, about 290 um in diameter, and 
slightly projected. Teleoconch of about 2 
rapidly-increasing whorls; dorsally and 
ventrally convex; completely smooth 
except for 5-6 somewhat oblique incised 
lines seen on the dorsum of the first 
teleoconch whorl; these fade on the 
periphery, which is rounded. A thick 
cordon marks the boundary of the 
umbilicus, which is not occluded by a 
callus. Aperture rounded, robust; thick 
and somewhat reflected columella, but 
without callus formation. Umbilicus not 
too wide or deep, flanked by the spiral 
cord. Aperture rounded, oblique, col- 
umellar margin not thickened, outer 
and inner lips sharp. 


Dimensions: Holotype 1.26 mm in 
diameter, similar to the paratypes. 

Animal and radula unknown. 

Habitat: This species has been col- 
lected between 15 and 35 m in coralline 
sand. 

Distribution: Only known from 
Cuba. 

Remarks: Vitrinella pseudoaristata 
spec. nov. has a glistening shell with a 
very attractive and characteristic sculp- 
ture which makes it unmistakable. V. 
anneliesae and V. aristata are the only 
species with a similar ornamentation, 
possessing slightly oblique cords near 
the suture. V. pseudoaristata spec. nov. 
differs from V. aristata spec. nov. by 
having fewer oblique lines and having 
them limited to the dorsum of the first 
teleoconch whorl; furthermore it has a 
spiral cord that delimits the umbilicus. 


Vitrinella pelorcei spec. nov. (Figures 100A-C) 


Type material: Holotype (Figs. 100A-B) in MNHN (24202). 


Type locality: Reef Bay Bronage, Antigua. 


Etymology: After Jacques Pelorce, the French malacologist who collected the holotype. 


Description: Shell (Figs. 100A-B) of 
small size, depressed, appearing 
fragile, with a sutural groove and 
characteristic large axials ribs. Proto- 
conch (Fig. 100C) of about 1 % whorls, 
placed on a higher plane than the 
teleoconch, with a smooth surface at 


176 


its beginning and 3-4 fine spiral 
cordlets of microtubercles in the subse- 
quent part, about 290 um in diameter. 
Teleoconch with almost 1 Y rapidly- 
increasing whorls; dorsally and ven- 
trally convex. Ornamentation formed 
by axial growth lines and strong axial 


RUBIO ET 412.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 99A-G. Vitrinella pseudoaristata spec. nov. A: holotype, 1.26 mm, Rancho Luna Beach, 
Cuba (MNCN); B: paratype, 1.2 mm, Rancho Luna Beach (MNHN); C: paratype, 1.2 mm, 
Rancho Luna Beach (1ES); D: protoconch; E: microsculpture. 

Figuras 99A-G. Vitrinella pseudoaristata spec. nov. A: holotipo, 1,26 mm, Playa Rancho Luna, Cuba 
(MNCN); B: paratipo, 1,2 mm, Playa Rancho Luna (MNHN); C: paratipo, 1,2 mm, Playa Rancho 
Luna (IES); D: protoconcha; E: microescultura. 


ribs, nine on first teleoconch whorl; the 
last Y whorl loses its axial sculpture 
leaving only very faint spiral cordlets. 
The suture is very evident in the proto- 


conch, and it becomes a wide groove 
along the  teleoconch. Aperture 
rounded, oblique, columellar margin 
not thickened, outer and inner lips 


AZ 


Iberus, 29 (2), 2011 


Figures 100A-C. Vitrinella pelorceí spec. nov. A-B: holotype, 1.02 mm, Reef Bay Bronage, Antigua 
(MNHN); C: protoconch. 
Figuras 100A-C. Vitrinella pelorcei spec. nov. A-B: holotipo, 1,02 mm, Reef Bay Bronage, Antigua 
(MNAN); C: protoconcha. 


sharp. Umbilicus wide and deep 
exposing the previous whorls. No 
sculpture except for axial growth lines. 

Dimensions: Holotype 1.02 mm in 
maximum diameter. 

Animal and radula unknown. 

Habitat: The only shell known was 
collected on a coralline bottom at 1 m. 

Distribution: Only known from 
Antigua, the type locality. 

Remarks: Vitrinella pelorcei spec. 
nov. has a characteristic ornamentation 


formed by strong and widely- 
spaced axial ribs and an evident 
suture in the protoconch which in 
the teleoconch evolves into a sutural 
sulcus continuing to the aperture. 
These characters make this species 
unmistakable. Vitrinella floridana is 
the only species with a similar 
sutural sulcus, but it is narrower 
and deeper; Furthermore the latter 
has no other sculpture, axial or 
spiral. 


Vitrinella aguayoi (Corgan, 1968) (Figures 101A-E) 


Vitrinella tenuisculpta Aguayo «€ Borro, 1946b. Rev. Sociedad. Malac. “Carlos de la Torre,” 4(2): 
43-44, pl. 3, figs. 4-6. [Type locality: Matanzas, Cuba], preoccupied by Vitrinella tenuisculp ta 


Carpenter, 1865. 
Solariorbis aguayo1 Corgan, 1968. 


178 


RUBIO £7 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 101A-H. Vitrinella aguayoi Corgan, 1968. A-D: shells, 1.45, 1.4, 1.4, 1.2 mm, Cienfue- 
gos, Cuba (MHNS); E: protoconch. 

Figuras 1O1A-H. Vitrinella aguayoi Corgan, 1968. A-D: conchas, 1,45, 1,4, 1,4, 1,2 mm, Cienfue- 
gos, Cuba (MHNS); E: protoconcha. 


Type material: Holotype represented in AGUAYO éz BORRO (1946, figs. 4-6). Deposited in Museo 
Poey, Havana, Cuba (12005). Described as fossil of the Upper Tertiary found in the formation 
“Yumurí,” Upper Miocene of Cuba. Not examined. 


179 


Iberus, 29 (2), 2011 


Other material examined: Cuba: 6 s, Rancho Luna Beach, 12 m (MHNS); 51 s, Rancho Luna Beach, 
10-20 m (MHNS); 1 s, Rancho Luna Beach, 35 m (MHNS); 1 sp, 5 s Faro de los Colorados, 56 m 


(MHNS). 


Description: The original descrip- 
tion (AGUAYO é BORRO, 1946b) is as 
follows: “Concha diminuta, depri- 
mida, plano convexa por encima, 
ampliamente umbilicada por debajo. 
Provista de 3 vueltas redondeadas, la 
ultima ligeramente aquillada. Las pri- 
meras 1 Y (nucleares), lisas, las restan- 
tes de escultura reticulada. Escultura 
espiral formada por numerosas líneas 
salientes (unas 15 en la última vuelta); 
escultura axial formada por líneas mas 
elevadas que las espirales, formando 
con estas cuadrículas que comunican a 
la concha un aspecto granuloso. Perife- 
ria subangulosa. Ombligo amplio, for- 
mando con la base de la concha un 
ángulo muy marcado que lo bordea 
por completo. Abertura semilunar, con 
el borde unido por un leve callo parte- 
pal 

Shell (Figs. 101A-C) subconical, 
depressed, solid, whitish and formed 
by 3 % rapidly-increasing whorls. Pro- 
toconch (Fig. 101H) of about 1 % 
whorls, about 290 um in diameter, has 
a rough surface at its beginning and 
fine oblique threads in the subsequent 
part, slightly overlapped by the subse- 
quent whorl. Teleoconch of about 2 Y 
whorls, sculpture of 38-39 spiral 
cordlets, 24 on the dorsum and 14-15 
ventrally, spiral cordlets crossed by 75- 
80 fine axial ribs, forming small quad- 
rangular hollows. Dorsally convex 
with a subangulated periphery. Ven- 
trally slightly concave with one promi- 
nent spiral cord which limits a wide 
infundibulum of the deep umbilicus 
with smooth walls. Aperture oval 
almost circular, columella and inner lip 
very thick and reflected outward and 
the upper part of the external lip sharp 
and advanced. There is no columellar 
thickening or encroachment on the 
umbilicus. At junction of the columella 
and umbilical cord the aperture is 
slightly angulated. 

Dimensions: Holotype 1.19 mm in 
diameter and 0.5 mm in height. We 


180 


have shells reaching 1.45 mm in 
maximum dimension 

Habitat: The type is a fossil speci- 
men collected in the Upper Miocene of 
Cuba, from the “Yumuri” Formation. 
Our recent material was collected 
between 10 and 56 m. 

Distribution: Cuba: Cienfuegos. 
Since its description this species has 
not been recorded from any other 
Caribbean locality. Thus it could be 
endemic to Cuba. 

Remarks: Solariorbis aguayoi was 
proposed by CORGAN (1968) as a 
replacement name for Vitrinella tenuis- 
culpta Aguayo € Borro, 1946, which is 
preoccupied by V. tenuisculpta Car- 
penter 1965: The renamed species, 
known only from the Miocene of 
Cuba, was placed in the vitrinellid 
genus Solariorbis Conrad, 1865 by 
Corgan on the basis of reticulate 
microsculpture. 

AGUAYO € Borro (1946b) men- 
tioned that V. tenuisculpta. (=V. 
aguayo1) is more similar to Vitrinella 
multistriata (A.E. Verrill), and it can be 
distinguished by being smaller, having 
weaker sculpture, the axial lines being 
proportionately more prominent than 
the spiral ones, the umbilicus wider, 
and the periphery angled angulate. 

Vitrinella aguayoi and V. cal- 
liglypta are very similar, and they 
were considered as morphotypes of 
the one species in the past. They live in 
the same type of marine bottom, and 
maintain their distinguishing charac- 
ters without intergradation. The proto- 
conchs are identical in size and orna- 
mentation and, as with other species 
included in the genus Vitrinella (V. 
annelisae, V. contracta, V. funiculus), 
the protoconch is slightly overlapped 
by the first whorl of the teleoconch. 

Vitrinella aguayoi may be distin- 
guished from V. calliglypta by its 
angled periphery and by the greater 
number of spiral cordlets and axial 
ribs. 


RUBIO ET ALz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Vitrinella calliglypta Aguayo, 1949 (Figures 102A-E) 


Vitrinella (Delphinoidea) calliglypta Aguayo, 1949. Rev. Sdad. Malac. “Carlos de la Torre,” 6: 94, 
pl. 4, fig. 4, 6. [Type locality: Gibara, Eastern, Cuba]. 


Type material: Holotype deposited in the Museo Poey, Havana, Cuba (12901). Collected by P.J. 
Bermudez and C.G. Aguayo. Not examined. 

Other material examined: Cuba: 2 s, Cienfuegos Bay, 20-30 m (MHNS) 1 s, Cienfuegos Bay, 30 m 
(MHNS); 48 s, Rancho Luna Beach, 10-20 m (MHNS); 3 s, Rancho Luna Beach, 40 m (MHNS); 8 s, 
Rancho Luna Beach, 45 m (MHNS); 3 s, Los Laberintos, Rancho Luna Beach, 35 m (MHNS); 3 s, 


Faro los Colorados, 56 m (MHNS). 


Description: The original descrip- 
tion (AGUAYO, 1949) is as follows: 
“Concha diminuta, deprimida, planor- 
boide, blanca, lustrosa, translúcida, 
ampliamente  umbilicada.  Espira 
deprimida, sutura profunda pero 
estrecha. Provista de 3 Y% vueltas, de 
las cuales 1 Y son lisas, y las restantes 
con una escultura entrecruzada, 
formada por numerosas costillitas 
planas axiales, algo más estrechas que 
sus intervalos y muchas costillas espi- 
rales (unas 15 en la última vuelta) más 
bajas que las axiales, cuyo entrecruza- 
miento le da un aspecto de numerosas 
depresiones cuadrangulares. Periferia 
redondeada. Base aplanada, con un 
ámplio ombligo de un 40% del diáme- 
tro de la concha. Abertura subcircular, 
peristoma cortante”. 

Shell (Figs. 102A-D) subconical 
depressed, solid, whitish, and com- 
prised of 3 % rapidly-increasing 
whorls. Protoconch (Fig. 102E) project- 
ing slightly upward, about 1 Y whorls 
and about 290 um in diameter. It has a 
rough surface at its beginning and fine 
oblique threads in the subsequent 
part, slightly overlapped by the subse- 
quent whorl. Teleoconch of about 2 
whorls, with sculpture formed by 32- 
34 spiral cordlets: 20 on the dorsum 
and 12-14 ventrally, each crossed by 
55-60 fine axial ribs, forming small 


quadrangular  hollows.  Dorsally 
slightly convex with a biangulated 
periphery. Ventrally the shell is 
slightly concave with one spiral cord 
which limits a wide infundibulum of 
the deep umbilicus. Aperture oval, 
almost circular, columella and inner 
lip very thick and reflected outward, 
and the upper part of the external lip 
sharp and advanced. At the junction 
between columella and the periumbili- 
cal cord, the aperture is slightly angu- 
lated. 

Dimensions: Holotype 1.0 mm in 
diameter and 0.6 mm of height. We 
have shells reaching about 1.23 mm in 
maximum dimension. 

Habitat: The type was dredged off 
Gibara, East of Cuba, at 30 fathoms in 
depth. Our material was collected 
between 10 and 56 m. 

Distribution: Cuba: Holguin, 
Gibara, Cienfuegos. Since its descrip- 
tion this species has not been recorded 
from any other Caribbean locality, so it 
could be endemic to Cuba. 

Remarks: AGUAYO (1949) men- 
tioned that V. calliglypta shows a little 
similarity to V. tenuisculpta Aguayo é 
Borro, and it can be distinguished by 
the biangulate periphery and the 
fewer axial ribs and spiral cords. V. 
multistriata Bush is smaller and has 
stronger sculpture. 


Vitrinella cupidinensis Altena, 1966 (Figures 103A-F) 


Vitrinella (Striovitrinella) cupidinensis Altena, 1966. Zoologische Mededelingen, 41: 235-236, 
figs. 2a-d. [Type locality: Cupido, river Maratakka, Surinam]. 


Type material: Represented in ALTENA (1966b, 1975). Not examined. 


181 


Iberus, 29 (2), 2011 


Beach, Cuba (MHNS); E: protoconch. 


Figures 102A-E. Vitrinella calliglypta Aguayo, 1949. A-D: 1.4, 1.3, 1.1, 1.2 mm, Rancho Luna 


Figuras 102A-E. Vitrinella calliglypta Aguayo, 1949. A-D: 1,4, 13, LL, 1,2 mm, Playa Rancho 


Luna, Cuba (MANS); E: protoconcha. 


Other material examined: Guatemala: 2 s, Livingston, 3 m (MHNS). Trinidad and Tobago: Tobago, 
1 s, Courland Bay, drift (CHL); 1 s, Scarborough (CHL). Venezuela: 1 f, Juan Griego, Isla Margarita 


(CHL); 2 s, Isla Margarita (CHL). 


Description: Shell (Figs. 103A-C) 
lenticular, depressed, whitish, with 3 Y 
spiral whorls. Protoconch (Fig. 103D) 1 
1% whorls, diameter of about 350 um; the 


182 


first Y whorl is smooth, and the next 
whorl has small granules on the periph- 
eral area. Two varices mark the end of 
each stage. Teleoconch of about 1 % 


RUBIO ET 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 103A-E. Vitrinella cupidinensis Altena, 1966. A-C: shells, 1.7, 1.7, 1.35 mm, Livingston, 
Guatemala (MHNS); D: shell, 1.7 mm, Tobago (CHL); E: protoconch, Guatemala; F: Proto- 
conch, Tobago. 

Figuras 103A-E Vitrinella cupidinensis Altena, 1966. A-C: conchas, 1,7, 1,7, 1,35 mm, Livingston, 
Guatemala (MHNS); D: concha, 1,7 mm, Tobago (CHL); E: protoconcha, Guatemala; F: protocon- 
cha, Tobago. 


Iberus, 29 (2), 2011 


whorls, its surface totally covered by 
spiral cords of similar size and very fine 
axial striae which cross the sulci 
forming a characteristic sculpture. 
Dimensions: Holotype 1.8 mm in 
diameter and 0.9 mm in height. We have 
shells reaching 1.7 mm in maximum 
diameter and 1.06 mm in height. 
Habitat: We found no mention of the 
habitat of this species in the literature. 
Our material was collected in muddy 
bottom with turbid waters at 3 m. 
Distribution: Colombia (Díaz éz 
PUYANA, 1994). Brazil: Recife (Altena, 


1966), Pernambuco (Rios, 1994) and Liv- 
ingston, Guatemala. 

Remarks: Vitrinella cupidinensis was 
described from fossil shells in the 
Holocene of Surinam. The distinguish- 
ing Characteristic of this species is the 
teleoconch sculpture of fine spiral cords 
crossed by very fine axial striae. ALTENA 
(1966) indicated that this species is very 
similar to Vitrinella (Striovitrinella) 
elegans Olsson £ McGinty, 1958, but the 
latter is slightly larger and, at the same 
number of whorls, the spiral sculpture is 
smaller and the radial more prominent. 


Vitrinella filifera Pilsbry €££ McGinty, 1946 (Figures 104A-D) 


Vitrinella filifera Pilsbry € McGinty, 1946. The Nautilus, 60: 15, pl. 2, figs. 2-2b. [Type locality: 
Biscayne Bay at Baker's Haulover, Miami, Florida]. 


Type material: Holotype in ANSP (n* 181879) not figured by MOORE (1964). Not examined. 
Material examined: Virgin Islands: 1s, Magens Bay, N St. Thomas, (CHL). 


Description: This is the original 
description in PIiLsBRY é MCGINTY 
(1946): “The shell is depressed, umbili- 
cate, the width of umbilicus contained a 
little more than 4 times in the diameter; 
thin, white (dead), smooth. The upper 
surface is convex with slightly promi- 
nent apex, the whorls convex, the last 
whorl having a cord a short distance 
below the suture and parallel to it, 
becoming weaker near the aperture; the 
periphery is broadly rounded; base 
convex, a little impressed along the cord 
around the umbilicus; which in its last 
turn enlarges to about double its former 
width and is bounded by a cord which 
becomes weaker near the aperture. The 
aperture is rounded, somewhat oblique, 
the peristome thin, upper margin is 
strongly arched forward, retracted to 
the upper insertion, the basal margin 
straightened or a little curved forward 
in a basal view, and there is a slight 
angle at the termination of the umbilical 
cord. The columella is slanting, near 
straight, rather thick. Parietal callus 
thin. Diameter 1.25 mm; height 0.7 mm; 
3 Y whorls”. 

We add the following: The proto- 
conch (Fig. 104D) projects slightly, has 


184 


1 % spiral whorls, is bulbous, mea- 
sures about 280 um in diameter, and is 
sculptured with microtubercles of 
varying size distributed irregularly, 
the larger ones near the subsutural 
area. On the last segment there are 
some spiral threads near the suture. 
The teleoconch has 1 Y whorls com- 
pletely covered by dense, fine axial 
ribs, which cross the spiral cordlets, 
more evident on the periphery. The 
axial sculpture is predominant on the 
dorsum. 

On the base two more prominent 
spiral cords can be observed; one of 
them borders the periphery, and the 
other delimits the umbilical zone. The 
umbilicus is wide, with convex walls 
on which axial and spiral cordlets can 
be seen. Aperture rounded, columella 
thickened and reflected outward. 

Habitat: This species has not yet 
been found alive (PILSBRY €: MCGINTY, 
1946). MOORE (1964) did not comment 
on its ecology. 

Distribution: The species is known 
from the USA: recorded from Biscayne 
Bay at Baker”s Haulover, Miami, 
Florida (PILsBRY éz MCGINTY, 1946b); 
from both sides of the Florida penin- 


RUBIO E7 41.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Virgin Islands (CHL); D: protoconch. 


Figures 104A-D. Vitrinella filifera Pilsbry 82 Mc 


Ginty, 1946. A-C: shell, 0.92 mm, St. Thomas, 


Figuras 104A-D. Vitrinella filifera Pilsbry * McGinty, 1946. A-C: concha, 0,92 mm, St. Thomas, 


Virgin Islands (CHL); D: protoconcha. 


sula from Palm Beach to St. Petesburg 
(MOORE, 1964); from Florida to Texas 
and Colombia (ODÉ, 1987). Colombia, 
(Díaz MERLANO 6 PUYANA HEGEDUS, 
1994). Brazil: Cabo Frio, Rio de Janeiro 
(Rios, 1994). 

Remarks: PILSBRY é€  MCGINTY 
(1946) described Vitrinella filifera, but 


in spite of some similarities with V. 
thomasi (Pilsbry, 1945), they didn't 
compare the two species. MOORE (1964) 
considered the two synonymous, indi- 
cating that V. filifera was described from 
a somewhat eroded adult shell which 
may have lost some sculptural charac- 
ters Furthermore, he considered Vit- 


183 


Iberus, 29 (2), 2011 


rinella filifera as the valid nominal taxon 
since “Cyclostrema” thomasi was a sec- 
ondary junior homonym of Vitrinella 
thomasi Bartsch, 1918 (see below). The 
strong thread which follows the suture 
and the strongly convex outline of the 
upper margin of the aperture, seen in 


apical or basal view, are distinctive fea- 
tures of this species, which has not yet 
been found alive (PILSBRY € MCGINTY, 
1946). The projected protoconch, the fine 
and dense axial ribs, which cross the 
spiral cordlets, distinguish it from V. 
solaris. 


Vitrinella solaris nom. nov. (Figures 105A-C) 


“Cyclostrema” thomasi Pilsbry, 1945b. The Nautilus, 59: 60, pl. 6, figs. 7-7b. [Type locality: North 
Inlet of Lake Worth, Palm Beach, Florida]. 
[non Vitrinella thomasi Bartsch, 1918]. 


Type material: The holotype of “Cyclostrema” thomasiin ANSP (181309). The drawings in the orig- 
inal description are quite distinctive. 

Other material examined: Cuba: 1 s, Cienfuegos Bay, 8 m (MHNS); 1 s, Cienfuegos Bay, 10-20 m 
(MHNS); 4 s, Cañon of Cienfuegos Bay, 8 m (MHNS); 1 s, Rancho Luna Beach, 10-20 m (MHNS); 
2 s, Cayo Carenas, Cienfuegos Bay, 10 m (CFG). 

Etymology: The specific name is based on the image of a child's drawings of the sun, rounded with 


radial lines. 


Description: See PILSBRY (1945b). 

The holotype is 1 mm. 

Habitat: Species living in shallow 
waters. Bathymetric range: 1-20 m. 

Distribution: USA: Florida (PILSBRY, 
1945b); Texas (MOORE, 1964); ODÉ, 
1987b). Cuba in our material. 

Remarks: Vitrinella solaris nom. 
nov. and Vitrinella filifera are two very 
close species. For this reason, they 
were placed in synonymy by MOORE 
(1964). While it is true that the shells 


on which V. filifera was based were 
very eroded, and some characters were 
lost by abrasion, it is nonetheless a dis- 
tinct species. The shells figured here 
show that there are two species 
involved, a little similar in profile but 
perfectly distinct in SEM micropho- 
tographs. Vitrinella filifera has narrow 
and curved axial ribs on the dorsum, 
and the protoconch has a spiral row of 
tubercles which are not seen in V. 
solaris nom. nov. 


Genus Vitrinorbis Pilsbry €: Olsson, 1952 


Vitrinorbis callistus Pilsbry €: Olsson, 1952. Type species by original designation. 


Remarks: This genus was proposed 
for a group of minute, few-whorled dis- 
coidal species with a low, flat or concave 
spire, the base with a wide, open 
umbilicus and strongly  carinated 
periphery. The surface is delicately 
sculptured with microscopic spiral 


threads which give it a soft, satiny 
luster. The two previously known 
species are from the eastern Pacific 
(Panamic Province), but the following 
one from the Caribbean appears to 
belong to this genus (PILSBRY $: OLSSON, 
1952). 


Vitrinorbis elegans Olsson £ McGinty, 1958 (Figures 106A-E) 


Vitrinorbis elegans Olsson €: McGinty, 1958. Bulletin of American Paleontology 39: 31-32, pl. 4, 
figs. 3-3a. [Type locality: Bocas Island, Panama]. 


186 


RUBIO ET 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


(MHNS). C: protoconch. 


Figures 105A-C. Vitrinella solaris nom. nov. A-B: shells, 0.95, 0.95 mm, Cienfuegos Bay 


: 100 um 


Figuras 105A-C. Vitrinella solaris nom. nov. A-B: conchas, 0,95, 0,95 mm, Babía de Cienfuegos 


(MANS). C: protoconcha. 


Type material: Holotype (211881) and one paratype (211882) in ANSP. Not examined. 
Other material examined: Panama: 2 s, Portobello (CHL); 3 s, Bocas Island, topotypes (CHL). 


Description: The original description 
(OLSssSON é£ MCGINTY, 1958) is rather 
complete: “The shell is small (greater 
diameter, 1.4 to 1.7 mm.), thin, white or 
subtranslucent, depressed, with a large 
peripheral keel. The spire is slightly ele- 
vated, composed of 2 Y to 3 whorls of 
which the nuclear portion of 1 Y whorls 
is relatively large, smooth, helicoid in 
shape. The postnuclear whorls are a little 
convex or vaulted by a large, angular, 


submedial ridge; on the inner side of this 
rmáage “the. surface 15 flattened to 
depressed, the outer side a little wider 
and slopes convexly towards the outer 
suture or towards the peripheral keel. 
The basal section of the shell is more 
depressed than the upper and likewise 
carries a submedial ridge or angle; 
within this ridge the surface has the 
shape of a vortex which dips into a deep 
umbilicus showing the inner volutions of 


187 


Iberus, 29 (2), 2011 


Figures 106A-E. Vitrinorbis elegans Olsson 82 McGinty, 1958. A-C: shells, 1.6, 1.3, 1.6 mm, Bocas 
Island, Panama (CHL); D: detail of the sculpture; E: protoconch. 

Figuras 106A-E. Vitrinorbis elegans Olsson e McGinty, 1958. A-C: conchas, 1,6, 1,3, 1,6 mm, lsla 
Bocas, Panamá (CHL); D: detalle de la escultura; E: protoconcha. 


the spire whorls. Surface on both the 
dorsal and ventral sides has a delicate 
satiny texture produced by a sculpture of 
fine, microscopic spiral threads minutely 
cancellated by still finer lines of growth; 
in the sutural areas, the growth lines are 
heavier and may develop into axials 
nearly as large and strong as the spiral 
threads; aperture subovate, strongly 
oblique, attached weakly to the body 
whorl between the basal ridge and the 


188 


peripheral keel; no parietal callus. Holo- 
type greater diameter 1.4 mm”. 

Maximum reported size: 1.7 mm. Of 
the figured specimens, the larger is 
about 1.60 mm in diameter and the other 
about 1.27 mm in diameter and 0.59 in 
height. 

Habitat: Nothing is known about the 
habitat of this species. The type material 
was obtained by sorting beach drift. 


Depth: 0 m. 


RUBIO ET AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Distribution: Costa Rica (HOUBRICK, 
1968); Panama (OLsson € MCGINTY, 
1958). 

Remarks: This is the only species con- 
sidered to be in the genus described in 


the Caribbean, the two previously 
known species are from the Panamic 
Province. This is a very characteristic 
species easily recognized by its delicate 
ornamentation and characteristic profile. 


SPECIES NOT REPRESENTED IN OUR MATERIAL 


Anticlimax schumoi (Vanatta, 1913) 


Discopsis schumo1 Vanatta, 1913. Proc. of the Acad. of Nat. Sci. of Philadelphia, 65: 24-25, pl. 2, 
figs. 2, 7. [Type locality: Monkey River, British Honduras]. 


Type material: In ANSP (76581). 


Descrip tion: In VANATTA (1913). 

Maximum reported size: 2.5 mm. 

Distribution: British Honduras: 
Belize (VANATTA, 1913) Panama 


(RADWIN, 1969), Venezuela: unlocalized 
(PRINCZ, 1982). 

Remarks: This species is not Teinos- 
toma schumo1i Vanatta, 1913. 


Cochliolepis surinamensis Altena, 1966 


Cochliolepis surinamensis Altena, 1966. Zoologische Mededelingen, 41: 236-237, figs. 3a-d. [Type 
locality: shell ridge at the “Kerkplein” (Church square) in Paramaribo, Surinam, at 1 m depth]. 


Type material: In RNHL. 


Description: In ALTENA (1966). 

Maximum reported size: 2.7 mm. 

Distribution: Venezuela: unlocalized 
(PriNcCz, 1982). 


Remarks: Described as 
Holocene fossil (ALTENA, 1966, 
1975); recorded as living by PRINCZ 
(1982). 


Discopsis omalos (de Folin, 1870) 


Adeorbis omalos Folin, 1870. Les Fonds de la Mer 1: 190-191, pl. 23, figs. 6-7. [Type locality: 


Pointe-a-Pitre, Guadeloupe]. 


Discopsis omalus (sic): Error for D. omalos by JOHNSON (1934). 


Description: In De FOLIN (1870). 
Maximum reported size: 1.7 mm. 
Distribution: Guadeloupe. 
Remarks: PILSBRY 2 OLSSON (1945b) 
wrote that “D. omalos has not been 
found, to our knowledge, since the 
original collection”. However the name 
Discopsis omalos has been used for 
Europe by NORDSIECK (1968), and West 
Africa, albeit with reservations, 
because the species was supposed to 
be Caribbean. The claim of PILSBRY éz 


OLssoN (1945b) and the absence of 
specimens of this species in the mater- 
ial studied for this work, leads us to 
consider the possibility that the species 
is not really from the Caribbean and 
may have a limited distribution in the 
eastern Mediterranean and West 
Africa. De Folin was the captain of 
Bayonne harbour and received anchor 
mud from ships worldwide, which 
makes the possibility of mixing locali- 
ties quite likely. 


189 


Iberus, 29 (2), 2011 


Pleuromalaxis pauli Olsson £z McGinty, 1958 


Pleuromalaxis pauli Olsson £ McGinty, 1958. Bulletins of American Paleontology, 39: 30, pl. 3, 


figs. 3-3a. [Type locality: Bocas Island, Panama]. 


Type material: In ANSP (211902). 


Description: In OLSSON £ MCGINTY 
(1958). 
Maximum reported size: 1 mm. 


Distribution: Panama; Trinidad « 
Tobago: Tobago (OLSSON éz MCGINTY, 1958). 
Depth: 0 m. 


Solariorbis hondurasensis (Vanatta, 1913) 


Teinostoma hondurasensis Vanatta, 1913. Proceedings of the Academy of Natural Sciences of 
Philadelphia, 65: 26, pl. 2, figs. 8, 12. [Type locality: Belize and Monkey River, British Hon- 


duras]. 


Type material: In ANSP (76535). 


Description: In VANATTA (1913). 
Maximum reported size: 1.8 mm. 
Distribution: British Honduras: Belize 


(VANATTA, 1913); Panama (RADWIN, 1969). 
Remarks: MOORE (1964) placed this 
species in Solariorbis. 


Solariorbis petitii (P. Fischer, 1857) (Figures 107A-B) 


Skenea petitii P. Fischer, 1857c. Journal de Conchyliolo g ie, 6: 288. [Type locality: Guadeloupe]. 


Type material: Syntypes in MNHN (23240), the better preserved one (Fig. 107) is hereby designated 


the lectotype. 


Description: In P. FISCHER (1857). 

Maximum reported size: 2.5 mm. 

Distribution: Guadeloupe  (P. 
FISCHER, 1857). 

Remarks: MOORE (1964: 110-111): 
“De tupe o) ts species comidos 
be found in the Paul Fischer type 
collection in the Laboratoire de 
Malacologie in Paris. However, in 
the general collection of the Labora- 
toire there is a card bearing a glass 
tube containing four specimens. The 
card bears the following informa- 
on Ad ecorbis petite cl 
(86)”. In his discussion of his new 
species, Fischer states that soft parts 
and operculum resemble Skenea 
planorbis, while the shell has more 
the aspect of an Adeorbis. The shells 
fit the description of Skenea petiti 


190 


with only one discrepancy, the 
diameter is 3 Y mm instead of 
Fischer's figure of 2 Y mm. Fischer 
apparently wrote Adeorbis on the 
card while working on the collection 
from the Antilles and did not bother 
to change it later. Thus the writer 
considers the four specimens to be 
the syntypes of Fischer's lost 
species. It has never been figured. 

We have examined the photo- 
graph of one of the best preserved 
syntypes in MNHN and saw that 
the poor state of conservation pre- 
cludes morphological comparison. 
Some small spiral cordlets on the 
external margin of the lip and the 
shape of the umbilicus suggest some 
similarity with Solariorbis multis- 
triatus. 


RUBIO ET A4L.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 107A-B. Solariorbis petitii (P. Fischer, 1857). A-B: Lectotype (MNHN). 
Figuras 107A-B. Solariorbis petitii (P Fischer, 1857). A-B: Lectotipo (MNAN). 


Teimostoma avunculus Pilsbry, 1953 
Teinostoma (Pseudorotella) avunculus Pilsbry, 1953. Monographs of the Academy of Natural 
Sciences of Philadelphia, 18: 413-414, pl. 49, fig. 3-3d. [Type locality: Plio-Pleistocene of North 
St. Petersburg, Florida]. 


Type material: In ANSP (18917). 


Description: In PILsBRY (1953). Depth: 11 to 18 m. 

Maximum reported size: 2.3 mm. Remarks: Fossil species described by 

Distribution: USA: Florida, Texas PILSBRY (1953); Recent species by ODÉ 
(ODÉ, 19876b). (1987b, 1988). 


Teinostoma morlierei Jousseaume, 1872 


Range: 14.5%N, 61%W. 
Maximum reported size: 3.5 mm. Distribution: Martinique. 


19 


Iberus, 29 (2), 2011 


Teinostoma parvum (Stimpson, 1851) 


Rotella parva Stimpson, 1851 (dubious name). 


Range: SANO 990 NOS WWEStO 
782W. 
Distribution: USA: North Carolina. 


References: KURTZ (1860); PORTER 
(1974). 


Vitrinella anomala (d4'Orbigny, 1842) (Figures 108A-C) 


Trochus (Rotella) anomala d'Orbigny, 1842. Mollusques. Histoire Physique, Politique et Natu- 
relle de líle de Cuba 2: 64, pl. 18, figs. 32-34. [Type locality: Saint-Thomas]. 


Type material: In NAMUK. 


Description: D'ORBIGNY (1842) 
in Sagra: “Concha orbicular muy 
deprimida, translúcida, muy lisa, 
brillante, ligeramente aquillada en 
su contorno; ombligo abierto sin 
encostramiento calcareo, verifican- 
dose el engrosamiento calcareo por 
dentro del borde columelar. Espira 
casi horizontal, compuesta de cinco 


vueltas deprimidas. Boca oblonga, 


oval, celbbordescolunielaimbuv 
grueso. Color blanco vítreo”. 

Maximum reported size: 
Oman: 


Distribution: Colombia (Díaz 
MERLANO é PUYANA HEGEDOS, 
1994); Virgin Islands: St. Thomas 
(D'ORBIGNY, 1842). 


Vitrinella carinata (d'Orbigny, 1842) 


Trochus (Rotella) carinata d'Orbigny, 1842. Mollusques. Histoire Physique, Politique et Naturelle 
de l'ile de Cuba 2: 62-63, pl. 18, figs. 26-28. [Type locality: Saint-Thomas]. 


Type material: The material in USNM was not found (Yolanda Villacampa, pers. comm.). The mate- 


rial in NAMUK apparently had the labels changed and was a different species. 


Descrip tion: In D'ORBIGNY (1842). 

Maximum Reported Size: 1.5 mm 

Distribution: USA: North Carolina 
(PORTER, 1974); Virgin Islands: St. 


Thomas (D'ORBIGNY, 1842b). Depth: 
ZA 

Remarks: Nomen dubium according 
to MOORE (1964). 


Vitrinella hemphilli Vanatta, 1913 


Vitrinella hemphilli Vanatta, 1913. Proceedings of the Academy of Natural Sciences of Philadel- 
phia 65: 24, pl. 2, figs. 1, 3. [Type locality: Cedar Keys, Florida]. 


Type material: In ANSP (10236). 


Description: In VANATTA (1913). 

Maximum reported size: 2.5 mm. 

Distribution: USA: Florida: West 
Florida (VANATTA, 1913); Texas (ODÉ, 
1987c); Colombia (Díaz MERLANO éz 


192 


PUYANA HEGEDUS, 1994). Depth: 0.6 to 
73 m (alive at 51 m). 

Remarks: MOORE (1964):  “V. 
hemphilli has a dorsal spiral cord which 
disappears after one turn. In this, it is 


RUBIO E7 41.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 108A-G. Syntypes of Rotella. A-C: Rotella anomala (d1'Orbigny, 1842), syntypes in 
NHMUK; D-F: Rotella striata (d'Orbigny, 1842), syntypes in NHMUK; G: Rotella diaphana 
('Orbigny, 1842), syntype in NHMUK. 

Figuras 108A-G. Sintipos de Rotella. A-C: Rotella anomala (4'Orbigny, 1842), sintipos en NAMUK; 
D-F: Rotella striata (4'Orbigny, 1842), sintipos en NHMUK; G: Rotella diaphana (4'Orbigny, 
1842), sintipo en NHAMUK. 


like V. helicoidea, but the pointed spire the umbilicus separates it from that 
and absence of a strong carina around species”. 


Vitrinella pusilla (L. Pfeiffer, 1840) (Figs. 108G, 109A-E) 


Rotella pusilla Pfeiffer, 1840. Archiv fúr Naturgeschichte, 6(1): 255. Not figured [Type locality: 
Cuba (Nordkiiste, L. PFEIFFER 1839: 349; Matanzas Bay or Cardenas, based on localities cited 
by L. PFEIFFER, 1854]. 

Trochus (Rotella) diaphana d'Orbigny, 1842. Mollusques. Histoire Physique, Politique et Natu- 
relle de lle de Cuba 2: 62, pl. 18, figs. 23-25. [Type locality: Saint-Thomas]. 

Pseudorotella pusilla (L. Pfeiffer, 1840). 


Type material: One syntype in NHAMUK (Fig. 108G]. In USNM (35431) (secondary type collection: 
as Rotella diaphana). 


195 


Iberus, 29 (2), 2011 


Aedo 1897 


% / 
DL, e CLOS, 


Mus. 3543 / U. S Fish don 
4ne 


Sta. O, AU 


4 


* 1013031354] 


Smithsonian 


Q) 


€ 


Figures 109A-E. Vitrinella diaphana (4'Orbigny, 1842b). A-B: specimens identified by Bush, from 


off Cape Hatteras; 1.27 mm, USNM (35431); C: protoconch; D-E: labels. 
Figuras 109A-E. Vitrinella diaphana (4'Orbigny, 1842b). A-B: ejemplares identificados por Bush, pro- 
cedentes de frente a Cape Hatteras; 1,27 mm, USNM (35431); C: protoconcha; D-E: etiquetas. 


Description: Original description of 
Rotella pusilla in PFEIFFER (1840): 
“Testa discoidea nitide alba; anfract. S; 
basi concaviuscula, medio callosa: 
aperturasorbiculari. Diam. 4 al eS 
lora 

Original description of Trochus 
(Rotella) diaphana in D'ORBIGNY, 1842: 
“Coquille orbiculaire, deprimee, dia- 
phane, trés lisse, tres polie, convexe du 
cóte de la espire, legérement concave 
du cóte de la bouche, la callosite étant 
peu etendue. Spire peu elevée, tres 
obtuse, composée de quatre tours 


194 


convexes, á pourtour arrondie. Bouche 
ovale, oblique. Couleur: blanc transpa- 
rentcomme du verre. 

Maximum reported size: 1.5 mm. 

Distribution: Cuba: North Havana 
Province (ARANGO, 1880); USA: North 
Carolina (Bush, 1897); Panama: Bocas 
Island (OLsson éz MCGINTY, 1958). 

Remarks: The types of Rotella 
pusilla Pfeiffer, 1840 are untraceable. L. 
PFEIFFER (1840) offered a short and 
ambiguous text and no figure. Conse- 
quently it may be confused with 
several taxa. The type material is from 


RUBIO £7 AZz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Cuba, but we have not found any 
museum or shell collection with mate- 
rial labeled R. pusilla, which could 
serve as a reference. Therefore we con- 
sider Rotella pusilla L. Pfeiffer, 1840 as 
a nomen dubium until the type mater- 
ial is located. Under these conditions 
the synonymy established for Vit- 
rinella diaphana can not be accepted. 

The types of Vitrinella diaphana 
Orbigny, 1842 in NHMUK are in very 
poor condition (Fig. 108G). In the sec- 
ondary type collection of the USNM 
(35431) are deposited 2 shells labeled 
by Bush (1897) as V. diaphana 
(d'Orb.), from Cape Hatteras, North 
Carolina. 


This taxon has been included in the 
following genera: Parkeria, Adeorbis 
and Teinostoma. 

We have some doubts about the 
accuracy of the identification made by 
BusH (1897) due to its great similarity to 
Solariorbis terminalis. The differences 
between them are quite minor: The shell 
labeled as V. diaphana lacks the subsu- 
tural sulcus, its periphery is more 
rounded, and some wide depressed 
spiral cords are visible on the whole 
surface of the shell. At present we shall 
keep the taxa separate until topotypes of 
V. diaphana become available. Such 
material should help in the correct 
assignament of both nominal taxa. 


Vitrinella tryoni Bush, 1897 


Vitrinella helicoidea auct. non C.B. Adams, 1850. 


Vitrinella tryoni Bush, 1897. Transactions of the Connecticut Academy of Arts and Sciences, 10: 
123, pl. 22, figs. 11-11a. [Type locality: USFC sta. 2278, off Cape Hatteras, North Carolina, 16 


fms (29 m)]. 
Type material: Holotype in USNM (41561). 


Descrip tion: BusH (1897). 

Maximum reported size: 2 mm. 

Distribution: USA: North Carolina 
(Bush, 1897); Cuba: North Havana 
Province, North Matanzas (AGUAYO € 
JAUME, 1936). 

Depth: 29 m. 


Remarks: The species was misidenti- 
fied by TRYON (1888: 102, pl1.34, figs. 40, 
41) as V. helicoidea C.B. Adams. The 
type of this species (USNM 41561) is 
only a fragment and canot be identified 
at the species level (Yolanda Villacampa 
and Jerry Harasewych, pers. comm.). 


FOSSIL OR SUPPOSEDLY FOSSIL SPECIES, NOT RECENT 


Many of the fossil species described 
in some papers (for example, PILSBRY, 
1953) have been recorded in other works 
as also occurring in the recent. Other 
species, already known as fossil, are 


added in this paper to the recent fauna 
for the first time. Others yet, only 
known as fossil species, are not the main 
topic of this work. Of the last group, we 
can mention the following; 


Anticlimax athleenae (Pilsbry £ McGinty, 1946) 


Climacia athleenae Pilsbry ££ McGinty, 1946. The Nautilus, 59: 78-79, pl. 8, figs. 3-3a. [Type local- 


ity: Boca Ciega Bay]. 


Type material: In ANSP (181291). 


195 


Iberus, 29 (2), 2011 


Figures 110A-F. Cyclostremiscus fargoi Pilsbry, 1953. A-C: shell, 1.6 mm, Pliocene of La Belle, 
Florida (CHL); D-F: shell, 1.4 mm, Pliocene of La Belle, Florida (CHL). 
Figuras 110A-E Cyclostremiscus fargoi Pilsbry 1953. A-C: concha, 1,6 mm, Plioceno de La Belle, 
Florida (CHL); D-E: concha, 1,4 mm, Plioceno de La Belle, Florida (CHL). 


196 


RUBIO £7 41.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Description: In PiLsBRY € MCGINTY Remarks: MOORE (1964) men- 
(1946). tions: “This species is known from a 
Maximum reported size: 2.6 mm. single. specimen taken Miome $11 


Distribution: USA: Florida: West Florida dredged from Boca Ciega Bay. It is 
(Pilsbry € McGinty, 1946a); Colombia (DÍAZ here comsidered to be a fossil of 
MERLANO éz PUYANA HEGEDUS, 1994). unknown age”. 

Aorotrema gardnerae Pilsbry, 1953. 
Aorotrema gardnerae Pilsbry, 1953. Monographs of the Academy of Natural Sciences of 
Philadelphia, 18: 424, pl. 39, fig. 13. [Type locality: Upper Miocene, Natural Well, Duplin Co., 
North Carolina]. 


Type material: In ANSP (19550). 


Cyclostremiscus fargoi Pilsbry, 1953 (Figures 110A-F) 


Cyclostremiscus fargo1 Pilsbry, 1953. Monographs of the Academy of Natural Sciences of Philadel- 
phia, 18: 424-425, pl. 56, figs. 1-1c. [Type locality: Plio-Pleistocene, St. Petersburg, Florida]. 


Type material: ANSP (18399). 
Other material examined: USA: 2 s, Pliocene of La Belle, Florida (CHL). 
Cyclostremiscus gunteri (Mansfield, 1930) 


Circulus gunteri Mansfield, 1930. Florida State Geol. Survey Bull. 3: 132, pl. 20, figs. 16-18. [Type 
locality: Upper Miocene, Leon County, Florida]. 


Type material: In USNM (370493). 


Cyclostremiscus olssoni Pilsbry, 1953. 


Cyclostremiscus olssoni Pilsbry, 1953. Monographs of the Academy of Natural Sciences of 
Philadelphia, 18: 426, pls. 54, figs. 6-6c. [Type locality: Plio-Pleistocene, Shell Creek, Florida]. 


Type material: In ANSP (18453). 


Parviturboides avitus Pilsbry, 1953. 


Parviturboides avitus Pilsbry, 1953. Monographs of the Academy of Natural Sciences of 
Philadelphia, 18: 436, pl. 56, figs. 3-3a. [Type locality: Plio-Pleistocene, St. Petersburg, Florida]. 


Type material: In ANSP (18460). 


Solariorbis eugenes Pilsbry, 1953 (Figures 111A-D) 


Solariorbis eugenes Pilsbry, 1953. Monographs of the Academy of Natural Sciences of Philadel- 
phia, 18: 418-419, pl. 54, figs. 1, 1a-b. [Type locality: Plio-Pleistocene, St. Petersburg, Florida]. 


197 


Iberus, 29 (2), 2011 


Figures 111A-D. Solariorbis eugenes Pilsbry, 1953. A-C: shell, 4.3 mm (CHL); D: protoconch. 
Figuras 111A-D. Solariorbis eugenes Pilsbry 1953. A-C: concha, 4,3 mm (CHL); D: protoconcha. 


Type material: In ANSP (18413). 


Other material examined: USA, Florida: 1 s, fossil Pleistocene, ST. Petesburg (CHL). 


Description: This is the original 
description in PILsBRY (1953): “The shell 
is solid, somewhat lens-shaped, with 
low spire, rounded periphery and a 


198 


rather narrow, deep umbilicus. There 
are 3 Y whorls, the first 1 Y relatively 
large, convex and smooth, the next 
whorl with about 8 or 9 narrow and 


RUBIO £7 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


shallow spiral grooves more or less dis- 
tinctly interrupted by retractive radial 
striae crossing the grooves. On the last 
whorl this sculpture is nearly or wholly 
lost, leaving it almost smooth except for 
a distinct subsutural margin defined by 
a spiral groove. The aperture is oblique, 
subcircular but with a grooved upper 
angle. The peristome is rather thin. The 
columella is widened by a flat, triangu- 
lar callus at the end of the rounded 
ridge which surrounds and contracts the 


umbilicus. The parietal callus is thin, its 
outer edge indistinct. Diameter 4.2 mm, 
height 2.5 mm”. 

Remarks: Plio-Pleistocene  fossil 
species of St. Petersburg, Florida. We do 
not know any current reference for this 
species. PILSBRY (1953) commented: 
“This appears to be a typical Solariorbis, 
having sculpture on the penult whorl 
like the type, becoming almost smooth 
at the last whorl”. The shell figured does 
not appear to be a fossil shell. 


Teinostoma caloosaense Dall, 1892 


Teinostoma caloosaense Dall, 1892. Trans. Wagner Free Inst. Sci., 3: 413, pl. 23, fig. 8. [Type local- 
ity: Plio-Pleistocene, Caloosahatchie River, Florida]. 


Type material: In USNM (113110). 


Teinostoma tectispira Pilsbry, 1953 


Teinostoma tectispira Pilsbry, 1953. Monographs of the Academy of Natural Sciences of 
Philadelphia, 18: 417, pl. 50, figs. 6-6c. [Type locality: Plio-Pleistocene, St. Petersburg, Florida]. 


Type material: In ANSP (18406). 


SPECIES FROM NEIGHBOURING GEOGRAPHIC AREAS 


Neusas marshalli (Sykes, 1925) (Figures 112A-C) 


Homalogyra (?) marshalli Sykes, 1925: 193, pl. 9, fig. 9-9a [Type locality: off Portugal, 3942'N, 
09%43'W, 1092-1993 m, Porcupine Expedition sta. 17]. 


Description: SYKES (1925): “Mor- 
phology: Shell planispiral, resem- 
bling a planorbid, with rounded and 
almost smooth whorls with a deep 
suture. Protoconch tall-spired and 
obliquely inserted, with slightly 
more than mo mwhoris. smooth. 
Teleoconch with about three slightly 
irregularly coiled whorls. Operculum 
corneous, multispiral, round with 
central nucleus”. 


Size: Shell diameter up to 2.06 
mm. 

Distribution: See WARÉN € 
BOUCHET (2001). Mid-Atlantic Ridge: 
Menez Gwen hydrothermal vent, 
Azores (37.84 N; 31.522W to 9.722W). 
Depth: 870 to 860 m (collected alive). 

One shell (Fig. 112 D) of Neusas 
s.p (Absaláo, pers. com.) showed the 
wide distribution of this genus in 
south Atlantic. 


199 


Iberus, 29 (2), 2011 


Figures 112A-D. Neusas marshalli (Sykes, 1925). A-C: shells, 2 mm, Menez Gwen hydrothermal 
vent, Azores, Atlantic Ocean (with authorization of A. Warén); D: Neusas sp., 1.1 mm, Campos 
Basin, Rio de Janeiro, Brazil (with authorization of R. Absaláo). 

Figuras 112A-D. Neusas marshalli (Sykes, 1925). A-C: conchas, 2 mm, Menez Gwen hydrothermal 
vent, Azores, Atlantic Ocean (con la autorización de A. Warén); D: Neusas sp., 1.1 mm, Campos 
Basin, Rio de Janeiro, Brazil (con la autorización de R. Absaláo). 


Ponderinella xacriaba Absaláo, 2009 (Figures 113A-C) 


Ponderinella xacriaba Absaláo, 2009. American Malacological Bulletin, 27: 138, figs. 2E-H. 
[Type locality: BC Sul L, sta. 73, Campos Basin, Rio de Janeiro State, Brazil, 22%41'35”S, 
40%00'45”W, 1950 m]. 


Description: ABSALAO (2009). Distribution: Brazil: Rio de Janeiro. 
Maximum reported size: 1.23 mm. Depth: 1030 to 1950 m. 


Teinostoma abnorme E.A. Smith, 1890 


Teinostoma ?abnorme E.A. Smith, 1890. Proceedings of the Zoological Society of London, 1890: 
293, pl. 24, fig. 5. [Type locality: St. Helena]. 


Description: In SmITH (1890). Distribution: Eastern Atlantic: St. 
Maximum reported size: 1 mm. Helena. Only known from its type locality. 


200 


RUBIO E7 4Z.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 113A-C. Ponderinella xacriaba Absaláo, 2009. (Photographs authorized by R. Absalio) 
Figuras 113A-C. Ponderinella xacriaba Absaláo, 2009. (Fotografías autorizadas por R. Absaláo) 


SPECIES RECORDED ERRONEOUSLY FROM THE CARIBBEAN 


Adeorbis elegans (A. Adams, 1850) 


Cyclostrema elegans A. Adams, 1850. Proceedings of the Zoological Society of London, 18: 44. 
[Type locality: Sibonga, island of Cebu, Philippines, 10 fathoms (18 m)]. 


Remarks: It has been reported from its presence in the Caribbean is very 
St. Thomas but this is a species improbable. 
described from the Philippines and so 


Vitrinella regularis (C.B. Adams, 1852) 


Renmiuanes:. Holotype.. n= MCEZ cating “Jamaica”. Probably it is an error, 
(156374). With the shell is a label indi- because it is a Pacific (Panamic) species. 


201 


Iberus, 29 (2), 2011 


SUPPLEMENT 


DUBIOUS SPECIES, APPARENTLY NOT TORNIDAE 


These taxa are not the subject of the 
present work. Nevertheless, some of 
them have been recorded dubiously or 


positively in Vitrinellidae. As we 
obtained photographs, we present some 
of them as information for the reader. 


“Aorotrema” erraticum Pilsbry £ McGinty, 1945 (Figure 114A) 


Aorotrema erraticum Pilsbry £ McGinty, 1945a. The Nautilus, 59: 1, pl. 11. [Type locality: 1.5 


miles off Cape Florida, 12 fms (22 m)]. 


Material examined: 1 s, Cienfuegos, Cuba (MHNS). 


Remarks: MOORE (1964: 189) said: 
“the examination of the holotype of A. 
erraticum informs that it is a juvenile of 
Turbo castaneus Gmelin due to the coin- 


cidence of all the details, protoconch, 
shape and sculpture”. We accept this 
opinion and represent a sample of this 


shell. 


“Vitrinella tincta” C.B. Adams, 1850 (Figs. 114B-D) 
Vitrinella tincta C.B. Adams, 1850. Monograph of Vitrinella: 8. 


Type material: The lectotype in MCZ (156257) (figured in CLENCH 6: TURNER, 1950, plate 35 fig. 6) 
and 1 second specimen (also figured in CLENCH éz TURNER, 1950, plate 35 fig. 3) labeled as paratype, 
in MCZ (186189). 

Other material examined: Virgin Islands: 1 s, beach at Magens Bay, N coast of St. Thomas, Virgin 


Is. (CHL). 


Remarks: This species is repre- 
sented by some drawings of the types, 
and the description is commented on 
in CLENCH é TURNER (1950). The 


appearance is not that of a valid 
species but rather a juvenile Tegula as 
suggested by its colored spots aligned 
spirally. 


“Vitrinella” carinifex Dall, 1927 (Figures 115A-D) 


Vitrinella? Carinifex Dall, 1927. Proceedings of the United States National Museum, 70(2667): 


126. [Type locality: Off Georgia]. 


Type material: Lectotype in USNM (108399) (Figs. 115A-D). 


Description: In DaLL (1927). 

Distribution: Only known from the 
type material, from Georgia, USA. 
Range: 30.739; 79.432W. Depth: 805 m. 

Remarks: DALL (1927) comments: 
“This is probably not a true Vitrinella 
and may be immature but certainly is 


202 


not the young of any of the species enu- 
merated in this discussion”. 

The lectotype of Vitrinella carinifex 
in USNM is not a Vitrinella or a tornid. 
In our opinion perhaps it is a young 
naticid (see the notch in the columella). 
Maximum reported size: 3.7 mm. 


RUBIO ET 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figure 114A. “Aorotrema” erraticum Pilsbry 82 McGinty, 1945. A: shell, 5.1 mm, Cienfuegos, Cuba 
(MHNS). Figures 114B-D. Vitrinella tincta C.B. Adams, 1850. Lectotype (labeled as syntype) 
(MCZ 156257). 
Figura 114A. “Aorotrema” erraticum Pilsbry € McGinty 1945. A: concha, 5,1 mm, Cienfuegos, Cuba 
(MANS). Figuras 114B-D. Vitrinella tincta C.B. Adams, 1850. Lectotipo (etiquetado como sintipo) 
(MCZ 156257). 


Vitrinella cyclostomoides (L. Pfeiffer, 1840) 


Helix (?) cyclostomoides Pfeiffer, 1840. Archiv fúr Naturgeschichte 6(1): 251, [Pfeiffer, 1850, pl. 
85, figs. 24-26]. [Type locality: Cuba [Nordkúste (L. PFEIFFER, 1839: 349); Matanzas (L. PFEIFFER, 
1854c)]. 


Description: L. PFEIFFER (1840). Remarks: This species has been 
Maximum reported size: 3.2 mm. treated as both terrestrial and marine. 
Distribution: Cuba: North Matanzas: L. PFEIFFER (1840) named it a Helix 
L. PFEIFFER (1840); L. PFEIFFER (1852). but found his specimen(s) at the 
Depth: 0 m. “Seestrande”. He later expressed doubt 


203 


Iberus, 29 (2), 2011 


Figures 115A-D. *Vitrinella” carinifex Dall, 1927. A-C: lectotype, 3.5 mm in diameter (USNM 
108399); D: protoconch. (SEM's by Yolanda Villacampa, USNM). 

Figuras 115A-D. “Vitrinella” carinifex Dall, 1927. A-C: lectotipo, 3,5 mm de diámetro (USNM 
108399); D: protoconcha. (Fotografías al MEB de Yolanda Villacampa, USNM,). 


that it was a landsnail (L. PFEIFFER, 
1854: 179). ARANGO (1880) treated it as 
the valid name for Adeorbis adamesii 
and ESPINOSA ET AL. (1995) regarded it 
as a valid species of Vitrinella. TRYON 
(1887: 100), however, treated it as a 
Microphysa and RICHARDSON (1986) 
placed it in Hojeda (Sagdidae). It is 


not listed among the Cuban Hojeda 
by ESPINOSA éz ORTEA (1999), and the 
only Cuban Hojeda with a range that 
includes Matanzas is Hojeda boothi- 
ana (L. Pfeiffer, 1839), a taxon that 
Pfeiffer clearly did not consider con- 
specific with his elo 
cyclostomoides. 


Vitrinella hyalina C.B. Adams, 1850 


Vitrinella hyalina C.B. Adams, 1850e. Monograph of Vitrinella, a New Genus of New Species of 
Turbinidae: 5, Not figured. [Type locality: Port Royal, Jamaica]. 


Type material: Lost (CLENCH éz TURNER, 1950). 


Description: C.B. Adams (1850). 
Maximum reported size: 1.7 mm. 
Distribution: Jamaica: C.B. ADAMS (1850). 


Remarks: Possibly this is a naticid 
(see MOORE, 1964: 51). 


204 


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VITRINELLID SPECIES TRANSFERRED TO OTHER FAMILIES 


Family TURBINIDAE Rafinesque, 1815 


Some genera of this family are com- 
posed of small, lenticular or depressed shells 
with a low spire, and for these reasons they 


may be confused with Tornidae. Occasion- 
ally, some of them have been placed in this 
family so we make reference to them here. 


Subfamily SKENEINAE Clark, 1851 


Genus Cirsonella Angas, 1877 


Cirsonella Angas, 1877: 38. Type species, by monotypy, Cirsonella australis Angas, 1877 (Fig. 


11A), southern Australia. 


Tharsis Jeffreys, 1883: 93 (not Giebel, 1847). Type species, by monotypy, Oxistele romettensis 


Granata, 1877, Mediterranean. 


Tharsiella Bush, 1897: 113. Replacement name for Tharsis Jeffreys, 1883 not Giebel, 1847. 
Porcupinia Cossmamn, 1900: 43. Replacement name for Tharsis Jeffreys, 1883. 


Porcup ina Cossmamn, 1925: 287. Misspelling. 


Diagnosis: In WARÉN (1991: 159): 
“Small skeneimorph gastropods with 
almost globular, smooth shell, almost 
round, prosocline aperture, with thicken- 
ing at umbilicus. Protoconch finely and 
irregularly spirally striated. Operculum 
sturdy, yellowish with long growth 
zone. Radula with four to five undiffer- 
entiated lateral teeth and well developed 
basal plate on innermost marginal tooth. 
Propodial penis not present”. 


The operculum of Cirsonella differs 
from the typical species of Skeneidae, 
having its last 1/3 whorl slowly taper- 
ing, while in Skeneidae, it ends 
abruptly with an oblique edge cover- 
ing about 1/20 of the whorl. Another 
differential characteristic is that species 
of Cirsonella retract the operculum 
only very slightly, or not at all, behind 
the peristome, contrary to most 
skeneids. 


Cirsonella floridensis (Dall, 1927) (Figures 116A-C) 


Pseudorotella floridensis Dall, 1927. Proceedings of the United States National Museum 
70(2667): 126. [Type locality: Albatross sta. 2668, off Fernandina, Florida [actually off Georgia], 


30%58'N, 79*38'W, 294 fathoms (529 m)l. 


Type material: Lectotype in USNM (108133). Examined by SEM microphotographs. 


Original description: In DaALL (1297): 
sshelle small, —porcelaneous- =whtte, 
smooth, polished, turbinate, with four 
well-rounded whorls; suture distinct, 
not deep; aperture nearly circular, the 
margin simple, continuous over the 
body with a marked deposit of enamel 
behind the body lip and over the umbil- 
ical region, completely closing the 
umbilicus and in the completely adult 
sometimes produced in subangulate 
form above and below. Height, 2 mm; 
maximum diameter, 3 mm”. 


We add the following: The proto- 
conch measures 260 um in maximum 
diameter, smooth and % of a whorl. 
There is no sculpture. There are two fine 
spiral cordlets in the umbilicus and 
there is no umbilical cord bordering it. 
In the figured specimen the columellar 
callus reduces the umbilicus to a fine 
fissure, however, as is explained in 
DaLL (1927), in other shells the callus 
totally covers the umbilicus. In the inner 
part of the aperture and along the col- 
umella and the inner lip there is a wide 


205 


Iberus, 29 (2), 2011 


Figures 116A-C. Cirsonella floridensis (Dall, 1927). A-B: lectotype, 2.2 mm in diameter (USNM, 
108133); C: protoconch. (SEM's by Yolanda Villacampa, USNM). Figures 116D-E Cirsonella 
georgiana (Dall, 1927); D-E: lectotype, 1.95 mm, (USNM, 329375); F: protoconch. (SEM's by 


Yolanda Villacampa, USNM). 

Figuras 116A-C. Cirsonella floridensis (Dall, 1927). A-B: lectotipo, 2.2 mm de diámetro (USNM, 

1 08133); C: protoconcha. (Fotografías al MEB deYolanda Villacampa, USNM). Figuras 116D-E Cir- 

sonella E iana (Dall, 1927); D-E: ed 1.95 mm, (USNM, 329375); F: protoconcha. (Foto- 
a 


grafías al MEB de Yolanda Villacampa, USNM,). 


206 


RUBIO ET 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


fold which is like an “opercular stop”, 
characterístic of the species of the genus 
Cirsonella. 

The studied shell is 2.2 mm in diame- 
ter and 2.2 mm in height. 

Habitat: Dredged from 678 meters 
depth, in gray sand and broken coral 
bottom. Abundant. 

Distribution: From off Fernandina 
(actually off Georgia) (DALL, 1927); from 
North Carolina (PORTER, 1974); ODÉ 
(1987a). 

Remarks: Actually this species has 
been placed in the genus Teinostoma. 
DaLL (1927) mentions: “This is much like 
Pseudotorella solida, but larger, more ele- 
vated, and with the umbilical pad heavier 


and more simple”. As in Lydipnis mar- 
garitiformis, the morphologic characters 
of the shell make one think it is a skeneid, 
most probably in the genus Cirsonella 
Angas, 1877. The shell studied is very 
similar to Cirsonella extrema Thiele, 1912 
(USNM 613041), in WARÉN (1991: 212, fig. 
11B), being distinguished from it by the 
lower number of spiral striae around the 
umbilicus. It is also similar to Cirsonella 
romettensis (Granata, 1877) even to the 
number of the spiral fillets around the 
umbilicus, but from that species it differs 
by its smooth vs spirally striated, proto- 
conch. Based on all the above, we 
propose the new combination Cirsonella 
floridensis (Dall, 1927). 


Cirsonella georgiana (Dall, 1927) (Figs. 116D-F) 


Vitrinella georgiana Dall, 1927. Proceedings of the United States National Museum 70(2667): 109. 


[Type locality: Off Georgia]. 


Type material: Lectotype and six paralectotypes in USNM (329375). The lectotype examined by 


SEM microphotographs. 


Original description: “Shell minute, 
glassy white, low turbiniform, with 
about three and a half smooth, well- 
rounded whorls; nucleus minute, 
suture distinct, not deep; aperture sub- 
circular, the peristome interrupted by 
the body whorl, not reflected, sharp; 
base roundly convex, with a minute 
perforate umbilicus, partly shadowed 
by the inner lip. Height 1.6; diameter, 2 

etlectotype is 1.95 mm in 
maximum diameter and 1.95 mm in 
height. 

Habitat: Dredged from 805 meters 
depth, on a broken coral, coarse sand, 
and broken shell bottom. 


Distribution: Only know from the 
USA: Georgia, the type locality. 

Remarks: As in other described 
species, no spiral fine cordlets have been 
observed in the umbilicus; but there is a 
strong cord delimiting it. Due to the 
poor condition of the studied material, 
we can not determine if this is a previ- 
ously known species. We believe that 
Vitrinella georgiana, the present species, 
as well as Pseudorotella floridensis and 
Lydipnis margaritiformis, all described 
by DaLL (1927), are not tornids, but they 
may be placed in the subfamily 
Skeneinae Clark, 1851, genus Cirsonella. 
Thus the new combination should be 
Cirsonella georgiana (Dall, 1927). 


Cirsonella margaritiformis (Dall, 1927) (Figures 117A-E) 


Lydiphnis margaritiformis Dall, 1927. Proceedings of the United States National Museum 
70(2667): 123-124. [Type locality: Albatross sta. 2668, off Fernandina, Florida [actually off 
Georgia], 30%58'N, 79238'W, 294 fathoms (529 m)]l. 


Circulus margaritiformis (Dall, 1927). 


Type material: Lectotype (Figs. 117A-C) and a paralectotype in USNM (108146). 


207 


Iberus, 29 (2), 2011 


Description: This is the original 
description in DALL (1927: 123-124). 
“Shell small, white, smooth, with nearly 
four whorls including a minute globular 
nucleus, having a general form much 
resembling Margarites helicinus; whorls 
moderately rounded, the last much the 
largest, suture distinct, not deep; surface 
smooth except for faint incremental 
lines, and on the base two strong widely 
spaced threads around the umbilical pit 
and a few faint spiral striae behind 
them; base rounded, umbilicus minutely 
perforate; aperture rounded with a 
small angulations above, the peristome 
continuous over the body, thin, and not 
reflected. Height, 3 mm; diameter, 4 
mm ?. 

Habitat: Dredged from 678 meters 
on a gray sand and broken coral bottom. 

Distribution: Only known from the 
type material. 

Remarks: DALL (1927) Remarked: 
“This differs from typical Lydipnis in 
being less depressed and without 
carinae, but seems most nearly allied to 
that genus”. 


In our opinion this species is not a 
tornid. Its general appearance is that of 
a skeneid. The  convex  whorls, 
impressed sutures of the protoconch, 
smooth shell, prosocline aperture, and 
continuous peristome with a thickening 
near the umbilicus place it close to the 
genus Cirsonella Angas, 1877. Cir- 
sonella characteristically has the opercu- 
lum only very slightly, or not at all, 
behind the peristome. The margin of the 
inner lip edge of this species has a ledge 
that probably serves as an opercular 
stop. The spiral threads around the 
umbilicus place this species close to Cir- 
sonella extrema Thiele, 1912 from the 
Antartic and to C. australis Angas, 1877 
(WARÉN, 1992). 

By its general form this species 
could be considered closely related to 
species of the genus Skenea Fleming, 
1825, e.g., Skenea larseni Warén, 1993 
and Skenea trochoides (Friele, 1876), but 
the shape of the protoconch and the 
umbilical thickening distinguish it. Thus 
this species is treated as Cirsonella mar- 
garitiformis (Dall, 1927). 


Genus Mikro Warén, 1996 
Mikro cerion (Dall, 1927) (Figures 118A-C) 


Vitrinella cerion Dall, 1927. Proceedings of the United States National Museum 70(2667): 125. 
[Type locality: Off Georgia, 30.732; 79.432W. Depth: 805 m]. 


Type material: Lectotype (Fig. 100A-C) in USNM (108433). 


Description: In DaLL (1927). 
Maximum reported size: 1.8 mm. 
Distribution: Only known from the 
type material, from Georgia, USA. 
Remarks: DALL (1927) mentions: 
“The general form recalls that of Helix 
chersina Say”. After the examination of 
the photographs of the lectotype of Vit- 
rinella cerion deposited in USNM, we 
reached the conclusion that it is not a 
Vitrinella. The smooth, bulbous and 
short protoconch (barely Y whorl), the 
teleoconch apically keeled on the first 
whorl and angled apically on the fol- 
lowing, the rounded aperture, the 
orthocline outer lip, and the strongly 


208 


prosocline inner lip impel us to con- 
sider its placement in Mikro Warén, 
1996, which is characterized by “Very 
small, skeneid-like gastropods with 
smooth protoconch, apically keeled 
first teleoconch whorl and intraumbili- 
calce els 

The bathymetric range of the 
species included in this genus is 
between 200 and 1100 m, and its distri- 
bution is known in Southern Iceland 
and the Mediterranean. The genus 
Mikro was placed provisionally in 
Skeneidae by WARÉN (1996), and its 
systematic position is not known with 
certainty, but it was placed in 


RUBIO £7 AZ.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 117A-E. Cirsonella margaritiformis (Dall, 1927). A-C: lectotype, 4 mm in diameter 
(USNM, 108146); D-E: protoconch. (SEM's by Yolanda Villacampa, USNM). 

Figuras 117A-E. Cirsonella margaritiformis (Dall, 1927). A-C: lectotipo, 4 mm de diámetro 
(USNM, 108146); D-E: protoconcha. (Fotografías al MEB de Yolanda Villacampa, USNM). 


209 


Iberus, 29 (2), 2011 


Figures 118A-C. Mikro cerion (Dall, 1927). A-B: lectotype, 1.6 ao la diameter (USNM, 


108433); C: protoconch. (SEM's by Yolanda Villacampa, USNM). 
Figuras 118A-C. Mikro cerion (Dall, 1927). A-B: lectotipo, 1,6 mm de diámetro (USNM, 108433); 
C: protoconcha. (Fotografías al MEB de Yolanda Villacampa, USNM). 


Archaeogastropoda. This appears well 
founded, based on the presence of sen- 
sorial papillae on the cephalic tentacles. 
For these reasons we consider that the 
correct name is Mikro cerion (Dall, 


1927). Other species known in the 
genus Mikro are: Mikro giustii (Bogi € 
Nofroni, 1989) from the Isle of Capri, 
Mediterranean Sea and Mikro globulus 
(Warén, 1996) from southern Iceland. 


Genus Xyloskenea Marshall, 1988 
Xyloskenea rhyssa (Dall, 1927) (Figures 119A-D) 


Vitrinella rhyssa Dall, 1927. Proceedings of the United States National Museum 70(2667): 125. 
[Type locality: Albatross sta. 2668, off Fernandina, Florida [actually off Georgia], 30%58'N, 


79238'W, 294 fathoms (529 m)]. 
Type material: Lectotype in USNM (1081276b). 
Descrip tion: In DaLL (1927). 
Dimensions: lectotype, 2.1 mm. 


Distribution: Only known from the 
type material: 30.98%N; 79.64-W. 


210 


Remarks: The morphological charac- 
ters of Vitrinella rhyssa are very similar 
to those of the genus Xyloskenea Mar- 
shall, 1988. This genus was provision- 


RUBIO E7 4Z.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 119. A-D. Xyloskenea rhyssa (Dall, 1927). A-C: lectotype, 2.1 mm in diameter (USNM, 
108127b); D: protoconch. (SEM's by Yolanda Villacampa, USNM). 

Figuras 119. A-D. Xyloskenea rhyssa (Dall, 1927). A-C: lectotipo, 2,1 mm de diámetro (USNM, 
108127b); D: protoconcha. (Fotografías al MEB de Yolanda Villacampa, USNM). 


ally placed in Skeneidae, having many 
species in bathyal depths, world wide, 
and is usually associated with sunken 
wood. Conchologically close species 
include Xyloskenea costulifera Marshall, 


1988 from New Zealand (the type 
species of the genus) and Xyloskenea 
naticiformis (Jeffreys, 1883) from off 
southwestern Iceland to off Portugal 
(WARÉN, 1996: 202, figs. 3A-D). 


Xyloskenea translucens (Dall, 1927) (Figures 120A-F) 


Lydiphnis translucens Dall, 1927b. Proceedings of the United States National Museum 70(2667): 


124. [Type locality: Off Georgia]. 
Circulus translucens (Dall, 1927). 


Type material: Lectotype and paralectotype in USNM (108434). 


Description: The original description 
in DALL (1927: 124) is as follows: “Shell 
minute, translucent white, the spire flat- 
tened, with three moderately rounded 
whorls; axial sculpture of incremental 
lines only; spiral sculpture of a thread 


slightly in front of the suture and 
another beyond the periphery on the 
base, tMHese “are. .hardly. prominent 
enough to be called carinae; the umbili- 
cal pit is wide and shallow, the umbili- 
cus itself minutely perforate; aperture 


Za l 


Iberus, 29 (2), 2011 


rather wide, rounded, oblique, the 
margin thin, continuous, straight near 
the umbilicus. Height, 1.5 mm; diame- 
Er 2 Me 

The protoconch (Figs. 102E-F) is 
about 210 um in maximum diameter 
and is % of a whorl, of which the first 
half has 4-5 spiral ribs the remaining 
part is smooth. A strong varix separates 
the protoconch from the teleoconch. 

Habitat: Dredged from 805 meters 
depth on a broken coral, coarse sand, 
and broken shell bottom. 

Distribution: Only known from the 
type material. 

Remarks: DALL (1927) commented: 
“Notwithstanding its small size 1t has 
the shell" characters cof the cenus 
(Lydiphnis). The incremental lines are 
rather pronounced”. 

In our examination of the shell of 
Lydipnis translucens we carefully noted 
the depressed spire and the bicarinate 
shell as well as the protoconch sculp- 
ture. 

Based on these characters, L. translu- 
cens Closely resembles species of 
Xyloskenea Marshall, 1988 such as 
Xyloskenea naticiformis (Jeffreys, 1883) 
and species of Ventsia Warén «€ Bouchet, 
1993 such as Ventsia tricarinata Warén éz 
Bouchet, 1993 in both protoconch and 
teleoconch characters. These two genera 


can be distinguished only by the radula, 
being very similar in shell and soft parts 
and considered closely related. Another 
related genus is Trenchia Knudsen, 1964, 
type species T. wolffi Knudsen, 1964, the 
species of which have a similar shell 
and radula, but their protoconch and 
teleoconch differ in being smooth 
(WARÉN é BOUCHET, 1993). It must be 
mentioned that Xyloskenea naticiformis, 
the species with the greatest similarity, 
was tentatively referred to Tranchia by 
MACLEAN (1992). WARÉN (1996) prefers 
a placement in Xyloskenea based on the 
similarities in protoconch sculpture and 
size. 

Taking all the above, as well as the 
characters of the protoconch and teleo- 
conch, into consideration, we consider 
that Lydipnis translucens must be 
placed in the genus Xyloskenea, thus 
the combination Xyloskenea translucens 
(Dall, 1927). 

The genus Xyloskenea is comprised 
of numerous species in abyssal and 
bathyal depths, world wide, and in 
every case where the substratum is 
known, it is associated with sunken 
wood. Closely related genera are 
Trenchia Knudsen, 1964 (on sunken 
wood, southwestern Pacific) and Ventsia 
Warén € Bouchet, 1993 (hydrothermal 
vents off Fiji) (WARÉN, 1996). 


“Vitrinella” massarita Dall, 1927 


Vitrinella massarita Dall, 1927. Proceedings of the United States National Museum 70(2667): 125. 
[Type locality: Albatross sta. 2668, off Fernandina, Florida [actually off Georgia], 30%58'N, 


79%38'W, 294 fathoms (529 m)!l. 


Type material: In USNM (108137) (three specimens). 


Description: DALL (1927). 
Maximum Reported Size: 1.7 mm. 


CONCLUSIONS AND COMMENTS 


In the present work, which is focused 
on the recent fauna of the family 
Tornidae, subfamilies Circulinae, Teinos- 
tomatinae, Torninae and Vitrinellinae, the 
following 13 genera were studied: 


212 


Distribution: USA: Georgia: DALL (1927) 
Depth: 538 m. 


Circulus (4 species), Teinostoma (27 
species), Tornus (2 species), Anticlimax 
(8 species) Aorotrema (2 species), 
Cyclostremiscus (14 species), Cochli- 
olepis (9 species), Episcynia (1 species), 


RUBIO E7 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Figures 120A-E. Xyloskenea translucens (Dall, 1927). A-D: lectotye, 2.0 mm in diameter, (USNM, 
108434); E-F: spire and protoconch. (SEM's by Yolanda Villacampa, USNM). 

Figuras 120A-E Xyloskenea translucens (Dall, 1927). A-D: lectotipo, 2,0 mm de diámetro, (USNM, 
108434); E-F: espira y protoconcha. (Fotografías al MEB de Yolanda Villacampa, USNM). 


Parviturboides (1 species), Pleuroma- 
laxis (1 species), Solariorbis (16 species), 
Vitrinella (15 species), and Vitrinorbis (1 
species). 

In total, about 2700 specimens and 
shells were examined. This is a large 
number of shells, but if we consider that 
the number of species included in the 


work is about 100, it may seem less 
imposing. Furthermore, if we consider 
the fact that some of them are common, 
and that we have studied large lots such 
as Cochliolepis parasitica (more than 
340 shells), Vitrinella anneliesae (about 
290 shells) or Parviturboides interruptus 
(about 250 shells), it is clear that many 


ES 


Iberus, 29 (2), 2011 


other species were available only in 
small numbers. For example, almost 20 
species were known from only 1 shell, 
and, in a little more than half of the 
total (57 species), each was represented 
by less than 9 shells. 

This scarcity may be due to various 
causes: 1-the small size of most of the 
species; 2- the habitat (unknown for 
most of them) which could be in 
crevices among rocks or symbiotic with 
other organisms; 3- the small number 
of malacologists who give attention to 
micromolluscs; 4- the difficulty of 
finding these minute specimens in old 
collections, many times lost or suffer- 
ing from problems in conservation due 
to fungi, humidity, acidic glass, etc. 

Nonetheless we were able to study 
101. species of Tornidae in the 
Caribbean and neighboring regions. Of 
this total, 86 were previously known, 
and 23 are described as new to science; 
2 more are presented as “sp.” because 
of doubts about their taxonomic valid- 
ity or inadequacy of material. 

A lectotype is here designated for 
each of the following species: Teinos- 
toma reclusum, Teinostoma solidum, 
and Parviturboides interruptus, all in 
the MCZ, Solariorbis petitii in the 
MNHN, Episcynia inornata in the 
NHMUK, and Cochliolepis parasitica 
in the USNM. 

A new name is proposed: Vitrinella 
solaris nom. nov. pro “Cyclostrema” 
thomasi Pilsbry, 1945 non Vitrinella 
thomasi Bartsch, 1918. 

Three neotypes are designated: for 
Teinostoma megastoma and Teinos- 
toma semistriatum in MCZ, and for 
Circulus orbignyi in the MNHN. 

Some types from the Dall and K.J. 
Bush collections, all in USNM, are 
imaged for first time by SEM: Teinos- 
toma minuscula (holotype); Vitrinella 
diaphana (d'Orbigny, 1842); “Vit- 
rinella” carinifex; Pseudorotella flori- 
densis, Vitrinella georgiana, Lydiphnis 
margaritiformis, Vitrinella cerion, Vit- 
rinella rhyssa, Lydiphnis henderson1, 
and Lydiphnis translucens. Also 
Cochliolepis parasitica and Vitrinella 
tincta. 


214 


From the examination of the types 
of DALL (1927), we concluded that “Vit- 
rinella” carinifex, Vitrinella cerion, Vit- 
rinella rhyssa, Vitrinella georgiana, 
Pseudorotella floridensis, Lydiphnis 
margaritiformis and Lydiphnis translu- 
cens are not tornids but skeneids 
included in the genera Cirsonella, 
Micro and Xyloskenea, thus creating the 
new combinations: Cirsonella floriden- 
sis, Cirsonella georgiana, Cirsonella 
margaritiformis, Mikro cerion, Xyloske- 
nea rhyssa, and Xyloskenea translucens. 
We examined types of all these species 
except Vitrinella massarita and could 
not suggest a generic placement only 
for “Vitrinella” carinifex. 

Many types of species belonging to 
different families were also studied in 
order to ascertain that they could not be 
placed in the Tornidae. Sometimes shell 
morphology is sufficient to allow place- 
ment in genus and family. On other 
occasions this determination is not 
easily made. For this reason, we have 
presented an annotated list of species 
which can be included in the Tornidae 
and of others which were excluded 
because they were fossil species or 
probably from other groups. This ancil- 
lary information is presented to provide 
future researchers with all the possible 
information on the species of this group 
and taxa with which they have been 
confused. 

The following species, previously 
considered in synonymy, are treated as 
valid species: 

Teimostoma nessaeum and Teinos- 
toma obtectum are not synonyms of 
Teinostoma biscaynense. 

Cyclostremiscus trilix is not a 
synonym of Cyclostremiscus pen- 
tagonus. 

Cyclostrema thomasi is not a 
synonym of Vitrinella filifera. 

On the other hand, Teinostoma 
clavium is herein considered a 
synonym of Teinostoma semistriatum. 

Two species previously placed in 
the genus Cyclostremiscus: Cyclostrem- 
iscus caraboboensis and C. schrammil 
are placed in the genus Tornus on the 
basis of the similarity of the morpholog- 


RUBIO £7 AZ.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


ical characters of their shells with 
species of this genus in the West coast 
of Africa. 

In relation with the bathymetric 
range, the tornids usually do not live in 
very deep water, most species inhabit- 
ing bottoms between the subtidal level 
and the first 30 meters of the infralitoral 
level. But in relation to the depth data 
of empty shells collected and in most 
bibliographic records, the conclusions 
could be different, appearing in deeper 
water, probably due to the orography of 
the bottom and marine currents. Of the 


species studied, 65 have been found 
between 0 and 60 m, and 18 more have 
been found also in circalitoral and 
bathyal levels, one of them (Teinostoma 
reclusum) having been recorded from 
1170 m. For many species the bathyme- 
try is unknown due to the dearth of 
bottom samples and predominance of 
beach drift in collections. The group of 
species of the genus Cyclostremiscus 
including C. dalli, C. pentagonus, C. 
trilix, and C. hendersoni seem to have a 
deeper bathymetric range, all of them 
reaching between 500 and 800 m. 


APPENDIX 


SPECIES THAT WERE INCLUDED IN TORNIDAE AND NOW BELONG 
TO OTHER FAMILIES 


Family LIOTIDAE Gray, 1850 
Genus Cyclostrema Marryat, 1818 


Cyclostrema cancellatum Marryatt, 1818 
Cyclostrema fulgidum Jeftreys, 1883 
Cyclostrema rugulosum G.O. Sars, 1878 
Cyclostrema valvatoides Jeffreys, 1883 
Cyclostrema pompholyx Dall, 1889 (is now placed in the genus Parviturbo) 
Cyclostrema turbinum Dall, 1889 


Family TURBINIDAE Rafinesque, 1815 
Subfamily SKENEINAE Clark, 1851 
Genus Ganesa Jeffreys, 1883 


Ganesa proxima Tryon, 1888 
Ganesa bushae Dall, 1927 
Ganesa conica Dall, 1927 

Ganesa depressa Dall, 1927 
Ganesa valvata Dall, 1927 

Ganesa diaphana A.E. Verrill, 1884 
Ganesa striata Bush, 1897 
Ganesa convexa Bush, 1897 
Ganesa verrilli Tryon, 1888 
Ganesa ornatam A.E. Verrill, 1884 
Ganesa dalli A.E. Verrill, 1882 
Ganesa abyssicola Bush, 1897 


2D 


Iberus, 29 (2), 2011 


Figures 121A-E. Dillwynella modesta Dall, 1889. A-E: syntypes, 1.4, 1.6 mm (MCZ 007657). 
Figuras 121A-E. Dillwynella modesta Dall, 1889. A-E: sintipos, 1,4, 1,6 mm (MCZ 007657). 


Genus Granigyra Dall, 1889 


Granigyra limata Dall, 1889 
Granigyra radiata Dall, 1927 
Granigyra spinulosa Bush, 1897 


Genus Dillwynella Dall, 1889 
Dillwynella modesta (Dall, 1889) (Figures 121A-E) 


Dillwynella modesta Dall, 1889. Bull. MCZ, 18: 362-363, pl. 21, figs. 3-3a. [Type locality: Blake sta. 
215, off St. Lucia, 226 fathoms]. 


Type material: Three syntypes, MCZ (007657). 


Genus Molleriopsis Bush, 1897 


Molleriopsis abyssicola Bush, 1897 
Molleriopsis sincera Dall, 1889 


Genus Leptogyra Bush, 1897 


Leptogyra verrilli Bush, 1897 
Leptogyra inconspicua Bush, 1897 
Leptogyra eritmeta Bush, 1897 


PS 


RUBIO ET 42Z.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Family LEPETELLIDAE Dall, 1881 


Genus Choristella Bush, 1897 


Choristella leptalea Bush, 1897 
Choristella brychia Bush, 1897 


Family ODOSTOMIIDAE Pelseneer, 1928 
Genus Cyclostremella Bush, 1897 


Cyclostremella humilis Bush, 1897 


ACKNOWLEDGEMENTS 


The authors offer thanks to Dr. Jerry 
Harasewych and Yolanda Villacampa 
of the Smithsonian Institution (USNM) 
for their help in the study of Cochli- 
olepis parasitica and many other types 
deposited in this institution. To Adam 
J. Baldinger, for the loan of material of 
Tornidae, including some types from 
MCZ, and his cooperation with the use 
of this material. To Marlo Krisberg of 
Merrit Islands for the loan of material 
from his collection. To Colin Redfern of 
Boca Raton for the same. To Flaviano 
for sending material from Itaparica, 
Brazil. To Jacques Pelorce, from Paris, 
for sending material collected on his 
trips to the Caribbean. Also to Dr. 
Harry G. Lee, of Jacksonville, Florida, 
who put his collection at our disposal, 
authorizing the photography of all the 
necessary material, and donating 
several types of the new species unique 
to his collection. Also for his revision 
during the editing period of this work. 

The type material of Cyclostremis- 
cus  multiliratus was obtained 
through support by the National 


BIBLIOGRAPHY 


ABBOTT R.T. 1954. American seashells. D. Van 
Nostrand Company, Inc., Princeton, NJ, 
xiv + 541 pp, 40 pls. 

ABBOTT R.T. 1974. American seashells: the 
marine Mollusca of the Atlantic and Pa- 
cific coasts of North America, 2nd ed. Van 
Nostrand Reinhold Company. New York, 
663 pp, 24 pls. 


Science Foundation under Grant No. 
0515995: 

To Jesus Méndez and Inés Pazos of 
the Centro de Apoyo Científico y Tec- 
nológico a la Investigación (CACTI) 
of the University of Vigo for most of 
the SEM images; also to Ramiro Bar- 
reiro Pérez of the Centro de Apoyo 
Científico y Tecnológico of the Uni- 
versity of Santiago de Compostela 
(CACTUS) for some micrographs. 

To the Museo de Historia Natural 
“Luis Iglesias”, of the University of 
Santiago de Compostela, its director 
Dr. María Isabel Fraga Vila, for coop- 
eration during the entire study sup- 
porting all the necessary material, 
microphotography, correspondence, 
etc. 

To Ricardo Absaláo and Petrobras 
S.A. (Brazilian Oil company) which 
authorized the use of photographs 
from Campos Basin, Rio de Janeiro. 

To Anders Warén who authorized 
the use of Neusas photographs. 

Javier Conde made the corrections 
to the English language. 


ABBOTI R.I. € DANCE-S. P. 1982. Com.- 
pendium of seashells: a color guide to more 
than 4,200 of the world's marine shells. E. 
P Dutton, New York, x -I- 411 [+ 1] pp. 

ABSALAO R.S. 2009. New small deep-sea 
species of Gastropoda from the Campos 
Basin off Brazil. American Malacological 
Bulletin, 27: 133-140. 


ZA 


Iberus, 29 (2), 2011 


ABSALAO R.S. € PIMENTA A.D., 2005. Moluscos 
marinhos da Apa do Archipelago de San- 
tana, Macaré, RJ. Ciencia Moderna, Rio de Ja- 
neiro. 84 pp. 

ADAM W. 4: KNUDSEN J. 1969. Quelques genres 
de Mollusques prosobranches marins in- 
connus ou peu connus de l'Afrique occi- 
dentale. Bulletin Institut Royal des Sciences 
Naturelles de Belgique, 44(27): 1-69. 

ADAMS A. 1850. Monographs of Cyclostrema, 
Marryat, and Separatista, Gray, two genera 
of gasteropodous mollusks. Proceedings of 
the Zoological Society of London 1850: 41-45. 

ADAMS C.B. 1850. Monograph of Vitrinella, a 
New Genus of New Species of Turbinidae 10 
pp. Author, Amherst, Massachusetts. 

ADAMS A. 1866. Monographs of the genera Cy- 
clostrema, Adeorbis, and Teinostoma. The- 
saurus Conchyliorum.Sowerby, G.B. (ed.) 3: 
249-262. 

AGUAYO C.G. 1949. Tres nuevos moluscos ma- 
rinos de las costas de Cuba. Revista de la So- 
ciedad Malacológica “Carlos de la Torre” 6: 
93-96, pl. 4. 

AGUAYO C. 8 BORRO P. 1946a. Nuevos molus- 
cos del Terciario Superior de Cuba. Revista 
de la Sociedad Malacologica “Carlos de la To- 
rre” 4(1): 9-12, pl. 1. 

AGUAYO C. BORRO P. 1946b. Algunos mo- 
luscos Terciarios de Cuba. Revista de la So- 
ciedad Malacologica “Carlos de la Torre” 
4(2): 43-49, pl. 3. 

AGUAYO C. €: JAUME M.L. 1936. Sobre algunos 
moluscos marinos de Cuba. Memorias de la 
Sociedad Cubana de Historia Natural “Felipe 
Poey” 10(2): 115-122. 

ALTENA C.O. van R. 1966. Vitrinellidae (ma- 
rine Mollusca Gastropoda) from holocene 
deposits in Surinam (Dutch Guiana). Zoolo- 
gische Mededelingen, 41: 233-241. 

ALTENA C.O. van R. 1975. The marine mollusca 
of Suriname (Dutch Guiana) Holocene and 
Recent. Part Il. Gastropoda and 
Cephalopoda. Zoologische Verhandelingen, 
139: 1-104, pls. 12. 

ANDREWS J. 1977. Shell and Shore of Texas. 
University of Texas Press. Austin, United 
States. Journal of Molluscan Studies, 47: 248- 
Z 


ANDREWS J. 1994. A field guide to shells of the 
Florida coast. Gulf Publishing, Houston. 182 


pp- 

ARANGO R. 1880. Contribucion a la Fauna Ma- 
lacologica Cubana. 137-280, 1-35. G. Mon- 
tiel: Habana. 15 Jul. 

BIELER R. 6: MIKKELSEN P.M. 1988. Anatomy 
and reproductive biology of two western 
Atlantic species of Vitrinellidae, with a case 
of protandrous hermaphroditism in the 
Rissoacea. The Nautilus, 102(1): 1-29. 


218 


BOUCHET P. éz ROCROIJ.P. (Ed.); ErydaJ., Haus- 
dorf B., Ponder W., Valdés Á. € Warén A. 
2005. Classification and nomenclator of gas- 
tropod families. Malacolo gía, 47(1-2): 397 pp. 
ConchBooks, Hackenheim, Germany. 

BoYKoO C.B. ££ CORDEIRO J.R. 2001. Catalog of Re- 
cent type specimens in the division of inver- 
tebrate zoology , American Museum of Na- 
tural History . V . Mollusca, part 2 (class Gas- 
tropoda [exclusive of Opisthobranchia and 
Pulmonata ], with supplements to Gastro- 
poda [ Opisthobranchia ], and Bivalvia. Bu- 
lletin of the American Museum of Natural 
History, 262: 3-158. 

Bush K.J. 1885. List of the shallow-water Mol- 
lusca dredged off Cape Hatteras by the “Al- 
batross” in 1883. Annual Reportof the United 
States Commissioner of Fish and Fisheries, 
1883: 579-595. 

Bush K.J. 1897. Revision of the marine gas- 
tropods referred to Cyclostrema, Adeorbis, 
Vitrinella, and related genera; with descrip- 
tions of some new genera and species be- 
longing to the Atlantic fauna of Amer- 
ica. Transactions of the Connecticut Acad- 
emy of Arts and Sciences, 10: 97-144, pls. 
22-23. 

CAMPBELL L.D. 1993. Pliocene molluscs from 
the Yorktown and Chowan river formations 
in Virginia. Virginia Division of Mineral Re- 
sources, Publication 127, 1-259, 

CLARKE H.C. 1962. On the composition, zoo- 
geography, origin and age of the deep-sea 
mollusk fauna. Deep Sea Research and 
Oceanographic Abstracts, 9(7-10): 291-306. 

CLENCH W.J. 6: TURNER R.D., 1950. The western 
Atlantic marine mollusks described by C.B. 
Adams. Occasional Papers on Mollusks, 1(15): 
233-403 incl. pls. 29-49. 

COOLEY N.P. 1978. An inventory of the estuar- 
ine fauna in the vecinity of Pensacola, Florida. 
Florida Marine Research Publications, 33: 
1i+119. 

CORGAN J.X. 1968. New name for Vitrinella 
tenuisculpta Aguayo y Borro (Gastropoda). 
Nomenclatural Notes. Journal of Paleontol- 
ogy, 43: 1298. 

COSEL R. VON 1986. Moluscos de la región de la 
Ciénaga Grande de Santa Marta (costa del Ca- 
ribe colombiano) Anales del Instituto de In- 
vestigaciones Marinas de Punta Betín, 15-16: 
79-370 

DaLL W.H. 1889a. Reports on the results of 
dredgings, under the supervision of Alexan- 
der Agassiz, in the Gulf of Mexico (1877-78) 
and in the Caribbean Sea (1879-80), by the U. 
S. Coast Survey Steamer “Blake”. Bulletin of 
the Museum of Comparative Zoology, 18: 1- 
492, pls. 10-40. 


RUBIO £7 AL.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


DaALL W.H. 1889b. A preliminary catalogue of 
the shell-bearing marine mollusks and bra- 
chiopods of the south-eastern coast of the 
United States, with illustrations of many of 
the species. Bulletin of the United States Na- 
tional Museum, 37: 221 p., pps. 1-74. 

DaLL W.H. 1892. Contributions to the Tertiary 
fauna of Florida, with especial reference to the 
Miocene silex-beds of Tampa and the Pliocene 
beds of the Caloosahatchie River. Part II. 
Streptodont and other gastropods, con- 
cluded. Transactions of the Wagner Free In- 
stitute of Science of Philadelphia, 3 [i-viil: 
201-473, 1 fold-out map, pls. 13-22. 

DaLL W.H. 1889. Report on the results of dred- 
ning by the U. S. Coast survey steamer 
“Blake”. XXIX. Report on the Mollusca, part 
II. Gastropoda and Scaphopoda. Bulletin of 
the Museum of Comparative Zoology, 18: 1- 
492, pls. 10-40. 

DaLL W.H. 1903. Contributions to the Tertiary 
fauna of Florida with especial reference to 
the Silex Beds of Tampa and the Pliocene 
Beds of the Caloosahatchie River including in 
many cases a complete revision of the generic 
groups treated of and their American Ter- 
tiary species. Part VI. Concluding the work. 
Transactions of the Wagner Free Institute of 
Science of Philadelphia, 3(4): i-xiv, 1219- 1654, 
pls. 48-60. 

DaLL W.H. 1927. Small shells from dredgings 
off the southeast coast of the United States by 
the United States Fisheries steamer Albatross 
in 1885 and 1886. Proceedings of the United 
States National Museum, 70(2667): 1-134. 

Díaz MERLANO J.M. € PUYANA HEGEDUS M. 
1994. Moluscos del Caribe colombiano. Un ca- 
tálogo ilustrado. Colciencias. Fundación Na- 
tura. Invermar. Santa Fé de Bogotá, 291 pp, 
74 pls. 

EMERSON W.K. éz JACOBSON M.R. 1976. The 
American Museum of Natural History, Guide 
to shells. Alfred A. Knopf, New York. 482 
pp, 47 pls. 

ESPINOSA J., FERNÁNDEZ-GARCÉS R. 6 ROLÁN E. 
1985. Catálogo actualizado de los moluscos 
marinos de Cuba. Reseñas Malacológicas, 9: 
1-90. 

ESPINOSA J. 6 ORTEA J. 2001. Moluscos del mar 
Caribe de Costa Rica: Desde Cahuita hasta 
Gandoca. Avicennia (Supl. 4): 1-77. 

FABER MJ. 1995. New data on mollusk species 
described by Gordon W. Nowell-Usticke. De 
Kreukel 31: 59-69. 

FABER M.J. 2007. Marine gastropods from the 
ABC Islands and other localities 20. Sola- 
riorbis semipunctus Moore, 1965 (Gastro- 
poda: Vitrinellidae), first records for Florida, 
USA and Curacao. Miscellanea Malacologica, 
2(4): 84. Figs. 1-3. 


FISCHER P. 1857. Études sur un groupe de co- 
quilles de la famille des Trochidae. Journal de 
Conchyliolo g1e, 6: 42-53, 168-176, 284-288, pl. 
10. 

FIscHER P. 1880-1887. Manuel de conchyliolo- 
gie et de paléontologie conchyliologique ou 
histoire naturelle des mollusques vivants et 
fossiles suivi d'un appendice sur les Bra- 
chiopodes. Savy, Paris, xxiv + 1-1569 + 23 
pls., facing pl. explanations. [1-112 (1880); 
113-304 (1881); 305-416 (1882); 417-608 (1883); 
609-688 (1884); 689-896 (1885); 897-1008 (1886); 
1009-1369 (1887)]. 

FOLIN L. DE 1870. Vera-Cruz et Carmen. Les 
Fonds de la Mer 1: 181-191, pls. 23, 25-56 
Savy: Paris. 

FRETTER V. 1956. The anatomy of the proso- 
branch Circulus striatus (Philippi) and a re- 
view of its systematic position. Proceedings 
ofthe Zoological Society of London, 126: 369- 
381. 

FRETTER V. 1984. Prosobranchs. In: Wilbur et 
al. (eds.). The Mollusca, Vol. 7, Reproduc- 
tion. Academic Press, New York and Lon- 
don, p. 1-45. 

FRETTER V. € GRAHAM A. 1962. British proso- 
branch molluscs: their functional anatomy 
and ecology. Ray Society, London, xvi -1- 755 


pie: 

FRETTER V. é: GRAHAM A. 1964. Reproduction. 
In: Wilbur, K. M. and C. M. Yonge (eds). 
Physiology of Mollusca, Vol. 1. Academic 
Press, New York and London, p. 127-164. 

FRETTER V. 82 GRAHAM A. 1978. The prosobranch 
molluscs of Britain and Denmark. Part 4 — 
Marine Rissoacea. Journal of Molluscan Stud - 
1es, Supplement 6: 153-241. 

GABB W.M. 1873. On the Santo Domingo 
Miocene and its Fossils. Proceedings Amer- 
ican Philosophy Society, 12: 571-573. 

GABB W.M. 1881. Descriptions of new species 
of fossils from Pliocene Clay Beds between 
Limon and Moen, Costa Rica, together with 
notes on previously known species from there 
and elsewhere in the Caribbean area. Journal 
of the Academy of Natural Sciences of 
Philadelphia, 8: 349-380, 4 pls. 

GARCIA E.F. 2002. More discoveries from a col- 
lecting expedition off the Louisiana coast. 
American Concholo gist 30(1): 6-7, 10. 

GARCÍA-CUBAS A. 1970. Ecologia y distribución 
de los micromoluscos recientes de la Laguna 
Madre, Tamaulipas, Mexico. Universidad 
Autonoma de Mexico, Instituto de Geología, 
Boletin 86: 1+1v+1-44, 2 foldouts tables, 8 pls. 

GARCÍA-CUBAS A. 1971. Estudios oceanográfi- 
cos y biológicos de reconocimiento en la Ba- 
hía de Guaymas. Instituto de Biología, Uni- 
versidad Nacional Autónoma de México. 87 
p., tablas, figs. y mapas. 


219 


Iberus, 29 (2), 2011 


GARCÍA-CUBAS A. € REGUERO M., 1990. Mo- 
luscos del sistema lagunar Tupilco-Ostión, 
Tabasco, México: Sistemática y Ecología. 
Anales del Instituto de Ciencias del Mar y 
Limnología, Universidad Nacional Autó- 
noma de México, 17(2): 309-343. 

GARCÍA-CUBAS A., ESCOBAR-DE LA LLATA F., 
GONZÁLEZ L. V. 6 REGUERO M. 1990. Mo- 
luscos de la laguna Mecoacán, Tabasco, Mé- 
xico: Sistemática y Ecología. Anales del Ins- 
tituto de Ciencias del Mar y Limnología, 
Universidad Nacional Autónoma de Mé- 
xico IAN SO: 

GARDNER J. 1948. Mollusca from the Miocene 
and lower Pliocene of Virginia and North 
Carolina. Part 2. Scaphopoda and Gas- 
tropoda. U.S. Geological Survey Professional 
Paper, 199-B: 1-111, 179-310, pls. 24-38. 

GOFAS S., PINTO AFONSO J. £ BRANDAO M. 
1985. Conchas e moluscos de Angola. Uni- 
versidad de Agostinho Neto / Elf Aquitaine. 
Angola. 139 pp. 

GRAHAM A. 1982. Tornus subcarinatus (Proso- 
branchia, Rissoacea), anatomy and rela- 
tionships. Journal of Molluscan Studies, 48: 
144-147. 

GurrY R.J.L. 1866. On the Tertiary Mollusca of 
Jamaica. Quarterly Journal of the Geological 
Society of London, 22(1): 281-295, pls. 16-18. 

HICKMAN C.S. £ MCLEAN J.H. 1990. System- 
atic revision and suprageneric classification 
of trochacean gastropods. Natural History 
Museum of Los Angeles County, Science 
Senes od 11Óo. 

HOLMES F.S., 1858-1860 [published in parts]. 
Post-Pleiocene Fossils of South Carolina. 
Russell and Jones, Charleston, 1-98 p, 14 pls. 
[R. E. Petit offers a collation of this rare se- 
rial work: Title page (dated 1860); verso: 
name of printer and engraver. Dedication 
page; verso: Preface and acknowledgements. 
[1] - xii - Introduction, [i]-v - Index; v verso: 
Errata, 1-64, pls. 1-10 - Cover for Nos. 1 é€z 2 
with printed date of 1858. “3, 4 + 5” plus “2 
plates to a number” added in pen; 65-98, 
pls. 11-14 - Cover for “Nos. 6 and 7” with 
printed date of 1859.]. 

HOUBRICK J.R. 1967. Notes on Cyclostremis- 
cus schrammii. The Nautilus, 80(4): 131-133. 

HOUBRICK J.R. 1968. A survey of the litoral ma- 
rine molluscs of the Caribbean coast of Costa 
Rica. Veliger, 11: 4-23. 

Jacor A.P. 1921. Some marine shells of Beau- 
fort and vicinity. Journal of the Elisha 
Mitchell Scientific Society, 36: 129-145, pls. 
11-13. 

JAUME M.L. € BORRO P. 1946. Novedades en 
moluscos marinos Cubanos. Revista de la So- 
ciedad Malacologica “Carlos de la Torre”, 
4(1): 13-22, pl. 2. 


220 


JEFFREYSJ.G. 1865. British conchology, or an ac- 
countof the Mollusca which now inhabit the 
British Isles and the surrounding seas. Vol. 
II. Marine shells, comprising the remaining 
Conchifera, the Solenoconchia, and Gas- 
teropoda as far as Littorina.]. van Voorst, 
London, 393 (+ 1) p., 8 pls. 

JerFREYS J.G. 1883 On the Mollusca procured 
during the “Lightning” and “Porcupine” ex- 
peditions 1868-70. Part VI. Proceedings of 
the Zoological Society of London, 1882: 88- 
149. 

JONG K.M. DE é£ COOMANS H.E. 1988. Marine 
gastropods from Curacao, Aruba and 
Bonaire. Studies on the Fauna of Curagao 
and other Caribbean Islands, 69. E.J. Brill. 
261 pp, 47 pls. 

JOHNSON C.W. 1934. List of marine Mollusca of 
the Atlantic coast from Labrador to Texas. 
Proceedings of the Boston Society of Natural 
History, 40(1): 1-204. 

JOHNSON R.I. 1989. Molluscan taxa of Addison 
Emery Verrill and Katharine Jeannette Bush, 
including those introduced by Sanderson 
Smith and Alpheus Hyatt Verrill. MCZ Pub- 
lications on Mollusks, Occasional Papers on 
Mollusks, 5: 1-143. 

KEEN A.M. 1960. In R. C. Moore (ed.), Treatise 
on Invertebrate Paleontology, Part 1, Mol- 
lusca 1. Mollusca - general features, 
Scaphopoda, Amphineura, Monoplacophora, 
Gastropoda - general features, Archaegas- 
tropoda, mainly Paleozoic Caenogastropoda 
and Opisthobranchia. Geological Society of 
America éz Univ. Kansas Press, Boulder, CO 
and Lawrence KS, xxiii + pp. 1-351. 

KURTZ J.D. 1860. Catalogue of Recent marine 
shells, found on the coasts of North and South 
Carolina. Portland, David Tucker.ME 9 pp. 

LANGE-DE-MORRETES F. 1949. Ensaio de catálogo 
dos moluscos do Brasil. Archos Mus. Paraná, 
7(1): 3-216. 

LEE H. G. 2009. Marine shells of northeast 
Florida. Jacksonville Shell Club, Jacksonville, 
204 pp., 19 pls. 

Lyons W.G. 1989. Nearshore marine ecology at 
Hutchinson Island, Florida: 1971-1974. XI. 
Mollusks. Florida Marine Research Publica- 
tions, 47: i+iv, 1-131. 

MARSHALL B.A. 1988. Skeneidae, Vitrinellidae 
and Orbitestellidae (Mollusca: Gastropoda) 
associated with biogenic substrata from ba- 
thyal depths off New Zealand and New South 
Wales. Journal of Natural History, 22: 949- 
1004. 

MAURY CJ. 1922. Recent Mollusca of the Gulf 
of Mexico and Pleistocene and Pliocene 
species from Gult states. Part 2: Scaphopoda, 
Gastropoda, Amphineura, Cephalopoda. Bul- 
letins of American Paleontolo gy, 9(38): 33-172. 


RUBIO £7 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


MELLO R.L.S. € PERRIER, L.L. 1986. Polypla- 
cophora e Gastropoda do litoral sul de Per- 
nambuco, Brasil. Cadernos Omega da Uni- 
versidade Federal Rural de Pernambuco 2: 
107-144. 

MIKKELSEN P.M., MIKKELSEN P.S. 6: KARLEN 
D.J. 1995. Molluscan biodiversity in the In- 
dian River Lagoon, Florida. Bull. Mar. Sci., 
57: 94-127. 

MOORE D.R. 1964. The family Vitritiellidae in 
South Florida and the Gulfof Mexico. Ph. D. 
dissertation. University of Miami, Coral 
Gables, FL, xi + 235 (+ 2) pl. 

MOORE D.R. 1965. New species of Vitrinellidae 
from Gulf of Mexico and adjacent waters. 
The Nautilus, 78(3): 73-79, pls. 7-8. 

MOORE D.R. 1969. Cyclostrema miranda 
Bartsch, a synonym of Tornus subcarinatus 
Montagu. The Veliger, 12(2): 169-170. 

MOORE D.R. 1972. Cochliolepis parasitica, a 
non parasitic marine gastropod, and its place 
in the Vitrinellidae. Bulletin of Marine Sci- 
ence, 22(1): 100-112. 

MORRIS P.A. 1973. A field guide to shells of the 
Atlantic and Gulf coasts and the West In- 
dies. Hougton Mifflin Co., Boston. 330 pp, 76 
pls. 

NORDSIECK, F. 1968. Die europdischen Meeres- 
Geháuseschneken. G. Fischer, Sttutgart. 273 
pp: 

NOWwELL-UsTICKE G.W. 1959. A Check Listof the 
Marine Shells of St. Crotx, U.S Virgin Is- 
lands. Lane Press, Burlington. 90 pp., 4 pls. 

NOWELL-USTICKE G.W. 1971. A Supplementary 
Listing of New Shells (Illustrated) Revised 
Edition. to be Added to the Check Listof the 
Marine Shells of St. Croix. Author: San Croix, 
Virgin Islands, 32 pp., 6 pls. 

ODÉ H. 1969. Cochliolepis parasitica Stimpson. 
Texas Conchologist 5(7): 73-74. 

ODÉ H. 1987a. Distribution and records of the 
marine Mollusca in the northwest Gulf of 
Mexico, families Skeneidae, Vitrinellidae. 
Texas Conchologist, 23(4): 110-121. 

ODÉ H. 1987b. Distribution and records of the 
marine Mollusca in the northwest Gulf of 
Mexico, family Vitrinellidae. Texas Con- 
chologist, 24(1): 21-40. 

ODÉ H. 1988. Distribution and records of the ma- 
rine Mollusca in the northwest Gulf of Mex- 
ico, family Vitrinellidae. Texas Concholo- 
gist, 24(2): 59-72. 

OLIVER J.D. € ROLÁN E. 2011. The family 
Tornidae (Gastropoda, Rissooidea) in the 
East Atlantic, 2. Circulinae. Iberus, 29(1): 9- 
99: 

OLSSON A.A. é££ HARBISON A. 1953. Pliocene 
Mollusca of southern Florida. Monographs 
of the Academy of Natural Sciences of 
Philadelphia, 18: 1-457 + 65 pls. 


OLssoN A.A. 8 MCGINTY T. L. 1958. Recent 
marine mollusks from the Caribbean coast of 
Panama with the description of some new 
genera and species. Bulletins of American 
Paleontology, 39: 1-58, pls. 1-5. 

ORBIGNY A. D' 1842. Mollusques Histoire Phy- 
sique, Politique et Naturelle de lle de Cuba, 
2. Arthus Bertrand, Paris. pp 1-112, pls. 1-7. 

ORTIZ-CORPS E. 1985. An annotated checklist of 
the Recent marine Gastropoda (Mollusca) 
from Puerto Rico. Memorias del Quinto Sim- 
posto de la Fauna de Puerto Rico y el Caribe 
1220 pp: 

PARKER R.H. 1959. Macro-invertebrate assem- 
blages of central Texas coastal bays and La- 
guna Madre. AAPG Bulletin, 43: 2100-2166. 

PARKER R.H. € CURRAY J.R. 1956. Fauna and ba- 
thymetry of banks on continental shelf, north- 
west Gulf of Mexico. Bulletin of the Ameri- 
can Association of Petroleum Geologists, 
40(10): 2428-2439. 

PFEIFFER L. 1839. Bericht ber die Ergebnisse 
meiner Reise nach Cuba im Winter 1838- 
1839. Archiv fiir Naturgeschichte 5(1): 346- 
358. 

PFEIFFER L. 1840. Uebersicht der im Januar, Fe- 
bruar und Márz 1839 auf Cuba gesammelten 
Mollusken. Archiv fir Naturgeschichte, 6(1): 
250-261. 

PFEIFFER L. 1850. Die Schnirkelschnecken nebst 
den zunáchst verwandten Gattungen. Sy- 
stematische Conchylien-Cabinet 1: 1-32, pls. 
12*,:85, 87, 89, 93, 96, 102-106, 110-112,114. 

PFEIFFER L. 1854. Zur Molluskenfauna der In- 
sel Cuba. Malakozoologische Blátter 1: 170- 
215: 

PIÉSBRY HA. 1939. .A new Floridan Cy- 
clostrema. The Nautilus, 53: 53, pl. 8. 

PiLsBRY H.A. 1946. The type specimens of C.B. 
Adams's Jamaican species of Vit- 
rinella. MNotulae Naturae of the Academy 
of Natural Sciences of Philadelphia, 1162: 1- 
5, figs. 1-6. 

PiLsBRY H.A. 1953. Pliocene Mollusca of South- 
ern Florida. Part II-A. The Vitrinelli- 
dae. Monographs of the Academy of Natural 
Sciences of Philadelphia, 18: 411-438, pls. 49- 
56. 

PirsBRY H.A. £ AGUAYO C.G. 1933. Marine 
and freshwater mollusks new to the fauna 
of Cuba. The Nautilus, 46: 116-123, pl. 6. 

RIESBRY TL:AS Se MEGINTY EL. 19454. “Cy= 
clostrematidae” and Vitrinellidae of Florida. 
Part 1. The Nautilus, 59(1): 1-13, pls. 1, 2. 

PILSBRY H.A. €: MCGINTY T.L. 1945b. “Cy- 
clostrematidae” and Vitrinellidae of Florida. 
Part 2. The Nautilus, 59(2): 52-59, pl. 6. 

PiLSBRY FLA. € MCcGINTY T.L. 1946a. “Cy- 
clostrematidae” and Vitrinellidae of Florida. 
Part 3. The Nautilus, 59(3): 77-83, pl. 8. 


ZLl 


Iberus, 29 (2), 2011 


PiLsBRY H.A. €: MCGINTY T.L. 1946b. Vitrinelli- 
dae of Florida, Part 4. The Nautilus, 60(1): 12- 
15 7pli2: 

PiLsBRY H.A. 6: MCGINTY T.L. 1950. Vitrinellidae 
of Florida: Part 5. The Nautilus, 63(3): 85-87, 
plo! 

PiLsBRY H.A. 6: OLSSON A.A. 1945. Vitrinellidae 
and similar gastropods of the Panamic 
Province. Part I. Proceedings of the Academy 
of Natural Sciences of Philadelphia, 97:249-278, 
pls. 22-30. 

PiLsBRY H.A. 8: OLSSON A.A. 1952. Vitrinellidae 
of the Panamic Province: II. Proceedings of 
the Academy of Natural Sciences of Philadel- 
phia, 104: 35-88, pls. 2-13. 

PONDER W.F. 1988. The Truncatelloidean (=Ris- 
soacean) radiation- A preliminary phylogeny: 
Prosobranch phylogeny. Malacological Re- 
view, Supplement, 4: 129-164. 

PONDER W.F. 1994. The Malacofauna of Hong, 
Kong and Southern China II (ed. B. Morton). 
Proceedings of the Third International Work- 
shop on the Malacofauna of Hong Kong and 
Southern China, Hong Kong, 13 April-1 May 
1992. Hong Kong University Press, 1994. 

PONDER W.F. 82 W.ARÉN A. 1988. Classification of 
the Caenogastropoda and Heterostropha —a 
list of the family-group names and higher taxa. 
Malacological Review, Supplement 4: 288-317. 

PORTER H.J. 1974. The North Carolina marine 
and estuarine mollusca: An atlas of Occur- 
rence. University North Carolina Institute of 
Marine Science, Morehead City, North Car- 
olina, 351 pp. 

Prinez D. 1973. Los moluscos gasterópodos y pe- 
lecípodos del estado de Nueva Esparta, Ve- 
nezuela. Memoria, Sociedad de Ciencias Na- 
turales La Salle, 33: 169-222. 

Princz D. 1977. Notas sobre algunos micromo- 
luscos de la plataforma de Guyana. Memoria, 
Sociedad de Ciencias Naturales La Salle, 
36(108): 283-293. 

PrINcz D. 1978. Los moluscos marinos del Golfo 
de Venezuela. Memoria, Sociedad de Cien- 
cias Naturales La Salle, 109: 51-76. 

PrINCZ D. 1982. Lista y bibliografia de los gaste- 
rópodos marinos vivos de los mares de Ve- 
nezuela, Trinidad e Islas de Sotavento. Bole- 
tín de la Sociedad Venezolana de Ciencias Na- 
turales, 37: 103-147. 

Princz D. 1986. Bionomie des micromollusques 
benthiques du nord -est du Venezuela. Tesis de 
Doctorado, Université de Bretagne Occiden- 
tale, Brest, 176 pp. 

RADWIN G.E. 1969. A recent molluscan fauna 
from the Caribbean coast of southeastern 
Panama with special reference to those from 
north Saint Petersburg. Monographs of the 
Academy of Natural Sciences of Philadelphia, 
8: 411-438, pls. 49-56. 


ZZZ 


REDFERN C. 2001. Bahamian seashells. A thou- 
sand Species from Abaco, Bahamas. Ba- 
hamiansehlls.com Inc. Boca Raton, 280 pp, 
124 pls. 

REED J.K. 82 MIKKELSEN P.M. 1987. The mol- 
luscan community associated with the scle- 
ractinian coral Oculina varicosa. Bulletin of 
Marine Science, 40: 99-131. 

REGUERO M., GARCÍA-CUBAS A. 6z ZÚNIGA G., 
1991. Moluscos de la laguna Tampamachoco, 
Veracruz, México: Sistemática y Ecología. 
Anales del Instituto de Ciencias del Mar y 
Limnología, Universidad Nacional Autó- 
noma de México, 18(2): 289-328. 

RICE W.H. €: KORNICKER L.S. 1965. Mollusks 
from the deeper waters of the northwestern 
Campeche Bank, Mexico. Publications of the 
Institute of Marine Science, 10: 108-172. 

RICHARDSON C.L. 1986. Polygyracea: Catalogue 
of species (Parts 1, Polygyridae; 2, Corilli- 
dae; 3 Sagdidae). Tryonia, 13: 1-139, 1-4, 1- 
38. 

Rios E.C. 1975. Brazilian marine mollusks ico- 
nography. Museu Oceanografico, Fundacáo 
Universidade do Rio Grande, Rio Grande, 331 
pp, 91 pls. 

RIOS E.C. 1994. Seashells of Brazil. Fundacao Ci- 
dade do Rio Grande, Rio Grande, 345 pp, 
113 pls. 

Rios E.C. 2009. Compedium of Brazilian Sea 
Shell. Universidade Federal Rio Grande, 668 


pp- 

ROBINSON D.G. 1991. The systematics and pa- 
leoecology of the prosobranch gastropods 
of the Pleistocene Moín formation of Costa 
Rica. Doctoral dissertacion, Tulane Univer- 
sity, New Orleans, Louisiana. 748 pp. 

ROBINSON D.G. € MONTOYA M. 1987. Los mo- 
luscos marinos de la costa Atlántica de Costa 
Rica. Revista Biología Tropical, 35: 375-400. 

ROLÁN E., FERNÁNDEZ-GARCÉS R. € RUBIO 
F. 1997. Una nueva especie de Anticlimax 
(Gastropoda: Vitrinellidae) de Cuba. Iberus, 
15: 31-34. 

ROLÁN E. 4 RUBIO F. 2002. The family Tornidae 
(Gastropoda, Rissooidea) in the East Atlantic. 
Supl. Reseñas Malacologicas, SEM: 1-98. 

RUBIO F. €££ RODRÍGUEZ BABÍO C. 1991. Sobre la 
posición sistemática de Pseudorbis granu- 
lum Brugnone, 1873 (Mollusca, Archaeo- 
gastropoda, Skeneidae) y descripción de 
Pseudorbis jameoensis n. sp. procedente de 
las Islas Canarias. Iberus, 9(1-2): 203-207. 

RUBIO F. € ROLÁN E. 1998. Una nueva especie 
de Tomura (Gastropoda, Heterobranchia, 
Cornirostridae) del Caribe. Iberus, 16(1): 119- 
128% 

RUBIO F. € ROLÁN R. 2009. Sobre la posición sis- 
temática de Teinostoma azorica Dautzen- 
berg éz Fischer, 1896. Noticiario SEM, 52: 55- 
56. 


RUBIO £7 ALz.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


SAGRA R. DE LA. 1831. Historia Económico-Po- 
lítica y Estad ística de la Isla de Cuba. Arazoza 
y Soler. 12 Tomos. Segunda parte, Historia 
Natural. Atlas de Zoología, Tomo VIII. Pa- 
ris. Librería de Arthus Bertrand. 

SARASÚA H. 1970. Prosobranquios marinos nue- 
vos para la fauna de Cuba (Mollusca: Pro- 
sobranchia). Poeyana, 72: 1-20. 

SCHWENGEL J.S. £ MCGINTY T.L. 1942. Some 
new and interesting marine shells from north- 
west Florida. The Nautilus, 56: 13-18, pl. 3. 

SMITH E.A,, 1871. A list of species of shells from 
West Africa, with description of those hith- 
erto undescribed. Proceedings of the Zoo- 
logical Society of London, 1871: 727-739. 

SMITH E.A. 1890. Report on the marine mol- 
luscan fauna of the island of St. He- 
lena. Proceedings of the Zoological Society 
of London 1890: 293, pl. 24, fig. 5. Type lo- 
cality: St. Helena] 

SMITH M. 1937. Further notes upon Tertiary 
and Recent mollusks from Florida together 
with descriptions of new species. The Nau- 
tilus, 51(2): 65-66, pi. 6. 

STIMPSON W. 1858. On Cochliolepis parasiti- 
cus. Proceedings of the Boston Society of 
Natural History, 6: 307-309. 

STIMPSON C.T. 1887. Contributions to the Mol- 
lusca of Florida. Proceedings of the Daven- 
port Academy of Natural Sciences, 5: 45-72. 

SYKES E.R. 1925. On the Mollusca procured dur- 
ing the “Porcupine” Expeditions, 1869-70. 
Supplemental notes, part V. Proceedings of 
the Malacological Society of London 16: 181- 
195, plo. 

THIELE J. 1929-1935. Handbuch der Systemati- 
schen Weichtierkunde. 2 vols. 1154 p., 584 
figs. 

TREECE G.D. 1980. Bathymetric records of ma- 
rine shelled Mollusca from the Northeast- 
ern shelf and upper slope of Yucatan, Mex- 
ico. Bulletin of Marine Science, 30(3): 552- 
570. 

TRYON G.W. 1887. Manual of Conchology (sec- 
ond series). 3 Helicidae:- vol. 1. Academy of 
Natural Sciences: Philadelphia. xii + [1]-313 
+ 63 pls. 

TRYON G.W. 1888. Monograph of the families 
Neritidae, Neritopsidae, Adeorbiidae, Cy- 
clostrematidae, and Liotiidae. Manual of 
Conchology 10. Academy of Natural Sci- 
ences: Philadelphia. 3-160, pls. 1-36. 

TUNNELL J.W. €z CHANEY A.H. 1970. A check- 
list of the mollusks of Seven and One-Half 
Fathom Reef, Northwestern Gulf of Mexico. 
Cont. Marine Science University Texas., 15: 
192-203. 

VANATTA E.G. 1904. A list of shells collected in 
western Florida and Horn Island, Missis- 
sippi. Proceedings of the Academy of Natural 
Sciences of Philadelphia, 55: 756-759. 


VANATTA E.G. 1913. Descriptions of new species 
of marine shells. Proceedings of the Acad- 
emy of Natural Sciences of Philadelphia, 65: 
22-27, pL.2. 

VERRILL A.E. 1882. Catalogue of marine Mol- 
lusca added to the fauna of the New England 
region, during the past ten years. Transactions 
of the Connecticut Academy of Arts and Sci- 
ences, 5: 451-587, pls. 42-44, 57-58. 

VERRILL A.E. 1884. Second catalogue of Mollusca 
recently added to the fauna of the New Eng- 
land coast and the adjacent parts of the Atlantic, 
consisting mostly of deep-sea species, with 
notes on others previously 
recorded. Transactions of the Connecticut 
Academy of Arts and Sciences, 6: 139-294, pls. 
28-32. 

VOKES H.E. € VOKES E.H. “1983” [1984]. Distri- 
bution ofshallow-water marine Mollusca, Yu- 
catan Peninsula, Mexico. Mesoamerican Ecol- 
ogy Institute Monograph 1, Middle Ameri- 
can Research Institute Publication, 54: viii, 183 
pp, 50 pls. 

WARÉN A. 1991. New and little known 
“Skeneimorph” gastropods from the Mediter- 
ranean Sea and the adjacents Atlantic Ocean. 
Bolletino Malacologico, 27(10-12): 149-248. 

WAREN A. 1996. New and little known mollusca 
from Iceland and Scandinavia. Part 3. Sarsia, 
81: 197-245. 

WARÉN A. 6: BOUCHET P. 1993. New records, 
species, genera and a new family of gastropods 
from hydrothermal vents and hydrocarbon 
seeps. Zoologica Serip ta, 22: 1-90. 

WARÉN A. 6 BOUCHET P. 2001. Gastropoda and 
Monoplacophora from Hydrotermal vents 
and hydrocarbon seeps; new taxa and records. 
The Veliger, 44(2): 116-231. 

WARMKE G.L. € ABBOTT R.T. 1961. Caribbean 
seashells: a guide to the marine mollusks of 
Puerto Rico and other West Indian Islands, 
Bermuda and the Lower Florida Keys. Liv- 
ingston Publishing Company, Wynnewood, 
PA. 348 pp., 44 pls. 

WAass M.L. 1972. A check list of the biota of lower. 
Chesapeake Bay. Va. Inst. Mar. Sci. Spec. Sci. 
Rept. No. 65, 290 p. 

WEISBORD N.E. 1962. Late Cenozoic gastropods 
from northern Venezuela. Bulletins of Amer- 
ican Paleontology, 42(193): 1-672, 48 pls. 

WILEY G.N., CIRCÉ R.C. 82 TUNNELL J.W. JR. 1982. 
Mollusca of the rocky shores of east central Ve- 
racruz State, Mexico. The Nautilus, 96: 55-61. 

WOODRING W.P. 1928. Miocene mollusks from 
Bowden, Jamaica. Part II: Gastropods and dis- 
cussion of results. Carnegie Institution of Wash- 
ington. Publication, 395: vii + 564 p., 40 pls. 

WOODWARD M.F. 1898. On the anatomy of Ade- 
orbis subcarinatus, Montagu. Proceedings 
of the Malacological Society of London, 3: 140- 
146, pl. 8. 


LS 


RUBIO £7 4£.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


ALPHABETICAL INDEX OF GENERA AND SPECIES 


Plate numbers, page numbers 


abnorme, Teinostoma 
abyssicola, Ganesa 
abyssicola, Molleriopsis 
adamsil, Cochliolepis 
Adeorbis 

Adeorbis adamsii 
Adeorbis beauli 
Adeorbis elegans 
Adeorbis infracarinata 
Adeorbis nautiliformis 
Adeorbis omalos 
Adeorbis orbignyi 
Adeorbis supranitidus 
aguayoi, Vitrinella 
altum, Teinostoma 
anastomosis, Teinostoma 
Anticlimax annae 
anneliesae, Vitrinella 
Annulicallus 

anomala, Vitrinella 
Anticlimax 

Anticlimax annae 
Anticlimax athleenae 
Anticlimax crassilabris 
Anticlimax decorata 
Anticlimax glabra 
Anticlimax hispaniolensis cratera 
Anticlimax locklin: 
Anticlimax pilsbryl 
Anticlimax proboscidea 
Anticlimax schumoi 
antillensis, Solariorbis 
Aorotrema 

Aorotrema cistrontum 
Aorotrema erraticum 
Aorotrema gardnerae 
Aorotrema pontogenes 
aristata, Vitrinella 
athleenae, Anticlimax 
avitus, Parviturboides 


avunculus, Teinostoma 


SL, 
62, 


112, 


101, 
23, 
12, 
40, 
90, 


108, 


40, 


35, 
37, 
36, 
41, 
39, 
38, 
37, 


73, 


42, 


114, 


98, 


200 
215 
216 
108 
201 
108 

84 
201 
139 


162 


192 


195 


135 


202 
197 


173 
195 
197 
191 


Plate numbers, page numbers 


baldingeri, Ternostoma 
balesi, Parviturbo1des 
bartschi, Cyclostremiscus 
beauti, Cyclostremiscus 
biscaynense, Teinostoma 
blake1, Solariorbis 
brychia, Choristella 
bushae, Ganesa 

calliglyp ta, Vitrinella 
caloosaense, Teinostoma 
canaliculata, Vitrinella 
Canimarina 
caraboboensis, Tornus 
carinata, Vitrinella 
carinicallus, Teinostoma 
carinifex, “Vitrinella” 
cerion, Mikro 
Choristella 

Choristella brychia 
Choristella lep talea 
cienfuegosense, Teinostoma 
Circulus 

Circulus bicarinatus 
Circulus cubanus 
Circulus dalli 

Circulus gunteri 
Circulus hendersoni 
Circulus liratus 

Circulus margaritiformis 
Circulus orbignyl 
Circulus quadricristatus 
Circulus semisculp tus 
Circulus stirophorus 
Circulus striatus 
Circulus suppressus 
Circulus texanus 
Circulus translucens 
Circulus trilix 
Cirsonella 

Cirsonella extrema 


Cirsonella floridensis 


10, 24 
AL ASS 
49, 95 
43, 84 
14, 29 
74, 137 
102, 181 
77 T/3 
39, 63 
32, 6l 


115, 202 
118, 208 


29, 56 


106, 197 


116, 205 


229 


Iberus, 29 (2), 2011 


Plate numbers, page numbers 


Cirsonella margaritiformis 117, 
ciskae, Teinostoma 5, 
cistrontum, Aorotrema 42, 
Climacia 

Climacia athleenae 

Climacia tholus 

Climacina 

Cochliolepis 

Cochliolepis adamst 57, 
Cochliolepis differens 66, 
Cochliolepis holmesti 58, 
Cochliolepis nautiliformis 59, 
60,61,62, 
Cochliolepis patriciol 65, 


Cochliolepis parasitica 


Cochliolepis planispiralis 64, 
Cochliolepis sp. 67, 
Cochliolepis striata 63, 
Cochliolepis surinamensis 

cocolitoris, Teinostoma 26, 
conica, Ganesa 

contracta, Vitrinella 91, 
convexa, Ganesa 

crassilabris, Anticlimax 35, 
cubanus, Cyclostremiscus 54, 
cupidinensis, Vitrinella 103, 
cyclostomoides, Vitrinella 

Cyclostrema 

Cyclostrema angulatum 

Cyclostrema bartschi 

Cyclostrema bicarinatum 

Cyclostrema cistrontum 

Cyclostrema elegans 

Cyclostrema gordana 

Cyclostrema interrup tum 

Cyclostrema pontogenes 

Cyclostrema sanibelense 

Cyclostrema schrammii 

Cyclostrema zacalles 

“Cyclostrema” thomasi 

Cyclostremella humilis 
Cyclostremiscus 

Cyclostremiscus bartschi 49, 
Cyclostremiscus beauil 43, 


Cyclostremiscus caraboboensis 


226 


207 
5 
80 
67 

195 
74 
67 

108 

108 

123 

109 

112 

ap2 

120 

118 

123 

118 

189 
50 

Zi 

164 

ZIS 
68 

103 

181 

203 

215 
84 
95 
84 
80 

201 
86 

129 
81 

128 
64 

129 

186 

ZN 
83 
99, 
84 
63 


Plate numbers, page numbers 


Cyclostremiscus crassilabris 
Cyclostremiscus cubanus 
Cyclostremiscus dalli 
Cyclostremiscus diminutus 
Cyclostremiscus euglyptus 
Cyclostremiscus fargoi 
Cyclostremiscus gunteri 
Cyclostremiscus hendersont 
Cyclostremiscus jeannae 
Cyclostremiscus major 
Cyclostremiscus microstriatus 
Cyclostremiscus multiliratus 
Cyclostremiscus olssoni 
Cyclostremiscus pentagonus 
Cyclostremiscus puntagordensis 
Cyclostremiscus salvatierrensis 
Cyclostremiscus suppressus 
Cyclostremiscus trilix 
Cyclostremiscus vanbruggeni 
dalli, Cyclostremiscus 

dalli, Ganesa 

decorata, Anticlimax 
Delphinoidea 

Delpinula depressa 
diaphana, Ganesa 

differens, Cochliolepis 
Dillwynella 

Dillwynella modesta 
diminutus, Cyclostremiscus 
Discopsis omalos 

Discopsis schumoi 

elegans, Adeorbis 

elegans, Solariorbis 

elegans, Vitrinorbis 
Ellipetylus 

Episcynia 

eritmeta, Leptogyra 
erraticum, Aorotrema 
Ethalia multistriata 

Ethalia reclusa 

Ethalia solida 

eugenes, Solariorbis 
euglyptus, Cyclostremiscus 


expansum, Teinostoma 


54, 
45, 
53, 
55, 
110, 


48, 
50, 


56, 
44, 


46, 
51, 
47, 
52, 


45, 


E 


66, 


121, 
53, 


75, 
106, 


114, 


111, 
55, 
16, 


68 
103 
88 
100 
105 
197 
197 
93 
97 
86 
105 
86 
197 
10,88 
103 
108 
97 
Sil 
100 
88 
ZAS 
70 
181 
18S 
215 
123 
215 
216 
100 
189 
189 
201 
199 
186 
50 
LS) 
216 
202 
145 
21 
48 
197 
105 
34 


RUBIO ET 42.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Plate numbers, page numbers 


fargo1, Cyclostremiscus 
filifera, Vitrinella 
floridana, Vitrinella 
floridensis, Cirsonella 
funiculus, Vitrinella 
Ganesa 

Ganesa bushae 

Ganesa conica 

Ganesa convexa 
Ganesa dalli 

Ganesa depressa 
Ganesa diaphana 
Ganesa ornatam 
Ganesa proxima 
Ganesa valvata 

Ganesa verrilli 
gardnerae, Aorotrema 
georgiana, Cirsonella 
glabra, Anticlimax 
goniogyrus, Teinostoma 
Granigyra 

Granigyra limata 
Granigyra radiata 
Granigyra spinulosa 
gutanensis, Solariorbis 
gunteri, Cyclostremiscus 
helicinum, Teinostoma 
helicoidea, Vitrinella 
Helix cyclostomoides 
hemphilli, Vitrinella 


hendersoni, Cyclostremiscus 


110, 
104, 
96, 
116, 
92; 


19, 


27, 
94, 95, 


48, 


hispaniolensis cratera, Anticlimax 41, 


holmesti, Cochliolepis 
Homalogyra marshalli 
hondurasensis, Solariorbis 
humilis, Cyclostremella 
hyalina, Vitrinella 
Idioraphe 

incertum, Teinostoma 
inconspicua, Leptogyra 
infracarinatus, Solariorbis 
mornata, Episcynia 
Interruptus, Parviturboides 


jeannae, Cyclostremiscus 


58, 


167, 


197 
184 
pz 
205 
165 
24119) 
Z15 
ZAD 
215 
215 
Z15 
SAUS) 
215 
Z15 
215 
ZAS 
197 
207 

70 

16 
216 
216 
216 
216 
189 
197 

53 
195 
203 
192 

93 

78 
109 
199 
190 
287 
204 


24, 29, 33 


11, 


76, 77, 
68, 69, 
70, 71, 

50, 


24 
216 
189 
126 
129, 

7 


Plate numbers, page numbers 


lenticulare, Teinostoma 
leptalea, Choristella 
Leptogyra 

Leptogyra eritmeta 
Leptogyra inconspicua 
Leptogyra verrilli 

lerema, Teinostoma 
limata, Granigyra 
lineopunctatus, Solariorbis 
liratus, Circulus 
lituspalmarum, Teinostoma 
locklini, Anticlimax 
lunense, Teinostoma 
Lydiphnis dalli 

Lydiphnis hendersoni 
Lydiphnis margaritiformis 
Lydiphnis translucens 
margaritiformis, Cirsonella 
marshalli, Neusas 
massarita, “Vitrinella” 
megacallum, Teinostoma 


megastoma, Teinostoma 


microstriatus, Cyclostremiscus 


Mikro 

Mikro cerion 
minusculum, Teinostoma 
modesta, Dillwynella 
Molleriopsis sincera 
mooreanus, Solariorbis 
morlierei, Teinostoma 
multiliratus, Cyclostremiscus 
multistriatus, Solariorbis 
nautiliformis, Cochliolepis 
Nemafera compressa 
nesaeum, Teinostoma 
Neusas 

Neusas marshalli 
obtectum, Teinostoma 
olssoni, Cyclostremiscus 
Omalaxis funiculus 
Omalaxis lirata 

omalos, Discopsis 
opsitelotus, Vitrinella 


orbigny1, Circulus 


7, 


117, 
112, 


31, 
28, 
56, 
118, 
20, 
121, 
78, 
44, 
79, 
59 


17, 


112, 
15, 


93, 


E) 
216 
216 
216 
216 
216 


Z15 
155 


Za 


Iberus, 29 (2), 2011 


Plate numbers, page numbers 


ornatam, Ganesa 
Paludinella helicoides 
panamense, Teinostoma 
parasitica, Cochliolepis 
parvicallum, Temnostoma 
Parviturbo 

Parviturbo zacalles 
Parviturboides 
Parviturboides avitus 
Parviturboides balesi 
Parviturboides interruptus 
parvum, Teinostoma 
patricio1, Cochliolepis 
pauli, Pleuromalaxis 
pelorcei, Vitrinella 
pentagonus, Cyclostremiscus 
petiti, Solariorbis 
pilsbryi, Anticlimax 
planispiralis, Cochliolepis 
Pleuromalaxis 
Pleuromalaxis pauli 
Ponderinella 
Ponderinella xacriaba 
pontogenes, Aorotrema 
Porcupinia 

proboscidea, Anticlimax 
proxima, Ganesa 
pseudoaristata, Vitrinella 
Pseudomalaxis 
Pseudomalaxis balesi 
Pseudorbis granulum 
Pseudorotella carinicallus 
Pseudorotella floridensis 
Pseudorotella minúscula 
Pseudorotella pusilla 
Pseudorotella semistriata 
punctostriatus, Solariorbis 
pusilla, Vitrinella 

radiata, Granigyra 
reclusum, Teinostoma 
regularis, Vitrinella 
rhyssa, Xyloskenea 
Rotella carinata 


Rotella cryptospira 


228 


13, 
60,61,62, 
30, 


72, 
70, 71, 


65, 


100, 
46, 
107, 
38, 
64, 


ZS 
1159 

29 
Jo 

DY 
132 
129 
128 
197 
133 
129 
192 
120 
190 
176 

88 
190 

ZO 
118 
182 


133, 190 


113, 


3, 


99, 


87, 
108G, 109, 


8, 9, 


119, 


200 
200 
81 
205 
73 
ZO 
176 
132 
SS 
132 
61 
205 
41 
193 
38 
157 
193 
215 
Z il 
201 
210 
16 
43 


Plate numbers, page numbers 


Rotella lenticularis 
Rotella pusilla 

Rotella umbellicata 

ruris, Solariorbis 
schrammii, Tornus 
schumoi, Anticlimax 
schumoi, Solariorbis 
semipunctus, Solariorbis 
semisculptus, Circulus 
semistriatum, Teinostoma 
sincera, Molleriopsis 
Skenea larseni 

Skenea petiti 

Skenea sulcata 

Skenea trilix 

Skenea trochoides 
Solariorbis 

Solariorbis aguayol 
Solariorbis antillensis 
Solariorbis bassilisus 
Solariorbis blake1 
Solariorbis corylus 
Solariorbis decipiens 
Solariorbis elegans 
Solariorbis eugenes 
Solariorbis euzonus 
Solariorbis gutanensis 
Solariorbis hondurasensis 
Solariorbis infracarinatus 
Solariorbis lineopunctatus 
Solariorbis mooreanus 
Solariorbis multistriatus 
Solariorbis opsitelotus 
Solariorbis petitii 
Solariorbis punctostriatus 
Solariorbis ruris 
Solariorbis schumo1 
Solariorbis semipunctus 
Solariorbis solidus 
Solariorbis sp. 
Solariorbis terminalis 
Solariorbis truncatus 
solaris, Vitrinella 


Solarium inornatum 


85, 
34, 


79, 


74, 


75, 
111, 
139, 

73, 


76, 77, 
86, 
78, 
79, 


107, 
87, 

85, 

8l, 

82, 

88, 89, 
86, 

3, 84, 
80, 
105, 


RUBIO £7 41.: The family Tornidae (Gastropoda, Rissooidea) in the Caribbean 


Plate numbers, page numbers 


solidum, “Teinostoma” 
solidus, Solariorbis 

sp. , Cochliolepis 

sp., Solariorbis 

spinulosa, Granigyra 
striata, Cochliolepis 
striata, Ganesa 
Striovitrinella 

suppressus, Cyclostremiscus 
surinamensis, Cochliolepis 
tectispira, Teinostoma 
Teinostoma 

Teinostoma (Ethalia) suppressa 
Teinostoma altum 
Teinostoma anastomosis 
Teinostoma avunculus 
Teinostoma baldingeri 
Teinostoma bartschi 
Teinostoma biscaynense 
Teinostoma caloosaense 
Teinostoma carinicallus 
Teinostoma cienfuegosense 
Teinostoma ciskae 
Teinostoma clavium 
Teinostoma cocolitoris 
Teinostoma expansum 
Teinostoma funiculus 
Teinostoma goniogyrus 
Teinostoma helicinum 
Teinostoma hondurasensis 
Teinostoma incertum 
Teinostoma lenticulare 
Teinostoma lerema 
Teinostoma lituspalmarum 
Teinostoma lunense 
Teinostoma megacallum 
Teimostoma megastoma 
Teinostoma millepunctata 
Teinostoma minusculum 
Teinostoma morlierei 
Teinostoma nesaeum 
Teinostoma obtectum 
Teinostoma opsitelotus 


Teinostoma panamense 


24, 
88, 89, 
67, 
86, 
63, 


139, 
51, 


139, 


20, 


17, 
15, 


13, 


48 
159 
123 
159 
ZN 
118 
215 
181 

97 
189 
199 

14 

97 

46 

2h 
191 

24 
142 

29 
199 

61 

56 

15 

38 

50 

34 
165 

16 

D9 
190 

24 

19 

43 

61 

50 

59 

54 

15 

41 
19] 

36 

33 
167 

2) 


Plate numbers, page numbers 


Teinostoma parvicallum 
Teimostoma parvum 
Teinostoma pilsbryi 
Teimostoma politum 
Teinostoma proboscidea 
Teinostoma reclusum 
Teinostoma schumo1 
Teinostoma semistriatum 
“Teinostoma” solidum 
Teimostoma tectispira 
Teinostoma umbilicatum 
terminalis, Solariorbis 
Texanus, Circulus 
Tharsiella 

Tharsis 

tincta, Vitrinella 

Tornus 

Tornus caraboboensis 
Tornus schramm il 
translucens, Xyloskenea 
trilix, Cyclostremiscus 
Trochus (Rotella) anómala 
Trochus (Rotella) carinata 
Trochus (Rotella) diaphana 
Trochus (Rotella) semistriata 
Trochus inornatum 
truncatus, Solariorbis 
tryon1, Vitrinella 
umbilicatum, Teinostoma 
valvata, Ganesa 
vanbruggeni, Cyclostremiscus 
verrilli, Ganesa 

verrilli, Leptogyra 
Vitrinella 

Vitrinella aguayol 
Vitrinella anneliesae 
Vitrinella anomala 
Vitrinella aristata 
Vitrinella blakei 
Vitrinella calliglyp ta 
Vitrinella canaliculata 
Vitrinella carinata 
Vitrinella cerion 


Vitrinella contracta 


307 97 
192 

23 

14 

73 

90) 32 
147 
18,19, 38 
24, 48 
199 

22, 44 
83,84, 150 
11 

205 

205 

114, 202 
63 

33, 63 
34, 64 
1207-2111 
47, 91 
192 

192 

193 

38 

126 

80, 145 
195 

22, 44 
Z15 

52, 100 
Z15 

216 

159 

101, 178 
90, 162 
108, 192 
98, 173 
197 

102, 181 
IS ALZO 
192 

208 

91, 164 


229 


Iberus, 29 (2), 2011 


Plate numbers, page numbers 


Vitrinella cupidinens 
Vitrinella cyclostomoides 
Vitrinella diaphana 
Vitrinella elegans 
Vitrinella filifera 
Vitrinella floridana 
Vitrinella funiculus 
Vitrinella georgiana 
Vitrinella helicoidea 
Vitrinella hemphilli 
Vitrinella holmesti 
Vitrinella hyalina 
Vitrinella interrup ta 
“Vitrinella” massarita 
Vitrinella megastoma 
Vitrinella mooreana 
Vitrinella multicarinata 
Vitrinella opsitelotus 
Vitrinella panamensis 
Vitrinella pelorcei 
Vitrinella pentagonus 
Vitrinella praecox 


Vitrinella pseudoaristata 


230 


94, 95, 


103, 157, 181 


203 

139) 

59 

104, 184 
IS 17d 
O2AAIOS 
207 

LO USO 
192 

109 

204 
125429 
ZA 

54 

143 

126 

OS 1O67 
DO YY 
100, 176 
88 

167 
OO 


Plate numbers, page numbers 


Vitrinella pusilla 
Vitrinella regularis 
Vitrinella rhyssa 
Vitrinella semisculp ta 
Vitrinella solaris 
Vitrinella tenuisculp ta 
Vitrinella terminalis 
Vitrinella texana 
Vitrinella thomasi 
Vitrinella tincta 
Vitrinella truncata 
Vitrinella tryoni 
Vitrinella tryoni 
“Vitrinella” carinifex 
Vitrinorbis 

Vitrinorbis callistus 
Vitrinorbis elegans 
xacriaba, Ponderinella 
Xyloskenea 
Xyloskenea costulifera 
Xyloskenea naticiformis 
Xyloskenea rhyssa 


Xyloskenea translucens 


108G, 109, 


105, 


115, 


106, 
113, 


119, 
120, 


NORMAS DE PUBLICACIÓN 


e La revista Iberus publica artículos de fondo, notas y monografías que versen sobre cualquiera de los aspectos 
relacionados con la Malacología. Se entiende por artículo un trabajo de investigación de más de 5 páginas de 
texto, incluidas láminas, gráficos y tablas. Las notas son trabajos de menor extensión. Las monografías son tra- 
bajos sobre un tema único, de extensión superior a las 50 páginas de la revista y que serán publicadas, si 
procede, como un suplemento de Iberus. Los autores interesados en publicar monografías deberán ponerse 
previamente en contacto con el Editor de Publicaciones. Se entiende que el contenido de los manuscritos no ha 
sido publicado, ni enviado simultáneamente a otra revista para su consideración. 

e Los manuscritos, así como toda la correspondencia relacionada con los mismos, deberán ser remitidos a: 
Serge Gofas, Editor de Publicaciones, Departamento de Biología Animal, Universidad de Málaga, Campus de 
Teatinos, s/n, 29071, Málaga, España y / o al correo electrónico <sgofasWuma.es>. 

e El texto del trabajo podrá estar redactado en español, inglés, italiano, francés o portugués. 

e Los artículos, notas y monografías deberán presentarse sobre DIN A-4, por una sola cara a doble espacio 
(incluyendo referencias, pies de figura y tablas), con al menos 3 centímetros de margen por cada lado. Los tra- 
bajos se entregarán por triplicado (original y dos copias) y se incluirá una versión en un CD, o bien remitida 
por correo electrónico, utilizando procesadores de texto en sus versiones de corrientes de Windows (pero no 
en el formato .docx de Word 2007, el habitual de Windows Vista) o Macintosh. En caso de autoría compartida, 
uno de los autores deberá hacerse responsable de toda la correspondencia. 

e Junto con el trabajo debe incluirse una lista de al menos 4 posibles revisores del mismo, sin perjuicio de los 
que el propio Editor considere oportunos. 

e Los manuscritos se presentarán de acuerdo al siguiente esquema: 

Primera página. Deberá incluir un título conciso, pero sugerente del contenido del trabajo, así como una tra- 
ducción al inglés del mismo (si el artículo no está escrito en inglés). Cuando sea preciso, deberá incluir referen- 
cia a familia o taxones superiores. A continuación figurarán, por este orden, el nombre y apellidos completos 
del autor o autores, las direcciones completas de los mismos, y un resumen del trabajo y su traducción al 
inglés. Dicho resumen deberá sintetizar, en conjunción con el título, tanto los resultados como las conclusiones 
del artículo; se sugiere una extensión de 100 a 200 palabras. 

Páginas siguientes. Incluirán el resto del artículo, que debe dividirse en secciones precedidas por breves enca- 
bezamientos. Siempre que sea posible, se recomienda seguir el siguiente esquema: Introducción, Material y 
métodos, Resultados, Discusión, Conclusiones, Agradecimientos y Bibliografía. Si se emplean abreviaturas no 
habituales en el texto, deberán indicarse tras el apartado de Material y Métodos. 

e Las notas breves deberán presentarse de la misma forma, pero sin resumen. 

e Deberán evitarse notas a pie de página y referencias cruzadas. Deberán respetarse estrictamente los Códigos 
Internacionales de Nomenclatura Zoológica y Botánica (últimas ediciones). Cuando un taxón aparezca por 
primera vez deberá citarse su autor y fecha de su descripción. En el caso de artículos sistemáticos, cuando se 
den las sinonimias de los taxones, éstas deberán citarse COMPLETAS, incluyendo en forma abreviada la 
publicación donde fueron descritas, y la localidad tipo si es conocida entre corchetes, según el siguiente 
esquema (préstese especial cuidado a la puntuación): 

Dendrodoris limbata (Cuvier, 1804) 

Sinonimias 

Doris limbata Cuvier, 1804, Ann. Mus. Hist. Nat. Paris, 4 (24): 468-469 [Localidad tipo: Marsella]. 

Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop.-Car., 10: 275. 

Dichas referencias no deberán incluirse en la lista de Bibliografía si es la única vez que se nombran en el texto. 
Si se incluyen una lista completa de referencias de un taxón inmediatamente tras éste, deberá seguirse el 
mismo esquema (sin incluir en Bibliografía las referencias que no se mencionen en otro lugar del texto). 

e Sólo los nombres en latín y los de taxones genéricos y específicos deberán llevar subrayado sencillo o prefe- 
rentemente ir en cursiva. En ningún caso deberá escribirse una palabra totalmente en letras mayúsculas, ni 
siquiera el Título. Las unidades a utilizar deberán pertenecer al Sistema Métrico Decimal, junto con sus correc- 
tas abreviaturas. En artículos escritos en castellano, en los números decimales sepárese la parte entera de la 
decimal por una coma inferior (,), NUNCA por un punto (.) o coma superior ('). 

e Las referencias bibliográficas irán en el texto con minúsculas o versalitas: Fretter y Graham (1962) o FRETTER 
Y GRAHAM (1962). Si son más de dos autores se deberán citar todos la primera vez que aparecen en el texto 
[Smith, Jones y Brown (1970)] empleándose et al. las siguientes veces [Smith et al. (1970)]. Si un autor ha publi- 
cado más de un trabajo en un año se citarán con letras: (Davis, 1989a; Davis, 1989b). No deberá emplearse op. 
cit. La lista de referencias deberá incluir todas las citas del texto y sólo éstas, ordenadas alfabéticamente. Se 
citarán los nombres de todos los autores de cada referencia, sea cual sea su número. Los nombres de los 
autores deberán escribirse, en letras minúsculas o Versalitas. No deberán incluirse referencias a autores 
cuando éstos aparezcan en el texto exclusivamente como autoridades de un taxón. Los nombres de las publica- 


ciones periódicas deberán aparecer COMPLETOS, no abreviados. Cuando se citen libros, dése el título, editor, 
lugar de publicación, n” de edición si no es la primera y número total de páginas. Deberán evitarse referencias 
a Tesis Doctorales u otros documentos inéditos de difícil consulta. Síganse los siguientes ejemplos (préstese 
atención a la puntuación): 

Fretter V. y Graham A. 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp. 

Ponder W.F. 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. En Ponder, W.F. 
(Ed.): Prosobranch Phylogeny. Malacological Review, suppl. 4: 129-166. 

Ros J. 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Mis- 
celánea Zoológica, 3 (5): 21-51. 

e Las gráficas e ilustraciones deberán ser originales y presentarse preferentemente en formato electrónico al 
formato de caja de la revista o proporcional a éste. Este formato es de 57 mm (una columna) o 121 mm (dos) de 
anchura y hasta 196 mm de altura, si bien se recomienda utilizar el formato a dos columnas. Es importante que 
todas las figuras sean remitidas en su formato original (por ejemplo, las fotografías en .j¡pg de alta calidad o .tif, 
las gráficas en hojas de cálculo Excel o documentos de CorelDraw), puesto que las ilustraciones insertadas en 
el manuscrito WORD son inservibles en la fase de imprenta. Las imágenes digitales deben ser formateadas en 
su tamaño de impresión con una resolución mínima de 300 ppp para imágenes en color o escala de grises y de 
600 ppp para las de blanco y negro. Considérese la reducción que será necesaria a la hora de decidir el tamaño 
de las escalas o letras en las figuras, que no deberán bajar de los 2 mm. En figuras compuestas, cada parte 
deberá etiquetarse con letras mayúsculas, el resto de las letras deberán ser minúsculas. Las escalas de dibujos y 
fotografías deberán ser gráficas, utilizando unidades del sistema métrico decimal; no deberán hacerse referen- 
cias a los aumentos de una determinada ilustración, ya que éstos cambian con la reducción. En su caso, se 
recomienda la utilización de mapas con proyección UTM. Cada figura, gráfica o ilustración deberá presentarse 
en hojas separadas y con numeración arábiga (1, 2, 3,...), sin separar “Figuras” y “Láminas”. Los pies de 
figura, en una hoja aparte, deberán acompañarse de su traducción al inglés (o español si el inglés es la lengua 
del trabajo). Utilícese el esquema siguiente: 

Figura 1. Neodoris carvi. A: animal desplazándose; B: detalle de un rinóforo; C: branquia. 

Las abreviaturas empleadas en las ilustraciones deberán incluirse en los pies de figura. 

Los autores interesados en incluir láminas en color deberán consultar con el editor y sufragar el sobrecoste con 
una contribución de 30 € por página. Por lo demás, deberán ajustarse a los mismos requisitos indicados para 
las figuras. 

Si se pretende enviar gráficas o ilustraciones en impresión de papel es imprescindible presentar originales de 
buena calidad. Las imágenes en semitonos deben estar bien contrastadas y ajustarse al tamaño definitivo de 
impresión; al componer fotografías sobre una hoja, procúrese que los espacios entre ellas sean regulares y que 
estén debidamente alineadas. Téngase en cuenta que incluir fotografías de distinto contraste en una misma 
página conlleva una pobre reproducción final. Las gráficas de ordenador deberán imprimirse con impresora 
láser sobre papel de buena calidad. 

e Las Tablas se presentarán en hojas separadas, siempre con numeración romana (l, II, IIL...). Las leyendas se 
incluirán en una hoja aparte acompañándose de una traducción al inglés. Deberán evitarse las tablas particu- 
larmente complejas. Se recomienda reducir el número y extensión de ilustraciones, láminas o tablas al mínimo 
necesario. 

e Los artículos que no se ajusten a las normas de publicación serán devueltos al autor con las indicaciones de 
los cambios necesarios. 

e El Comité Editorial comunicará al autor responsable del trabajo la fecha de recepción del trabajo y la fecha 
de envío a revisión. Cada original recibido será sometido a revisión por al menos dos investigadores. El 
Comité Editorial, a la vista de los informes de los revisores decidirá sobre la aceptación o no de cada manus- 
crito. El autor recibirá en cada caso copia de los comentarios de los revisores sobre su artículo. En caso de 
aceptación, el mismo Comité Editorial, si lo considera conveniente, podrá solicitar a los autores otras modifica- 
ciones que considere oportunas. Si el trabajo es aceptado, el autor deberá enviar una copia impresa del mismo 
corregida, acompañada por una versión en un CD, utilizando procesadores de texto en sus versiones corrien- 
tes de Windows (pero no el formato .docx generado por Word 2007, el habitual de Windows Vista) o Macin- 
tosh. La fecha de aceptación figurará en el artículo publicado. 

e Las pruebas de imprenta serán enviadas por correo electrónico al autor responsable, exclusivamente para la 
corrección de erratas, y deberán ser devueltas en un plazo máximo de una semana. Se recomienda prestar 
especial atención en la corrección de las pruebas. 

e De cada trabajo se entregarán gratuitamente 50 separatas, además de una versión electrónica en formato 
«pdf. Aquellos autores que deseen un número mayor, deberán hacerlo constar al devolver las pruebas de 
imprenta, y nunca posteriormente. El coste de las separatas adicionales será cargado al autor. 


INSTRUCTIONS TO AUTHORS 


e Iberus publishes research papers, notes and monographs devoted to the various aspects of Malacology. 
Papers are manuscripts of more than 5 typed pages, including figures and tables. Notes are shorter papers. 
Monographs should exceed 50 pages of the final periodical, and may be published as Supplements. Authors 
wishing to publish monographs should contact the Editor. Manuscripts are considered on the understanding 
that their contents have not been published or simultaneously submitted for publication elsewhere. 

e Manuscripts and correspondence regarding editorial matters must be sent to: Serge Gofas, Editor de Publica- 
ciones, Departamento de Biología Animal, Universidad de Málaga, Campus de Teatinos, s/n, 29071, Málaga, 
Spain and / or to the e-mail <sgofasVuma.es>. 

+ Manuscripts may be written in Spanish, English, Italian, French or Portuguese. 

+ Manuscripts must be typed double spaced (including the references, figure captions and tables) on one side 
on A-4 (297x210 mm) with margins of at least 3 cm. An original and two copies must be submitted, together 
with a CD or e-mail remittance containing the article written with a current Windows (but not .docx format 
generated by Word 2007, mainly used with Windows Vista) or Macintosh word processor. When a paper has 
joint authorship, one author must accept responsibility for all correspondence. 

e The authors must include a list of at least 4 possible referees; the Editor can choose any others if appropriate. 

e Papers should conform the following layout: 

First page. This must include a concise but informative title, with mention of family of higher taxon when 
appropriate, and its Spanish translation. It will be followed by all authors” names and surnames, their full 
address(es), an abstract (and its Spanish translation) not exceeding 200 words which summarizes not only con- 
tents but results and conclusions. 

Following pages. These should content the rest of the paper, divided into sections under short headings. 
Whenever possible the text should be arranged as follows: Introduction, Material and methods, Results, Dis- 
cussion, Conclusions, Acknowledgements and References. Unusual abbreviations used in the text must be 
grouped in one alphabetic sequence after the Material and methods section. 

e Notes should follow the same layout, without the abstract. 

e Footnotes and cross-references must be avoided. The International Codes of Zoological and Botanical 
Nomenclature must be strictly followed. The first mention in the text of any taxon must be followed by its 
authority including the year. In systematic papers, when synonyms of a taxon are given, they must be cited IN 
FULL, including the periodical, in an abbreviate form, where they were described, and the type localities in 
square brackets when known. Follow this example (please note the punctuation): 

Dendrodoris limbata (Cuvier, 1804) 

Synonyms 

Doris limbata Cuvier, 1804, Ann. Mus. Hist. Nat. 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. If a 
full list of references of the taxon is to be given immediately below it, the same layout should be followed (also 
excluding from the Bibliography list those which are not cited elsewhere). 

Only Latin words and names of genera and species should be underlined once or be given in italics. No word 
must be written in UPPER CASE LETTERS. SI units are to be used, together with their appropriate symbols. In 
Spanish manuscripts, decimal numbers must be separated with a comma (,), NEVER with a point (.) or upper 
comma ('). 

e References in the text should be written in small letters or Small capitals: Fretter and Graham (1962) or 
FRETTER AND GRAHAM (1962). The first mention in the text of a paper with more than two authors must 
include all of them [Smith, Jones and Brown (1970)], thereafter use et al. [Smith et al. (1970)]. If an author has 
published more than one paper per year, refer to them with letters: (Davis, 1989a; Davis, 1989b). Avoid op. cit. 

The references in the reference list should be in alphabetical order and 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 tax- 
onomic name. Titles of periodicals must be given IN FULL, not abbreviated. For books, give the title, name of 
publisher, place of publication, indication of edition if not the first and total number of pages. Keep references 
to doctoral theses or any other unpublished documents to an absolute minimum. See the following examples 
(please note the punctuation): 

Fretter V. and Graham A. 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp. 

Ponder W.F. 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. In Ponder W.F. 
(Ed.): Prosobranch Phylogeny. Malacological Review, suppl. 4: 129-166. 

Ros J. 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Mis- 
celánea Zoológica, 3 (5): 21-51. 


e Figures must be original and provided preferably in electronic format and adjusted to page format and 
column size. These should be one column (57 mm) or two columns (121 mm) wide and up 196 mm high, or be 
proportional to these sizes. Two columns format is recommended. It is essential that all figures be supplied in 
their original format (e.g. photographs as high-grade .jpg or as .tif files, graphs as Excel spreadsheets or Corel- 
Draw files), as the files inserted into WORD documents cannot be used for printing. Digital images must be 
given their final printing size with a resolution at least 300 dpi for colour and halftones, and at least 600 dpi for 
black / white. 

Take into account possible reduction in lettering drawings; final lettering must be at least 2 mm high. In com- 
posite drawings, each figure should be given a capital letter; additional lettering should be in lower-case 
letters. A scale line, labelled with SI units, must be used to indicate size; magnification ratio must be avoided 
as it may be changed during printing. UTM maps are to be used if necessary. Figures must be submitted on 
separate sheets, and numbered with consecutive Arabic numbers (1, 2, 3,...), without separating “Plates” and 
“Figures”. Legends for Figures must be typed in numerical order on a separate sheet, and a Spanish translation 
must be included. Follow this example (please note the punctuation): 

Figure 1. Neodoris carvi. A: animal crawling; B: rhinophore; C: gills. 

If abbreviations are to be used in illustrations, they should be included in the figure captions. 

Authors wishing to publish illustrations in colour should consult with the editor and will be charged a contri- 
bution of 30 € per page. They should otherwise follow the same standards as black and white prints. 

If the authors want to send Figures in printed format, it is essential to supply good quality originals. Half-tone 
images must be of good contrast, and should be submitted in the final printing size. When mounting pho- 
tographs in a block, ensure spacers are of uniform width. Remember that grouping photographs of varied con- 
trast results in poor reproduction. Computer graphics must be printed on high quality white paper with a 
laser printer. 

e Tables must be numbered with Roman numbexss (1, II, TIL...) and each typed on a separate sheet. Headings 
should be typed on a separate sheet, together with their English translation. Complex tables should be 
avoided. As a general rule, keep the number and extension of illustrations and tables as reduced as possible. 

e Manuscripts that do not conform to these instructions will be returned for correction before reviewing. 

e Authors submitting manuscripts will receive an acknowledgement of receipt, including receipt date, and the 
date the manuscript was sent for reviewing. Each manuscript will be critically evaluated by at least two refer- 
ees. Based of these evaluations, the Editorial Board will decide on acceptance or rejection. Anyway, authors 
will receive a copy of the referees” comments. If a manuscript is accepted, the Editorial Board may indicate 
additional changes if desirable. Acceptable manuscripts will be returned to the author for consideration of 
comments and criticism; a finalized manuscript must then be returned to the Editor, together with a CD con- 
taining the article written with current Windows (but not a .docx format generated by Word 2007, mainly used 
under Windows Vista) or Macintosh word processor. Dates of reception and acceptance of the manuscript will 
appear in all published articles. 

e Proofs will be e-mailed to the author for correcting errors and must be returned corrected within one week. 
At this stage no stylistic changes will be accepted. Pay special attention to references and their dates in the text 
and the Bibliography section, and also to numbers of Figures and Tables appearing in the text. 

e Fifty reprints per article and a .pdf file will be supplied free of charge. Additional reprints must be ordered 
when the page proofs are returned, and will be charged at cost. NO LATER orders will be accepted. 


La SocieDAD ESPAÑOLA DE IMALACOLOGÍA 


Junta Directiva desde el 11 de julio de 2011 


Presidente Jesús S. Troncoso 
Vicepresidente Gonzalo Giribet 
Secretario Ramón Álvarez Halcón 
Tesorero Luis Murillo Guillén 
Editor de Publicaciones Serge Gofas 
Bibliotecario Rafael Araujo Armero 
Vocales José Templado González 
Carmen Salas Casanovas 
Alberto Martínez Ortí 


José Ramón Arrébola Burgos 
Benjamín Gómez Moliner 


Editor invitado para vol. 29(2) Harry G. Lee 


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 2012: 


DESDE ABRIL 

Socio numerario (en Europa): 40 euros 45 euros 

(fuera de Europa): 50 euros 55 euros 
Cuota reducida estudiante 30 euros 34 euros 
Socio Familiar: (sin recepcion de revista) 5 euros 10 euros 
Socio Protector: (mínimo) 50 euros 60 euros 
Socio Corporativo (en Europa): 50 euros 60 euros 

(fuera de Europa): 60 euros 70 euros 
Cuota Conjunta SEM+SIM: 73 euros 78 euros 


INSCRIPCIÓN: 6 euros, además de la cuota correspondiente. 


A los socios residentes en España se les aconseja domiciliar su cuota. Todos los abonos deberán enviarse al 
Tesorero (dirección reseñada anteriormente) el 1 de enero de cada año. Los abonos se harán sin recargos para la 
sociedad y en favor de la Sociedad Española de Malacología y no de ninguna persona de la junta directiva. Aque- 
llos socios que no abonen su cuota anual dejarán de recibir las publicaciones de la Sociedad. Los bonos de ins- 
cripción se enviarán junto con el abono de una cuota anual al Tesorero. 

A los residentes en el extranjero se les ruega que abonen su cuota mediante giro postal en euros (internatio- 
nal postal money orders in euros sent to the Treasurer). Members living in foreing countries can deduce 6 euros 


if paid before 15 April. 


Cada socio tiene derecho a recibir anualmente los números de /berus, Reseñas Malacológicas y Noticiarios que 
se publiquen. 


ITHSONIAN INSTITUTION LIBRARI 


CC, 


3 9088 01663 890 


Iberus 29 (2) 2011 


RUBIO F., FÉERNÁNDEZ- GARCÉS, R. and ROLÁN E. The family Tornidae (Gastropoda, Rissooidea) 
in the Caribbean and neighboring areas. 


La familia Tornidae (Gastropoda, Rissovidea) en el Caribe y áreas VeciMAs. ocn. 1-240 


ISSN 0212-3010 


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