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Vol. 26 (1)

REVISTA DE LA

SOCIEDAD ESPAÑOLA

DE MALACOLOGÍA

Oviedo, junio 2008

Iberas

Revista de la

Sociedad Española de Malacología

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Editor di Publicaciones (Editor-in-Chief)

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Iberus

REVISTA DE LA

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DE MALACOLOGÍA

Vol. 26 (1)

Oviedo, junio 2008

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BIOSIS.

Dep. Leg. B-43072-81

ISSN 0212-3010

Diseño y maquetación: Gonzalo Rodríguez

Impresión: LOREDO, S. L. - Gijón

© Sociedad Española de Malacología Iberas, 26 (1); 1-4, 2008

"(w

Re-discovery of living specimens of Heliacus ( Gyriscus ) jef-

freysianus (Tiberi, 1867) (Gastropoda: Architectonicidae)

Redescubrimiento de ejemplares vivos de Heliacus ( Gyriscus ) jef-

Jreysian i/s (Ti be ri , 1867) (Gastropoda: Architectonicidae)

Constantine MIFSUD* and Panayotis OVALIS**

Recibido el 17-VII-2007. Aceptado 9-X-2007

ABSTRACT

Heliacus (Gyriscus) ¡effreysianus (Tiberi, 1 867), a rare architectonicid species associated

with Corallium rubrum (L., 1758), has been rediscovered alive off Crete, eastern Mediter-

ranean.

RESUMEN

Heliacus ( Gyriscus ) ¡effreysianus (Tiberi, 1 867), una especia rara de architectonicido aso-

ciado a Corallium rubrum (Linnaeus, 1 758), ha sido redescubierta con ejemplares vivos,

frente a la isla de Creta en el Mediterráneo oriental.

KEY WORDS: Mollusca, Gastropoda, Architectonicidae, Heliacus j effreysianus, Crete.

PALABRAS CLAVE: Mollusca, Gastropoda, Architectonicidae, Heliacus jeffreysianus, Creta.

INTRODUCTION

Heliacus ( Gyriscus ) jeffreysianus was

described by Tiberi (1867) from three

specimens collected from a red coral

substratum off Sardinia. He described

the species a year after its discovery, as

stated by Coen (1932).

SYSTEMATICS

Original description: Cochlea turbinata,

elato-conica, turrita, modice umbilicata, inteo-

fulvescens; apex obtusiusculus, laevigatus,

vértice intorto, subperf orato, spiraliter invo-

luto; anfr. 7 convexi, sutura profunda divisim

transverse cingulati, cingulis confortis, alter-

natim majoribus (num.fere 15 totidemque

minaribus in ultima anfractu; num. 6 in

penúltimo), eleganter granulosis, submonili-

formibus; anfr. Ultimus rotundatus, subin-

flatus, basi paululum depressus; umbilicus

mediocris, pervius, superne circolariter crenu-

latus; apert. Subcircularis, effusa, intus haud

margaritacea, marginibus acutis, callo pari-

etali junctis; margine columellari sinuato,

reflexo, umbilici partem occultante. - Diam.

Maj. 9, min. 8, alt 10? mili. - Operculum

corneum, superne núcleo centrali depresso

lamellaque erecto-crenulata multispiratum,

inferné processu centrali styliformi paucispi-

rato praeditum, limbo peripherico incrassa-

tum- Animal hucusque incognitum.

Hab. Infundís coralligenis maris Sar-

diniam meridionalem ambientis.

* 5, Triq ir-Rghajja, Rabat RBT 2486, Malta

** Agisilaou 37-39, Tzitzifies/Kallithea, 17674 Athens, Greece

1

Iberus , 26 (1), 2008

Figure 1. Heliacus (Gyriscus) jeffreysianus (Tiberi, 1867), off Crete Island on Corallium rubrum

(Linnaeus, 1758) in 120m. Size: 9x8 mm. Photo P. Ovalis.

Figura 1. Heliacus (Gyriscus) jeffreysianus (Tiberi, 1867), frente a la isla de Creta sobre Corallium

rubrum (Linnaeus, 1758) en 120m. Size: 9x8 mm. Foto P. Ovalis.

Tiberi dedicated his species to the

illustrious malacologist John Gwyn Jef-

freys (1809-1885). Two of the specimens

are presently in the Coen collection at

The Hebrew University, (Department of

Zoology) Jerusalem, Israel and the other

specimen is in the Jeffreys collection at

the Smithsonian Institution, Washington

(USA) (Melone and Taviani 1984).

New material: Two live specimens of

Heliacus ( Gyriscus ) jeffreysianus were

recently recovered from a living colony

of Corallium rubrum (Linnaeus, 1758)

from off the island of Crete at a depth of

120 m. Figure 1 shows one of the two

similar specimens, measuring h= 9 mm

x w= 8 mm which is about the same size

as those found by Tiberi. The specimens

were hand picked from a coral colony

by a professional scuba diver from Crete

using specialized diving equipment. It is

not known whether the molluscs were

actually feeding on the coral polyps.

However, the species seems to have a

cióse connection with C. rubrum. Several

species from the family Architectonici-

dae are known to be parasitic on coráis

(Bieler 1993). Judging from the two

presently known records, the species

has always been found with these red

coráis, and as it is well known, it was

never found in any other Mediterranean

habitat notwithstanding the large

amount of research carried out in the

last hundred years.

Since its original description, the

species had never been found again

until recently, when living specimens of

Heliacus jeffreysianus were collected in

the Adriatic Sea (Stanic and Schiapar-

elli, 2007). It is very curious, but not

surprising, that both this and our dis-

coveries were recorded at the same time

and at distantly separated places. This is

certainly due to the more recent interest

being undertaken both by amateur and

professional researchers in the depths of

the red and white coral substratum of

the Mediterranean Sea. The authors are

also aware of other unrecorded species

living in this part of the Mediterranean,

and originally described from deep

water in the Atlantic.

Otherwise, Heliacus jeffreysianus has

only been cited in the literature in a few

species lists and catalogues pertaining

to the Mediterranean malacofauna and

it was even thought to be extinct. More-

over, no fossil specimens of the species

have ever come to light (Melone and

Taviani 1984).

Monteros ato (1880) had com-

mented about the species of Tiberi "II G.

jeffreysianus, é una delle nostre piú rare

gemme, la cui scoperta, come di tante

altre raritá coralligene, si deve al Dr.

Tiberi. Soltanto tre esemplari ne sono

conosciuti sinora. Beati possidentes! lo

voglio ammettere che nella mente di un

naturalista il Gyriscus abbia piú valore

2

Mifsud AND OvaliS: Re-discovery of living specimens of Heliacus (Gyriscus) jeffreysianus

Figure 2. Map of the Mediterranean showing the three locadons cited for H. jeffreysianus. 1: Sardi-

nia Island; 2: Adriatic Sea; 3: Crete Island.

Figura 2. Mapa del Mediterráneo mostrando las tres localidades citadas para H. jeffreysianus. 1: isla

de Cerdeña; 2: mar Adriático; 3: isla de Creta.

del piú grosso diamente della corona

d'Inghilterra, ma francamente il prezzo

al quale lo vende il Dr. Tiberi, per una

conchiglia cosí piccola, sará riguardato

tanto dai naturalisti quanto dai mercanti

come assai esagerato"

In transía tion: "The G. jeffreysianus is

one of our most rare gems, its discovery,

like many other coralligeneous rarities,

is owed to Dr. Tiberi. Only thee speci-

mens are known up to now. Blessed are

the possessors\ I must admit that in the

minds of the naturalist the Gyriscus has

more valué than the largest diamond

from the British crown, but frankly the

price which Dr. Tiberi has asked, for

such a small shell is regarded by natu-

ralists and also by merchants as being

exaggerated."

H. jeffreysianus is one of the most

beautiful species from the family Archi-

tectonicidae. Its sculpture of small-

beaded spiral chords and its trochoid

shape make it outstanding and very dis-

tinguished among the other species of

the family in the Mediterranean. In fact,

Tiberi (1867) created a genus for it,

Gyriscus, which is nowadays a subgenus

of Heliacus. A very similar Pacific species

is H. ( Gyriscus ) asteleformis (Powell,

1965) from New Zealand.

Coen (1932) figured the operculum

from one of the two specimens in his

collection (also figured in Melone and

Taviani, 1984). Melone and Taviani

(1984) described and figured a syntype

from the Coen collection. They also

figured the radula (as redrawn after

Merril, 1970). Other descriptions and

diagnoses of the genus and the species

can also be found in Coen (1932),

Melone and Taviani (1984) and in

Bieler (1993).

CONCLUSIONS

This very long elapsed period of

o ver 140 years for the re-discovery for

H. jeffreysianus is in all probability due

to its rarity and the very particularly

restricted and difficult to sample

habitat. Red coral was traditionally har-

vested by fishermen utilizing a particu-

lar gear called the Cross, which consists

of two large beams tied together in the

form of a cross and armed with dan-

gling ropes and pieces of netting to

enable the coral to entangle during

dredging. This heavy equipment,

although it seemed adequate for the

purpose, besides being very destructive

to the substratum, constantly shakes the

entangled coráis and the molluscs seem

to always fall off before the gear is

brought aboard the vessel. The modern

method of manual harvesting through

SCUBA is more selective, and therefore

much less destructive. However, it is

much more risky and dangerous.

Curiously, this species had not been

discovered on red coral before, although

3

Iberus , 26 (1), 2008

the method of manual harvesting of coral

by specialized deep diving equipment

(using a mixture of gases) has now been

in practice for many years, especially by

Spanish, Italian and Greek coral divers.

More probably, it may take the keen eyes

of a biologist, a naturalist or a shell col-

lector to notice and pick out the small

mollusc shells in situ. Therefore any spec-

imens brought up by chance by the fish-

ermen or the divers are probably either

thrown back into the sea with the other

rubbish as in the case of the fishermen or

in the case of the diver, they fall off the

coral unnoticed during the long decom-

pression process. Moreover, the diver

would be even more concerned for his

"treasure" rather than a few "worthless"

shells. Although Mediterranean Corallium

rubrum is also found at great depths in the

Mediterranean, the two existing records

of H. jeffreysianus are both from shallow

BIBLIOGRAPHY

Bieler, R., 1993. Architectonicidae ofthe Indo-Pa-

cific. Gustav Fischer Verlag, pp. 377. Stuttgart.

Coen, G. S., 1932. Sul genere Gyriscus, Tiberi,

1867. Bollettino della Societd Veneziana di

Scienze Naturali, 1 (1): 9-13.

Melone, G. and Taviani, M., 1984. Revisione

delle Architectonicidae del Mediterráneo.

Atti del Simposio: Sistemática dei Prosobranchi

del Mediterráneo, Bologna 24-26 Setiembre 1992,

(B. Sabelli, ed.) Lavori S.I.M., 21: 149-192.

Merrill, A. S., 1970. The family Architectonici-

dae (Gastropoda: Mollusca) in the western and

eastern Atlantic. Unpubl. Ph.D. Thesis, Univ.

Delaware (Univ. Microfilms Int. Inc., Ann

Arbor, Michigan, U.S.A., Nr. 7144444).

water living colonies. Stanic and Schia-

parelli (2007) did not mention the species

on which they found their specimens.

Finally, this record extends the species dis-

tribution to quite a larger area of the

Mediterranean (Fig. 2) and it is expected

that more specimens which could con-

tribute to the study of the biology of the

species are likely to turn up.

ACKNOWLEDGEMENTS

We would like to thank Agios Nico-

laos, a specialized deep sea coral diver

for donating one of the two specimens

which he collected to one of the authors

(P. O.) for this study. Thanks are also

due to Sophie Valtat (Belgium) for pro-

viding important literature and to S.

Gofas (Spain) for revising and enhanc-

ing the manuscript.

Monterosato, T. A., 1880, Nota sopra alcune

conchiglie coralligene del Mediterráneo. Bol-

lettino della Societd Malacologica Italiana, 6:

243-259.

Stanic, R. and Schiaparelli, S., 2007. New

finding of living specimens of the rare ar-

chitectonicid Heliacus (Gyriscus) jeffreysianus

(Tiberi, 1867). Bollettino Malacologico, 43 (9-12):

143-146.

Tiberi, N., 1867. Diagnose du nouveau genre

méditerranéen Gyriscus. Journal de Conchy-

liology, 15: 303. Paris.

4

© Sociedad Española de Malacología

Notes on West African Perrona (Gastropoda: Clavatulidae),

with the description of a new species

Notas sobre Perrona (Gastropoda: Clavatulidae) en Africa

Occidental, con la descripción de una nueva especie

Emilio ROLÁN*, Peter RYALL** and Juan HORRO***

Recibido el 31 -VI 1-2007. Aceptado el 19-X-2007

ABSTRACT

The type species of the genus Perrona is poorly known and given an erroneous type local-

ity. One of the purposes of this paper is to clarify its status and correct spelling as well as

illustrating the species and defining its distribution area. A new species from Angola

belonging to this genus is described. The shell, protoconch, operculum and radula are

illustrated, and the reasons for assigning this new species to this genus are discussed. All

known species of Perrona are figured.

RESUMEN

La especie tipo del género Perrona es poco conocida y la localidad tipo dada es errónea.

El objetivo de este trabajo es clarificar su situación, su correcto nombre, además de repre-

sentar la especie y clarificar su área de distribución. Se describe una nueva especie de

este género de Angola. Se muestran concha, protoconcha, opérculo y rádula de la nueva

especie y se discuten las razones para asignarla a este género y no a otros con similar

morfología. Se ilustran todas las especies conocidas de este género.

KEY WORDS: Clavatulidae, Perrona, West Africa, Angola, new species.

PALABRAS CLAVE: Clavatulidae, Perrona, África occidental, Angola, nueva especie.

INTRODUCTION

Classical malacological works like

Bruguiére (1792), Lamarck (1801, 1816

and 1822), Kiener (1840) and Reeve

(1843, 1845) and publications dealing

with Turridae (s.l.) such as Récluz

(1851), Petit de la Saussaye (1851), von

Maltzan (1883, 1884), Sykes (1905),

Dautzenberg (1912), Strebel (1912,

1914) have described numerous West

African species assigned to the family

Clavatulidae Gray, 1853.

Their generic placements and syn-

onymies are still not clear. Some modem

works like Knudsen (1952, 1956),

Ardovini (2004), Boyer and Hernán-

dez (2004), Nolf (2006), Nolf and Ver-

straeten (2006) and Boyer and Ryall

(2006) have described new species or

revised others, placing most of them in

the genus Clavatula.

Among our material from Angola,

dredged over many years by Xico Fer-

* Museo de Historia Natural, Campus Universitario Sur, E 15782 Santiago de Compostela, Spain

** St. Ulrich 16, A-9161 Maria Rain, Austria

*** Montero Ríos, 30-3° 36201 Vigo, Spain

5

Iberas, 26 (1), 2008

nandes, a new small unidentified

species was found. Its description is one

of the subjects of this work. The generic

attribution to the genus Perrona (Schu-

macher, 1817) requires a review of the

type species because it is a rather poorly

known species. This paper therefore

also comments on the type species.

The identity of Perrona perron

Martini and Chemnitz (1788: 278,

pl. 164, figs. 1573, 1574) illustrated a

shell they named "Der Perron ". Later

Gmelin (1791), referring to Chemnitz's

figures, described it as Murex perron and

indicated "in Océano australi" as its

geographic origin. This type locality is

obviously wrong. These figures, or the

shell they represent, can be considered

to be the lectotype. Its present where-

abouts, as noted below, are unknown.

When Lamarck (1816) described

Pleurotoma spirata he also made refer-

ence to the same illustrations, but sepa-

rated his species from them by stating

"La figure citée de Chemnitz offre sur la

base du dernier tour ; des sillons dont notre

coquille est absolutement dépourvue".

Schumacher (1817) also referred to

the same illustrations of Chemnitz when

he described the genus Perrona and nomi-

nated Perrona tritonium as the type of this

genus. P. tritonium is therefore a júnior

objective synonym of Perrona perron

(Gmelin). Again no locality was given.

The species was subsequently

described for a third time by Reeve (1843)

who referred once more to Chemnitz's

illustrations. He gave it the ñame Pleuro-

toma perronii, drawing his own figure

from his own shell. He likewise did not

State a locality or habitat. Reeve cited

Lamarck 's spirata as a different species,

writing "1 have a shell before me ofvery dif-

ferent character (Pleurotoma Perronii,

nobis), answering exactly to the figure in

question". Reeve (1843) did not cite

Gmelin and therefore appeared to con-

sider his ñame as a valid new species.

However it is clearly a júnior synonym of

Gmelin's species and it is concluded that

this was an omission in Reeve's work.

Marrat (1877) was the first to give

an indication of the true range of the

species when he included it in a list of

North-west African shells which he

stated carne "from Madeira to the Gulf

of Guinea".

Tryon (1884), although stating his

Opinión that this species was an inter-

medíate form between lineata and

spirata , mentioned it as Clavatula (sub-

genus Perrona ) perron Chemnitz and

provided a detailed description.

Strebel (1912), in his revisión of

genus Perrona, treated as clearly differ-

ent species both spirata and perron. He

was able to examine Chemnitz's speci-

men from which the famous drawings

were made and reillustrated it. He

referred immature specimens of the

species to Reeve's description, calling

them perron var. reevei. He also correctly

indicated part of the correct habitat

range of the species, citing specimens he

had examined from Pointe Noire,

Congo to Ambriz, northem Angola.

Knudsen (1952) employed the ñame

"Clavatula perronii (Reeve)" noting that

the "Atlantide expedition" could not

obtain any additional material and that

Chemnitz's type could no longer be

traced at the ZMUC.

Powell (1966), who referred cor-

rectly to the type species of Perrona, did

not figure Perrona perron but illustrated

Perrona spirata (Lamarck, 1816).

More recently, Cernohorsky (1974)

figured a syntype from Chemnitz (in

ZMUC), commenting that "the species

most probably Uves in West Africa but this

locality has been not substantiated by either

literature records or authentic specimens.

Another possibility is that Murex perron is

conspecific with Pleurotoma spirata

Lamarck, 1816, as suggested by Nordsieck

(1968: 154) although the two species appear

quite distinct".

Finally, Bernard (1984, fig. 182)

figured under the erroneous ñame

"Clavatula kraepelini" three specimens of

this species. He unknowingly extended

Strebel's habitat range northwards,

quoting the localities of both Cape Este-

rias and Mayumba in Gabon, and indi-

cating a depth of -20 to -50 meters.

After examining Reeve's type speci-

men, the available literature and some

ó

RoláN ET AL.: Notes on West African Perrona with the description of a new species

shells in the collection of one of the

authors (PR), it was decided to record

some additional information about this

poorly known group.

Abbreviations:

AMNH American Museum of Natural

History, New York

ANSP Academy of Natural Sciences,

Philadelphia

BMNH The Natural History Museum,

London

MCZ Museum of Comparative Zoology,

Harvard university

MHNS Museo de Historia Natural "Luis

Iglesias" Universidad, Santiago de

Compostela

MNCN Museo Nacional de Ciencias

Naturales, Madrid

MNHN Muséum national d'Histoire

naturelle, París

ZMUC Zoologisk Museum, Copenhagen

USNM National Museum of Natural

History, Washington

ZSM Zoological Staatmuenchen

Museum, Muenchen

CJH collection of Juan Horro, Vigo

CPR collection of Peter Ryall, Maria

Rain

SYSTEM ATIC PART

Family Clavatulidae Gray, 1853

Genus Perrona Schumacher, 1817

Type species: Perrona tritonium Schumacher, 1817 = Murex perron Gmelin, 1791 = Pleurotoma

perronii Reeve, 1843.

Perrona perron (Gmelin, 1791) (Figs. 1-13)

Der Perron Martini and Chemnitz, 1788: 278, pl. 164. figs. 1573, 1574 (Fig. 1).

Murex perron Gmelin, 1971: 3559.

Perrona tritonium Schumacher, 1817: 218.

Pleurotoma perronii Reeve, 1843: pl. 11, sp. 94 (Fig. 2).

Type material: Figured syntype of Der Perron, whereabouts presently unknown; this specimen is

reported as examined by Strebel (1912) but could not be traced by Knudsen (1952) who never-

theless mentions another shell labelled in the handwriting of Chemnitz. Pleurotoma perronii: holo-

type BMNH 1900.2.8.26.

Other material examined: 8 shells (Figs. 4-10), Pointe Noire, Congo (CPR).

Type locality: Erroneous for Der Perron (in Océano australi). Not mentioned in the original descrip-

tion of any of the other taxa. Here designated at Pointe Noire, Congo.

Description: Shell (Figs. 3-12) see the

original descriptions as well as Tryon

(1884) and Strebel (1912). The follow-

ing data can be added: the protoconch

(Fig. 13) consists of about 2 1 ¡i shiny,

smooth whorls, honey brown in colour;

the first whorl slightly darker and of 750

pm diameter. From the protoconch

whorls a series of opisthocline axial ribs

start to form, first as thin riblets extend-

ing from the top to the bottom of the

whorl, becoming stronger and shaped

as an inverted C in the second teleo-

conch whorl where their bases become

more globose. Between them small

striae can be seen. In the subsequent

whorl they detach themselves entirely

from the upper subsutural cord which

also increases in prominence. In the

fourth teleoconch whorl these vestiges

of the axial ribs become closer and

closer to the lower suture, declining in

prominence until they entirely disap-

pear. The following whorls are smooth

with an irregular, raised subsutural

collar and the whorls are traversed by s-

shaped striae. At the same time as the

axial riblets form in the third whorl.

7

Iberus , 26 (1), 2008

about 9 radial striae also start to emerge

becoming slightly stronger in the fol-

lowing whorls but then decreasing in

intensity in the fourth or fifth whorls

and thereafter are hardly noticeable.

The shell is a light honey-brown

throughout. On the fourth whorl the

subsutural colar starts to become lighter,

being entirely white in the fifth whorl.

In subsequent whorls it can be interrup-

ted by patches of light brown; in the

final whorl the two major cords are

white interspaced with small brown

patches the same colour as the whorl

itself and the cords on the columnella

are also white.

Size : the shell can reach up to 37.2

mm (Bernard, 1984).

Remarks : The comparison of juvenile

specimens of P. perron with species of

the genus Clavatula (type species: Clavat-

ula coronata Lamarck, 1801) show much

similarity in the aspect of the opistho-

cline axial ribs, which tend to disappear

with maturity in the subsequent whorls.

For this reason it is concluded that there

is a stronger relationship between these

two genera, than with other genera

within this family.

Whilst some authors synonymized P.

perron with P spirata Lamarck, 1816 it

must be noted that not only are they

morphologically different, but the latter

inhabits an area further to the south,

being restricted to the Luanda area

where it has been found at -5/50 meters.

Distribution : This species inhabits the

West African infralittoral coast from Cap

Esterias (Gabon) to Ambriz (northern

Angola).

Special remarks: The type locality is

always related with a holotype or a lec-

totype. In the present case, the original

figures are sufficient to determine the

identity of the species and it is not nec-

essary to designate a neotype; Pleuro-

toma perronii was described lacking type

locality as confirmed by the label of this

holotype in BMNH. If the type locality

of the first description is erroneous, and

no other is mentioned in the description

of the synonymized taxa, it is necessary

to give more detailed information.

Pointe Noire (Congo) is designated as

the type locality being in the center of

the known distribution area, as well as a

locality mentioned by Strebel (1912)

and the source of our specimens. Our

material agrees very well with the

figures and the known holotype.

Concerning the correct spelling for

the ñame of this species, it must be

perron, which is the first ñame employed

by Gmelin (1791) and which is appa-

rently a ñame not dedicated to a person.

Tryon (1884) stated that Chemnitz

employed the common Dutch ñame

"perron" (= flight of stairs, in reference

to the profile) and that Reeve (1843),

misinterpriting Chemnitz's ñame to be a

personal ñame, Latinized it to read

"perronii”.

Perrona micro spec. nov. (Figs. 14-19, 21, 23, 24-28)

Material examined: Holotype (Figs. 14, 15), in MNCN (15.05/47050). Paratypes in the following

collections: MNHN (1, Fig. 16); BMNH (1, Fig. 17); AMNH (1); USNM (1); ZSM (1); ANSP (1); MCZ

(1); CPR (11); CJH (6); MHNS (16, Fig. 18).

Type locality: Palmeirinhas, south of Luanda, Angola, between 3 and 20 meters depth.

Etymology: The specific ñame refers to the small size of the species.

Description: Shell (Figs. 14-18) small

for the genus, very solid, lanceolate.

Protoconch (Figs. 21, 23, 24-26) with a

little more than one whorl, of 530 pm in

diameter and white in colour. The teleo-

conch begins with three grooves that

sepárate 4 cords, later increasing to four

and then five spiral grooves; there are

six in the subsequent whorl; the two

cords immediately below the suture as

well as the lowest cord are more promi-

nent. This gives the middle part of the

whorl a concave appearance whilst the

upper and lower parts protrude before

their separation at the suture. There is

no axial sculpture except for growth

8

RoláN ET AL.: Notes on West African Perrona with the description of a new species

Figures 1-13. Perrona perron. 1: figure from MARTINI AND CHEMNITZ (1788); 2: figure of the

holotype of Pleurotoma perronii from Reeve (1843); 3: holotype (see below Figs. 11, 12) of Pleuro-

toma perronii (BMNH) at the same comparative size; 4-6: shell (27.6 mm), Pointe Noire, Congo

(CPR); 7: shell, 28.6 mm, Pointe Noire (CPR); 8, 9: juveniles, 18.6 and 14.2 mm, Pointe Noire

(CPR); 10: shell, 27.2 mm, Pointe Noire (CPR); 11, 12: holotype of P. perronii , 25 mm (BMNH);

13: spire and protoconch, Pointe Noire (CPR).

Figuras 1-13. Perrona perron. 1: figura de MARTINI Y CHEMNITZ ( 1 788); 2: figura del holotipo de Pleurotoma

perronii, de Reeve ( 1 843); 3: holotype ( véase abajo Figs. 1 1, 12) de Pleurotoma perronii (BMNH) con el mismo

tamaño relativo; 4-6: concha, 27,6 mm, Pointe Noire, Congo (CPR); 7: concha, 28,6 mm, Pointe Noire (CPR);

8, 9: juveniles, 18,6 y 14,2 mm, Pointe Noire (CPR); 10: concha, 27,2 mm, Pointe Noire (CPR); 11, 12: holotipo

de P. perronii, 25 mm (BMNH); 13: spire and protoconch, Pointe Noire (CPR).

9

Iberus, 26 (1), 2008

Figures 14-19. Perrona micro spec. nov. 14, 15: holotype, 10.3 mm (MNCN); 16: paratype, 10.6

mm (MNHN); 17: paratype, 8.5 mm (BMNH); 18: paratype, 7.6 mm (scanning electrón micro-

scope) (MHNS); 19: operculum (SEM). Figures 20-23. Comparison of the spire and protoconch

of Clavatula quinteni (Figs. 20, 22) and Perrona micro (Figs. 21, 23).

Figuras 14-19. Perrona micro spec. nov. 14, 15: holotipo, 10,3 mm (MNCN); 16: paratipo, 10,6 mm

(MNHN); 17: paratipo, 8,5 mm (BMNH); 18: paratipo, 7,6 mm (microscopio electrónico de barrido)

(MHNS); 19: opérenlo ( MEB). Figuras 20-23. Comparación de la espira y protoconcha de Clavatula

quinteni (Figs. 20, 22) y Perrona micro (Figs. 21, 23).

10

ROLÁN ETAL.: Notes on West African Perrona with the description of a new species

Figures 24-27. Perrona micro. 24: detail of the spire; 25, 26: detail of the protoconch; 27: radular

tooth; 28: radula. Figures 29-33. Perrona obesa. 29: holotype, 34.0 mm (BMNH); 30: shell, 31.2

mm, 10 m, Namibe, Angola (CER); 31: shell, 35.7 mm, Namibe, Angola (CPR); 32, 33: shells,

40.2, 41.6 mm, Sacomar, Namibe, Angola (CPR).

Figuras 24-27. Perrona micro. 24: detalle de la espira; 25, 26: detalle de la protoconcha; 27: dientes de

la rádula; 28: rádula. Figuras 29-33. Perrona obesa. 29: holotipo, 34. 0 mm (BMNH); 30: concha,

31,2 mm, 10 m, Namibe, Angola (CER); 31: concha, 35,7 mm, Namibe, Angola (CPR); 32, 33:

conchas, 40.2, 41.6 mm, Sacomar, Namibe, Angola (CPR).

Iberus, 26 (1), 2008

lines which form an U-shape in the

concave central portion; this is reversed

on both the upper and lower protruding

parts forming an overall S-shape on the

whorl. There are about 6 whorls on the

teleoconch, increasing gradually in

width and height. The last whorl

becomes elongated towards the base,

the siphonal canal is widely open and

almost similar in length to the lowest

part of outer lip. The aperture is ovoid,

with an U-shape sinus in the upper part,

a sharp external lip, and a small callus

on the columella. There are 20 spiral

grooves from the periphery to the base.

The colour is typically light brown or

orangish, with white and dark blotches

alternating on the subsutural cords, but

in a few specimens the colour is darker

and the white pattern is absent.

Soft parts: Study of a retracted speci-

men preserved in alcohol reveáis a light

coloured animal; the operculum (Fig.

19) is ovoid with the nucleus centrally

placed on the interior. The radula (Fig.

28) is formed by two rows of about 70

elongate and sharply pointed marginal

teeth (Fig. 27) at the center of which is a

very small central tooth reduced to a

cusp.

Dimensions : Flolotype is 10.3 mm.

The largest paratype is 12 mm.

Distribution: Only known from the

type locality, where it is probably

endemic.

Discussion: There is no known

species from West Africa with which it

can be confused. Only juvenile forms of

Clavatula quinteni Nolf and Verstraeten,

2006, which are much larger when

adult, are similar in profile and coloura-

tion; they both lack axial sculpture (see

comparison in Figs. 20 and 21, 22 and

23) but C. quinteni has a much more

elongated siphonal canal, a larger proto-

conch (more than double the diameter)

and lacks spiral grooves.

There was some hesitation about the

generic assignment of this new species.

Comparison with morphologically

similar species in some genera was

made, in particular Microdrillia Casey,

1903; Suavodrillia Dalí, 1918; Parecuneus

Laseron, 1954, and Maoritomella Powell,

1942, Tomopleura Casey, 1904, and Viridi-

turris Powell, 1964. In all cases opercu-

lum and radula do not agree.

Perrona micro spec. nov., in spite of

its small size, belongs to Clavatulidae.

This is confirmed by the operculum

with a medium-lateral nucleus and the

wishbone-like marginal teeth with a

vestigial rachidian one in the middle.

Powell (1966) reviewed the Turri-

dae (s. 1.) and defined the subfamily

Clavatulinae H. and A. Adams, 1858

with four recent and two fossil genera;

the recent genera being Clionella, Clavat-

ula, Perrona (with subgenus Tomellana)

and Pusionella. Kilburn (1985), who

dealt only with South African species,

noted 4 genera; Toxiclionella, Bentho-

clionella , Clavatula and Clionella. Tucker

(2004) considers 492 fossil and 97 recent

species in this group. Later Bouchet

and Rocroi (2005) elevated Clavatuli-

dae to family status.

The genus Clavatula Lamarck, 1801 is

relatively abundant along the West

Africa coast. Powell (1966) mentions 11

species. Most of them, including the

type species ( Clavatula coronata Lamarck,

1801) are of large size. Other species

recently illustrated and/or described, as

detailed in our introduction, are also rel-

atively large. Although Boyer and

Ryall (2006) noted that Knudsen (1952,

1956) classified other genera of Clavatul-

idae under the genus Clavatula, subse-

quent authors seem to ignore these

genera and their paramaters as estab-

lished by Kilburn (1985). A revisión

could conclude that some of them

belong to other genera.

The genus Clavatula is described by

Powell (1966): "shell modera tely large

to large sized, 15 - 55 mm, very solid,

buccinoid, coarsely axially and spirally

sculptured; with a tall, often coronated

spire, and a trúncate body-whorl, termi-

nated in a relatively short reflected and

deeply notched anterior canal.... Oper-

culum ovate-lunate with a medio-lateral

nucleus. Radula consisting of a pair of

stout narrowly pointed marginal teeth

of modified wishbone type, the proxi-

mal extremity sepárate and superim-

posed upon the larger element. A small

12

RoláN ET AL. : Notes on West African Perrona with the description of a new species

Figures 34-37. Perrona spirata. 34-36: shell covered by organic material, 31.5 mm, Luanda, Angola

(CPR); 37: operculum. Figures 38-41. Perrona subspirata. 38: shell, 31.5 mm; 39-41: shell, 32.0

mm, Namibe, Angola (CPR); 42: operculum.

Figuras 34-37. Perrona spirata. 34-36: concha recubierta por material orgánico, 31,5 mm, Luanda,

Angola (CPR); 37: opérculo. Figuras 38-41. Perrona subspirata. 38: concha, 31,5 mm; 39-41: concha,

32,0 mm, Namibe, Angola (CPR); 42: opérculo.

13

Iberus, 26 (1), 2008

but we 11 formed unicuspid central tooth

is present also." According to Kilburn

(1985) the main features of Clavatula are:

deep anal sinus, realtively long siphonal

canal, distict parietal tubercle and proto-

conch that is bluntly domed with about

2 1/2 whorls.

The species P. micro spec. nov. has

some shell characters of Clavatula but

has no axial sculpture and is smaller

than all of them. Clavatula cossignanii

Ardovini, 2004, the smallest Clavatuli-

dae known in West Africa, is different in

most features including the fewer

number of protoconch whorls and is

endemic to Senegal. The holotype is 13.2

mm but it can reach a size of 15.9 mm

(CPR).

According to Powell (1966) the

genus Clionella Gray, 1847 differs from

Pusionella Lamarck, 1801 by its clavi-

form shape with tall, flat-sided spire

whorls; simple long flexuous axials ribs

without either subsutural or peripheral

processes and a truncated body-whorl.

Kilburn (1985) defines Clionella with an

anal sinus; a slight notch; siphonal canal

short with no distinct parietal tubercle;

protoconch somewhat conical, of about

2 whorls, the first one rounded and

tilted. The genus is usually restricted to

South Africa. P. micro spec. nov. lacks

axial sculpture, its sinus is deep and

does not have a truncated body-whorl:

therefore it does not agree with Clionella.

The new species was also ruled out

of the genus Pusionella Gray 1847

because many specific characters are

absent in P. micro ; the flat-sided whorls;

the outer lip bordered by a broad, sub-

sutural, very slight sinus and the

smooth and polished surface.

Finally, the genus Perrona Schu-

macher, 1817 is described by Powell

(1966) as: "shell moderately large, 25 -

40 mm, rather narrowly fusiform, with a

tall spire of rapidly increasing whorls

ACKNOWLEDGEMENTS

The authors wish to thank the Euro-

pean Synthesys Program which allowed

one of the authors (ER) to examine the

and a narrow body-whorl... surface

smooth, or nearly so... with a conspicu-

ous narrowly carinate subsutural

collar.... operculum as in Clavatula...

radula with a pair of elongate margináis

and a narrow-based unicusp central

tooth."

The known West African species of

Perrona, all figured in the plates in the

present work, are the following: P.

perron (Gmelin, 1791) (Figs. 1-13), P.

spirata (Lamarck, 1816) (Figs. 34-37),

both already mentioned, P. obesa

(Reeve, 1843) (Figs. 29-33) and P subspi-

rata (von Martens, 1903) (Figs. 38-41),

both endemic to South Angola. Other

authors have placed other species in

this genus, for example Perrona nifat

(Bruguiére, 1789) by Abbott and

Dance (1986), but this generic place-

ment is erroneous in our opinión.

Powell (1966) also mentions another

West African species belonging to this

genus but placed in the subgenus

Tomellana Wenz, 1843: Perrona ( Tomel -

lana) lineata Lamarck, 1816; but this

shell has a short, concave-outlined spire

and a deep slit-like anal sinus and does

not resemble our species. Strebel

(1912) referred other species to Tomel-

lana; they also do not correspond with

the species currently reviewed.

Perrona micro spec. nov. is more

similar to Perrona perron than to any

other type species and the new species

is placed in this genus as being the

closest. Also noticed is a similarity in the

colouration of the subsutural cord as

well as the S-shaped axial striae. We

find this at present the better solution

rather than to create a new genus.

Whilst the similarity noted with

Clavatula quinteni should not create con-

fusión, a result of this paper must also

be to assign this species to the genus

Perrona although this has not been the

reason for this work.

type material of Pleurotoma perronii and

other types; Jesús Méndez of the Centro

de Apoyo Científico y Tecnológico a la

14

ROLÁN ET AL. : Notes on West African Perrona with the description of a new species

Investigación (CACTI) of the University

of Vigo for the SEM photographs; Jesús

S. Troncoso for allowing us to use the

Photographic apparatus of the Departa-

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Iberus, 26 (1): 17-28, 2008

© Sociedad Española de Malacología

Pattern of reserve storage of the two mussel species Perna

perna and Mytilus galloprovincialis living on Moroccan

coasts: annual variation and effect of pollution

Patrones de almacenamiento de reservas en dos especies de mejillón

Perna perna y Mytilus galloprovincialis de las costas de Marruecos:

variación anual y efecto de la contaminación

Abdellatif MOUKRIM*1, Mohamed ID HALLA*, Abderazak KAAYA*,

Abdellah BOUHAIMI*, Soumaya BENOMAR* et Michel MATHIEU**

Recibido el 23-XI-2005. Aceptado el 30-X-2007

ABSTRACT

The pattern of reserve storage and ¡ts ¡mportance ¡n the reproductive cycle of the two mus-

sel species living in Moroccan coasts, Perna perna and Mytilus galloprovincialis, were

studied comparatively, during two annual cycles. Study was conducted at a polluted and

an unpolluted site in Agadir bay. Stereological analysis shows that P. perna presents only

one type of storage cells (vesicular cells) storing glycogen. In M. galloprovincialis, two

types of storage cells are present: vesicular cells and adipogranular cells (containing

glycogen, lipids and proteins). In both species, seasonal variations of reserve tissue vol-

ume are conversely proportional to those of the germinal tissues. Reserve tissue appeared

in spring (May) and increased in summer. It decreased quickly from August onwards, and

disappeared between January and April. In M. galloprovincialis, adipogranular cells dis-

appeared before vesicular cells and reappeared first. In the polluted site, seasonal varia-

tion of reserve and germinal tissues is comparable to that in unpolluted site but the máxi-

mum valué of reserve tissue percentage is less ¡mportant. Moreover, during the year, this

tissue did not disappear. Biochemical analysis shows that mean valúes of glycogen and

proteins quantities are significantly higher in P. perna than in M. galloprovincialis. How-

ever, total lipids quantity mean is higher in M. galloprovincialis. Seasonal variations of the

three biochemical parameters present a similar profile in the two species. Compared to

unpolluted site animáis, in polluted one, molluscs possess low glycogen and high lipids lev-

éis. Seasonal variations of these parameters show a perturbed profile.

RESUMEN

Se ha estudiado y comparado el patrón de almacenamiento de reservas y su importancia

en el ciclo reproductor en dos especies de mejillón de las costas de Marruecos, Perna

perna y Mytilus galloprovincialis, durante dos ciclos anuales. Este estudio se llevó a cabo

en un sitio contaminado y otro no contaminado de la bahía de Agadir. El análisis estere-

ológico demostró que P. perna presenta un solo tipo de células de reserva (células vesicu-

lares) almacenando glucógeno. En M. galloprovincialis, hay dos tipos de células de

* Laboratory Aquatic Ecosystems: Marine and Continental Field, Biology Department, Sciences Faculty, Ibn

Zohr University, BP 8106, 80000, Agadir, Morocco.

** Laboratory Marine Biology and Biotechnology, I. B. B. A., Caen University, France.

1 Corresponding author.

17

Iberus , 26 (1), 2008

reserva: células vesiculares y células adipogranulares (conteniendo glucógeno, lípidos y

proteínas). En ambas especies, las variaciones estacionales del volumen de tejido de

reserva son inversamente proporcionales a las del tejido germinal. El tejido de reservas

apareció en la primavera (mayo) y se incrementó en verano. A partir de agosto, dismin-

uyó rápidamente y desapareció entre enero y abril. En M. galloprovincialis, las células

adipogranulares desaparecieron antes que las células vesiculares y son las primeras en

aparecer en el siguiente ciclo. En el sitio contaminado, la variación estacional de tejidos

de reserva y germinales es similar a la del sitio no contaminado, salvo que el porcentaje

máximo de tejido de reserva es menor. Además, este tejido no desapareció a lo largo del

año. Los análisis bioquímicos muestran que las cantidades promedias de glucógeno y de

proteínas son significativamente más altas en P. perno que en M. galloprovincialis. Sin

embargo, el promedio de lipidos totales fue mayor en M. galloprovincialis. Las varia-

ciones estacionales de los tres parámetros presentan un perfil similar en las dos especies.

Comparados con los del sitio no contaminado, los animales del sitio contaminado poseen

niveles bajos en glucógeno y altos en lípidos. Las variaciones estacionales de estos

parámetros muestran un perfil perturbado.

KEY WORDS: Agadir, Morocco, mussels, Mytilus galloprovincialis, Perna perna, pollution effects, reproductive

cycle, reserves strategy.

PALABRAS CLAVE: Agadir, Marruecos, mejillones, Mytilus galloprovincialis, Perna perna , efectos de la conta-

minación, ciclo reproductor, estrategias de reserva.

INTRODUCTION

Two sympatric mussel species are

found along the Moroccan coast: the

African mussel Perna perna and the

Mediterranean mussel Mytilus gallo-

provincialis. The first species is located in

the north of its geographical distribu-

tion; the second one in its Southern

limit. In Agadir bay, these local popula-

tions of mussels have important com-

mercial valúes (Id Halla, 1997) and

environmental interest (Najimi,

Bouhaimi, Daubéze, Zekhnini, Pel-

LERIN, NARBONNE, AND MOUKRIM, 1997;

Kaaya, Najimi, Ribera, Narbonne, and

Moukrim, 1999; Moukrim, Kaaya,

Najimi, Roméo, Gnassia-Barelli, and

Narbonne, 2000) as sentinel species.

The few studies carried out on

mussel biology in this area, concern

only some aspects of the reproduction

cycle (Shafee, 1989; Id Halla,

Bouhaimi, Zekhnini, Narbonne,

Mathieu, and Moukrim, 1997). They

showed that P. perna and M. galloprovin-

cialis present synchronous reproductive

cycles, and breed throughout the year

with a principal spawning period in

spring. The only differences observed

by Id Halla et al. (1997), regard the

duration of the principal spawning

period (more important for P. perna ) and

the genital activity rhythm in summer

(reduced in this species).

In view of the importance of

reserves in the reproductive cycle

(Bayne, Bubel, Gabbott, Livingstone,

Lowe and Moore, 1982; Lowe, Moore

and Bayne, 1982; Pipe, 1987) and of the

particularities of reserve strategy in

Mytilidae (strategy essentially based on

the glycogen, presence of specific

storage cells and existence of a direct

relationship between reserve metabo-

lism and reproductive cycle), it was

interesting to conduct a comparative

study of P. perna and M. galloprovincialis

in order to describe the reserves strat-

egy for these two species living in the

same environmental conditions. The

two molluscs present an important dif-

ference regarding the cells involved in

the reserve storage. As indicated by

Lunetta (1969) P. perna presents only

one type of storage cells (vesicular cells

or glycogen cells), storing especially the

glycogen ; however, in M. galloprovin-

18

Moukrim ET AL.: Reserves strategy of mussels in Moroccan coasts

cialis (Herlin-Houtteville, 1974; Dan-

ton, Kiymoto, Komaru, Wada, Awaji

and Mathieu, 1996), two types of

storage cells are present, vesicular cells

(storing glycogen) and adipogranular

cells (containing glycogen, lipids and

proteins).

Furthermore, beyond the determina-

tion of pattern of reserve storage and its

importance in the reproductive cycle of

the two mussels living in Moroccan

coasts, this work studies the impact of

pollution on this pattern. Therefore, a

comparative study of two sites (unpol-

luted and polluted) was conducted.

MATERIAL AND METHODS

Sampling

This study was conducted during

two annual reproductive cycles

(October 1994 to August 1996). Two

types of sites, representative of the

Agadir marine bay were considered: i) a

reference site (unpolluted). Cap Ghir,

located 50 km north of Agadir City and

far from any human activity, and ii) a

polluted site (Anza) located 5 km north

of Agadir and receiving the industrial

and domestic untreated waste waters of

Anza zone. Many studies, conducted

during the same period of the present

study, showed that this site is contami-

nated by heavy metáis and PAHs either

accumulated by mussels or in sediment

(Id Halla, 1997; Najimi, 1997;

Moukrim, Kaaya, Najimi, Roméo,

Gnassia-Barelli and Narbonne 2000;

Kaaya, 2002), with some perturbations

in the physical and Chemical parameters

of seawater (Id Halla, 1997).

During this study, for each month,

ninety individuáis of 35 mm for each

species are collected at random in each

of the two sites. 30 individuáis/

species/sites were subject to stereologi-

cal analysis and 60 individuáis/

species/sites were dedicated to the bio-

chemical analysis.

Stereological analysis

In order to follow the seasonal vari-

ation of the glycogen according the

reproduction cycle in Mytilus gallo-

provincialis and Perna perna, an histo-

chemical study (stereological analysis)

was conducted in the mantle. Thirty

adult mussels were collected and

directly fixed in Gendre liquid for 3

days. Animáis were then removed from

shells and a central portion of mantle

tissue was correctly excised and embed-

ded in paraffin. Sections of 6 pm were

cut and stained by the periodic acid of

Schiff (PAS). Stereological analysis was

applied according the method of

Weibel, Kistler, and Scherle (1966)

which quantified the volume occupied

by the reserve and germinal tissues.

The cell types were determined accord-

ing the definition of Lubet (1959): i)

adipogranular cells (20-25 pm for

length, 4.5-5 pm for length of nucleus,

green coloration, contain lipids, pro-

teins and glycogen), and ii) vesicular

cells (50-80 pm for length, 3-5 pm for

length of nucleus, pink coloration,

contain only glycogen)

Biochemical analysis

Biochemical analysis (glycogen,

lipids and proteins) was carried out on

mussel mantle which in Mytilidae, con-

stitutes an important organ in storage of

reserves. Monthly, sixty adults were ran-

domly sampled, quickly transferred to

the laboratory in isotherm conditions

and frozen at -30°C. Glycogen, total

lipids and proteins were respectively

measured according the methods of

Duchateau and Florkin (1959),

Folch, Lees and Sloane-Stanley

(1957) and Lowry (1951). Rates of these

compounds were expressed as mg/g

fresh weight (mg/ g FW).

Statistical analysis

Stereological analysis was expressed

as a percentage of the total volume of

mantle tissue ; biochemical results as

means ± standard deviations. The statis-

tical significance of difference between

samples was evaluated by the "t" test

using the Statistica software (Release

4.5 A StatSoft Inc. Ed. 1993). A "P" valué

of less than 0.05 was considered as sta-

tistically significant.

19

Iberus , 26 (1), 2008

Figure 1. Cells implicated in the reserve storage in Perna perna (A) (only one type of cells: ve:

vesicular cells) and Mytilus galloprovincialis (B) (two types of cells, ve: vesicular cells; adge: adi-

pogranular cells). Scale bars 10 pm.

Figura 1. Células implicadas en la acumulación de reservas en Perna perna (A) (solo un tipo de células:

ve: células vesiculares) y Mytilus galloprovincialis (B) (dos tipos de células, ve: células vesiculares; adge:

células adipogranulares). Escalas 10 pm.

RESULTS

Stereological analysis

Figure 1A shows that the mantle of

Perna perna presents only one type of

storage cells: vesicular cells (VC). In

Mytilus galloprovincialis (Fig. IB), two

types of storage cells are presentí adi-

pogranular cells (ADGC) and vesicular

cells (VC).

For P. perna, the respective volumes

of germinal and reserve tissues are con-

versely proportional (Fig. 2A). The sea-

sonal profile is similar during the two

studied annual eyeles. The volume of

germinal tissue increases from October

to February, when the máximum is

reached (respectively 72% and 77% in

the first and second year). From March

onwards, the germinal tissue volume

decreases to reach the minimum in

August (respectively 8% and 6% in the

first and second year). The reserve

tissues appeared in spring (May) and

increased until reaching maximal valúes

in summer (August). They decreased

quickly and disappeared between

January and April. In the polluted site

(Fig. 3A), the seasonal variation of

reserve and germinal tissues is similar

to that in the reference site but the

máximum valué of reserve tissues per-

centage is lower (57% and 62% respec-

tively in the first and second year).

Moreover, during the year, this tissue

did not disappear. The minimum valué

recorded was 7-10% in February.

For Mytilus galloprovincialis (Fig. 2B),

the volume of germinal tissue increases

in autumn and in the beginning of

winter to reach a máximum in February

(75%). Thereafter, we observe a reduc-

tion of the surface occupied by this

tissue until October where the

minimum (12%) is reached. During the

second eyele a similar evolution is

recorded. Otherwise, as with P. perna,

the volume of germinal tissue is

inversely proportional to reserve tissue.

Nevertheless, in this case, the máximum

is reached in October (64%), whereas the

disappearance takes place from January

to March for the two studied yearly

eyeles. In the polluted site (Fig. 3B), the

reserve tissue presents a seasonal evolu-

tion similar to the reference site. Besides,

this tissue never disappears as in P.

perna. The occupied minimal volume is

5% reached in February.

Moreover, for M. galloprovincialis, in

the reference site, the adipogranular cells

disappeared before the vesicular cells

and then reappeared first. In the polluted

site the two cellular categories persist

20

Moukrim ET AL. : Reserves strategy of mussels in Moroccan coasts

A

Reserve tissue ^ Germinal tissue O Others

B

100%

90%

OND JFMAM J JASOND JFMAM J JA

ADGC

VC

Germinal tissue

I | Others

Figure 2. Seasonal variation of germinal and reserve tissues in Perna perna (A) and Mytilus gallo-

provincialis (B) in the unpolluted site. (VC: vesicular cells storing glycogen; ADGC: adipogranular

cells, containing glycogen, lipids and proteins).

Figura 2. Variación estacional de los tejidos germinal y de reservas en Perna perna (A) y Mytilus gallo-

provincialis (B) en el sitio no contaminado. (VC: células vesiculares almacenando glucógeno; ADGC:

células adipogranulares, conteniendo glucógeno, lípidos y proteínas ).

during all the year. However, an oscilla-

tion can be observed in the volumes

occupied by the two categories of cells.

Biochemical analysis

Glycogen: The mean amount of

glycogen is significantly (F= 11.19 and

P= 0.027) higher in Perna perna than in

Mytilus galloprovincialis (respectively

59.6 ± 7.36 and 48.7 ± 4.78 mg/g FW).

For the two species, the seasonal varia-

tions are significant (F= 4.95 ; P= 10"6 for

Perna perna and F= 2.65 ; P= 4.7xl0'6 for

Mytilus galloprovincialis) and exhibit a

similar profile (Fig. 4A). The glycogen

increases in spring (from April), reaches

a maximal valué in August (219.4 and

191.9 mg/g FW in Perna perna and

Mytilus galloprovincialis respectively),

then decreases during autumn and

winter. The minimal valúes are noted in

February (0.39 and 0.34 mg/g FW in

Perna perna and Mytilus galloprovincialis

respectively).

21

Iberus, 26 (1), 2008

ioo%-

90% -

80% -

70% '

60% -

50% -

40% '

30% '

20% '

10%'

o%-

i

¡i

i I

li

I

l

i

i ¡ I

p

II

lli

Wm

i i

11.

li

,1

H

1*

"

L

i

i

■I

i»

1

OND JFMAM J JA SOND JFMAM J JA

Reserve tissue ^ Germinal tissue Q Others

100%

90%

80%

70%

60%

50%

40%

30%

20%

10%

0%

B

Figure 3. Seasonal variation of germinal and reserve tissues in Perna perna (A) and Mytilus gallo-

provincialis (B) in the polluted site. (VC: vesicular cells storing glycogen; ADGC: adipogranular

cells, containing glycogen, lipids and proteins).

Figura 3. Variación estacional de los tejidos germinal y de reservas en Perna perna (A) y Mytilus gallo-

provincialis (B) en el sitio contaminado. (VC: células vesiculares almacenando glucógeno; ADG: células

adip ogranulares, conteniendo glucógeno, lípidos y proteínas).

ONDJFMAMJ JASONDJFMAMJ JA

¡H ADGC H VC ^ Germinal tissue Q Others

In the polluted site (Fig. 4B), the dif-

ference between the mean amount of

glycogen in the two species (39.75 and

23.44 mg/g FW in P. perna and Mytilus

galloprovincialis respectively) is not sig-

nificant (F= 6.257 and P= 0.130). Other-

wise, their seasonal variation are signifi-

cantly different (F= 4.03 and P= 10'6 in

P. perna and F= 9.78 and P= 10~7 in M.

galloprovincialis). The seasonal profile is

completely different for the two mol-

luscs: For P. perna , it presents three

peaks, the first in June (96.05 mg/ g FW),

the second in November (118.87 mg/g

FW) and the third in May of second year

(159.02 mg/g FW). For M. galloprovin-

cialis, only two peaks are recovered

65.03 and 153.64 mg/g FW respectively

in May and November of the first year.

The comparison of valúes for refer-

ence and polluted site animáis shows a

significant difference. The valúes in the

polluted site are less important (39.8 ±

13.36 instead of 59.6 ± 7.36 mg/g FW in

22

Moukrim ET AL Reserves strategy of mussels in Moroccan coasts

P perna ▲ M. galloprovincialis

Month

Figure 4. Seasonal variation of glycogen in the mantle of Perna perna and Mytilus galloprovincialis

in the reference site (A) and the polluted site (B)

Figura 4. Variación estacional del glucógeno el el manto de Perna perna y Mytilus galloprovincialis en

el sitio de referencia (A) y en el sitio contaminado (B)

the reference site, for Perna perna). More-

over, the seasonal profile of glycogen

contení (Fig. 4B) presents some pertur-

bations compared to the reference site.

Lipids : A significant difference was

noted between the mean amounts of total

lipids in the two mussels species (92.86

and 125.04 mg/g FW in P. perna and

Mytilus galloprovincialis respectively).

However, the difference of their seasonal

variation are significant with F= 4.42 and

P= 10'6 in P. perna and F= 3.6 and P=

4.5xl0'5 in M. galloprovincialis. In the first

year, the seasonal profile (Fig. 5A) is similar

for the two molluscs. The lipid contení

increased in autumn, with a maximal valué

in December (respectively 354.3 and 316.9

mg/ g FW in Perna perna and Mytilus gal-

loprovincialis), and decreased in winter and

spring. However, in the second year, a

slight increase was noted in summer (June

- July) only in P. perna.

In the polluted site, the two species

present a significant difference between

total lipids mean valúes (respectively

231.4 ± 21.47 and 135.5 ± 16.93 mg/g

FW in Perna perna and Mytilus gallo-

provincialis). Besides, the difference in

their seasonal variation (Fig. 5B) is sig-

nificant with F= 2.59 and P= 0.002 in P.

perna and F= 3.60 and P= 4.5xl0"4 in M.

galloprovincialis.

Ótherwise, the valúes are generally

higher for the polluted site than the ref-

erence site for the two species. More-

over, compared to the unpolluted site,

the seasonal variation (Fig. 5B) shows a

perturbed profile with several peaks

during the annual cycle and a máximum

valúes is reached rather in August

(777.03 mg/ g FW in P. perna and 659.95

mg/g FW in M. galloprovincialis).

Proteins : The mean contení of protein

is significantly higher in Perna perna (28.9

23

Iberus, 26 (1), 2008

P. perna a M. galloprovincialis

Month

Figure 5. Seasonal variation of lipids in the mantle of Perna perna and Mytilus galloprovincialis in

the reference site (A) and the polluted site (B).

Figura 5. Variación estacional de los lípidos en el manto de Perna perna y Mytilus galloprovincialis en

el sitio de referencia (A) y en el sitio contaminado (B).

± 3.51 mg/g FW) compared to Mytilus

galloprovincialis (20.2 ± 3.17 mg/g FW).

However, the seasonal profile (Fig. 6A) is

similar for the two species. The maximal

valúes were generally recorded in

summer and autumn in the first year

and in winter and beginning spring in

the second year, whereas the minimal

valúes were reached at the end of

autumn and the beginning of winter.

In the polluted site (Fig. 6B), the

mean amount of protein is higher in P.

perna (30.11 ± 7.742 mg/g FW) than in

M. galloprovincialis (19.74 mg/g FW).

However, the mean valúes and the sea-

sonal profile do not show any signifi-

cant difference compared with those

observed from the reference site (F= 1.08

and P= 0.30 for Perna perna ; F= 0.31 and

P= 0.57 for Mytilus galloprovincialis ). For

the two species, the máximum valúes

were recorded in summer and autumn.

DISCUSSION

The comparative study of the

reserve pattern of the two mussels

species living in the Moroccan coasts,

Perna perna and Mytilus galloprovincialis,

sampled in the same site (Cap Ghir,

Agadir bay), shows that, in spite of the

difference in their reserve tissues (only

one cell type, vesicular cells, in Perna

perna ; two cell types, adipogranular

cells and vesicular cells, in Mytilus gallo-

provincialis), the two molluscs have a

similar fluctuation.

This strategy presents cycles (of re-

serve compounds and cells involved in

the storage) which, compared to the cy-

cle of reproduction (determined in these

same animáis by Id Halla et al., 1997)

are inversely proportional to it. The ac-

cumulation of reserves in these cells is

related to the period of reduced sexual

24

Moukrim ET AL. : Reserves strategy of mussels in Moroccan coasts

P. perna a M. galloprovincialis

Figure 6. Seasonal variation of proteins in the mantle of Perna perna and Mytilus galloprovincialis

in the reference site (A) and the polluted site (B).

Figura 6. Variación estacional de las proteínas el manto de Perna perna y Mytilus galloprovincialis en

el sitio de referencia (A) y en el sitio contaminado (B).

activity (in summer). The low levels of

reserve are obtained in autumn and par-

ticularly in winter (gametogenesis and

spawning periods). Similar results have

been reported in Perna perna from

Brazilian coasts (Lunetta, 1969) and in

Mytilus galloprovincialis from the French

coasts (Herlin-Houtteville, 1974;

Danton, Kiymoto, Komaru, Wada,

Awaji and Mathieu, 1996). Also, in the

Ria de Vigo (Spain), the profile of the

variations of the gonadic Índex and so-

matic Índex in cultured mussels show

clearly their inverse and gradual fluctu-

ations (Cáceres-Martínez and

Figueras, 1998). This association be-

tween gonad and storage tissue cycles

in mussels is well known. Decline of the

ADG cells occurs during gametogenesis

by a lysosomal autophagic mechanism

(Bayne et al., 1982; Lowe, Moore and

Bayne; 1982; Pipe, 1987). The energy

used in gonad restoration following

spawning during spring and summer

probably derived directly from feeding

since ADG cells disappeared from the

mantle in early spring (Villalba, 1995).

Furthermore, the reserve accumula-

tion in the two mollusc species is related

to the proliferation of the phytoplanc-

tonic biomass linked to upwelling cur-

rents which take place, between Febru-

ary and August, in the Agadir bay and

more precisely in Cap Ghir (Belveze,

1984 ; Agoumi and Orbi, 1992).

Cáceres-Martínez and Figueras

(1998) reported that this increase in food

availability for mussels in the area

favours the accumulation of reserves

during this period. These results also

confirm the cióse dependence, described

by Mathieu (1987), between the reserve

storage and environmental conditions in

marine ecosystems. As suggested by

25

Iberus, 26 (1), 2008

this author, the disappearance of reserve

tissue in mussels seems to be under

endocrine control via cerebroid ganglia

which provoke disappearance of adi-

pogranular cells and vesicular cells in

the mantle after liberation of their

reserves which are indispensable for

gametogenesis and spawning process.

The synchronic character of this disap-

pearance, in Perna yerna and Mytilus gal-

loyrovincialis, could be explained by the

existence of the same mechanism for the

control of reserves in both species.

The comparison of the reserve com-

pounds in the two molluscs shows that

the glycogen, total lipids and proteins

contents are higher in Perna yerna (in

spite of the presence of only one type of

reserve storage cells, vesicular cells)

compared to Mytilus galloyrovincialis.

These results could explain the differ-

ences observed by Id Halla et al.

(1997) between the reproductive cycles

of these bivalves. According to these

authors, in Perna yerna, compared to

Mytilus galloyrovincialis, the main-

spawning period in spring is longer and

the sexual activity in summer is

reduced. Otherwise, the important

reserve levels in this species could be

attributed to the spatial distribution of

each species: Perna yerna lives essen-

tially at the infra-littoral level and is

more immersed than Mytilus galloyrovin-

cialis (Id Halla et al., 1997) and, conse-

quently, has access to more nutrients,

thus allowing the synthesis and storage

of more reserves. Seed (1976) showed

that the gonadal development was

faster in mussels from the low intertidal

zone, than in those from the upper zone

and related these results to food avail-

ability. Other studies associated local

variations in gonadal cycle with envi-

ronmental conditions (Ferrán, 1991;

Villalba, 1995). According to Cáceres-

Martínez and Figueras (1998), there is

no influence of locality and depth in the

gonadal development of cultured

mussels.

The seasonal profile of the biochemi-

cal reserves studied shows that the first

peak of glycogen and lipids, recorded in

summer, is related to the occurrence of

upwelling currents which provide food

availability, ensuring an abundant

planktonic food supply for mussels.

According to Cáceres-Martínez and

Figueras (1998), massive spawns occur

in spring coinciding with an increase in

temperature and chlorophyll-a concen-

traron in the area providing favorable

conditions for larval growth. In winter,

the second peak of lipids coincides with

the maturity period of gametes. This

peak takes place just after the disap-

pearance of the glycogen peak. It is

probably a result of the glycogen trans-

formaron. The metabolic conversión of

glycogen to lipids has been reported by

Zaba and Davies (1980), using 14C-

glucose. According to Gosling (1992),

the mantle is considered as the organ of

many and extensive metabolic transfor-

mations during the sexual cycle. The

reserves, particularly the glycogen,

accumulated during summer, are used

in autumn and winter for the gametoge-

nesis. Similar results were reported by

Shafee (1989) in Perna yicta of Temara

(North of Morocco).

In the polluted site (Anza), many

perturbations in the metabolism of

reserves were noted comparatively to

the reference site (Cap Ghir). The

reserve tissues, which disappeared in

winter and at the beginning of spring, in

mussels of Cap Ghir, persist throughout

the sexual cycle in Anza mussels. This

could probably be a result of the pollu-

tion effect on the cerebroid ganglions

neurosecretions which are, according to

Lubet, Herlin, Mathieu, and Collin

(1976) and Mathieu (1987) involved in

the control of reserve cells.

The analysis of the seasonal profile

of the reserve levels in mussels sampled

in the polluted site shows some pertur-

bations as compared to the reference

site. For example, the glycogen content

is low in summer (August), in spite of

the availability of phytoplanctonic

biomass in this period. This is probably

linked to the stress caused by pollution

of industrial and domestic waste waters

discharged directly in this site, without

any treatment. This fact was indicated

by Deslous-Paoli, Wolowicz, and

26

Moukrim ET AL.: Reserves strategy of mussels in Moroccan coasts

Boromthanarat (1991) who reported

that, in Mytilus edulis, reserves could be

used both in reproductive process and

to overeóme the hard environmental

conditions. According to Thompson

(1972) the reserves are used in order to

reach the basal level of energy necessary

for stressed animáis.

Contrary to the glycogen, the total

lipids are more important in Perna yerna

and Mytilus galloprovincialis living in

Anza (polluted site). This could be

explained by an eventual direct assimi-

lation of lipids from the organic matter

of waste waters and/or a change of the

reserve storage process. According to

Gosling (1992) the lipid storage process

in molluscs is considerably linked to the

environmental conditions, particularly

to the presence of pollutants.

As a general conclusión, in the two

species of mussel P. yerna (with only

vesicular cells) and M. galloprovincialis

(with adipogranular cells and vesicular

cells), the respective germinal and

reserve tissues clearly show their

inverse and gradual profile. Their sea-

sonal fluctuations are similar in the two

molluscs. In the polluted site, many per-

turbations of the reserve metabolism

were noted comparatively to the refer-

ence site. Then, contrary to animáis of

this latter site, which presented a glyco-

BIBLIOGRAPHY

Agoumi, A. and Orbi, A., 1992. Evolution

météorologique et upwelling le long de la cote

atlantique marocaine. Hydroécologie Appliquée,

2 (4): 149-158.

Bayne, B. L., Bubel, A., Gabbott, P. A., Liv-

INGSTONE, D. R., LOWE, D. M. AND MOORE,

M. N., 1982. Glycogen utilisation and game-

togenesis in Mytilus edulis L. Marine Biology

Letters, 3: 89-105.

Belveze, H., 1984. Biologie et dynamique des pop-

ulations de la sardine (Sardina pilchardus Wal-

baum) peuplant les cotes atlantiques du Maroc.

Thése de Doctorat d'Etat, Université de Bre-

tagne Occidentale, Brest: 229 pp.

Cáceres-Martínez, J. and Figueras, A., 1998.

Long-term survey on wild and cultured mus-

sels ( Mytilus galloprovincialis Lmk.) repro-

ductive eyeles in the Ria de Vigo (NW. Spain).

Aquaculture, 162: 141-156.

genic strategy, a lipidie strategy takes

place in molluscs of the polluted site.

The results obtained in this work

constitute a contribution to the knowl-

edge of the reserve strategy in the two

species living along Moroccan coasts, the

African mussel Perna yerna and the

Mediterranean mussel Mytilus gallo-

yrovincialis. So, several faets are reported:

i) the type, the seasonal variation and the

eyele of the reserve strategy in the two

molluscs, ii) cells involved in the storage,

iii) the relationship between the reserve

strategy and the reproduction eyele, and

iv) the response of this strategy to the

environmental conditions. Nevertheless,

it would be interesting to identify the

mechanism and the control process of

the reserve strategy in these molluscs.

ACKNOWLEDGMENTS

We are grateful to Miss Joanne

Preston (Southampton University, UK)

and Mrs Barbara Picot (Angers Univer-

sity, France) for their Language correc-

tions. We thank the IFS (Sweden),

AUPELF (Cañada), Ministére des

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28

Iberus, 26 (1): 29-42, 2008

© Sociedad Española de Malacología

Reproduction of the cockle Cerastoderma edule (Linné,

1758) in the estuary of Oued Souss (southwestern Morocco)

Reproducción del berberecho Cerastoderma edule (Linné, 1758) en

el estuario del Oued Souss (suroeste de Marruecos)

Hafida BERGAYOU**, Abdellatif MOUKRIM*1, Michel MATHIEU** and

Jean-Pierre GIMAZANE**

Recibido el 14-11-2006. Aceptado el 3 l-X-2007

ABSTRACT

Field and laboratory investigations were carried out from 2001 to 2003 on a population

of Cerastoderma edule living ¡n the estuary of Oued Souss, ¡n order to determine the

impact of domestic wastewater discharges on the biology of this species. During waste-

water discharges, several episodes of partial spawning occurred from November to

March, followed by another in April-May. The main spawning occurred nevertheless in

August. The gonadic Índex was always higher than 1, so that the period of sexual rest

was short. The period of recruitment (from June to October) was mainly correlated with

spawning peaks. The renewal of the population was ensured by the recruitment of June-

July, whereas young cockles recruited from August to October only represented 9.2% of

the population at the end of autumn. Most 1-year-old cockles disappeared during the sum-

mer of the second year following their recruitment. After wastewater pollution stopped, an

important recruitment of young cockles occurred. The spatial distribution of this species

progressed upstream and the biomass of cockles had strongly increased. Because of its

reproduction which takes place throughout the year, C. edule , as an opportunistic

species, succeeded in colonizing this site.

RESUMEN

Se realizaron, entre 2001 y 2003, estudios de campo y de laboratorio sobre una pobla-

ción de Cerastoderma edule del estuario del Oued Souss, con el objetivo de determinar el

impacto de los vertidos de aguas residuales sobre la biología de esta especie. Coinci-

diendo con los vertidos, se produjeron varios episodios de puesta parcial entre Noviem-

bre y Marzo, seguidos de otro en Abril-Mayo. La puesta principal, sin embargo, se pro-

dujo en Agosto. El índice gonádico fue siempre superior a 1 , indicando que el periodo de

reposo sexual fue corto. El periodo de reclutamiento (de Junio a Octubre) estuvo esencial-

mente relacionado con picos de puesta. La reposición de la población fue asegurada por

el reclutamiento de Junio-Julio, mientras berberechos jóvenes reclutados entre Agosto y

Noviembre representaban tan solo el 9,2% de la población a finales de otoño. La mayor

parte de los berberechos con un año de edad desaparecieron durante el verano del

segundo año después de su reclutamiento. Al finalizar los vertidos se produjo un impor-

tante reclutamiento de berberechos jóvenes. La especie se extendió río arriba y la bio-

* Laboratory Aquatic Ecosystems : Marine and Continental Field, Biology Department, Sciences Faculty, Ibn

Zohr University, BP 8106, 80000, Agadir, Morocco.

** Laboratory of Marine Biology and Biotechnology, University of Caen, F- 14000, Caen, France.

1 Corresponding author

29

Iberus , 26 (1), 2008

masa de berberechos aumento fuertemente. Por su reproducción que se extiende sobre

todo el año, C. edule, como especie oportunista, fue exitosa en la colonización de este

lugar.

KEY WORDS: Cerastoderma edule, estuary, Oued Souss river, pollütion, recruitment, reproductive cycle, waste-

water discharges.

PALABRAS CLAVE: Cerastoderma edule, estuario, Oued Souss, contaminación, reclutamiento, ciclo reproductor,

INTRODUCTION

In marine environment, many biotic

and abiotic factors may influence the life

cycle of bivalves. If temperature, salin-

ity, food supply, and tidal exposure are

the most important causes known to

modulate development in mytilids

(Seed, 1975), other elements, such as

intraspecific competition or environ-

mental contaminants, can result in great

variations in growth rate of bivalves

(Seed and Suchanek, 1992). These

factors also have an effect on the repro-

duction, settlement, recruitment, and

production of these molluscs so that

their variations directly affect the devel-

opment of beds for each species of

bivalve (Widdows and Donkin, 1992).

Contrary to numerous ecological

reports on the effects of contaminants on

different marine bivalves, the recovery

of mollusc populations after the disap-

pearance of pollütion has been less

investigated.

The implantation of a wastewater pu-

rification plant since November 2002

along the estuary of Oued Souss, at 4 km

from the mouth of the estuary, had re-

sulted in the fact that the fresh water still

running in the river and the decanted

wastewater were no longer discharged in

the estuary but diverted to another

Coastal site: M'Zar, located at 3 km south.

For this reason, the estuary, upstream to

the purification plant, was only swept by

sea tide at the present time.

As there existed a population of

Cerastoderma edule living in the estuary

of the Oued Souss river, it was interest-

ing to determine the physiological State

of these cockles during the pollütion

period by wastewater and the changes

which have occurred after this contami-

nation stopped. In view of these objec-

tives, the following two questions aróse:

Had the discharge of wastewater before

November 2002 caused repercussions

on the reproductive cycle of C. edule and

on the dynamics of this population?

What consequences did the presence of

sea water in the estuary after November

2002 have on the spatial distribution

and biomass of C. edule? To answer the

first question, monthly investigations

from January 2001 to December 2002

were carried out in the habitat of C.

edule i) to follow the different stages of

sexual maturity in males and females,

and ii) to specify the development of the

different generations and cohorts of

cockles. To tackle the second question,

two surveys in 2002 and 2003 during

summer months were performed in the

estuary of Oued Souss river.

This study complements other

reports made by our research team in

the same ecosystem (Mouneyrac, Pel-

LERIN, MOUKRIM, AlT ALLA, DUROU AND

Viault, 2005; Ait Allá, Mouneyrac,

Durou, Moukrim and Pellerin, 2005;

Ait Allá, Gillet, Deutsch, Moukrim

and Bergayou, 2005; Bergayou and

Moukrim, 2003 and Gillet, Gorman,

Tallec, Moukrim, Mouloud, Anajjar,

Ait Allá, Bergayou and Kaaya, 2003).

MATERIALS AND METHODS

The estuary of Oued Souss is located

on the Atlantic coast, in southwestern

Morocco and is subjected to an arid

climate. The mouth is swept by an

intense marine hydrodynamism,

responsible of the presence of great

30

Bergayou ET AL.: Reproduction of Cerastoderma edule in Southwestern Morocco

Figure 1 . Location of sampling sites along the estuary of Oued Souss in the Bay of Agadir, south-

western Morocco.

Figura 1. Situación de las localidades de maestreo a lo largo del estuario del Oued Souss, en la bahía de

Agadir, suroeste de Marruecos.

sandbanks in the estuary, with currents

linked to high or low tides, and with

high salinity (from 31.2 to 34.2 %o).

Reproductive cycle

The study of reproductive cycle and

population dynamics for C. edule were

carried out at station 2 (30° 21.97'N

north, 9o 35.98'W west), (Fig. 1). Selected

because of its high biomass of the cockle

population, this station was character-

ized by fine silty sand 2-5% of organic

matter in the substratum and 340-440

mg/1 of suspended matter in the water.

Two methods for studying the repro-

ductive cycle of C. edule were used. The

first was a classical histological study of

gonads and was performed from

January 2001 to March 2002. The second

was a complementary study of condi-

tion Índex and microscopic examination

of gonad smears (Guillou, Bachelet,

and Glémarec, 1991) and was per-

formed from August 2001 to August

2002. For the first and second study.

respectively, 30 and 100 cockles (length,

20 to 30 mm) were collected by hand at

low tide, at monthly intervals (respec-

tively, a total of about 450 and 1300

accumulated cockles was examined).

For the first study (histological study

of gonads), the shells of C. edule were

opened and soft masses were prefixed

in the Gendre's fixative for 24 h. In the

laboratory, the shell of each bivalve was

removed and small pieces of soft masses

were post-fixed in a new solution of

Gendre's fixative for 48 h before being

dehydrated through a graded series of

ethanol and butanol, and finally embed-

ded in cytoparaffin (56°-58°C). Serial

sections (thickness, 5 pm) were made

before being stained with Gabe's

trichrome, hemalun-eosin, or Mann-

Dominici's method (Gabe, 1968). The

maturity of gonads was determined

using the scale proposed by Lubet

(1959) for Mytilus edulis (Table I).

The gonadic Índex (Seed, 1975) indi-

cates the State of gonad maturity for

31

Iberus, 26 (1), 2008

Table I. Terminology used by LUBET (1959) and LUCAS (1965) to study morphologically and his-

tologically the gonadic developmental stages in bivalves.

Tabla I. Terminología empleada por Lubet (1959) y Lucas (1965) para el estudio morfológico y his-

tológico de estadios de desarrollo gonadal en bivalvos.

32

Bergayou ET AL.: Reproduction of Cerastoderma edule in Southwestern Morocco

size class (20-30 mm) was considered

and 30 bivalves were randomly chosen

to determine this condition Índex. The

índex selected was that proposed by

Lucas and Beninger (1985): IC = [(dry

weight of soft masses) / (dry weight of

valves) x 1000]. The dry weight was

obtained using a dehydration of soft

masses (or shell) in a desiccator (60°C,

24 h).

Population dynamics

To study the dynamics of the popu-

lation and to specify the period of juve-

nile recruitment, samplings of C. edule

were performed from January to

December 2002. This study was also

carried out in the station 2. Monthly,

sixteen sediment samples (surface,

0.0625 m2, height, 20 cm) were collected

at low tide according to the method of

quadrats used by Elliott and Decamps

(1973, in Bayed, 1982). These samples

were sieved (square meshes, 1 mm) to

recover and count cockles. The antero-

posterior length of each cockle was mea-

sured using a calliper rule (precisión, 0.1

mm). For mollusc sizes less than 5 mm,

a stereomicroscope equipped with a

micrometric ocular was used. The indi-

vidual valúes recorded for the length of

bivalves and sampling dates were com-

pared using the FISAT software

(Gayanilo, Sparke and Pauly, 1996) to

draw size histograms in relation to the

frequency of animáis and to make a

modal analysis based on the algorithm

according to the method by Battacharya

(1967). This comparison, with the use of

the x2 test (at P<0.05) allows to distin-

guish the different size classes in each

monthly sample, to determine mollusc

densities, and to specify recruitment

periods.

Spatial distribution, density and

biomass

To determine the spatial distribu-

tion, density, and biomass (ash-free dry

weight, AFDW) of cockles, two surveys

were carried out in 2002 and 2003

(during summer period). The replication

of this study during these two periods

(in 2002, when the estuary received the

wastewater, and in 2003, when the dis-

charge stopped) gives insights on the

wastewater pollution impact and the

reestablishment of the ecosystem when

this pollution stopped. These investiga-

tions were made in a total of 8 stations

(27 sampling sites) located at 400-500 m

intervals from the estuary mouth up to

the wastewater outlet (Fig. 1). In each

site, four to six sediment samples

(surface, 0.0625 m2, height, 20 cm) were

collected at low tide according to the

method of quadrats used by Elliott

and Decamps (1973, in Bayed, 1982).

These samples were sieved (square

meshes, 1 mm) to recupérate and count

cockles, as above.

Ash-free dry weight (AFDW) was

determined by dipping molluscs in a

solution of 10% HC1 until the shell was

completely dissolved. Then, the soft

parts were dried for 48 h at 80°C,

weighed, placed at 600°C for 2 h and

weighed again. The weight loss at 600°C

was considered to represent the AFDW

of individuáis and was expressed in

g/m2 (Bachelet, Bouchet and

Lissalde, 1980-1981).

RESULTS

Effects of pollution on the biology

of the species

Reproductive cycle: The study of

gonad smears from August 2001 to

August 2002 had allowed to recognize

sex in 1138 bivalves (87.5%). Within this

group, the sex ratio of C. edule was

51.1% ± 2.6%, and was well-balanced at

1:1 (x2 = 2.5, P < 0.05). No hermaphro-

dite individuáis were found in this pop-

ulation. In September, the A stage

(undetectable sex) was observed in 75%

of cockles and might correspond to a

previtellogenic phase rather than a

sexual rest, as the percentage of C. edule

showing the C stage (gonad maturation)

increased in October to reach about 50%

in November. The frequency of C stage

was above 50% between November and

April (Fig. 2A), and subsequently

decreased in May (from 75 to 20%), thus

corresponding to a slight decrease of the

33

Iberus, 26 (1), 2008

Figure 2. Distribution of sexual-development stages (A) according to the scale of LUCAS (1965) and

annual cycle of condition indexes (B) in the populations of C. edule studied from August 2001 to

August 2002. The cumulated frequencies of the different sexual-development stages corresponded to

the total of bivalves studied (100%). For details of each stage, see Materials and Methods (Table I).

Figura 2. Distribución de los estadios de desarrollo sexual (A) según la escala de LUCAS (1965) y ciclo

anual de índices de condición (B) en las poblaciones de C. edule estudiadas entre agosto 2001 y agosto

2002. Las frecuencias acumuladas de distintos estadios de desarrollo sexual corresponden al total de

bivalvos estudiados ( 1 00%). Véase Material y Métodos (Tabla L) para los detalles de cada estadio.

condition Índex (Fig. 2B). An episode of

pardal spawning had thus occurred

during this last period. In May, the fre-

quency of the D stage (80%) proved that

gametogenesis had resumed. In June

and July, the C stage was the most fre-

quent (65 to 75%) and was followed in

August by a more marked decrease of

the condition Índex (from 80 to 50 %o)

thus indicating an important release of

gametes. It may be concluded that two

periods of gamete maturation (C stage),

the first ranging from November to

April and the other occurring in June-

July, were differentiated.

The histological study was carried

out over a longer period of time (January

2001- March 2002) than the analysis of

gonad smears. Mature gonads (IIIA

stage) were often observed on more than

50% of cockles studied from January to

April (Fig. 3). Episodes of pardal spawn-

ing occurred from January to March; fol-

lowed by another spawning in April and

May (in May, the IIIB stage was noted in

60% of males, Fig. 3A). The reconstitu-

tion of the gonad (IIIC stage) is more

marked in females (Fig. 3B) in May

(80%) and seems more precocious in

males. In both sexes, this stage devel-

oped in parallel from May to July so that

pardal spawning occurred during these

months. In August, the IIIB stage was

preponderant in both sexes (80% of

34

Bergayou ET AL.: Reproduction of Cerastoderma edule in Southwestern Morocco

A

B

2001 Months 2002

2001 Months 2002

Figure 3. Distribution of gonadic-development stages over 2001-2002 in the males (A) and females

(B) of C. edule. The cumulated frequencies of the different developmental stages corresponded to

the whole bivalves studied (100%). For details of each stage, see Materials and Methods (Table I).

Figura 3. Distribución de los estadios de desarrollo gonadal sobre el periodo 2001-2001 en machos (A) y

hembras (B) de C. edule. Las frecuencias acumuladas de distintos estadios de desarrollo sexual corre-

sponden al total de bivalvos estudiados (100%). Véase Material y Métodos (Tabla I) para los detalles de

cada estadio.

males, 60% of females), proving that the

main spawning episode had happened.

The resorption of gonads (IIID stage)

was observed in a few individuáis after

the spawning of May and was less than

25% from June to August. Several

females in IIID stage were noted during

the other months, but they never

exceeded 5%. In September, gametogen-

esis started in both sexes (I and II stages

were found in 95% of cockles). The mat-

uration of the gonad (IIIA stage) was

observed in the first bivalves from

October to December and partial spawn-

ing occurred from November to March.

The highest valúes (2.7) of the

gonadic Índex (Fig. 4) were noted

between January and April, thus con-

firming the presence of ripe gonads

during this period. The lowest valúes

(1.7) were found from June to August. In

both sexes, the gonadic Índex was

always higher than 1 so that the period

of sexual rest was short.

Gametogenesis in the population of C.

edule: Figure 5 gives the results of stere-

ological analysis. From January to

March, the percentages of mature

gametes increased to peak in April.

Another episode of gametogenesis

began in May, as demonstrated by the

high frequencies of young germinal cells

during this month in both sexes. In

males, the effect of this developing sper-

35

Iberas, 26 (1), 2008

—°"“a — b

Figure 4. Annual cycle of the gonadic Índex in the males (A) and females (B) of C. edule over

2001-2002.

Figura 4. Ciclo anual del índice gonádico en machos (a) y hembras (b) de C. edule en el periodo 2001-

2002.

matogenesis was high rates of sper-

matids and spermatozoa in June or

August (grouped frequencies, 75% and

85%, respectively). In females, an impor-

tant vitellogenesis was observed in May

and June so that the highest frequency

of mature oocytes (55%) was noted in

July and that of degenerated oocytes

(60%) in August. In September, the

highest percentages of spermatogoniae

(29%) in males and of oogoniae (37%) in

females were noted, thus indicating the

starting of another gametogenetic wave.

From September to December, the

gonadic tissue of males was important,

as the grouped frequencies of spermato-

goniae and of spermatocytes were more

than 47%. The same finding was noted

in females during this period (grouped

rates of oogoniae and of vitellogenic

oocytes, >70%). Low percentages of

atresic oocytes (5-25%) were always

observed in females throughout the

year.

From the above study, it can be con-

cluded that three successive gametoge-

netic waves were differentiated: the first

one from January to April, the second

one from May to July, and the third one

from September to March. Ripe cockles

for both sexes were predominant from

January to April, in July, and from

January to March.

Population dynamics: The lengths of

C. edule , measured from January to

December 2002, are given in Figure 6.

The period of recruitment spanned from

the end of spring to the onset of

autumn. The first settlement of juvenile

cockles occurred in June. A second

cohort appeared in July, a third cohort in

August, and a fourth one in October. At

the end of autumn, the individuáis

recruited in June-July had a length of

18.3 ±1.8 mm and constituted 71.6% of

the population. By contrast, those origi-

nating from the grouped cohorts of

August and October only had a length

of 11.1 ± 1.1 mm and represented 9.2%

of the population. In addition to these

recruitments; a few young cockles (2-

3%) were found for each date of sam-

pling. In summer, there was a strong

decrease in the number of cockles mea-

suring 24.6 ±1.9 mm in length. This

diminution is followed by the almost

complete disappearance of adult cockles

just after the period of recruitment

(August 2002) so that the population of

C. edule in 2002 was mainly composed of

young individuáis which are being

renewed each year.

36

BERGAYOU ET AL. : Reproduction of Cerastoderma edule in Sourhwestern Morocco

■ Spz

Hfl Spc

EH3 Spg

B

JFMAMJ JASONDJFM

2001 Months 2002

Figure 5. Frequencies of different cell categories in the male (A) and female (B) gonads of C. edule

collected between January 2001 and March 2002. Male line: Spc (primary and secondary sperma-

tocytes), Spg (protogoniae and spermatogoniae), Spz (spermatids and spermatozoa) . Female line:

D (degenerated oocytes), Og (oogoniae), R (mature oocytes), V (vitellogenic oocytes). The cumu-

lated frequencies of the different cell categories corresponded to the whole bivalves studied (100%).

Figura 5. Frecuencia de distintas categorías de células en gónadas de machos (A) y hembras (B) de C.

edule recolectados entre enero 2001 y marzo 2002. En machos: Spc (Espermatocitos primarios y secun-

darios), Spg (protogonias y espermatogonias), Spz (espermatidios y espermatozoos); en hembras: D (ovoc-

itos degenerados), Og ( oogonias ), R (ovocitos maduros), V (ovocitos vitelogénicos). Las frecuencias acu-

muladas de distintos estadios de desarrollo sexual corresponden al total de bivalvos estudiados (1 00%).

Spatial distribution, biomass, and

structure of population of C. edule

before and after wastewater discharges

The spatial distribution of C. edule

along the estuary, its density and its

biomass are given in Table II for the

summer surveys (2002 and 2003).

During the period of wastewater dis-

charge, the species was confined down-

stream (stations 1 and 2). By contrast, in

2003, its distribution had extended with

an upstream penetration of cockles up

to the station 5. The density of C. edule

did not exceed 320 individuals/m2 in

2002, and strongly increased in 2003 to

reach valúes higher than 4,900

cockles/m2 in several places (these

valúes were found during the periods of

recruitment). The mean biomass

(AFDW) calculated on the cockle

samples collected during summer

surveys, increased from 2.54 g/m2 in

2002 to 14.87 g/m2 in 2003.

Table III shows the different groups

of cockles constituting the population in

summer surveys (2002 and 2003).

37

Iberus , 26 (1), 2008

Length (mm)

0 10 20 30

N=171

N=182

N=207

N=242

N=300

N=244

N=1 14

N=119

N=256

N=92

N=119

Figure 6. Size distribution of C. edule in the estuary of Oued Souss from January to December

2002 and the principal normal components. Month and numbers of individuáis measured (N) are

given for each sampling.

Figura 6. Distribución de tallas de C. edule en el estuario del Oued Souss entre enero y diciembre 2002

y componentes normales principales. Meses y número de ejemplares medidos (N) están indicados para

cada muestra.

During the wastewater discharges in the

estuary (July 2002), individuáis are dis-

tributed among four groups: two classes

with lowest lengths (6.81 ± 1.53 mm and

9.97 ± 0.83 mm), deriving from a recruit-

ment in May and June; and two classes

of oíd cockles (24,72 ±2.17 mm and 27.5

± 0,76 mm). In July 2003, the most fre-

quent group (89.2%) was composed of

young individuáis (length, 6.55 ± 1.17

mm; 11.76 ± 1.63 mm) originating from

a recruitment in May and June. Another

group of médium size (36.25 ± 1.73 mm)

appeared for the first time and probably

comprised cockles more than one year

oíd, as the absence of growth lines on

cockle valves did not allow to easily

identify the different age subgroups

constituting this last size group.

DISCUSSION

The results reported in the present

study on the gonochorism of C. edule

agree with the reports of several authors

(Gimazane, 1971; Kingston, 1974).

Several cases of accidental hermaphro-

dism (4%0) were also noted by Fernan-

dez Castro, Guillou, Le Pennec and

Cárdenas López (1989). In the estuary

of Oued Souss, the sex ratio of cockles

was well-balanced so that pollution did

not have an influence on the distribu-

tion of males and females. This finding

agrees with studies that some authors

have performed in other populations of

C. edule at different latitudes, such as the

report by Kingston (1974) along the

coasts of Kent (UK), that of Ivell (1981)

38

BERGAYOU ET AL.: Reproduction of Cerastoderma edule in Southwestern Morocco

Table II. Spatial distribution of C. edule along the estuary, its density and its Biomass (AFDWg

/ m2) during wastewater contamination (summer 2002) and after stopping of pollution (summer

2003).

Tabla II. Distribución espacial de C. edule a lo largodel estuario , su densidad y su biomasa (AFDW/m2)

durante vertidos de agua contaminada ( verano 2002) y después de poner fin a la contaminación ( vera-

no 2003).

in the Limfjord (Denmark), that of

Mejuto (1984) in Ria de Pasaxe (Spain),

or still that of Fernandez Castro,

Guillou, Le Pennec and Cárdenas

López (1989) at Brouennou (France).

Through methods used for repro-

ductive cycle analysis, complementary

and concordant results were noted. In

the year, two successive gametogenetic

waves, the first occurring from Septem-

ber to April and the second in May-July,

were differentiated in this population

and spawning periods staggered over

time. These phenomena can be

explained by an effect of latitude (Sola,

1997; Rodriguez-Rua, Prado, Romeo

and Bruzon, 2003) and, in particular,

by water temperature (Hugues, 1971).

When temperature was less than 10°C, it

induced early spawning, followed by

gonad reconstitution and a second

period of spawning.

The presence of atresic oocytes

observed throughout the year during

vitellogenesis, as mentioned by Lubet

(1991) is apparently a frequent phenom-

enon in bivalves noted in the start of

gametogenesis (first mature oocytes

degenerate), after partial spawnings and

in the end of a reproductive cycle.

The long period of cockle recruit-

ment (from June to October) in 2002 can

39

Iberus, 26 (1), 2008

Table III. Distribution of shell lengths for C. edule during summer surveys (2002 and 2003): prin-

cipal normal components.

Tabla III. Distribución de longitudes de conchas de C. edule durante muéstreos de verano (2002 y

2003): principales componentes normales.

be easily explained by spawning which

occurred from April to August and this

time was perfectly consistent with the

gonadic-development stages deter-

mined by the histological study of

gonads and the examination of gonad

smears. Contrary to juveniles recruited

in June-July which had a better develop-

ment, those settled from August to

October showed a great mortality. This

last result would not be related to

wastewater pollution, in agreement

with the reports by Madani (1989),

Sauriau (1992), Bachelet, Guillou

and Labourg (1992a), or with that by

Bachelet, Desprez, Ducrotoy,

Guillou, Labourg, Rybarczyk,

Sauriau, Elkaim and Glémarec

(1992b). According to these authors,

high mortalities of cockles were noted in

the recruitments which occurred at the

end of summer, in autumn, and in

winter. This mortality affected small-

sized (<10 mm) cockles and may be

explained by the almost complete

absence of energetic reserves (Sauriau,

1992) whereas the metabolism of young

cockles was changing (Gabbott, 1976).

By contrast, the disappearance of

medium-sized molluscs in 2002 coin-

cided with the period of summer

recruitment. This finding might be

explained, either by a great predation of

cockles by Haematopus ostralegus (this

bird only tackled on médium sizes as

reported by Atkinson, Clark, Bell,

Daré, Clark and Ireland, 2003; John-

stone and Norris, 2000), or by a high

rate of mortality for these bivalves, as

their vulnerability became more

marked with increasing age and was

also dependent on the quality of waters

and/or an eventual eutrophication in

the estuary, as demonstrated by Ducro-

toy and Ibanez (2002). Indeed, as sea

waters in winter and spring had high

loads of mineral seston in the absence

of pollution, the filter-feeders produced

numerous pseudo-faeces and had a

marked energetic expense for selective

sorting of particles, mucus secretion,

and cleaning of gills for Mytilus edulis

(Widdows, Fieth and Worrall, 1979)

or for Crassostrea gigas (Héral, 1986).

According to Foster-Smith (1975a, b)

and Newell (1977), the behaviour of

adult cockles was different, as they

adapted their pumping activity in the

presence of high concentrations of

mineral seston, with a passage by a rel-

ative State of dormancy (Savari, Lock-

wood and Sheader, 1991). However,

such a behaviour did not seem to exist

during wastewater discharges in

summer, as cockles were in reproduc-

tion and had to live in waters double

loaded with matters in suspensión (pol-

lution) and the phyto-planktonic bloom

(with the increase of temperature),

therefore placing these bivalves under

stress conditions and inducing a high

mortality.

Since November 2002, the discharge

of decanted wastewater in another site

40

Bergayou ET AL. : Reproduction of Cerastoderma edule in Southwestern Morocco

beyond the estuary and the conversión

of this ecosystem into a marine environ-

ment had resulted, for C. edule, in a

wider spatial distribution, a strong

increase of density, and the appearance

of médium sizes in this site. To

comment on these changes, it is neces-

sary to take into account the reproduc-

tion of cockles living in this estuary, as it

is stretched over all the year. Indeed,

according to Gordo (1982), the repro-

ductive cycle of Spanish and Portuguese

populations of C. edule showed a period

of sexual rest during summer months

and the author explained it by the mean

temperature of sea water which were

more than 28°C and would inhibit the

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42

© Sociedad Española de Malacología

Iberus , 26 (1): 43-46, 2008

Xylodiscula wareni n. sp., una nuova specie per le coste

orientali della Sicilia

Xylodiscula wareni n. sp., una nueva specie para las costas orientales

de Sicilia

Cesare BOGI* e Stefano BARTOLINI**

Recibido el 27-X-2007. Aceptado el 15-XII-2007

RIASSUNTO

Si descrive un piccolo gasteropode planispirale, che non corrisponde a nessuna delle spe-

cie attualmente viventi nel Mar Mediterráneo, trovato in campioni di sedimento raccolti a

profonditá cómprese tra i 35 e i 52 m lungo le coste siciliane. La forma generóle della

conchiglia ci ha indotto ad ascrivere questa specie, solo prowisoriamente, al genere Xylo-

discula Marshall, 1988 anche se alcune caratteristiche della protoconca, non iperstrofica,

lo escluderebbero.

Questa specie si descrive come nuova, con il nome di Xylodiscula wareni anche se proba-

bilmente non appartiene a questo genere, evitando di descrivere al momento un nuovo

genere scarsamente definito.

ABSTRACT

A small planispiral gastropod, which could not be ascribed to any known species from the

Mediterranean or adjacent seas, was found in sediment samples collected in depths

between 35 and 52 m in the seas around Sicily.

The general shape resmbles the genus Xylodiscula Marshall, 1 988, but the protoconch ¡s

not heterostrophic and has a distinct sculpture not described from any gastropod. There-

fore we describe this new species as Xylodiscula wareni, although it probably does not

belong to the genus, instead of creating a new, poorly defined genus.

RESUMEN

Se describe un pequeño gasterópodo planiespiral recolectado entre 35 y 52 m de profun-

didad en las costas Sicilianas, que no pudo ser asignado a ninguna especie conocida del

Mediterráneo ni de mares adyacentes. La forma general se asemeja al género Xylodiscula

Marshall, 1 988, aunque la protoconcha no sea heterostrófica y tenga una escultura dis-

tinta que no se conoce en ningún otro gasterópodo. Se propone como nueva especie

Xylodiscula wareni. Aunque probablemente no pertenezca a este género, se descarta de

momento la descripción de un género nuevo escasamente definido.

PAROLE CHIAVE: Gastropoda, Xylodisculidae, Xylodiscula wareni, nuova specie, Mar Mediterráneo, recente.

KEY WORDS: Gastropoda, Xylodisculidae, Xylodiscula wareni , new species, Mediterranean Sea, recent.

PALABRAS CLAVE: Gastropoda, Xylodisculidae, Xylodiscula wareni , nueva especie, mar Mediterráneo, reciente.

* Via delle Viole 7 1-57124, Livorno. e-mail bogicesare@tiscali.it

** Via E. Zacconi,l6 1-50137, Firenze. e-mail stefmaria.bartolini@libero.it

43

Iberus , 26 (1), 2008

INTRODUZIONE

Le nostre continué ricerche malaco-

logiche e Pesame di numerosi campioni

di sedimento raccolti lungo le coste

della Sicilia Nord-Orientale e le isole

antistanti, ci ha portato, questa volta, a

studiare alcuni micromolluschi, in tutto

quattro esemplari, isolati da piccole

quantitá di detrito raccolto a profonditá

variabili tra i 35 mt. e i 52 mt.

II piccolo gasteropode di forma pla-

norbide viene, per l'aspetto generale

della conchiglia, solo provvisoriamente

attribuito al genere Xylodiscula Marshall,

1988, in quanto le caratteristiche della

sua protoconca, non iperstrofica, lo

escluderebbero .

SISTEMATICA

Ordine Heterostropha Fischer P. , 1885

Famiglia Xylodisculidae Waren A., 1992

Genere Xylodiscula Marshall,1988

Xylodiscula wareni n. sp. (Fig. 1)

Materiale esaminato: 1 esemplare (olotipo, fig.l a-e) proveniente da detriti raccolti nel Luglio 2005

alia base della "Secca dei 6 metri" presso l'isola di Filicudi a -50 m. , 1 esemplare (paratipo A) rac-

colto nel Giugno 2006 a Cannizzaro, loe. Bellatrix, -35 m.di profonditá lungo una párete rocciosa,

2 esemplari (paratipi B e C) provenienti da campioni di detriti coralligeni raccolti a Scilla a -52 m.

Materiale tipo: L'olotipo (Diam.= 1.1 mm), (Fig. 1A-E ) é stato depositato nella collezione malaco-

logia del Museo di Storia Naturale del Mediterráneo di Livorno, Italia, con il numero: Malacologia

Vol. V, n. 734. 1 paratipi sono conservad nelle seguenti collezioni: il paratipo A (Diam.= 1.0 mm),

nella collezione di S. Bartolini (Firenze), il paratipo B (Diam.= 1,3 mm) , e il paratipo C (Diam.= 1,2

mm), nella collezione E. Perna (Napoli).

Locus typicus: Detriti raccolti alia base della "Secca dei 6 metri" presso Pisóla di Filicudi a -50 m.

di profonditá.

Etimología: La specie é stata dedicata al Dottor Anders Warén, malacologo di fama mondiale e

specialista, tra l'altro, nei piccoli "Skeneimorpha".

Diagnosi (olotipo): Conchiglia piccola,

fragüe, trasparente, di forma planor-

bide.La protoconca (fig.l e) é costituita

da 0.7 giri di spira di cui la parte iniziale

é scolpita da piccole e irregolari depres-

sioni.La teleoconca é costituita da circa 2

giri di spira attraversati da deboli linee

di accrescimento. I giri si toccano solo

per un breve tratto piú o meno alia peri-

feria del giro precedente e quindi la

sutura risulta profonda e leggermente

canaliculata. Spira arrotondata con la

protoconca di poco sporgente nella

parte superiore della stessa. Alia base

della conchiglia é ben visibile per tutta

la lunghezza della teleoconca, una evi-

dente carena posizionata internamente

all'ombelico. La bocea é di forma roton-

deggiante con la parte inferiore legger-

mente espansa in prossimitá della

carena. La larghezza dell'ombelico é

circa il 30% del diámetro delPintera con-

chiglia.

Dimensioni : Diámetro^ 1.1 mm.

Opercolo e partí molli ancora scono-

sciute.

Distribuzzione: La specie sembra

avere una distribuzione ristretta a poche

localitá delle coste siciliane, ed il suo

habitat non sembra essere mai superfi-

ciale.

Discussione : L'attribuzione di questi

esemplari al genere Xylodiscula é da rite-

nersi provvisoria in quanto solo lo

studio delle partí molli potranno dame

un'inquadramento tassonomico piú

congeniale. La forma generale della con-

chiglia in realtá corrisponde alie caratte-

ristiche tipiche di questo genere, ma la

protoconca differisce in quanto non ha

uno sviluppo iperstrofico típico della

famiglia Xylodisculidae Warén, 1992.

44

Bogi E BartolinI: Xylodiscula wareni n.sp., una nuova specie per le coste della Sicilia

Figura 1. Xylodiscula wareni n. sp., Filicudi -50 m, olotipo (Museo di Storia Naturale del Mediter-

ráneo di Livorno, Coll. Malac. N° Malacologia Vol. V n. 734). A: vista dall’alto; B: vista basale; C:

vista frontale; D: dettaglio dell’ombelico; E: dettaglio della protoconca.

Figura 1. Xylodiscula wareni n.sp., Filicudi -50 m, holotype (Museo di Storia Naturale del Mediterrá-

neo di Livorno, Coll. Malac. N° Malacologia Vol. V n. 734). A: apical view; B: basal view; C: frontal

view; D: detail ofthe umbilicus; E: detall ofthe protoconch.

II genere Xylodiscula é attualmente

rappresentato nel Mar Mediterráneo da

due specie: X. boucheti Warén, Carrozza

e Rocchini, 1992, e X. leus Warén, 1992.

Warén stesso nel suo articolo (Warén,

1992) evidenzia la difficoltá nel distin-

guere le due specie che differiscono

quasi esclusivamente per il diámetro

45

Iberus, 26 (1), 2008

dell'ombelico che é il 30-33 % del diáme-

tro deH'intera conchiglia in X. boucheti

mentre in X. lens é il 40%. La caratteri-

stica principale che distingue X. wareni

da queste due specie, come dalle altre

non mediterranee appartenenti a questo

genere, oltre alia diversa protoconca, é

la presenza di una carena ben visibile

alia base della conchiglia (Fig. IB).

Abbiamo dubitato che gli esemplari

potessero appartenere ad una specie

dulciacquicola o terrestre, ma le nostre

ricerche e l'opinioni di alcuni specialisti

hanno escluso questa possibilitá.

RINGRAZIAMENTI

Un particolare ringraziamento al

Dottor Ánders Warén del Museo Svedese

di Storia Naturale di Stoccolma, per aver

eseguito le foto al SEM e per i consigli dati

nella rilettura critica dell' articolo, e all'a-

BIBLIOGRAFIA

Marshall, B.A., 1988. Skeneidae, Vitrinelli-

dae and Orbitestellidae (Mollusca: Gastro-

poda) associated with biogenic substraía

from bathyal depths off New Zealand and

New South Wales. Journal of Natural History,

22: 949-1004.

La specie tipo del genere, Xylodi-

scula vitrea Marshall, 1988, come puré X.

eximia Marshall, 1988, originarle rispetti-

vamente dell'Australia e della Nuova

Zelanda, sono state tróvate su pezzi di

legno affondati ed anche le specie

descritte da Warén per il Mar Mediterrá-

neo sono state raccolte viventi in un

biotopo caratterizzato da fibre di Posido-

nia e frammenti di legno affondati.

Nei campioni di sedimento dove noi

abbiamo trovato i pochi esemplari di X.

wareni, non erano presenti frammenti di

Posidonia.

mico Edoardo Perna per aver messo a

disposizione i suoi esemplari nonché le

foto degli stessi. All'amico Ivano Niero

per aver studiato gli esemplari come even-

tuali appartenenti alia fauna continentale.

Warén A., 1992. New and little known "Ske-

neimorph" gastropods from the Mediterra-

nean Sea and the adiacent Atlantic Ocean. Bol-

lettino Malacologico, 27 (10-12):149-248.

46

© Sociedad Española de Malaca logia

Iberus , 26 (1): 47-51, 2008

The colour white diminishes weight loss during aestivation

in the arid-dwelling land snail Sphincterochila (Albea) can -

didissima

El color blanco disminuye la pérdida de peso durante la estivación en

el caracol de medios áridos Sphincterochila (Albea) candidissima

Gregorio MORENO-RUEDA*

Recibido el 14-V-2007. Aceptado el 2-1-2008

ABSTRACT

It has been suggested that white colour is beneficial for snails living ¡n arid environments.

In this work, shell coloration in the arid-dwelling land snail Sphincterochila (Albea) can-

didissima was manipulated during aestivation. Snails painted black lost more body weight

than did control ones, presumably as a consequence of higher heat absorption. This sug-

gests that light colour is advantageous for this land snail.

RESUMEN

Se ha sugerido que el color blanco es beneficioso para los caracoles en ambientes ári-

dos. En el presente trabajo se manipula la coloración de la concha en el caracol de

medios áridos Sphincterochila (Albea) candidissima durante la estivación. Los caracoles

pintados de negro perdieron más peso que los caracoles que sirvieron como control, pre-

sumiblemente como una consecuencia de una mayor absorción de calor. Esto sugiere que

los colores claros son ventajosos para este caracol.

KEY WORDS: Sphincterochila candidissima , arid environments, colouration.

PALABRAS CLAVE: Sphincterochila candidissima , medios áridos, coloración.

INTRODUCTION

It is well established that tegument

colour has important functions in camou-

flage, as well as in communication (e.g.,

Badayev and Hill, 2000; Théry, Debut,

Gómez and Casas, 2005; Exnerova,

SVADORA, BARCALOVA, LANDOVA,

Prokopova, Fuchs and Socha, 2006).

The colour of teguments depends on the

wavelengths that are reflected. Therefore,

colouration affects the energy that is

absorbed by the tegument, the amount

being higher as the colour darkens. In

this sense, animal colouration may also

have a role in thermoregulation, espe-

cially important for arid-dwelling

animáis (Cloudsley-Thompson, 1978).

Terrestrial molluscs are very susceptible

to dehydration (Prior, 1985; Luchtel

and Deyrup-Olsen, 2001), and therefore

need adaptations to survive in arid envi-

* Konrad Lorenz Institut für Vergleichende Verhaltensforschung, Ósterreischische Academie der

Wissenschaften, SavoyenstraEe la, A- 1160, Wien (Austria) and Departamento de Biología Animal, Facultad de

Ciencias, Universidad de Granada, E- 18071, Granada (Spain).

47

Iberus , 26 (1), 2008

ronments. White colour may favour the

success of land-snails in warm environ-

ments: White shells have a higher

reflectance of sunlight (Schmidt-

Nielsen, Taylor and Shkolnik, 1971),

and, as a consequence, land snails with

light-coloured shells register lower body

temperatures (Heath, 1975), thereby

improving survival under high tempera-

tures (Richardson, 1974). This translates

as a selective pressure for whiter shells in

arid environments and, in fact, some

studies have correlated shell colour with

environment temperature (Jones, 1973;

but there are exceptions: Heller, 1984).

The snails of the genus Sphincterochila

have white shells. In Sierra Elvira (SE

Spain), Sphincterochila (Albea) candidísi-

ma (Draparnaud, 1801) is the most abun-

dant gastropod (Moreno-Rueda, 2002).

Sierra Elvira has an arid environment,

and sheltering in protective microhabi-

tats may be a vital strategy against dehy-

dration (Steinberger, Grossman, Du-

binsky and Shachak, 1983; Arad,

Goldenberg and Heller, 1989; Cook,

2001). However, S. candidísima does not

use refuges during drought periods

(spring and summer) (Moreno-Rueda,

2007; Moreno-Rueda and Collantes-

Martín, 2007). Because the shell of S.

candidísima is puré white, I hypothe-

sized that shell colour might contribute

to the survival of S. candidísima in Sierra

Elvira, explaining why this snail does

not need refuges against dehydration. I

investigated this hypothesis by manipu-

lating the shell colour of S. candidísima

during aestivation (painting some

snails), and by examining the effect of

this manipulation on the amount of

body mass lost. About 80-90% of fresh

body weight (shell not included) of

Sphincterochila land snails is water, and,

therefore, a decrease in body weight sug-

gests a loss of water (Schmidt-Nielsen

et al., 1971; Yom-Tov, 1971; Steinberg-

er, Grossman and Dubinsky, 1981).

METHODS

This study was performed in Sierra

Elvira (SE Spain, 37° 15' N, 3o 40' W), a

small mountain range with a dry

mesomediterranean climate (UNESCO,

1963). The study area undergoes five

months of drought each year, with an

averáge annual precipitation of 600-1000

mm (Alonso, López- Alcántara, Rivas

and IbáÑez, 1985). It is, therefore, a dry

zone for land snails. Table I presents cli-

matic data during the study period,

measured from the meteorological

station of Pinos Puente, about three kilo-

metres from the study area, and approx-

imately at the same altitude (630 m.

a.s.l.).

Sphincterochila candidísima is the

only species of the genus Sphincterochila

in Sierra Elvira (Ruiz Ruiz, Cárcaba

Pozo, Porras Crevillén and

Arrébola Burgos, 2006). In the study

area, this species begins aestivation in

April-May (Moreno-Rueda, 2007;

Moreno-Rueda and Collantes-

Martín, 2007). This snail adheres to

rock or vegetation during aestivation.

For manipulation, snails were not sepa-

rated from the substrate, because this

could provoke dehydration in the snails

(Luchtel and Deyrup-Olsen, 2001).

For this reason, I could not measure

body mass before treatment, but I col-

lected 75 additional individuáis in

order to analyse the relationship

between shell morphology and body

mass. The experiment started on 26

June 2005. Each individual found was

sequentially assigned to the control

group (C), to the control of manipula-

tion group (CM), or to the experimental

group (E). Manipulation in control

group was only a mark for recognition.

Snails in the CM group were painted in

yellow with a marker. Paint covered

approximately 50% of shell surface. In

the experimental group, the shell was

painted black with a marker in the

same way as in CM group. In total, 52

snails were used in each group (n =

156). The study area was prospected

two months later (26 August 2005).

Snails found were collected and mea-

sured (shell height and width) with a

calliper (accuracy 0.01 mm.) and

weighed with a digital balance (accu-

racy 0.1 g.).

48

MORENO Rueda: Colour white and aestivation weight loss in Sphincterochila candidissima

Table I. Climatic data of the meteorological station of Pinos Puente, located near of the study area,

for the study period (06/26/2005 to 08/26/2005).

Tabla I. Datos climáticos de la estación meteorológica de Pinos Puente, próxima a la zona de estudio,

para el período de estudio (2610612005 a 26/08/2005).

average weight after statistically controlling for shell height and width. The last column shows the

results of ANOVA and ANCOVA. In brackets is the standard error.

Tabla II. Peso promedio el día 26/08/2005, y altura y anchura de la concha para los caracoles en el

grupo experimental (E), control (C) y control de la manipulación ( CM). La última fila muestra el peso

promedio después de controlar estadísticamente por la altura y anchura de la concha. La última

columna muestra los resultados de los tests de ANOVA y ANCOVA. Entre paréntesis el error estándar.

The variables had a distribution

similar to normal (Kolmogorov-Smirnov

test, p > 0.05), and parametric statistics

were used. An ANOVA was used to test

the effect of the treatment on body

weight, and an ANCOVA was used con-

trolling by snail body size (height and

width). For post hoc comparisons the

Fisher LSD test was used. The Chi-

square was used to test the probability

of survival according to treatment.

RESULTS

In August, I recaptured 34 snails

alive in the control group, 30 for the CM

group, and 29 of the experimental

group. The frequency of recaptures did

not differ significantly between the three

groups (x22= 1.12; p= 0.57). When indi-

viduáis were collected in August, there

were significant differences for body

mass between the treatments (Table II).

Individuáis of the experimental group

weighed less than those in the control

one (post hoc Fisher LSD, p = 0.01),

while the average weight in the CM

group was intermediate between the

other two groups (post hoc, CM vs. C, p

= 0.22; CM vs. E, p = 0.22). In the addi-

tional sample of 75 individuáis, body

mass was strongly predicted by shell

morphology (Múltiple Regression

Model; R2 = 0.81; F 2,72 = 149.0; p < 0.001;

equation: Body mass = -5.16 (SE = 0.46;

t72 = 11.2) + 0.28 (SE = 0.03; t72 = 10.2) x

Width + 0.14 (SE = 0.03; t72 = 5.1) x

Height). There were no significant dif-

ferences for body size (height and

49

Iberus , 26 (1), 2008

width) between the three groups

(MANOVA, Wilks = 0.94; F4, ízs = 1.46; p

= 0.22; Table II), suggesting that initial

body mass did not differ among groups.

When the analyses were repeated with

shell height and width as a covariate,

differences in weight between groups

were accentuated (ANCOVA, Vi, 88 =

6.28; p < 0.003; Height effect: Fi, 88 =

10.90; p = 0.001; Width: Fi,88 = 21.98; p <

0.001; Table II). Differences between the

control group and the experimental

group increased in this analysis (post

hoc, p < 0.001), while body weight in the

CM group remained intermediate

between the other two groups (CM vs.

C, p = 0.08; CM vs. E, p = 0.08).

DISCUSSION

The findings of this experimental

study show that shell colour alteration

in Sphincterochila candidissima during

aestivation had effects on weight loss,

snails with shells painted black suffer-

ing a quicker loss of weight than control

snails. The most probable mechanism

behind this result is that light reflection

was lower in shells painted in black,

and for this reason they trapped more

heat, as shown in other studies with

other species of snails (e.g., Heath,

1975). The higher the body temperature,

the higher the water loss, decreasing

body weight (Yom-Tov, 1971). Body

size may affect the interaction between

shell colour and heat absorption

(Slotow, Goodfriend and Ward,

1993), but there were no differences in

body size between the groups, and dif-

ferences in weight remained significant

after controlling statistically for shell

morphology. This weight loss presum-

ably harms fitness, increasing the risk of

mortality, especially for the smallest

individuáis (with less reserves), or in

very dry years (Richardson, 1974).

Moreover, the treatment lasted only two

months (although the warmest), but S.

candidissima aestivates for 5-7 months in

the study area (Moreno-Rueda and

Collantes-Martín, 2007), and there-

fore, the effect should be more accentu-

ated if the entire aestivation period is

considered.

The control of manipulation (CM)

group, with shells painted in yellow,

had weight valúes intermediate to the

other two groups. In fact, this group is

not a true control of manipulation, as

colour was altered with respect to

unmanipulated snails. Because their

shells were darker than shells in the

control group, but lighter than shells in

the experimental group, the results

support that weight loss is due to shell

colour. As weight for the CM group was

intermediate, if there was an effect of

paint on weight, this cannot completely

explain the differences between the

control and the experimental group.

Therefore, results presented here

and in the literature strongly suggest

that the white colour is advantageous

for survival in arid-dwelling snails. The

question arises as to why white colour is

not more widespread in arid environ-

ments (see, for example, Heller, 1984).

Other selective mechanisms besides

thermal selection act on shell colour,

such as predation (Jones, Leith and

Rawlings, 1977). Lighter shells, espe-

cially those with a puré white colour as

in Sphincterochila candidissima , are

usually easier to detect by predators

(Reed and Janzen, 1999). However,

predation on Sphincterochila candidissima

is rare due to its thick shell (Yanes,

Suárez and Manrique, 1991), and thus

the absence of a strong selection by

predators in this species would favour

the maintenance of puré white shells.

In conclusión, this study, applying

an experimental approach, supports the

hypothesis that shell colour affects

weight loss in arid-dwelling land-snails,

and, as a consequence, fitness, by a

mechanism mediated by thermoregula-

tion.

ACKNOWLEDGEMENTS

Carlos Marfil Daza and José Luís

Ros Santaella collaborated in the field

work. David Nesbitt improved the

English.

50

MORENO Rueda: Colour white and aestivation weight loss in Sphincterochila candidissima

BIBLIOGRAPHY

Alonso, M. R., López- Alcántara, A., Rivas,

P. and IbáÑEZ, M., 1985. A biogeographic

study of Iberus gualtierianus (L.) (Pulmonata:

Helicidae). Soosiana, 13: 1-10.

Arad, Z., Goldenberg, S. and Heller, ]., 1989.

Resistance to desiccation and distribution

pattems in the land snail Sphincterochila. Jour-

nal ofZoology, 218: 353-364.

Badyaev, A. V. and Hill, G. E., 2000. Evolu-

tion of sexual dichromatism: contribution of

carotenoid- versus melanin-based coloration.

Biological Journal of the Linnean Society, 69:

153-172.

Cloudsley-Thompson, J. L., 1978. Adaptive

function of the colour s of desert animáis.

Comparative Physiological Ecology, 1: 109-120.

Cook, A., 2001. Behavioural ecology: On doing

the right thing, in the right place at the right

time. In Barker, G. M. (Ed.): The Biology ofTer-

restrial Molluscs. CAB International. Walling-

ford. Pp. 447-487.

Exnerova, A., Svadora, K., Barcalova, S.,

Landova, E., Prokopova, M., Fuchs, R. and

Socha, R., 2006. Importance of colour in the

reaction of passerine predators to aposematic

prey: experiments with mutants of Pyrrhocoris

apterus (Heteroptera). Biological Journal ofthe

Linnean Society, 88: 143-153.

Heath, D. J., 1975. Colour, sunlight and inter-

nal temperatures in the land-snail Cepaea

nemoralis (L.). Oecologia, 19: 29-38.

Heller, J., 1984. Shell colours of desert land-

snails. Malacologia, 25: 355-359.

Jones, J. S., 1973. Ecological genetics and

natural selection in molluscs. Science, 182:

546-552.

Jones, J. S., Leith, B. H. and Rawlings, P.,

1977. Polymorphism in Cepaea: a problem

with too many Solutions? Annual Review in

Ecology and Systematics, 8: 109-143.

Luchtel, D. L., Deyrup-Olsen, I., 2001. Body

wall: form and function. In Barker, G. M.

(Ed.): The Biology ofTerrestrial Molluscs. CAB

International. Wallingford. Pp. 147-178.

Moreno-Rueda, G., 2002. Selección de hábitat

por Iberus gualtierianus, Rumina decollata y

Sphincterochila candidissima (Gastropoda: Pul-

monata) en una sierra del sureste español.

Iberus, 20: 55-62.

Moreno-Rueda, G., 2007. Refuge selection by

two sympatric species of arid-dwelling land

snails: Different adaptive strategies to achieve

the same objective. Journal of Arid Enviro-

ments, 68: 588-598.

Moreno-Rueda, G. and Collantes-Martín,

E., 2007. Ciclo anual de actividad del caracol

Sphincterochila (Albea) candidissima (Dra-

pamaud, 1801) en un medio semiárido. Iberus,

25: 49-56.

Prior, D. ]., 1985. Water-regulatory behaviour

in terrestrial gastropods. Biological Reviews,

60: 403-424.

Reed, W. L. and Janzen, F. J., 1999. Natural

selection by avian predators on size and

colour of a freshwater snail ( Pomacea flage-

llata ). Biological Journal of the Linnean

Society, 67: 331-342.

Richardson, A. M. M., 1974. Differential cli-

ma tic selection in natural population of land

snail Cepaea nemoralis. Nature, 247: 572-573.

Ruiz Ruiz, A., Cárcaba Pozo, A., Porras Cre-

villen, A. I. and Arrebola, J. R., 2006. Guía

de los caracoles terrestres de Andalucía. Fun-

dación Gypaetus, Seville.

Schmidt-Nielsen, K., Taylor, C. R. and

Shkolnik, A., 1971. Desert snails: problems

of heat, water and food. Journal of Experi-

mental Biology, 55: 385-398.

Slotow, R., Goodfriend, W. and Ward, D.,

1993. Shell colour polymorphism of the

Negev desert landsnail, Trochoidea seetzeni: the

importance of temperature and predation.

Journal of Arid Environments, 24: 47-61.

Steinberger, Y., Grossman, S. and Dubinsky,

Z., 1981. Some aspects of the ecology of the

desert snail Sphincterochila prophetarum in re-

lation to energy and water flow. Oecologia, 50:

103-108.

Steinberger, Y., Grossman, S., Dubinsky, Z.

and Shachak, M., 1983. Stone microhabitats

and the movement and activity of desert

snails, Sphincterochila prophetarum. Malaco-

logical Review, 16: 63-70.

Théry, M., Debut, M., Gómez, D. and Casas,

J., 2005. Specific color sensitivities of prey

and predator explain camouflage in differ-

ent visual Systems. Behavioral Ecology, 16 : 25-

29.

UNESCO, 1963. Recherches sur la zone aride.

Etude écologique de la zone méditerraéenne. Cañe

bioclimatique de la zone méditerranéenne. Notice

explicative. UNESCO. Paris.

Yanes, M., Suárez, F. and Manrique, J., 1991.

La cogujada montesina, Galerida theklae, como

depredador del caracol Otala lactea: com-

portamiento alimenticio y selección de presa.

Ardeola, 38: 297-303.

Yom-Tov, Y., 1971. Annual fluctuations in the

water contení of desert snails. Malacological

Review, 4: 121-126.

51

1

Iberus, 26 (1): 53-63, 2008

© Sociedad Española de Malacologta

Notes on the genus Anadema H. and A. Adams, 1854 (Gastropoda:

Colloniidae)

Notas sobre el género Anadema H. y A. Adams, 1854 (Gastropoda:

Colloniidae)

James H. MCLEAN* and Serge GOFAS**

Recibido el 15-1-2008. Aceptado el 23-IV-2008

ABSTRACT

Shell morphology and characters of the living animal of the poorly known, Atlantic Moroc-

can species Anadema mocandrewii (Mórch, 1 864) are described and illustrated, based

on beach collected specimens and a single live-collected specimen. The genus ¡s mono-

typic and is assigned to the Colloniidae rather than Turbinidae because of the dome-

shaped profile of the shell, open umbilicus, symmetrical tooth rows of the radula, lack of

cephalic lappets, and the non-bicarinate ¡uvenile shell. Within the Colloniidae, it unusual

for its relatively large mature shell, ¡uvenile shell with a keeled profile, and the lack of the

secondary flap above the rachidian tooth. The species is regarded as sexually dimorphic,

with the female shell having a raised periumbilical rim comparable to that of other tro-

choideans modified for brooding by means of an enlarged umbilical cavity.

RESUMEN

Se describe e ilustra la morfología de la concha y del animal vivo de Anadema macan-

drewii (Mórch, 1864), una especie poco conocida de la costa atlántica de Marruecos. El

género es monotípico y se asigna a la familia Colloniidae, en lugar de a los Turbinidae

por la forma abombada de la concha, el ombligo abierto, las filas de dientes radulares

simétricas, la ausencia de lóbulos cefálicos y por su concha juvenil no bicarenada. Entre

los Colloniidae, la especie es insólita por el tamaño relativamente grande de la concha

adulta, el perfil de la concha juvenil con una quilla y la ausencia de un repliegue secun-

dario sobre el diente raquídeo. Se considera que existe dimorfismo sexual en esta

especie, pues la concha de la hembra tiene un reborde periumbilical elevado comparable

con el de otros trocoideos modificados para incubar en una cavidad umbilical ampliada.

KEY WORDS: Trochoidea, Morocco, brooding, endemism.

PALABRAS CLAVE: Trochoidea, Marruecos, incubación, endemismo.

INTRODUCTION

The poorly known genus Anadema been regarded as a trochid, a liotiid, a

H. and A. Adams, 1854 has at times colloniid, and a turbinid. Here we

* Natural History Museum of Los Angeles County, California 90007, USA.

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

53

Iberus , 26 (1), 2008

review the taxonomy and update what

is now known about the type and only

known species of this genus.

Anadema was proposed for the com-

bination Omphalius (Anadema) caelata A.

Adams, 1854, an Atlantic Moroccan

species then thought to be a trochid in

the absence of data on a calcified oper-

culum. The species was not to be illus-

trated until 19 years later but it was

described in sufficient detail that its

identity has never been in question. In

his remarks that followed the Latin

description of the species, Adams (1855:

39) wrote: "The character of this shell is

so peculiar, on account of the internal

spiral callus of the umbilicus, and the

absence of characters which constitute

allied forms, that I propose to consider it

a subgenus of Omphalius , under the

ñame of Anadema." The generic ñame

Omphalius Philippi, 1847 now pertains

to the Tegulinae, as defined by

Hickman and McLean (1990).

Ten years after the introduction of

Anadema , Mórch (1864: 46) reported

that a living specimen had been col-

lected at the type locality. Morch stated

that: "Omphalius (Anadema) caelata is

provided with a calcareous operculum,

which proves that this species may be

removed to Turbo. As there is already a

T. caelata, L., I propose for this species

Turbo macandrewii." Morch's placement

of the species in Turbo predated most

work on the genera of turbiniform gas-

tropods, and was based on the assump-

tion that any turbiniform species with a

calcareous operculum could only be a

Turbo. This was unfair to Arthur Adams,

but the ICZN rules about secondary

homonymy forcé us to abandon the

original ñame and to use the replace-

ment ñame. This ñame honors the re-

discoverer Robert McAndrew, who

dredged a living specimen at Mogador

(now Essaouira), the type locality of the

species.

The first shell figure known to us

was provided still later by P. Fischer

(1873), who used the replacement ñame

Turbo macandrewii; this is a drawing that

has been copied by many subsequent

authors. The operculum has never been

figured and the repository of the oper-

culate specimen examined by Morch is

unknown.

Pilsbry (1888) ignored the replace-

ment ñame and called it Leptothyra

caelata, a not unreasonable choice, as it

hardly resembles a species of Turbo. Lep-

tothyra is now assigned to Colloniidae.

Wenz (1938: 340) recognized the

genus Anadema and placed it in the

turbinid subfamily Liotiinae, under a

broad definition in which he also

included genera with calcareous oper-

cula related to Homalopoma.

Keen (1960: 270) placed Anadema in

the more restricted Homalopomatinae, a

group with fully calcified operculum,

now subsumed under the turbinid sub-

family Colloniinae in the classification

of Hickman and McLean (1990).

Nordseick (1968: 33) overlooked

Mórch (1864) and placed it in Liotiinae:

"Deckel spiralig und mit Kalkbesatz,

nicht verdickt", apparently having

assumed that it must have the opercular

definition of the now understood Lioti-

idae, in which the operculum is multi-

spiral with calcareous beads on the

outer surface.

Resolution of the uncertainty is here

provided by a preserved immature

specimen with operculum, which was

collected by the second author in 1991 at

Essaouira (formerly Mogador), the type

locality. We therefore take this opportu-

nity to illustrate the species and confirm

its placement.

MATERIALS

This report is based on material col-

lected by the second author, which is

now in the malacology collection of the

Muséum National d'Histoire Naturelle,

Paris (MNHN). The external features of

the single live-collected specimen were

drawn with the animal fully extended.

Subsequent preservation of the speci-

men resulted in retraction within the

shell. It was later critical-point dried for

SEM examination, after which it was

rehydrated for extraction of the radula

for SEM analysis.

54

McLean AND GOFAS: Notes on the genus Anadema H. and A. Adams, 1854

SYSTEM ATICS

Superfamily Trochoidea Rafinesque, 1815

Hickman and McLean (1990)

divided the family Turbinidae into a

number of subfamilies, including Lioti-

inae, Colloniinae, and Turbininae.

However, Williams and Ozawa (2007)

have reported that their data toward a

molecular phylogeny of the family

Turbinidae indicates that there are two

well-supported groups within the previ-

ously defined Turbinidae, which there-

fore preeludes the placement of all of

the subfamilies in the same family. In

view of the ongoing reconsideration of

the relationships among these groups.

we here treat the Liotiidae, Colloniidae,

and Turbinidae at the family level

within the superfamily Trochoidea,

which simply raises the ranking in the

existing classification. This has already

been adopted by McLean and Kiel

(2007). Additionally, and without dis-

cussion, Warén and Bouchet in Bouchet

and Rocroi (2005: 245) have separated

Turbinoidea and Trochoidea at the

superfamily level, which also indicates

that the higher classification for trochi-

form vetigastropods is currently unset-

tled.

Family Colloniidae Cossmann, 1916

Subfamily Colloniinae Cossmann, 1916

Hickman and McLean (1990) pro-

vided an extensive treatment of the Col-

loniidae (then as Colloniinae), distin-

guishing them from Turbinidae (then as

Turbininae) on their smaller size, non-bi-

carinate juvenile shell, lack of cephalic

lappets, symmetrical tooth rows and inner

lateral teeth that are not greatly enlarged.

This is in contrast with the Turbinidae,

characterized by larger size, bicarinate ju-

venile shell, asymmetrical tooth rows and

enlarged inner lateral teeth of the radula.

McLean and Kiel (2007) distin-

guished two subfamilies within the Col-

loniidae, based on opercular morphol-

ogy: the basal and mostly extinct

Petropomatinae Cox in Knight et al.,

1960, having a calcified operculum that

is conical on the inner surface, with a

fully multispiral pattern, and the Col-

loniinae, in which the operculum is fíat

on the inner surface with a multispiral

pattern that changes to broadly pau-

cispiral on the final volution. There is

one living genus ( Liotipoma McLean and

Kiel, 2007) of Petropomatinae, whereas

there are numerous living and fossil

genera of Colloniinae.

Genus Anadema H. and A. Adams, 1854

Anadema H. and A. Adams, 1854: 430 [as subgenus of Omphalius Philippi, 1847]. Type species

(monotypy): Omphalius ( Anadema ) caelata A. Adams in H. and A. Adams, 1854 [= Turbo macan-

drewii Morch, 1864; not Turbo caelata Linnaeus, 1758].

Anadema macandrewii (Morch, 1864) (Figs. 1-23)

Omphalius (Anadema) caelata A. Adams, in H and A. Adams, 1854: 430 [as type of new subgenus].

A. Adams, 1855: 39 [more detailed description].

Turbo macandrewii Morch, 1864: 46 [new ñame for secondary homonym Omphalius (Anadema)

caelata, not Turbo caelata Linnaeus, 1758].

Turbo macandrewi. Fischer, 1873: 98, pl. 29, fig. 3 [first illustration]. Pasteur-Humbert, 1962: 132

[listed].

Leptothyra coelata [sic]. Pilsbry, 1888: 255, pl. 48, fig. 38 [figure after Fischer],

55

Iberus, 26 (1), 2008

Turbo (Anadema) macandrewi. Pallary, 1920: 63.

Anadema coelata [sic]. Wenz, 1938: 340, fig. 795 [figure after Fischer]. Nordesick, 1968: 33, fig.

17.00

Anadema caelata. Keen, 1960: 270 [no figure]. Trew, 1992: 19 [listing of species described by H.&

A. Adams].

Material examined: BMNH, 2 syntypes, BMNH 1968183, Mogador (now Essaouira), Atlantic

Morocco (height 12.3 mm, diameter 16.6 mm; height 12.1 mm, diameter 17.3 mm). MNHN, Essaouira

(formerly Mogador), Atlantic Morocco (31° 31' N, 9o 47' W), 1 live-collected male specimen and

several beach-worn shells, leg. Gofas, 23 September 1991. MNHN, El Jadida, Atlantic Morocco (33°

16' N, 8o 29' W), beach worn shells, leg. Gofas, 26 September, 1991. MNHN, Mohammedia (for-

merly Fedala), Atlantic Morocco (33° 43' N, 7o 21' W), 10 beach-worn shells, leg. Gofas, 1970-71.

Description : Because the genus is

monotypic, the description that follows

applies both to the genus and species.

Shell composed of 5 whorls with a low

spire; profile dome-shaped, broader than

high, suture not impressed; whorls weakly

rounded, periphery of immature shell

strongly projecting, forming a keel; periph-

ery spinose in early stages, but losing

spination at maturity; mature shell with

a subangulate base; axial sculpture of fine,

raised lamellae; spiral sculpture of low,

strongly beaded cords; cords between

suture and beaded basal cord increasing

from three in juvenile to 6 at maturity;

base slightly convex, basal cords of com-

parable strength and beading to those of

body whorl, increasing from three in juve-

nile to 8 at maturity; spiral sculpture of

both body whorl and base separated by

narro w interspaces; axial lamellae well

developed in interspaces but not

expressed on surface of nodular beads;

umbilicus open in juvenile shell, closed in

male shell at basal diameter of 9 mm;

remaining open in female shell; umbilicus

of mature female shell bordered by raised,

unbeaded peripheral cord that partially

obstructs final quarter whorl and connects

directly at base of aperture; aperture

oblique, thickened within, descending

slightly on final whorl; interior nacreous,

inner wall of aperture of female shell

smooth, edge with U-shaped sinus; shell

colour brick-red.

Shell dimensions. Mature female

shell (Figs. 1-3): height 13.0, diameter

17.1 mm; immature male shell (Figs. 4-

6): height 4.9, diameter 9.1 mm;

máximum dimensions possible for male

shell unknown; immature female shell

(Figs. 7-9), height 5.9, diameter 9.1 mm.

(Right page) Figures 1-15. Anadema macandrewii (Morch, 1864). 1-3: Mature, beach-worn female

shell, from Essaouira, Morocco (MNHN), 3 views, height 13.0 mm, diameter 17.1 mm; 4-6: live-

collected, immature male specimen with operculum in place, same locality (MNHN), 3 views,

height 4.9 mm, diameter 9.1 mm; 7-9: immature, beach-worn female shell, from Mohammedia,

Morocco (MNHN), 3 views, height 5.9 mm, diameter 9.1 mm; 10, 11: juvenile beach-worn shell,

from Essaouira, Morocco (MNHN), 2 views, height 2.1 mm, diameter 5.0 mm; 12,13: juvenile

shell with protoconch showing in umbilical view, from Essaouira, Morocco (MNHN), 2 views,

diameter 2.7 mm; 14, 15: operculum of specimen shown in Figures 4-6 and 16, exterior and inte-

rior views, máximum diameter 2.1 mm.

(Página derecha) Figuras 1-15. Anadema macandrewii (Morch, 1864). 1-3: Concha adulta,

explayada, de una hembra, de Essaouira, Marruecos (MNHN), 3 vistas, altura 13,0 mm, diámetro

17,1 mm; 4-6: ejemplar macho inmaduro, recolectado vivo, con el opérculo cerrando la concha; misma

localidad (MNHN), 3 vistas, altura 4,9 mm, diámetro 9,1 mm; 7-9: concha inmadura, explayada, de

una hembra, de Mohammedia, Marruecos (MNHN), 3 vistas, altura 5,9 mm, diámetro 9,1 mm; 10,

11: concha juvenil explayada, de Essaouira, Marruecos (MNHN), 2 vistas, altura 2,1 mm, diámetro

5,0 mm; 12, 13: concha juvenil con la protoconcha visible desde el ombligo, de Essaouira, Marruecos

(MNHN), 2 vistas, diámetro 2,7 mm; 14, 15: opérculo del ejemplar representado en las Figuras 4-6 y

16, vistas exterior e interior, diámetro máximo 2,1 mm.

56

McLean AND GOFAS: Notes on the genus Anadema H. and A. Adams, 1854

57

Iberus , 26 (1), 2008

Figure 16. Anadema macandrewii (Morch, 1864). Drawing of living specimen shown in Figures 4-6.

Figura 16. Anadema macandrewii (Morch , 1864). Dibujo del animal vivo, del mismo ejemplar de las

Figuras 4-6.

Juvenile shell (Figs. 10-13) not with

raised axial lamellae, of low profile,

exposing protoconch and early whorls

in basal view; profile not equally bicari-

nate, the upper carination spinose and

more strongly projecting than lower,

non-spinose carination; protoconch and

first teleoconch whorl positioned

slightly below level of second whorl.

Operculum (Figs. 14-15) calcareous,

thick, with a concavity at the base where

the white outer surface is coloured light

reddish-brown. There is a strong ridge

and groove of similar width leading to

the upper part, which bears strong pus-

tules in the región adj acent to the col-

umella; outer edge with a narrow

groove; inner surface becoming broadly

paucispiral in final volution. This oper-

culum is chipped or somehow reduced

at its inner edge, to the extent that the

outline is not oval; the chitinous layer of

the inner surface that shows at the lower

left of Figure 15 is missing at the upper

left.

Flead-foot (Figs. 16-19). Head pro-

portionally large, provided anteriorly

with a snout terminating in a broad.

(Right page) Figures 17-23. Anadema macandrewii (Morch, 1864). 17-19. Scanning electrón

micrographs of critical-point dried specimen, same as shown in Figure 16. 17: general view of

head-foot; 18: close-up of snout; note the absence of cephalic lappets; 19: close-up of right ante-

rior bundle of epipodial tentacle. 20-23. Scanning electrón micrographs of the radula, same speci-

men as Figure 16. 20: complete radula; 21: close-up of several complete rows; 22: detail of central

and lateral teeth; 23: detail of marginal teeth. Abbreviations, ct: cephalic tentacle; et: epipodial

tentacles; f: foot (pointing to propodium); sn: snout.

(Página derecha) Figuras 17-23. Anadema macandrewii (Morch, 1864). 17-19. Micrografías elec-

trónicas de barrido del ejemplar de la Figura 16 deshidratado por punto crítico. 17: vista general de la

cabeza y del pie; 18: vista aumentada del hocico; nótese la ausencia de lóbulos cefálicos; 19: vista

aumentada de un haz anterior derecho de tentáculos epipodiales. 20-23. Micrografías electrónicas de

barrido de la rádula, ejemplar de la Figura 16. 20: radula completa; 21: vista aumentada de algunas

filas completas; 22: detalle de los dientes centrales y laterales; 23: detalle de los dientes marginales. Abre-

viaturas, ct: tentáculo cefálico; et: tentáculos epipodiales; f: pie (señalado el propodio); sn: hocico.

58

McLean AND GOFAS: Notes on the genus Anadema H. and A. Adams, 1854

59

Iberusy 26 (1), 2008

flattened area surrounding the mouth,

laterally with two prominent, smooth

bulges containing rather large, black

eyes and, next to these on the anterior

side, two slender, villose cephalic tenta-

cles; cephalic lappets lacking. Neck

tobes present, rather symmetrical, undi-

vided. Foot rather small, less than half

the diametre of the shell when

extended; each side of the epipodium

provided in its anterior part with a

cluster of three tentacles, the foremost

on hardly more than a rounded bulge,

the next two tapering and villose,

hardly one-tenth of the cephalic tenta-

cles in size; in the posterior part with

two more tentacles similar in size and

shape to the latter.

Radula (Figs. 20-23) strikingly sym-

metrical, with broad rachidian having

lateral extensions, and moderately long,

tapered overhanging cusps; with four

pairs of similar lateral teeth, which are

elbowed like the rachidian and extend

above the shaft of the next tooth; shaft

of fifth lateral tooth not in cióse contact

with shaft of fourth lateral tooth; this

tooth broad on both sides, with a less

prominent cusp than those of the inner

lateral teeth or any of the marginal

teeth; innermost pair of marginal teeth

with short overhanging cusps, followed

by four pairs of marginal teeth with

longer cusps; outer marginal teeth with

longer shafts, overhanging cusps shorter

and deeply serrate on sides of the cusps.

The live-collected specimen (Figs. 4-

6, 16) reported here (diameter 9.1 mm)

has a nearly closed umbilicus and a gen-

erally unmodified base, with no indica-

tion of the incipient formation of the

DISCUSSION

Knowledge of the genus Anadema

has been slow to develop because the

species A. macandrewii lives in the sublit-

toral zone on rocky bottoms exposed to

strong surf, where there have appar-

ently been few efforts at collecting by

diving due to the exposure, low visibil-

ity, and extensive muddy bottoms off-

shore in relatively shallow water. The

projecting rim of mature female shells; it

is therefore identified as a male speci-

men. There are immature beach-worn

shells of about 10 mm in diameter (Figs.

7-9), which are considered to be female

shells, having the beginning of a project-

ing rim that will form the strong peri-

umbilical angulation of the large female

shell (Fig. 2). It is evident that male

shells are much less frequent among the

beach-worn shells. Because the male

shell with an operculum is larger than

any of the comparable beach-worn

shells considered to be male shells, and

because the lip is immature, there is no

indication from the material at hand as

to the possible size reached by male

shells. It may be that male shells can

reach a size similar to that of female

shells. If so, they would differ from

female shells in having a rounded

umbilical wall, rather than the project-

ing umbilical rim of female shells.

Both male and female shells seal the

umbilicus upon attaining a half-grown

diameter of about 10 mm. The female

shell then proceeds to form a secondary

umbilicus for the brooding cavity. The

initial sealing of the umbilicus may help

to protect the shell from exposure due to

erosión of the apical whorls, which is

also avoided in most marine gastropods

by shell deposition of a plug from

within.

There is variation in the peripheral

spination of immature stages. The

spination of the live-collected male

specimen (Figs. 4-6) is stronger than that

of any of the beach-worn shells of

similar size, whether identified as male

or female.

single live-collected specimen reported

here from the intertidal zone is an

unusual record for the species.

Systematic position: There is now no

doubt that Anadema should be assigned

to the family Colloniidae, on the basis of

shell and opercular characters, charac-

ters of the external anatomy, and the

radula. The fine lamellar sculpture had

60

McLean AND GOFAS: Notes on the genus Anadema H. and A. Adams, 1854

brought to mind a comparison with

Liotiidae, but that is ruled out by the

calcareous operculum.

There are many possibilities for

opercular morphology in the Colloni-

idae, just as there are in the Turbinidae

(see Vermeij and Williams, 2007).

There is no comparative work on the

colloniid operculum, but the operculum

of Anadema is within the range of possi-

ble expression for the family. Immature

shells of the Mediterranean turbinid

species Bolma rugosa (Linnaeus, 1767)

have a resemblance to Anadema macan-

drewii in having a somewhat similar

operculum and a spinose peripheral car-

ination, but mature specimens of Bolma

are much larger, have a higher profile

and an impressed suture, a closed

umbilicus and an expansive columellar

callus that forms a columellar shield

nearly as broad as the aperture. For a

review of Bolma , see Beu and Ponder

(1979). Anadema macandrewii differs from

all illustrated species of Bolma in its low,

dome-shaped profile and open umbili-

cus at maturity.

The keeled early juvenile shells indi-

cate that early sculpture is not evenly

bicarinate, which is a defining feature

for Turbinidae, according to the

restricted definition provided by

Hickman and McLean (1990: 55) at the

previously recognized subfamily level.

Most colloniids have the juvenile shell

with even spiral cords, but there are

exceptions to that generalization. The

keeled early stage brings to mind the

recently described Liotipoma McLean

and Kiel, 2007, in the basal subfamily

Petropomatinae, which also has an early

keel.

The lack of cephalic lappets is consis-

tent with the assignment to Colloniidae,

in which lappets are lacking (Hickman

and McLean, 1990). This is in contrast

to the Turbinidae, in which the lappets

are well-developed (Hickman and

McLean, 1990), albeit they are small in

the Mediterranean Bolma rugosa.

The radular morphology of Anadema

provides convincing evidence that it is

colloniid rather than turbinid because

the tooth rows are perfectly symmetri-

cal, and the inner margináis are not

greatly enlarged. In turbinids, the asym-

metrical tooth row causes the tooth

alignment to be skewed, because the

large inner laterals must altérnate in

zipper fashion when the radula is longi-

tudinally folded (as detailed by

Hickman and McLean, 1990).

Anadema is highly unusual within

Colloniidae for its large size and its

keeled early stage. With its máximum

shell diameter of 17 mm, it may well be

the largest known colloniid. All of the

colloniid radulae illustrated by

Hickman and McLean (1990) have a

secondary flap that projects above the

rachidian tooth; the radula of Anadema

is unusual in not having the secondary

flap. In addition, the morphology of the

fifth lateral of Anadema seems also to be

unusual for the family.

Larval brooding: Brooding of larvae

within the umbilical cavity has been

broadly reported among the Trochoidea

(Hickman, 1992: 254). In the Trochidae

it is known in Margantes vorticiferus, as

shown by Lindberg and Doberteen

(1981). In Liotiidae, it has been reported

for Arene socorroensis by Shasky (1968)

and again by Hertz (1998), and for

"Munditia" subquadrata it has been

reported by Burn (1976). In the Colloni-

idae, it is known in the recently

described genus Liotipoma, as reported

by McLean and Kiel (2007). Its occur-

rence in Anadema is therefore the second

known example. In each of these cases,

the umbilical rim of the female shell is

raised to increase the volume of the

umbilical cavity. Such a modification

can be the only explanation for the

strongly raised umbilical rim of what

we interpret as the female shells of

Anadema macandrewii.

The size reached by the shells of

males remains to be discovered; it is

possible that mature male shells are

smaller than female shells. In support of

that possibility, the operculum illus-

trated here shows the expansión of the

final volution that is characteristic of a

mature operculum. However, this shell

is somewhat immature because the final

lip is not thickened. All beach-worn

61

Iberus, 26 (1), 2008

male shells in the material on hand

seem to be immature. The immature

male shells are represented by fewer

specimens than the larger shells attrib-

uted to, for reasons unknown.

ACKNOWLEDGMENTS

We thank Philippe Bouchet and Vir-

ginie Heros of the MNHN for arranging

the loan of the Anadema specimens.

Illustrations of shells (from photos by

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Iberus, 26 (1): 65-68, 2008

© Sociedad Española de Malacología

A new species of Mangelia (Turridae: Mangeliinae) from the

Mediterranean Sea

Una nueva especie de Mangelia (Turridae: Mangeliinae) del

Mediterráneo

Charles CACHIA* and Constantine MIFSUD**

Recibido el 17-VII-2007. Aceptado el 12-III-2008

ABSTRACT

A new species Mangelia melitensis nov. sp., from Malta, is introduced and compared with

other similar congeneric species.

RESUMEN

Se describe una nueva especie Mangelia melitensis, de Malta, y se compara con especies

congenéricas de aspecto similar.

KEY WORDS: Mollusca, Conoidea, Turridae, Mangeliinae, Mangelia nov. sp., Mediterranean.

PALABRAS CLAVE: Mollusca, Conoidea, Turridae, Mangeliinae, Mangelia nov. sp., Mediterráneo.

INTRODUCTION

The genus Mangelia is represented in

the Mediterranean Sea by about 28

species (CLEMAM 2007). The shells are

slender, fusiform in shape, with a sculp-

ture consisting of axial, and often spiral

sculpture. The siphonal canal is short

and open and the columella is smooth

without any processes. The mollusc has

a wide foot, long tentacles with eyes sit-

uated on the sides of the stalks and at

about 2/ 3 their height, a long siphon and

the body is mostly of a transparent

white colouration, but sometimes with

opaque coloured streaks or blotches,

especially on the siphon. The radula

consists of hollow marginal teeth, rarely

barbed, with an irregular basal swelling

(McLean, 1971). The animal lacks an

operculum. The diet consists mainly of

* 1, Alley 1, St. Catherine Street, Qormi QRM, Malta

** 5, Triq ir-Rghajja, Rabat RBT 2486, Malta

polychaete worms, sometimes larger

than the mollusc itself. (C. M. pers. obs.)

Certain authors (Bouquoy,

Dautzenberg and Dollfus, 1883;

Powell, 1966; van Aartsen and Fehr

de Wal, 1978) have proposed that the

Mediterranean species classification

should be separated into two genera,

Mangelia (Risso, 1826) type species Man-

gelia attenuata (Montagu, 1803) and

Mangiliella (B.D.D., 1826) type species

Mangiliella multilineolata (Deshayes,

1836). The proposition is based on the

type of development of the protoconch.

Mangelia has a multi-spiral or plank-

totrophic type of protoconch, usually

consisting of 2 l/i or more whorls while

Mangiliella has a paucispiral or direct

type of protoconch consisting of 1 1 /lio

65

Iberus, 26 (1), 2008

2 whorls. Although this arrangement is

very convenient for separating the

species, Bouchet (1990) advised against

such methods for creating genera and

subgenera in this already problematic

family.

The Mediterranean species have

been dealt with by several authors

(Nordsieck, 1977; van Aartsen and

Fehr de Wal, 1978; Sabelli, Giannuzzi-

-Savelli and Bedulli, 1990; Cachia,

Mifsud and Sammut, 2001; Gruppo

Malacologico Livornese and Gruppo

Malacologica Romagnolo, 2005).

During our ongoing research on the

marine mollusca which inhabit the

coasts of the Maltese Islands, we have

come across many specimens of a dis-

tinct Mangelia species which is new to

Science. It will be introduced herein.

SYSTEMATICS

Superfamily Conoidea Vaught, 1989

Family Turridae Swainson, 1840

Subfamily Mangeliinae Vaught, 1989

Genus Mangelia Risso, 1826

Mangelia melitensis nov. sp. (Figs. 1, 2)

Material studied: All studied material is from the Maltese Islands. Holotvpe. collection number

BMNH 20070312 (H= 6mm, W= 2.5mm), and two paratypes, collection number BMNH

20070313, off Gnejna Bay, in sand and algae from 40 m., x-2006, The Natural History Museum,

London. Three paratypes, off Ras il-Wahx, 35 m., ix-1992, collection number MNHN 9988,

Muséum National d'Histoire Naturelle, París. Three paratypes, off Rdum id-Delli, 50 m., ix-2006,

collection number. TAU MO 57001, National Collections of Natural History, Department of

Zoology, Tel Aviv University. Four paratypes, off Ras il-Wahx, 25-30 m., ix-1992, and three

paratypes, St Thomas Bay, 3-4 m., viii-1987, National Museum of Natural History (Mdina,

Malta). Other Paratypes: Salina Bay, 4 m viii-2000, 21 shells; off Rdum id-Delli, 60 m, viii-2006, 5

shells; off Cirkewwa Point, 34 m, viii-1996, 14 shells, in prívate collection of Charles Cachia. [Coll.

No 1185]. Off St. Paul's Bay, 40-50 m., ix-1986, 3 shells; Qammieh, Cumnija, 50 m., viii-1986, 4

shells; St. Thomas Bay, 3-4 m., viii-1987, 11 shells; Bahar ic-Caghaq, beached, i-1991, 5 shells;

Gozo Island, Dwejra, Coral cave, in sand, 25 m., 1 shell, 9-vi-2001 (leg. Anthony Sammut); off

Golden Bay, 40 m, iii-1991, 9 shells; off Ras il-Wahx, 25-30 m., ix-1992, 31 shells off Rdum id-

Delli, 50 m., ix-2006, 9 shells, in prívate collection of Constantine Mifsud [coll. No. M1601]. Off

Ras il-Wahx, 25-30 m., ix-1992, 2 shells, in prívate collection of Charles Sammut. Salina Bay, 4 m.,

viii-1998, 2 shells (C. Cachia leg.), in prívate collection of Gert Lindner.

Type locality: Gnejna Bay, Maltese Islands.

Etymology: Named after the type locality.

Description : Shell small, rather fragüe,

fusiform, typical of the genus. Proto-

conch consists of 2 1 ¡i, rounded, trans-

parent white, glassy whorls, the last with

a few small, close-set axial riblets crossed

by spiral micro-striae. Teleoconch con-

sists of up to four whorls which are sub-

angulated at their adapical third and

curving regularly towards the suture.

Sculpture of 8-9 narrow, flexuous, slight-

ly opisthocline, axial ribs which continué

to the base. Their interspaces are about

twice as wide. The whole surface of the

shell is covered with dense micro-spiral

striae of about the same thickness and

close-set axial growth lines, producing

an iridescent pattern. There are about 45-

50 of these micro-spiral striae at the

penultimate whorl. There are also 5-6

well spaced, thicker spiral chords at the

base of the shell, near the siphonal canal.

Aperture about half height of shell. Out-

er lip sharp, thin and fragüe, thickened

internally, with its topmost part clearly

curved. Lip varix present in some speci-

mens. Sinus shallow. Columella smooth.

66

Cachia AND MlFSUD: A new species of Mangelia from the Mediterranean Sea

Figure 1. Mangelia melitensis nov. sp., shells. A: holotype from off Gnejna Bay, in sand and algae,

40 m (height 6 mm); B: paratype from the type locality (height 6 mm); C, D: paratype from off

Ras il-Wahx, 25-30 m (height 6.1 mm).

Figura 1. Mangelia melitensis nov. sp., conchas. A: holotipo, frente a Gnejna Bay, en arena y algas, 40

m (altura 6 mm); B: paratipo de la localidad tipo (altura 6 mm); C, D: paratipo, frente a Ras

il—Wahx, 25-30 m (altura 6,1 mm).

Siphonal canal short and wide. The shell

colour is white or beige. Rarely, some

specimens are decorated with four, faint,

brownish spiral bands or with tiny,

brown sub-sutural blotches on the

penultimate whorl, while others have a

dark blotch at the middle of the exterior

of the outer lip. A single specimen is

completely brown.

DISCUSSION

M. melitensis nov. sp. had already

been introduced as Mangelia sp. A. in

Cachia et al. (2001). It differs from all

the other Mediterranean forms of the

M. unifasciata (Deshayes, 1835)

complex, to which it is very similar, by

the lack of bold spiral cords on its

whorls. It is similar in this respect to M.

costulata Risso, 1826 [= M. smithii

(Forbes, 1840)], but in that species the

spiral striae are fewer, flatter, much

bolder, and continuous, not crossed by

growth lines, while their interspaces

are deeply incised. Mangelia costulata

never forms a thickened rib on the

The animal is transparent white with

large black eyes at about two-thirds of

the height of the tentacle stems. The foot

has opaque white spots, while the

siphon has small opaque yellowish

spots. There is no operculum.

The species is found near or in Posi-

donia oceánica (Linnaeus) Delile, substra-

tum at depths of 3 to 50 metres.

aperture, unlike the present species:

Moreover, in M. costulata the last proto-

conch whorl has a bold reticulate type

of sculpture, whilst in M. melitensis nov.

sp. this whorl has only close-set flex-

uous axials with spiral micro-striae. M.

costulata usually has a continuous wide

brown band on the ultimate whorl and

the animal has a bright orange spotted

siphon.

Mangelia melitensis nov. sp., also

offers similarities to Mangelia costata

(Donovan, 1804). In that species

however the whorls are very evenly

convex not somewhat subangulated as

67

ikrus, 26 (1), 2008

Figure 2. Mangelia melitensis nov. sp., scanning electrón micrographs, same paratype as Figures

1C-D. A: protoconch; B: detail of the sculpture on a spire whorl.

Figure 2. Mangelia melitensis nov. sp., vistas en microscopía electrónica, mismo paratipo que las Figura

1 C-D. A: protoconcha; B: detalle de la escultura en una vuelta de la espira.

in Mangelia melitensis nov. sp. The shell

is also smaller, rather glossy, differently

coloured, of a more fusiform aspect and

with a slightly different protoconch. The

animal is completely white.

Distribution: The Maltese Islands.

The authors have not come across any

specimens of M. melitensis nov. sp. from

any other Mediterranean locality.

BIBLIOGRAPHY

Buquoy, E., Dautzenberg, P. and Dollfus,

G., 1882-1886. Les Mollusques marins du Rous-

sillon. I: Gastropodes. París, Bailliére, 570 pp.

Bouchet, P., 1990. Turrid genera and mode of

development: The use and abuse of proto-

conch morphology. Malacologia, 32 (1): 69-77.

Cachia, C., Mifsud, C. and Sammut, P., 2001.

The Marine Mollusca of the Maltese Islands,

Part 3 Neogastropoda, Backhuys Publishers,

266 pp.. Leiden.

CLEMAM, Unitas Malacologica; Checklist of

European Marine Mollusca: Web Page:

http: / / www.somali.asso.fr /clemam/bio-

taxis.php [accessed 5-vi-2007]

Gruppo Malacologico Livornese and

Gruppo Malacologico Romagnolo, 2005.

Nota sulle Mangelia Mediterranee. Notiziario

S.I.M., 23 (9-12): 22-33.

MacLean, J. H., 1971. A revised classification

of the Family Turridae, with the proposal

of new Subfamilies, genera and subgenera

from the eastem Pacific. The Veliger, 14 (1):

114-130.

ACKNOWLEDGEMENTS

We would like to thank our colleague

P. Sammut (Rabat, Malta) for revising the

manuscript and G. Lindner (Germany) for

his kind disposition. The scanning elec-

trón micrographs for this paper were pro-

duced at the University of Málaga with

the help of Gregorio Martín Caballero.

Mifsud, C., 1998. The molluscan species living

in Maltese Posidonia meadows. La Conchiglia,

30 (287): 37-48, 61.

Nordsieck, F., 1977. Turridae of the European

Seas. "La Pirámide" for La Conchiglia, 131 pp.

Rome.

Nordsieck, F., 1982. Die europaischen Meeres-

Gehauseschnecken. 2nd edition. 539 pp. Gus-

tav Fischer, Stuttgart

Powell, A. W. B. 1966. The Molluscan families

Speightiidae and Turridae. Bulletin of the

Auckland Institute and Museum, 5, 184 pp. +

23 plates. New Zealand.

Sabelli, B., Gianuzzi-Savelli, R. and Bedulli,

D., 1990. Catalogo annotato dei molluschi

marini del Mediterráneo, S. I. M. Librería Nat-

uralística Bolognese, Vol. 1. xiv, 1-348. Bologna.

van Aarsten, J. J. and Fehr de Wahl, M.C.,

1978. The sub family Mangeliinae Fischer,

1887 in the Mediterranean. Conchiglie, 14 (3-

6): 97-110.

68

© Sociedad Española de Malacologí

Iberas , 26 (1): 69-80, 2008

Catálogo de los bivalvos marinos del sector central del Golfo

de Valencia (España)

Checklist of the marine bivalves in the central sector of the Gulf of

Valencia (Spain)

Jorge C. TAMAYO GOYA*

Recibido el 12-11-2007. Aceptado el 31-III-2008

RESUMEN

Se presenta una lista con 1 84 especies de Bivalvos marinos citados en la literatura y rea>

lectados por el autor entre 1998 y 2008, en la costa del Golfo de Valencia, sector cen-

tral: 39° 42' N, 0o 12' W (Playa de Canet de Berenguer, Sagunto) - 38° 55' N, 0o 03#

W (Playa de Oliva), Mediterráneo occidental. Cari fervensis se cita por primera vez en ¡a

zona y se confirma la existencia de poblaciones establecidas de Eastonia rugosa, Fulvia

fragilis y Lentidium mediterraneum.

ABSTRACT

A list is presented with 1 84 species of marine Bivalves cited in the literature or collected

by the author between 1 998 and 2007, along the coast of Gulf of Valencia, central sec-

tor: 39° 42' N, 0o 12' W (Playa de Canet de Berenguer, Sagunto) - 38° 55' N, 0o 03'

W (Playa de Oliva), Western Mediterranean. Gari fervensis is cited for the first time in the

area and the occurrence of established populations is confirmed for Eastonia rugosa, Ful-

via fragilis and Lentidium mediterraneum.

PALABRAS CLAVE: Catálogo, moluscos, bivalvos, Golfo de Valencia, Mar Mediterráneo.

KEY WORDS: Checklist, molluscs, bivalves, Gulf of Valencia, Mediterranean Sea.

INTRODUCCIÓN

En este trabajo se presenta una actua-

lización de la lista faunística de moluscos

bivalvos de la Bahía de Valencia. Estos

complementan un catálogo reciente-

mente publicado (Oliver Baldoví, 2007)

donde se aporta una actualización de

Gasterópodos marinos testáceos de la

zona sur del Golfo de Valencia.

La gran mayoría de las conchas reco-

lectadas son explayadas, por lo que no se

puede indicar su hábitat originario. En

mucha menor medida, se han obtenido

buceando a pulmón-con tubo (entre -2 y

-3 m) o recogidas por una embarcación

"marisquera" faenando a escasos metros

de la costa (zona infralitoral). Las recogi-

das de muestras se han ido realizando a

lo largo de todas las estaciones entre los

años 1998 y 2008 y no se ha seguido una

metodología concreta.

Los sustratos sedimentarios que

constituyen el hábitat de las comunida-

des presentes en el piso infralitoral son,

las correspondientes a: arenas finas

* C/Polo y Peyrolón, 37, izda 16, 46201, Valencia.

69

Iberus , 26 (1), 2008

Figura 1 . Localidades de muestreo en el Golfo de Valencia.

Figure 1. Sampling sites in Gulf of Valencia.

superficiales, arenas finas bien calibra-

das (entre -2/ -3 m. y -20/ -25 m.), arenas

finas fangosas (entre -20/-25 m. y -35/-

40 m.), arenas con proporciones peque-

ñas de finos (<15%) (consideradas como

transición entre ambos sustratos) y los

fondos detríticos.

El sustrato rocoso está presente en

mucha menor medida, lo mismo que las

praderas de Posidonia oceánica (en

mucha mayor proporción de mata

muerta que degradada, y que suele

extenderse en una distribución batimé-

trica entre -5 m y -20 m), así como la

Cymodocea nodosa y la Caulerpa prolifera.

Las arenas gruesas y gravillas, ocupan

franjas a profundidades variables y

sujetas a corrientes marinas.

A continuación en profundidad, en

la plataforma continental, está el piso

circalitoral (entre -40 m. y -100 m.), de

escasa pendiente, en el que están pre-

sentes los fondos detríticos costeros. Los

fondos fangosos terrígenos (a partir de -

100/-150 m.) se extienden por zonas

muy amplias del fondo marino.

El litoral en estudio, eminentemente

sedimentario, se caracteriza por la suce-

sión casi continuada de playas de arena

fina, sólo interrumpidas por las instala-

ciones portuarias, las desembocaduras

fluviales (áreas de fondos fangosos) y el

Cabo de Cullera (acantilado mediano)

Las poblaciones en cuyas playas se

han efectuado la recogida de ejemplares

son colindantes entre sí y han sido agru-

padas, de norte a sur, en las siguientes

subzonas (Fig. 1), siendo la B y la D en

donde mayor número de muéstreos se

han realizado:

A: (Canet d'En Berenguer), Sagunto,

Puzol y El Puig. Predominan las arenas

finas bien calibradas, en mucha menor

medida las de arenas finas fangosas y

las extensiones de mata muerta de Posi-

donia. El puerto de Sagunto y el puerto

deportivo de Canet de Berenguer, la

planta siderúrgica de Sagunto, el emisa-

rio de Canet de Berenguer, y la desem-

bocadura del río Palancia, son agentes

determinantes en la configuración sedi-

mentaria de esta zona del litoral.

70

Tamayo GOYA: Catálogo de los bivalvos marinos del sector central del Golfo de Valencia

B: Puebla de Farnals, Massamagrell,

Meliana, Alboraya y las playas al norte

de Valencia. Las arenas finas fangosas

predominan sobre las arenas finas bien

calibradas. Es de mencionar el sustrato

rocoso frente al Puerto de Valencia y las

extensiones de mata muerta de

Posidonia, en mayor superficie que las de

la subzona A. Él puerto deportivo de

Port sa Playa, el Barranco de Carraixet y

el emisario de Vera (al norte del Puerto

de Valencia), son los agentes de esta

subzona.

C: Playas al sur de Valencia. Hay

mucho mayor predominio de las arenas

finas bien calibradas sobre las arenas

finas fangosas. El Puerto de Valencia, la

desembocadura del río Turia, el emisa-

rio del sur del Puerto y las Golas del

Pujol y de El Perellonet (provenientes de

La Albufera), son los agentes a destacar.

D: Sueca y Cullera . Es la subzona

con mayor proporción de arenas finas

bien calibradas y con menor de finas

fangosas en el área de estudio. A

reseñar también, la presencia de mata

muerta de Posidonia en el afloramiento

rocoso frente a Cullera. Cabe mencionar

en esta subzona, la desembocadura del

Jucar y las Golas de El Perelló y la del

Estany.

E: Tavernes de la Valldigna, Xeraco,

Xeresa, Gandía, Daimuz, Guardamar,

Miramar, Piles y Oliva. Las arenas finas

fangosas predominan sobre las finas

bien calibradas. Frente a Gandía existe

otro enclave rocoso con presencia de

Posidonia. Los puertos de Gandía y

Oliva, constituyen lo más destacable

como agentes determinantes.

SISTEMÁTICA

En la Tabla I se presenta el listado de

especies encontradas en el área de

estudio, junto con el número de especí-

menes encontrados, sus sectores de dis-

tribución y citas bibliográficas. Esta lista

ha sido ordenada de acuerdo con la cla-

sificación propuesta por CLEMAM.

Check List of European Marine

Mollusca: chttp:// www.somali-

asso.fr/ demarro .

DISCUSIÓN

Glycymeris violacescens (Lamarck, 1819)

Es una especie muy presente en gran

parte del litoral de Valencia que con fre-

cuencia se muestra con un polimorfismo

y policromía muy característicos (Fig. 2),

predominando el contorno de la concha

menos subcuadrangular, muchas veces

más oblicua; el color gris negruzco-

azulado, principalmente en la mitad

superior y umbo, combinado con los

colores tierra (beige y marrón) y con

mucha mayor superficie de mancha en

el interior de la valva; generalmente, los

umbos más separados y con mayor área

cardinal; cohabitando en proporciones

similares con los ejemplares más

comunes-tipo, más frecuentemente

divulgados (Fig. 3).

Fulviafragilis (Forsskal in Niehbur, 1775)

Se confirma (Fig. 4) la presencia en la

zona de esta especie, previamente citada

en Cullera por Zenetos et al. (2004) a

partir de ejemplares recolectados en

1991. Ello indica que esta especie está

formando poblaciones perennes en la

zona, al igual que sucede en otras partes

del Mediterráneo (Crocetta, 2005;

Vardala-Theodorou , 1999).

Eastonia rugosa (Helbling, 1779)

Especie presente ocasionalmente en

este sector del Golfo de Valencia, salvo en

la subzona B en la que si es habitual

hallarla. En la playa de Meliana (subzona

B) es donde se concentra la gran mayoría

de los individuos de esta especie encon-

trados en esta zona del litoral (Fig. 5).

Decenas y decenas de conchas de indivi-

duos jóvenes y adultos son depositadas

todos los años en esta playa. En esta

subzona B, también se localiza una impor-

tante concentración de ejemplares, aunque

en menor medida, de Lutraria magna (da

Costa, 1778) y Venericardia antiquata (Linné,

1758). Es de notar que esta especie mani-

fiesta una tendencia a ampliar su exten-

sión en el Mediterráneo, siendo reciente-

mente observada en abundancia en las

costas italianas en localidades donde no

se conocía anteriormente (La Valle, Vani,

Liboni y Smriglio, 2007).

71

Iberus, 26 (1), 2008

Tabla I. Listado de especies encontradas en el área de estudio. Ej: concha recolectada explayada;

Ej*: entre 1 y 3 ejemplares recolectados; Ej*: entre 1 y 3 ejemplares recolectados, deteriorados; L:

especie presente en la colección de referencia del laboratorio de Biología Marina de la Universidad

de Valencia (1982-1989, García Carrascosa, no publicado) o citada en la bibliografía reseñada, sin

ejemplar recolectado por el autor. Las letras (ABCDE) corresponden a los sectores de la Figura 1

en los que se han recolectado ejemplares. Las referencias a las citas se indican como sigue: (1)

Colección de referencia del laboratorio de la Universidad de Valencia; (2) ApáRICI SEGUER,

Rowland, Taylor y García Carrascosa, 1996; (3) Bonnin y Rodríguez Babio, 1990; (4)

Costa, García Carrascosa, Monzó, Peris, Stubing y Valero, 1984; (3) Generalitat

Valenciana, Conselleria de Agricultura y Medio Ambiente, 1994-1997; (6) Giner

Ponce, 1989; (7) Hidalgo, 1917; (8) Montero Agüera, 1971; (9) Ramón Herrero, 1993;

(10) ROSELLÓ, 1910; (11) SÁNCHEZ Diana, 1980. Los nombres subrayados corresponden a las

especies que están comentadas en este trabajo.

Tal? le I. List of the species found in the studied urea. Ej: shell collected on the beach; Ej*: 1 to 3 speci-

mens found; Ej*: 1 to 3 specimens found, damaged; L: species found in the reference collection deposited

in the Marine Biology laboratory ofthe University of Valencia (1982-1989, García Carrascosa, unpu-

blished data) or cited in literature, no specimens collected by the author. Letters (ABCDE) refer to

Figure 1 sectors. Literature cited as follows: (1) reference collection deposited in the laboratory ofthe

University of Valencia; (2) APARICI SEGUER, R.OWLAND, TAYLOR AND GARCIA CARRASCOSA, 1996;

(3) Bonnin y Rodríguez Babio, 1990; (4) Costa, García Carrascosa, Monzó, Peris,

Stubing and Valero, 1984; (5) Generalitat Valenciana, Conselleria de Agricultura y

Medio Ambiente, 1994-1997; (6) Giner Ponce, 1989; (7) Hidalgo, 1917; (8) Montero

Agüera, 1971; (9) Ramón Herrero, 1993; (10) Roselló, 1910; (11) Sánchez Diana, 1980.

Underlined ñames are discussed in this paper.

72

Tamayo Goya: Catálogo de los bivalvos marinos del sector central del Golfo de Valencia

Tabla I. Continuación.

Table I. Continuation.

73

Iberus , 26 (1), 2008

Tabla I. Continuación.

Table I. Continuation.

74

TAMAYO GOYA: Catálogo de los bivalvos marinos del sector central del Golfo de Valencia

Tabla I. Continuación.

Table I. Continuation.

75

Iberus, 26 (1), 2008

Tabla I. Continuación.

Table I. Continuation.

76

Tamayo Goya: Catálogo de los bivalvos marinos del sector central del Golfo de Valencia

Tabla I. Continuación.

Table I. Continuation.

Gari fervensis (Gmelin, 1791)

Esta epecie tampoco está citada en

los trabajos anteriormente relacionados,

pero también habita de forma perenne

en la Bahía de Valencia (Fig. 6). No es

frecuente su presencia en el litoral exa-

minado, aunque tampoco lo es de forma

ocasional. Se pueden localizar, año tras

año, individuos jóvenes y adultos en las

mismos emplazamientos y con similar

frecuencia y abundancia que su congé-

nere, Gari depressa (Pennant, 1777).

Lentidium mediterraneum (Costa O.G.,

1829)

Diversos trabajos escalonados en el

tiempo, (recientes: Aparici et al., 1989;

García Carrascosa, 1988; y menos

reciente. Montero Agüera, 1971)

además de los ejemplares localizados

por el autor en Puebla de Farnals, Valen-

cia, Sueca y Cullera, entre 2004 y 2007

(Fig. 7), evidencian la presencia no de

forma esporádica, de esta especie a lo

largo de todo el litoral en estudio.

AGRADECIMIENTOS

Mi más sincera gratitud y reconoci-

miento por su generosa y desinteresada

colaboración, a Celso Rodríguez Babío y

Manuel García Carrascosa, ambos de la

Facultad de Ciencias, Universidad de

Valencia, por su permanente ánimo,

apoyo y por la mucha bibliografía pro-

porcionada. Mención especial merece el

artículo escrito por Bonin y Rodríguez

(1991) que ha sido punto de referencia

obligado para la elaboración de este

trabajo. A Carmen Salas Casanova y

Serge Gofas, ambos de la Facultad de

Ciencias, Universidad de Málaga, por su

decisiva colaboración, ayuda y correc-

ciones, sin las cuales no hubiera sido

posible este artículo. A Javier Segarra

Julve y colaboradores, Francisco Ruiz

Sánchez y Emilio Valero Alfaro, todos

ellos de la Generalitat Valenciana, Con-

sellería de Agricultura Pesca y Alimen-

tación, por su colaboración en propor-

cionar los documentos relacionados en

77

Iberus , 26 (1), 2008

Figura 2. Glycymeris violacescens (Lamarck, 1819), ejemplares hallados en Mareny Blau (Sueca).

Longitud de la mayor concha encontrada: 67 mm. Figura 3. Glycymeris violacescens (Lamarck,

1819), ejemplares hallados en Cullera. Longitud de la mayor concha encontrada: 64 mm.

Figure 2. Glycymeris violacescens ( Lamarck , 1819), specimens collected in Mareny Blau (Sueca).

Length ofthe biggest shell found: 67 mm. Figure 3. Glycymeris violacescens (Lamarck, 1819), speci-

mens collected in Cullera. Length ofthe biggest shell found : 64 mm.

78

Tamayo GOYA: Catálogo de los bivalvos marinos del sector central del Golfo de Valencia

Figura 4. Fulvia fragilis (Forsskal in Niehbur, 1775), ejemplares hallados en Cullera (izquierda) y

Oliva (derecha). Longitud de la mayor concha encontrada: 42 mm. Figura 5. Conchas de Eastonia

rugosa recolectadas en Meliana. Longitud de la mayor concha encontrada: 80 mm. Figura 6. Gari

fervensis , ejemplar hallado en Cullera. Longitud de la mayor concha encontrada: 35 mm. Figura 7.

Conchas de Lentidium mediterraneum recolectadas en Mareny Blau (Sueca). Longitud de la mayor

concha encontrada por el autor de esta especie: 8 mm.

Figure 4. Fulvia fragilis (Forsskal in Niehbur, 1775), specimens collected in Cullera (left) and Oliva

( ñght). Length ofthe biggest shell found: 42 mm. Figure 5. Eastonia rugosa shells collected in Meliana.

Length ofthe biggest shell found: 80 mm. Figure 6. Gari fervensis, specimen collected in Cullera.

Length ofthe biggest shell found: 35 mm. Figura 7. Lentidium mediterraneum shells collected in

Mareny Blau (Sueca). Length ofthe biggest shell found: 8 mm.

79

Iberus, 26 (1), 2008

la bibliografía. A Rafael Araujo Armero,

del Museo de Ciencias Naturales de

Madrid, Alberto Martínez-Ortí, del

Museo de Historia Natural de Valencia;

Margarita Belinchón García, del Museo

de Ciencias Naturales de Valencia y

Francesc Uribe Porta, del Museo de

Ciencias Naturales de la Ciutadella

(Barcelona), por su colaboración en faci-

litar la consulta de ejemplares de las res-

pectivas colecciones y, en algún caso,

bibliografía. A Isabel Morón Marchante

y Mercedes González de Quevedo, de la

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La Valle, P., Vani, D., Liboni, A., y Smriglio,

C., 2007. Eastonia rugosa (Helbling, 1799) (Bi-

valvia, Mactridae) occurring along the Latium

coasts. Basteria, 71(1-3): 93-95.

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meradas.

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blaciones de Chamelea gallina (Linnaeus, 1758)

y Donax trunculus (Linnaeus, 1758) (Mollusco:

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Occidental). Tesis doctoral, Univ. Barcelona:

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del Reino de Valencia. Instituto de Estudios

Alicantnos. Diputación Provincial de Ali-

cante. 154 p.

Vardala-Theodorou, G. E., 1999. The occur-

rence of the Indo-Pacific molluscan species

Fulvia fragilis (Forsskal, 1775) and Bulla am-

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Hellenic Zoological Society, 31: 10-11.

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]., 2004. CIESM Atlas ofexotic species in the Me-

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80

Iberus, 26 (1): 81-170, 2008

© Sociedad Española de Malacología

New data on the Caribbean Triphoridae (Caenogastropoda,

Triphoroidea) with the description of 26 new species

Nuevos datos sobre los Triphoridae ( Caenogastropoda , Triphoroidea)

del Caribe , con la descripción de 26 nuevas especies

Emilio ROLÁN* and Raúl FERNÁNDEZ-GARCÉS**

Recibido el 24-IX-2007. Aceptado el 31-III-2008

ABSTRACT

In the present work 68 taxa of the family Triphoridae from the Caribbean and adjacent

areas are studied. Comments are also provided on one invalid taxon and on another that

has been erroneously reported from this area. Among these, 30 were previously known

species and 26 are described as new ( Metaxia propinqua , M. propria, M. prompto, Isot-

riphora guanahacabibes, Eutriphoro ouffenbergi , Marshallora ostenta , Marshallora apex-

diversus, Inella pseudolongissima , I. noduloides, I. apexbilirata , I. harryleei, I. undeber-

muda, I. pseudotorticula, I. differens, I. slapcinskyi , I. faberi , Sagenotriphora candidula,

"T." inaudita, "T." pseudonovem, " T. " yociusi, " T. " guadaloupensis, "T." amicorum, "T."

turtlebayensis, "T." grenadensis, "T." abacoensis a nd "T." portoricensis). Twelve species

remain unnamed or are tentatively identified, due to scarcity or ¡nadequacy of available

material, or doubtful specific assignment. New information is reported for Monophorus oli-

vaceus (Dalí, 1889) and Sagenotriphora osclausum (Rolán and Fernández-Garcés, 1995)

already treated in previous works.

Lectotypes are designated and figured for the following species Inella bigemma (Watson,

1880) , Inella ¡nflata (Watson, 1880), Inella longissima (Dalí, 1881), Inella trlserialis (Dalí,

1881) , Inella intermedia (Dalí, 1881), Inella ibex (Dalí, 1881), Inella torticula (Dalí,

1881), Inella colon (Dalí, 1881), Inella compsa (Dalí, 1927), Inella enopla (Dalí, 1927),

Inella meteoro (Dalí, 1927), Inella pompona (Dalí, 1927), Inella dinea (Dalí, 1927), Inella

sentoma (Dalí, 1927), Inella gaesona (Dalí, 1927), "Triphora" cylindrella (Dalí, 1927),

"Triphora" abrupta (Dalí, 1881), "Triphora" caracca Dalí, 1927, "Triphora" georgiana

Dalí, 1927, "Triphora" indígena Dalí, 1927, "Triphora" lilacina (Dalí, 1889), "Triphora"

pyrrha Henderson and Bartsch, 1914, Triphora" atlántica (E. A. Smith, 1890) and for the

western Atlantic species Triphora aspera Jeffreys, 1 885.

A list of the ñames employed for Caribbean species of Triphoridae is given, mentioning

mistakes and synonymies.

RESUMEN

En el presente trabajo se estudian 68 taxones de la familia Triphoridae del Caribe y aguas

próximas. También se aporta información sobre un taxon que carece de validez y otro erró-

neamente mencionado para el área. Entre ellos, 30 corresponden a especies previamente

conocidas, 26 se describen como nuevas para la ciencia [Metaxia propinqua, M. propria,

M. prompta, Isotriphora guanahacabibes, Eutriphora auffenbergi, Marshallora ostenta , Mars-

hallora apexdiversus, Inella pseudolongissima, I. noduloides, I. apexbilirata, I. harryleei, I.

* Museo de Historia Natural, Campus Universitario Sur, 15782, Santiago de Compostela, Spain

** Centro de Estudios Ambientales de Cienfuegos (CEAC), Grupo de Gestión Ambiental (GGA), calle 17,

esquina Ave. 46, Cienfuegos, Cuba.

81

Iberus , 26 (1), 2008

undebermuda , I. pseudotorticula, I. differens, I. slapcinskyi , I. faberi, Sagenotriphora candi-

dula, "T." inaudita, "T." pseudonovem, “T." yociusi, "T." guadaloupensis, "T." amicorum, "T."

turtlebayensis, "T." grenadensis, "T" abacoensis y "T." portoricensis) y o 1 2 de ellas no se les

asigna un nombre definitivo debido a la escasez del material, o al mal estado del mismo, y

también a la existencia de dudas sobre su determinación. Además, se aporta nueva infor-

mación sobre Monophorus olivaceus (Dalí, 1 889) y Sagenotriphora osclausum (Rolán y Fer-

nández-Garcés, 1 995), especies ya citadas en trabajos previos.

Se designan y figuran lectotipos de las siguientes especies: Inella bigemma (Watson,

1 880), Inella inflata (Watson, 1 880), Inella longissima (Dalí, 1881), Inella triserialis (Dalí,

1881), Inella intermedia (Dalí, 1881), Inella ibex (Dalí, 1881), Inella torticula (Dalí,

1881), Inella colon (Dalí, 1881), Inella compsa (Dalí, 1927), Inella enopla (Dalí, 1927),

Inella meteoro (Dalí, 1927), Inella pompona (Dalí, 1927), Inella dinea (Dalí, 1927), Inella

sentoma (Dalí, 1927), Inella gaesona (Dalí, 1927), "Triphora" cylindrella (Dalí, 1927),

"Triphora" abrupta (Dalí, 1881), "Triphora" caracca Dalí, 1927, "Triphora" georgiana

Dalí, 1927, "Triphora" indígena Dalí, 1927, "Triphora" ¡Hacina (Dalí, 1889), "Triphora"

pyrrha Henderson y Bartsch, 1914 y también de la especie atlántica Triphora aspera Jef-

freys, 1885.

Al final se aporta una lista de los nombres empleados para las especies de Triphoridae

del Caribe, señalando los errores habidos y las sinonimias.

KEY WORDS: Triphoridae, Caribbean, new species.

PALABRAS CLAVE: Triphoridae, Caribbean, new species.

INTRODUCTION

Triphoridae J. E. Gray, 1847 is a very

diverse family of marine gastropods

that occurs world-wide, with about 600

ñames proposed for the Recent species

(Marshall, 1983). Triphorids live on

rocky substrates from the intertidal

fringe to a depth of more than 1000 m. A

complete general information on the

family Triphoridae is given by Mar-

shall (1983) and Wells (1998), who

noted that it is a highly species-rich

group, most species being sinistral, with

a high spire, numerous whorls, a short

to long anterior canal and a posterior

apertural notch or canal. The teleoconch

sculpture of triphorids is notably vari-

able, and can be beaded, spinose or

reticulate, and the shells are small

(usually less than 10 mm, although

some can be up to 40 mm or more).

Marshall (1983), in his revisión of the

Indo-Pacific species, emphasized the

importance of the shape and sculpture

of the protoconch for taxonomy, and

pointed out that "under absolutely no

circumstances should further new

species be proposed unless a complete,

unworn protoconch can be illustrated".

The family has its máximum diver-

sity in the tropical Indo-Pacific where,

according to Marshall (1983), there are

more than 1000 species. An impression

of the wealth of species in this area is

afforded by the 174 species found

during an extensive sampling in the

coral reef lagoon of Koumac, New Cale-

donia (Bouchet et al., 2002), Triphori-

dae being the second richest family of

molluscs, after Turridae.

Although notably less diverse than in

the Indo-Pacific, this family contains

many species in the Atlantic Ocean,

poorly known in most areas. Bouchet

and Guillemot (1978) and Bouchet

(1985, 1997) revised the species of this

family occurring in the Mediterranean

and neighbouring Atlantic, where

CLEMAM data base recognised 19

species in 10 genera. Some additional

species were described by Fernándes

and Rolán (1988, 1993) and Van der

Linden (1998) in the Cape Verde Islands,

82

ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

and Rolán and Peñas (2001) from the

Canary Is. and the Mediterranean.

Information on triphorids from

many Caribbean areas is contained in a

number of works, such as those for

Jamaica (C. B. Adams, 1850a, 1850b,

most species represented in Clench

and Turner, 1950), Virgin Islands

(Nowell-Usticke, 1959 and 1971),

Yucatán Península, México (Vokes and

Vokes, 1983), Puerto Rico and nearby

Caribbean areas (Warmke and Abbott,

1961), Curasao, Aruba and Bonaire (De

Jong and Coomans, 1988), Oceanic

Islands off Brazil (Leal, 1991), Colom-

bian Caribbean (Díaz Merlano and

Puyana Hegedus, 1994), the Atlantic

and Gulf coasts and West Indies

(Morris, 1973), and Brazil (Ríos, 1994).

Rolán and Fernández-Garcés (1993a,

1993b, 1994 and 1995) and Rolán and

Espinosa (1994) showed the known

species from Cuba and described some

new species. Besides, other authors

described new species from other

Caribbean areas, such as De Jong and

Coomans (1988), Moolenbeek and

Faber (1989), Faber and Moolenbeek

(1991), Rolán and Crúz-Ábrego (1996)

and Rolán and Luque (1999). More

recently a large new species has been

described from Brazil (Simone, 2006).

Many other species were recorded from

deep water, and described by Watson

(1880, 1886), Dall (1881, 1889, 1927) and

other authors.

Fossil species were mentioned in

Olsson and Harbison (1953).

Colour photographs of 33 species,

including most of the known shallow-

water species and those described

during the last 20 years in previously

mentioned papers, are provided in a

recent paper (Rolán and Fernández-

Garcés, 2007) together with a list of

ñames assigned to the Caribbean

Triphoridae.

Henee, the present work has the fol-

lowing objectives: 1- To give available

information on the deeper water

species, showing the type material

whenever possible; 2- To provide addi-

tional colour photographs of the

Caribbean species which were not

figured in our previous work (Rolán

and Fernández-Garcés, 2007); 3- To

supply additional information that has

recently been obtained on other species

already known from shallow water; 4-

To describe some new species which

have been collected from shallow water;

5- To present an updated list of all the

taxa mentioned for the Caribbean and

nearby areas, making corrections to our

previous list.

MATERIAL AND METHODS

Some of the species presented here

are only known from the type material,

loaned by several museums (mentioned

in Abbreviations and Acknowledge-

ments), or viewed on photographs, pro-

vided by their Curators or other person-

nel. In the collected material, the collec-

tor ñame and the data of collection were

referred when both were known.

Besides, material loaned by Harry G.

Lee from his collection, mainly from

deep water, is included in this study.

Finally, a few species were collected by

the authors and persons who have

cooperated with them. Most of the early

described species, some of which had

never been figured previously or had

only been represented by drawings, are

photographed here in colour, and when-

ever possible these figures inelude the

types and also illustrate intraspecific

variability. Some photos taken by Sally

Diana Kaicher (found in ANSP) were

included.

The order of presentation is basically

a grouping based on similarity. It was

also the authors' intention to inelude

most of the available information on

protoconchs, radulae and opercula, but

unfortunately this was only possible

with a few species. Most of the type

material excluded this possibility, and

much of the studied material consisted

of shells without soft parts. An attempt

has been made also to solve taxonomic

problems relevant to some of the

studied taxa.

None of Dall's descriptions included

a designated holotype, although some

83

Iberus , 26 (1), 2008

seem to have been based on a single

shell. Under these circumstances, Dall's

material placed in some museums must

be considered as syntypes. After exami-

nation of available material from these

museums, it was therefore decided to

designate as lectotypes the shells that

were closest to the original description

or the original figure (shown in Dall,

1889).

In the description of shells, the spiral

beaded cords are frequently referred to

as "spiral 1, 2, 3 etc.", spiral 1 being the

subsutural cord on the adapical part of

the whorl, with subsequent cords on the

teleoconch whorls referred to as spirals

2 and 3, folio wing Marshall (1983).

Other important characters in the

description are, those of the protoconch,

such as the diameter, the shape and size

of the nucleus (elevated, depressed,

large, small, ...), the number of spiral

cords (none, 1 or 2), and the presence or

absence of axial ribs. We have followed

the method of Verduin (1976) for count-

ing the number of whorls of the proto-

conch. On the teleoconch, the number

and size of the spiral cords is noted on

the first and last whorls, where they are

frequently different. Also noteworthy is

the position of spiral 2 (closer to spiral 1,

equidistant between spirals 1 and 3) and

its size (nodules smaller, elongate, etc.).

It is important to point out that the

nodules, which are formed by the inter-

section of spirals and axial ribs, are

sometimes simply spherical, while other

times they are concave above and

convex below; in this case they are

crossed by a line that represents the

spiral cord and appears to cut the

nodule: these are referred to as cut

nodules or nodules that are cut in the

middle.

Due to the fact that the radula of

most species was unavailable, and as

this character is very important for

generic assignment, it was necessary in

many cases to employ generic ñames

very much in a sensu lato as "Triphora"

or "Inella". For generic arrangement we

mainly follow the works of Laseron

(1958), Marshall (1983) and Bouchet

(1985).

Abbreviations:

AMNH: American Museum of Natural

History, New York

ANSP: Academy of Natural Sciences,

Philadelphia

BMSM: Bailey-Matthews Shell Museum,

Sanibel Is., Florida

FLMNH: Florida Museum of Natural

History, Gainesville, Florida

IES: Instituto de Ecología y Sistemática,

La Habana

MCZ: Museum of Comparative

Zoology, Cambridge

MHNS: Museo de Historia Natural,

Santiago de Compostela

MNCN: Museo Nacional de Ciencias

Naturales, Madrid

USNM: United States National

Museum, Washington

CCR: collection of Colin Redfern

CHL: collection of Harry G. Lee

CFG: collection of Raúl Fernández-

Garcés

CMK: collection of Mario Krisberg

sp: specimen with soft parts

s: shell

j: juvenile

f: fragment

RESULTS

I. Taxonomic part

Family Triphoridae Gray, 1847

Subfamily Metaxiinae Marshall, 1977

Genus Metaxia Monterosato, 1884

Type species: Cerithium rugulosum C. B. Adams, 1850. According to Marshall (1983), this sub-

family is formed by dextral triphorids.

84

ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Metaxia propinqua spec. nov. Rolán and Fernández-Garcés (Figs. 1C-H)

Type material: Holotype (Figs. 1C, ID) in FLMNH (154988). Paratypes: 4 s, Louisiana, 27.98568°

N 92.6472° W, 65-91 m (FLMNH UF 291348); 5 s, Hillsborough Co., SW Egmont Key, Florida 73-91

m (collector Steger) (FLMNH 238632); 2 s (Figs. 1E, 1F), Palm Beach Co., Florida, off Boynton Inlet,

84-106 m (M. Glockstein, 1980) (FLMNH 47382).

Other material examined: 2 s (with broken protoconch), Louisiana, 27.98568° N, 92.6472° W, 65-

91 m (FLMNH UF 291348); 4 s (protoconch lost or eroded), Hillsborough Co., SW Egmont Key,

Florida 73-91 m (Steger) (FLMNH 238632); 1 s (protoconch eroded), 2 miles off Virginia Key, Florida,

9 m (FLMNH UF 365086); 1 s (protoconch eroded) (FLMNH).

Type locality: Florida, Monroe Co, WNW Dry Tortugas, 76 m.

Etymology: The specific ñame derives from the Latín word propinquus -a -um which means “cióse,

similar", alluding to the similarity with other species of this genus.

Description: Shell conical, very elon-

gate, relatively solid. Protoconch (Figs.

1G, 1H) white, paucispiral, with a little

more than two whorls and about 300 pm

in diameter; several spiral threads begin

at the apex. These are well defined, a

little irregular, four on the first whorl

decreasing to two on the second, where

numerous axial ribs run from the suture

to the upper thread. The teleoconch can

be white or light brown. Four spiral

cords are present from the beginning,

with the subsutural upper one less

prominent than the other three. Axial

ribs are present on the entire shell, about

10-11 on the first whorls and eighteen on

the final one. Suture depressed with a

very small spiral thread above. On the

base, the profile is slightly concave and

one additional cord is present. Aperture

almost round with an open and short

siphonal canal.

Dimensions : The holotype measures

6.3 mm.

Distribution : Known from Florida

and Louisiana, USA.

Remarks: In order to facilítate com-

parison of the protoconchs of the Metaxia

species mentioned here, they have all

been presented together in Figure 3.

M. rugulosa has a protoconch (Figs. 3A-

3D) with 2 1/i whorls (see Rolán and

Redfern, 1996), sometimes reaching three

whorls, with a spiral sculpture on the first

whorl formed by undulating or zigzag-

ging lines; the second protoconch whorl

has axial ribs which termínate at a single

spiral thread (only exceptionally double);

the teleoconch is white. M. excelsa and M.

taeniolata have multispiral protoconchs.

Metaxia propria spec. nov. Rolán and Fernández-Garcés (Figs. 2A-C)

Type material: Holotype (Figs. 2A, 2B) in FLMNH (UF 393603).

Type locality: Florida, Key Largo, 228 m.

Etymology: The specific ñame derives from the Latín word proprius -a -um which means “special,

characteristic", alluding to the differential characters of the protoconch that distinguish it from

other species of this genus.

Description : Shell conical, very elon-

gate, relatively solid. Protoconch white

(Fig. 2C) with a little more than 2 3A

whorls and about 400 pm in diameter;

two well defined spiral cords are crossed

by 13-15 axial ribs per whorl which are

continuous with those of the subsequent

whorl. Teleoconch white. Four spiral

cords are present from the beginning, the

subsutural upper one being less promi-

nent than the other three. Axial ribs

present on the entire shell, with about 10-

11 on the first whorls and 13 on the final

one. Suture depressed with a very small

spiral thread above. On the base, the

profile is slightly concave and one addi-

tional cord is present. Aperture almost

round, with an open siphonal canal.

Dimensions : The holotype measures

5.9 mm.

85

Iberus, 26 (1), 2008

Distribution : Only known from

Florida.

Remarks : Characters of the proto-

conch provide the main differences

between this species and those previ-

ously known:

M. rugulosa has a protoconch (Figs.

3A-3D) with 2 l/i whorls, sometimes

reaching 3, with a spiral sculpture on

the first whorl formed by undulating or

zigzagging lines; the second protoconch

whorl has axial ribs which terminate at

a single spiral thread (only exception-

ally double).

M. propincua spec. nov. lacks axial

ribs on the first whorl, and they reach

only to the upper spiral cord on the

second whorl.

Metaxia prompta spec. nov. Rolán and Fernández-Garcés (Figs. 2D-J)

Type material: Holotype (Figs. 2D, 2E) and 2 paratypes (Figs. 2G, 21) (FLMNH UF 359136).

Type locality: Bermuda, Hamilton Parish, Shelly Bay.

Etymology: The specific ñame derives from the Latin word promptus -a -um which means

"evident, available", alluding to the characters of the protoconch that are different from other

species of this genus.

Description : Shell conical, very elon-

gate, relatively solid. Protoconch (Figs.

2F, 2H, 2J) white, paucispiral with about

2 V2 whorls and between 300 and 400

pm in diameter; a single spiral cord is

poorly defined on the first whorl and a

little stronger on the second. The axial

ribs are numerous but not continuous;

they descend from the upper suture and

almost disappear just before reaching the

spiral cord, reappearing and strengthen-

ing on the cord before fading again

towards the lower suture. Teleoconch

white or light brown. Four spiral cords

are present from the beginning, the sub-

sutural upper one being less prominent

than the other three. Axial ribs are

present on the entire shell, varying in

number on the early whorls and increas-

ing to about 18 on the final one. Suture

depressed with a very small spiral

thread above. On the base, the profile is

slightly concave. Aperture almost round

with an open siphonal canal.

Dimensions : The holotype measures

4.1 mm.

Distribution : Only known from

Bermuda Archipelago.

Remarks : Characters of the proto-

conch easily differentiate this species

from related ones:

M. rugulosa has more rounded

whorls with zigzagging spiral lines on

the first whorl and numerous axial ribs

on the second, which only reach the

middle of the whorl.

M. espinosai has more angulate,

wider whorls, with prominent elongate

nodules and is always milky white.

M. propincua has rounded whorls,

with straight spiral cords beginning at

the apex and two cords on the second

whorl, with axial ribs absent from the

lower half of the whorl.

M. propria has almost three whorls,

rounded with two spiral cords and with

a few widely separated and prominent

axial ribs on the first whorls.

Metaxia sp. 1 (Figs. 1A-B)

Material examined: 1 s. Barbados, 183 m (USNM, labeled "Cerithiopsis abrupta Watson, 1880").

Remarks : The shell from Barbados

in USNM (without any type label) has

a paucispiral protoconch of 450 pm in

diameter, differing from M. abrupta

(Watson, 1880), which has a proto-

conch that is more depressed and also

has axial ribs (figured in Bouchet,

1985).

Dimensions : The shell studied mea-

sures 5.0 mm.

86

RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Bouchet (1985) was unable to

confirm the existence of M. abrupta in the

western Atlantic. Probably this species is

endemic to the Azores Archipelago. The

species studied here could be the one

referred to by Dall (1889) as Cerithiopsis

abruptus (Fig. 36L), but we are not sure.

This species is different from the

other two Caribbean species with pau-

cispiral protoconch: M. espinosai

(described by Rolán and Fernández-

Garcés, 1993a) has a paucispiral proto-

conch (Fig. 3E) with a prominent nodu-

lous keeled border; and M. rugulosa (C.

B. Adams, 1850), which has a narrower

protoconch (Figs. 3A-3D) with a differ-

ent sculpture (Rolán and Redfern,

1996). However, it must be pointed out

here that the protoconch of the taxon M.

rugulosa is not known, as the apex is

missing from the holotype of this

species, represented in Clench and

Turner (1950). Another fragment in

BMNH with the label "figured speci-

men" had no protoconch.

In several works it has been

assumed that the protoconchs from

Cuba and the Bahamas figured in

Rolán and Redfern (1996) represent

the protoconchs of M. rugulosa , probably

due to the proximity of Cuba to Jamaica,

which is the type locality. Similar proto-

conchs were figured in Redfern (2001).

The other two known species of

Caribbean Metaxia ( M . excelsa and M.

taeniolata) have multispiral protoconchs.

The present species is probably

unknown and unnamed, but as only one

shell has been examined, and its proto-

conch is not in very good condition, it is

kept unnamed pending the availability

of more material in the future.

Subfamily Triphorinae Gray, 1847

Genus Monophorus Granata-Grillo, 1877

Type species (by monotypy): Trochus perversus Linnaeus, 1758. Recent, Mediterranean.

Remarks: The main characters (shown

in Marshall, 1983, Bouchet, 1985 and

Rolán and Fernández-Garcés, 1994)

are the following: multispiral protoconch,

teleoconch whorls flat-sided or shallowly

convex, well defined spiral cords and axial

ribs, nodular intersections and a typical

radula with central tooth with 3-5 cusps,

lateral with 5-6 and numerous margináis

with only 2-3 short cusps.

Monophorus olivaceus (Dall, 1889) (Figs. 4B-F, 4F4-K)

Triforis decorata var. olivácea Dall, 1889. Bull. M. C. Z., 18: 243. [Type locality: Gulf of México, west

of Florida, 91 m, Key West, Hemphill].

Triphora ornata auct. non Deshayes (1832).

Type material: One syntype, fragment of 10 mm, here designated as lectotype (Figs. 4C, 4D) (MCZ

7379).

Other material examined: México: 1 s, Cancun, Quintana Roo (Heilman, March, 1963) (ANSP

285517). Florida: 3 s, WSW of John's Pass, 50 m (Steger, 1956) (ANSP 306254); 1 s, Palm Beach Co.,

Bath and Tennis (McGinty coll/ Apr. 1951), 36-55 m (FLMNH 176649); 4 s, Collier Co., SW of Naples,

26 20' N (Powlus/Aug., 1960), 55 m (FLMNH 158202); 9 s, Palm Beach Co., off Palm Beach, 100 m

(FLMNH 154886); 3 s, Palm Beach, WOE Radio Towers to Bath and Tennis (McGinty, May/1951)

36-55 m (FLMNH 250132); 1 s, Palm Beach Co., (R/V Tritón, Apr. 1950) 40-50 m (FLMNH 204877);

1 s, off Marathón (M. and S. Snyder, Jul. 1966) (ANSP 309744). Bahamas: 1 s, reef NE of North Point,

Elbow (Little Guana) Cay, Abaco (Robertson, Aug. 1955) (ANSP 298409); 4 s, Grand Bahama Island,

26° 31' 00" N, 78° 46' 30" W (J.N. Worsfold) (ANSP 373939); 1 s, Grand Bahama Island, 26° 29' 45"

N, 78° 37' 15" W (G. Taylor and D. Nilsen) (ANSP 273605); 2 s, Tamarind, Grand Bahama Island

26° 30' 45" N, 78° 36' 00" W (J. Worsfold) (ANSP 368896); 1 s, Bootle Bay, Grand Bahama Island 26°

87

Iberus , 26 (1), 2008

Figure 1. A, B. Metaxia sp. 1. A: shell, 5 mm, Barbados, 183 m (USNM); B: protoconch. C-H.

Metaxia propinqua spec. nov. C, D: holotype, 6.3 mm (FLMNH); E, F: paratype, 3.4 mm

(FLMNH); G: protoconch of the paratype; H: detail of the protoconch of the holotype.

Figura 1. A, B. Metaxia sp. 1. A: concha , 5 mm, Barbados, 183 m (USNM); B: protoconcha. C-H.

Metaxia propinqua spec. nov. C, D: holotipo, 6,3 mm (FLMNH); E, F: paratipo, 3,4 mm (FLMNH);

G: protoconcha del paratipo; H: detalle de la protoconcha del holotipo.

88

RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 2. A-C. Metaxia propria spec. nov. A, B: holotype, 5.9 mm (FLMNH); C: protoconch of

the holotype. D-J. Metaxia prompta spec. nov. D, E: holotype, 4.1 mm (FLMNH); F, G: proto-

conch and shell of paratype, 3.3 mm (FLMNH); H, I: protoconch and shell of paratype, 4.0 mm

(FLMNH); J: protoconch of holotype.

Figura 2. A-C. Metaxia propria spec. nov. A, B: holotipo, 5,9 mm (FLMNH); C: protoconcha del

holotipo. D-J. Metaxia prompta spec. nov. D, E: holotipo, 4,1 mm (FLMNH); F, G: protoconcha y

concha de un paratipo, 3,3 mm (FLMNH); H, I: protoconcha y concha de un paratipo, 4,0 mm

(FLMNH); J: protoconcha del holotipo.

89

Iberus, 26 (1), 2008

39' 30" N, 78° 57' 00" W (J. Worsfold) 80 m (ANSP 371846); 1 s, Schooner Cays, NW of Powell Point,

Eleuthera 24° 56' 45" N 76° 25' 00" (R. Robertson) (ANSP 363405); 3 s (G. Buchanan) (BMNH

90.11.24.428-34). Venezuela: 4 s, distrito Federal (Berthier, 1935) (ANSP 264204). Grenada: 1 s, mouth

of St. George Harbour (Ostheimer, Jan. 1964) 44-66 m (ANSP 297306). Virgin Islands: 3 s, St. Croix

(R.E. Griffith) (ANSP 18201). Dutch Antilles: 5 s, (Gray coll.) Saint Vincent (BMNH). No exact local-

ity: 3 s (McAndrew coll.) (BMNH Acc n° 1563); 2 s (R. Sykes coll.) (BMNH acc. n° 1825).

Description : See Dall (1889). It is

important to point out some characters:

the large size of the beads, spiral 2 being

absent from the early whorls and

smaller on most of the shell, the colour

forming blotches, predominantly on

spirals 1 and 2. Spiral 3 is white, but the

cord between the beads is usually dark.

Dimensions : The lectotype measures

10 mm.

Distribution: Known throughout the

Caribbean.

Remarks: There is considerable con-

fusión regarding this taxon: the species

was referred to as Triphora ornata by

many authors (Warmke and Abbott,

1961; Morris, 1973; Ríos, 1994) until

Moolenbeek and Faber (1991) showed

the correct ñame to be M. olivaceus (Dall,

1889). The type material of Triphora

ornata was examined by Faber (pers.

comm.) in 1983 from MNHN (Ecole des

Mines collection where the Deshayes

types should have been). This lot con-

sists of several poor shells, none of

which matches the Caribbean species,

and one of them in better condition cor-

responds to a Red Sea species.

The only known type specimen of

Triforis decor ata var. olivácea is an atypical

and faded shell. It can be confused with

some species with axial colour blotches:

Latitriphora albida (Fig. 4A) has very

characteristic smaller nodules, and no

confusión is possible; Nototriphora deco-

rata may be more similar and requires

careful comparison. We have compared

a typical shell of Nototriphora decorata

(Fig. 4G) with representative shells of

M. olivaceus (Figs. 4D-F) and with the

lectotype of the latter species (Figs. 4B,

C). It can be seen that the early whorls

of N. decorata are narrower than those of

the type of M. olivaceus and have smaller

nodules, whereas the early whorls of

representative M. olivaceus are very

similar to those of the type specimen. As

it is necessary for nomenclatural stabil-

ity to designate a lectotype, we propose

as such the only known syntype.

The species M. olivaceus is very vari-

able, with large, wide shells sometimes

found in the same population as small,

narro w ones. Also, the dark colour may

vary from almost black to light brown

and can be variably distributed, but it is

always present on spiral 1 and on a few

nodules of spiral 2.

Examination of a shell from Florida

(Figs. 4H-4J) revealed a dark subsutural

spiral and a dark base. It appeared

slightly different from average examples

of M. olivaceus (like Fig. 4K), as the

colour is not only darker but is also

slightly differently distributed; spiral 1

has only isolated white beads and the

brown color does not form blotches,

spiral 2 is mostly white and spiral 3 is

totally white, without any brown colour

between the beads. Also the beads on

spiral 2 become comparatively larger

several whorls before the final whorl.

However, it is considered to be only a

variety.

Genus Isotriphora Cotton and Godfrey, 1931

Type species (by original description) Triforis tasmanica T. Wood, 1875. Recent, South Australia,

Tasmania.

Description : The most important cióse to that of the genus Eutriphora

character of this genus is the paucispiral Cotton and Godfrey, 1931, but the latter

protoconch with nodular spiral cords, has axial riblets. Operculum multispiral.

90

RoláN AND FERNÁNDEZ-GARCÉS: New data on the Caribbean Triphoridae

Figure 3. Protoconchs of Metaxia species. A-D. M. rugulosa (C. B. Adams, 1830) from Cuba (A)

and Bahamas (B-D). E. M. espinosai Rolán and Fernández-Garcés, 1993, Cuba.

Figura 3. Protoconchas de especies de Metaxia. A-D. M. rugulosa (C. B. Adams, 1850) de Cuba (A) y

Bahamas (B-D). E. M. espinosai Rolán y Fernández-Garcés, 1993, Cuba.

Isotriphora guanahacabibes spec. nov. Rolán and Fernández-Garcés (Figs.

5A-H)

Type material: Holotype (Fig. 5A) in MNCN (15.05/15.05/47054) and paratypes in the following

collections: MHNS, 2 s (Fig. 5B); IES, 1 s (Fig. 5C); CFG (3 s, 8 f), all from the type locality; 1 f and

1 j, from Cienfuegos, Cuba (MHNS); 1 s, outer slope of beach, W of Georgetown, Cayman Is. 100-

115 fms (Tyler, Oct. 1964) (ANSP 300623).

Other material examined: 2 f, Cienfuegos Bay, Cuba 40 m; 1 j, S shore of island, 3 miles E of South-

west Point, Cayman Brac Island (ANSP 397055).

Type locality: From sediment at Cueva de Pipo, Maria la Gorda, Guanahacabibes, Pinar del Río,

Cuba, at 35 m.

Etymology: The specific ñame comes from the type locality.

Description : Shell (Figs. 5A-5C)

whitish, cortical elongate, solid. Proto-

conch (Figs. 5D-5F) paucispiral with

about 300 pm in diameter, the number of

whorls not appreciable due to indetermi-

nate separation from the teleoconch.

The protoconch begins with a very

narrow nucleus and the following two

whorls have two prominent nodular

spirals, with another very small one below

the suture. Teleoconch with about 10

whorls and with a straight profile; each

91

Iberus, 26 (1), 2008

whorl with two widely separated nodu-

lous spiral cords (spirals 1 and 3), the

lower one more prominent; about the 6-

7^ whorls, a small cord (spiral 2) appears

closer to spiral 1 (Fig. 5G). Spiral 2 con-

tinúes onto the subsequent whorls, becom-

ing narrow, undulating and slightly nodu-

lous towards the end. Spiral 2 is always

smaller than spiral 1 and 3 and is closer

to the one above. On the last two whorls

an additional very small, smooth spiral 4

appears at the suture. The aperture is

ovoid, the outer lip fine and prominent.

Near the end of the final whorl there is a

small hole, located cióse to the apertural

border on the upper part of the whorl.

Siphonal canal closed, curving towards

the rear. The colour is totally white, but

some specimens can have small areas of

light brown (Fig. 5C). Under high magni-

fication spiral striae (Fig. 5H) can be seen

between the spirals.

Dimensions : The holotype measures

5.5 x 1.6 mm.

Distribution: Known from the type

locality and from the Cayman Islands,

down to 183-210 m.

Remarles: This species must be com-

pared with the other congeneric species

with a similar protoconch:

Isotriphora peetersae (Moolenbeek and

Faber, 1989) is dark brown, except for

the protoconch and first whorls of the

teleoconch, and the shell is more obese

and with fewer whorls. Furthermore,

the anal sinus is at the end of the spire

and is not closed.

Isotriphora taenialba Rolán and

Espinosa, 1994, is similar to I. peetersae

but a little larger, and always has dark

and lighter bands of colour; also the

anal sinus is open at the end of the spire.

At first sight it might be confused

with species of the genus Iniforis

Jousseaume, 1884 such as I. immaculata,

which is totally white, but that species

has a protoconch with only one smooth

fine cord.

Genus Eutriphora Cotton and Godfrey, 1931

Type species (o. d.): Triphora cana Verco, 1909, Recent, Australia.

Description: After Marshall (1983),

similar to the genus Isotriphora but with

multispiral protoconch with axial

riblets. Radula with the rachidian tooth

having three cusps, lateral teeth with 4-5

and numerous marginal teeth with an

elongate central cusp. Operculum pau-

cispiral.

Eutriphora auffenbergi spec. nov. Rolán and Lee (Figs. 6A-I)

Type material: Holotype (Figs. 6A-6B) in FLMNH. Paratypes: 1 s (Fig. 6C) (BMSM); 1 s (USNM);

1 s (CHL), all from type locality and ex CHL. Other paratypes: 3 s, 1 j, off Egmont Key, Hillsbor-

ough Co., Florida, 76 m (Jim Moore /Cavalier, 1962) (CHL); 1 j, 65 miles E St. Augustine, St. Johns

Co., Florida (T. Yocius, Jun. 1972), 54 m (CHL); 1 j, 32 miles E St. Augustine, St. Johns Co., Florida

(T. Yocius, Jun. 1981), 30 m (CHL); 4 s (Fig. 6D), Florida, (MCZ 356088); 1 s (Fig. 6E), Florida, 5 m

(MCZ 356096); 1 s, Hillsborough Co., W of Egmont Key, Florida, 274 m (FLMNH 169703); 1 sp,

Palm Beach Co., E of Palm Beach, Florida, 183-578 m (FLMNH 257207).

Other material examined (in bad condition): 1 s, 10 f, Pinellas Co., WSW John's Pass, Florida, (D.

Steger) 68 m (FLMNH 238662).

Type locality: West of Dry Tortugas, Monroe Co., Florida, USA (Jim Moore/ Cavalier, 1972), 90 m.

Etymology: The species is named after Kurt Auffenberg, former Collections Manager in Malacol-

ogy. Florida Museum of Natural History, USA.

Description : Shell (Figs. 6A-6E) spiral, with about 4 whorls, the surface

sharp-pointed, elongate, solid, brown or of the apex with small tubercles and the

light brown. Protoconch (Fig. 6G) multi- subsequent whorls with two small

92

RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

spiral cords crossed by numerous fine

axial ribs that have a slight S-shaped

curve. Teleoconch with about 17-18

whorls in large specimens, beginning

with spirals 1 and 3, crossed by proso-

cline axial ribs which form small

nodules at the intersection points. On

whorls 4-5, spiral 2 appears cióse to

spiral 1; on the subsequent whorls, the

three spirals are similar in size but spiral

3 is a little more prominent. An addi-

tional spiral cord, appearing on about

whorls 10-12, is slightly nodulous,

always smaller, and is located just below

the suture (Fig. 6F). This cord is more

evident on the final whorl, where it

occurs near the periphery. Below it there

are two more smooth spiral cords, the

lower one on the base of the siphonal

canal (Fig. 6C). Aperture rounded, cól-

umella curved, siphonal canal elongate

and recurved, closed at its base by an

extensión of the aperture.

Dimensions: The holotype measures

21.8 mm, with 20 whorls. Other shells

are smaller.

Operculum (Fig. 6H): paucispiral,

subcircular, yellowish, rather transpar-

ent, with the nucleus subcentral.

Radula (Fig. 61): relatively large,

with a formula 16-1-1-1-16, the rachid-

ian tooth having three prominent cusps;

the lateral has 5 cusps of which cusps 2

and 4 are smaller. The marginal teeth

have a filiform central elongate cusp,

with the other two short and similar in

size.

Distribution : The species has been

found only in Florida.

Remarks: The generic assignment is

based on the similarity of the radula,

operculum and protoconch to the type

species of the genus Eutriphora : E. armil-

lata (Verco, 1909) from Australia (see

Marshall, 1983).

The holotype of E. auffenbergi spec.

nov. resembles the figure of T. triserialis

in Dall (1889: pl. 20, fig. 6a) (Fig. 36E),

but the following characters of the

present species are not coincident: the

shell is brown and the protoconch also is

darker, typically with 4 whorls and with

tubercles on the apex; the siphonal canal

is elongate. All these characters are dis-

similar from DalTs species (see below).

Comparison and differentiation

must be made with the following

Caribbean species described here:

"Triphora" abrupta has a shorter shell,

with the nodules less spherical, spiral 1

is very small and cióse to the suture,

almost similar to spiral 4, while in T. auf-

fenbergi spiral 1 is similar to the other

main two.

"T. " georgiana has a very small spiral

1 which remains reduced on the final

whorls, and the protoconch probably

has fewer whorls.

"T. " caracca has two main spirals and

between them a very narrow spiral 2;

the axial ribs are very prosocline.

"T. " indígena has a more convex

profile to the whorls; the spiral cords are

wider and the nodules at the intersec-

tion points are very small, resulting in

rectangular pits in the interspaces

between cords and ribs.

E. auffenbergi is somewhat similar to

Sagenotriphora osclausum (Rolán and Fer-

nández-Garcés, 1995) (see below), but

the latter species is shorter (usually with

no more than 9 teleoconch whorls), and

the siphonal canal is also very short and

curved.

Other brown species are Marshallora

nigrocincta (C. B. Adams, 1839) and M.

modesta (C. B. Adams, 1850), but these

species have a shorter spire (not more

than 10 whorls), a more convex profile

and a very short siphonal canal.

Genus Marshallora Bouchet, 1985

Type species: Murex adversus Montagu, 1803 (o.d.).

Diagnosis: After Bouchet (1985),

protoconch of 4-5 whorls with two

spiral threads, granules on its apical

part, teleoconch with the second cord

appearing later, operculum paucispiral

and radula with central and lateral teeth

93

Iberus, 26 (1), 2008

with two groups of cusps. Caribbean Garcés (1995) and Rolán and Crúz-

species in Rolán and Fernández- Ábrego (1996).

Marshallora ostenta spec. nov. Rolán and Femández-Garcés (Figs. 7A-N, 8E, 8F)

Type material: Holotype (Figs. 7A, 7B) in MNCN (15.05/47055) and a paratype (Fig. 7C) in IES,

both from type locality. Other paratypes in the following institutions: 1 s (Fig. 7E), off Mayport,

Duval Co., Florida (Yergin, Jan. 99), 30 m (FLMNH, ex CHL); 2 s (Figs. 7F, 7G), off Big Pine Key,

Monroe Co., Florida, 30-50 m, (K. Sunderland, Aug/91) (BMSC ex CHL); 1 s, off Big Pine Key,

Monroe Co., Florida, 30-50 m (K. Sunderland, 1991) (CHL, Fig. 71); 3 s, off Egmont Key, Hillsbor-

ough Co., Florida (J. Moore/ Cavalier , 1962), 72 m (CHL); 1 s (Fig. 7H), 32 miles E. St. Augustine, St.

Johns Co., Florida (T. Yocius, 1972) 30 m (USNM ex CHL); 8 s, Palm Beach Co., Florida, off Boynton

Inlet (D. and H. Akers, Dec. 1969) 33 m (FLMNH 177040); 1 s (Fig. 7J) Miami, Florida, 73 m (FLMNH

UF363887); 1 s (Fig. 7K), Palm Beach Co., Florida, off Dodge Estate (R/V Tritón, Dec. 1951) (McGinty

coll.) (FLMNH 178201).

Other material examined: Cuba: 2 sp (Fig. 7D)(destroyed for radular study), Cienfuegos (MHNS).

USA: Florida: 1 sp (destroyed for radular study) and 2 s. Anclote Key, near Tarpon Springs, Pasco

Co. (CHL); 1 s, off Dry Tortugas, Monroe Co., Riley Black (ex J. Dawley, 1/4/1983), 55 m (CHL)

(the last two lots not in good condition); 4 s, off Destin, 25 m (ANSP 252157); 1 s, Palm Beach Co.,

off Yamato Rocks, (F. B. Lyman/ Apr. 1939) 11 m (FLMNH 10242); 3 s, St. Augustine (FLMNH

286017).

Type locality: Cienfuegos Bay, Cuba.

Etymology: The specific ñame derives from the Latín past participle ostentus -a, -um which

means "shown", indicating that the review of material from several collections allowed us to rec-

ognize this species.

Description : Shell (Figs. 7A-7K) light,

solid, conical elongate. Protoconch (Figs.

8E, 8F) dark brown, multispiral, with

between 3 l/i and 4 whorls; the apex is

covered by small tubercles; fine axial

ribs are crossed by two spiral cords on

the lower whorls and sometimes by a

single cord on the first whorl. Teleo-

conch with 7-10 whorls, the first 4-5

with only two beaded cords (spirals 1

and 3); spiral 2 appears between them

on about the fifth or sixth whorl,

increasing in size until the three spirals

are the same size on the final whorl.

Another very small, smooth spiral

occurs at the suture. The colour is

usually white at the beginning of the

teleoconch, sometimes including the

first 1-2 whorls; subsequently the colour

of the shell is more variable: frequently

uniform cream or light brown, darker at

the suture; the beaded cords may be dif-

ferent, whitish, cream or light brown,

and either the lower or upper cord can

be lighter. Aperture rounded; base

brown, sometimes only light brown, the

darker colour extending to the col-

umella; siphonal canal short and open.

Dimensions: The holotype measures

4.2 mm; some paratypes reach 5.5 mm.

The soft parts (examined in only two

specimens) are whitish, slightly translu-

cent, with milk-white spots on the head

and a few on the tentacles.

Operculum (Fig. 7N) rounded-

ovoid, multispiral, with the nucleus

somewhat deviated from the centre.

Radula (Figs. 7L, 7M) studied in one

specimen from Cuba and another from

Florida, has a formula n-l-l-l-n. The

rachidian tooth has two parts with 5-6

sharp cusps at each side; the lateral ones

are very similar to the central one; the

marginal ones are sharp pointed, very

elongate and narrow.

Distribution: Known from Cuba and

Florida.

Remarks : In spite of the amount of

material examined, it was not possible to

arrive at a description that represented

all the shells, suggesting that this must

be a variable species. There are some dif-

ferences in the protoconch (number of

whorls, one or two spiral threads on the

first whorl), but the main differences

concern the very variable colour distrib-

94

RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 4. A. Latitriphora albida (A. Adams, 1854) (BMNH). B, C. Monophorus olivaceus (Dalí,

1889), lectotype, 10 mm (USNM). D-F. M. olivaceus typical shells from Cuba, all for comparison

with the shell of the lectotype. G. Nototriphora decorata (C. B. Adams, 1850), Cuba. H-K.

Monophorus olivaceus (Dalí, 1889). H-J: shell, 7.0 mm, Peanut Island, Florida (MHNS); K: shell,

8 mm, Florida (FLMNH).

Figura 4. A. Latitriphora albida (A. Adams, 1854) (BMNH). B, C. Monophorus olivaceus (Dalí,

1889), lectotipo, 10 mm (USNM); D-F. M. olivaceus, conchas típicas de Cuba, todas para compara-

ción con la concha del lectotipo. G. Nototriphora decorata (C. B. Adams, 1850), Cuba. H-K.

Monophorus olivaceus (Dalí, 1889). H-J: concha, 7,0 mm, Peanut Island, Florida (MHNS); K:

concha, 8 mm, Florida (FLMNH).

95

Iberus , 26 (1), 2008

ution on the teleoconch and the initial

appearance of spiral 2 between the 5th

and 7th teleoconch whorls. All of this

suggested to us that more than one

species might be involved in the studied

material. Anyway, some shells with clear

differences ha ve been separated from the

initial lot, and these are described below

(the following species). The remainder

were considered to represent a single

variable species, which future study of

live collected material may confirm

through observation of the colour of the

soft parts and examination of radulae.

Similiphora intermedia (C. B. Adams,

1850) always has the spiral cords white

and brown, with a rather uniform color

pattern of a darker upper cord and a

dark or light brown middle one; the soft

parts are black. The radula has numer-

ous teeth all similar in size.

Marshallora modesta (C. B. Adams,

1850) and M. nigrocincta (C. B. Adams,

1839) have similar shells but they are

darker brown and never whitish. The

protoconchs are somewhat different,

being wider and shorter (as can be seen

in Figures 8H and 81).

Marshallora apexdiversus spec. nov. Rolán and Lee (Figs. 8A-D, 8G)

Type material: Holotype (Fig. 8A) and 3 s (Figs. 8B, 8C), paratypes (FLMNH UF363887) from type

locality; 1 s, 1 s (Fig. 8D), Palm Beach Co., Florida, off Dodge Estate (R/V Tritón, Dec. 1951) (McGinty

coll.) (FLMNH 178201).

Type locality: Miami, Florida, 73 m.

Etymology: The specific ñame derives from the Latin words diversus, which means "different", and

apex, referring to an important character for separation from other cióse species.

Description : Shell (Figs. 8A-8D) light,

solid, conical elongate. Protoconch (Fig.

8G) light brown, with 5 whorls; the apex

is covered by small tubercles and is very

narrow; the whorls with fine axial ribs that

are crossed by two spiral cords, the upper

cord almost in the middle of the whorl

and the lower one closer to the suture be-

low. Teleoconch with 8-11 whorls, the first

2-3 with only two beaded cords (spirals 1

and 3); spiral 2 appears between them on

about the fourth whorl, increasing in size

until the three cords are of equal size on

about the fifth or sixth whorls. Another

very small, smooth spiral occurs at the su-

ture on the final whorls. The colour is usu-

ally white on the first 2 whorls of the

teleoconch; on subsequent whorls spiral

1 is light brown, and on some shells (but

not on the holotype)this colour also reach-

es the suture. The nodules are always

lighter than the ground colour. Aperture

rounded-ovoid; base cream, siphonal

canal short and closed at the aperture by

an extensión of the outer lip.

Dimensions: The holotype measures

6.2 mm.

Distribution : Only known from

Florida.

Remarks : The present species is tenta-

tively included in Marshallora due to sim-

ilarity with the other Caribbean species in

this genus. At the beginning of this study

the shells included in this taxon were

placed in the variable M. ostenta spec. nov.

Finally, both species were considered dif-

ferent on the basis of the following char-

acters: M. ostenta almost always has a dark

brown base, the protoconch has fewer

whorls (see Figs. 8E, 8F in comparison

with Fig. 8G), the apex is wider, the spiral

threads on the protoconch are in the

middle of the whorl; on the teleoconch,

spiral 2 never begins before the fifth whorl,

sometimes even later.

Similiphora intermedia (C. B. Adams,

1850) always has very dark colour on

spiral 1, while spiral 2 is light or dark

brown.

Marshallora modesta (C. B. Adams,

1850) and M. nigrocincta (C. B. Adams,

1839), have similar shells but in both

cases they are darker brown and never

whitish or cream. The protoconchs of

both species are rather different, being

wider at the apex and shorter (as can be

seen in Figures 8H and 81, in compari-

son with Fig. 8G).

96

ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 5. Isotriphora guanahacabibes spec. nov. A: holotype, 5.5 mm (MNCN); B: paratype, 4.7

mm (MHNS); C: paratype, 4.35 mm (IES). D-F: protoconchs of paratypes (MHNS); G: detail of

the end of the last whorl, paratype in figure B; H: detail of the microsculpture.

Figura 5. Isotriphora guanahacabibes spec. nov. A: holotipo, 5,5 mm (MNCN); B: paratipo, 4,7 mm

(MHNS); C: paratipo, 4,55 mm (IES). D-F: protoconchas de paratipos (MHNS); G: detalle del final

de la última vuelta, paratipo de la Figura B; H: detalle de la microescultura.

Genus Inella Bayle, 1879

Ino Hinds, 1843, p. 17. Type species (subsequent designation of Jousseaume 1844: 230): Triforis

(Ino) gigas Hinds, 1843. Recent, New Guinea.

Inella Bayle, 1879, p. 35. New ñame for Ino Hinds (not Ino Samsuelle, 1817).

Remarks : The type species (see

Marshall, 1983) has a blunt and wide

protoconch with between two and

three whorls, each with two spiral

cords; the shells included in this genus

are elongate and usually have two

spiral beaded cords at the beginning of

the teleoconch (three exceptionally) and

three on the subsequent whorls,

frequently with a fourth cord

appearing at the suture. Marshall

(1983) mentioned that the limits of the

genus are uncertain, and the radula has

numerous small and similar teeth

except the rachidian, that has symmet-

rical cusps.

97

Iberus , 26 (1), 2008

Inella bigemma (Watson, 1880) (Figs. 9A-B, 36B)

Cerithium (Triforis) bigemma Watson, 1880. Prelim. Report., pt 5. Journ. Linnean Soc. London, 15:

101, sp. 2. [Type locality: 18° 38' 30" N, 65° 5' 30" W, Off Culebra Island, 713 m].

Type material: Syntype (BMNH 1887.2.9.1762) (Fig. 9A) here designated as lectotype.

Other material examined: 1 s (Fig. 9B), (BMNH 1915.12.31.199) from Irish Fishery Board.

Description : See Watson (1881, 1886).

The protoconch is unknown. Two spiral

cords are present on the first whorls,

subsequently with an additional

smaller, less prominent cord appearing

above them. About 17 whorls on the lec-

totype.

Dimensions: The lectotype measures

13.5 mm.

Distribution : Only known with cer-

tainty from the lectotype. Culebra Island

(between Puerto Rico and Virgin

Islands).

Remarks: The shell mentioned in the

original description as having 0.6 inch

(=15 mm) and the original figure (Fig.

36B) are coincident in size and number

of whorls with the shell studied by us,

and probably it is the same.

The second shell studied (Fig. 9B) has

a different profile and could be a different

species (Bouchet and Warén, 1993 are

of the opinión that this shell is Strobiligera

brychiá). Without protoconchs it is not pos-

sible to make a correct comparison. This

species is very similar to I. torticula (see

below, in Remarks for that species).

This taxon is mentioned in Dall

(1981, 1989), who suggested that the

species Triforis torticula could be a

variety of T. bigemma. As the shells are

not in perfect condition and lack proto-

conchs, not enough information was

obtained to decide if they represent dif-

ferent species, but the whorls seem to

expand more slowly on the type of T.

torticula, although not on the other

shells studied.

Inella inflata (Watson, 1880) (Figs. 9C, 36G)

Cerithium (Triforis) inflatum Watson, 1880. Prelim. Report., pt 5. Journ. Linn. Soc., 15: 103. [Type

locality: 18° 38' 30" N, 65° 5' 30" W, St. Thomas, N of Culebra Island, Danish West Indies, 713

m; indicating: Habitat.- Yucatán Strait, 1170 m (Dalí).].

Triforis inflatum Watson: Dall, 1881. Bull. Mus. Comp. Zool., 9: 6.

Triforis (Sychar) inflata Watson: Dall, 1889. Bull. Mus. Comp. Zool., 18: 249.

Type material: Syntype (BMNH 1887.2.9.1766) (Fig. 9C) here designated as lectotype.

Description : See Watson (1881, 1886).

The protoconch is pupoid and perhaps

with about two whorls, the first one

wider than the second. It may be about

500 pm in diameter. The teleoconch is in

bad condition, but it is possible to see

the two main beaded spirals, and one

more near the suture. The nodules are

concave above and pointed.

Dimensions: The lectotype measures

4.8 mm.

Distribution: Only known from the

type locality.

Remarks: No other syntype of this

species was found in other museums

and the shell is nearly identical to that

represented in the original figure of I.

Inflata (Fig. 36G), for which 13 whorls

are mentioned (really 12 including the

protoconch) and the correct size of 4.8

mm. Therefore, probably it could be the

only type specimen but, as no holotype

was mentioned in the original descrip-

tion, this shell is here designated as lec-

totype. Its protoconch is very different

from most others from the Caribbean

area, and is therefore a differential char-

acter.

Dall (1889) placed his species T. ibex

as a variety of I. inflata Watson. Both

98

RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 6. Eutriphora aujfenbergi spec. nov. A, B: holotype, 21.8 mm, off Dry Tortugas, Florida

(FLMNH); C: paratype, 8.4 mm, same locality (BMSM); D: paratype, 13.7 mm, Fia., USA

(station 4)(MCZ 336088). E: paratype, 12.1 mm, 14 whorls, Fia., USA, 5.5 m (MCZ 356096).

F: detail of sculpture of the holotype; G: protoconch of the paratype in fig. C; H: operculum (of a

specimen from Palm Beach, Florida, 100-300 fms (FLMNH); I: radula (from the same specimen).

Figura 6. Eutriphora auffenbergi spec. nov. A, B: holotipo, 21,8 mm, de fuera de Dry Tortugas,

Florida (FLMNH); C: paratipo, 8,4 mm, la misma localidad (BMSM); D: paratipo, 13,7 mm. Fia.,

USA (estación 4) (MCZ 356088). E: paratipo, 12,1 mm, 14 vueltas, Fia., USA, 5,5 m (MCZ

356096). F: detalle de la escultura del holotipo ; G: protoconcha del paratipo en la fig. C; H: opérculo

(de un ejemplar de Palm Beach, Florida, 100-300 brazas (FLMNH); I: rddula (del mismo ejemplar).

99

Iberus, 26 (1), 2008

taxa do not appear to be similar: I. ibex

lacks a protoconch and the type, with 14

whorls, is 10.4 mm; the description of I.

inflata mentions 13 whorls and 4.8 mm,

but the shell is less mature than the type

specimens of I. ibex. Comparison of the

types shows a different sculpture, the

suture is deeper on I. ibex and the

sutural threads are less prominent.

It is more difficult to distinguish it

from other species of Inella and this

shall be done below.

Inella longissima (Dalí, 1881) (Figs. 10A-G, 36D)

Triforis (Ino) longissimus Dalí, 1881. Bull. Mus. Comp. Zool., 9: 80. [Type locality: Not stated].

Triforis (Inella) longissima Dalí, 1889. Bull. Mus. Comp. Zool., 18: 246, pl. 20, fig. 10. [Habitat:

Sigsbee knoll, off Havana, in 320-823 m].

Type material: One syntype, here designated as lectotype (Fig. 10A). Off Havana, Cuba 22° 9' N

82° 21' 30W, 175 m. Blake 1877-1878 Exped. Sta. 56 (MCZ 7381).

Other material studied: Bahamas: 1 f, Tamarind, Grand Bahama Island 26° 30' 45"N, 78° 36' 00"

W (J. Worsfold) (ANSP 368279); Florida: 1 j (Fig. 10F) with protoconch, off Alligator Reef Light,

lower Florida Keys (J. Moore, 1965) (ANSP 312592); 1 s (Fig. 10B), 1 j, SW of Egmont Key, (D. Steger,

1966) 366-420 m (ANSP 306391); 1 s, Monroe Co., 205° off Sombrero Light, (McGinty, Jul. 1951),

335-367 m (FLMNH 249518).

Description: See Dall (1881). Shell

(Fig. 10A) elongate, solid, whitish. The

protoconch is not known from the syn-

type, but a juvenile (Fig. 10F), which

could be this species, shows a paucispi-

ral protoconch of about 400 pm, with a

pupoid apex and three whorls with two

spiral cords. On this juvenile, the begin-

ning of the teleoconch has only two spi-

rals, but after a few whorls the third

very small spiral appears between them.

The teleoconch in large shells can reach

up to 27 or more whorls (Fig. 10B). The

profile of the whorls is straight, with

four spirals: spirals 1 and 3 are almost

similar in size and between them is spi-

ral 2, which is initially slightly smaller

but is subsequently of similar size; very

cióse to the suture there is one more

smooth spiral cord (Fig. 10D). Spiral 3 is

a little more prominent than spirals 1-2.

The nodules are always rounded.

Dimensions: The lectotype measures

26.7 mm.

A larger specimen, supposedly of

this species, has a more prominent

spiral 3 on the final whorls (Fig. 10B)

and the nodules are a little larger.

Distribution: Known from Bahamas,

Florida and Cuba.

Remarks: The syntypes with this

label from USNM and MCZ were exam-

ined. The problem is that the two lots

apparently represent two different

species. In this situation, after the

description of Dall (1881) was carefully

revised, it was decided to desígnate as

lectotype the syntype in MCZ, due to

the fact that the characters of this shell

more closely matched the original

description. In this description, the

author pointed out that: "all the shells

were decapitated" (which is not the case

in the USNM syntype), and he only

mentioned three spiral cords; further-

more, the shell designated as lectotype

has characters that are more similar to

those of the figure of I. longissima repre-

sented in Dall (1889: pl. 20, fig. 10)(Fig.

36D), and is cióse to 26 mm in size. The

syntype in USNM (with the label T.

longissimus ) has a size of 22 mm, differ-

ent to the dimensions mentioned by

Dall (1881), has a perfect protoconch

and has only two spirals on the first 6-7

whorls of the teleoconch; additionally, a

third and fourth cord appear, but these

differ from the figure in Dall (1889).

Also the siphonal canal is larger. For all

these differences it was considered to be

a sepárate species, and will be described

below as new ( I . pseudolongissima). The

shell from ANSP (Fig. 10B) may be the

same species, but spiral 2 is very small

100

ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 7. Marshallora ostenta spec. nov. A, B: holotype, 4.2 mm, MNCN; C: paratype, 3.6 mm

(IES); D: shell, 3.9 mm, specimen destroyed for radular studies; all from Cienfuegos, Cuba; E:

paratype, 3.9 mm, off Mayport, Florida (FLMNH); F, G: paratypes, 3.3 and 5.4 mm, off Big Pine

Key, Florida (BMSM); H: paratype, 3.5 mm, off St. Augustine, St. Johns Co., Florida (USNM); I:

paratype, 5.5 mm, off Big Pine Key (CHL); J: paratype, 5.5 mm, Florida (FLMNH); K: paratype,

5.3 mm, Florida (FLMNH); L: radula from a specimen from Cienfuegos; M: radula from a speci-

men from Florida; N: operculum, from Cienfuegos.

Figura 7. Marshallora ostenta spec. nov. A, B: holotipo, 4,2 mm, MNCN; C: paratipo, 3,6 mm (IES);

D: concha, 3,9 mm, ejemplar destruido para estudios radulares; todos de Cienfuegos, Cuba; E: paratipo,

3,9 mm, fuera de Mayport, Florida (FLMNH); F, G: paratipos, 5,5 y 5,4 mm, fuera de Big Pine Key,

Florida (BMSM); H: paratipo, 3,5 mm, fuera de St. Augustine, St. Johns Co., Florida (USNM); I:

paratipo, 5,5 mm, fuera de Big Pine Key (CHL); J: paratipo, 5,5 mm, Florida (FLMNH); K: paratipo,

5.3 mm. Florida (FLMNH); L: rádula de un ejemplar de Cienfuegos; M: radula de un ejemplar de

Florida; N: opérculo, de Cienfuegos.

101

Iberus, 26 (1), 2008

from the first teleoconch whorls, and on

the final whorls spiral 3 is more promi-

nent than on the lectotype. However, as

these minor differences probably only

indícate some variation in the species,

the shell has been kept in this taxon.

In spite of there being no information

about the protoconch of the lectotype of

this species, it has been considered to be-

long to the genus Inella on the basis of the

protoconch present on a juvenile, which

probably represents the same species.

Inella pseudolongissima Rolán and Femández-Garcés spec. nov. (Figs. 11 A-H)

Type material: Holotype (Fig. 11 A), a paratype (Fig. 11D) and a fragment (USNM 87316: syntypes

of Triforis longissimus ) from the type locality; 1 paratype (Fig. HE), 26° 40' N (M.E. Powlus, July

1960) (McGinty coll.) 77 m, (FLMNH 171183); 1 paratype (Fig. 11G), Hillsborough Co., Tampa,

Florida, West of Egmont Key (J. Moore, ]un. 1962) (McGinty coll.) 366 m (FLMNH 158180).

Type locality: Cuba, off Havana, 823 m.

Etymology: The specific ñame alludes to the similarity with 1. longissima, with which this species

was confused.

Description : Shell (Figs. 11A, 11D, 11E,

IIG) very elongate, irregularly white or

creamish. Protoconch (Figs. 11B, 11F,

IIH) multispiral with nearly 4 whorls,

almost 1 mm in height and with a diam-

eter of about 500 pm, sculptured with 2

spiral cords that are more evident on the

last whorls; these two cords are unap-

parent or only insinuated on the first two

whorls. Teleoconch with between 18 and

24 whorls, the first whorls with two

beaded spiral cords (spirals 1 and 3), the

nodules being rounded; below, on

whorls 3-4, a small, very narrow cord

(spiral 2) appears between them; this

new cord is always smaller than the

other two and closer to the upper one;

spiral 3 subsequently increases in size

and is very clearly the largest on the last

4-5 whorls, the other two being smaller;

also an additional small, unbeaded cord

appears at the suture. Axial ribs are

almost not appreciable; the growth lines

are prosocline. Aperture rhomboid,

siphonal canal short and open.

The colour of the holotype is totally

white. Paratypes are cream with some

areas of light brown.

Dimensions : The holotype measures

22.0 mm.

Distribution : Known from Florida

and Cuba.

Remarles: I. pseudolongissima may be

distinguished from I. longissima by teleo-

conch sculpture, the latter species

having three cords of equal size on most

of the spire. Also spiral 2 appears on the

first whorls of I. pseudolongissima but on

about the seventh whorl in I. longissima.

The protoconch of I. longissima has three

whorls with two well defined cords,

while the protoconch of I. pseudolongis-

sima has four smooth whorls, and the

apex is smaller.

For differentiation from other cióse

species, see below.

Inella noduloides spec. nov. Rolán and Fernández-Garcés (Figs. 12 A, B)

Type material: Holotype (Figs. 12A, 12B) in ANSP (374588).

Type locality: Tamarind, Grand Bahama Island, Bahamas 26° 30' 45"N, 78° 36' 00" W, 500 m (J.

Worsfold, May, 1985).

Etymology: The ñame alludes to the prominent nodules of the whorls.

Description: Shell (Figs. 12A, 12B) small,

subcylindrical, elongate, whitish and solid.

Protoconch with a pupoid apex of about

380 pm diameter and 3 whorls, each with

two smooth spiral cords; the lower cord

becomes more prominent near the end,

where another smooth, less prominent

cord appears above the other two, very

102

RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 8. A-D. Marshallora apexdiversus spec. nov. A: holotype, 6.2 mm, Florida (FLMNH); B-D:

paratypes, 6.9, 6.5, 4,6 mm, Florida (FLMNH). E-I. Protoconchs of Marshallora species. E, F: M.

ostenta , Florida; G: M. apexdiversus , Florida; H: M. modesta, Cuba; I: M. nigrocinta, Cuba.

Figura 8. A-D. Marshallora apexdiversus spec. nov. A: holotipo, 6,2 mm, Florida (FLMNH); B-D:

paratipos, 6,9, 6,5, 4,6 mm, Florida (FLMNH). E-I. Protoconchas de diferentes especies de Marshal-

lora. E, F: M. ostenta. Florida; G: M. apexdiversus. Florida; H: M. modesta, Cuba; I: M.

nigrocinta, Cuba.

103

IberuSy 26 (1), 2008

cióse to the suture; under high magnifi-

cation numerous tubercles can be seen. On

the first two whorls of the teleoconch there

is one prominent cord in the middle and

another smaller one below the suture; both

ha ve prominent nodules numbering about

10-11 per whorl. On the subsequent whorls

there is a small nodulous cord below the

suture with another small, slightly nodu-

lous one below it; these are followed by

the most prominent cord, with rather

sharply pointed nodules, and one more

below it, cióse to the lower suture. The

aperture is rounded but irregular, as the

shell is juvenile.

Dimensions : The holotype measures

2.1 mm.

Distribution : Only known from the

type locality.

Remarks: No other species has these

characters, with one spiral cord with

very prominent nodules and 3 weaker

spirals. The closest species may be the

new species I. harryleei (see below).

Inella apexbilirata spec. nov. Rolán and Fernández-Garcés (Figs. 12C-E)

Type material: Holotype (Fig. 12C) (ANSP 367841); 4 paratypes (ANSP 367840) (Fig. 12D). All from

the type locality.

Type locality: Lucaya, Grand Bahama Island, Bahamas 26° 29' 45" N, 78° 37' 15" W (J. Wors-

fold), 300 m.

Etymology: The specific ñame alludes to the beginning of the protoconch, with two spirals com-

mencing from the nucleus.

Description : Shell (Figs. 12C, 12D)

small, subcylindrical, elongate, shiny,

whitish and solid. Protoconch (Fig. 12

E), shining white, with a very small

nucleus, a diameter of nearly 400 pm

and a height of about 660 pm, 3 whorls

with two smooth spiral cords which

begin just on the nucleus. The teleo-

conch has two main beaded cords, the

lower one slightly larger; one smooth

smaller cord occurs at the suture. The

nodules on the main spirals are pointed

and cut at the middle. The axial ribs

which connect the nodules are scarcely

evident and are slightly prosocline.

There are about 13 on the first whorls

and 17 on the last. Aperture ovoid,

siphonal canal very short and open.

Dimensions : The holotype measures

3.5 mm.

Distribution : Only known from the

type locality.

Remarks: The species with two main

spirals are the following:

I. bigemma and I. inflata have whorls

that increase quickly in width, and the

latter species has a pupoid protoconch.

I. harryleei spec. nov. (see below) has

spirals 1 and 3 at the beginning, spiral 2

appearing later and being smaller.

I. ibex has the same number of

nodules in spite of being a more devel-

oped shell. Besides the two main spirals

there is only one more near the suture.

I. torticula has three spirals on the

first whorls and 4-5 on the last ones.

I. colon has two spirals but equidis-

tant from both sutures.

I. compsa also has two main spirals,

with spiral 1 being smaller; its proto-

conch is unknown, but the whorls on

this shell increase rapidly in width.

I. enopla and I. meteora have a very

different wide, pupoid protoconch.

Inella pinarena Espinosa, Ortea and Fernández-Garcés, 2007 (Figs. 12F-J)

Inella pinarena Espinosa, Ortea and Fernández-Garcés, 2007. Avicennia, 19: 73, fig. 46. [Type local-

ity: Yemayá, Maria la Gorda, Guanahacabibes, Pinar del Rio, Cuba],

Description: See Espinosa, Ortea and 12F. 12G), are shown in SEM pho-

Fernández-Garcés (2007). Shells (Figs. tographs. We add some informations on

104

ROLÁN AND FERNÁNDEZ-GARCÉS: New data on the Caribbean Triphoridae

Figure 9. A. Cerithium bigemma Watson, 1880. lectotype, 13.5 mm (BMNH). B. Inella bigemma ,

shell, 13.0 mm (BMNH). C. Cerithium inflatum Watson, 1880, lectotype, 4.8 mm (BMNH).

Figura 9. A. Cerithium bigemma Watson, 1880. lectotipo, 13,5 mm (BMNH). B. Inella bigemma,

concha, 13,0 mm (BMNH). C. Cerithium inflatum Watson, 1880, lectotipo, 4,8 mm (BMNH).

the protoconch (Figs. 12H, 121): small

tubercles are present on the protoconch

surface between the spiral ribs. The

microsculpture of the teleoconch (Fig.

12J) is formed by very small spiral striae,

more evident on the subsutural part.

Distribution : Only known from the

type material.

Remarks : This species was recently

described but, in the original descrip-

tion, only one optical photo was pre-

sented. More details on the shell, proto-

conch and microsculpture are here illus-

trated, showing that there is no similar-

ity with other species of the group in the

Caribbean.

Inella harryleei spec. nov. Rolán and Fernández-Garcés (Figs. 13A-K)

Type material: Holotype (Fig. 13A) and 2 (s) paratypes in FLMNH; one more in BMSM (Fig. 13B)

all ex CHL; another in CHL (Fig. 13C); all from the type locality. Other paratypes: 9 j and f, 28° 4.57'

N, 90° 59.99' W, 87.9 m (E.F. Garcia /Pelican, 9/1-5/01) (CHL); 1 j, from the same locality (USNM);

2 j more 26° 06.866' N, 91° 02.418' W, off Louisiana, 59-65 m (E.F. Garcia /Pelican, 6/30/01)(CHL);

105

Iberus, 26 (1), 2008

1 f, off Alligator Reef Light, Lower Florida Keys (J. Moore, 1965) (ANSP 312593); 3 s, SW of Egmont

Key, Florida (D. Steger, 1966) 366-420 m (ANSP 306391); 3 j, W of Tampa Bay, Florida (Richardson)

500 m (ANSP 335492); 1 s, Hillsborough Co., 110 miles SW of Egmont Key, Florida (D. Steger, 1953)

165 m (FLMNH 151609); 1 s (Fig. 13D), Hillsborough Co., SW of Egmont Key, Florida (J. Moore,

1967) 46 m (FLMNH 158178).

Other material examined: 2 f, Stn. 15, Florida (MCZ 356092) which are not in sufficiently good con-

dition to examine all their characters and attribute them with certainty to this species; 1 f (A.M.

Norman coll.) (Blake Expedition), West Indies (BMNH acc. n° 2283).

Type locality: off Dry Tortugas, Monroe Co., Florida, 90 m (J. Moore/ Cavalier, 1992).

Etymology: The species is named after Harry G. Lee, American malacologist who lent us the mate-

rial on which this species is based.

Description : Shell (Figs. 13A-13D)

elongate, very large for the genus, solid,

well sculptured. Protoconch (Figs. 13G-

13K) with between 3 and 3 V4 whorls,

bulbous, white, with barely discernible

separation from the teleoconch, the first

whorl smooth and the next with two

spiral cords that are sometimes a little

irregular; on the last whorl the lower

cord is more prominent, and the upper

one may divide in two. The teleoconch

has many whorls, 22 in the holotype

and 17 in a paratype (Fig. 13B). The first

whorls have two spiral nodulous cords

(spirals 1 and 3), connected by very

narrow prosocline ribs; below, around

the fourth or fifth whorls, spiral 2

appears between those cords and

remains very narrow for several whorls;

after the tenth whorl spiral 3 becomes a

little larger, and on the subsequent

whorls it is more clearly different and

prominent (Fig. 13E). Spiral 2 strength-

ens but continúes to be the smallest cord

until the end. The suture is very deep.

The axial ribs are slightly prosocline.

Aperture (Fig. 13F) ovoid, siphonal

canal narrow and large, dark in colour.

Shell colour light brown or white with

irregular brown axial flammules.

Dimensions : The holotype measures

18.3 mm. The paratypes are smaller.

Distribution : Only known from the

type locality.

Remarks: The present species must be

compared to and differentiated from the

following:

I. longissima is uniformly white and

more cylindrical, having three equal spi-

ral cords throughout most of the spire,

and another smaller one on the suture

which is relatively shallower; the shell

sculpture is very similar from the begin-

ning to the end with only a little more

prominence of the lowermost cord.

I. pseudolongissima spec. nov. is white

with 4 cords, the upper one smooth at

the suture, the following two very small,

and only the lowermost is clearly

prominent; the suture is not deep; the

protoconch has more whorls.

I. noduloides spec. nov. has one

prominent beaded spiral cord with three

more smaller ones which are absent

from the first whorls of the teleoconch

of I. harryleei.

Other elongate deep water species

are Inella colon Dalí, 1881, and I. ibex

Dalí, 1881; both have only two spiral

beaded cords per whorl.

Inella aff. harryleei (Figs. 14A, B)

Material examined: 1 s, Manatee Co., West of Anna María Key, Florida (J. Moore, 1961) (McGinty

coll.) 55 m (FLMNH 158177).

Remarks : The specimen studied is in

perfect condition and appears rather

similar to I. harryleei spec. nov., but it

differs in the following characters: a

more pointed protoconch, with about

two whorls and an angulation on the

second whorl, without visible cords; on

the teleoconch, spiral 1 is not larger than

spiral 2, but is less prominent; on the

base, below spiral 3, there is only one

106

RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 10. Inella longissima (Dalí, 1881). A: lectotype, 26.7 mm, off Havana, 175 m (MCZ 7381);

B: shell, 27.9 mm (ANSP); C, D: detail of the spire of lectotype; E: fragment of first whorls, 6.3

mm (ANSP); F: juvenile with protoconch (ANSP 312592); G: detail of last whorls of the shell

from Figure B.

Figura 10. Inella longissima (Dalí, 1881). A: lectotipo, 26,7 mm, fuera de La Habana, 175 m (MCZ

7381); B: concha, 27,9 mm (ANSP); C, D: detalle de la espira del lectotipo; E: fragmento de las

primeras vueltas, 6,3 mm (ANSP); F: juvenil con protoconcha (ANSP 312592); G: detalle de las

últimas vueltas de la concha de la Figura B.

107

Iberus , 26 (1), 2008

smooth cord and another very small one

immediately below. The siphonal canal

is shorter.

Dimensions : The only shell studied

measures 28.8 mm.

All these differences indícate that

this shell may represent a séparate

species, but they are not enough to

justify naming it. Additional material is

required.

Inella triserialis (Dalí, 1881) (Figs. 14C-E, 36E)

Triforis (Inella) triserialis Dalí, 1881. Bull. Mus. Comp. Zool., 9: 84. [Yucatán Strait, 640 fms (1171 m).

Off Cape San Antonio, 640 fms (1171 m). Station 2, 805 fms (1472 m)].

Triforis (Inella) triserialis Dalí, 1881. In Dalí, 1889. Bull. Mus. Comp. Zool., 18: 249, pl. 20, figs. 5a, 6a.

Triphora aspera auct. non Jeffreys, 1885.

Type material: Two syntypes (USNM 87319), one of them (Fig. 14C) measuring 6.4 mm is here des-

ignated as lectotype; the other shell (5.6 mm) would be a paralectotype (Fig. 14D). Both from Cape

San Antonio, Cuba, 1170 m; probably, 1 syntype (BMNH Acc. n° 2283), off Havana, 823 m (in very

bad condition).

Other material studied: 1 j, SW of Egmont Key, Florida (Steger, 1966) 366-421 m (ANSP 306391).

Description : See Dall (1881). Proto-

conch paucispiral (Fig. 14E), wide, with

about 3 whorls which have two spiral

cords, the apex depressed and smooth.

The teleoconch begins with three spiral

nodulous cords, spiral 2 being smaller

and closer to spiral 1. On the following

whorls spiral 1 increases in size, with

nodules that are larger than those on the

other cords and are cut at the middle.

These differences continué up to the end

of the shell.

Dimensions : The lectotype measures

6.4 mm.

Distribution: Known from Yucatán

(México) and Cuba.

Remarks: As with other taxa, the

syntype material found in USNM and

MCZ has been examined. The material

from USNM is coincident in protoconch

and teleoconch sculpture with the

description of Dall (1881) and with one

of the figures in Dall (1898: píate 20,

fig. 5a) (Fig. 36E) but not with the other

figure (Dall, 1898: píate 20, fig. 6a),

which seems to be a different species

with a sharply pointed apex. The size of

the lectotype is slightly smaller than that

mentioned in the original description

(8.25 mm), but it may be that this type

could be partially broken. The material

from MCZ seems to be what Dall

named as var. intermedia (see below).

Dall (1989: 246) considered this

taxon in relation with Triphora aspera Jef-

freys, 1885 (preoccupied, renamed

Triphora brychia Bouchet and Guillemot,

1978; assigned to genus Strobiligera Dall,

1924 by Bouchet and Warén, 1985). He

therefore referred to Triforis (Inella) trise-

rialis var. aspera Jeffreys, 1885, which

perhaps can correspond with the figure

previously mentioned (Fig. 36E). A

syntype of Jeffreys' taxon (Fig. 14F,

BMNH 85.11.5.2673) has been examined,

and is here designa ted as lectotype. It is

a larger and sharply pointed shell with a

multispiral protoconch, and there is no

indication that this species occurs in the

Caribbean.

I. triserialis is different from most of

the Caribbean species referred to in this

work, mainly due to its spiral 1 having

larger nodules. Comparison will be

made in the Remarks of some species

described below (such as I. sarissa).

Inella intermedia (Dall, 1881) (Figs. 14G-J, 36F)

Triforis intermedius Dall, 1881. Bull. Mus. Comp. Zool., 9: 85. [Yucatán Strait, 1170 m; off Cape San

Antonio, 1170 m; station 2, 1472 m].

108

RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 11. Inella pseudolongissima spec. nov. A: holotype, 22.0 mm, off Havana, 823 m (USNM);

B: protoconch of the holotype; C: detail of the spire; D: paratype, 10.4 mm (USNM); E:

paratype, 18.5 mm (FLMNH); F: protoconch of this paratype; G: paratype, 23.4 mm (FLMNH);

H: protoconch of this paratype.

Figura 11. Inella pseudolongissima spec. nov. A: holotipo, 22,0 mm, fuera de La Habana, 823

(USNM); B: protoconcha del holotipo ; C: detalle de la espira; D: paratipo, 10,4 mm (USNM);

paratipo, 18,5 mm (FLMNH); F: protoconcha de este paratipo; G: paratipo, 23,4 mm (FLMNH);

protoconcha de este paratipo.

109

Iberas, 26 (1), 2008

Tñforis triserialis var. intermedia Dalí, 1881. In Dall, 1889. Bull. Mus. Comp. Zoo/., 18: 287, píate 20,

fig. 8. [Habitat: 805 fms (1473 m). Barbados, 183 m].

Triphora clenchi Aguayo, 1935. Unnecessary replacement ñame pro Triforis intermedius Dall, 1881.

Type material: Syntype (Fig. 14G) here designated as lectotype (MCZ 7384) (off Cuba, Yucatán

Strait and Cape San Antonio. Depth: 1171 m. Blake 1877-1878 Exped.); one paralectotype (Fig.

14H)(MCZ 7382) (Havana, off Morro Light. Depth: 1473 m. Blake 1877-1878 Exped.). Both shells

were labeled T. triserialis.

Description : See Dall (1881). No pro-

toconch was known and the author

refers to "nucleus missing". Two spirals

of similar strength (spirals 1 and 3) are

present on the first whorls of the shell;

spiral 2 appears below and is clearly

smaller up to near the end of the spire,

where the three spirals are almost similar

in size (Fig. 141). Axial ribs prosocline,

and the nodules are spherical.

Dimensions : The lectotype measures

7.9 mm.

Distribution : From Yucatán, México,

Barbados and Cuba.

Remarks: The lectotype here selected

is very similar to the shell represented in

Dall (1989: pl. 20, fig. 8) (Fig. 36F) but it

is not coincident with the description,

which refers to a size of 11 mm and 23

whorls; also the figure does not agree

with the description, because in spite of

the fact that a height of 11 mm is men-

tioned, there are not more that 15-16

whorls. The shell here selected as lecto-

type is only 7.9 mm and has 12 whorls,

and it may be the original shell which

lost some part of the spire. Anyway, it

must be kept as lectotype in order to

attain nomenclatural stability. The para-

lectotype is 7.2 mm and has about 13

whorls (MCZ 7382).

The ñame Triphora triserialis clenchi

Aguayo, 1935 is an unnecessary replace-

ment ñame for Triforis intermedius Dall,

1881 (which is the same taxon as Triforis

triserialis var. intermedia Dall, 1881 (in

Dall, 1889)). This is due to the fact that

this ñame is not preoccupied by

Cerithium intermedium C. B. Adams,

1850; initially both belonged in different

genera and finally the first one is placed

in Inella and the second was recently

placed by Rolán and Fernández-

Garcés (1995) in the genus Similiphora

Bouchet, 1985.

Lack of knowledge of the proto-

conch is a problem when comparing this

species to others from the Caribbean.

Basing this comparison on the teleo-

conch, the following species must be

taken into account:

Inella longissima has a more cylindri-

cal, more elongate shell, spiral 2 is of

similar size to the other two spirals on

most of the spire, while in I. intermedia it

is always smaller.

Inella pseudolongissima spec. nov. has

spiral 3 more prominent and with larger

nodules.

Inella harryleei spec. nov. has a typi-

cally deep suture, spiral 3 being the

most prominent, and spiral 1 is also

larger than spiral 2, these differences

being maintained to the end of the shell.

Inella triserialis has spiral 1 more

prominent and with larger nodules.

Inella ibex (Dall, 1881) (Figs. 14K-M, 36G)

Triforis ibex Dall, 1881. Bull. Mus. Comp. Zool., 9: 86. [Type locality: off Cape San Antonio, 1171 m;

Yucatán Strait, 1171 m; Sigsbee, off Havana, 823 m].

Triforis (Sychar) inflata var. ibex Dall, 1889. Bull. Mus. Comp. Zool, 18: 249, pl. 20, fig. 12b.

Type material: A syntype (f) of 10.4 mm here designated as lectotype (Fig. 14K), off Cuba, Yucatán

Strait and Cape San Antonio, 1171 m (MCZ 7391); another syntype, now paralectotype, f of 7.7 mm,

off Havana, Cuba 823 m (MCZ 7392). Another paralectotype (Fig. 14L) (USNM 87313), off San

Antonio, Cuba, 1189 m.

110

RoláN AND FernÁNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 12 A, B. Inella noduloides spec. nov. holotype, 2.1 mm, Grand Bahama Island (ANSP). C-

E. Inella apexbilirata spec. nov. C: holotype, 3.5 mm, Grand Bahama Island (ANSP); D: paratype,

3.2 mm, Grand Bahama Island (ANSP); E: protoconch of the paratype. F-J. Inella pinarena

Espinosa, Ortea and Fernández-Garcés, 2007. F: paratype, 6.0 mm (CFG); G: shell, 5.0 mm

(MHNS); H: protoconch of the paratype; I: detail of the protoconch; J: microsculpture.

Figura 12. A, B. Inella noduloides spec. nov. holotipo, 2, 1 mm, Grand Bahama (ANSP). C-E. Inella

apexbilirata spec. nov. C: holotipo, 3,5 mm, Grand Bahama (ANSP); D: paratipo, 3,2 mm, Grand

Bahama (ANSP); E: protoconcha del paratipo. F-J. Inella pinarena Espinosa, Ortea and Fernández-

Garcés, 2007. F: paratipo, 6,0 mm (CFG); G: concha, 5,0 mm (MHNS); H: protoconcha del

paratipo; I: detalle de la protoconcha; J: microescultura.

111

Iberus, 26 (1), 2008

Type locality: The three localities mentioned in the original description are written on the label of

the lectotype, and henee there is no exact information about which locality applies to this type.

Description : See Dall (1881). The orig-

inal description mentions a nucleus twice

as large as that of I. colon , smooth, inflated,

rounded on top, larger than the succeed-

ing nuclear whorl, which has two incon-

spicuous narrow keels which are wavy

and almost tuberculate at first, and merge

almost imperceptibly into the usual sculp-

ture of the shell. The type material studied

has no protoconch, which obviously was

lost. The most important characters of the

teleoconch are: shell (Figs. 14K, 14L) with

two rows of nodules throughout the spire

and with one more, smaller and not nodu-

lous, occurring at the suture at the end the

spire (Fig. 14M). The nodules are cut at

the middle. The axial ribs are scarcely

appreciable and slightly prosocline.

Dimensions : The lectotype measures

10.4 mm.

Distribution: Known from Yucatán

(México) and Cuba.

Remarles : Triforis ibex is the type

species of the genus Strobiligera Dall,

1924. The problem is that we have no

image or a good description of the pro-

toconch of this species, and the original

description allows us to think that the

protoconch is almost smooth, the first

whorl wider than the second and with

two whorls, which is typical of a proto-

conch of an Inella species. In this case;

Strobiligera could be a synonym of Inella.

The assignment of this species to the

genus Inella is tentative, The lectotype

here designated is similar to the shell

represented in Dall (1889, pl. 20, fig.

12b) (Fig. 36G) and also its dimensions

are closer to those referred to in the orig-

inal description.

Comments on the differences of I.

ibex from other species with two spiral

beaded cords can be seen below in I.

colon, which is the most similar species.

Inella torticula (Dall, 1881) (Figs. 15, 36H)

Triforis torticulus Dall, 1881. Bull. Mus. Comp. Zoo/., 18: 82. [Type locality: Yucatán Strait, 1170 m],

Triforis (Sychar) torticula Dall, 1881. In Dall, 1889. Bull. Mus. Comp. Zool., 18: 249, pl. 20, fig. 11b.

Type material: One syntype (Fig. 15A) (f) here designated as lectotype, 6.7 mm, off Cuba, Yucatán

Strait, 1171 m, Blake 1877-1878 Exped. (MCZ 7390); the syntypes in USNM have been lost (Ellen

Strong, pers. comm.).

Other material examined: 1 f (Fig. 15B) (labeled as Triphora bigemma Watson, 1880), 9.7 mm, 9

whorls, Yucatán Strait, 1171 m (MCZ 7388); 1 s (Figs. 15C, 15D), 25.0 mm, Straits of Florida, 1000-

1200 m (CHL).

Description: See Dall (1881). The

most important characters are: 4 spirals,

the lowermost spiral 4 very narrow and

very cióse to the suture; spiral 1 smaller

than spirals 2 and 3, which are equal in

size and prominence. Colour whitish.

The study of an almost complete adult

shell (without protoconch) with 17.5

whorls (Figs. 15C, 15D) (CHL), allows

us to add some information: at the

beginning of the shell there are only

three spirals, almost similar in size, but

the uppermost slightly smaller; later, the

differences between spiral 1 and spirals

2-3 are more evident, and at same time

spiral 4 appears; on about whorl 13 (on

this shell) the uppermost spiral 1 is

divided in two, there being 5 spirals at

the end of the shell (Fig. 15E). The ribs

are orthocline.

Dimensions: see below.

Distribution: Known from deep

water off Florida and Cuba.

Remarks: The placement of this

species in the genus Inella is tentative, as

the protoconch and the aperture are not

known.

The syntype in MCZ (Fig. 15A), here

designated as lectotype, is a shell frag-

ment that seems to be the one figured in

112

RolAn and FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 13. Inella harryleei spec. nov. A: holotype, 18.3 mm, off Dry Tortugas, 90 m (CHL); B:

paratype 11.7 mm (BMSM); C: paratype, 12.8 mm (CHL); D: paratype, 14.6 mm (FLMNH). E:

detail of the spire of the holotype; F: detail of the aperture of the paratype in figure B; G-K: proto-

conchs of several paratypes (CHL).

Figura 13. Inella harryleei spec. nov. A: holotipo, 18,3 mm, fuera de Dry Tortugas, 90 m (CHL); B:

paratipo 11,7 mm (BMSM); C: paratipo, 12,8 mm (CHL); D: paratipo, 14,6 mm (FLMNH). E:

detalle de la espira del holotipo; F: detalle de la abertura del paratipo de la figura B; G-K: protoconchas

de varios paratipos ( CHL).

113

Iberus , 26 (1), 2008

Dall (1889, pl. 20, fig. 11b) (Fig. 36H),

although a little more cylindrical,

having only 4 whorls instead 6 and

measuring 6.7 mm instead of 10.5,

which could be due to the loss of two

whorls of the shell. As it is compatible

with the description, it was thought that

it should be designated as lectotype in

order to maintain nomenclatural stabil-

ity. The other shell examined (Fig. 15B)

measured 9.7 mm and had 9 whorls, the

label without any indication of "type"

and with the ñame Triphora bigemma; it

is undoubtedly the same species.

The differences with I. bigemma Wat-

son are very small, bearing in mind that

although the latter species was de-

scribed as having only two rows of

beads, the figure and the type clearly

show three on the final whorls. How-

ever, this is a juvenile, and to make a de-

cisión it would be necessary to examine

the protoconchs and to compare several

samples of both species.

The other shell studied (Figs. 15C,

15D) has a profile slightly different from

the lectotype, with the whorls expand-

ing more rapidly and with 4-5 spirals on

the final whorls. This could represent a

different species, but it is rather coinci-

dent in some characters with the shell

figured by Dall (1889) (Fig. 36H) and

for this reason it is considered to be an

example of the variability of this taxon,

which has differences between the first

and the last whorls.

Inella colon (Dall, 1881) (Figs. 16, 361)

Triforis colon Dall, 1881. Bull. Mus. Comp. Zool., 9: 86. [Type locality: off Havana, Cuba 450 fms

(823 m); off Cape San Antonio, 1171 and 1834 mi.

Triforis (Inella) colon Dall, 1881. In Dall, 1889. Bull. Mus. Comp. Zool., 18: 249, pl. 20, fig. 12.

Type material: One syntype (Fig. 16A) (f) here designated as lectotype, 12.7 mm, with the label "off

Havana, 823 m" (USNM 87315), and another syntype (Fig. 16B) (f) 7.8 mm, now paralectotype, from

the same lot. A syntype (Fig. 16C) (f) 8.6 mm, now paralectotype, from Cape San Antonio, 1171-

1834 m (MCZ 7387).

Other material examined: 2 f of 16.2 and 5.6 mm, Egmont Key, Florida, Gulf of México, (J. Moor e/Cav-

alier, 1968), 180 m (CHL); 1 s (Fig. 16E), Yucatán, 21° 60' N, 86° 80' W, ENW of Contoy Fight, off

Cape Catoche (Riley Black, Apr. 1967) (McGinty coll.) 366-412 m (FLMNH 170221); 1 s (Fig. 16F),

24° 15' 28"N, 81° 57' 89"W, Monroe Co., Straits of Florida, 236-255 m (FLMNH 26660).

Type locality: With the designation of the lectotype, the type locality is restricted to off Havana,

Cuba 823 m.

Description: See Dall (1881). In the

original description the protoconch is

mentioned as follows: nucleus and first

nuclear whorls white, polished, smooth,

flattened on top. The most important char-

acters of the teleoconch are: shell almost

cylindrical (Figs. 16A-16C), white, with

two main very narrow spiral beaded cords

and two smaller, smooth, slightly wavy

cords located near the suture; one of them

above and the other below the suture, this

being almost imperceptible. The nodules

are small and slightly prominent; the

uppermost a little larger. The axial ribs

almost inapparent, a little prosocline;

whorls very short and fíat.

Dimensions: The lectotype measures

12.7 mm.

Distribution: Known from deep

water of Florida, Yucatán and Cuba.

Remarks: The lectotype (Fig. 16 A) is in

accordance with the original description

and it seems to be the shell figured in

Dall (1889: pl. 20, fig. 12) (herein Fig. 361),

having the same number of whorls and

almost the same dimensions. A paralec-

totype in the same lot as the lectotype (Fig.

16B) seems to be a different species,

because this shell has three spirals instead

of two. The paralectotype in MCZ (Fig.

16C) is a little different from the lectotype,

but it has the typical distribution of the

cords (Fig. 16D) and is probably the same

species. Some shells from the Gulf of

México (Figs. 16E and 16F) have been

included in this taxon but they could rep-

1 14

ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 14. A, B. Inella aff. harryleei, shell, 28.8 mm and protoconch (FLMNH). C-E. Inella triseri-

alis (Dalí, 1881). C: lectotype, 6.4 mm (USNM); D: paralectotype, 5.6 mm (USNM); E: proto-

conch of the paralectotype. F. Triphora aspera, lectotype, 13.0 mm (BMNH) ( =Strobiligera brychia).

G-J. Inella intermedia (Dalí, 1881). G: lectotype, 7.9 mm, off Cuba, Yucatán Strait and Cape San

Antonio, 1171 m (syntype of T. triserialis, MCZ 7384). H: paralectotype, 7.2 mm, (Havana, off

Morro Light, 1473 m (syntype of T. triserialis , MZS 7382); I: detail of the sculpture of the paralec-

totype. J: shell photographed by Kaicher (ANSP, unknown size and other data). K-M. Inella ibex

(Dalí, 1881). K: lectotype, 10.4 mm, off Cuba, Yucatán Strait and Cape San Antonio, 1 171 m (MCZ

7391); L: paralectotype, 9.2 mm (USNM 87313); M: detail of the spire of the lectotype.

Figura 14. A, B. Inella aff. harryleei, concha, 28,8 mmy protoconcha (FLMNH). C-E. Inella triserialis

(Dalí, 1881). C: lectotipo, 6,4 mm (USNM); D: paralectotipo, 5,6 mm (USNM); E: protoconcha del

paralectotipo. F. Triphora aspera, lectotipo, 13, 0 mm (BMNH) ( =Strobiligera brychia). G-J. Inella inter-

media (Dalí, 1881). G: lectotipo, 7,9 mm, Juera de Cuba, Estrecho de Yucatán y Cabo de San Antonio,

1171 m (sintipo de Y. triserialis, MCZ 7384). H: paralectotipo, 7,2 mm, (La Habana, Juera de Morro

Light, 1473 m (sintipo de T. triserialis, MZS 7382); I: detalle de la escultura del paralectotipo. J: concha

fotograjiada por Kaicher (ANSP, tamaño desconocido así como otros datos). K-M. Inella ibex (Dalí, 1881).

K: lectotipo, 10,4 mm, fuera de Cuba, Estrecho de Yucatán y Cabo de San Antonio, 1171 m (MCZ 7391);

L: paralectotipo, 9,2 mm (USNM 87313); M: detalle de la espira del lectotipo.

115

IberuSy 26 (1), 2008

resent a different species, because the

spirals are more prominent and are more

widely separated than on the lectotype.

However, the lack of protoconch and the

differences of the sculpture are not very

important, and so provisionally they ha ve

been kept in this species awaiting more

material in the fu ture.

Dall (1881) referred to the differ-

ences between I. íbex and I. colon, proba-

bly because he considered both species

very similar. Unfortunately at present

their protoconchs are not known, but

Dall was of the opinión that the proto-

conch of I. íbex was twice as large as that

of I. colon. In the teleoconch characters, I.

íbex has more rapidly expanding whorls.

the nodules are larger and are cut at the

middle, and fewer whorls occupy the

same space.

The other Caribbean species may

have two spirals at the beginning of the

teleoconch, but finally have three or

more spirals. This ineludes I. pompona

and I. meteora, although spiral 1 is very

small and sometimes difficult to see.

Inella unicornium Simone, 2006, from

Brazil, has a larger shell (up to 50 mm,

the biggest known species of Triphori-

dae), which according to the original

description may be differentiated from 1.

colon not only by its larger size, but also

by its proportionally longer whorls, taller

spiral cords and projecting outer lip.

Inella undebermuda spec. nov. Rolán and Lee (Figs. 17A-E)

Type material: Holotype (Fig. 17A) in FLMNH; a paratype (Fig. 17B), 12.3 mm, in USNM. Both

from type locality (J.R.H. Lightboum and A.T. Guest, 1988; ex J.R.H. Lightbourn 9/8/93; ex CHL).

Type locality: S. of Castle Roads, Bermuda, 90 m.

Etymology: The specific ñame derives from the Latín word unde "origin" and the ñame of the arch-

ipelago where the species was collected.

Description : Shell (Figs. 17A, 17B) elon-

gate, whitish and solid. Protoconch (Fig.

17C) white, without a clear separation from

the teleoconch, with about two whorls,

the nucleus smooth, one spiral cord

appearing in the middle of the first whorl

and becoming two on the second; the

diameter of about 380 pm. The first whorls

of the teleoconch have two spiral beaded

cords (spirals 1 and 3), and a very small

smooth and narrow thread very cióse to

the suture; between the 6-7* whorls, spiral

2, which is also narrow and smooth,

appears between the other two, closer to

spiral 1. On the subsequent whorls, spiral

2 increases slowly in size, becoming

slightly undulating and with nodules

appearing at the end of the spire (Fig. 17D),

but always remaining nearer to spiral 1.

The nodules of spirals 1 and 3 are larger

and cut at the middle. Aperture (Fig. 17E)

rhomboid, columella almost straight,

siphonal canal very short and curved.

Dimensions : The holotype measures

7.8 mm, and the paratype 12.3 mm.

Distribution : Only known from the

type locality.

Remarles : The holotype is the smallest

shell, but has a protoconch in good con-

dition. The paratype lacks the first proto-

conch whorl. The most important differ-

ential character is the smaller size of

spiral 2 and its location closer to spiral 1.

This species must be compared with

and separated from the following:

Inella longissima is more elongate and

cylindrical and has the three spirals similar

in size and distribution. In juveniles

assumed to belong to this species, the pro-

toconch has three whorls, and when spiral

2 appears on the first whorls of the teleo-

conch, it is in the middle between spirals

1 and 3 and not closer to the upper one.

Inella pseudolongissima spec. nov. has

a protoconch with four whorls and the

spirals of the teleoconch are different,

spiral 3 being larger and more promi-

nent than the others. Spiral 2 is only the

smaller one at the beginning.

Inella harryleei spec. nov. has three

whorls in the protoconch, and in the

teleoconch spiral 2 becomes nodulous

very early (between whorls 5-6) and is

not closer to spiral 1, and the suture is

1 16

ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 15. Inella torticula (Dalí, 1881). A: lectotype, 6.7 mm (MCZ 7390); B: shell, (labeled as T.

bigemma ), 9.7 mm, 9 whorls, Yucatán Strait, 1171 m (MCZ 7388); C, D: shell, 25.0 mm, Straits

of Florida, 1000-1200 m (CHL); E: detail of sculpture. F: shell photographed by Kaicher (ANSP,

lacking size and data).

Figura 15. Inella torticula (Dalí, 1881). A: lectotipo, 6,7 mm (MCZ 7390); B: concha, (etiquetada

como T. bigemma,), 9,7 mm, 9 vueltas, Estrecho de Yucatán, 1171 m (MCZ 7388); C, D: concha,

25,0 mm, Estrechos de Florida, 1000-1200 m (CHL); E: detalle de la escultura. F: concha fotografiada

por Kaicher (ANSP, careciendo de información sobre su tamaño y otros datos).

1 17

Iberus , 26 (1), 2008

more evident. Furthermore, the colour

consists of axial brown flammules and

brown siphonal canal, more elongate.

Inella intermedia has a shorter shell

and spiral 2 is strongly nodulous on the

early whorls, being almost the same size

as the other two and located at a similar

distance between them.

Inella ibex and I. colon have only two

main spirals.

Inella torticula can have up to five

spirals on the final whorls.

Inella sarissa (Dalí, 1889)

Triforis (Inella) sarissa Dalí, 1889. Bull. Mus. Comp. Zoo/., 18: 247. [Type locality: near Barbados, 13°

11.54' N, 59° 38.45' W, 134 m].

Type material: The lot in USNM labeled as "87314 Holotype" of this species is empty (E. Strong,

pers. comm.).

Description : See Dall (1889).The fol-

lowing characters must be pointed out

from the original description: three rows

of strong tubercles, the uppermost being

the largest and spiral 3 the next in size; spi-

ral 2 is the smallest and is cióse to spiral 1.

It is also mentioned that one specimen had

24 whorls, thus being an elongate shell.

Dimensions : 15.5 mm is mentioned

in the original description.

Distribution: Only know from its

type locality.

Remarks: Apparently only I. triseri-

alis has the uppermost spiral with

larger nodules, but the other two

spirals have the nodules rather similar

in size; none of the other species here

described have these characters.

Anyway I. triserialis has a shorter

shell.

Inella compsa (Dall, 1927) (Figs. 17F, G)

Triphora ( Strobiligera ) compsa Dall, 1927. Proc. U. S. N. M., 70: 96. [Type locality: Off Georgia].

Type material: Lectotype, here designated (Fig. 17F) 12.0 mm, and one paralectotype (Fig. 17G) 9.4

mm (USNM 333518).

Description : See Dall (1927).

Distribution : Only known from the

type locality.

Remarks : The description of the

species seems to have been based on the

characters of both lectotype and paralec-

totype, in spite of the fact that they may

be different species. The lectotype (Fig.

17F) is in accordance with the descrip-

tion with the exception that it has not

two but three prominent beaded cords

with almost orthocline ribs, but the

dimensions are those indicated in the

original description. Ffowever it is a

fragment of a shell with a broken aper-

ture and lacking a protoconch, and it is

therefore difficult to define exactly. The

paralectotype (Fig. 17G) has only two

spirals, with a third appearing on the

final whorls cióse to the undulating

thread near the suture, and it could be a

different species. Due to the lack of

additional material apart from these

mentioned types, it is preferable to wait

for more suitable material to study in

the future.

Inella pseudotorticula spec. nov. Rolán and Lee (Figs. 17H-L)

Type material: Holotype (Fig. 17H) in FLMNH; a paratype (Fig. 171) in USNM (both vyJ.R.H. Light-

boum, 9/8/93; ex CHL).

118

ROLÁN AND FERNÁNDEZ-GARCÉS: New data on the Caribbean Triphoridae

Figure 16. Inella colon (Dalí, 1881). A: lectotype, 12.7 mm, off Havana, 823 m (USNM); B: para-

lectotype, 7.8 mm, off Havana, 823 m (USNM); C: paralectotype, 8.6 mm, Cape San Antonio,

1171-1834 m (MCZ 7387); D: detail of the sculpture; E: shell, 20.3 mm, Yucatán (FLMNH); F:

shell, 29.4 mm, straits of Florida (FLMNH).

Figura 16. Inella colon (Dalí, 1881). A: lectotipo, 12,7 mm, fuera de La Habana, 823 m (USNM);

B: paralectotipo, 7,8 mm, fuera de La Habana, 823 m (USNM); C: paralectotipo, 8,6 mm, Cabo de

San Antonio, 1171-1834 m (MCZ 7387); D: detalle de la escultura; E: concha, 20,3 mm, Yucatán

(FLMNH); F: concha, 29,4 mm, estrechos de Florida (FLMNH).

119

Iberus , 26 (1), 2008

Type locality: South of Castle Roads, SE Bermuda, (J.R.H. Lightboum and A.T. Guest, 1988) 100 m.

Etymology: The specific ñame alludes to the similarity with I. torticula.

Description : Shell (Figs. 17H, 171)

very elongate, pointed, with blunt apex,

not straight in profile, solid. Protoconch

(Fig. 17J) with about 2 l/i whorls, with a

wide nucleus and two spiral cords, of

which the upper one is smaller and less

prominent; diameter of about 400 pm.

The teleoconch has about 23 whorls (on

the holotype) with spirals 2 and 3

present from the beginning, very similar

in size, and with the nodules cut in the

middle. On the early whorls spiral 1 is

very small and is located cióse to the

upper suture, with spiral 4 appearing

like a fine thread near the lower suture;

on about the ninth whorl, spiral 1 is

more clearly defined on the upper part

of the whorl, below the suture, but is

smaller and less prominent than the

other two, while spiral 2 is the most

prominent and slightly larger; on about

the seventeenth whorl, spiral 1 contin-

úes to be closer to spiral 2 (Fig. 17K). On

about the twenty-first whorl, the three

spirals are almost similar in size. Aper-

ture (Fig. 17L) ovoid with a short

siphonal canal.

The shell has some white whorls

alternating with entirely light brown

whorls. The cords are often light brown

between the nodules.

Dimensions : The holotype measures

16.8 mm.

Distribution: Only know from the

type locality.

Remarks : This species must be com-

pared with the following ones that have

elongate shells:

Inella longissima has spirals 1 and 3 from

the beginning, with spiral 2 appearing

below; at the end of the spire the spirals

are of almost equal strength, but spiral 2

is slightly smaller and spiral 3 a little more

prominent on most of the shell.

I. pseudolongissima spec. nov. has

spirals 1 and 3 from the beginning, with

spiral 2 smaller; on the final whorls

spiral 3 is noticeably larger than the

others; the colour is whitish. The proto-

conch has four whorls.

I. harryleei spec. nov. has spirals 1

and 3 from the beginning; spiral 2

appears after the first few whorls but

remains smaller throughout the shell; at

the end, spiral 3 is very large in compar-

ison with the others; the colour is white

or light brown with brown axial flam-

mules.

I. triserialis (Dalí, 1881) has a shorter

shell with three spirals, spiral 1 being

larger with more inflated nodules. The

apex is more depressed.

I. intermedia (Dalí, 1881) has a shorter

shell with spirals 1 and 3 present at the

beginning; spiral 2 is smaller and

appears later.

I. ibex (Dalí, 1881) and I. colon have

only two prominent beaded cords.

I. torticula (Dalí, 1881) has spiral 1

less prominent on the early whorls; sub-

sequently the final whorls are wider and

have up to 5 spirals.

I. undebermuda spec. nov. has spirals

1 and 3 at the beginning of the teleo-

conch, with spiral 2 appearing between

the other two, always remaining smaller

and closer to spiral 1.

I. unicornium, described by Simone,

2006 from Brazil, has some general simi-

larity with I. pseudotorticula, but the

holotype is a shell with 40 whorls that

measures almost 50 mm in height; also

it is mentioned in the original descrip-

tion that the protoconch has only one

whorl and the teleoconch whorls have

only two beaded spiral cords, with an

additional smooth cord appearing on

the final whorl.

Inella enopla (Dalí, 1927) (Figs. 18A-E)

Triphora enopla Dalí, 1927. Proc. U. S. N. M., 70: 95. [Type locality: Off Fernandina, Florida, 294

fms (538 m)].

120

RoláN AND FERNÁNDEZ-GARCÉS: New data on the Caribbean Triphoridae

Figure 17. A-E. Inella undebermuda spec. nov. A: holotype, 7.8 mm, S. of Castle Roads, Bermuda,

90 m (FLMNH); B: paratype, 12.3 mm, same locality (USNM); C: protoconch of the holotype;

D: detail of the sculpture, holotype; E: detail of the aperture, holotype. F, G. Inella compsa (Dalí,

1927). F: lectotype, 12.0 mm, off Georgia, 803 m; (USNM); G: paralectotype, 9.4 mm, off

Georgia, 805 m (USNM). H-L. Inella pseudotorticula spec. nov. H: holotype, 16.8 mm, SE

Bermuda (FLMNH); I: paratype, 8.5 mm, S. of Castle Roads, Bermuda (USNM); J: protoconch,

holotype; K: detail of the sculpture, holotype; L: detail of the aperture.

Figura 17. A-E. Inella undebermuda spec. nov. A: holotipo, 7,8 mm, S. de Castle Roads, Bermuda, 90

m (FLMNH); B: paratipo, 12,3 mm, la misma localidad (USNM); C: protoconcha del holotipo; D:

detalle de la escultura, holotipo; E: detalle de la abertura, holotipo. F, G. Inella compsa (Dalí, 1927).

F: lectotipo, 12,0 mm, fuera de Georgia, 805 m; (USNM); G: paralectotipo, 9,4 mm, fuera de

Georgia, 805 m (USNM). H-L. Inella pseudotorticula spec. nov. H: holotipo, 16,8 mm, SE Bermuda

(FLMNH); I: paratipo, 8,5 mm, S. de Castle Roads, Bermuda (USNM); J: protoconcha, holotipo; K:

detalle de la escultura, holotipo; L: detalle de la abertura.

121

Iberus, 26 (1), 2008

Type material: Lectotype (2 f, labeled as "type"), here designated (Figs. 18A, 18B) and 5 paralec-

totypes (3 j, 2 f) (USNM 108074).

Other material examined: 2 f (with the label " Triforis colon"), off Havana, Cuba 823 m (Blake Expe-

dition, Sta. 243) (BMNH acc. n° 2283).

Description : Dall (1927). Protoconch

(Figs. 18C, 18D) with the apex smooth,

pupoid, wide (about 500 pm), followed

by about 2 whorls with three very cióse

cords, the uppermost smaller and the

lowest one the largest and most promi-

nent. Teleoconch whorls with two spiral

nodulous cords and another two very

fine cords that are separated by the

suture (Fig. 18E); the subsutural one is

very small and has elongate nodules,

while the one just above the suture is

smooth. The nodules on the main spi-

rals are pointed and cut at the middle.

Dimensions : The lectotype consists of

fragments measuring 7.2 and 6.6 mm.

Distribution : Only known from

Florida and Cuba.

Remarks : The original description men-

tions 15 whorls and dimensions of 11.5 x

2 mm. The glass tube labeled "type" con-

tained two fragments (7.2 and 6.6 mm

respectively), but there were no apparent

indication thar they were previously unit-

edtherefore it is doubtful, although pos-

sible, that they carne from the same shell.

However, they give a good representation

of the specific characters, and as it is not

certain whether or not they are from the

same broken shell, it is better to keep both

fragments together and consider both as

parts of the lectotype.

The large pupoid protoconch is the

most important differential character.

The most similar species are:

I. inflata has the protoconch whorls

with two scarcely visible spiral cords,

the nodules of the teleoconch smaller

and sharply pointed, the shell with

rapidly enlarging whorls.

Inella bigemma has the shell with more

rapidly enlarging whorls, spiral 1 more

developed and more clearly nodulous.

I. ibex is not so cylindrical and has

more pointed nodules, with only one

small thread near the suture, lacking the

small upper spiral on the final whorls.

I. harryleei, I. compsa, I. pseudotorticula

spec. nov. and I. intermedia have three

spiral cords.

I. colon has a more cylindrical shell,

with spiral cords less prominent, the

nodules very small, and the small cords

at both sides of the suture almost imper-

ceptible.

I. undebermuda has three main

spirals, spiral 2 being small and cióse to

spiral 1; only one small thread near the

suture.

I. pompona has a much larger space

between spiral 2 and spiral 3, the

nodules are smaller and the protoconch

is wider and more prominent at the

apex, and has two spiral cords.

Inella meteora (Dalí, 1927) (Figs. 18F-H)

"Triphora" meteora Dall, 1927. Proc. U. S. N. M., 70: 95. [Type locality: Off Fernandina, Florida].

Type material: Lectotype (labeled as "type", in three fragments), here designated (Figs. 18F, 18G)

(USNM 108081) and 3 paralectotypes.

Description : Dall (1927). The proto-

conch (Fig. 18G) is pupoid with about 2

whorls and a diameter of about 600 pm;

below the nucleus there are two main

smooth spiral cords and another smaller

subsutural one.

The whorls of the teleoconch have

three spiral cords from the beginning

(Fig. 18H), the lower two more similar

in prominence, the nodules slightly

larger on spiral 2; the uppermost is

smaller from the beginning of the teleo-

conch and also is less prominent on the

final whorls. The nodules are uniformly

rounded on spiral 1 and larger and cut

at the middle on the other two. Above

122

RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 18. A-E. Inella enopla (Dalí, 1927). A, B: two fragments labeled as “type”, 7.2, 6.6 mm,

538 m (USNM); C, D: protoconch of a paralectotype; E: detail of the sculpture of a paralectotype.

F-H. Inella meteora (Dalí, 1927). F, G: two fragments labeled as “type” 4.0 and 5.1 mm, off Fer-

nandina, 538 m (USNM); H: detail of the sculpture.

Figura 18. A-E. Inella enopla (Dalí, 1927). A, B: dos fragmentos etiquetados como “tipo”, 7,2, 6,6

mm, 538 m (USNM); C, D: protoconcha de un paralectotipo; E: detalle de la escultura de un paralec-

totipo. F-H. Inella meteora (Dalí, 1927). F, G: dos fragmentos etiquetados como “tipo” 4,0 y 5,1 mm,

fuera de Fernandina, 538 m (USNM); H: detalle de la escultura.

123

Iberus , 26 (1), 2008

the suture there is a small undulating

thread.

Dimensions : See below.

Distribution : Only know from its

type locality.

Remarks : The original description gives

dimensions of 9 x 2 mm; the fragments in

the tube labeled "type" measure 6.2, 5.1

and 4.0 mm; they are of the same species

but probably not from the same shell.

The 3 paralectotypes in the other

tube appear to be a different species.

The differences with the closest

species are:

1. bigemma has two spirals at the

beginning of the teleoconch.

I. pompona and I. inflata have two

spirals at the beginning of the teleo-

conch and a wider, pupoid protoconch.

I. longissima has two spirals at the

beginning, subsequently three regular

spirals of similar size, with spherical

nodules, and the protoconch has three

whorls.

I. pseudolongissima spec. nov. has a

protoconch with 4 whorls, and spirals 2

and 3 are more evident at the beginning

of the teleoconch.

I. harryleei has a more depressed

apex on the protoconch; on the teleo-

conch the lower spiral is larger, with

spiral 2 smaller.

I. ibex and I. enopla have only two

spirals on the protoconch.

I. torticula can have up to 5 spirals at

the end.

I. colon is more cylindrical and has

only two main spirals on the teleoconch

with small nodules.

I. undebermuda spec. nov. has two

main spirals at the beginning of the

teleoconch and one smaller spiral 2 in-

between, located closer to spiral 1.

I. compsa is more conical, having

more rapidly enlarging whorls; the

nodules are less prominent and spheri-

cal.

I. pseudotorticula spec. nov. has a pro-

toconch with a smaller diameter and

with 2 well defined cords; only spirals 2

and 3 are present at the beginning of the

teleoconch, spiral 1 appearing later.

Inella pompona (Dalí, 1927) (Figs. 19A-C)

Triphora (Strobiligera) pompona Dalí, 1927. Proc. U. S. N. M., 70: 94. [Type locality: Off Georgia].

Type material: In the material under this ñame from USNM (Figs. 19A-19C) there is a label men-

tioning "type", but in the three glass tubes there is no other indication. The shell measuring almost

20 mm with 21 whorls is here designated as lectotype (Fig. 19A); it is referred to in the original

description as the "larger but imperfect specimen" because the apex is missing. Numerous para-

lectotypes (USNM 108339), most of them fragments (6 in one tube and 48 in another); the largest

fragment measures 23.5 mm.

Description : Dall (1927). The proto-

conch (Fig. 19B) is pupoid and wide, the

nucleus and the first whorl wider than

the subsequent ones, with the diameter

of the first whorl between 650 and 750

pm; in the original description it is

described as having 1 1/i whorls, but it

can have more than 2 whorls, the

second one with two spiral cords. Teleo-

conch with two main spirals (spirals 2

and 3); spirals 1 and 4 are very cióse to

the suture, which separates them (Fig.

19C). Spiral 1 has very small nodules,

while spiral 4 is smooth. The nodules on

the main spirals are not very prominent

and only a little higher than the cords,

and they are slightly cut at the middle.

Dimensions : The lectotype measures

almost 20 mm, and some shells are

somewhat larger.

Distribution : Only know from the

type locality.

Remarks : The species with a pupoid

and wide protoconch are:

I. enopla has a slightly narrower pro-

toconch of three whorls, with three

scarcely visible spiral cords.

I. ibex and I. colon each have informa-

tion on the protoconch in the original

description, but shells with a complete

124

ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 19. A-C. Inella pompona (Dalí, 1927). A: lectotype, 19.5 mm, 804 m (USNM); B: proto-

conch of a paralectotype; C: detail of the sculpture of lectotype. D-I. Inella dinea (Dalí, 1927). D:

lectotype, 7.9 mm, 804 m (USNM); E-G: paralectotypes, 6.6, 6.0. 4.7 mm, same locality; H, I:

protoconchs, lectotype and paralectotype.

Figura 19. A-C. Inella pompona (Dalí, 1927). A: lectotipo, 19,5 mm, 804 m (USNM); B: protocon-

cha de un paralectotipo; C: detalle de la escultura del lectotipo. D-I. Inella dinea (Dalí, 1927). D: lec-

totipo, 7,9 mm, 804 m (USNM); E-G: paralectotipos, 6,6, 6,0, 4,7 mm, la misma localidad; H, I:

protoconchas, lectotipo y paralectotipo.

125

Iberus , 26 (1), 2008

apex are not known; on the teleoconch

there are similarly two spiral beaded

cords, but on 1. colon the sutural area is

narro wer and the nodules are smaller; I.

ibex has smooth spirals at the beginning

of the teleoconch.

I. inflata has a similar protoconch,

but clearly narrower; there is a greater

increase in the width of the whorls and

the suture is more prominent.

I. undebermuda spec. nov. has a nar-

rower protoconch; in the teleoconch.

between the main spirals 1 and 3, there

is a small spiral 2 closer to spiral 1.

1. dinea (see below).

The species with unknown proto-

conch:

I. compsa has rapidly enlarging

whorls and three almost similar spirals.

I. sarissa has spiral 1 larger, with

spiral 2 very small and cióse to spiral 1.

I. bigemma has small nodules on

spiral 1 and fewer whorls, never elon-

gated.

Inella dinea (Dalí, 1927) (Figs. 19D-I)

" Triphora " pompona dinea Dalí, 1927. Proc. U. S. N. M., 70: 94. [Type locality: Off Georgia, 440 fms

(805 m)].

Type material: Lectotype (Fig. 19D), here designated, and 11 paralectotypes (Figs. 19E-19G)

(USNM 333517).

Description : Dall (1927), comparing

shells to typical I. pompona, only men-

tioned that the beading is less promi-

nent and the shells are more slender and

compact, measuring 10 mm for 15

whorls, with a máximum diameter of

1.75 mm. Now it can be added that the

protoconch (Figs. 19H, 191) has between

2 !A - 2 1 ¡2 smooth whorls and is a little

variable in size, but it can reach up to

700 pm and is pupoid at the apex, with

a large nucleus and with two or three

depressed smooth cords on the second

whorl. On the teleoconch, the suture has

a smooth fine cord on each side. The

axial sculpture is formed by small ribs

on some shells; on others the ribs are

almost absent; there are small orthocline

growth lines. There are three spirals,

equal in size on the lectotype but with

spiral 1 less prominent on other shells.

Dimensions: The lectotype measures

7.9 mm.

Distribution : Rosenberg (2005) gives

coordinates of 30.73° N, 79.43° W, with a

depth of 805 m.

Remarks: In the material studied

there was no shell measuring 10 mm,

but the designated lectotype has 14

whorls including the protoconch, and is

almost 8 mm. This shell was chosen

because the number of whorls is closest

to the number given in the description.

Also because it has three similar cords

on each whorl, very different from T.

pompona. Not all the paralectotypes

seem to be the same species.

Although the present taxon was pub-

lished as a variety of T. pompona, compar-

ison of the types of both species allowed

us to confirm that they are different

species. T. pompona has two nodulous

cords per whorl and a very small cord,

lacking nodules, on either side of the

suture. T. dinea has a similar pupoid pro-

toconch but the shell is variable, with

some shells having three cords of similar

size; other shells have only two spiral

nodulous cords and the third, near the

suture, is depressed. These could repre-

sent characters of two different species

but, as the studied material did not

inelude enough complete shells, it is better

to keep these two morphs within T. dinea.

I. sentoma has a fíat profile, lacks

nodules and is also a larger shell.

I. gaesona has depressed spirals

almost without nodules and the proto-

conch is narrower, with a depressed

apex.

I. inflata has a narrower protoconch,

with 2 poorly defined cords. The teleo-

conch has more prominent spirals and

nodules.

126

Rolan AND FernÁNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 20. A, B. Inella sentoma (Dalí, 1927). A: lectotype, 28 mm, off Fernandina, 538 m

(USNM): B: paralectotype, 4.4 mm, same locality (USNM). C. Inella gaesona (Dalí, 1927), lecto-

type, 9 mm, off Georgia, 805 m (USNM). D-H. Inella differens spec. nov. D: holotype, 7.6 mm,

off Egmont Key, Hillsborough Co., Florida (FLMNH); E: paratype, 6.2 mm; F, G: protoconchs of

holotype and paratype; H; detail of the sculpture.

Figura 20. A, B. Inella sentoma (Dalí, 1927). A: lectotipo, 28 mm, fuera de Fernandina, 538 m

(USNM): B: paralectotipo, 4,4 mm, la misma localidad (USNM). C. Inella gaesona (Dalí, 1927),

lectotipo, 9 mm, fuera de Georgia, 805 m (USNM). D-H. Inella differens spec. nov. D: holotipo , 7,6

mm, fuera de Egmont Key, Hillsborough Co., Florida (FLMNH); E: paratipo, 6,2 mm; F, G: protocon-

chas del holotipo y paratipo; H; detalle de la escultura.

127

Negro

Iberus , 26 (1), 2008

Inella sentoma (Dalí, 1927) (Figs. 20A-B)

Triphora sentoma Dalí, 1927. Proc. U. S. N. M., 70: 96. [Type locality: Off Fernandina, Florida, 805 m].

Type material: Lectotype, here designated (Fig. 20A), 28 mm and a paralectotype (Fig. 20B), 4.4

mm (USNM 108072).

Description : See Dall (1927). The pro-

toconch is unknown. Both the lectotype

and paralectotype are in very poor con-

dition, so the most important characters

from the original description are men-

tioned here: two low flattened spiral cords,

faintly undulated, one at the periphery of

the whorl, the other in front of it; the suture

linear with a minute threadlet on each side

of it, the interspaces shallow.

In reality it can be said that there are

grooves and orthocline growth lines, but

no prominent sculpture.

Dimensions: The lectotype measures

28 mm.

Distribution : Only know from its

type locality.

Remarks : This species is very elongate

and with depressed sculpture. There is no

Information about the protoconch or aper-

ture, but in the original description it is

mentioned that one of the fragments had

a protoconch of a whorl and a half. For

this reason and the almost cylindrical shell

form it is included in the genus Inella.

The paralectotype (Fig. 20B) is more

similar to the lectotype of I. gaesona (see

below).

No other Inella species in the studied

area has such depressed sculpture.

Inella gaesona (Dall, 1927) (Figs. 20C)

Triphora (Strobiligera) gaesona Dall, 1927. Proc. U. S. N. M., 70: 95. [Type locality: Off Georgia, 805 m].

Type material: Lectotype, here designated (Fig. 20C) with 16 whorls, 9 mm (USNM 1088341). Para-

lectotypes: 15 s and f (not examined) (USNM).

Description: See Dall (1927). Proto-

conch paucispiral, pupoid and de-

pressed, with the first whorl wider than

the following one. The spiral sculpture

is formed more by grooves which sepá-

rate the whorls into three parts than by

prominent spirals. The axial sculpture

consists only of orthocline growth lines.

Dimensions: The lectotype measures 9 mm.

Distribution: Only know from its

type locality.

Remarks: Species very different from

any other due to the pupoid and

depressed protoconch and the fíat spiral

cords without nodules. It only has some

similarity to I. sentoma, but a complete

comparison could not be made because

of the poor condition of the lectotype of

this species and the lack of sufficient

material. For this reason they are kept as

different valid taxa, awaiting more

material for study.

Inella differens spec. nov. Rolán and Lee (Figs. 20D-H)

Type material: Holotype (Fig. 20D) in FLMNH. Paratype (Fig. 20E) in BMSM (both ex CHL).

Other material examined: Off Louisiana: 3 f without protoconch, 28° 05.61' N, 91° 02.245' W by

28° 05.524' N, 91° 02.036' W, 58 m (E. F. Garda/ Pelican 06/09/00) (CHL).

Type locality: Off Egmont Key, Hillsborough Co., Florida, USA, (J. Moor e/Caualier, 1962), 135 m.

Etymology: The specific ñame alludes to the shell having different characters from others in the area.

Description: Shell subcylindrical toconch (Figs. 20F, 20G) pupoid with 1

(Figs. 20D, 20E) solid, light brown. Pro- 3A -2 whorls and a diameter of about

128

ROLÁN and FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae

Figure 21. A-D. Inella slapcinskyi spec. nov. A, B: holotype, 2.6 mm (FLMNH); C: protoconch of

the holotype; D: detail of the fragment (ANSP). E-I. 1: Inella faberi spec. nov. E: holotype, 3.7

mm; F, G: paratypes, 4.4 and 3.0 mm. Louisiana (FLMNH); H, I: protoconch and detail of teleo-

conch sculpture, paratype in figure F.

Figura 21. A-D. Inella slapcinskyi spec. nov. A, B: holotipo , 2,6 mm (FLMNH); C: protoconcha del

holotipo ; D: detalle del fragmento (ANSP). E-I. 1: Inella faberi spec. nov. E: holotipo, 3.7 mm; F, G:

paratipos, 4.4 y 3.0 mm. Louisiana (FLMNH); H, I: protoconcha y detalle de la escultura de la telo-

concha, paratipo de la figura F.

129

Iberus, 26 (1), 2008

600 pm, the nucleus smooth and the

whorls with two smooth spiral cords.

The teleoconch begins when the two

spiral cords become nodulous; these

beaded spirals are equal in size and

both are closer to the suture and more

widely separated in the middle of the

whorl. About the fourth or fifth whorl,

spiral 2 appears below and very cióse to

spiral 1, beginning as a fine thread

undulating beneath the nodules (Fig.

20H). Immediately it becomes nodulous

but with a smaller nodular size. About

the eleventh whorl the nodules reach

almost the size of those on spirals 1 and

3. There are three additional smooth

cords on the base. Aperture ovoid elon-

gate; siphonal canal short and open.

Dimensions : The holotype measures

7.6 mm.

Distribution: Only know from its

type locality.

Remarks : This species is kept in the

genus Inella due to the characteristic

protoconch.

It may be differentiated from most of

the other closely similar species by its

short protoconch with only 1 3A - 2

whorls and its brown colour:

I. dinea has more weakly sculptured

whorls and the protoconch has a wider

pupoid form.

1. meteora, I. undebermuda spec. nov., I.

enopla and I. pompona have a similar

number of protoconch whorls but the apex

is more inflated and pupoid, and the shells

are larger and white; furthermore, I. meteora

has three spiral cords on the teleoconch,

spiral 1 being smaller; I. undebermuda has

spiral 2 beginning on about the tenth whorl

(instead of on the fourth in I. differens), and

it is very small and closer to spiral 1; I.

enopla and I. pompona have two main spiral

cords on the teleoconch with an additional

cord on each side of the suture, the proto-

conch being very much wider.

I. triserialis has a rather similar shell

but the protoconch has 3 whorls, the

apex is more depressed, the nodules of

spiral 1 are larger and cut at the middle.

I. sarissa is white, spiral 2 is smaller

and very cióse to spiral 1.

I. compsa has rapidly enlarging

whorls, and spiral 2 is not smaller.

I. intermedia has a larger shell that is

lighter, whitish and more conical, the

spirals almost equal in size.

See also I.faberi below.

Inella slapcinskyi spec. nov. Rolán and Fernández-Garcés (Figs. 21A-D)

Type material: Holotype (juvenile, Figs. 21A, 21B) (FLMNH UF350382).

Other material studied: 1 f (ANSP).

Type locality: Cayman Islands, Little Cayman, Jackson's Bight (P. Watson, Dec/1992), 35 m.

Etymology: The species is named after John Slapcinsky, Curator of Molluscs of the FLMNH, for

his help in the examination of the material housed in this museum.

Description: Shell (Figs. 21A, 21B)

small, almost cylindrical, whitish. Proto-

conch paucispiral (Fig. 21C) with a

diameter of about 400 pm and almost

three whorls, with two narrow spiral

cords on the first whorl, three on the fol-

lowing, well defined, the lower one

more prominent; each spiral cord bears

a row of small tubercles on its upper

surface. The nucleus of the protoconch

is very small. The teleoconch has four

spirals; spiral 1 has more prominent

nodules; spiral 4 is the next in size and

spirals 2 and 3 have smaller, elongated

nodules (Fig. 21D). The axial ribs are

wide and strongly prosocline. On the