e

Bull. Mus. natl. Hist, nat., Paris, 4 sér., 14, 1992,

section A , n° 2 : 293-381.

Taxonomy of Tropical West African Bivalves

IV. Arcidae
by P. G r a h a m OLIVER and R u d o VON COSEL

Abstract. — Twenty species of Arcidae are described from tropical West Africa, defined here as
between 23° N and 17° S. The Arcinae are represented by four genera and include four new taxa : Area
avellana turbatrix n. subsp., Barbatia gabonensis n. sp., B. ionthados n. sp., B. (Nipponarca) allocostata n.
sp. The anatomy of the latter is described and confirms the value of the subgenus which is redefined. The
Anadarinae, other than Senilia and Bathyarca are all included in the genus Anadara because the use of the
ligament orientation and inequivalve condition are abandoned as phyletic characters. Two new species are
described : Anadara eborensis n. sp. and A. camerunensis n. sp. A. polii (Mayer) is the species often cited as
"A. diluvii (Lamarck)" in publications concerning the Mediterranean. Variation within species is
frequent, notably in Anadara polii and A. senegalensis. Ecological factors and geographical clines are
invoked to explain some of this variation but local genetic isolation could not be excluded. The
relationships of the shallow water West African species are analysed and compared to the faunas of the
Mediterranean, Caribbean, Panamic and Indo-Pacific regions. Only one genus is endemic to West Africa,
namely Senilia. There are three species common to the Mediterranean but none to the Caribbean
although there are sibling species. Overall similarity is greatest with the Indo-Pacific but the Anadarinae
in isolation have affinity to the Panamic fauna. A general discussion on the zoogeographical patterns for
all the Arcoidea is presented in the concluding part of this study which is to be published in the next issue
of the Bulletin du Muséum.
Résumé. — Description de vingt espèces d'Afrique occidentale tropicale, ici définie entre 23° N et
17°S. Les Arcinae sont représentés par quatre genres; quatre taxa nouveaux y sont inclus : Area avellana
turbatrix n. subsp., Barbatia gabonensis n. sp., B. ionthados n. sp., B. (Nipponarca) allocostata n. sp.
L'anatomie de la dernière est décrite, confirmant la valeur du sous-genre ici redéfini. Outre Senilia et
Bathyarca, toutes les Anadarinae sont incluses dans le genre Anadara, car l'utilisation de l'orientation du
ligament et l'inégalité des deux valves sont abandonnées en tant que caractères phylétiques. Deux espèces
nouvelles, Anadara eborensis n. sp. et A. camerunensis n. sp. sont décrites. Dans des publications
concernant la Méditerranée, A. polii (Mayer) a été souvent citée comme « A. diluvii (Lamarck) ». Une
variabilité intraspécifique est fréquente, surtout chez A. polii et A. senegalensis. Des facteurs écologiques
ainsi que des clines géographiques sont évoqués pour l'explication d'une partie de cette variabilité, mais
on ne peut pas exclure d'une isolation génétique locale. Les espèces du plateau continental ouest-africain
sont analysées sur le plan de leurs relations, et comparées avec les faunes de la Méditerranée, des
Caraïbes, du Pacifique panaméen et de l'Indo-Pacifique. Senilia est l'unique genre d'Arcidae endémique
de l'Afrique occidentale. Trois espèces vivent aussi en Méditerranée, mais aucune n'est présente dans les
Caraïbes bien qu'il y ait des espèces jumelles. Les Arcidae dans leur ensemble présentent le maximum de
similitude avec celles de l'Indo-Pacifique ; cependant les Anadarinae montrent une affinité plus marquée
avec celles du Pacifique panaméen. Une discussion générale sur la répartition zoogéographique de tous les
Arcoidea se trouve dans une deuxième partie de cette étude, qui sera publiée dans le prochain numéro du
Bulletin du Muséum.

P. G . OLIVER, National Museum of Wales, Cardiff, U.K.

R. VON COSEL, Muséum national d'Histoire naturelle. Laboratoire de Biologie des Invertébrés marins et Malaco-
logie. 55, rue Buffon, 75005 Paris.
 - 294 —

INTRODUCTION

This paper on the Arcidae and the forthcoming second part on the Noetiidae (which also
will contain the concluding discussion) are parts of a revision of the Bivalvia of tropical West
Africa being undertaken by the second a u t h o r and m a d e possible by the availability of recent
collections through O R S T O M (with main part of the fieldwork by the second author) and
Serge GOFAS ( M N H N ) . The region is defined here as lying between Rio de O r o (southern part
of West Sahara, 23° N ) and Baia dos Tigres (Southern Angola, 17° S). Previous studies of the
region are sparse with only the work of DAUTZENBERG (1909, 1913, 1927) and NICKLES (1950,
1955) of note. These papers illustrate the concept of shared taxa with the Caribbean and
Mediterranean regions but the recognition of pan Atlantic species and local species appears to
be inconsistent. F o r example the Barbatia candidalcomplanata complex was usually considered
to be a single pan Atlantic species but the equally confusing Arcopsis afra\adamsi complex (cf.
second part, OLIVER and COSEL, in print) has always been regarded as two distinct species.
Interestingly A. afra has also been widely cited from the Indo-Pacific. The actual relationships
of such apparent circumtropical species complexes have never been elucidated. The aim of this
review is therefore twofold : (1) T o describe and illustrate all species of Arcoidea from tropical
West Africa and (2) To examine the relationships of the west African species to those from the
Caribbean, Mediterranean and Indo-Pacific.
In both parts, data on the morphometries of the shells are frequently presented as box
plots. The box represents the range of values of the first thirty percent of the sample either side
of the mode, the projecting lines the next ten percent and the outlying points the remainder of
the values. We have used these as we feel they give a rapid visual understanding of the range of
values and the consequent variation. T h e box plots and all other descriptive and comparative
statistics are derived from analyses using Statview SE™.

The general discussion on zoogeographical patterns is presented in the second and concluding part of this
paper which will be published in the following issue of this Bulletin.

Abbreviations used in the text

BMNH : British Museum (Natural History) (now : The Natural History Museum), London, U.K.
MNHN : Museum national d'Histoire naturelle, Paris, France.
NMWZ : National Museum of Wales, Dept. of Zoology, Cardiff, U.K.
SMF : Natur-Museum und Forschungsinstitut Senckenberg, Frankfurt/M., Germany.
USNM : United States National Museum, Smithsonian Institution, Washington, D.C., USA.
ZMB : Zoologisches Museum der Humboldt-Universität Berlin, Berlin, Germany.
ZMC : Universitets Zoologisk Museum, Copenhagen, Denmark.
leg. : legit, collected by; sh. : shell, shells; spm. : specimen, specimens; sta. : sampling station;
v. : valve; RV : right valve; LV : left valve.
 — 295 -

Family ARCIDAE

Subfamily ARCINAE

Genus ARCA Linné, 1758

TYPE SPECIES : Area noae Linné, 1758 (SD Schmidt, 1818, ICZN Opinion 189).

Area noae Linné, 1758

(Pl. I, 1A-1D, 2 ; fig. 1; m a p 1)

Area noae Linné, 1758 : 693.

Area despecta P. Fischer, 1876 : 258-259; pi. 8, fig. 1.

TYPE MATERIAL : One marked specimen in the Linnean Collection in London (DODGE, 1952 : 143).
The holotype of Area despecta Fischer is in MNHN.

TYPE LOCALITY : " M . rubro, Mediterraneo, Indico" but now taken to be the Mediterranean. For A.
despecta Fischer " littora Africae occidentalis " but the precise locality is not recorded.

DESCRIPTION

Shell to 90 m m in length. Equivalve. Inflated. Inequilateral, beaks in the anterior quarter.

Outline narrowly elongate, subtrapezoidal. Posterior area long a n d narrow usually
slightly sulcate; posterior angle initially carinate, becoming rounded but remaining distinct;
posterior margin sloping outwards, usually auriculate, m a y be subtruncate. Median sinus
distinct in early stages but becoming o b s c u r e ; ventral margin straight or sinuous usually
indented at byssal gape. Anterior margin sloping inwards, more or less straight or gently
curved. Dorsal margin very long, straight.
Dorsal area very long and m a y be extremely wide, more or less flat; u m b o n a l separation
great, u m b o s low. Ligament initially restricted to a triangular area between the beaks but
developing to fill most of the dorsal area. Chevrons initially symmetrical but becoming
irregular with numerous insertion sites.
Hinge plate very n a r r o w ; teeth in two series, separation obscure but just behind the line
of the beaks. Anterior set u p to 40 teeth, posterior set u p to 60 teeth. Teeth minutely serrated,
more or less straight a n d vertical.
Sculpture primarily radial of narrow ribs, riblets a n d raised threads. In juveniles there are
30-35 more or less uniform riblets. In adults posterior area with 2-3 subobsolete ribs; posterior
angle becoming s m o o t h ; postero-median zone with 5-8 n a r r o w ribs which are n o t much
stronger than the 10-15 median riblets; anterior area with 4-6 riblets; all median and anterior
zones with interspace raised threads.
Periostracum usually worn remaining a r o u n d byssal gape a n d margins, composed of
 — 296 —

fragile semidiaphanous straw coloured concentric lamellae with very weak radial thickenings;
carinal bristles b r o a d , irregular, laminar, rarely persistent.
External background colour cream to white overlain with closely spaced buff to rust
brown oblique somewhat zigzag bands.
A d d u c t o r scars unequal, the posterior a b o u t twice the size of the anterior. Byssus
retractor elongate, triangular, extending along three quarters of the posterior hinge plate.

SELECTED SHELL MEASUREMENTS : For ratios of length to height, length to tumidity and length to
anterior to beak length, see figure 3 under Area bouvieri.

DISTRIBUTION : Eastern Atlantic from southern Portugal (Lagos, Algarve (rare)) south to Senegal
M'Bour, 14°N); Canary Islands, Cape Verde Islands; throughout the Mediterranean.

MATERIAL EXAMINED : Mediterranean Morocco : M'diq, on beach, 2 v., leg. GOFAS : Ceuta, Ensenada
de la Almadraba, 20-26 m, 1 juv. v., leg. BOUCHET; south of Ceuta, 35°53'N/05°19'W, 50 m, 1 juv. v., leg.
VON COSEL, both V.1986; Melilla, 1 sh., 2 v., coll. Staadt, all MNHN. Algeria : Beni Saf, 1 spm.; Oran,
6 spm., both coll. LOCARD, 1892; 3 sh., 1 v., coll. PALLARY, 1904; 3 v., old coll.; Alger, 1 sh., 2 v., old
coll.; all MNHN. Tunisia : Sfax, 2 v., coll. LOCARD, 1892; NW of Bou Grara Sea, Gulf of Gabes, 1 spm.,
some juv. v.; Borj Djillidj, Djerba Island, 10 spm., Djerba, north coast, on beach, many v.; Canal d'Ajim,
Djerba, 10-32 m, many spm., all leg. BOUCHET & WAREN, 1982; all MNHN. Greece : Euboea Channel,
1 spm., coll. CHAPER, MNHN. Italy : Venice, 1 sh., coll. MONTEROSATO, 1906; Palerme, 1 sh., coll.
ALLERY, 1872; Naples, 3 spm., coll. LOCARD, 1892; 2 sh., coll. PETIT, 1873; all MNHN. Mediterranean
France : Cannes, Alpes-Maritimes, 1 spm., 2 sh., old coll. M N H N ; St. Tropez, Var, 3 sh.; Porquerolles,
Var, 1 sh.; Sanary-sur-Mer, Var, 1 sh.; Marseille, 3 sh.; St. Henri, Bouches-du-Rhone. 1 sh., all coll.
LOCARD, 1892, MNHN. Corsica : St. Florent, 1 spm.; Bastia, 1 spm., both coll. LOCARD, 1892;
Propriano, on beach, 1 v., leg. DELAUNAY; Ajaccio, 2 spm., coll. LOCARD, 1882; all MNHN. Balearic
Islands : Mahon, Menorca, 5 spm., MELVILL-TOMLIN colln., NMWZ. Palma de Mallorca, 12 v., coll.
SOYER, 1969, MNHN. Mediterranean Spain : Cadaques, Costa Brava, 3 sh., 1 v., leg. CHERRIERE,
29.VIII.1966; 5 spm., both old coll. MNHN. Atlantic Morocco : Tanger, 1 spm., 1 sh., coll. PALLARY,
MNHN. Canary Islands : Las Palmas, Playa Alcaravaneras, 1 sh., 3 v., in the nets of fishermen; Playa
Las Canteras, 5 v., both leg. VON COSEL, 1965, MNHN ex coll. VON COSEL. West Sahara : off Cap Barbas,
21°05'N/17°14'W, 43-45 m, 2 spm., 1 v., " Calypso "-Sta. 1, leg. MARCHE-MARCHAD, MNHN. Maurita­
nia : Port-Etienne (now Nouadhibou), 2 spm., 2 sh., Mission GRUVEL ; Pointe des Maures (20°55' N),
intertidal, 1 spm., leg. BOUCHET, V.1983; Pointe Cansado, 2 v., Mission GRUVEL, 5.IV.1908; all MNHN.
Cape Verde Islands : (without precise locality) 1 v., coll. DE CESSAC, 1877; Santo Antao, Punta do Sal,
5 v., leg. CADENAT; Sao Vicente, Baia Matiota, Mindelo, 3m, 3 juv. v., leg. VON COSEL, 16.XII.1978;
Razo, on beach, 1 v., leg. SCHLEICH, VIII.1981; Sao Nicolau, Tarrafal, 1 v., leg. GROH, XII.1978; Ilha do
Sal, Santa Maria, on beach, several juv. v.; Boavista, Sal Rei, 1 v.; Boavista, 1,4 miles W of Pta. Areia,
22 m, 4 v.; Boavista, off Praia Corralhino, 10 m, 1 v.; Sao Tiago, northern part of Baia de Sta. Clara, 15-
35m, l s p m . ; Sao Tiago, 14°54'N/23°30,5'W, 15m, 1 v.; 15°16'N/23°47'W, 55-60m, 1 v.;
15°13,2'N/23°46,3'W, 1 juv. v.; Fogo, SW of Sta. da Encarnacao, 20-25m, 1 juv. spm., 10m, 1 juv. ch.
all dredged R/V "Calypso", leg. MARCHE-MARCHAD, XI. 1959; all MNHN. Senegal : Banc de Seminole,
43-45m, 1 juv. spm., 3 v., leg. MARCHE-MARCHAD; Dakar, Bel-Air, Plage de la Marine, 2 sh., 3 v., leg.
VON COSEL, 16.X.1988; Dakar-Hann, on beach, 1 sh., 3 v., R/V "President Theodore Tissier", littoral
station, 1936; Goree, lspm., leg. GUILBERT, 7.II.1958; SE of Goree, 14°41'N/17°23,3'W, 17-19m,
1 spm., 5 v., dredged R/V "Louis Sauger", leg. VON COSEL, 24.111.1988; Cap Rouge, several juv. sh.,
Mission GRUVEL, III-IV.1909 ; M'Bour, Petite Cote, on beach, 1 sh., 2 v., leg. VON COSEL, 22.111.1988; off
M'Bour, 14°26,5'N/17°05,5'W, 10-11 m, 4 spm., 6 v.; off Saloum estuary, 14°2,5'N/17°09'W, 24m, 2 v.,
both dredged R/V "Commandant Henri Gomis", leg. BODARD, 8.XII.1966; all MNHN. Guinea : Off
Dubreka Estuary, 9°42'N/15°33'W, 35m, 1 old valve which still shows colours; off Conakry,
9°30'N/15°09,6'W, 45 m, 1 old, apparently subfossil v.; off the border to Sierra Leone, 9°06'N/14°35'W,
52m, 1 old, app. subfossil v.; all dredged R/V "Andre Nizery", leg. VON COSEL, 29-30.X.1988, all
MNHN.
 — 297

BIOTOPE : Area noae lives mainly on hard grounds and is byssally attached on rocks, to the undersides
of rocks and in crevices. On soft bottoms it can be found attached to isolated stones or other hard objects
including the shells of larger gastropods. The bathymétrie range is from low water to about 50 m.

REMARKS : See under Area bouvieri.

1 2
FIG. 1-2. — 1 : Internal view of right valve of Area noae L. Western Sahara. 2 : Internal view of right valve of Area
bouvieri Fischer. Angola. (Scales bars = 10 m m . )

Area bouvieri P. Fischer, 1874

(Pl. I, 3A-D, 4 ; fig. 2 ; m a p 1)

Area bouvieri P. Fischer, 1874 : 206.

Area sanctahelenae Smith, 1890 : 305, pi. 22, figs. 8-8b.

TYPE MATERIAL : The holotype (coll. BOUVIER) has not been isolated in the MNHN although
ROCHEBRUNE (1881 : 250) cites it as " Sta. Vincent, Bouvier 1870, Mus. Paris. " Two paratypes (complete
specimens) from coll. PETIT DE LA SAUSSAYE are present and among them is the specimen figured by
FISCHER (1876 : 239-240, pi. 8, fig. 2).

TYPE LOCALITY : Cape Verde Islands.

DESCRIPTION

Shell to 70 m m in length. Solid. Equivalve. Inequilateral, beaks in the anterior third.

Outline subtrapezoidal, oblong, height relatively great. In large specimens : Posterior area
relatively large; posterior angle r o u n d e d ; posterior margin long, almost vertical, often straight
some weakly auriculate. Median area as a shallow sinus, ventral margin more or less straight
or with a shallow indentation at byssal gape. Anterior margin broadly rounded. Dorsal margin
long, straight. In small specimens : Posterior area n a r r o w ; posterior angle c a r i n a t e ; posterior
margin subacute, sloping inwardly, auriculate at junction with posterior carina. Median sinus
m a r k e d ; ventral margin indented. Anterior margin sloping inwardly, weakly indented. Dorsal
margin relatively very long, lateral junctions acute.
 — 298 -

Dorsal area long, very wide and almost flat rising gently to low u m b o s . Ligament initially
restricted to triangular area between beaks b u t eventually covering most of t h e dorsal area
except for a n a r r o w posterior strip a n d a very n a r r o w anterior strip. Chevrons developing
symmetrically at first b u t in gerontic examples becoming irregular a n d often very numerous.
Hinge plate very narrow. Teeth in t w o series, junction just behind the beaks a n d
indistinct. Anterior set u p to 40, posterior set u p to 50 in number. All teeth finely serrated,
postero-median teeth slightly chevron shaped, remainder more or less straight a n d vertical.
Sculpture primarily radial of ribs, riblets a n d raised threads. Juveniles which a b o u t 30-35
more or less uniform riblets except for those o n the posterior area which are much larger. In
large specimens : posterior and postero-median area with 8-10 rounded ribs with u p to 6 raised
threads in the interspaces. Median area with n u m e r o u s riblets a n d raised threads. Anterior
area with 4-6 ribs.
Periostracum mostly worn, retained a r o u n d margins only in adult examples. C o m p o s e d of
very thin, subdiaphanous, straw coloured concentric lamellae, very weakly reinforced by radial
thickenings a n d breaking down over ribs to give an a p p a r e n t radial arrangement. In juveniles
the carinal bristles are occasionally preserved, these are greatly expanded, irregularly
spathulate a n d extremely fragile.
The external background colour of the shell is cream and is variously overlain with b r o a d
oblique, sometimes zigzag bands from brick red to purplish red in colour. Larger shells
frequently have an orange tinge to the posterior area. Internal colour is mostly white except
for a brickred marginal zone.
The adductor muscle scars a r e subequal, the posterior a little larger. The byssus retractor
scar lies under the posterior part of the dorsal area, is a n elongate triangle extending along two
thirds of the posterior hinge plate.

SELECTED SHELL MEASUREMENTS : See in Remark section.

DISTRIBUTION : Senegal (Cape Verde Peninsula) to southern Angola (Mocamedes); Cape Verde
Islands, Sâo Tomé, Principe, Annobon ; Ascension, St. Helena.

MATERIAL EXAMINED : Senegal : Dakar, SW of Madeleines Islands, 45-46 m, 3 juv. spm., 11 juv. v.,
leg. MARCHE-MARCHAD, 9.1.1954; Dakar, on Gouvernment Beach, 2 v., R/V "Président Théodore
Tissier", littoral station, 1936; Dakar, SW of Cap Manuel, 50 m, 2 v., dredged R/V "Gérard Tréca", leg.
MARCHE-MARCHAD, 20.11.1956; S of Gorée, 110-112m, 1 juv. v., dredged R/V "Gérard Tréca", leg.
MARCHE-MARCHAD, 18.11.1954; Gorée, 15-25 m, 11 spm., leg. PIN, 1987; Baie de Rufisque, 18-20 m,
1 juv. spm., 5 juv. v., Mission GRUVEL, II-IV.1909; off M'Bour, 14°2,5'N/17°09'W, 24m, 6 spm., 1 v.,
dredged R/V "Commandant Henri Gomis", leg. BODARD, 7.XII.1966; M'Bour, on beach, 1 v., leg. VON
COSEL, 22.111.1988; S-Casamance, off Cap Skirring, 12°25'N/17°17'W, 25 m, fine sand with stones,
1 spm., dredged R/V " Louis Sauger", leg. VON COSEL, 27.III. 1988 ; all MNHN. Cape Verde Islands : (no
precise locality), 1 sh., coll. DE CESSAC, 1874; Sao Vicente, 1 sh., 1 v., coll. MABILLE, 1905; S of Santa
Luzia, 16°44'N/24°44,5'W, 52 m, coarse sand with shells and stones, 1 juv. spm., 2 v., dredged R/V
"Princess Alice I I " , 1901 ; Ilha do Sal, Pedra Lume, on beach, 1 juv. sh., coll. CADENAT; Sào Tiago,
Praia, 14°53,7'N/23°30,4'W, 25-30m, 4juv. v., dredged R/V "Calypso", 17.XI.1959; Brava, SW of
PuntaTantào, 20 m, 1 juv. v., dredged R/V "Calypso", 21.XI.1959 ; all MNHN. Guinea : off Rio Nunez,
10°27'N/15°37,5'W, 39 m, mixed sand with shells and gravel, 2 spm. ; off Tannah Island,
9°12'N/14°16'W, 41 m, coarse sand with stones, 4 spm., several v., both dredged R/V "André Nizery",
leg. VON COSEL, 28-30.X.1988, both MNHN. Côte d'Ivoire : off Abidjan, 5°01,5'N/3°23,5'W, 70m, sand
 — 299

MAP 1. — Distribution of Area noae (circles) and A. bouvieri (squares). The empty circle indicates a record of subfossil
valves only.

with shells, 1 juv. v.; off Grand Bassam, 4°57'N/2°42'W, 40m, 1 spm., 1 v., both dredged R/V " L a
Rafale", Guinean Trawling Survey, leg. CHERBONNIER, 19-22.III.1964, both MNHN. Ghana : Cape
Coast, 26-31 m, 1 juv. spm., leg. LE LOEUFF, 10.11.1968; 4*36,5'N/1*31'W, 50m, 1 v., dredged R/V
"Calypso", leg. MARCHE-MARCHAD, 24.V.1956, both MNHN. Nigeria : off Niger Delta, 4°00'N/6"11'E,
34m, 3 spm.; 4°03'N/6° 12'E, 32m, 10 juv. v., both dredged R/V "Calypso", 26.V.1956, leg. MARCHE-
MARCHAD, both MNHN. Ilha do Principe : Between Pta. da Mina and I. Santa Ana, 10-12m, 1 spm.,
1 sh.; 1°42,5' N/7°28' E, 21m, algues calcaires, 1 spm.; 1°43' N/7°29' E, 37 m, 1 spm.; l°43'N/7°28,4' E,
73m, 1 spm., 1 v.; r33'N/7°31'E, 90m, 1 juv. spm.; all dredged R/V "Calypso", leg. MARCHE-
MARCHAD, 24.VI-1.VII.1956, MNHN. Sao Tome : off Baia de Ana de Chaves, 5m, calcareous algae,
1 spm.; off Praia Lagarto, 5-6m, 1 spm.; off Diego Nufies, 0°23'N/6°43'E, 30m, 1 spm.; north coast,
 — 300

0°25,5' N/6°40' E, 50 in, 1 juv. spm., all dredged R/V "Calypso", leg. MARCHE-MARCHAD, 11-21.VI.1956;
all MNHN. Annobon : (without precise locality), 60 m, 3 juv. sh., coli. CAVELIER DE CUVERVILLE, 1886;
north of Santo Antonio, 23 m, calcareous algae, 2 spm., dredged R/V " Calypso ", leg. MARCHE-
MARCHAD, 4.VII.1956; all MNHN. Equatorial Guinea: Bata, 1 sh., coli. POBEGUIN, 1891, MNHN.
Gabon : Pont-Gentil, Ile aux Pigeons, 3 m, on mangrove roots and Pinna, 5 spm., leg. CHEVALIER, 1984-
89, MNHN. R. P. Congo : Pointe-Noire, plage Sauvage, 2 v., leg. VON COSEL, 10.XII.1985, MNHN.
Angola : Barra do Dande, Bengo province, on rocks at low tide, 1 spm. ; Cacuaco, Bengo province, on
rocks at low tide, 1 spm.; 5-10m, 9 juv. spm., 8 juv. v.; Ilha de Luanda, Luanda province, 40-60m,
1 spm.; Corimba, Luanda province, 10-20m, numerous spm.; Palmeirinhas, Luanda province, on rocks
at low tide, 1 spm.; Cabo Ledo, Luanda province, 3 juv. spm., 3 juv. v.; Baia de Canoco, Benguela
province, 2 juv. spm.; Praia Amelia, Moçâmedes, 40-60 m, 2 spm., all leg. GOFAS, 1981-86, MNHN.

BIOTOPE : Byssally attached on hard substrates such as rocks, stones, shell agglomerations and
secondary hard substrates like shells or small stones on sand, mostly with calcareous algae. Bathymétrie
range from sublittoral to 60 m.

REMARKS

Area bouvieri and A. noae have overlapping ranges only in the Senegal a n d C a p Verde
regions. They can be separated at all sizes by the differences in shell shape (fig. 3) and
sculpture. Area bouvieri is less elongate, m o r e compressed and less inequilateral than A. noae.
The radial ribs of A. bouvieri are m u c h stronger especially over the posterior area.

Area bouvien
2.8.

2.6. • Area noae

c
2.4 X-2.22
o X = 2.11^ o X = 2.11
2.2. o o
-X
2
T
181 0 T To
1.6. X - 1.76
X-1.87

1.2.
L/H UT L/A-Bk

FIG. 3. — Box plots of the ratios of T o t a l Shell Length to Height ( L / H ) , Tumidity (L/T) and Anterior to Beak Length
(L/A-Bk) of Area bouvieri (n = 20) and Area noae (n = 20).

Area bouvieri belongs to the A. noae group of species which is represented in the
Caribbean by A. zebra, in the Indo-Pacific by A. navicularis and in the Panamic Pacific by A.
pacifica. A. bouvieri differs from all of these in its relatively short q u a d r a t e outline and the
large difference between the median and lateral sculpture. In these respects it differs most from
A. noae and is closer to A. zebra and A. navicularis.
We can find no reason to regard A. sanctahelenae Smith as distinct from A. bouvieri. It
 301 -

would appear that SMITH (1890) overlooked FISCHER'S species as he compares the St. Helena
material only with A. noae and A. navicular is. LAMY (1907 : 24) also considered A.
sanctahelenae to be a synonym.

Area a veil an a turbatrix n. subsp.

(PI. II, 1A-D, 2 ; fig. 7 ; m a p 2)

TYPE MATERIAL : Holotype (28.3mm), MNHN, Santo Antonio, Benguela province, Angola;
intertidal, rocks; leg. GOFAS. Paratypes MNHN : 2 spm. as holotype; 2 spm. Lucira, Mocamedes
province, Angola; NMWZ : 8 spm. Corimba, Mocamedes province, Angola; all leg. GOFAS.

TYPE LOCALITY : Santo Antonio, Benguela province, Angola.

DESCRIPTION

Shell to 35 m m in length. Equivalve. Inflated. Inequilateral, beaks in the anterior third.

Outline subtrapezoidal, posterior area demarcated by a strong posterior angle which is
often carinate. If small and not distorted then : longer than h i g h ; posterior margin obliquely
truncate usually slightly auriculate, posterior ventral junction a c u t e ; ventral margin straight,
gently curving or slightly indented by byssal g a p e ; anterior margin broadly r o u n d e d ; dorsal
margin long straight, dorsal lateral junctions angular. Posterior area well demarcated by
carinate posterior ridge, median sinus not apparent. If large then usually deformed to some
degree and then : some remaining longer than high others almost as high as l o n g ; posterior
margin obliquely subtruncate, posterior ventral junction angular or narrowly r o u n d e d ; ventral
margin variously indented byssal gape which may be extensive; anterior margin rounded,
straight or irregular.
Dorsal area large, broadening rapidly with wide imbonal separation. Ligament extensive,
covering dorsal area, chevrons irregular, developing rapidly and numbering u p to 10 in
gerondic specimens.
Hinge plate narrow, teeth small in two series but separation obscure, anterior set u p to
38 teeth, posterior set u p to 43 teeth. All teeth minutely serrated and with a frontal g r o o v e ;
posterior marginal teeth becoming chevron shaped, others straight and vertical.
Sculpture on posterior area of larger shells of 5-8 ribs, the first three from the posterior
carina most prominent and usually at least one bifid. T o w a r d s dorsal margin ribs smaller,
more variable in size, some as interspace riblets; of smaller shells usually of 4-5 simple ribs.
Median area frequently divided by a weak posterior median ridge, posterior section with 15-20
nodulose riblets, anterior section and anterior area with 25-30 radial riblets variously with
interspace radial threads. In most specimens the radial element is d o m i n a n t but in more
elongate examples the concentric element is equally expressed especially on the posterior
median zone.
Periostracum rarely well preserved but usually present a r o u n d byssal gape and remnants
on posterior carina. Posterior carina with a single row of large, spathulate bristles which are
smooth on the anterior edge and coarsely serrated on the posterior edge, they are thin, fragile
a n d straw coloured (pi. II, 1A). On the posterior area short lamellar elements are present
 — 302 —

between the ribs and are reinforced by weak slightly emergent hairs. On the remainder it is
concentrically lamellar but breaks down into radially arranged segments which are weakly
reinforced emergent hairs.
Shell generally pale in colour usually with mauve-brown tinges along the posterior border,
occasionally extending t h r o u g h o u t the posterior third, over the hinge plate and onto the
anterior area, some darker brown on posterior area. M a n t l e pigmentation absent except for
the black eyespots.

SELECTED SHELL MEASUREMENTS : See figure 4.

DISTRIBUTION : From Guinea (Conakry area) south to Cameroon and Angola (Luanda).

MATERIAL EXAMINED : Senegal : S of Cap Skirring, S-Casamance, 2 worn v., on beach, leg. VON
COSEL, 3-6.III.1988, MNHN. Guinea : Roume, lies de Los, beach on the N-side, 2 v., leg. VON COSEL,
29.V.1988; Kassa, Iles de Los, E-Side, N of village, on the undersides of boulders and stones with very
thin layer of mud on them, low tide, 2 spm., leg. VON COSEL, 12.XI.1988, both MNHN. Liberia :
5°21,5'N/9°54,5'W, 73-80m, 1 v., dredged R/V "Calypso", leg. MARCHE-MARCHAD, 20.V.1956, MNHN.
Säo Tome : Praia Emilia, under stones at low tide, 1 spm., leg. GOFAS, XI. 1983, MNHN. Cameroon :
Victoria (now Limbe), Morton Bay, 2 spm., 1 v ; Mondoleh Island, Ambas Bay, 6 spm., all under stones
at low tide, leg. VON COSEL, IV.1969 and XI.1985, MNHN. Gabon : Cap Esterias-Pointe Idolo, on stones
in sand at low tide, 3 spm., 1 v., leg. VON COSEL, M N H N ; (no precise locality), 6 spm., coll. H. FISCHER,
MNHN. R.P. Congo : Pointe-Noire, Plage ORSTOM, 3-6 m, many v. ; Plage Mondaine, on beach,
many v.; Songolo, 3-5m, 3 v.; Plage Sauvage, on beach, 3 v., all leg. VON COSEL, XI-XII.1985; Pointe-
Noire, 6 spm., 6 v., coll. Office Pte.-Noire; Pointe-Noire, 5 spm., coll. AUBERT DE LA R U E ; all MNHN.
Angola : Ambrizete, Zaire province, 1 spm., 1 v. ; Barra do Dande, Bengo province, 6 spm. ; Cucuaco,
Bengo province, 1 spm.; Praia Etambar, Corimba, Luanda province, 10 spm.; Caotinha, Benguela
province, 4 spm., all leg. GOFAS, MNHN.

BIOTOPE : Byssally attached to stones and rocks, mostly on the undersides. This species appears to
prefer more or less turbid waters and is also found in areas with slightly reduced salinities during the rainy
season. Bathymétrie range from low intertidal to sublittoral.

DERIVATIO NOMINIS : turbatrix, Latin, f. the "disturber" or " trouble maker" making reference to the
systematic confusion prevalent in this group of species.

REMARKS

This is the tropical West African species referred to as Area imbricata Bruguiere of
authors. Its identity has never been examined in any detail and it is assumed that it was
identified as that species on geographical considerations alone. The distinctions between the
Caribbean form k n o w n as A. imbricata and its Indo-Pacific counterpart. A. avellana Lamarck
are similarly obscure. We have examined these in relation to the West African form and make
the following observations :
 — 303 —

CHARACTER INDO-PACIFIC WEST AFRICAN CARIBBEAN

Size To 5 0 mm. To 3 5 mm. To 6 5 mm.

Posterior sculpture Ribs often irregular, Ribs often irregular, Ribs mostly regular,
occasionally bifid. usually bifid. rarely bifid

Periostracal insertion marks Lacking Lacking Present

Ligament chevron № Medium High Low

Llmbonal separation Wide Very wide Moderate

Carinal bristles Large, fragile Large, fragile Very large, robust

straw coloured straw coloured greyish

Internal shell colour Usually partly white Mostly white Usually dark overall

Mantle colour Patterned shades of White except for eye Patterned shades of
brown spots brown

Area avellana (Indo-Pacific) (pi. II, 4) : This form has a larger m a x i m u m size but the
general form of the shell, sculpture and periostracum are so similar that we are unable to
identify significant differences. Only the u m b o n a l separation and relative n u m b e r of ligament
chevrons show a difference (figs. 5 & 6). The colouration pattern is similar but there does seem
to be a greater incidence of dark coloured specimens from the Indo-Pacific and this may also
be reflected in the greater pigmentation of the mantle.
Area imbricata (Caribbean) (pi. II, 3A-B) : The Caribbean form attains a much larger
m a x i m u m size and in general is much less distorted. The shells are frequently more elongate
and the ribs on the posterior area are rarely bifid but bear short grooves which represent the
insertion points of the periostracal bristles. U m b o n a l separation is relatively small and the
n u m b e r of ligament chevrons is relatively fewer. T h e internal colouration is frequently darker
with most specimens coloured t h r o u g h o u t and many being chocolate brown or black. Most
significantly however are the carinal periostracal bristles which are approximately twice the
size, more strongly reinforced and more persistent (pi. IX, B).
F r o m these comparisons it would appear that the West African form is more closely allied
to that of the Indo-Pacific than it is to that of the Caribbean. However we have not addressed
the possibility that these differences are merely ecophenotypic or of regional significance only.
We suggest that as the total range of variation is not seen in all three groups that there is at
least some regional significance. Even so this may reflect the absence of certain habitats in a
region and in West Africa the exclusion of coral reef communities may be important. The
habitats described here of the West African population are unusual in that the preference
seems to be for turbid intertidal areas which are in contrast to the coral reef association with
clear water in the tropical regions of the Caribbean and Indo-Pacific. Of the characters cited
we view the carinal bristles to be most relevant as it is difficult to envisage the functional
significance of the differences noted. The umbonal separation and ligament chevrons are
related to growth with both having an allometric relationships to shell size (THOMAS, 1976;
 5.5

Shell Length mm K Shell Length mm

FIG. 4-6. — 4 : Box plots of the ratios of T o t a l Shell Length to Height ( L / H ) , T u m i d i t y (L/T) a n d Anterior to Beak
Length (L/A-Bk) of Area avellana lurbatrix (n = 25). 5 : G r a p h s of U m b o n a l separation plotted against Shell
Length for four samples of the Area imbricata complex. F l o r i d a and G u a d e l o u p e are A. imbricata, Seychelles is A.
avellana and West Africa is A. a. turbatrix. 6 : G r a p h s of ligament chevron n u m b e r plotted against shell length for
four samples of the Area imbricata complex. Florida and G u a d e l o u p e are A. imbricata, Seychelles is A. avellana and
West Africa is A. a. turbatrix.

OLIVER and A L L E N , 1980). T h e differences observed in these could be related to growth rate
and if the West African population was restricted to less favourable conditions the allometry
would then occur in the smaller size classes. The only meaningful way forward would be to
carry out studies related to the genetic m a k e u p of each population but this is beyond the
scope of the material at hand.
Resorting to traditional techniques we conclude that the West African form is most
closely allied to that of the Indo-Pacific but the differences observed may be due to regional
environmental conditions. However b o t h populations are geographically isolated as the Indian
Ocean populations d o not spread beyond the province of Natal and the West African form is
restricted to the Gulf of G u i n e a n o r t h of Namibia. Consequently we have given subspecific
status to the West African form as A. avellana turbatrix but not in keeping with tradition
regard it as distinct from the Caribbean form A. imbricata. If one views the differences to be
insignificant then one c a n n o t regard the Caribbean and Indo-Pacific forms to be distinct either
 — 305 —

and neither would t h e distinction of the P a n a m i c Pacific form k n o w n as A. mutabilis be

justifiable. The whole complex would then have t o be regarded as a single cosmopolitan species
taking the earliest n a m e of A. imbricata Bruguiere. This is the position adopted by LAMY
(1907 : 26-38) b u t he accepted three varieties within the Indo-Pacific a d West African ranges,
namely var. arabica Philippi, var. avellana Lamarck, a n d var. martensi D u n k e r . H e also
isolated the Caribbean form as t h e nominate species A. imbricata.

FIG. 7-8. — 7 : Internal view of right valve of Area avellana turbatrix n. ssp. Angola. 8 : Internal view of right valve of
Area retragona Poli. Ireland. (Scales bars = 5 m m . )

Area tetragona Poli, 1795

(PI. I I , 5A-B, 6 ; fig. 8 ; m a p 2)

Area tetragona Poli, 1795 : 137.

TYPE MATERIAL : Not located.

TYPE LOCALITY : Sicily.

DESCRIPTION

Shell t o 40 m m in length. Equivalve. Inflated if growth is unhindered b u t m a y be

compressed if constricted in crevices. Inequilateral, beak position variable, in the anterior third
in regular specimens b u t only just in front of the mid line in some distorted examples.
Outline subtrapezoisal, angular, juveniles elongate, adults irregular oblong. Posterior area
narrow, well demarcated by t h e posterior angle which remains sharply c a r i n a t e ; posterior
margin oblique, more o r less straight, some very weakly auriculate. M e d i a n area n o t
depressed, ventral margin gently curved or sinuate at byssal gape. Anterior margin rounded,
variable. Dorsal margin long, straight.
Dorsal area wide t o very wide, usually flat but m a y be cleft in distorted examples; u m b o s
very low. Ligament with unusual growth p a t t e r n ; initially with two strips of fibrous ligament
radiating from the beaks, between these a n d t o either side of them t h e dorsal area is free of
ligament material. Lamellar chevrons a r e added mostly within the anterior strip then t o the
posterior strip and finally new sites of ligament growth appear along the hinge line between the
 — 306

original strips so that in the largest shells most of the dorsal area is covered. T h e chevrons are
very narrow and may reach 25 in number.
Hinge plate narrow, teeth in two series, separation obscure at a point behind the umbonal
line; anterior set u p to 30 teeth, posterior set u p to 15 teeth. Teeth minutely serrated anterior
teeth small, some chevron shaped, mostly vertical but marginal teeth oblique ; posterior teeth
straight but becoming progressively oblique towards the margin where they are subparallel.
Sculpture of radial riblets and raised threads, initially with 4-5 riblets on the posterior
area and 30-35 finer riblets on the remainder, carinal riblet bifid often imbricate. Subsequently
posterior riblets divide and are added to by interspace threads. Median area with closely
spaced, somewhat acute riblets contrasting slightly with the more rounded and slightly broader
riblets on the anterior area.
Periostracum usually lost, persistent a r o u n d margins and on the posterior area only.
Composed of concentric lamellae reinforced by bristles arising from the riblet interspaces,
most often the lamellar part is worm and the arrangement appears simply radial. The bristles
become spicate on the postero-median and posterior areas. The carinal bristles are large,
lanceolate but with irregular margins. All the bristles are dark coloured from brown to black.
Shell colouration is weak, mostly greyish white but with some brown tinges internally
over the posterior area.
A d d u c t o r scars small, subequal ; both with myophoric ridges extending into the u m b o n a l
cavity. Byssal retractor small, triangular restricted to the posterior half of the shortened
posterior hinge plate.

DISTRIBUTION : Shetlands to Morocco (Essaouira/Mogador), the Azores and throughout the

Mediterranean.

MATERIAL EXAMINED : Ireland : St. Johns PL, Donegal, 5 spm., leg. P. G. OLIVER, Berehaven, Bantry
Bay, 9 spm., leg. J. T. MARSHALL; both NMWZ. England : Falmouth, leg. J. T. MARSHALL, 8 spm.,
NMWZ. Celtic Sea : 8 stations between 49° 15' and 48°33'N and 05°09' and 05°28,5'W, 102-112 m, 15 v.,
all dredged R/V "Thalassa", leg. VON COSEL, XII. 1983, MNHN. Atlantic France : Brest, Finistère, 2 sh. ;
Lorient, Morbihan, 3 spm.; Belle-Ile, Morbihan, 1 sh. ; Le Croisic, Loire-Atlantique, 4 spm.; He d'Yeu,
Vendée, 2 spm.; Ile d'Oléron, Charente-Maritime, 3 spm., all. coll. LOCARD, 1982; Golfe de Gascogne
(Biscay), 18 spm., coll. DENIS; Arcachon, Gironde, 1 spm., coll. ROCHE, 1894; St-Jean-de-Luz, Basses-
Pyrénées, numerous sh. and v., coll. H. FISCHER, 1898; all MNHN. Atlantic Spain : Ceuta, Strait of
Gibraltar, Punta Almina, 25-40 m, 6 spm., 1 v., leg. BOUCHET, GOFAS & LOZOUET, V.1986, MNHN.
Algeria : Oran, 40m, 2 spm., 2 sh., coll. PALLARY; 60m, 3 spm., coll. LOCARD, 1892; Alger, 4 juv. sh.,
1 v., old coll., all MNHN. Italy : Palermo, Sicily, 11 spm., 3 sh., coll. JOUSSEAUME, 1916; 3 spm., 2 sh.,
coll. ALLERY, 1872; 7 spm., coll. MONTEROSATO, 1906; Naples, 5 spm., 4 sh., coll. PETIT, 1873; 4 spm.,
2 sh., old coll.; 1 v., coll. MONTEROSATO, 1906; Capri, 3 spm., coll. LOCARD, 1892; all MNHN.
Mediterranean France : Nice, Alpes-Maritimes, 1 spm. ; St. Raphaël, Var, 3 spm. ; St. Tropez, Var,
2 spm., all coll. LOCARD, 1892; Toulon, Var, 1 spm., old coll.; Marseille, Bouches-du-Rhône, 9 spm.,
coll. LOCARD, 1892; 1 spm., coll. VAYSSIÈRE, 1907; Sète, Hérault, 2 spm., coll. LOCARD, 1892; Banyuls-
sur-Mer, Pyrénées-Orientales, 8 spm., coll. Lab. Arago, 1904; 4 spm., old coll., all MNHN. Corsica : St.
Florent, N-coast, 2 spm., coll. LOCARD, 1892, MNHN. Atlantic Morocco : Mogador (now Essaouira),
6 spm., coll. PALLARY, MNHN. Azores : Ponta Delgada, Sào Miguel, 10-20 m, 2 spm., 5 v., leg. BOUCHET,
9-15.VII.1983, MNHN. 25 lots with numerous spm., sh. and v., from Sâo Miguel, Terceira, Faial channel
and Flores, between 39 and 225 m, empty shells and valves washed down on the steep slope to 1675 m, all
dredged R/V "Jean Charcot", BIACORES-Expedition, leg. MÉTIVIER, X.1971, all MNHN.

BIOTOPE : Byssally attached to rocks in crevices or in more open situations in tranquil sites such as
those at depth. Bathymétrie range from low in the intertidal to 100 m.
 — 307 —

I |20° |0° [20° I

MAP 2. — Distribution of Area avellana turbatrix (circles), A. a. avellana (shaded area) and A. tetragona (squares).
 — 308 —

REMARKS : This species probably cannot be considered to be a c o m p o n e n t of the tropical

West African fauna but as it can be confused with Area avellana turbatrix and juveniles of
Area noae we have included it here. Even when distorted it can be easily separated on the form
of the ligament and periostracal bristles.

Genus ACAR G r a y , 1857

TYPE SPECIES : Area gradata Broderip and Sowerby, 1829 (SD Woodring, 1925).

Acar cf. plicata (Dillwyn, 1817)

(PI. II, 7 A - B ; fig. 10; m a p 3)

Area plicata Dillwyn, 1817 : 227-228.

TYPE MATERIAL : Based on the figure in CHEMNITZ ( 1 7 9 5 : 2 4 4 , pi. 2 0 4 , fig. 2 0 0 8 ) .

TYPE LOCALITY : Red Sea.

DESCRIPTION

Shell to 26 m m in length. Equivalve. Initially slightly compressed but soon becoming

rather tumid to very tumid in gerontic specimens (the onset of the gerontic expansion can
occur in a wide range of sizes). Inequilateral, beaks in the anterior 1/4 rather low.
Outline subrectangular-subtrapezoidal, anteriorly usually slightly n a r r o w e d ; frequently
distorted especially in gerontic examples. Posterior area narrow, demarcated by a persistent
posterior carinal ridge; posterior margin oblique, posterior ventral junction subacute. Median
area flat to weakly sulcate; ventral margin more or less straight but often s i n u o u s ; byssal gape
obsolete or very narrow. Anterior area small, demarcated by a rounded ridge; anterior margin
broadly rounded. Dorsal margin straight.
Dorsal area usually very narrow, wider anterior to the beaks but becoming wide overall in
gerontic specimens; u m b o n a l separation usually slight. Ligament restricted to well behind the
beaks, chevrons very narrow, numbering up to 8 in normal specimens but as many as 20 in
gerontic examples.
Hinge plate narrow to moderate, teeth in 2 series but separation obsolete except in old
specimens where ligament growth has eroded the small central teeth; anterior set up to
10 teeth, posterior set u p to 15 teeth. Teeth strong, anterior laminar, posterior chevron
shaped, all coarsely serrated, marginal teeth becoming very oblique. In gerontic specimens the
teeth may be irregular, often divided medially.
Sculpture imbricate, of u p to 30 rather broad concentric ridges dissected by 25-35 radial
ribs these dividing during growth to number u p to 50. The resulting vesicular nodes are
smoothly rounded except on the posterior angle where they may be incomplete and therefore
slightly plicate. The posterior area bears 5-7 strong ribs on which the nodes are greatly swollen
 — 309 —

giving the cross section of the posterior margin a strongly corrugated appearance. This is a
little less so for the Cape Verde Islands material.
Periostracum very thin, rarely retained.
Shell whitish, exterior rather dirty c r e a m ; interior of Cape Verde, Sâo T o m é and
A n n o b o n specimens mostly flushed with pale orange/pink, the Angolan material rarely so.
Adductor scars more or less equal, raised with myophoric flanges along their inner edges.
Inner lateral margins crenulate, ventral margin weakly so or irregularly denticulate.

SELECTED SHELL MEASUREMENTS : See figure 9.

5
T
4 5.
X • 3.88 o
4-
3.5-

3-

2.5-
X= 1.93
2- o X-1.77

1.5-
0
li 1

L/A-Bk

FIG. 9. — Box plots of the ratios of Total Shell Length to Height ( L / H ) , Tumidity (L/T) and Anterior to Beak Length
(L/A-Bk) of Acar cf. plicata from West Africa (n = 24).

DISTRIBUTION : This species occurs from the Cape Verde Islands south to Angola but it is most
common on the Cape Verde Islands, the islands in the Gulf of Guinea and on the coast of Angola.

MATERIAL EXAMINED : Cape Verde Islands : (no precise locality), numerous spm. and v., leg. RANG,
1837 ; 4 spm., coll. DE CESSAC; 5 spm., coll. BOUVIER ; Santo Antäo (no precise locality), 5 v. ; Säo Vicente,
Baia Matiota, Mindelo, on the undersides of stones, 1-2,5 m, 3 spm., 2 v. ; Säo Vicente, Baia Porto
Grande, 2 sh., 10 v., both leg. VON COSEL, XII. 1978 ; Ilha do Sal, (no precise locality), numerous spm., sh.
and v., leg. CADENAT, 1950; Ilha do Sal, Palmeira, undersides of rocks at low tide, 7 spm. ; Ilha do Sal,
Baia Algodoeiro, on beach, 4 v. ; Ilha do Sal, Santa Maria, underside of boulders 2-3 m, 3 spm., all leg.
VON COSEL, 15.XII.1978-1.1.1979; Ilha do Sal, 16°35'N/22°55'W, calcareous algae, 15m, 1 spm., dredged
R/V "Calypso", 26.XI.1959; Boavista, Gâtas (Punta Rodrigo), on the undersides of stones on mixed
sand, l,5-2m, 6 spm., leg. VON COSEL, 27.XII.1978; Säo Tiago, Praia, 8 v., leg. CADENAT, 1950; Säo
Tiago, Porto de Praia, four lots with 3 spm. and 7 v., from 2 to 30m, all dredged R/V "Calypso", 17-
19.XI.1959; Brava SW of Punta Tantäo, 20m, 3 v. ; Fogo, SW of Santa da Encarnacäo, 20-25m,
12spm. ; Fogo, Punta da Araia, 2.5m, 2 spm. both dredged R/V "Calypso", 20.XI.1959; all MNHN.
Senegal : Dakar, SW of Cap Manuel, 50m, 1 v.; Dakar, 129-150m, 1 v., both dredged R/V "Gérard
Tréca", leg. MARCHE-MARCHAD, 1956 and 1958, MNHN, Liberia : 5°21,5'N/9°54,5'W, 73-80m, 4 v.,
dredged R/V "Calypso", leg. MARCHE-MARCHAD, 20.V.1956, MNHN. Côte d'Ivoire : Abidjan region (no
precise locality), 2 v., leg. LE LOEUFF, MNHN. Ilha do Principe : W-coast, 1°37'N/7°22'E, 30m, 1 v.;
Baia de Santo Antonio, between Punta da Mina and Punta de Novo Destino, 6m, 1 v.; Baia Santo
Antonio, between Punta da Mina and Ilheu Santa Ana, 10-12m, 1 spm.; both dredged R/V "Calypso",
leg. MARCHE-MARCHAD, VI. 1956, MNHN. Säo Tome : Punta Diogo Vaz, west coast, 30m, 1 spm.;
r38,4'N/7°22,l'E, 31m, 1 spm.; off S. Tomé city, 0°20' N/6°45' E, 10m, 2 spm., 5 v.; off Praia Lagarto,
 — 310 —

MAP 3. — Distribution of Acar cf. plicata (circles), A. plicata (shaded area) and A. pulchella (squares).
5-6m, 5 spm., all dredged R/V "Calypso", leg. MARCHE-MARCHAD, 11-27,VI, 1956; all MNHN.
Annobon : N of Santo Antonio, 23m, 1 spm.; 1°25,2'S/5°36,l'E, 20m, 1 spm., both dredged R/V
"Calypso", leg. MARCHE-MARCHAD, VI and VII. 1956; 1°24'S/5°37,5'E, 20-40m, 1 spm., 1 v., leg.
CROSNIER, 11.XII.1965; 1°28,5'S/5°37,5'E, 35-55m, 3 spm., leg. POINSARD, 16.VI.1967; all MNHN. R.P.
Congo : Pointe-Noire, Plage Mondaine, N of lighthouse, low tide, 7 v.; Plage Sauvage, on beach, 1 v.;
Plage ORSTOM, coarse sand and stones, 5-7m, 1 v., all leg. VON COSEL, XI-XII.1985, all M N H N ;
Loango, 4 v., leg. Office Pointe-Noire, 1969, MNHN. Angola : Cabo Ledo, Luanda province, 10-40 m,
2 spm.; Praia Etambar, Corimba, Luanda province, rocks, infralittoral, many spm.; Baia do Mussulo,
Luanda province, 5 spm.; Baia de Santa Maria, Benguela province, rocks, 0-2m, 18 spm.; Caotinha,
Benguela province, rocks, infralittoral, 7 spm.; Baia do Limagem, Benguela province, rocks, 0-2 m,
1 spm.; Baia de Lucira (Bissonga), Mocämedes province, intertidal, 4 spm.; Lucira (Praia do Cesar),
Mocämedes province, infralittoral, 3 spm.; Baia das Pipas, Mocamedes province, rocks, 2 spm.; Säo
Nicolau, Mocämedes province, intertidal, 3 spm.; Praia Amelia, Mocämedes province, rocks, infralittoral,
2 spm., all leg. S. GOFAS, 1982-1984, MNHN.

BIOTOPE : This species lives in crevices and under rocks where it is attached by a weak byssus. It is
found from low in the intertidal to 40 m.

FIG. 10. — Internal view of right valve of Acar cf. plicata (Dillwyn). Sào T o m é . (Scale b a r = 5 mm.)

REMARKS

The genus Acar is distinctive with its coarse imbricate sculpture and raised muscle scars
but the species systematics are very complex. Traditionally the species are identified on their
geographic distribution such that A. plicata is Indo-Pacific, A. pulchella (Reeve, 1844) is
Mediterranean, A. domingensis (Lamarck, 1819) is Caribbean and A. gradata (Broderip &
Sowerby, 1829) is Panamic. The West African form has been k n o w n by all but the Panamic
name. ROST (1955) in discussing A. gradata mentioned the coarse and fine sculptured varieties
of that species a n d we have observed the same feature in the Caribbean A. domingensis but not
to any great extent in the West African form. Even in the coarse sculptured form of A.
domingensis the nodes never appear to be so inflated as in the West African material and the
posterior ribs are finer or lower. The orange/pink colouration is apparently absent from
Caribbean material. In sculptural coarseness the Indo-Pacific A. plicata is very similar but
generally as the name implies the concentric elements are more lamellar especially on the
posterior angle which often appears foliate. The posterior ribs are similar in development but
again the nodes are not inflated or entire but are rather the form of domed plications. As with
the West African form the frequency of orange/pink shells is rather great. Acar pulchella
(pi. II, 8A-B) from the Mediterranean is a small species rarely exceeding 15 m m and differs
from all others in that the sculpture is primarily of fluted, concentric, slightly elevated lamellae.
The distinctions rest therefore on the coarseness of the sculpture and the degree to which
the radial and concentric elements fuse into either vesicular nodes of fluted lamellae. Given the
great range in variation we are refraining from giving the West African material separate
nomenclatural status. F r o m our observations the similarities in coarseness of sculpture,
distinctiveness of the posterior ribs, size and colour we conclude that the West African form is
closer to that of the Indo-Pacific. MEYER (1868) in discussing the Tertiary forms of Acar
suggested that A. clathrata (Defrance, 1816) was replaced by the Recent A. pulchella. In
general form the Pliocene species is close to A. plicata a n d could also be the fossil equivalent of
the West African form. W e recognise the affinity of the West African and Indo-Pacific forms
by using the name A. plicata.

Genus BARBATIA Gray, 1842

TYPE SPECIES : Arca barbata Linné, 1758 (SD Gray, 1857).

Barbatia complanata (Bruguiére, 1789)

(PI. I l l , 1A-B, 2, 3, 4, 5 A - B ; fig. 6 ; m a p 4)

Area complanata Chemnitz, 1784 : 198, pi. 55, fig. 544.

Area complanata Bruguiére, 1789 : 100.
Arca stigmosa Dunker, 1853 : 46, pi. 9, figs. 8-11.

TYPE MATERIAL : The type is based on the figure given by CHEMNITZ (1784) and the specimen may be
in the ZMC, Copenhagen. The type of A. stigmosa is probably in ZMB Berlin.

TYPE LOCALITY : This is cited as " G u i n e a " by CHEMNITZ which refers to tropical west Africa.
BRUGUIÉRE cites Madagascar and O. Americano but this is erroneous and arises from the early belief that
there was only a single pan-tropical species.

DESCRIPTION

Shell to 50 m m in length but usually not exceeding 35 m m . Equivalve. Compressed.

Inequilateral, beaks in the anterior third.
Outline subrectangular-subtrapezoidal, often somewhat distorted (some to the extent of
being subtrigonal), the anterior usually a little narrower. Posterior area poorly demarcated by
a low weakly angled posterior ridged but more noticeably by the change in the size of the r i b s ;
posterior margin oblique, m o r e or less straight to gently curved, posterior ventral junction
roundly subacute. Median area weakly sulcate; ventral margin subparallel to dorsal margin to
inclined towards the anterior, more or less straight or indented or distorted especially at byssal
 — 313 —

gape which is rather long but not very wide. Anterior area small, demarcated by a weak
rounded ridge, anterior margin broadly rounded but often restricted and the oblique, slanting
inwards. Dorsal margin long, straight.
Dorsal area narrow to very narrow, becoming deeply cleft, u m b o n a l separation slight.
Ligament initially opisthodetic, rapidly the dorsal area and extending anteriorly well beyond
the b e a k s ; chevrons narrow, n u m e r o u s in gerontic specimens, numbering u p to 12.
Hinge plate rather narrow especially centrally, teeth in two series which are almost
inseparable, total teeth count u p to 45. Central teeth very small, often destroyed by ligament
incursion, lateral teeth laminar or chevron shaped, becoming strongly oblique.
Sculpture of numerous radial riblets and ribs; posterior area with 6-7 large ribs often with
1-3 small secondary riblets. In some, the primaries are bifid, not greatly elevated, well spaced
and evenly dissected giving the appearance of a twisted r o p e ; anterior and median areas with
55-60 primary riblets, the anterior and those adjacent to the posterior angle a little larger than
the more closely spaced median riblets, posterior median riblets bifurcating at an early stage.
All radial elements dissected by concentric element and therefore appearing nodulose,
concentric element also present as widely spaced low ridges caused by the enlargement of the
nodules at regular intervals, these correspond with the concentric rows of larger periostracal
bristles.
Periostracum with a primarily concentric arrangement of raised lamellae strengthened by
slightly emergent bristles arising from the interspaces; the lamellae usually break down into
rectangular sections to give a frilled a p p e a r a n c e ; lamellae pale straw coloured, emergent
bristles brown giving a very characteristic pattern.
Shell white.
A d d u c t o r scars subequal the posterior a little larger, posterior pedal/byssal retractor scar
small, elongate. Inner margin finely crenulate.

SELECTED SHELL MEASUREMENTS : See figure 11.

DISTRIBUTION : This species ranges from Guinea (Conakry area) and the Cape Verde Islands (where
it is rare) south to southern Angola and is also found in Sao Tome, Principe and Annobon.

MATERIAL EXAMINED : Cape Verde Islands : Ilha do Sal (no precise locality), 1 dwarf spm., leg.
CADENAT, IV-VI.1950, MNHN. Guinea : Kassa, lies de Los, E-side, N of village, on the underside of a
stone, at low tide, 1 spm., leg. VON COSEL, 12.XI.1988, MNHN. Liberia : 6°40'N/11°23"W, 51 m, 1 juv. v.
dredged R/V "Calypso", leg. MARCHE-MARCHAD, 19.V.1956, MNHN. Cote d'lvoire : off Tabou,
o
4°16,5'N/7 30'W, hard bottom, 40m, dredged R/V " L a Rafale", Guinean Trawling Survey, leg.
CHERBONNIER, 8.IV.1964; Abidjan region, "Palm Beach", on rocks, 6-10m, 1 spm.; Canal de Vridi,
Abidjan, 6 spm., 1 sh., 3 v.; Abidjan, SE of " 2 Poteaux", on sandstone, 37m, 1 juv. spm., all leg. LE
LOEUFF, 1967-1973; all MNHN. Ghana : Cape Coast, hard bottom, 26-31 m, 1 spm., leg. LE LOEUFF,
10.11.1968; 4°37'N/0°50'W, 90-100m, 3 v., dredged R/V "Calypso", leg. MARCHE-MARCHAD, 24.V.1956,
both MNHN. Togo : 6°10'N/r37'E, 19m, 2 juv. spm., leg. CROSNIER, 26.VII.1964, MNHN. Benin :
Ouidah, 6°10'N/2°05'E, 100m, 5 juv. v., leg. MARCHE-MARCHAD, 22.XI.1958, MNHN. Nigeria : off the
Niger delta, 4°00'N/6°H'E, 34m, on Chama, 1 juv. spm., dredged R/V "Calypso", leg. MARCHE-
MARCHAD, 26.V.1956, MNHN. Cameroon : Victoria (now Limbe), Morton Bay, E-side, between rocks at
low tide 14 spm., leg. VON COSEL, 1-3.XII.1985; Mondoleh Island, Ambas Bay, under and between stones
near low tide, numerous spm., leg. VON COSEL, IV. 1969; Kribi, on beach, 8 v., leg. NICKLES, 1947; 3 spm.,
leg. VON COSEL, IV. 1969; off Kribi, 2°36,8'N/9°46'E, 1 spm.; 2°33,3'N/9°42,6'E, 1 spm., both leg.
CROSNIER, XII.1962; all MNHN. Ilha do Principe : Baia de Santo Antonio, 6-15m, 4 spm., 1 sh., 8 v.;
Baia das Agulhas, 4-8 m, 1 spm.; 1°37'N/7°22'E, 30 m, 3 juv. v.; 1°42' N/7°28,8' E, 37 m, 1 spm.;
 — 314 —

L/A-bk

FIG. 1 1 . — Box plots of the ratios of T o t a l Shell Length to Height ( L / H ) , Tumidity (L/T) and Anterior t o Beak
Length (L/A-Bk) of Barbada complánala from West Africa (n = 1 4 ) .

r 120° 10° I
MAP 4 . — Distribution of Barbatia complanata.

1°20,7'N/7°17,6'E, 25-40m, 1 dwarf spm., all dredged R/V "Calypso", leg. MARCHE-MARCHAD, 25.VI-
1.VII.1956; all MNHN. Sao Tome : Esprainha, near Neves, 3 spm.; Praia Morro Peixe, 1 spm.; Praia
das Conchas, 2 spm.; Praia Emilia, several spm., all between and under rocks at low tide, leg. GOFAS,
XI. 1983; Punta Diego Vaz, 0,6 m, 3 spm.; Ilheu das Cabras, 1 sh.,offPta. Diogo Nunes, 4 m, 1 spm.; off
Praia Lagarto, 5-6m, 4 spm., 5 juv. v., all R/V "Calypso", leg. MARCHE-MARCHAD, 6-11.VI.1956; all
MNHN. Annobon : 1°24' S/5°37,5' E, 20-40m, 2 spm., leg. CROSNIER, 11.XII.1965; 1°24' S/5°36,7' E, 10m,
1 spm.; 1°25,2'S/5°36,l'E, 20 m, rocks with calcareous algae, 2 spm., both dredged R/V "Calypso", leg.
 — 315 -

MARCHE-MARCHAD, 13.VI.1956; all MNHN. Gabon ; off Port-Gentil, on oil platform anchor, 30m,
1 spm., leg. CHEVALIER, 1987; Port-Gentil, 2 spm., coll. Office Pte-Noire, both MNHN. R.P. Congo ;
Pointe-Noire, Plage Mondaine, on beach. 1 v., leg. VON COSEL, XII. 1985, MNHN. Angola : Cabo Ledo,
Luanda province, 10-40m, 1 sh.; Caotinha, Benguela province, 10 spm., 1 sh.; Baia Azul, Benguela
province, rocks at low tide, 4 spm., 1 sh.; Baia do Limagem, Benguela province, rocks, 0-2 m, 1 spm.;
Baia de Santa Maria, Benguela province, rocks 0-2m, 3 spm., 1 v.; Baia da Lucira (Bissonga),
Mocämedes province, rocks at low tide, 1 spm.; Lucira (Praia do Cesar), Mocämedes province, 3 spm.;
Chapeu Armado, Mocämedes province, rocks at low tide, 5 spm.; Praia Amelia, Mocämedes province,
rocks at low tide, 3 spm., all leg. GOFAS, 1982-85; all MNHN.

BIOTOPE : Byssally attached to rocks, stones, shells and other hard objects, from low tide about 40 m.

REMARKS

This species despite its variability should n o t be confused with other west African species
of Barbatia as it is the only species with the distinctive well spaced concentric ridges which give
the periostracum a frilled appearance. It is also the only species to be encountered in the rocky
lower intertidal a n d sublittoral.
We investigated the possibility that as in the Indo-Pacific there might be two closely
related species equivalent to B. foliata (Forsskál, 1775) a n d B, decussata (Sowerby, 1833).
Despite the variability in form of the west African specimens we could find n o indication of
pattern except that crevice dwelling examples were usually fore shortened o r distorted.
Barbatia complanata belongs to the g r o u p of species represented by B. candida (Helbling,
1779) in t h e Caribbean, by B. foliata (Forsskál, 1775) in the Indo-Pacific a n d by B. reeveana
(d'Orbigny, 1846) in the Panamic province. The west African species differs in having a much
finer sculpture with approximately twice as many anterior a n d median primary riblets a n d in
the posterior riblets narrower, more regular a n d numbering from 6-7 rather than the 3-4
typical of the Caribbean species. The periostracum of B. candida is heavier a n d the emergent
bristles a r e consequently much longer a n d more robust. T h e Indo-Pacific B. foliata is a n
altogether much larger species with a more coarse sculpture which lacks the regular, well
spaced concentric ridges. The west African species is therefore closer to that of the Caribbean.
American authors place these species in the subgenus Cucullearca C o n r a d , 1865. These
species are distinct from those of Barbatia sensu stricto such as B. barbata a n d B. gabonensis
(see below) b u t we are not certain of the affinities with other, mainly Indo-Pacific, groups and
refrain from using this taxon until a more thorough revision is completed.

Barbatia gabonensis n. sp.

(PI. I l l , 6 A - B ; fig. 14; m a p 5)

TYPE MATERIAL : Holotype (27.7 mm), MNHN, off Mayumba, Gabon, 3°25' S/9°56' E, 100 m, trawled
R/V "Thierry", Guiñean Trawling Survey, Sta. 58/6, 30.XI.1963. Paratypes : NMWZ, Ilha de Luanda,
Angola, 90m, 1 spm., leg. GOFAS; MNHN, off Grand Bassam, Cote dTvoire, 5°00'N/3°23'W 100m,
1 spm., trawled R/V " L a Rafale", Guiñean Trawling Survey, Sta. 26/6.

TYPE LOCALITY : Mayumba, Gabon.

FIG. 12. — Internal views of right valves of Barbatia complanata (Bruguière) : A , C ô t e d ' I v o i r e ; B, Angola. (Scale
bar = 1 0 m m . )

DESCRIPTION

Shell to 30 m m in length, not very thick for the genus. Equivalve, not inflated.
Inequilateral, beaks in the anterior quarter.
Outline roughly rectangular, longer t h a n high, length : height ratio from 2.2-2.6 : 1, dorsal
and ventral margins more or less parallel. Posterior area indistinct; posterior angle smoothly
r o u n d e d ; posterior margin oblique, gently curving to posterior ventral junction which is
narrowly rounded or subacute. Median area flattened or weakly depressed; byssal gape very
narrow, s h o r t ; ventral margin gently curved to straight, byssal sinus if present, weak. Anterior
margin in smaller shells broadly r o u n d e d ; later becoming m o r e straight, oblique with a
narrowly rounded anterior ventral junction. Dorsal margin moderately long, more or less
straight.
Dorsal area very narrow, cleft; u m b o n a l separation slight. Ligament restricted to area
between and behind the beaks, chevrons narrow, numbering u p to 7.
Hinge plate moderately thick, teeth in two series which are barely separable but the
junction lies below the beaks, anterior set u p to 10 teeth, posterior set u p to 16 teeth. Teeth
serrated, straight or slightly chevron shaped, becoming a little oblique towards the margins of
the plate.
Sculpture fine of nodulose radial riblets, 90-110 in n u m b e r m o r e or less uniform in size
with the exception 13-18 pairs of larger riblets which border distinct radial grooves.
Periostracum mostly lamellar except for long, subtubular, semispathulate bristles arising
from the radial grooves. Lamellae straw coloured, bristles dark brown.
Shell mostly white, some with faint tinges of brown or brownish pink colour.
A d d u c t o r scars subequal, the posterior a little longer, posterior pedal/byssal retractor
long and very narrow.
 SELECTED SHELL MEASUREMENTS

Tumidity Anterior
Length Height
margin to beak Rib No. Groove No.
(mm) (mm) (mm)
(mm)

26.9 16.1 12.1 8.3 92 13

29.9 16.9 13.5 7.6 105 17
22.0 11.8 8.7 5,6 100 16
23.5 13.2 10.2 6.4 98 16
26.2 15.6 10.7 7.7 108 15
21.5 15.6 8.4 5.9 109 18

DISTRIBUTION : Barbatia gabonensis is an entirely tropical species with a restricted distribution from
Liberia south to northern Angola.

FIG. 1 3 . — Box plots of the ratios of T o t a l Shell Length to Height ( L / H ) , T u m i d i t y (L/T) a n d Anterior to Beak
Length (L/A-Bk) of Barbatia gabonensis from West Africa.

MATERIAL EXAMINED : The type material; Liberia : 4°37'N/10°50'W, 90-100m, 2 v., dredged R/V
"Calypso", leg. MARCHE-MARCHAD, 24.V.1956, MNHN. R.P. Congo : off Pointe-Noire (no precise
locality), 1 spm., leg. MARCHE-MARCHAD, MNHN.

BIOTOPE : Barbatia gabonensis is a deep water form with the present data indicating a range around
100 m. The byssus is functional although small and this suggests that this species lives attached to rock,
stones or other hard substrates. The byssus is much weaker than in related forms from the littoral and
sublittoral zones further suggesting that the local conditions are tranquil or sheltered. This could be
explained by the lack of wave action at the depths from which B. gabonensis has been recovered.

DERIVATIO NOMINIS : gabonensis from Gabon, the country where the type locality is situated.

REMARKS : The immediate distinctive feature of this species is the series of radial grooves
and the long periostracal bristles arising from them. This distinguishes it from the Barbatia
 — 318 —

LOE
O
È: O
CM

O
' ' À o
1
% .

* 0

— 1 o

*
r

120° |0°

MAP 5. — Distribution of Barbatia gabonensis (circles) and B. ioníhados (triangles).

candida-complanata-foliata complex in which the bristles are not differentiated. Some species
have differentiated bristles but none so marked or in such widely separated rows as B.
gabonensis. Barbatia sculpturata T u r t o n , 1932 is also much smaller and has only a few broad
ribs on the posterior area, it is restricted to the sublittoral zone of the coasts of Natal and Cape
Province (S. Africa) (KILBURN, 1983). Barbatia barbata (Linné, 1758) from the Mediterranean
has a similar bristle pattern when small but later there is much less distinction between the
primary and secondary bristles and in the largest specimens the whole of the dorsal area is
thatched. F u r t h e r m o r e in B. barbata the ligament fills the whole of the dorsal area. Barbatia
pistacia (Lamarck, 1819) from south eastern Australia has a similar rib pattern but the bristles
are poorly differentiated and the rows of larger bristles are much more numerous.

Barbatia ionthados n. sp.

(Pl. I l l , 7 A - B ; fig. 15; m a p 5)

TYPE MATERIAL : Holotype (20.6 mm), MNHN, South of Bassam, Côte d'Ivoire, 200 m, leg. MARCHE-
MARCHAD, 13.X.1960. Paratypes : MNHN, off Cape three Points, Ghana, 4°43,5'N/2°45,5'W, 100m,
1 spm., trawled R/V " L a Rafale", Guiñean Trawling Survey II, Sta. 27/6, 20.111.1964; NMWZ, as
previous, 1 spm.

TYPE LOCALITY : South of Bassam, Côte d'Ivoire.

 — 319

FIG. 1 4 - 1 5 . — 1 4 : Internal view of right valve of Barbatia gabonensis n. sp. G a b o n . 1 5 : Internal view of right valve of
Barbatia ionthados n. sp. Cote dTvoire. (Scales bars = 5 m m . )

DESCRIPTION

Shell to 21 m m in length, very fragile. Equivalve. A little compressed. Very inequilateral,

beaks in the anterior fifth.
Outline elongate rectangular, length to height ratio 2.6-2.7 : 1, dorsal and ventral margins
more or less parallel. Posterior area not expanded but rather long, posterior angle rounded but
distinct, posterior margin oblique to broadly rounded, posterior ventral junction rounded.
Median area lacking sinus, ventral margin more or less straight, byssal gape minute. Anterior
margin broadly rounded. Dorsal margin long, m o r e or less straight.
Dorsal area very narrow, slightly cleft, u m b o n a l separation slight. Ligament situated
entirely behind the beaks, chevrons n a r r o w numbering u p to 4.
Hinge plate long, n a r r o w ; teeth in two series, separated by a very small gap at a point
below the b e a k s ; anterior set u p to 9 teeth, posterior set u p to 31 teeth. Teeth small, more or
less straight, finely serrated; central teeth more or less vertical, lateral teeth slightly oblique.
Sculpture very fine, of weakly nodulose radial riblets, 88-96 in number but those on the
anterior margin are difficult to distinguish and some shells m a y have more riblets. The riblets
show no pattern except that they become progressively finer towards the anterior.
Periostracum of long very fine dark straw coloured suberect bristles which are longest on
the posterior angle and most erect on the posterior slope. There is no trace of inter bristle
lamellae.
Shell uniformly white.
Inner margin weakly serrate corresponding to radial riblets. A d d u c t o r scars subequal,
posterior pedal/byssus retractor narrowly elongate. Byssus a thin n a r r o w strap.
 — 320 —

SELECTED SHELL MEASUREMENTS

Anterior
Length Height Tumidity
margin to beak Rib No.
(mm) (mm) (mm)
(mm)

20.6 9.7 7.5 4.0 88

19.1 9.3 7.3 3.8 96
11.1 5.6 4.2 2.1 90

DISTRIBUTION : Living specimens have been taken only off the northern coast of the Gulf of Guinea.
A number of worn valves from off Senegal suggests a wider distribution and one might expect this species
to occur throughout tropical West Africa.

MATERIAL EXAMINED : The type material; Senegal : off Goree, 80 m, 8 v., poorly preserved, dredged
R/V "Gerard Treca", leg. MARCHE-MARCHAD, 20.11.1956, M N H N .

BIOTOPE : From the limited material available B. ionthados is apparently a deep water species living
from 100-200 m. The functional byssus and general shell form suggest that it is epibyssate living attached
to hard substrates. As with its Indo-Pacific counterpart Barbatia tenella Reeve, 1844 it may live in crevices
or under stones where there is shelter but in the deep waters preferred by B. ionthados such protection
may not be required.

DERIVATIO NOMINIS : ionthados, Greek, "long haired" or "shaggy", referring to the form of the
periostracum.

REMARKS : Only one other species resembles B. ionthados, that is B. tenella Reeve, from
the Indo-Pacific which shares the fragile shell, posterior ligament, fine sculpture and an entirely
spicate periostracum. D A L L , BARTSCH and REHDER (1938) erected the genus Barbarca and
IREDALE (1939) the genus Opularca, for species close to this form. As they are clearly linked to
Barbatia they would only warrant subgeneric status of which the earliest n a m e is Barbarca.

Barbatia legumen (Lamy, 1907)

(PI. I l l , 8, 9 A - B , 10; fig. 16-18; m a p 6)

Area legumen Lamy, 1907 : 74-76, pi. 1, fig. 3-4.

TYPE MATERIAL : M N H N , 16 syntypes from three different localities: 5 spm. Gabon, coll. DIBOWSKY,
1893; 5 spm., no locality, coll. LARGENTIERE, 1887; 2 spm., 4 v., " Bata Congo" ( = Bata in Equatorial
Guinea), coll. POBEGUIN, 1891. Among the latter lot is the specimen figured by LAMY (1907) and this is
here selected as the lectotype.

TYPE LOCALITY : Bata, Equatorial Guinea. On the original label of the POBEGUIN collection this
locality is always cited as " Bata Congo ", at that time it belonged to the French Congo and did not
become Spanish until 1900.
 DESCRIPTION

Shell to 52 m m in length. Equivalve. Compressed. Inequilateral, beaks in the anterior

third.
Outline of adult submytiliform, often arcuate, anterior part narrow, posterior part
expanded. Posterior area small ; posterior angle broad, rounded and forming greater part of
the posterior half of the shell ; posterior margin long, oblique, sloping steeply, dorsal auricule
indistinct or lacking. Median area distinctly sinuate, ventral margin long, arcuate. Anterior
area small, demarcated by a rounded anterior angle; anterior margin narrowly rounded.
Dorsal margin moderately long, straight. Outline of juveniles similar but posterior margin
shorter and distinctly auriculate, ventral margin less arcuate and anterior area larger but
anterior angle indistinct. The increasing arcuate development with age is illustrated in
figure 17.
Dorsal area very narrow, deeply cleft ; umbonal separation slight, ligament barely visible
in joined valves. Ligament filling the whole of the dorsal area including the anterior p a r t ;
ligament almost entirely lamellar, chevrons very strong numbering u p to 5.
Hinge plate moderately thick ; teeth probably in two series but the central teeth are
usually worn giving the appearance of distinct anterior and posterior sets. W h e n well preserved
u p to 50 teeth in all. Teeth slightly chevron shaped, finely serrated, becoming oblique laterally.
Sculpture of very n u m e r o u s radial riblets, usually worn but best preserved in median
sinus.
Periostracum initially with broad appressed bristles on posterior angle but later consisting
primarily of thick concentric lamellar layers with scarcely emergent bristles. Colour a deep
tanned dark brown.
Shell external colour white to cream, internally bluish in juveniles white in adults.
A d d u c t o r muscle scars unequal, the posterior twice the size of the anterior. Posterior
pedal/byssus retractor small, narrowly oval.

SELECTED SHELL MEASUREMENTS : See figure 16.

DISTRIBUTION : Barbatia legumen occurs only in the southern part of tropical West Africa, from
Equatorial Guinea to southern Angola (Santo Antonio). The greater part of this range lies within the
influence of the cold Benguela Current which in the southern winter can reach the coast of Gabon. Here
and in Equatorial Guinea it seems to be very rare.

MATERIAL EXAMINED : The type material ; R.P. Congo : Pointe Indienne, rocky shore 1 sh. ; Pointe-
Noire, Plage Sauvage, on beach, 3 v. ; Plage Mondaine, on beach, numerous sh. and v. ; off Plage
ORSTOM, 3-7 m, numerous sh. and v., all leg. VON COSEL, XI-XII.1985 ; several sh. and v., leg. CROSNIER
(Office Pte.-Noire); all MNHN. Angola : 10km S of Ambrizete, Zaire province, intertidal, on beach,
many fresh sh. and v. ; Barra do Dande, Bengo province, between rocks at low tide, 2 spm. ; Baia Sào
Tiago, Bengo province, between rocks at low tide, 4 spm. ; Cacuaco, Bengo province, between rocks, 0-
1 m, 14 spm. ; Corimba, Luanda province, Praia Etambar, between rocks at low tide, 5 spm., 1 sh. ; Cabo
Ledo, Luanda province, 10m, 11 juv. spm., 3 juv. sh. ; Santo Antonio, Benguela province, between rocks
at low tide, 1 spm. ; Baia das Pipas, Moçâmedes province, on rock platform, 1 juv. spm. ; Porto
Alexandre, Moçâmedes province, 2m, 1 juv. v., all leg. GOFAS, 1982-85; all MNHN.

BIOTOPE : This species lives byssally attached to rocks, on the undersides of stones and to secondary
hard substrates on soft sediments. The bathymétrie range is from the intertidal to 1-2 m in the sublittoral.
 — 322 —

4.25, 1 1 1 L

3.75.

FIG. 1 6 - 1 7 . — 1 6 : Box plots of the ratios of Total Shell Length to Height ( L / H ) , Posterior Height ( L / P H ) , Tumidity
(L/T) and Anterior to Beak Length (L/A-Bk) of Barbatia legumen from West Africa. 1 7 : G r a p h of the ratio Total
Shell Length : Difference in M e d i a n and Anterior Height plotted against T o t a l Shell Length on log axis, showing
the increasing arcuate form of the shell with age in Barbatia legumen.

REMARKS

Barbatia legumen belongs to a g r o u p of species which are submytiliform in shape and are
typical inhabitants of the littoral and shallow sublittoral t h r o u g h o u t the Indo-Pacific. T h e
most widespread of these related species are B. obliquata (Gray, 1837) [Indian Ocean] and B.
virescens (Reeve, 1844) [Indo-West Pacific]. B. obliquata is more expanded posteriorly, not
auriculate and the ligament is restricted to behind the beaks. B. virescens can be auriculate and
the ligament is on both sides of the beaks but the outline is much shorter, the sculpture more
pronounced and the periostracum more spicate.
Some submytiliform species have been given generic r a n k ; Savignyarca Jousseaume, 1898
(type S. savignyarca = Barbatia obliquata); Obliquarca Sacco, 1898 (type, Area modiliformis
Deshayes, 1831) and Barbatirus Iredale, 1939 (type B. minulus Iredale, 1939). H A B E (1977)
regards these as synonyms of Savignyarca but this presupposes that the submytiliform species
have a c o m m o n ancestry. We d o not regard the c o m m o n adaptive form as sufficient evidence
to conclude monophyly because the incidence of convergent radiations within the Arcoidea is
 — 323 -

well documented (STANLEY, 1972). In recent forms the convergent radiations within the
Arcidae and Noetiidae illustrate the point.

FIG. 1 8 . — Internal view of right valve of Barbatia legumen Lamy. Angola. (Scale b a r = 1 0 mm.)

Subgenus N I P P O N A R C A H a b e , 1951

It has not been our intention to introduce subgenera within Barbatia but in this case we
have both anatomical and shell characters which allow us to confirm that Nipponarca is a
useful division.

TYPE SPECIES : Area bistrigata Dunker, 1858 : 87, pi. 30, fig. 4, see also HABE, 1951, fig. 49.

REVISED DIAGNOSIS : Shells to 50 mm. Equivalve. A little compressed. Inequilateral, beaks in the
anterior third. Outline subrectangular, distinctly longer than high, dorsal and ventral margins subparallel.
Hinge plate long, straight, teeth small. Dorsal area flat or almost so. Sculpture of few radial ribs;
posterior ribs rather broad, flat and weakly sculptured; median ribs bifurcate, nodulose. Periostracum
with long semi-erect bristles arising from the interspaces of the posterior angle. Labial palps very large
with numerous sorting ridges. Byssate species inhabiting inshore areas where the conditions are turbid
and often under estuarine influences.

SPECIES INCLUDED : Barbatia (Nipponarca) bistrigata (Dunker, 1858) [Indo-Pacific, Pakistan to

Japan]. Barbatia (Nipponarca) allocostata this paper [tropical west Africa, Senegal to Sierra Leone].
Probably Barbatia signata (Dunker, 1858) [Indo-Pacific, India to China].

Barbatia (Nipponarca) allocostata n. sp.

(PI. IV, 1A-B, 2 ; fig. 19, 2 0 ; m a p 6)

TYPE MATERIAL Holotype (28.4 mm), MNHN, Mouth of Rio Geba, Guinea-Bissau,
ir57,5'N/16"27'W, 7m, agglomerations of live and dead Crassostrea gasar, on oyster shells, trawled R/V
"Andre Nizery", leg. VON COSEL, 9.X.1988. Paratypes as holotype; MNHN, 9 spm.; NMWZ, 1990.017,
5 spm.

TYPE LOCALITY : Mouth of Rio Geba, 11°57,5'N/16°27'W, Guinea-Bissau.

 - 324 -

DESCRIPTION

Shell to 35 m m in length. Equivalve. A little compressed, most inflated across the anterior
u m b o n a l region and narrowing posteriorly. Inequilateral, beaks in the anterior third.
Outline subrectangular, much longer than high. Posterior area long, weakly demarcated,
posterior angle rounded and becoming obsolete; posterior margin initially rounded, more or
less vertical becoming oblique and some auriculate; posterior ventral junction rounded.
Median area sulcate, ventral margin more or less straight, usually indented at byssal gape
which is relatively short and narrow. Anterior area short, demarcated by weak anterior ridge;
anterior margin broadly rounded. Dorsal margin long, straight.

FIG. 19-20. — 1 9 : Barbatia (Nipponarca) allocostata n. sp. A , internal view of right valve; B, external view of left
valve with periostracum removed. Guinea-Bissau. 20 : Size series of dorsal views of Barbatia (Nipponarca)
allocostata n. sp. Guinea-Bissau. (Scales bars = 5 mm.)

Dorsal area narrow, weakly cleft to flat, u m b o s well separated. Ligament initially behind
the beaks but developing anteriorly in larger specimens; chevrons few, numbering u p to 3.
Hinge plate narrow, teeth in two series but separation is indistinct, total number 50. Teeth
small, weakly serrated, more or less straight, vertical centrally becoming a little oblique
laterally.
 — 325 -

Sculpture of relatively few, 26-28, radial ribs. Posterior 10-12 ribs broad, flat, more or less
s m o o t h ; median ribs deeply bifurcate, n o d u l o s e ; anterior ribs simple, irregularly nodulose.
Periostracum of very long semierect bristles arising from the rib interspaces over the
posterior angle, elsewhere the pattern is the same but the bristles are much shorter and fragile.
Bristles dark brown, remainder paler.
Shell mostly white but externally u m b o s grey and some with brown blotches on the
posterior a r e a ; internally the u m b o n a l cavity is suffused with green or greyish green tints.
Adductor muscle scars unequal, the posterior half as big again as anterior scar. Posterior
pedal/byssus retractor scar narrowly elongate.

Anatomy (the following account is based on preserved specimens only)

The mantle edge (fig. 21) is entirely unfused and thickened posteriorly where it is thrown
into small folds. Pigmentation is weak with only a few brown patches on the exterior of the
outer fold in the posterior ventral region. Eyespots are absent but there is a single anterior
dorsal pigment spot.
The gills (fig. 21) are typically arcoid and the gill axis is contorted suggesting that it is
muscular and capable of extending the gills to the margin of the shell. The labial palps (fig. 21)
are very large, semicircular in outline and with a b o u t 90 sorting ridges. These palps are very
large for the genus, most being n a r r o w with a b o u t 15-20 ridges.

MEP

FIG. 2 1 . — G r o s s a n a t o m y of Barbatia (Nipponarca) allocostata n. sp. viewed from the left side after removal of the
left valve and mantle. (Scale bar = 5 m m . )
A A , anterior a d d u c t o r muscle; B, b y s s u s ; C T , c t e n i d i u m ; F , foot; LP, labial p a l p s ; M E , folded mantle e d g e ;
M E P , m a n t l e edge p i g m e n t a t i o n ; PA, posterior a d d u c t o r m u s c l e ; P P R , posterior pedal retractor muscle.

The gut is typical of the genus with a short oesophagus entering the stomach on its
anterior dorsal face; the mid gut exits from the ventral floor of the stomach and passes deeply
into the foot where it is weakly folded before forming a large anterior loop, then it passes
dorsally across the right hand side of the stomach before bending posteriorly as the hind gut.
T h e anterior loop is m o r e prominent than in other species of Barbatia. T h e stomach is
typically arcoid but the food sorting area is restricted to the anterior dorsal food sorting
caecum which is endowed with prominent sorting ridges. The right and anterior ventral
 — 326 -

embayments have openings to the digestive diverticula and ridges are restricted to the rejection
tract which follows the major typhlosole.
The foot (fig. 2 1 ) is large with a well developed heel a n d toe. The byssus groove is
centrally placed and produces a relatively large but narrow sheet byssus (fig. 2 1 ) typical of the
genus. The size of the byssus is reflected in the large posterior pedal/byssus retractor muscles
which are narrowly elongate in section.
The a n a t o m y of this species is similar in most ways to that of Barbatia as a whole in that
it reflects an epibyssate habit. Significantly different is the size of the labial palps which are
more typical of infaunal forms such as some Anadara species (HEATH, 1 9 4 1 and LIM, 1 9 6 6 ) and
a new genus to be described in Part two of this revision. The biotope described below does
appear to be one where resuspended matter is dense and this would explain the adaptive value
of this epibyssate species having a large sorting capacity. A little surprisingly are the
apparently weak mantle edge folds, these are usually more developed in species which inhabit
m u d d y biotopes. The lack of pigmentation and eyespots on the mantle edge are not
characteristics of Barbatia but again can be explained if the water column is so full of
suspended matter that light is virtually excluded.

SELECTED SHELL MEASUREMENTS : See figure 22.

DISTRIBUTION : This species seems to have a limited distribution to the northern part of tropical West
Africa. It is actually known from the extreme south of Senegal (southern Casamance) to the border
between Guinea and Sierra Leone. Most probably it goes further south into Sierra Leone but we have
never seen material from south of 9°N.
3.75.

3.5.

3.25.

3.

2.75.
V)
o
1 2.5. X = 2.27
OC
2.25.

2.

1.75.

1.5.
UT L/A-Bk

FIG. 22. — Box plots of the ratios of Total Shell Length to Height ( L / H ) , Tumidity (L/T) and Anterior to Beak
Length (L/A-Bk) of Barbatia (Nipponarca) allocostata from West Africa.

MATERIAL EXAMINED : The type material; Senegal : Karabane Bolon, Casamance, off Karabane, 4m,
7 v.; Casamance, creek off Elinkine, 3 m. 3 v.; Ourong Bolon, S-Casamance, 2-4 m, 2 v.; Cap Skirring-
Diembering, on beach, 1 v.; Essoukoudiak Bolon, on the frontier to Guinea-Bissau, 4-6 m, coarse sand
with pebbles, 1 spm., many v., all leg. VON COSEL, 5-17.III. 1988, all MNHN. Guinea-Bissau : Mouth of
Rio Geba, H°57,5'N/16°27'W, 7m, on oysters (from type lot), 48 associated spm., MNHN. Guinea :
Kaporo, NW coast of Conakry Peninsula, in front of ORSTOM, low tide, undersides of rocks, 1 spm.;
Kaporo Port, Conakry Peninsula, rocks at low tide, 1 v.; Dixinn Port, NW coast of Conakry Peninsula,
gravel, low tide, 1 v.; Conakry, Corniche Sud, in front of the " People's Palace ", rocks at low tide, 4 v.;
Kassa, lies de Los, E-side, N of village, on the underside of a stone at low tide, 1 spm., all leg. VON COSEL,
V and XI. 1988, all MNHN.
 — 327 -

MAP 6. — Distribution of Barbatia legumen (circles) and B. allocostata (triangles).

BIOTOPE : Barbatia (Nipponarca) allocostata occurs on hard bottoms such as stones and rocks and is
byssally attached. It is frequently found on offshore oyster beds consisting of Crassostrea gasar
(Dautzenberg, 1891) where it is attached in crevices and between oyster shells. In Guinea-Bissau it was
taken in trawls from sandy bottoms where it was found on shells of living Cymbium pepo (Lightfoot,
1786) which were also covered by small oysters. It was likewise found on living or hermit crab inhabited
shells of Pugilina morio (L., 1858) and even on oyster covered valves of large bivalves like Macoma
cancellata (Sowerby, 1873) and Mactra rostrata Spengler, 1802. B. allocostata seems to be limited to
murky, sediment and seston laden water in regions with estuarine influences and seasonal salinity changes.
On the rocky shores of the Conakry peninsula it was found on the sides and undersides of stones at low
tide and is a rare but regular member of the Crassostrea gasar community. B. allocostata is not common
in the intertidal and is most frequently found offshore to about 15 m.

DERIVATIO NOMINIS : allocostata, Greek, compounded from allasso meaning " to change" and
costatus meaning " t o bear ribs". This refers to the abrupt change in the form of the radial ribs from the
median to posterior areas.

REMARKS : Barbatia (Nipponarca) allocostata is very closely allied to B. (N.) bistrigata

(Dunker, 1858). The latter species is heavier shelled, reaches a larger size, the marginal
crenulations are stronger, the anterior most, posterior ribs develop u p to 4 ridges which d o not
bisect the rib and the posterior angle periostracal bristles are shorter and more stout.
 - 328 -

Genus BENTHARCA Thiele, 1931

TYPE SPECIES : Area nodulosa Müller, 1776.

Bentharca asperula (Dali, 1881)

(PI. VIII, 6 A - B ; fig. 23)

Macrodon asperula Dali, 1881 : 120-121.

Area profundicola Verril & Smith in VERRILL, 1885 : 439-440, pl. 44, flg. 23-23a.
Area (Barbatia) pteroessa Smith, 1885 : 844.

TYPE MATERIAL : In USNM.

TYPE LOCALITY : Off S. E . United States, " B l a k e " sta. 33, 2875m.

DESCRIPTION

Shell to 12 m m in length. Equivalve. Somewhat compressed. Inequilateral, beaks in the

anterior quarter.
Outline subtrapezoidal. Anterior area greatly restricted, posterior roundly expanded
giving an overall wedge-shaped appearance. Posterior margin broadly r o u n d e d ; ventral
margin sloping to narrow anterior margin, the latter slightly indented by byssal sinus. Median
byssal sinus present at all sizes but byssal gape indistinct. Dorsal margin long and straight.
Dorsal area long, narrow initially but expanding rapidly with age. Ligament restricted to
the posterior area but filling area between beaks with maturity.
Hinge plate very n a r r o w ; teeth in two series, separated by a wide edentulous g a p . Teeth
numbering up to five in each set; posterior teeth becoming subparallel to dorsal margin.
Sculpture very weak, of faint radial striae only. In other regions the sculpture can be a
little stronger with faint raised riblets.
Periostracum usually worn a n d preserved at margins o n l y ; of very slender appressed
bristles.
Colour white, periostracum a dirty olive brown.
A d d u c t o r muscle scars indistinct but unequal, the posterior a b o u t twice the size of the
anterior. Byssus retractor muscle small, reflecting the weak byssus which consists of a single
narrow strap.

DISTRIBUTION : A cosmopolitan species restricted to the abyssal zone from 2000-5000 m.

MATERIAL EXAMINED : Based on the data in OLIVER and ALLEN (1980).

BIOTOPE : Bentharca asperula lives attached to small stones, gravel and grit particles.

REMARKS : T w o species of Bentharca occur in the Atlantic Ocean but the second B.
nodulosa (Müller, 1776) has not yet been recorded from tropical West Africa. B. nodulosa lives
 - 329 -

at shallow depths in the subarctic region and undergoes increasing submersion towards the
south of its range reaching 3000 m off Madeira. It is a larger shell with a more coarse sculpture
resembling that of the genus A car.

FIG. 23. — Internal view of right valve and dorsal view of Bentharca asperula (Dall). Angola, 3975 m. (Scale bar =
2 mm.)

Subfamily ANADARINAE

Genus ANADARA Gray, 1847

TYPE SPECIES : Area antiquata Linne, 1758 (OD).

DIAGNOSIS : The generic systematics of most of the Anadarinae is presently confusing, especially
concerning the distinction between Anadara, Scapharca and Cunearca. The distinctions as detailed by
NEWELL in MOORE (1969) rest on the form of the ligament and on the equality of size of right and left
valves. The ligament is referred to as amphidetic or opisthodetic, however in many species the ligament
initially develops behind the beaks and eventually grows to fill the whole of the dorsal area. The degree of
overlap of the right valve also has an ontogenetic relationship as well as being variable between species of
obvious similarity in general form. These characters therefore are unreliable when used in isolation and
NEWELL'S definitions are consequently too broad. Other genera and subgenera have been revived or
introduced mainly by OLSSON (1961) for panamic forms and by IREDALE (1939) for Australian forms.
OLSSON (1961) uses a wider range of characters yet there is still reliance on some that do have ontogenetic
changes and he does not compare his subgenera with those from the Indo-Pacific. Consequently a few
divisions appear to be stable such as Rasia Gray, 1857; Larkinia Reinhart, 1935 and Caloosarca Olsson,
1961. In other cases such as OLSSON'S (1961) use of Diluvarca Woodring, 1925 there remains a wide
interpretation with some overlap into Anadara and Scapharca forms.

Our conclusion is to use Anadara as a b r o a d generic group with subgeneric designations

reserved for those cases where there is a clear resemblance in overall form to the type species of
the subgenus concerned. There is clearly the need for a revised generic systematics on a world
wide scale but this is beyond the scope of this paper and we refrain from introducing revised
concepts in isolation.
 — 330 -

Anadara geissei (Dunker, 1891)

(Pl. IV, 3A-B, 4A-B, 5, 6 A - B ; fig. 2 4 ; m a p 7)

Area setigera Dunker, 1853 : 45, pi. 9, figs. 16-18 (non Reeve, 1844).
Area (Anomalocardia) dunkeri Kobelt, 1891 : 162, pi. 41, figs. 2-4.
Area {Anomalocardia) geissei Dunker in KOBELT, 1891 : 163-164, pi. 41, figs. 5-6.

TYPE MATERIAL : In SMF Frankfurt.

TYPE LOCALITY : Senegal.

DESCRIPTION

Shell t o 75 m m in length. Slightly inequivalve in juveniles b u t adults more or less

equivalve. N o t inflated, tumidity approximately 1/2 to 1/3 of the length. Inequilateral, beaks in
the anterior third.

FIG. 24. — Internal view of right valve of Anadara geissei ( D u n k e r ) . Angola. (Scale b a r = 10 mm.)

Outline oblong, subrectangular, distinctly longer than high, juveniles distinctly expanded
posteriorly, adults symmetrical. Posterior area indistinct; posterior angle marked in juveniles
but becoming obsolete early in o n t o g e n y ; posterior margin obliquely subtruncate a n d slightly
auriculate in juveniles, becoming simply subtruncate and finally rounded but more acute than
anterior margin. Median area initially weakly sulcate, this noticeable t o about 30 m m , then
becoming progressively obsolete; ventral margin rather long a n d more o r less straight, larger
examples with a narrow anterior ventral byssus gape. Anterior margin broadly rounded.
Dorsal margin rather long, straight.
 — 331 —

Dorsal area long, rather n a r r o w except in the largest specimens, u m b o n a l separation

moderate. Ligament initially opisthodetic but developing anteriorly at a b o u t 6 m m , apparently
amphidetic by 15 m m and remaining with a single lateral chevron bordering a large fibrous
sheet until a b o u t 40 m m when new chevrons are added these totalling 4-5 in the largest
specimens.
Hinge plate rather n a r r o w to m o d e r a t e depending on the degree of encroachment by the
ligament; teeth in two series, separation distinct, anterior set u p to 30 teeth, posterior set u p to
35 teeth. Teeth small, median series more or less straight and vertical; lateral series slightly
chevron shaped, slightly oblique.
Sculpture of 29-32 radial ribs of which u p to 18 anteriorly become progressively bifurcate,
posterior ribs wider and more flat, anterior ribs very weakly ridged by low closely spaced cross
bars.
Periostracum a thick coating of appressed laminae with bristles developing from the
primary interspaces a n d from the bifurcate interspaces; anterior and median bristles short,
lanceolate and soft, posterior bristles coarse, long, rigid, semi-erect; colour a deep reddish
brown except for posterior bristles which are black; juveniles have a distinctive maculate
pattern over the posterior area formed by differential thickening of the periostracum.

X = 3.12

e x = 2.os

L/A-Bk

FIG. 25-26. — 25 : Box plots of the ratios of T o t a l Shell Length to Heigth ( L / H ) , Tumidity (L/T) and Anterior to Beak
Length (L/A-Bk) of Anadara geissei from West Africa (n = 27). 26 : Bar chart of R i b N u m b e r for 22 shells of
Anadara geissei.
 - 332 —

Shell white.
Adductor scars subequal, the posterior a little larger; posterior pedal/byssal retractor
elongate oval. Inner margin deeply crenulate.

SELECTED SHELL MEASUREMENTS : See figure 25.

DISTRIBUTION : This species ranges from the Cape Verde Islands and Mauritania (Cap Blanc) to
southern Angola (Benguela).

MATERIAL EXAMINED : Mauritania : Off Port Etienne, 20°20'N/16°22' W, 10m, 3 v., coll. MARCHE-
MARCHAD, 8.V.1965, MNHN. Cape Verde Islands: Sta. Lucia, 52m, 1 spm. + 1 v., dredged R/V
"Princess Alice" (sta. 1152), coll. H.FISCHER, 1901; off Boa Vista, 15m, 1 spm., dredged R/V
"Calypso", 1959; SW of Boa Vista, 30 m, 2 v., 23.X. 1948; Säo Vicente, Baia Porto Grande, beach, 11 v.,
leg. VON COSEL, 19.XII.1978; Säo Vicente, Mindelo, beach, 3 v., leg. VON COSEL, 17.XII.1978; all MNHN.
Senegal : No precise locality, 1 spm., coll. VAYSSIERE ; Rufisque, 18-20 m, many spm.; Pte. Cansado, 2 v.;
W of Cap Rouge, 6 v., all coll. Mission GRUVEL, IV. 1909; Goree, 1 spm., coll. BAVAY; Goree, 1 spm.,
coll. GERET; Cap Vert, 1 v., coll. MAURY; off Dakar, 50m, 4 v.; Banc de Seminole, 43-45m, 1 v.; off
Goree, 95-98m, many v., all dredged R/V "Gerard Treca", leg. MARCHE-MARCHAD, 1954-1956; Dakar,
Anse Bernard, 1 spm., leg. MARCHE-MARCHAD; SE of Goree, 14°41'N/17°23,2'W, 17m, fine sand and
mud, many v., dredged R/V "Louis Sauger", leg. VON COSEL, III.1988; M'Bour, Petite Cote, on beach,
1 v., leg. VON COSEL, 22.III. 1988 ; S. Casamance, Essoukoudiak Bölon, 4-6 m, shelly sand, 1 v.; Karabane
Bolon, 3-4m, shelly sand, 1 v., both leg. VON COSEL, III.1988; N. Casamance, Abene, 13°03'N/17°03'W,
20m, very fine sand, 1 v.; Kafountine, 12°57,5'N/17°16,8'W, 35m, medium sand and rocks, 1 v.;
12°56,9'N/17°06,8"W, 22m, fine sand, 3 v.; 12°44,5'N/17°27,3'W, 40m, fine sand, 8 v.;
12°32'N/17°28,8'W, 45m, fine sand, 3 v.; off Diembering, 12°29,6' N/17°24,3'W, 35m, fine sand, 1 v., all
dredged R/V "Louis Sauger", leg. VON COSEL, III. 1988, all MNHN. Guinea-Bissau : Ilhas Bissagos, 7 v.,
leg. Mission GAIN, 1913, MNHN. Guinea: 10°30'N/15°43,5' W, 21m, 2 v.; 10° 12' N/13°06' W, 20m,
0
2 spm.; 9°50'N/14°14'W, 15m, 2 spm. + 5 v.; 9°30'N/13°53'W, 19m, 1 spm.; 9 18'N/13°45"W, 21 m,
1 spm.; 9°12'N/14°15,1'W, 33m, 1 spm.; 9°05,7'N/13°38'W, 24m, 1 spm.; all dredged R/V "Andre
Nizery", leg. VON COSEL, IV-V.1988; 9°25'N/13°48,5'W, 1 v., dredged R/V "President Theodore
Tissier"; all MNHN. Sierra Leone: 7°15,5'N/12°51'W, 64m, 1 v., dredged R/V "Calypso", leg.
MARCHE-MARCHAD, 19.V.1956, MNHN. Nigeria : 4°00' N/6°l 1' E, 34m, 3 spm.; 4°03' N/6° 12' E, 32m,
many v., both dredged R/V "Calypso", leg. MARCHE-MARCHAD, 1956, both MNHN. Gabon :
0°14'S/9°14'E, 15-16m, 2 v., dredged R/V "Thierry", Guinean Trawling Survey, leg. CHERBONNIER,
16.XI.1963, MNHN. R.P. Congo: Pointe-Noire, off plage ORSTOM, 3-7 m, 1 spm., 1 v.; plage
ORSTOM, on beach, 1 v., plage Mondaine, on beach, 6 v., all leg. VON COSEL, XI-XII.1985; all MNHN.
Angola : Baia de Corimba, Luanda province, 10-20m, many spm.; Cacuaco, Bengo province, 5-10m,
many spm.; Ponta das Lagostas, 5-20m, 1 spm.; Santo Antonio, Benguela province, intertidal, 1 v., all
leg. GOFAS, 1981-85, all MNHN.

BIOTOPE : Anadara geissei inhabits fine and mixed sand with calcareous algae and shell debris. The
juveniles are byssally attached to hard objects. It is a subtidal species ranging from 3-30 m.

REMARKS

This species was first described by D U N K E R in 1853 as Area setigera but that name is
preoccupied by Area setigera Reeve, 1844. KOBELT (1891 : 162) renamed this A. dunkeri but he
also then described (1891 : 163-164) A. geissei using a manuscript name from D U N K E R . The
two are undoubtedly synonymous, A. geissei representing a young specimen and A. dunkeri an
adult, but as the name A. geissei has been applied in all previous regional faunal studies we
have retained this name.
 1.9.

1.8.

1.7. ~|~X . 1.61

1 X-1.39

1.4.
T
1.3-

1.2.
Angola - L/H Guinea - L/H Cape Verdes - L/H

FIG. 27. — Box plots for Length to Height ratios of three samples of Anadara geissei from Angola, G u i n e a and the
C a p e Verde Islands.

M A H 7. — Distribution of Anadara geissei.

Anadara geissei is not greatly variable t h r o u g h o u t its range but the data displayed in
figure 27 shows that there is variation in the Length/Height ratio, with some examples from
the Cape Verde Islands being much higher and therefore m o r e q u a d r a t e a n d in which the
posterior margin remains subauriculate. The choice of samples used suggests clinal variation
with latitude but it could also be ecophenotypic. The material and ecological d a t a are limited
 — 334 —

and at this time we consider all forms to be nothing more than individual varieties of A.
geissei. It should be noted that the unusual specimens approach the Caribbean A. notabilis
(Roding, 1798), and LOCARD (1898) and LAMY (1907) record, in our view erroneously, this
species from Cape Verde Islands and Senegal.
W e consider that A. geissei, A. floridana and the panamic species A. formosa (Sowerby,
1833) to be closely related and that the west African and Panamic species are most similar. The
subgenus Rasia Gray, 1857 is probably a useful taxon within the Atlantic/Panamic provinces
but we are not certain of its relationship with Indo-Pacific forms. A. geissei is a semi-infaunal
species forming extensive beds reminiscent of Modiolus a n d it is consequently rather different
from the truly infaunal burrowing forms of more typical A n a d a r i n a e . The type species of
Anadara, A. antiquata is similar in being much longer than high and one of us has (OLIVER,
pers. obs.) noted that this species is also semi-infaunal although to a lesser degree. It is
therefore inappropriate at this time to m a k e a decision on the value of the subgenus Rasia
world wide.

Anadara polii (Mayer, 1868)

(Pl. V, 1-9; fig. 3 0 ; m a p 8)

Area antiquata Poli (non Linné, 1758), 1795 : 146, pl. 25, figs. 14-15.
Area weinkauffi Crosse, 1862 : 324-325. Nomen dubium.
Area diluvii auct. (non Lamarck, 1819 : 45).
Area polii Mayer, 1868 : 75.
Area diluvii var. inflato-subglobosa Aradas and Benoit, 1870 : 80.
Area diluvii var. elongato-depressa Aradas and Benoit, 1870 : 80.
Area (Anadara) sphaerica Kobelt, 1891 : 53-54, pi. 15, figs. 3-4.
Area talismani Locard, 1898 : 308, pi. 8, figs. 21-24.
Area polii var. minor Locard, 1898 : 305.
Area polii var. ventricosa Locard, 1898 : 305.
Area polii var. curta Locard, 1898 : 306.
Area polii var. transversa Locard, 1898 : 306.
Area polii var. obliqua Locard, 1898 : 306.
Area polii var. angulosa Locard, 1898 : 306.

TYPE MATERIAL : The type material of Area weinkauffi is uncertain : it was cited by CROSSE as being in
the collection of WEINKAUFF, but later WEINKAUFF stated that it was never in his collection but was a
single specimen in the Algerian Museum which had been stolen and replaced by a monstrosity of Area
diluvii (WEINKAUFF, 1880 : 200). Given this uncertainty we regard A. weinkauffi as a nomen dubium.
Anadara polii was erected by MAYER in order to distinguish the recent Mediterranean species from
the Miocene A. diluvii Lamarck, 1819. His name was based on figures of the recent species in a number of
early publications of which the earliest is that of POLI, 1795. Consequently we regard the figure of A.
antiquata Linné in POLI to be the type.
The holotype of A. sphaerica is in SMF Frankfurt but it bears no locality data.
The type material of the varieties inflato-subglobosa and elongato-depressa are cited as in the
collection of ARADAS and BENOIT but its present location is not known.
The holotype of A. talismani is in MNHN and has a slightly broken hinge plate which has been
misrepresented in the original figure in LOCARD (1898).

TYLE LOCALITY : Sicily.

 — 335 —

GENERAL DESCRIPTION

We have encountered great difficulty in assessing the significance of the large range of
morphological variation seen within this taxon. Individual populations or samples show little
variation but between samples the extremes can be such that it is difficult to imagine that they
are part of a single taxon. M o r e over there is n o apparent pattern to this variation such that
we are unable to definitely invoke clinal, bathymetric or ecological causes to explain it. This
species is restricted to rather deep water and we have mostly small samples of live collected
material with the larger samples of dead valves only. This restricts our ability to assess the
distribution of the variants and also introduces the concern that some of the samples are of
subfossil origin, notably that of the large shells from Senegal.
In conclusion we have refrained from separating the m o r p h s into nominal taxa and
instead we present a general diagnosis which highlights the c o m m o n features and also
individual descriptions of the m o r p h s . W i t h further collection it may be possible to explain this
variation or even to conclude that this group is a complex of closely related species.
Shell to 55 m m in length. R a t h e r heavy. Slightly inequivalve when small, more or
less equivalve in adult, LV a- little larger. Inflated to greatly inflated, tumidity a little less
than or a little greater than height. Inequilateral, beaks in the anterior 1/4 to 1/5. Outline
oblong-subrectangular to suboval to ovate, usually longer than high but some only slightly
so. Length/Height ratio varying from 1 : 1 to 1.3 : 1 (fig. 28). Posterior area weakly defined
by rounded posterior angle; posterior margin oblique, curved to subtruncate. Median area
deeply sulcate in post larval shell, otherwise r o u n d e d ; ventral margin gently to deeply
curved. Anterior area short to severely constricted. Length/Anterior to Beak Length varying
from 2.9-4.7 (fig. 28); anterior margin broadly rounded.
Dorsal area moderately wide to wide. Ligament initially opisthodetic, developing
anteriorly to fill dorsal area, chevrons widely spaced, few in number. Teeth n u m e r o u s in two
series.
Sculpture of 24-28 (these include the small anterior and posterior dorsal riblets when
apparent) radial ribs (fig. 29), these rather narrow, high and equal to or narrower than
interspaces except down posterior angle where they m a y be wider. Rib sculpture discrepant;
LV with closely spaced, rather small raised cross bars, RV similar but much weaker.
Periostracum reddish brown, with bristles in the interspaces, these are all rather soft and
fragile, anterior and median areas with short sharply pointed triangular bristles, posterior
angle with very long, narrow, linear bristles.

MORPH DESCRIPTIONS

Anterior constricted morph (pi. V, 3B, 7, 8, 9) : Live collected shells to 25 m m . Inflated,

tumidity only slightly less to slightly greater than height. Outline obliquely suboval, anteriorly
constricted. Posterior margin oblique, gently curved, posterior ventral junction indistinct,
rounded. Ventral margin curved, inclined towards the anterior and merging without a distinct
junction into the broadly rounded anterior margin. Anterior area very small, sloping very
steeply towards the u m b o . Ligament extending anteriorly but not filling area completely.
 — 336 -

25

§15
O

5.

22 24 26 28 30
29 Number of Ribs

F I G . 2 8 - 2 9 . — 2 8 : Plots of Total Shell Length against Height and Anterior to Beak Length for 7 5 shells of Anadara
polii from t h r o u g h o u t its range. 2 9 : N u m b e r of Ribs for 7 5 shells of Anadara polii from t h r o u g h t its range.

West African "subfossil" globose morph (pi. V, 3C, 4B) : Shells to 40 mm. Similar in
outline to the above but some large shells are subspherical (probably gerontic). Sculpture of
24-28 ribs, usually 25-26 but this may be accounted for by wear of the posterior dorsal area.
Ribs very noticeably narrower than interspaces. Ligament filling most of the dorsal area in
large specimens.
Oblong morph (pi. V, 1A-B, 3A, 4A, 5, 6 ) : Shells to 55 m m in length. Inflated but tumidity
distinctly smaller than height. Outline oblong-subrectangular; posterior and anterior ventral
junctions distinct, especially posterior; ventral margin more or less parallel to dorsal margin.
Sculpture of 26-27 ribs, the posterior a little wider; rib sculpture on LV prominent especially
on small shells.
F I G . 30. — Internal view of right valve and dorsal view of Anaclara polii (Mayer). Anteriorly constricted m o r p h from
C a m e r o o n . (Scale bar = 5 m m . )

DISTRIBUTION : This species is recorded from Portugal (Setubal), throughout the Mediterranean
south to Angola (Mocamedes).

MATERIAL EXAMINED : Mediterranean, Algeria : No precise details, 2 spm. + 1 v., MNHN. Italy :
Sicily, Palermo, 1 spm., leg. MONTEROSATO, 1914; S. Andrea, 2 spm., leg. del Prete, 1928-1929, coll.
Staadt; Circeo-Tirreno, leg. TITTONI, 2.IV.1957, coll. Staadt; Naples, 5 spm., coll. PETIT, 1873; Naples,
1 spm., old coll.; all MNHN. France : St. Raphael, 1 spm., coll. LOCARD, 1892; off Banyuls-sur-Mer,
130-190m, 9 spm.; both MNHN. Atlantic Morocco : between 37°01'N and 30°06'N, 20-180m, numerous
spm., sh. and v. from 25 dredging stations of R/V "Vanneau", 1923-1929; between 36°49'N and
33°43'N, 144-332m, numerous sh. and v. from 6 stations of R/V " C r y o s " , BALGIM, VI. 1984; all
MNHN. Mauritania : SW of Nouakchott, 17°44'N/16°27'W, 100 m, 2 v., dredged "Leon Coursin", leg.
MARCHE-MARCHAD, 3.II.1957, MNHN. Cape Verde Islands: S. Vicente, 1 v.; off Sao Tiago,
15°16,5'N/23°47,5'W, 50-65m, 1 v.; 15°26,5'N/23°14'W, 100m, 1 spm., all dredged R/V "Calypso", leg.
MARCHE-MARCHAD, 1959; all MNHN. Senegal : Off Cap Vert, 14 v.; off Cap Vert 200-300 m, 1 v.; off
Kayar, 110-120m, 12 v., all leg. MARCHE-MARCHAD, 1967; 15°38'N/17°00"W, 130m, 1 v., dredged
"Leon Coursin", leg. MARCHE-MARCHAD, 1.II.1957; 14°50,1'N/17°29,3'W, 150m, many v., dredged
"Tenace", leg. MARCHE-MARCHAD, 15.111.1967; 14"23,5'N/17°24,5'W, 65-70m, 1 spm. + 8 v.; off
Dakar, 50m, 1 spm.; 200m, 1 spm.; 129-150 m, 9 v.; S of Goree, 110-112m, 13 v., all dredged R/V
"Gerard Treca", leg. MARCHE-MARCHAD, 1954-1958; Casamance, 12°32'N/28°8'W, 45m, fine sand,
2 v., dredged R/V "Louis Sauger", leg. VON COSEL, 28.111.1988; 14° 15,8' N/17°28' W, 100m, leg. LEUNG
TACK; Thiaroye, NNE of Dakar, 7-8m, leg. MARCHE-MARCHAD, 6.VI.1956; all MNHN. Liberia:
4°34,5'N/8°31'W, 64m, 2 v.; 5°21,5'N/9°54,5'W, 73-80m, 13 v., both dredged R/V "Calypso", leg.
MARCHE-MARCHAD, 20.V.1956, all MNHN. Cote d'lvoire : Off Abidjan, 1 spm. + 9 v.; off Abidjan,
60 m, 1 spm. ; off Abidjan, 70 m, 4 spm., all leg. LE LOEUFF (ORSTOM); off Abidjan, 2 v., leg. MARCHE-
MARCHAD; 4°50'N/5°57'W, 70m, 7 v.; 5°01'N/5°17"W, 40m, 1 spm., both dredged R/V " L a Rafale",
Guinean Trawling Survey, IV. 1964; all MNHN. Nigeria : 4°00'N/6°H'E, 2 v., dredged R/V "Calypso",
leg. MARCHE-MARCHAD, 1956, MNHN. Cameroon: 3°54,5'N/8°53'E, 62-64m, mud, 1 spm.;
3°54' N/8°50' E, 65-70 m, mud, 1 spm.; 2*51* N/9°41' E, 80 m, mud, 1 spm.; 2°39' N/9°40' E, 60-65 m, mud,
2 spm., all leg. CROSNIER, 22.VIII.1963; all MNHN. Gabon : Cap Lopez, on beach, 1 v., leg. CHEVALIER,
1984-1988; 0°32,8'S/8°43,1'E, 125-145m, 1 v., dredged R/V "Calypso", leg. MARCHE-MARCHAD, 1956,
MNHN. Angola : Off Ambrizete, Zaire province, 80m, 4 v.; off Mussulo, Luanda province, 90-100m,
muddy ground, 12 v.; Ilha de Luanda, Luanda province, 120 m, many v.; Ilha de Luanda, 40-60 m, 2 v,;
Ilha de Luanda, 90m, 1 v.; Praia Amelia, Mocamedes province, 40-60m, 1 v., all leg. GOFAS, 1981-1984;
all MNHN.

BIOTOPE : This species is restricted to offshore in depths from 50-200 m.

 — 338 —

MAP 8. — Distribution of Anadara polii.

REMARKS

Of the m o r p h s described the most c o m m o n in tropical West Africa is the " anteriorly
constricted " type. In West Africa living specimens are as yet all smaller t h a n 25 m m which is
half the size of those living in the Mediterranean. Considering the apparently subfossil samples
the size ranges are more similar with a valve from G a b o n reaching 4 0 mm. The most c o m m o n
type in the Mediterranean is the " o b l o n g " m o r p h but we have only 8 specimens and 1 valve
from West Africa. N o w h e r e are the two extremes mutually exclusive and we c a n n o t suggest
that there is clinal variation. The globose m o r p h represents m a n y of the shells collected by the
R / V " V a n n e a u " from off M o r o c c o and is only rarely found in the Mediterranean. The large
examples closely resemble the gerontic specimens of the subfossil samples from Senegal and
 — 339 -

such specimens are the basis of both LOCARD'S A. talismani and probably KOBELT'S A.
sphaerica.
The various m o r p h s are shown to overlap in their distribution and there is a merging of
one into the other when all samples are observed collectively. This degree of overlap can be
seen in figures 31-32 which display the ranges of Length/Height and Length/Anterior/Beak
Length ratios for shells from three parts of the range. These plots also show that the range of
variation is greatest in the tropical West African shells. There is therefore no value in giving
the m o r p h s even varietal status and all are considered to be variants of A. polii.

Medrterranean

FIG. 31-32. — 31 : Box plots of Shell Length to Height ratios of shells of Anadara polii from three parts of its range,
Mediterranean, M o r o c c o and tropical West Africa. 32 : Box plots of Length to Anterior to Beak Length of shells
of Anadara polii from three parts of its range, M e d i t e r r a n e a n , M o r o c c o a n d tropical West Africa.

We have chosen to use the name A. polii Mayer for this species and not the more often
used A. diluvii Lamarck. We d o not consider that from our examination of the types of A.
diluvii this species to be the Miocene/Pliocene counterpart of the Recent form because (i) the
rib number is 33 not 24-28, (ii) the beaks are situated much more closely to the midpoint,
(iii) the ribs are equal in width to the interspaces.
 - 340 -

Anadara eborensis n. sp.

(Pl. VI, 1 A - D ; fig. 3 3 ; m a p 9)

TYPE MATERIAL : Holotype (31.9 mm), MNHN, off Grand Bassam, Côte d'Ivoire, 5°07' N/3°22' W,
20m, dredged R/V " L a Rafale", Guiñean Trawling Survey, leg. CHERBONNIER, 21.III.1964. Paratypes :
MNHN, off Grand Lahou, Côte d'Ivoire, 5°00' N/5°28.5' W, 27 m, 1 spm., dredged R/V "Calypso", leg.
MARCHE-MARCHAD, 1956; NMWZ, Guinea, 9°36'N/14°12'W, 21m, 1 spm., dredged R/V "André
Nizery". leg. VON COSEL, 19.V.1988.

TYPE LOCALITY : Côte d'Ivoire, off Grand Bassam.

DESCRIPTION

Shell to 32 m m in length. Heavy. Slightly inequivalve, LV a little larger. Inflated, almost

as tumid as high, strongly u m b o n a t e . Inequilateral, beaks almost in the anterior quarter.
Outline subquadrate, only slightly longer than high. Posterior area broad sloping steeply,
set off by strong but rounded posterior angle; posterior margin obliquely subtruncate, more or
less straight; posterior ventral junction distinct, sharply rounded. Median area narrow, more
or less flat except in very earliest stages in which it is deeply sulcate, ventral margin gently
curved or more or less straight. Anterior margin broadly rounded. Dorsal margin straight,
anterior junction almost 90°, posterior junction oblique.
Dorsal area broad, slightly cleft or more or less flat, umbonal separation wide. Ligament
initially developing with a single posterior chevron, then adding anteriorly towards the beaks
and eventually in front of the beaks but chevron number remaining low with u p to 5 growth
points on the hinge line, chevrons narrow.
Hinge plate moderately thick, teeth in two series, separation slight with posterior set
pushing over the anterior, anterior set u p to 24 teeth, posterior set u p to 27 teeth. Teeth
laminar, narrow, vertical.
Sculpture of 24-27 (usually 25) radial ribs, these narrow and only half the width of the
interspaces, steeply angled with narrower, rather flat t o p ; RV weakly nodulose, LV almost
smooth.
Periostracum rust brown with slightly darker, short, t h o r n like, recurving bristles arising
from the interspaces of the anterior and median a r e a s ; those on the posterior angle all worn.
Shell white.
Adductor muscle scars unequal, the posterior considerably larger. Internal margin deeply
crenulate.

SELECTED SHELL MEASUREMENTS : See in Remarks section under Anadara subglobosa.

DISTRIBUTION : This species has a restricted distribution from Guinea to Cote d'lvoire and Gabon.

MATERIAL EXAMINED : The type material; Guinea : 9°40'N/14°05'W, 18m, many v., dredged R/V
"Calypso", leg. MARCHE-MARCHAD, 1956; 9°12'N/13°43,5'W, 24m, 1 spm.; 9°01'N/13°30'W, 12m,
1 spm., both dredged R/V "Andre Nizery", leg. VON COSEL, 27.IX.1988; all MNHN. Liberia:
 — 341 —

4°34,5'N/8°31'W, 64m, 1 v., dredged R/V "Calypso", leg. MARCHE-MARCHAD, 1956, MNHN. Cote
o
d'lvoire : 5°03'N/5 25'W, 20-25m, 1 v., dredged R/V "Calypso", leg. MARCHE-MARCHAD, 21.V. 1956;
off Abidjan, 2 spm., dredged R/V " Reine Pokou ", leg. LE LOEUFF (ORSTOM); both MNHN. Nigeria :
4°34,5'N/8°31'E, 64m, 1 v.; 4°03'N/6° 12'E, 32m, 1 v., both dredged R/V "Calypso", leg. MARCHE-
MARCHAD, 1956, MNHN. Gabon : Cap Lopez, 3 v., leg. CHEVALIER, 1984-1988, MNHN.

BIOTOPE : This species has been dredged from mud and mixed muddy substrates offshores from 18 to
64 m.

DERIVATIO NOMINIS : eborensis, Latin, derived from eboris, "ivory" with the suffix ensis denoting
place referring to the type locality of the Còte d'lvoire.

REMARKS : The heavy subquadrate appearance of this shell is reminiscent of other species
placed in the subgenus Larkinia but those species have a true amphidetic ligament. The only
Caribbean species with any similarity is Anadara brasiliana but it also has a truly amphidetic
ligament, the ribs are as b r o a d as the interspaces and the rib sculpture is more prominent.
Most akin to A. eborensis is perhaps A. (Esmerarcd) reinharti Lowe, 1935, both in the general
shape, the form of the ligament and the weak discrepant rib sculpture. The posterior margin of
A. reinharti by comparison is rounded, there is a slight posterior ventral sinuation and the ribs
are as wide or wider than the interspaces.
FIG. 33. — Internal view of right valve a n d dorsal view of Anadara eborensis n. sp. Cote d'lvoire. (Scale b a r = 5 mm.)

Anadara subglobosa (Dunker, 1891)

(PI. VI, 2 A - B ; fig. 3 4 ; m a p 10)

Area (Anomalocardia) subglobosa Dunker in KOBELT, 1891 : 99, pi. 26, figs 7-8.

TYPE MATERIAL : Most probably in ZMB Berlin. In the MNHN we have located a single specimen in
the BAVAY collection which is marked as coming from VON MALTZAN, collected at Goree, Senegal. This
agrees with the data available with the type as cited by DUNKER and also agrees with his figure (1891 :
pi. 26, fi.g. 8).

TYPE LOCALITY : Goree, Senegal.

DESCRIPTION

Shell to 19 m m in length. Slightly inequivalve in juveniles, more or less equivalve in adults.

Inflated to strongly inflated, usually with tumidity less than height but some adults with
tumidity greater than height, u m b o n a t e but n o t greatly. Inequilateral, beaks in the anterior 1/3
to 1/4.
Outline subrectangular, adults always distinctly longer than high with the anterior half of
the shell a little more expanded than the posterior, juveniles less so. Posterior area relatively
short, sloping rather steeply a n d demarcated by a distinct, acutely rounded posterior angle but
this is n o t so m a r k e d in the juveniles; posterior margin oblique, moderately long, more or less
straight; posterior ventral junction distinct, roundly acute. Median area strongly sulcate in
post larval shell a n d to a lesser degree in juveniles to a b o u t 8 m m , otherwise gently curved;
ventral margin gently curved, subparallel to dorsal margin. Anterior area small, anterior
margin broadly rounded, anterior ventral junction obsolete. Dorsal margin straight.
Dorsal area rather wide, more so anterior to the beaks, u m b o n a l separation m o d e r a t e to
wide. Ligament initially opisthodetic but very rapidly developing anteriorly a n d appearing
 — 343 -

secondarily amphidetic except for a small anterior ligament free a r e a ; chevrons very few, 2-3
in number, very narrow.

FIG. 34. — Internal view of right valve a n d dorsal view of Anadara subglobosa ( D u n k e r ) . M a u r i t a n i a . (Scale b a r =
5 mm.)

Hinge plate moderately thick; teeth in 2 series, the anterior set pushing below the
posterior at their j u n c t i o n ; anterior set u p to 18 teeth, posterior set u p to 24 teeth. Teeth
laminar, all teeth vertical except for a few slightly oblique posterior marginals.
Sculpture of 28-31 radial ribs, these narrower than interspaces in adults with steeply
angled sides a n d a rather flat topped appearance. Rib sculpture discrepant, stronger on LV of
rather long, sparse nodules, R V more or less smooth except for weak nodules on anterior ribs.
Interspace sculpture distinct of widely spaced concentric, crescentric notches.
Periostracum rarely preserved in a good s t a t e ; generally rust brown in colour with sparse
bristles in the interspaces; anterior a n d median bristles thorn-like recurved dorsally; on
posterior angle longer, narrower.
Shell white.
A d d u c t o r scars unequal, the posterior twice the size of anterior, posterior q u a d r a t e ;
anterior rounded, often a little raised. Inner margin deeply crenulate.

SELECTED SHELL MEASUREMENTS : See below in Remarks section.

DISTRIBUTION : From Mauritania (17°N) south to Cote d'lvoire.

MATERIAL EXAMINED : Mauritania : 17°30'N/16°15'W, 41m, 1 spm., dredged R/V " N ' D i a g o " ,
leg. RICHER DE FORGES, 1981, MNHN. Senegal : Goree, 1 spm., coll. BAVAY ex coll. VON MALTZAN;
Goree, 1 spm., coll. JOUSSEAUME; SE of Goree, 14°41'N/17°23,2'W, 17m, fine sand and mud, 2 v.,
dredged R/V "Louis Sauger", leg. VON COSEL, 24.111.1988; S of Goree, 65m, many v.; 110-112m,
1 v.; 95-98m, many v.; SW of Cap Manuel, 50m, 1 v., all dredged "Gerard Treca", leg. MARCHE-
MARCHAD, 11.1954; 14°50'N/17°29'W, 150m, dredged "Tenace", leg. MARCHE-MARCHAD, 15.III.I967;
14°31,9'N/17°13,5'W, 28m, 1 spm., coll. LEUNG TACK, 1983-1984; Casamance, Abene, 13°01,8'N/
17°25,5'W, 53m, hard ground and fine sand, 1 v.; 12°46,6'N/17°19,2'W, 32m, carbonate and fine
sand, 1 v., all dredged R/V "Louis Sauger", leg. VON COSEL, 28-29.111.1988; 14°2,5'N/17°09,rW,
24m, 2 v., dredged "Cdt. Henri Gomis", leg. BODARD, 7.XII.1966; 13°01'N/17°24'W, 51-55m,
1 spm. + 2 v., dredged, R/V "Calypso", leg. MARCHE-MARCHAD, 1956; all MNHN. Ghana :
4°57'N/2°42'W, 40m, muddy ground, 1 spm. -I- 1 v., dredged R/V " L a Rafale" Guinean Trawling
 344 -

Survey, leg. CHERBONNIER, 19.III.1964; MNHN. Cote d'lvoire : 4°52,5'N/5°57,5' W, 40m, hard ground
and calcareous algae, 2 v.; 5°05'N/3°22'W, 30 m, mud, sand and shell gravel, 2 spm. + 4 v.;
5°07'N/3"22"W, 20m, 1 spm.; all dredged R/V " L a Rafale", Guinean Trawling Survey, leg. CHER­
BONNIER, III and IV. 1964; all MNHN.

BIOTOPE : This species has been collected primarily in deep water from 17-200 m.

REMARKS

This species is clearly related to Anadara eborensis (this paper) by the similarity in growth
of the ligament a n d the shape a n d sculpture of the radial ribs. It differs from that species in
being rectangular rather than quadrate, that is with a larger Length/Height ratio (fig. 3 5 ) ; a n d
also less tumid (fig. 35).
Although there is a degree of overlap with these characters the rib number is consistently
different between the two species A. subglobosa having more ribs, 28-31 as against 24-27 for A.
eborensis (fig. 36).

1
• jr. -'- CO

o
CM

0
o

% o
o

|20° lo-

M A P 10. — Distribution of Anadara subglobosa (circles) a n d A. camerunensis (triangles).

The distribution of these species overlap, a n d we have not found intermediate samples in
the material at hand. It is of interest to note that it does seem possible to distinguish non
overlapping morphologies in some groups yet not in others. T h e degree of plasticity in shell
form within the Anadara group has never been explained but is well illustrated by referring to
 - 345 —

any of the major works on the Tertiary faunas of E u r o p e or the Caribbean where a multitude
of forms are described. Until we have understanding of this plasticity the species systematics
will remain fraught with difficulty so long as we are limited to traditional conchological
methods. With the Recent fauna recourse to electrophoretic and genetic methodology could be
advantageous.

1.6
Anadara eborensis
B
Anadara subghbosa
1.34

X - 1.21
X= 1.17°

35 Length : Height Length : Tumidity

I I Anadara subglobosa
[ | Anadara eborensis

20 22 24 28 30
36 Rib Number

F I G . 35-36. — 35 : Box plots of Length to Height and Length to Tumidity ratios for Anadara eborensis (n = 1 4 ) and
A. subglobosa (n = 20). 36 : Bar chart of rib n u m b e r in Anadara eborensis (mode 25) and A. subglobosa (mode 3 1 ) .

Anadara senegalensis (Gmelin, 1791)

(PI. VI, 3A-D, 4A-B, 5 A - B ; pi. VII, 1-4; fig. 3 9 ; m a p 11)

"Le Robet" Adanson, 1757 : pi. 2 4 8 , fig. 6.

Area senegalensis Gmelin, 1791 : 3 3 1 2 .
Area pertusa Reeve, 1844 : Sp. 2 8 , pi. V .

TYPE MATERIAL : Lectotype (here selected), MNHN, from coll. ADANSON, a left valve, 2 0 mm in
length. This is not the valve cited by FISCHER-PIETTE (1942) as the figured specimen in ADANSON (1757)
but we argue that the original figure is so poor that it does not adequately represent any of the specimens
labelled by ADANSON as " Le Robet". Furthermore the two valves cited by FISCHER-PIETTE are not a pair,
 - 346 —

the right valve does not fit into the left valve and the widths of the dorsal areas are different.
Paralectotype as lectotype, 2 valves.

TYPE LOCALITY : Senegal.

DESCRIPTION

Shell to 25 m m in length. Inequivalve, LV larger than RV. Greatly inflated, tumidity

almost equal to or a little greater than height; strongly u m b o n a t e . Inequilateral, beaks in the
anterior third.
Outline rhomboidal-oval, initially a little longer t h a n high, later almost as high as long.
Posterior area very short and very steep; posterior angle distinct but r o u n d e d ; posterior
margin long, a little oblique, almost vertical, gently curving in juveniles but with a subtle
angulation in adults. Median area curved except in post larval shells which are sulcate; ventral
margin curved, posterior ventral junction slightly angulate, anterior ventral junction not
marked but curving into anterior margin which is short, weakly curved and almost vertical.
Dorsal margin straight, rather short.
Dorsal area slightly cleft, rather n a r r o w , u m b o n a l separation slight. Ligament initially
opisthodetic, later apparently amphidetic but with a somewhat larger anterior ligament free
area, chevrons widely spaced, numbering u p to 4.
Hinge plate thick, teeth in two series, separation slight, anterior set u p to 22 teeth,
posterior to 20 teeth. Teeth laminar, some weakly chevron shaped, all more or less vertical.
Sculpture discrepant, of 23-24 radial ribs, LV with anterior 15-16 ribs strongly nodulose
formed by thickened crossbars, R V with anterior 6-10 ribs more weakly nodulose.
Periostracum a thin rust brown coating except for spicate bristles arising from interspaces,
on posterior area of closely spaced, thin, long, black, erect bristles; on median area of shorter
wider bristles directed posteriorly.
Shell white.
A d d u c t o r scars unequal, posterior twice as large as anterior, pedal retractor scars small
indistinct. Inner margin strongly crenulate corresponding to radial sculpture.

SELECTED SHELL MEASUREMENTS : See figure 37.

Casamance variety (pi. VII, 3-4) : This larger (to 29 m m ) variant differs from the typical
form in being distinctly longer than high a n d is therefore oblong-rhomboidal in outline. The
sculpture is slightly weaker but the nodules are still large if not so raised. The shell is thinner
and consequently the hinge plate is weaker and the teeth smaller. This is the form represented
by ADANSON'S type specimens.

DISTRIBUTION : This species has been recorded from Guinea south to Angola (Bengo province).

MATERIAL EXAMINED : Senegal : No precise locality, 4 spm., coll. H. FISCHER; Dakar, 2 v., coll.
CHEVALIER, 1900; N. Casamance, Kafountine, beach to the south, 4 v., Abene-Kafountine, beach, many
v.; S. Casamance, Cap Skirring-Diembering, on beach, many v.; Cap Skirring, beach to the south, many
v.; Bolon opposite Elinkine, 3 m, 8 v.; off Kafountine, 3-6 m, 1 v.; Katakalous Bolon, 2-4 m, 1 v.; S of
Cap Skirring, 3-5 m, many v., all leg. VON COSEL, 5-17.III. 1988; off Cap Skirring, 12°22,5'N/17°03'W,
 — 347 -

20m, 2 v.; off Cap Skirring, 12°23'N/16"52,8"W, 13m, 5 v.; 12°46,9'N/17°29,9'W, 45m, many spm., all
dredged R/V "Louis Sauger", leg. VON COSEL, 27-29.III. 1988, all MNHN. Guinea-Bissau : Casamance
border, Essoukoudiak Bolon, 5-6m, 2 v., leg. VON COSEL, 7.III.1988; 11°58,5'N/16°59"W, 14m, 3 spm.;
0 0 o o
ll 53'N/16 56,5'W, 11m, 2 spm.; 12 05'N/17 02,5'W, 14m, many spm., all dredged R/V "Andre
Nizery", leg. VON COSEL, 7.X.1988; all MNHN. Guinea : No precise locality, 2 spm., coll. PARFAIT, 1889;
Cap Varella, 3 v., coll. MAUNY, 1964; Conakry, many v., coll. DUFOSSE, 1905; all MNHN. Cote d'lvoire :
Off Abidjan, 1 spm., leg. MARCHE-MARCHAD ; Sassandra, Batebre beach, many v., coll. AUBERT DE LA
RUE, 1950; off Abidjan, 4°50'N/5°57'W, 70m, 8 v., dredged R/V " L a Rafale", Guinean Trawling
Survey, leg. CHERBONNIER, 4.IV.1964; off Bassam, 20m, 1 spm.; Abidjan region, 20-26m, 27 spm. +
1 sh. from 4 stations, all dredged R/V "Reine Pokou", leg. LE LOEUFF (ORSTOM), 1966-1967; all
MNHN. Togo : Lome, 14 v., MNHN. Gabon : No precise locality, 1 sh., coll. H. FISCHER; Cap Lopez,
beach south of lighthouse point, 6 v., leg. CHEVALIER, 1984-1988, both MNHN. R.P. Congo : Off
Conkouati, 4°00'S/10°59'E, 19m, many v.; 4°10'S/l 1°15'E, 19m, 1 spm., both trawled " K o u n d a " , leg.
VON COSEL, XII. 1985; Pointe-Noire, Plage Mondaine, many v.; Plage Raffmerie, 6 v., Plage ORSTOM,
2 v.; Baie de Pointe Noire, off Plage ORSTOM, 5-7m, many spm., dredged from muddy fine sand; 3-
4 m, 5 spm. + many v., dredged from fine sand and gravel; off Plage Mondaine, 5 m, 3 v., dredged from
muddy fine sand; off Songolo, 5-6m, 1 v., dredged from muddy fine sand, off Plage Raffmerie, 1 m, 3 v.,
all leg. VON COSEL, X-XII.1985; all MNHN. Angola : Cacuaco, Bengo province, 0-10m, 1 spm. +
5 sh. + 6 v.; Mocamedes, Mocamedes province, 0-2m, 1 v.; Ponta do Mussulo, Luanda province, on
beach, 1 v., all leg. GOFAS, 1981-1985, all MNHN.
3.25.
3.
2.75.
2.5. X-2.81 e
2.25.
I 2,
C
1.75,
1.5
1.25

L/H L/A-Bk
37
1.7
I'-'^l Diembering
1.6
1 .5
|<sl Cap Skirring
I ] Abene 1
<o 1-4 1 1
| 1.3

1 .2
T
1.1
T T
1 T T
o

38 Length : Height Length : Tumidity

FIG. 37-38. — 37 : Box plots of the ratios of T o t a l Shell Length to Height ( L / H ) , Tumidity (L/T) and Anterior to Beak
Length (L/A-Bk) of Anadara senegalensis typical form from Guinea-Bissau, C o t e d'lvoire and R.P. C o n g o (n =
59). 38 : Box plots of the ratios of Total Shell Length to Height and Tumidity of three samples of Anadara
senegalensis from the C a s a m a n c e region of West Africa (n = 20 per sample).
 — 348 —

BIOTOPE : Live collected material of this species has been collected in shallow depths from the
sublittoral to 30 m.

REMARKS

This is the most frequently encountered Anadara in the West African material examined,
and for most of its range it is well defined and not variable. There are, however, a number of
samples principally from the Casamance district of Senegal but also some from the R.P.
C o n g o and the northern provinces of Angola that include a rather different form. This form is
described above as the C a s a m a n c e variant.
We are fortunate to have a few large samples of beached valves from Casamance
(map 12) and the Length/Height and Length/Tumidity ratios are plotted in figure 38. This
shows that the sample (Abene) from the northern part, close to the G a m b i a River, contains
more " Casamance " variants t h a n those from further south, south of the Casamance River,
but that the two forms are not mutually exclusive. T h e fact that the « Casamance » variant is
 — 349 -

also to be found a r o u n d the C o n g o river indicates that this may not be clinal variation but is
perhaps linked to environmental conditions close to the estuaries of large rivers. M u c h more
detailed collecting would be needed to assess whether or not the " Casamance " variant is truly
linked to these ecological conditions.

One factor ANOVA table for Length to Height ratios of three samples of Anadara senegalensis
from the Casamance region.
One factor ANOVA, 2 degrees of freedom, F = 15.934, p = .001.

Group Count Mean Std. Dev

Dimbering 30 1.149 .067
Cap Skirring 30 1.17 .086
Abene 22 1.273 .095

Comparison Mean Diff. Scheffe F-test

Diembering vs Cap Skirring - .021 .51
Diembering vs Abene -.125 14.56*
Cap Skirring vs Abene -.103 9.979*

* Significant at 9 5 % .

F I G . 39. — Internal view of right valve and dorsal view of Anadara senegalensis (Gmelin). Typical adult shell from
Cote dTvoire. (Scale bar = 5 mm.)

The A N O V A table for the Length to Height ratios shows a statistically significant
difference between the Abene sample (from the N o r t h ) and those from C a p Skirring and
Diembering (from the South). However over the total range the Casamance variant and the
typical form grade into each other. The use of such analyses for the separation of taxa can be
misleading and in general for Anadara species other non continuous variables such as rib
number should always be considered.
Anadara senegalensis would from its general form be considered to belong the, subgenus
Cunearca but it differs in the form of the ligament which has many chevrons and is secondarily
 - 350 —

amphidetic. The ligament of the type species of Cunearca {Area incongrua Say, 1822) consists
of an amphidetic sheet of fibrous ligament with a single outer lamellar chevron. This form of
ligament is also found in Anadara chemnitzi Philippi, which is the closest Caribbean
counterpart to A. senegalensis, and in Anadara pilula, a similar Indo-Pacific species. The
significance of such differences in ligament structure have never been considered within the
Anadariinae, and it is not known whether it should be given any phylogenetic credence.

Anadara camerunensis n. sp.

(PI. V, 6 A - C ; fig. 4 3 ; m a p 10)

TYPE MATERIAL : Holotype (15.1mm), MNHN, Cameroon, Victoria (now Limbe)-Bota, 8-10 m, leg.
VON COSEL, 4.XII.1985. Paratypes : as holotype, 3 spm.; NMWZ, Cameroon, Victoria-Bota, 5-6 m, leg.
VON COSEL, 4.XII.1985, 2 spm.

TYPE LOCALITY : Cameroon, Victoria (now Limbe).

DESCRIPTION

Shell to 16.4 m m in height. Inequivalve, LV larger than RV. Greatly inflated, tumidity
almost as great as length; strongly u m b o n a t e . Inequilateral, beaks in the anterior third.
Outline obliquely-oval, subrhomboidal, higher than long. Posterior area short, very
steeply sloping; posterior angle prominent but r o u n d e d ; posterior margin very long, slightly
oblique, very gently curved. Median area rounded except in post larval shells which are
sulcate; ventral margin curving into anterior margin which is broadly rounded, posterior
ventral junction roundly angulate. Dorsal margin straight.
Dorsal area slightly cleft, wider in front of beaks, u m b o n a l separation moderate.
Ligament initially opisthodetic, later apparently amphidetic but ligament free area a little large
anteriorly, chevrons few, widely spaced, numbering u p to 3.
Hinge plate thick, teeth in two series, separation indistinct, anterior set u p to 17 teeth,
posterior set u p to 17 teeth. Teeth laminar or slightly chevron shaped, more or less vertical.
Sculpture of 21-22 radial ribs, discrepantly n o d u l o s e ; R V with the anterior 9-11 ribs
nodulose, LV with anterior 13-14 ribs strongly nodulose, all nodules as raised cross bars across
whole of rib.
Periostracum a thin coating, dark rust brown in colour except for interspaces which bear
erect bristles, on the posterior area these are very long rigid black hairs, elsewhere much
shorter and wider.
Shell white.
A d d u c t o r scars unequal, posterior twice the size of the anterior, pedal retractor scars
indistinct. Inner margin deeply crenulate corresponding to radial ribs.
 — 351 —

SELECTED SHELL MEASUREMENTS

Length Height Tumidity Anterior to Hinge line

(mm) (mm) (mm) beak to ventral Rib No. Locality
(mm) (mm)

15.1 16.4 14.9 5.4 12.3 [9/13] 21 Cameroon, 8-10 m

12.6 13.2 12.6 4.2 9.8 [11/14] 22
12.4 13.2 11.7 4.0 10.3 [10/14] 22
11.6 12.0 10.1 3.8 9.1 [10/14] 22
11.1 12.7 11.3 3.9 9.7 21 Cameroon, 24 m
12.4 13.2 12.2 3.7 10.6 21 Cameroon, 5-6 m

(The n u m b e r s in square brackets m a r k the nodulose ribs in RV and LV.)

1.5,

Abene Diembering Abidjan

O Anadara camerunensis
• • o •
• Anadara senegalensis typical •
• • • • • • •
• 5 -n [~1 • c • •
• •
DC i • • • •
• •
•
o o •
O o

o
14 16 18 22 24
41 Shell Length mm

F I G . 4 0 - 4 1 . — 4 0 : Box plots of Length to Height ratios for samples of Anadara senegalensis [Abene (n = 2 0 ) ,
Diembering (n = 2 0 ) , Abidjan (n = 2 0 ) a n d C o n g o (n = 2 3 ) ] and A. camerunensis [ C a m e r o o n (n = 6 ) ] . 4 1 :
G r a p h of Length to Height ratios plotted against Shell Length for Anadara senegalensis (n = 1 3 0 ) and A.
camerunensis (n = 6 ) . Overlapping points are shown as increasing symbol sizes.
 - 352 —

DISTRIBUTION : This species is restricted to the coast of Cameroon.

MATERIAL EXAMINED : Cameroon : Victoria/Limbe-Bota, 8-10 m, 5 v., leg. VON COSEL; off Victoria,
3°44'N/9°11'E, 24 m, 1 spm; between Wouri estuary and Cap Nachtigal, 3°44'N/9°22'E, 13 m, 1 v., both
trawled "Campo Star", leg. VON COSEL, X I I . 1985; all MNHN.

BIOTOPE : This species is infaunal and inhabits muddy fine sand from 5 to 24 m.

DERIVATIO NOMINIS : camerunensis after the Republic of Cameroon, the type locality.

REMARKS

We have considered at length that this species might be an ecomorph of Anadara

senegalensis, however we can find no environmental parameters that would give rise to the
changes observed. In A. camerunensis the adult size is only 1 6 m m as compared to 3 5 m m for
A. senegalensis. F o r most of its ontogeny A. camerunensis is higher than long which is the case
for only the occasional specimens of A. senegalensis (fig. 40-41).
The rib number, although n o t greatly different, is totally consistent : that of A.
camerunensis from 21-22 a n d for A. senegalensis 23-24 (fig. 42).

• Anadara senegalensis

Q Anadara camerunensis

20 40 60
Percentile

F I G . 4 2 . — Rib N u m b e r frequency d a t a for Anadara senegalensis (n 130) and A. camerunensis (n = 6) shown as

percentile line plots.

Comparing the adults of each, A. camerunensis is more oblique, the posterior ventral
margin being more attenuated. Overlaps with A. senegalensis have not yet been found, a n d we
have consequently given this form specific status.
Anadara camerunensis has its closests counterpart in the Panamic province, A. nux
(Sowerby, 1833), but differs in the form of the ligament. In A. nux there is a single outer
chevron (fig. 43C) bounding a large median sheet, a n d the growth is amphidetic. This type of
ligament is found in none of the West African A n a d a r i n a e included in this paper but is not
u n c o m m o n in the Panamic, Caribbean a n d Indo-Pacific. The shape of the shell of A. nux a n d
A. camerunensis would place them in Cunearca. However, should the adaptive form of the shell
 — 353 —

be given greater phyletic weight than the form of the ligament? This problem requires a
complete cladistic analysis of the A n a d a r i n a e and highlights t h e inconsistency in the current
generic systematics.

FIG. 4 3 . — A-B, Internal view of right valve and dorsal view of Anadara camerunensis n. sp. C a m e r o o n : C . Dorsal
view of Anadara nux (Sowerby). Ecuador. (Scale b a r = 3 m m . )

Anadara corbuloides ( M o n t e r o s a t o , 1878)

(PI. VII, 5 A - D ; fig. 4 6 ; m a p 12)

Area corbuloides Monterosato, 1878 : 67.

TYPE MATERIAL : Not located.

TYPE LOCALITY : Algeria.

DESCRIPTION

Shell t o 70 m m in length. Inequivalve, LV larger than R V , most noticeable in small

specimens, obsolete in the larger examples. Inflated, tumidity normally a little greater than half
the length b u t in gerontic specimens much greater. Inequilateral, beaks in the anterior half.
Outline subelliptical, longer than high. Posterior area poorly defined; posterior angle
rapidly becoming obsolete; posterior margin evenly rounded except for a small indentation at
the posterior dorsal junction. Median area rounded b u t sulcate in post larval shell; ventral
margin curved lacking angles with anterior and posterior margins. Anterior margin broadly
rounded to obliquely curved, some becoming rather straight resulting in a subacute anterior
dorsal junction. Dorsal margin relatively long, straight.
Dorsal area becoming wide, slightly cleft, umbonal separation moderate. Ligament
initially opisthodetic becoming secondarily amphidetic; chevrons few, widely separated,
numbering u p to 5.
Hinge plate rather narrow, teeth in t w o series, separation indistinct, anterior set u p t o
 — 354 —

37 teeth, posterior to 39 teeth. Teeth small, pointed, larger teeth slightly chevron shaped, most
vertical, a few lateral teeth a little oblique.
Sculpture of 30-35 radial r i b s ; rib sculpture discrepant, R V with a few anterior ribs with
very weak nodules or ridges, LV with all but posterior area ribs distinctly but not strongly
nodulose.
Periostracum a thin coating, rust to olivaceous brown in colour except for interspaces
which bear umbonally directed rigid spicate bristles.
Shell white.
Adductors scars subequal. The posterior a little larger. Inner margin deeply crenulate.

SELECTED SHELL MEASUREMENTS : See figure 44

4.5

4.

Mode = 33

24 26 28 32 34
45 Rib Number

FIG. 44-45. — 44 : Box plots of the ratios of T o t a l Shell Length to Heigth ( L / H ) , Tumidity (L/T) and Anterior to Beak
Length (L/A-Bk) of Anadara corbuloides (n = 1 2 ) . 45 : Bar chart of R i b N u m b e r for 1 2 shells of Anadara
corbuloides.

DISTRIBUTION : This species ranges throughout the Mediterranean and from Spain (Cadiz, HIDALGO,
1916) along the West African coast and south to northern Angola (Luanda), but is nowhere common.

MATERIAL EXAMINED : Algeria : No precise locality, 1 spm., leg. JOLY, 1921, MNHN. Tunisia : Gulf of
Gabes, N. of Djerba, beach, 1 v., leg. BOUCHET and WAREN, 1982, MNHN. Italy : Viaregio, 2 spm., coll.
DEL PRETE, MNHN. Mauritania : 17°42'N/16°16' W , 54m, 1 spm., dredged R/V " N ' D i a g o " , leg. RICHER
 — 355 —

DE FORGES, 1981, MNHN. Senegal : Kayar, north of Dakar, 110-120m, 1 v., leg. MARCHE-MARCHAD,
6.IV.1967; 14°30,6'N/17°18,9"W, 43m, 1 v., leg. LEUNG TACK, 1983-1984, both MNHN. Liberia:
4°34,5'N/8°31'W, 64m, 1 v., dredged R/V "Calypso", leg. MARCHE-MARCHAD, 1956, MNHN. Cote
d'lvoire : Off Abidjan, 1 v., leg. MARCHE-MARCHAD, MNHN. Ghana : 4°40'N/2°08'W, 50m, 1 spm.,
dredged R/V "Calypso", leg. MARCHE-MARCHAD, 1956, MNHN. Cameroon : Between Wouri estuary
and Cap Nachtigal, 3°43'N/9°13'E, 32-35m, 1 v., trawled " Campo Star", leg. VON COSEL, 22-29.XI.1985,
2°23'N/9°41'E, 45m, 1 v., dredged R/V "Calypso", leg. MARCHE-MARCHAD, 1956, both MNHN.
Angola : Off Ponta das Lagostas, Bengo, province, 30-50 m, 2 spm., leg. GOFAS, MNHN.

M A P 13. — Distribution of Anadara corbuloides.

BIOTOPE : Anadara corbuloides lives in fine muddy sand or mud, well offshore from 30 m to 60 m and
occasionally deeper.

REMARKS : Anadara corbuloides apparently has two counterparts in the Panamic Pacific
province, namely A. mazatlanica (Hertlein and Strong, 1943) and Scapharca (Caloosarcd)
biangulata (Sowerby, 1833). The fact that these species are placed in different genera by KEEN
(1971) shows how the use of the equivalve condition gives rise to a n o m a l o u s supraspecific
classification. In A. corbuloides the juveniles are distinctly inequivalve but this condition
becomes obsolete with ontogeny.
F I G . 4 6 . — Internal view of right valve and dorsal view of Anadara corbuloides ( M o n t e r o s a t o ) . Angola. (Scale bar =
1 0 mm.)

Genus SENILIA G r a y , 1842

TYPE SPECIES : Area senilis Linné, 1758 ( S D Gray, 1847).

Senilia senilis (Linné, 1758)

(Pl. VIII, 1A-B, 2, 3 ; fig. 4 9 ; m a p 13)

Area senilis Linné, 1758 : 694.

TYPE MATERIAL : One specimen in the Linnean collection, London (DODGE, 1952 : 151).

TYPE LOCALITY : "Jamaica" (10th edition) but later emended to " O . Africano" (12th edition).

DESCRIPTION

Shell to 135 m m in length. Equivalve at all sizes. Inflated a b o u t the u m b o s but wedge
shaped in cross section; moderately to strongly u m b o n a t e . Inequilateral, beaks in the anterior
third, strongly prosogyre.
Outline subrhomboidal-trigonal, anteriorly expanded, posteriorly to some degree attenua­
te. Posterior area not defined; posterior angle w e a k ; posterior margin oblique, slope variable
from 30° to 65°. Median area rather flat, never sulcate even in post larval sizes; ventral margin
more or less straight, posterior ventral junction subacute to attenuate. Anterior margin
broadly rounded. Dorsal margin relatively short, straight.
Dorsal area short, more or less flat to cleft, rhomboidal with anterior area much more
broad. Ligament initially opisthodetic, developing beyond the beaks 20 m m and secondarily
amphidetic in large or old shells; chevrons strong, numbering u p to 8.
 — 357 -

Hinge plate becoming massive, teeth in two series, separated by a narrow oblique g a p ;
anterior set u p to 15 teeth, posterior set u p to 16 teeth. Teeth long weakly chevron shaped but
often becoming divided or distorted, mostly more or less vertical.
Sculpture of 13-15 broad flattened smooth ribs, occasionally narrowed and more rounded
especially in the least attenuate shells.
Periostracum persistent, a thin coating with very fine concentric thickenings in the
interspaces. Colour initially olivaceous then reddish brown to black.
Shell white.
A d d u c t o r scars subequal. Inner margin with large crenulations.

3.5
3.25
3.
X-3.13
2.75
2.5
a
| 2.25
to

* 2,
1.75.
X = 1.44
1.5.
X = 1.54
1.25.

L/A-Bk

F I G . 4 7 - 4 8 . — 4 7 : Box plots of the ratios of Total Shell Length to Height (L/H), Tumidity (L/T) and Anterior to Beak
Length (L/A-Bk) of Senilia senilis (n 2 1 ) . 4 8 : Bar chart of R i b N u m b e r for 2 1 shells of Senilia senilis.

SELECTED SHELL MEASUREMENTS : See figure 4 7 .

DISTRIBUTION : This species ranges from West Sahara (Rio de Oro) south to Angola.

MATERIAL EXAMINED : Mauritania : Port Etienne (now Nouadhibou) to Nouakchott, many v., from
12 sta., all subfossil, Miss. GRUVEL, 1908, MNHN. Senegal : Dakar, Marigot de Hann, 2 sp., 1 v., Miss.
GRUVEL, V.1908; Sine-Saloun region, mangroves, 8 spm., leg. BOUCHET, 9.VIII.1973; S-Casamance :
Katakalous Bolon, muddy sand, 3-4 m, many juv. spm.; Ourong Bolon, on sandbank, low tide, 1 spm..
 — 358 —

CO

o
O
CM

i ^tt
1 Ip
0

— \• \ o

\1 0

|20° [0°

M A P 1 4 . — Distribution of Senilia senilis. T h e empty circles indicates records of valves with subfossil appearance.

1 v.; Karabane, sandbank, 2 sh., all leg. VON COSEL, III. 1988; all MNHN. Gambia: 1 sp., coll.
D'AUGUSTIN, 1835, MNHN. Sierra Leone : Sierra Leone River, 1 spm., leg. LONGHURST, 13.X.1954,
MNHN. Liberia : Cap des Palmes, 2 sh., old coll., MNHN. Côte d'Ivoire : Azuretti Lagoon, Grand
Bassam, 1 v., leg. AUBERT DE LA RUE, 1950, MNHN. Ghana : Accra, on beach, 3 v., Miss. GRUVEL,
26.1.1910, MNHN. Benin : Icheppê-Olokun, 3 v. ; Lac Abénué, 1 sh., both Miss. GRUVEL, 1910, MNHN.
Sào Tomé : Off Punta Oquedelrey, 5m, 3 sh., 3 v., all juv., dredged R/V "Calypso", leg. MARCHE-
MARCHAD, 1956, MNHN. Cameroon : Victoria/Limbe, on beach, 6 sh., leg. VON COSEL, XII.1985,
MNHN. Gabon : Baie de Cap Lopez, 3 sh., Miss. GRUVEL, 1910, MNHN. R.P. Congo : Cocobeach,
1 sh., coll. SOYER, 1969; Loango, 6 sh., Office Pointe-Noire, 1969; Pointe-Noire, Plage Mondaine, on
beach, 1 v., leg. VON COSEL, XII. 1985, all MNHN. Zaïre : Crique de Moanda, 1 sh., leg. Miss. GRUVEL,
1910, MNHN. Angola : Cabinda, 1 sh., leg. C. R. BOETTGER, 1909; Ponta do Mussulo, Luanda province,
on beach, many sh.; Baia dos Tigres, Moçâmedes province, 1 sh., both leg. GOFAS, 1981-1985, all
MNHN.

BIOTOPE : Senilia senilis is a typical marine-estuarine species with lives in sandy mud, fine muddy sand
or fine sand in estuaries, lagoons and creeks with regular tidal and seasonal salinity changes, often in the
vicinity of mangroves, where it may be found in great quantities. It tolerates reduced salinity as well hyper
salinity and lives predominantly in the lower intertidal zone.

REMARKS : This species is quite variable in outline with some shells distinctly attenuate
posteriorly and others rather shortened. In the latter this compression results in the ribs being
narrower, more rounded and more elevated (pl. VIII, 3 ) . The functional morphology of this
species was discussed by Y O N G E ( 1 9 5 5 ) a n d the recent and fossil ecology by DEBENAY and S Y
(1989).
 — 359 —

FIG. 49. — Internal view of right valve and dorsal view of Senilia senilis (L.). Angola. (Scale bar = 5 m m . )

G e n u s BATHYARCA Kobelt, 1891

TYPE SPECIES : Area pectunculoides Scacchi, 1834 ( O D ) .

DIAGNOSIS : The genus Bathyarca is placed in the Anadarine by NEWELL (in MOORE, 1969), but, as
shown by OLIVER and ALLEN (1980), the genus contains both epibyssate and endobyssate species thus
encompassing the range of morphology exhibited by both the Arcinae eg. Barbatia and the Anadarinae
eg. Scapharca, The unique feature of the genus is the mantle extension which is present to some degree
regardless of shell shape. This is one of the few examples in the Arcidae where a group of species can be
isolated by a synapomorphy and consequently it is a more rigid group than either the Arcinae or
Anadarinae. There may therefore be a case for raising the genus to subfamily level and creating genera to
separate the epibyssate species from the endobyssate species (OLIVER, in prep.).

Bathyarca grenophia (Risso, 1826)

(PI. VII, 4 ; fig. 50)
Area grenophia Risso, 1826 : 313.
Area pectunculoides Scacchi, 1834 : 82.

TYPE MATERIAL : Holotype (1 valve) of A. grenophia in MNHN Paris; type material of A.

pectunculoides not located.

TYPE LOCALITY : Not cited; here selected Nice, Mediterranean France.

DESCRIPTION

Shell to 6 m m in length. Inequivalve, R V fitting into LV, not greatly inflated.

Inequilateral, beaks in the anterior half.
Outline suboval, posteriorly expanded. Posterior area indistinct, posterior angle rapidly
becoming obsolete, posterior margin broadly rounded. Median area flattening or weakly
 — 360

sinuate, ventral margin rounded, usually with a n indentation at the byssal exit. Anterior
margin rounded but distinctly narrower than posterior. Dorsal margin straight.
Dorsal area narrow, u m b o n a l separation weak. Ligament of few chevrons restricted to
behind the beaks.
Hinge plate weak, teeth in two series separated by a distinctly edentulous space, anterior
set u p to 5 teeth, posterior set u p to 8 teeth. Teeth small, straight, lateral teeth becoming
strongly oblique.
Sculpture weak, subcancellate. Periostracum weakly haired. Shell white.

DISTRIBUTION : Throughout the Atlantic and in the subarctic from Norway to Angola in the east and
from Greenland to Bermuda in the west.

MATERIAL EXAMINED : Ibero-Maroccan Gulf : 36°46' N/07°07' W, 368-371 m, 5 spm. + 2 v. ;

36°36'N/07°24'W, 478-491 m, 1 spm. ; 36°15'N/08°0r W, 1340-1263 m, 15 spm. ; 35°54' N/06° 13' W, 133-
137 m, 1 v.; 35°54'N/06°14'W, 150m, 3 spm.; 36°41'N/07°19"W, 543-544 m, 4 spm.; 35°54'N/06°14'W,
293m, 6 spm.; 35°35'N/03°45'W, 480m, 5 spm. ; 35°31'N/07°26'W, 1209-1302m, 2 spm. ;
35°26'N/04°19'W, 170m, 1 spm. ; 34°24'N/07°31'W, 1203m, 1 spm. ; 34°21'N/07°24'W, 885-895m,
6 spm.; 33°49'N/08°24'W, 255-267m. 5 spm.; 33°46'N/08°30'W, 309m, many spm.; 33°45'N/08°32'W,
345 m, 1 spm., 6.VI.2984, all dredged R/V " C r y o s " , BALGIM campaign, all M N H N . Senegal : 230" off
Cap Manuel, 120-215m, many v. + 1 sh., in stomach of holothurian ; off Dakar, 14°27'N/17°33'W, 150-
250m, many v.; 14°53'N/17°30'W, 205-230m, 1 spm. + 3 v. ; 170-200m, 3 spm. + 19 v.; Baie de
Gorée, 80-250m, 14 v.; SW of Gorée, 150-250m, 5 v., all dredged "Gérard Tréca", leg. MARCHE-
MARCHAD, 1954-1958; all M N H N . Also based on the material used by OLIVER and ALLEN (1980).

BIOTOPE : This species lives attached to hard substrates and secondary hard substrates in soft
sediment areas such as small stones, dead shells, worm tubes etc. In the north of its range it is found in the
sublittoral zone as shallow as 2 m but further south it submerges to become a bathyal species and off the
west African coast is found from 100-1300 m.

REMARKS : Comparing the west African material a n d more typically that from the greater
end of the depth range with the northern shallow water material it is noted that the former
possesses a thinner almost translucent shell in which the sculpture is very weakly developed. In
these respect it resembles the form pellucida described by OLIVER and ALLEN ( 1 9 8 0 ) from
similar depths off the eastern American coast. Also amongst the B A L G I M material from the
Ibero-Maroccan gulf are many specimens of Bathyarca obliqua Philippi. This species is larger
and more elongate than B. grenophia with a resultantly longer hinge line. It also ranges further
north to boreal waters but as yet it has not been recorded from the west African coasts south
of Morocco.

Bathyarca inaequisculpta (Smith, 1885)

(PI. VIII, 5 ; fig. 51)

Area (Scapharca) inaequisculpta Smith, 1885 : 267, pi. 17, figs. 8a-d.
Bathyarca dakarensis Locard, 1898 : 321, pi. 8, figs. 25-28.
TYPE MATERIAL : Syntypes in B M N H . The syntypes of B. dakarensis are in M N H N .
TYPE LOCALITY : Off Culebra Islands, West Indies. The type locality of B. dakarensis is off Dakar.
Senegal.
 — 361 -

FlG. 50-51. — 50 : Internal view of left valve of Bathyarca grenophia (Risso). Senegal, 150-250 m. 51 : Internal view of
right valve and dorsal view of Bathyarca inaequisculpta (Smith). Angola, 3797 m. (Scales bars = 1mm.)

DESCRIPTION

Shell to 8 m m in length. Inequivalve. Inflated. Inequilateral, beaks in the anterior half.

Outline subcircular, as high or higher than long. Posterior area and posterior angle not
demarcated. Margins other than dorsal not differentiated, round. Dorsal margin relatively
long and straight. Median area curved, not sulcate.
Dorsal area narrow, u m b o n a l separation slight. Ligament with few chevrons restricted to
behind the beaks.
Hinge plate narrow, teeth in two series but separation obscure, anterior set u p to 8 teeth,
posterior set u p to 10 teeth. Teeth small, straight, lateral teeth becoming strongly oblique.
Sculpture weak, finely decussate, often a little stronger RV. Periostracum weak, sparsely
pilose. Shell white.
DISTRIBUTION : T h i s s p e c i e s is w i d e l y d i s t r i b u t e d in b a t h y a l a n d abyssal zones t h r o u g h o u t the
A t l a n t i c O c e a n f r o m 50 N t o 1 4 ° S .
 - 362 -

MATERIAL EXAMINED : The material examined is that used by OLIVER and ALLEN ( 1 9 8 0 ) .

BIOTOPE : This is a truly deep sea species and is confined to abyssal oozes in which it lives at or near
the surface anchored to sediment particles by a multi threaded byssus. Although it has been found as
shallow as 7 0 0 m it is more typically abyssal ranging from 1900-5000 m.

Acknowledgements

We would like to thank Chris MEECHAN for the line illustrations, Kevin THOMAS for the
photography and the National Museum of Wales for these and other facilities required to prepare this
paper. For loan of the major part of the material the first author is grateful to Philippe BOUCHET
(MNHN).
Sincere thanks go to our colleague Serge GOFAS (MNHN Paris) who contributed ecological data
and collected a considerable part of the material referred to in this paper during his time in Angola.
Most material from Mauritania was donated to M N H N by Bertrand RICHER DE FORGES (ORSTOM),
which is also gratefully acknowledged.
Logistic assistance during the fieldwork of the second author was rendered by several persons and
organisms : the centres of ORSTOM (Institut Français de Recherche Scientifique pour le Développe-
ment en Coopération) in Dakar (B. DALMAYRAC), Conakry (F. DOMAIN), Yaounde (Ph. MATHIEU),
Brazzaville ( J . L. FRÉZIL) and Pointe-Noire (M. BARRO), the Centre de Recherche Océanographique
Dakar-Thiaroye, Senegal (CRODT) (A. FONTANA), the Station de Recherches Halieutiques Limbe,
Cameroon ( J . C. NJOCK) and the Douala Research station of the Ministry of Higher Education and
Scientific Research of Cameroon (the Director). Assistance in the field during littoral work was
received from S. GILLES (ORSTOM Casamance, Ziguinchor), M. YANSANE (Fisheries Research
Institute Conakry) and P. BERNARD (Libreville). The captains and crews of the vessels " Louis Sauger"
(CRODT Dakar), "André Nizery " (ORSTOM), " C a m p o S t a r " (PECAM, Douala) and " K o u n d a "
(SOCIMPEX, Pointe-Noire) are thanked for their kind collaboration during work on board.

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208.

PLATE I

1A-D. — Area noae Linné : West Sahara, off C a p Barbas, 21°05'N/17°14'W, 43-45 m, M N H N . 73 m m . 1A,
periostracum removed.
2. — Area noae Linné : C a p e Verde Islands, Ilha do Sal, Santa M a r i a , on beach, M N H N . 11.5 mm.
3A-D. — Area bouvieri Fischer : Angola, C a c u a c o , Bengo province, 5-10m, M N H N . 6 0 m m . 3A, periostracum
removed.
4. — Area bouvieri Fischer : Angola, C o r i m b a , L u a n d a province, 10-20m, M N H N . 11mm.
PLATE I
 — 366

PLATE II

1A-D. — Arca avellana turbatrix n. ssp. : H o l o t y p e . Angola, Santo A n t o n i o , Benguela province, intertidal, rocks.
M N H N . 31 m m . 1A, periostracum removed.
2. — Arca avellana turbatrix n. ssp. : Paratype. Angola, Lucira, M o c à m e d e s province, M N H N . 18.7 m m .
3A-B. — Arca imbricata Bruguière : Florida, N M W Z . 50 m m . 3A, periostracum removed.
4. — Arca avellana Lamarck : Seychelles, M N H N . 30 m m , periostracum removed.
5A-B. — Arca tetragona Poli : Ireland, Berehaven, Bantry Bay, N M W Z . 21.3 m m .
6. — Arca tetragona Poli : England, F a l m o u t h , N M W Z . 26.6 m m .
7A-B. — Acar plicata (Dillwyn) : C a p e Verde Islands, Mindelo, Sào Vicente, M N H N . 7A, 19.1 m m ; 7B, 18.2mm.
8A-B. — Acar pulcinella (Reeve) : M e d i t e r r a n e a n , Algeria, M N H N . 18.2 m m .
 - 368 -

PLATE III

1A-B. — Barbatia complánala (Bruguière) : Côte d'Ivoire, Canal de Vridi, Abidjan, M N H N . 1A, 51 m m ; 1B, I 8 m m .
2. — Barbatia complánala (Bruguière) : Angola, C h a p e u A r m a d o , M o ç â m e d e s province, rocks at low tide, M N H N .
40 m m .
3. — Barbatia complánala (Bruguière) : Angola, C a o t i n h a , Benguela province, M N H N . 33 m m .
4. — Barbatia complánala (Bruguière) : Angola, Lucira (Praia d o Cesar), M o ç â m e d e s province, M N H N . 45 m m .
5A-B. — Barbatia complánala (Bruguière) : Angola, Praia Amelia, M o ç â m e d e s province, rocks at low tide, M N H N .
5A, 22 m m ; 5B, 16 m m .
6A-B. — Barbatia gabonensis n. sp. : Holotype. G a b o n : off M a y u m b a , 3°25'S/9°56'E, 100m, M N H N . 2 7 . 7 m m .
7A-B. — Barbatia ionlhados n. sp. : Holotype. Côte d'Ivoire, Sud Bassam, 200 m, M N H N , 20.6 m m .
8. — Barbatia iegumen (Lamy) : Angola, C o r i m b a , L u a n d a province, Praia E t a m b a r , between rocks at low tide,
MNHN. 41mm.
9A-B. — Barbatia Iegumen (Lamy) : Angola, 10 km. S of Ambrizete, Zaire province, intertidal, on beach, M N H N .
24 mm.
10. — Barbatia Iegumen (Lamy) : R.P. C o n g o , Pointe-Noire, Plage M o n d a i n e , M N H N . 53 m m .
PLATE III
 — 370 —

PLATE IV

1A-B. — Barbatia (Nipponarca) allocostata n. sp. : H o l o t y p e , Guinea-Bissau : M o u t h of Rio G e b a , 11°57,5'N/

16°27'W, 7 m , M N H N . 2 7 . 7 m m .
2. — Barbatia (Nipponarca) allocostata n. sp. : P a r a t y p e as holotype, M N H N , 28 m m , periostracum removed.
3A-B. — Anadara geissei ( D u n k e r ) : Angola, Baie de C o r i m b a , L u a n d a province, 10-20 m, M N H N . 7 1 m m .
4A-B. — Anadara geissei ( D u n k e r ) : Angola, C a c u a c o , Bengo province, 5-10m, M N H N . 4A, 3 3 m m ; 4B, 1 8 m m .
5. — Anacara geissei (Dunker) : G u i n e a , 9 ° 5 0 ' N / 1 4 ° 1 4 ' W , 15m, M N H N . 5 2 m m .
6A-B. — Anadara geissei ( D u n k e r ) : C a p e Verde Islands, Sào Vicente, Baia P o r t o G r a n d e , on beach, M N H N . 6A,
50 m m ; 6B, 48 m m .
PLATE IV
 - 372 -

PLATE V

1A-B. — Anadara polii (Mayer) : Mediterranean, Banyuls, M N H N . 3 4 m m , oblong m o r p h .

2. — Anadara polii (Mayer) : Mediterranean, Naples, M N H N . 39 m m , intermediate form.
3A-C. — Anadara polii (Mayer) : Atlantic M o r o c c o , M N H N . 3A, 30°24' N/09°54' W, 105 m, 28 m m , oblong m o r p h .
3B, 3 0 ° 2 6 ' N / 0 9 ° 4 4 ' W , 65 m, 23 m m , anteriorly constricted m o r p h . 3C, 3 0 ° 0 5 ' N / 0 9 ° 5 0 ' W , 110 m, 22 m m , globose
morph.
4A-B. — Anadara polii (Mayer) : Senegal, 14°50,1'N/17°29,3'W, 150m, M N H N . 4A, 3 1 m m , oblong m o r p h . 4B,
29 m m , globose m o r p h .
5. — Anadara polii (Mayer) : Côte d'Ivoire, 0 5 ° 0 1 ' N / 0 5 ° 1 7 ' W , 4 0 m , M N H N . 2 4 m m , oblong m o r p h .
6. — Anadara polii (Mayer) : Côte d'Ivoire, off Abidjan, 70 m, M N H N . 22 m m , oblong m o r p h .
7. — Anadara polii (Mayer) : Senegal, off D a k a r , 2 0 0 m , M N H N . 2 4 m m , anteriorly constricted m o r p h .
8. — Anadara polii (Mayer) : C a m e r o o n , 0 2 ° 3 9 ' N / 0 9 ° 4 0 ' E , 60-65 m, M N H N . 26 m m , anteriorly constricted m o r p h .
9. — Anadaripolii (Mayer) : G a b o n , 00°32,8'N/08°43,1 ' E , 125-145m, M N H N . 3 9 m m , anteriorly constricted m o r p h .
PLATE V
 — 374 —

PLATE VI

1A-D. — Anudara eborensis n. sp. : Holotype. C o t e d'lvoire, off G r a n d Bassam, 2 0 m , M N H N . 3 1 . 9 m m .

2A-B. — Anudara subglobosa ( D u n k e r ) : M a u r i t a n i a , 1 7 ° 3 0 ' N / 1 6 ° I 5 ' W , 100m, M N H N . 18.5mm.
3A-D. — Anadara senegalensis (Gmelin) : Cote d'lvoire, off Abidjan, 2 0 m , M N H N . 3A-C, 2 1 . 8 m m . 3 D , 2 3 . 5 m m .
4A-B. — Anadara senegalensis (Gmelin) : Guinea-Bissau, 12°05'N/17°02,5'W, 14m, M N H N . 2 1 . 8 m m .
5A-B. — Anadara senegalensis (Gmelin) : Angola, C a c u a c o , Bengo province, 0 - 1 0 m , M N H N . 15.8mm.
6A-C. — Anadara camerunensis n. sp. : Holotype. C a m e r o o n , Victoria/Limbe, 8-10m, M N H N . 15.1mm.
 — 376 —

PLATE VII

1. — Anadara senegalensis (Gmelin) : Senegal, South C a s a m a n c e , C a p Skirring-Diembéring, on beach, M N H N .

23.1 m m .
2. — Anadara senegalensis (Gmelin) : Senegal, South C a s a m a n c e , C a p Skirring-Diembéring, on beach, M N H N .
2 5 . 4 m m , C a s a m a n c e variant.
3. — Anadara senegalensis (Gmelin) : Senegal, N o r t h C a s a m a n c e , Abéné-Kafoutine, on beach, M N H N . 23.0 m m ,
intermediate form.
4. — Anadara senegalensis (Gmelin) : Senegal, N o r t h Casamance, Abéné-Kafoutine, on beach, M N H N . 2 6 . 2 m m ,
C a s a m a n c e variant.
5 A - D . — Anadara corbuloides (Monts.) : Angola, off P o n t a das Lagostas, Bengo province. 30-50m, M N H N . 6 9 m m .
PLATE VII
 — 378 —

PLATE VIII

1A-B. — Senilia senilis (Linné) : Angola, C a c u a c o , Bengo province, M N H N . 92 m m .

2. — Senilia senilis (Linné) : Angola, Pta. do Mussulo, L u a n d a province, M N H N . 50 m m .
3. — Senilia senilis (Linné) : G a b o n , C a p Esterias, Pointe Idolo, M N H N . 3 9 m m .
4. — Bathyarca grenophia (Risso) : M o r o c c o , 3 5 ° 3 5 ' N / 0 3 ° 4 5 ' W , 480 m, M N H N . 8.4 m m .
5. — Bathyarca inaequisculpta (Smith) : Angola, 14°40' S/09°54' E, 3797 m, N M W Z . 6.7 m m .
6A-B. — Bentharca asperula (Dall) : Angola, 14°31,8'S/09°46'E, 3 9 7 5 m , N M W Z . 10.7mm.
PLATE Vili
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PLATE IX

Scanning electron micrographs of the periostracal bristles from the carinal ridge of : A, Arca avellana lurba-
trix n. ssp. and B, Arca imbricata (Brug.) from Florida. (Scale bar = 0.5 m m . )
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PLATE IX