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SOCIETE BELGE DE MALACOLOGIE

Edgar WAIENGNIER, 1936 - 2010

Membre de la Société Belge de Malacologie depuis 1977, Edgar nous rejoignit au sein du comité en 1980.

Depuis lors ses connaissances en matière de malacologie, sa présence régulière à nos réunions, sa participation a

nos excursions et son implication dans tous nos projets ont toujours été très précieux et appréciés.

Il était également le spécialiste des mollusques terrestres, en particulier des Helicidae et nous a présente

plusieurs conférences sur ce sujet bien précis.

Nous saluons ici un collègue mais aussi un ami qui sera toujours parmi nous en pensée.

Ce numéro de Novapex est dédié à sa mémoire.

Member of the Belgian Malacological Society since 1977, Edgar joined us on the board of directors in 1980.

Since then his knowledge of malacology, his regular presence at our meetings, his participation in our tield trips

and his involvement in ail our projects were always highly valued and appreciated.

He was also the specialist on terrestrial molluscs, in particular of Helicidae, and he presented us with several

conférences on this subject.

We salute here not only a colleague but also a friend who will always be among us in thought.

This issue of Novapex is dedicated to his memory.

NOVAPEX est paru pour la première Ibis le 20 février 2000, remplaçant notre revue APEX qui en était à son

quatorzième volume. De 2000 à 2010 nous avons publié de nombreux articles dans les numéros ordinaires de

Novapex et dans 7 numéros Hors Série. Durant ces 1 I années nous avons fait appel à de nombreux référés. Leurs

noms sont mentionnés ci-dessous.

NOVAPEX was published for the first time on February 20, 2000 to replace APEX, then at its 14th volume.

From 2000 to the end of 2010 we published numerous papers in the normal issues of Novapex and in 7 Spécial

Issues. During these 11 years we asked for the help of many referees. They are listed below.

Reviewers for Novapex 2000-2010 - Référés pour Novapex 2000-2010

Warren ALLMON

Alan BEU

Arthur BOGAN

Philippe BOUCHET

Franck BOYER

Christiane DELONGUEVILLE

Bunjamin DHARMA

Henk DIJKSTRA

Doug EERNISSE

Marien FABER

Yves FINET

Koen FRAUSSEN

Emilio F. GARCIA

Olivier GARGOMINY

Serge GOFAS

Daniel GRAF

Roland HADORN

M.G. HARASEWYCH

Dai HERBERT

Greg HERBERT

Roland HOUART

Yuri K.ANTOR

t Kevin LAMPRELL

Harry G. LEE

Robert LIPE

James H. MC LEAN

Bruce MARSHALL

t Phillip A. MAXWELL

Tony McCLEERY

Didier MERLE

David MONSECOUR

Kevin MONSECOUR

Antonio MONTEIRO

Rob MOOLENBEEK

Igor MURATOV

Bruce NEVILLE

Robert PEUCHOT

Emilio ROLAN

Gary ROSENBERG

Richard SALISBURY

Roland SCAILLET

Luiz Ricardo LOPES SIMONE

Donn TIPPETT

t Hans TURNER

Georges VAUQUELIN

Geerat VERMEIJ

Claude VIL V ENS

Rudo von COSEL

t Edgar WA1ENGNIER

Andrew WAKEFIELD

Anders WAREN

John WOLFF

B. M. Landau & L. T. Groves

Novapex 12(1-2): 1-38, 10 mars 2011

Cypraeidae (Mollusca: Gastropoda) from the early Miocene

Cantaure Formation of northem Venezuela

Bernard M. LANDAU

Centre de Geologia da Universidade de Lisboa. Campo Grande, 1749-016 Lisboa, Portugal and

International Health Centres, Av. Infante de Henrique 7, Areias Sào Joâo, P-8200 Albufeira, Portugal.

bernielandau@sapo.pt

Corresponding author

Lindsey T. GROVES

Natural History Muséum of Los Angeles County, Malacology Department, 900 Exposition Boulevard,

Los Angeles, California, 90007, USA.

lgroves@nhm.org

KEYWORDS. Cypraeidae, Mollusca, Miocene, Cantaure Formation, Venezuela, new species.

ABSTRACT. This is the first account of the rich assemblage of Cypraeidae from the late early

Miocene, Burdigalian Cantaure Formation, Paraguanâ Peninsula, Falcôn Province, Venezuela.

Twelve species are recorded from these deposits, ten of which are new to science: Luria

cantaurana n. sp., L. isabellaprimitiva n. sp., Trôna ingrami n. sp., Propustularia longidentata n.

sp., P. paraguanensis n. sp., Zonaria pingata n. sp., Z. pseudotumulus n. sp., Pseudozonaria

pr 'aeaequinoctialis n. sp., P. primarobertsi n. sp. and P. falconensis n. sp. Comparative taxon

Muracypraea "henekeni ot Groves (1997) from the middle to late Miocene Gatun Formation ot

Panama and Angostura Formation of Ecuador is herein described as M woodringi n. sp. Cypraea

fossula (Ingram, 1947) is considered a junior subjective synonym of Trôna trinitatensis

(Mansfîeld, 1925). The type material of Jousseaumea joossi Schilder, 1939 and Pustularia

mejasensis Schilder, 1939 are in very poor condition and do not adequately show species

characteristics, and are therefore considered nomma dubia.

INTRODUCTION

A rich and varied cypraeid fauna is here reported from

the early Miocene (Burdigalian) Cantaure Formation

as defined by Hunter and Bartok (1974), Paraguanâ

Peninsula, Falcôn State, Venezuela. Schilder (1939)

first documented the family Cypraeidae from these

deposits with the report of Luria (Basilitrona)

patrespatriae (Maury, 1917) and Ingram (1947)

described Cypraea fossula from the sarne formation.

However, no further cypraeid taxonomie work on the

Cantaure Formation has been attempted. Jung’s

(1965) landmark systematic monograph on the

assemblage described and illustrated a specimen

identified as Cypraea aff. isabella Linnaeus, but made

no further mention of additional cypraeids in the

formation. Herein, these omissions are addressed.

Gibson-Smith & Gibson-Smith (1979) described

the presence of ‘upper’ and ‘Iower’ beds in the

Cantaure Formation. The basal unit is defined by a

breccia composed of Balanus barnacle fragments and

blocks of granité overlain by silty and gypsiferous

shales with sandy levels (Hunter & Bartok, 1974).

They also note rich molluscan levels within the Iower

unit. The upper level is sandier, and the transition

between the two units is unclear. Dr. Ernily Vokes,

who has also visited the deposits, did not recognize

the presence of an ‘upper’ and ‘Iower’ bed (personal

communication, BL). The gastropod assemblage

found in the ‘upper’ and ‘Iower’ beds is similar, with a

prédominance of filter-feeding turritellids in the

‘upper’ beds and a greater number of rocky-bottom

dwellers in the ‘Iower’ bed. Cypraeids are not

common in the Cantaure Formation, but are found far

more often in the ‘Iower’ beds, which are designated

as the type locality for the new species.

Material and Methods

The material described here is from the Gibson-Smith

collection housed in the Naturhistorisches Muséum

Basel (NMB coll.), Switzerland and the Bernard

Landau collection (BL coll.), now deposited in the

Naturhistorisches Muséum Wien (NHMW coll),

Vienna, Austria. Some specimens are deposited in the

Natural Histoiy Muséum of Los Angeles County,

Invertebrate Palaeontology Department (LACMIP

coll.). Ail shells were also examined under UV light

as described by Olsson & Petit (1968).

Abbreviations

Abbreviations used for institutional catalogue and/or

locality numbers are as follows: ANSP, Academy of

Natural Sciences of Philadelphia, USA; BL,

Collection of Bernard Landau (Collection now at

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northern Venezuela

NHMW); DFB coll., Dirk Fehse collection, Berlin,

Germany: LACMIP Natural History Muséum of Los

Angeles County, Invertebrate Palaeontology

Department, Los Angeles, California, USA; NHMW,

Naturhistorisches Muséum Wien, Vienna, Austria;

NMB, Naturhistorisches Muséum Basel, Switzerland;

PPP, Panama Palaeontologieal Project; TU, Tulane

University, New Orléans, USA (Tertiary collections

now at the National Muséum of Natural History,

Smithsonian Institution [USNM]); UCMP, University

of California, Muséum of Paleontology, Berkeley,

California, USA; and USGS, United States Geological

Survey, Reston, Virginia, USA.

Systematic Palaeontology

Cypraeids can be a notoriously difficult group to work

on, especially the fossils as the surface colour and

pattern are usually lost. A single major taxonomie

study has been undertaken in recent years on the

tropical American Neogene cypraeid assemblages;

Dolin (1991) revised the Cypraeoidea from the late

early Miocene Chipola Formation of northern Florida,

which is roughly contemporaneous with the Cantaure

Formation. For the ease of comparison of the two

faunas, the morphologie terminology of Dolin (1991:

fïg. 1) and Dolin & Lozouet (2004: fîgs. 2a-c) are

utilized.

Cypraeid systematics has been greatly enhanced

by the works of Meyer (2003, 2004) in particular his

molecular data-base for the family. The phylogenetic

implications of this new data outlined by Meyer

(2004), and the systematic arrangement suggested by

Lôpez Soriano (2006) based on Meyer’s work, are

followed here. Higher level systematics follows that

of Bouchet, et al. (2005).

The shell formula proposed by Schilder (1935:

327) has been given for each species. This formula is

derived from measurements taken from ail available

tully mature and normally formed specimens. It

consists of the following éléments: [L (W-H) LT: CT],

L: average length in mm, W: average width/ length

ratio in %, H: average height/ length ratio in %, LT:

normalized number ot labial teeth, CT: normalized

number of columellar teeth. The normalized number

of teeth - in relation to a shell of 25 mm length - is

calculated as follows: T = 7 + [(c-7) x V (25/L)], T:

normalized number of teeth, c: teeth counted, L:

length. This shell formula is useful to highlight

différences and similarities between species, but

should not be used on its own to distinguish between

species.

C lass GASTROPODA Cuvier, 1791

C lade HYPSOGAS TROPODA Ponder & Lindberg

1997

Clade CAENOGASTROPODA Cox, 1959

Clade LITTORINIMORPHA Golikov &

Starobogatov, 1975

Superfamily CYPRAEOIDEA Rafinesque, 1815

Family CYPRAEIDAE Rafinesque, 1815

Subfamily CYPRAEINAE Rafinesque, 1815

Genus Muracypraea Woodring, 1957

Type species: Cypraea mus Linnaeus, 1758, by

original désignation.

Description. Shell pyriform to triangular of small to

moderately large size (30 to 75 mm); posterior part of

dorsal surface smooth, roughened, warty, or

bituberculate; outer lip, wide, slightly constricted near

anterior end; fossula indistinct, wide, shallow, and

smooth.

Discussion. A consensus among specialists on the

generic assignment for this group of species is lacking

as is an agreement of the number of living and fossil

species (Woodring, 1959; Petuch, 1979, 1987;

Doneddu & Manunza 1996; Lorenz & Hubert, 2000;

Meyer, 2004). This may in part be due to the

inadéquate description of the type species by

Woodring (1957). Muracypraea was originally

proposed as a subgenus of Cypraea. Lorenz & Hubert

(2000) considered “ Muracypraea ” an informai group

within Siphocypraea Heilprin, 1887 which was

proposed as a subgenus within the genus Cypraea.

Some species of the S. problematica complex hâve a

strongly curled comma-shaped posterior channel that

conceals the spire. Muracypraea was proposed for the

Recent Cypraea mus Linnaeus, 1758 and the related

fossil species that possess a more normal posterior,

wide aperture, and commonly well-developed dorsal

callosities, dorsal tubercules, and occasionally a

central spike-like dorsal projection. Siphocypraea

ranges from the early Pliocène to Pleistocene and is

restricted to the southeastern United States whereas

Muracypraea ranges from the early Miocene to

Recent of the Caribbean Basin to Peru and

southeastern California. Doneddu & Manunza (1996)

and Fehse (in press, a) considered Muracypraea a

junior synonym of Barycypraea Schilder, 1927. They

drew attention to the long géologie history of the

genus with fossil représentatives in the Indo-Pacific

(see Dharma, 2005; Fehse (in press, b) and the

Caribbean Tertiary. Doneddu & Manunza (1996)

traced the origins of the M. mus complex to Cypraea

(Bernayia) [,?/c] saltoensis (Clark in Clark & Durham,

1946) ot the Eocene of Colombia. However, close

examination of the poorly preserved holotype makes it

clear that a conclusive generic assignment is not

possible, and détermination of a possible new genus is

beyond the scope of this paper. Kay (1996) and

Groves (1997) used Muracypraea as a full genus.

Woodring (1959) considered ail Miocene specimens

from the circum Caribbean basin to be Cypraea

(Muracypraea) henekeni (lectotype; Figs 144-145).

However, it is notable that specimens from the

Miocene Gatun Formation of Panama are different

from those of Venezuela, Trinidad, Colombia, and the

Dominican Republic. Most specimens from the Gatun

Formation examined by the senior author hâve an

B. M. Landau & L. T. Groves

NOVAPEX 12(1-2): 1-38, 10 mars 2011

axially striped colour pattern or blotching of dull

reddish brown or yellowish orange. Additionally, the

marginal callus is extremely thin and poorly delimited,

and does not extend onto the dorsum, and the shell

periphery of the Gatun shells feel corrugated, whereas

they are smooth in M. henekeni and M. hyaena. We

separate two distinct taxa within the Gatun

assemblages (see below). The early Pliocène

Muracypraea grahami (Ingram, 1947) t'rom Cubagua,

Venezuela (Figs 29-32) is again different (Landau &

Silva, 2010). Ingram (1947) described two

Muracypraea species from Cubagua; Cypraea nigosa

[junior homonym of C. nigosa Broderip, 1827] and

Cypraea grahami. Although the holotypes of the two

shells look quite different, Landau & Silva (2010)

illustrated intermediate forms and recognised a single

species in the Cubagua deposits. The shell is very

large and solid (63-73 mm in length) and their outline

is more triangular and their dorsum even higher than

M hyaena (Landau & Silva 2010). The tubercles are

staggered on the dorsum in a similar fashion to M.

hyaena. The margins are corrugated, as in

Muracypraea isthmica Schilder, 1927, but more

coarsely so. In the best preserved specimen from

Cubagua an indication of a striped colour pattern is

présent, akin to that seen in the Gatun Formation

shells. On the Pacific side of the Gatunian Province

Muracypraea cayapa (Pilsbry & Olsson, 1941) from

the early Pliocène Jama Formation of Ecuador is a

large poorly known species represented by a single

broken specimen. Many other taxa of Muracypraea

hâve been described from the Caribbean Neogene,

which awail validation. Species of the extant

Muracypraea mus complex are known to hâve an

intracapsular larval development (Ranson, 1967;

Hoeblich, 1979). This assumption cannot be validated

with the fossil taxa, however, this type of development

would favour a restricted distribution of species and a

high degree of variability between populations. This

helps explain the difficulty encountered with the

Caribbean Neogene Muracypraea populations and

may be analogous to the situation among Recent

species of the genus Zoila Jousseaume, 1884 in

Australia which also shows intracapsular

development. Numerous species of Zoila are not only

geographically restricted, but numerous intermediate

forms are common ofif the western and Southern coasts

of Australia (Lorenz, & Hubert, 2000; Lorenz, 2001).

Based on mitochondrial molecular data presented

by Meyer (2004), the two extant Barycypraea taxa B.

teulerei (Cazenavette, 1846) and B. fultoni (Sowerby,

1903) were placed as sister taxa to Zoila Jousseaume,

1884, reaffirming the (validity of the subfamilial clade

Bernayinae Schilder, 1927, whereas Muracypraea

mus was placed in the Cypraeinae Gray, 1824. Based

on morphological différences, Fehse (in press, a) will

reassign the two living Barycypraea species and the

fossil species B. zietsmani (Liltved & LeRoux, 1988)

[Pliocène, South Africa], B. schilderi (Dey in

Schilder, 1941) [Pliocène, India], B. gendingaensis

(Martin, 1899) [Pliocène, Indonesia], and B.

kendengensis (Schilder, 1941) [Pliocène, Indonesia] to

a new genus (Dirk Fehse personal communication

2010). Some of the shells of the lndonesian fossil

Barycypraea group are extremely similar to those of

Muracypraea in the Tropical American Neogene and

could represent a convergent evolutionary process. A

morphological feature that helps separate

Muracypraea from Barycypraea are the flattened

spatulate horizontal expansions on either side

produced from the abapical tips of the inner and outer

lips of M. henekeni and many related species.

Occasionally these spatulate expansions are hyper

developed in the Caribbean forms whereas they are

somewhat obsolète in the lndonesian species of

Barycypraea. Although the two généra may hâve a

common Tethyan ancestor, consistent shell différences

and the vast géographie séparation could justify

different généra. Many of the Muracypraea shells are

lightweight, less callused with a broad aperture and

less developed, widely spaced dentition, whereas the

shells of Barycapraea are heavy, strongly callused

with a narrow aperture and a distinct, close-set

dentition. The Muracypraea exceptions being M. mus

donmorei (Petuch, 1979), M mus bîcornis (Sowerby,

1870), and M. grahami (C. nigosa Ingram, 1947

junior homonym of C. rugosa Broderip, 1827; see

Landau & Silva 2010). The dorsal callosities of both

généra can be variable even among the same species.

However, in Muracypraea they tend to be restricted to

the posterior portion of the dorsal surface whereas in

Barycypraea there may be a second set located

slightly anterior of the dorsum centre. Because the

Miocene fossil record for the central Pacific islands is

incomplète, and until a conclusive answer to whether

two généra should be used for the two seemingly

geographically separated species-groups, the

classification of Schilder & Schilder (1971) for

Muracypraea is used for the Caribbean species.

Muracypraea hyaena (Schilder, 1939)

Figs 1-14

1939 Siphocypraea angustirirna hyaena Schilder,

p. 23, fig. 24.

1971 Siphocypraea (Muracypraea) angustirirna

hyaena Schilder and Schilder, p. 29.

Material. 8 specimens BL coll.; 1 specimen LACMIP

13648 (ex BL coll.).

Description. Shell small to medium, solid, rotund,

dorsum very strongly humped % distance from the

anterior end, in fully adult specimens bearing two

small tubercles, right tubercle larger and anterior to

left; spire depressed, covered by adapical callus; sides

rounded, strongly callused, with the callus ascending

progressively adapically to just below the apex

posterior to the dorsal hump; ventrum flattened,

slightly convex in profile; aperture almost straight.

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

anterior third weakly dilated; siphonal canal deep,

abaxially asymmetrical and abapically orthogonally

truncated, flanked on either side by flattened spatulate

horizontal expansions produced from the abapical tips

of the inner and outer lips; exhalant channel deeper,

limited by parallel lips; about 4 mm width weakly

concave channel between terminal ridge and first

columellar tooth, followed by 13-14 stout, short

columellar teeth, extending a short distance onto the

ventrum and into the aperture; terminal ridge obsolète,

merged into smooth abapical edge of fossula; fossula

small, concave, smooth; labral teeth heavy, 14-16 in

number, evenly distributed. Ventral and dorsal zones

spotted, marginal callus lighter coloured with larger

ocellated spots.

Shell formula. 39.4 (76.9 - 58.2) 16: 15

specimen

collection number

length

width

height

BL coll. 1; (Figs 5-6)

47.2

36.9

26.9

BL coll. 2; (Figs 1-4)

49.7

37.1

29.8

BL coll. 3; (Figs 7-8)

44.6

25.2

BL coll. 4

42.9

33.3

25.2

BL coll. 5

37.8

21.5

BL coll. 6

37.1

29.1

20.4

BL coll. 7

32.7

24.7

20.5

BL coll. 8

24.5

19.1

(Figs 9-12)

LACMIP 13648

30.4

23.3

17.7

Table 1. Dimensions and number of teeth; Muracypraea hyaena (Schilder, 1939).

Discussion. Muracypraea hyaena (Schilder, 1939)

(holotype; Figs 13-14) was described from younger

late Miocene beds in the Urumaco area, Falcôn Dept.,

Venezuela (Sânchez-Villagra & Aguilera, 2006). The

holotype is in very poor condition, but the general

shape, strength of the dorsal hump, disposition of the

teeth, and colour markings are consistent. There is

some variability as to the presence or absence of

tubercles on the dorsal hump; the tubercles usually

présent only in fully grown specimens, although our

largest specimen (Figs 1-4) is devoid of tubercles and

the smallest (Figs 9-12) has well developed little

tubercles.

Perilliat (1992) figured a fairly well preserved

specimen identified as Siphocypraea (Muracvpaea)

sp. cf. S. (M.) angustirima hyaena Schilder from the

middle Miocene Ferrotepec Formation, Michoacân,

Southern Mexico. This specimen is indeed a

Muracypraea somewhat similar to M. hyaena but has

coarser dentition and has a more produced anterior.

The Ferrotepec Formation shell is more likely an

undescribed species.

Geological and geographical distribution.

Lower Miocene: Cantaure Formation, Venezuela.

Middle Miocene: Urumaco, Venezuela (Schilder,

1939).

Muracypraea woodringi n. sp.

Figs 21-28

1951 Cypraea cf. C. henekeni Sowerby - Marks, p.

376.

1959 Cypraea (Muracypraea) henekeni Sowerby -

Woodring, p. 194, pl. 32, figs 1,4, 6, 9.

1964 Siphocypraea (Muracypraea) henekeni

(Sowerby) - Olsson, p. I 76, pl. 31, figs 3-3a.

1993 Cypraea (Muracypraea) henekeni Sowerby-

Pit&Pit, p. 3, pl. l.fig. 7.

1997 Muracypraea “henekeni” (Sowerby, 1850) —

Groves, p. 152, pl. 1, figs 4-6.

Figures 1-14

1-4. Muracypraea hyaena (Schilder, 1939) NHMW 2010/0036/0024 (NMMW; ex BL coll. no. 2), length 47.7

mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ

Falcon Stale ' Venezuela; 5-6. Muracypraea hyaena (Schilder, 1939) NHMW 2010/0036/0025

MM f „ e *' B X L C0lL no - len § th 47 - 2 mm - lower shel1 bed , 1 km Southwest of Casa Cantaure, about 10 km

i'q r o ?\iu'viu? -in PcnmsL1 ' a - Fa ' cbn State, Venezuela; 7-8. Muracypraea hyaena (Schilder,

fr ,, " 1 J6/0026 (NMMW; ex. BL coll.no. 3), length 44.6 mm. lower shell bed, 1 km Southwest

r asa antaure, about 10 km west of Pueblo Nuevo. Paraguanâ Peninsula, Falcôn State. Venezuela; 9-12.

she iThTe hyüena (Schllder ’ ,939) LACMIP '3648 (LACMIP coll.; ex BL coll.). length 30.4 mm. lower

State S0L ‘ 1 ^ st ° ( asa Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ Peninsula, Falcôn

31 0 mm nï .^cypraea hyaena (Schilder, 1939) Holotype NMB H11272 (NMB coll.), length

.0 mm. Urumaco area, Falcôn Dept., Venezuela, Urumaco Formation, early late Miocene.

B. M. Landau & L. T. Groves

Novapex 12(1-2): 1-38, 10 mars 2011

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

Dimensions and type material. Holotype; LACMIP

12431, length 42.1 mm (Figs 23-24); paratype I

ANSP 315087, length, 42.0 mm (Figs 21-22);

paratype 2 USNM 562582, length, 38.5 mm, paratype

3 NHMW 2010/0036/0001 (ex. BL coll.) length 43.4

mm (figs. 25-28).

Other Material. Panama, three incomplète specimens

PPP 02168 (NMB 18667); three incomplète

specimens PPP 02119 (NMB 17637); two specimens

PPP 00490 (NMB 17871); four specimens PPP 00487

(NMB 17868); one specimen PPP 1080 (NMB

18261); one specimen PPP 00224 (NMB 17642); two

specimens PPP 01034 (NMB 18258); one incomplète

specimen PPP 01078 (NMB 18325) ail NMB coll.;

one specimen locality ANSP 1731 (internai mould);

one specimen locality LACMIP 16936; five

specimens locality LACMIP 17908 (= TU 757); five

specimens locality LACMIP 17909 (= TU 958); two

specimens locality LACMIP 17910 (= TU 960); two

specimens locality LACMIP 17911 (= TU 1432); one

specimen from LACMIP 17912 (= TU 1433); Cativa,

Colon, BL coll. (Ail Gatun Formation); Colombia, one

specimen locality UCMP 154008 (fragment), Narino

Dept.; Ecuador, one incomplète specimen PPP 03391

(NMB 19122); one specimen AGC 99-102, NMB

coll.; Baja California Sur, Mexico, one specimen

locality USNM 1/1238 (ex USGS), San Ignacio

Formation!?).

Type locality. Angostura Formation (late Miocene),

Loc. LACMIP 16943 [= TU 1507], just east of Rio

Verde, approximately 30 km east of Rio Esmeraldas

Esmeraldas Province, Ecuador.

Type stratum. Angostura Formation (late Miocene).

Description. Shell medium-sized for genus, solid,

triangular-shaped, dorsum very strongly humped 2 / 3

distance from the anterior end, bearing two groups of

small warty tubercles, left group larger and slightly

anterior to right; spike-like projection at centre of apex

rises 2 mm above dorsal surface; spire depressed,

covered by adapical callus; sides rounded, strongly

callused, with the callus ascending progressively

adapically to just below the apex posterior to the

dorsal hump, callus edge poorly delimited; ventrum

flattened, slightly convex in profile; aperture almost

straight, anterior third weakly dilated; anterior canal

deep, abaxially asymmetrical and abapically

orthogonally truncated, flanked on either side by

flattened spatulate horizontal expansions produced

from the abapical tips of the inner and outer lips;

posterior channel deeper, limited by parallel lips;

about 4 mm width weakly concave channel between

terminal ridge and first columellar tooth, followed by

14 stout, becoming weak posteriorly, short columellar

teeth, extending a short distance onto the ventrum and

into the aperture; terminal ridge obsolète, merged into

smooth abapical edge of fossula; fossula small,

concave, smooth; labral teeth heavy, 14 in number,

evenly distributed, becoming weak posteriorly.

Ventral and dorsal zones striped, marginal callus

lighter coloured, dorsum with irregular stripes of dull

red-orange and tan.

Shell Formula. 45.3 (80.0-60.0) 21; 14

specimen

collection number

length

width

Height

holotype (Figs 23-24)

LACMIP 12431

43.5

34.5

27.1

paratype 1 (Figs 21-22)

ANSP 315087

42.1

34.1

24.6

paratype 2

USNM 562582

46.0

36.9

24.3

paratype 3 (Figs 25-28)

NHMW 2010/0036/0001

43.4

31.1

25.6

CAS 66589.06

37.5

29.7

22.6

USNM 562581

52.7

44.0

31.8

BL coll. 1

38.4

31.6

21.4

BL coll. 2

42.4

35.1

23.1

BL coll. 3

57.6

45.1

31.8

BL coll. 4

48.8

41.0

28.5

BL coll. 5

46.6

35.5

26.7

DF B coll.

No. 7406-1

48.5

39.8

26.3

DFB coll.

No. 7406-2

38.8

32.7

22.1

Table 2. Dimensions and number ot teeth; Muracypraea woodringi n. sp.

Discussion. Muracypraea woodringi is described

the forms noted by Groves (1997) as M. “heneke

hom the Gatun Formation of Panama, the Angosl

Formation of Ecuador, and Miocene strata

southwestern Colombia. A poorly preser

specimen from Southern Baja California Sur, Mex

may also represent M. woodringi. Mnracypn

woodringi is most similar to M. isthm

(Schilder, 1927) [= Cypraea henekeni Sowb. var

Brown & Pilsbry (1911:356-357, pi. 26, figs. 9-10)]

also from the Gatun Formation, which has a broad,

smooth rounded, poorly delimited dorsal gibbosity

centrally positioned (Figs 23-24). These shells were

illustrated by Woodring (1959; pi. 31, figs 6-10). The

specimen illustrated by Brown & Pilsbry (1911: pi-

26, figs. 9-10) clearly has a single dorsal gibbosity.

Untortunately, the holotype is missing. Brown &

Pilsbry (191 1: 356) noted that the “aperture is like that

B. M. Landau & L. T. Groves

NOVAPEX 12(1-2): 1-38, 10 mars 2011

of Santo Domingo C. henikeni [sic], except that the

teeth are more compressed and longer. In a specimen

42.5 mm. long there are 15 teeth on the inner, 19 on

the outer lip.” The length of the denticles seems

variable, but on average M. isthmicù has fewer

denticles on both the inner and outer lips than M.

henekeni or M. hyaena. Woodring (1959: pl. 32. figs.

1, 4, 6, 9) illustrated specimens that are considered

true M. woodringi. The two Gatun Formation taxa

cannot be separated by their dentition or fossula.

Landau and Silva (2010) discussed the presence of the

genus in the Venezuelan Pliocène assemblages and

stressed the need for a full révision of the genus. This

much needed révision is again outside the scope of

this worlc, however, at least half a dozen different

Muracypraea species may occur in the tropical

American Neogene.

Etymology. Named for Wendell P. Woodring in

récognition of his monumental work on the geology

and palaeontology of the Panama Canal Zone région.

Geologieal and geographical distribution.

Upper Miocene: Angostura Formation, Ecuador;

Gatun Fonnation, Panama.

Miocene (indeterminate): Narino Dept., Colombia.

Subfamily LUR1FNAE Schilder, 1932

Genus Luria Jousseaume, 1884

Type species: Cypraea lurida Linnaeus, 1758, by

original désignation.

Discussion. Dolin (1991) synonymised the

genera/subgenera Tessellata Jousseaume, 1884 (type

species Cypraea tessellata Swainson, 1822),

Basilitrona Iredale, 1930 (type species Cypraea

isabella Linnaeus, 1758) and Fossacypraea Schilder,

1939 (type species Cypraea hieroglyphica Schilder,

1923) with Talparia Troschel, 1863. Lorenz & Hubert

(2000) considered “ Basilitrona” and “ Tessellata" as

species groups within the genus Luria Jousseaume,

1884. Dolin & Lozouet (2004) transferred the tropical

American species included in Talparia by Dolin

(1991) to Luria (Tessellata). Meyer’s (2004)

molecular data supports the systematic arrangement of

Lorenz & Flubert (2000), on the basis of which Lôpez

Soriano (2006) fonnally synonymised Tessellata with

Luria. Shells in the genus Luria differ from those in

Talparia in having blunt extremities, short and deep

anterior and posterior canals, the anterior canal is

straight as opposed to slightly off-set in Talparia, and

the angulation is more prominent with the labral teeth

developed at the angulation running into the aperture

but not onto the ventrum. In Talparia the angulation is

developed and raised in the anterior portion but not in

the mid and posterior portion with the teeth extending

a short distance onto the ventrum. The character of the

fossula; spoon-like, smooth or weakly ribbed, with

denticles on the inner margin, is similar in both

généra.

Luria cantaurana n. sp.

Text-figure 1; Figs. 37-44

Dimensions and type material. Holotype; NHMW

2010/0036/0002, lenght 56.0 mm (Figs 37-40);

paratype 1 NHMW 2010/0036/0003, length, 42.0 mm

(Figs 41-44); paratype 2 NHMW 2010/0036/0004,

length, 38.5 mm.

Type locality. lower sheli bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Falcôn State, Venezuela (=locaiity GS12PGNA of

Gibson-Smith & Gibson-Smith, 1979).

Type stratum. Cantaure Formation (early Miocene:

Burdigalian).

Description. Shell medium-sized for genus, solid,

subcylindrical, posteriorly swollen, in latéral profile

dorsal curvature regularly attenuated abapically, spire

involute, covered by adapical callus; sides rounded,

callused, the callus extending to between Ft, and %

height; ventrum flattened, especially in the anterior

portion, slightly convex in profile in the posterior

portion; aperture parasigmoidal, anterior third wider,

dilated by hemispherical expansion on the labrum;

siphonal canal deep, abaxially asymmetrical and

abapically orthogonally truncated; exhalant channel

deep, limited by parallel lips; 28-33 very short

columellar teeth rnost clearly developed at the

angulation, running a short distance into the aperture,

but not onto the ventrum; angulation Sharp and ridge-

lilce in the anterior and mid-portions; terminal ridge

weakly developed, merged into abapical edge; fossula

well-delimited, spoon-like, concave, broad, smooth,

fringed with weak denticles at its inner edge;

columella peristome smooth; 24-29 labral teeth,

anterior third of labral teeth stretched out on

constricted, depressed, hemicircular area; remai nder

sharp, outer lip bevelled inwards, with teeth extending

across the bevelled inner portion and a very short

distance externally onto the labrum; no colour pattern

preserved, under UV light some axially elongated

blotches seen in two specimens, but no clear pattern.

Shell Formula. 45.5 (58.0 - 45.7) 20: 24

specimen

collection number

length

Width

Height

holotype (Figs 37-40)

NHMW 2010/0036/0002

56.0

33.3

25.6

paratype 1 (Figs 41-44)

NHMW 2010/0036/0003

42.0

23.8

18.9

paratype 2

NHMW 2010/0036/0004

38.5

22.1

17.9

Table 3. Dimensions and number of teeth; Luria cantaurana n. sp.

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

Text-figure 1 . Luria cantaurana n. sp. Holotype

NHMW 2010/0036/0002 (NMMW coll.; ex. BL coll.),

length 56.0 mm. lower shell bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Paraguanâ Peninsula, Falcôn State, Venezuela.

Discussion. Luria cantaurana n. sp. is by far the

largest Luria species of the Cantaure Formation. As

can be seen from the shell formulae, it differs from L.

fossula (shell formula; 43.0 (60.4 - 48.8) 22: 16) in

having significantly fewer columellar teeth, whereas

L. dominicensis (shell formula; 39.5 (59.4 -48.1) 30:

25) has a similar number of columellar teeth, but a far

greater number of labral teeth. The early Miocene L.

dockeryi from the Chipola Formation of Florida

cannot be separated from L. dominicensis on the basis

ofthe shell formula (see below).

Dolin (1991) synonymised Cypraea fossula

Ingram, 1947 with Cypraea dominicensis Gabb, 1873.

However, Cypraea fossula belongs within the genus

Trôna (see below). Luria dominicensis (LT 36, CT 29,

Pilsbry, 1922; shell formula 39.5 (58.7 - 48.1) 30: 24),

from an unknown locality in the Dominican

Republic). The species from the early Miocene

Chipola Formation of Florida originally identified as

Luria dominicensis by Dolin (1991) was renamed

Luria (Tessellata) dockeryi Dolin & Lozouet, 2004.

Luria dockeryi has numerous short teeth (LT 36, CT

29, Dolin, 1991; shell formula 39.5 (59.4 - 48.1) 30:

25). The shell formula is identical to that of the

holotype of Luria dominicensis. Unfortunately the

holotype (ANSP 3003) illustrated by Pilsbry (1922)

cannot be located. Dolin & Lozouet (2004) renamed

the Chipola species without mentioning how it

differed from the Dominican taxon. There is little to

distinguish the two, from the scant material available,

and they may be synonyms.

Jousseawnea joossi Schilder, 1939 (p. 23, fig. 23)

was synonymised with Luria dominicensis by Dolin

(1991). The holotype is an internai cast from the

Miocene of Trinidad representing an indéterminable

Luria sp., and it should be considered nomen dubium.

Etymology. Named for the Cantaure Formation.

Geological and geographical distribution.

Lower Miocene: Cantaure Formation, Venezuela.

Luria isabellaprimitiva n. sp.

Text-figure 2; Figs 45-53

1965 Cypraea aff. isabella Linné - Jung, p. 501,

pl. 67, figs 7-8.

Dimensions and type material. Holotype; NHMW

2010/0036/0005, length 27.7 mm (Figs 45-49);

paratype 1 NHMW 2010/0036/0006, length, 26.6 mm

(Figs 50-53); paratype 2 NHMW 2010/0036/0007,

length, 26.4 mm.

Other material. 4 specimens BL coll.

Type locality. lower shell bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Falcôn State, Venezuela (=locality GS12PGNA of

Gibson-Smith & Gibson-Smith, 1979).

Type stratum. Cantaure Formation (early Miocene:

Burdigalian).

Figures 15-28

15-17. Muracypraea henekeni (G. B. Sowerby I, 1850) NHMW 2010/0036/0027 (NMMW; ex. BL coll.), length

is TU1219 ’ lower Gurabo Formation, early Pliocène, Gurabo River, Dominican Republic-

l r W ( , G - B - S ° Werby 1 1850) NHMW 2010/0036/0028 (NMMW; ex. BL coll.), length

, ,, TU1219 ’ lower Gurabo Formation, early Pliocène, Gurabo River, Dominican Republic-

(middi euZTTr " n sp ' Paraty P e 1 ANSP 3 15087, length 42.1 mm. Lower Gatun Formation

sp Holotvne I TcMtp'mLu? ?" 'uT-iT’ Panama (P hoto Lindsey Groves); 23-24. Muracypraea woodringi n.

TU 1 5071^,.^ CM1P I243 F' englh 4 ~- m m. Angostura Formation (upper Miocene), Loc. LACM1P 16943 [=

(Photo 1LindS'Crlv5^s ie ; approx,mately ^ 30 k m east of Rio Esmeraldas Esmeraldas Province. Ecuador

ex BL coll 1 l?à 25 ’ 28 ; Murac yP raea woodringi n. sp. Paratype 3 NHMW 2010/0036/0001 (NMMW;

Boyd-Roo evélt Hthw mm t L °T r GatUn Formation < middle Miocene), Loc TU 961, roadcuts both sides of

yd Roosevelt H.ghway, just east of Cativa (= Woodring locality no. 138e), Colon Province, Panama.

B. M. Landau & L. T. Groves

NOVAPEX 12(1-2): 1-38, 10 mars 2011

r «V

iRSJ

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

Description. Shell small for genus, solid,

subcylindrical, slightly posteriorly swollen, in latéral

profile dorsal curvature regularly attenuated

abapically, spire involute, covered by adapical callus;

sides rounded, callused, the callus extending to l A

height; ventrum flattened, very slightly convex in

profile at the extremities; aperture straight, weakly

curved to the left in the posterior portion, anteriorly

somewhat dilated by hemispherical expansion on the

labrum; siphonal canal deep and narrow, abaxially

asymmetrical and abapically orthogonally truncated;

exhalant channel deep and narrow, limited by parailel

lips; 25-30 very short columellar teeth running deep

into the aperture from the angulation across the

columellar peristome, but not onto the ventrum;

angulation relatively sharp; terminal ridge weakly

developed, merged into abapical edge; fossula well-

delimited from the columellar peristome by an oblique

ridge, spoon-like, concave, strongly ridged, ridges

ending as denticles at its inner edge; columella

peristome very steep, ridged; 20-25 very short, sharp,

labral teeth, abapically teeth not stretched out onto

constricted, depressed, hemicircular area; teeth

extending across inner portion of lip, but not

extemally onto the labrum; colour pattern indicating

preserved spots on margins and rings on the dorsum

(Fig. 49).

Shell Formula. 25.8 (64.7 - 51.6) 27: 25

Specimen

collection number

Length

Width

height

holotype (Figs 45-49)

NHMW 2010/0036/0005

27.7

17.0

14.2

paratype 1 (Figs 50-53)

NHMW 2010/0036/0006

26.6

16.9

13.4

paratype 2

NHMW 2010/0036/0007

26.4

17.2

14.1

BL coll. 4

28.8

17.6

14.2

BL coll. 5

27.9

17.2

12.8

BL coll. 6

26.3

15.9

13.1

BL coll. 7

22.1

15.0

11.4

Table 4. Dimensions and number of teeth; Luria isabellaprimitiva n. sp.

Text-figure 2. Luria isabellaprimitiva n. sp. Holotype

NHMW 2010/0036/0005 (NMMW coll; ex. BL coll.),

length 27.7 mm. lower shell bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Paraguanâ Peninsula, Falcôn State, Venezuela.

Discussion. Luria isabellaprimitiva n. sp. differs from

its Cantaure Formation congener L. cantaurana n. sp.

in having a smaller, more subcylindrical shell, with

shorter extremities, and a smaller, deeper fossula. The

colour pattern is somewhat unusual in consisting of

circles, similar to that seen in the Recent Arestorides

argus (Linnaeus, 1758). This is not a colour pattern

présent in any living or fossil species of Luria. The

ocellated spots are not very clear and only visible on

part of the holotype, and not on any of the paratypes.

If this pattern were found on more specimens

placement within the genus Arestorides Iredale, 1930

might be supported, although there is no record so far

of this genus in the American Neogene to Recent

faunas.

Figures 29-44

29-32. Muracypraea grahami (Ingram, 1947) NHMW 2010/0036/0029 (NMMW; ex. BL coll.), length 63.2

taya Formation, Cubagua Group, early Pliocène, Canon de las Calderas, Cubagua Island, Nueva Esparta State,

Venezuela; 33-36. Troua trinitatensis (Mansfield, 1925). UCMP 35536, holotype of Cypraea fossula Ingram,

p engt p mm - 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ

tNMvrw’ t a c D ° i n State ’ Venezuela; 37-40. Luria cantaurana n. sp. Holotype NHMW 2010/0036/0002

fp ,, coll.), length 56.0 mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west

NHMW^nuwînumAnTH^ Peninsula, Falcôn State, Venezuela; 41-44. Luria cantaurana n. sp. Paratype 1

C™" 2 T , , „ (NM „ MW; «• BL co,1 -)> length 42.0 mm. lower Shell bed, I km Southwest ofCasa

. out 10 km west of Pueblo Nuevo. Paraguanâ Peninsula, Falcbn State, Venezuela.

B. M. Landau & L. T. Groves

NOVAPEX 12(1-2): 1-38, 10 mars 2011

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

In the character of its subcylindrical shape and very

short extremities Jung (1965) was correct to suggest

that L. isabellaprimitiva belonged within the L.

isabella- group. This is represented in the tropical

American fossil assemblages by L. patrespatriae

(Maury. 1917) (holotype; Fig. 146), which is found in

the Lower Pliocène Gurabo Formation of the

Dominican Republic. Schilder (1939) recorded L.

patrespatriae in the Cantaure Formation. Pilsbry

(1922) synonymised this taxon with the living L.

isabella (Linnaeus, 1758), which today is widespread

in the Indo-Pacific (Lorenz & Flubert, 2000). Ingram

(1947) reaffirmed the synonymy whilst Woodring

(1928) considered it a subspecies of L. isabella.

We hâve compared specimens of L. patrespatriae

from the early Pliocène Gurabo Formation of the

Dominican Republic with specimens L. isabella from

the Red Sea (BL coll.). Unfortunately, the Dominican

shells are fragile, including the holotype, and in

almost ail our specimens the columella is pushed

inwards and the teeth are uncountable. We hâve one

undamaged specimen in which the apertural characters

are well preserved (Figs 54-57). The shell formula for

L. patrespatriae [32.4 (54.0 - 44.0) 32: 29] is almost

identical to that of L. isabella [35 (52.9 - 44.0) 34:

29], One specimen from the Dominican Republic has

the colour pattern preserved, which shows the same

orange coloured tips seen in Recent L. isabella , but no

black striping. On ventral view the columella is

slightly more sinuous in L. patrespatriae and the

terminais, especially the posterior terminal, is slightly

more produced. The relationship between these

species within the L. isabella group is unclear, we

therefore consider them distinct at full species level.

Luria isabella has a long géologie history in the Indo-

Pacific, first recorded from the early Miocene of Fiji

(Ladd, 1977) as Cypraea (Talparia) isabella

lekalekana Ladd, 1934. This taxon was synonymised

with L. isabella by Lorenz & Hubert (2000).

This species group is represented today in the

tropical American Pacific by Luria isabellamexicana

(Stearns, 1893), which differs from L. isabella in

having a more pyriform, less subcylindrical shell

shape, fewer teeth on both inner and outer lips, more

protracted terminais and a somewhat different

colouration [shell formula for L. isabellamexicana

from île de Clipperton; ZMA coll. unnumbered lot;

39.7 (54.4 - 46.6) 29: 27], There are also radular

différences (Burgess, 1985, Lorenz, 2002).

It seems, therefore, that during the Pliocène L.

patrespatriae was présent in the Caribbean portion of

the Neogene Gatunian Province and L. isabella in the

Indo-Pacific. Following the closure of the Central

American isthmus, the L, isabel la-group disappeared

from the Caribbean, and its populations in the Pacific

became fragmented into a questionable number of

Indo-Pacific species/subspecies with restricted

géographie distributions (see Lorenz & Hubert, 2000;

Lorenz, 2002) and L. isabellamexicana restricted to

the tropical American Pacific, which is

unquestionably a valid taxon, lnterestingly, L.

isabellamexicana is présent and common in the Lower

Pleistocene Moin Formation of Costa Rica (BL

collection). At this time connections between the

Pacific and Caribbean, albeit intermittent, still existed

and is an example of a paciphile species (see Landau

et al., 2009). The L. isabella- group has not so far been

found in the neighbouring northem Atlantic

Caloosahatchian Province.

The early Miocene shells from the Cantaure

Formation were synonymised with L. patrespatriae by

Schilder (1939). However, the two populations are

different; the Cantaure shells are never as cylindrical

or elongated as those of L. isabella , and the teeth are

far less numerous and stronger than in L. isabella (see

shell formulae above). The Cantaure Formation shells

are doser in shape to those of L. isabellamexicana,

but differ in having less protracted terminais and again

fewer columellar and labral teeth. We therefore

consider the Cantaure shells a distinct species L.

isabellaprimitiva n. sp.

The genus Luria has also been recorded from the

mid-Atlantic middle Miocene Madeira Archipelago

(Lorenz & Groh, 1998). The shell of Luria santoensis

Lorenz & Groh, 1998 is not unlike that of L.

isabellaprimitiva in general shape, but with relatively

tewer columellar teeth, a wider aperture and smooth

fossula. Indeed, in the character of the fossula L.

isabellaprimitiva does not conform well within the

genus, which is characterised by shells with a smooth

fossula, or almost so.

Etymology. The name indicates an early member of

the L. isabella species-group.

Geological and geographical distribution.

Lower Miocene: Cantaure Formation, Venezuela.

Figures 45-57

45-49. Luria isabellaprimitiva n. sp. Holotype NHMW 2010/0036/0005 (NMMW; ex. BL coll.), length 27.7

mm. ower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ

n," S 'f, tC ’ Vene/ucla; 5 °- 53 - Luria isabellaprimitiva n. sp. Paratype 1 NHMW 2010/0036/0006

nf p ki \f[ X n° ’ ’ 26-6 mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west

Ot 1 ueblo Nuevo, Paraguanâ Peninsula, Falcôn State, Venezuela;

Tim'is ' M . na pa ' res P“ tr L ae ( Maur y, 1917) NHMW 2010/0036/0030 (NMMW; ex. BL coll.), length 30.0 mm.

l U1215, lower Gurabo Formation, early Pliocène, Gurabo River, Dominican Republic

B. M. Landau & L. T. Groves

Novapex 12(1-2): 1-38, 10 mars 2011

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

Genus Trôna .lousseaume, 1884

Type species: Cypraea stercoraria Linnaeus, 1758, by

original désignation.

Discussion. The généra Trôna and Macrocypraea

Schilder, 1930, traditionally considered to be closely

related and within the Cypraeinae (Lorenz & Hubert,

2000) were shown by Meyer (2004) to belong to

different subfamilies; Trôna within the Luriinae and

Macrocypraea within the Cypraeinae. The shell

characteristics of the two généra are quite different.

Macrocypraea has a rather light shell, as opposed to

the very heavy shell of Trôna. The shell of

Macrocypraea is elongate rather than globose, they

rarely hâve a marginal callus as opposed to the heavy

callus seen in most Trôna species, the base is far less

flattened than in Trôna , the apertures are wide with

spiny extremities, not seen in Trôna, and the shape of

the anterior expansion of the aperture is quite

different. The fossula in Macrocypraea is not as large

or spoon-shaped, nor as strongly sculptured as in

Trôna and the terminal ridge is much more prominent

in Macrocypraea.

The genus Trôna was previously represented by a

single subspecies in the Western Hemisphere. Dolin

(1991) described Trôna leporina calhounensis from

the Chipola Formation of Florida based on two

specimens. Trôna is also a characteristic component

within the European Miocene cypraeid fauna. Trôna

leporina (Lamarck, 1810) from contemporary late

early Miocene (Burdigalian) deposits in France

(Moulin de Cabannes, St Paul-les-Dax, Landes,

France; BL coll.) differs from T. leporina

calhounensis in having a less basally flattened shell, a

thicker marginal callus and thicker anterior channel

callus, a more rounded inner lip, a poorly developed

basal angulation with narrow adapical denticles,

fossula sculptured with altemate interrupted ridges

and rows of pustules, more rounded outer lip, and

shorter adapical denticles. Specimens from the early

Burdigalian of Le Peloua, France (BL coll), seem

slightly different, more ovate with rows of very small

pustules on the fossula. Dolin (1991) commented on

these small différences in fossular sculpture between

populations and postulated these may be an expression

of ecophenotypic variation or reproductive isolation

and opted to name the Chipola Formation shells T.

leporina calhounensis a subspecies of the European

taxon without implying a close phylogenetic link

between the two populations. This approach is

confusing and in view of the taxonomie problems still

to be resolved within the European populations and

the presence of additional species in the Caribbean

Neogene we consider this taxon distinct at full spécifie

rank. Trôna trinitatensis (Mansfield, 1925) is found

in the early and middle Miocene of Venezuela,

Trinidad, and Carriacou, Grenadine Islands, Lesser

Antilles. Trôna trinitatensis differs from T. ingrami

and T. calhounensis in having a smaller, more

elongated shell, in having much less sinuous and

narrower aperture, and somewhat more crowded teeth

on both labral and columellar sides (see below under

Trôna trinitatensis)

Trôna ingranti n. sp.

Text-figure 3; Figs 58-70

Dimensions and type material. Holotype; NMB

H18402, height 53.1 mm (Figs 58-60); paratype 1

NHMW 2010/0036/0008, height, 48.2 mm (Figs 61-

64); paratype 2 NHMW 2010/0036/0009, height, 47.2

mm (Figs 65-68); paratype 3 NHMW

2010/0036/0010, height, 49.7 mm (Figs 69-70).

Other material. 3 specimens NMB coll.; 18

specimens BL coll.; 1 specimen DFB, no. 6578.

Type locality. lower shell bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Falcôn State, Venezuela (=locality GSI2PGNA of

Gibson-Smith & Gibson-Smith, 1979).

Type stratum. Cantaure Formation (early Miocene:

Burdigalian).

Figures 58-77

58-6°. Trôna ingrami n. sp. Holotype NMB H18402 (NMB coll.), length 53.1 mm. lower shell bed, 1 km

Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ Peninsula, Falcôn State, Venezuela;

61-64. Trôna ingrami n. sp. Paratype 1 NHMW 2010/0036/0008 (NMMW; ex. BL coll. no. 9), length 48.2 mm.

ower shell bed 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ Peninsula,

a con 8tate, Venezuela; 65-68. Trôna ingrami n. sp. Paratype 2 NHMW 2010/0036/0009 (NMMW; ex. BL

co . no. 10), length 47.2 mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo

?nm/?trr!ailnn,77? U1 a ' Fa ' c ° n State > Venezuela; 69-70. Trôna ingrami n. sp. Paratype 3 NHMW

c . . , | ex - BL colb no - 15 )> length 49.7 mm. lower shell bed, 1 km Southwest of Casa

V , le ', a ’ l | t - km West of Puebl ° Nuevo, Paraguanâ Peninsula, Falcôn State, Venezuela; 71-73. Trôna

Casa Tanta'' a J^ s Ie d» 1925) NMB H18403 (NMB coll.), length 33.3 mm. lower shell bed, 1 km Southwest of

tri ni tnt rn, .P 7 West of Pllebl ° Nuevo, Paraguanâ Peninsula, Falcôn State, Venezuela; 74-77. Trôna

LTfr o m"" ! ÎV 925 P NHMW 2010/0036/0011 (NMMW; ex. BL coll.), length 33.0 mm. lower shell

Venezuela Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ Peninsula, Falcôn State,

B. M. Landau & L. T. Groves

NOVAPEX 12(1-2): 1-38, 10 mars 2011

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

Description. Shell medium-sized for genus, solid,

subcylindrical to rotund, posteriorly swollen, in latéral

profile dorsal curvature regularly attenuated

abapically, spire weakly projecting, covered by

adapical callus; sides rounded, weakly callused;

ventrum flattened, slighlly convex in profile; aperture

parasigmoidal, anterior third dilated by hemispherical

expansion; siphonal canal deep. abaxially

asymmetrical and abapically orthogonally truncated;

exhalant channel deeper, limited by parallel lips; four

or five anterior columellar teeth, most anterior

thinnest, strengthening adapically; other numerous

inner teeth parallel, extending deep within the

aperture, bituberculate, about 6 mm in length in the

columellar area, abrupt, extending a short distance

over the angulation of the basal and columellar planes

onto the ventrum where they are slightly coarser;

terminal ridge obsolète, merged into abapical edge,

fringed with denticles from fossula on inner margin;

fossula deep, concave, spoon-like, covered by narrow

ridges, sometimes interrupted, joining fossular

denticulation to anterior columellar teeth; 22-30 labral

teeth, anterior third of labral teeth stretched out on

constricted, depressed, hemicircular area; remainder

heavier, outer lip bevelled inwards, with teeth

extending across the bevelled inner portion, 3-4 ram in

length; ventral and dorsal zones spotted, marginal

callus lighter coloured, under UV light spire blotch

présent.

Shell Formula. 48.7 (63.0 - 50.1 ) 20: 18

Specimen

collection number

length

width

height

holotype (Figs 58-60)

NMB H18402

53.1

34.5

NMB lot 17516

35.3

22.4

18.2

NMB lot 17516

35.7

18.7

NMB lot 17516

26.8

paratype 1 (Figs 61-64)

NHMW 2010/0036/0008

48.2

32.6

25.7

paratype 2 (Figs 65-68)

NHMW 2010/0036/0009

47.2

29.7

23.2

paratype 3 (Figs 69-70)

NHMW 2010/0036/0010

49.7

30.4

26.3

BL coll. 1

57.9

36.7

28.3

BL coll. 2

56.1

34.6

26.9

BL coll. 3

55.8

35.4

28.2

BL coll. 4

58.9

36.3

29.7

BL coll. 5

57.4

39.4

31.1

BL coll. 6

55.9

34.3

27.6

BL coll. 7

55.6

29.7

BL coll. 8

60.2

37.5

29.4

BL coll. 1 1

48.7

30.9

25.1

BL coll. 12

47.8

31.8

BL coll. 13

45.1

30.6

25 3

BL coll. 14

44.2

28.2

BL coll. 16

44.8

27.2

20.9

dL coll. 1 7

51.5

31.4

25 2

BL coll. 18

49.5

32.8

25 3

BL coll. 19

40.1

24.3

18 9

BL coll. 20

37.1

17 3

If)

BL coll. subadult 21

DFB coll.

no. 6578

39.5

55.1

25.1

38.4

18.8

30.4

Table 5. Dimensions and number of teeth; Trôna ingrami n. sp

Text-figure 3 (opposite). Trôna ingrami n sp

Paratype 1 NHMW 2010/0036/0008 (NMMW coll.;

ex. BL coll. no. 9), length 48.2 mm. lower shell bed, 1

km Southwest of Casa Cantaure, about 10 km west of

Pueblo Nuevo, Paraguanâ Peninsula, Falcôn State

Venezuela ’

B. M. Landau & L. T. Groves

Novapex 12(1-2): 1-38, 10 mars 2011

Discussion. Trôna ingrami n. sp. has a rather variable

shell shape. The larger specimens tend to be more

rotund and broad (Figs 58-64), whereas the smaller

specimens hâve a more elongated aspect and a less

sinuous aperture, but intermediate specimens also

occur (see Text-figure 4). The single living

représentative of the genus Trôna stercoraria

(Linnaeus, 1758) distributed today along the coast of

tropical West Africa also has a variable shell shape

(Lorenz & Hubert, 2000; pl. 11 ). Dolin ( 1991 ) hints at

sexual dimorphism in the species without elaborating,

however, in Cypraeidae as well as in Ovulidae male

shells tend to be smaller than females.

height/length Trôna ingrami

♦ length/width

■ length/height

-Linear (length/height)

Linear (length/width)

30 35 40 45 50 55 60 65

length

Text-figure 4. Height/ Length; Trôna ingrami n. sp.

Dolin (1991) describes a light-weight shell with

uncallused margins, whereas the Cantaure shells are

very solid with a weakly developed marginal callus.

His description and figure clearly show the labral teeth

to be restricted to the inside of the aperture, whereas

the teeth in T. ingrami extend a short distance over the

basal plane onto the ventrum where they become

somewhat coarser. On the labral side the teeth

abapically are described as short, whereas in T.

ingrami the outer iip is bevelled inwards with the teeth

extending across the bevelled inner portion. The

fossula in T. ingrami has few elongated denticles

which tend to be continuous and rarely tubercles as

compared to the specimens of T. calhounensis figured

by Dolin (1991) (fig. 22a) which has the fossula

crowded with interrupted ridges and elongated

tubercles.

Etymology. Named for William Marcus Ingram in

récognition of his pioneering work on Caribbean

Neogene cypraeids.

Geological and geographical distribution.

Lower Miocene: Cantaure Formation, Venezuela.

Trôna trinitatensis (Mansfield, 1925)

Text-figure 5; Figs 71-77

1925 Cypraea trinitatensis Mansfield, p. 49, pl. 8,

fig. 10.

1939 Trôna (Macrocypraea) trinitatensis

Mansfield, 1925 - Schilder (partim?), p. 30, fig. 32.

1947 Cypraea fossula Ingram, p. 4 (128), pl. 1 (8),

fig. 3.

1971 Macrocypraea cervinetta fossula Ingram,

1947 - Schilder & Schilder, p. 33.

1971 Macrocypraea trinitatensis Mansfield, 1925

- Schilder & Schilder (partim?), p. 164.

71971 Macrocypraea aff. zébra (Linné) - Jung, p.

181, pl. 8, figs 1-3.

1993 Macrocypraea trinitatensis (Mansfield, 1925)

- Lorenz & Herbert, pl. 121, fig. 3.

2000 Macrocypraea trinitatensis (Mansfield, 1925)

- Lorenz & Herbert, pl. 121, fig. 3.

2004 Luria (Tessellata) fossula (Ingram, 1947) -

Dolin & Lozouet, p. 20.

Material. 1 specimen NMB H18403, NMB coll. (Figs

71-73); 1 specimen NHMW 2010/0036/0011, NHMW

coll. (ex. BL coll.) (Figs 74-77). Holotype of C.

fossula ; UCMP 35536.

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

Description. Shell small for genus, solid,

subcylindrical, weakly posteriorly swollen, spire

projecting, covered by adapical callus; sides angular to

rounded, callused; ventrum almost fiat; aperture

weakly parasigmoidal, anterior third dilated by

hemispherical expansion; siphonal canal deep,

abaxially asymmetrical and abapically orthogonally

truncated; exhalant channel deeper, Iimited by paraliel

lips; four anterior columellar teeth; other numerous

inner teeth paraliel, extending deep within the

aperture, bituberculate, abrupt, not extending over the

angulation onto the ventrum; terminal ridge obsolète,

merged into abapical edge. fringed with denticles from

fossula on inner margin; fossula deep, concave,

spoon-like, covered by narrow ridges, sometimes

interrupted, joining fossular denticulation to anterior

columellar teeth; 22-27 labral teeth, anterior third of

labral teeth stretched out on constricted, depressed,

hemicircular area; remainder heavier, outer lip

bevelled inwards, with teeth extending across the

bevelled inner portion; no colour pattern preserved,

under UV light spots présent on marginal callus.

Shell Formula. 33.2 (55.8 - 44.0) 22: 20

specimen

collection number

length

Width

height

Figs 71-73

NMB II18403

33.3

18.8

15.3

Figs 74-77

NHMW 2010/0036/0011

33.0

18.3

13.9

Figs 33-36

UCMP 35536

43.0

26.0

21.0

Table 6. Dimensions and number of teeth; Trôna trinitatensis n. sp.

Text-figure 5. Trôna trinitatensis (Mansfïeld, 1925)

NMB H18403 (NMB coll.), length 33.3 mm. lower

shell bed, 1 km Southwest of Casa Cantaure, about 10

km west of Pueblo Nuevo, Paraguanâ Peninsula,

Falcôn State, Venezuela.

Discussion. The holotype of Trôna trinitatensis

(Mansfïeld. 1925) is an internai niould collected from

the Guaracara Limestone Member of the late early to

early middle Miocene Tamana Formation of Trinidad

and is difficult to identify with any certainty.

According to Jung (1971), ail the other specimens

from Trinidad recorded by Schilder (1939) are also

internai moulds. Schilder (1939) also identified

specimens from the middle Miocene Grand Bay

Formation of Carriacou as Trôna (Macrocypraea)

trinitatensis. According to Jung (1971) it was unlikely

that the Carriacou shells were conspecific with those

from Trinidad and that the larger Carriacou shells

were doser to the living Macrocypraea zébra.

According to Jung they differed ‘morphologically’

from the shells from T. trinitatensis from Trinidad, but

he did not specify what these différences were.

Figures 78-95

78-81. Propustularia longidentata n. sp. Flolotype NFIMW 2010/0036/0012 (NMMW; ex. BL coll.), length 15.2

mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ

Peninsula, Falcôn State, Venezuela; 82-85. Propustularia longidentata n. sp. Paratype 1 NHMW

2010/0036/0013 (NMMW; ex. BL coll.), length 14.5 mm. lower shell bed, 1 km Southwest of Casa Cantaure,

about 10 km west of Pueblo Nuevo, Paraguanâ Peninsula, Falcôn State, Venezuela; 86-87. Propustularia

longidentata n. sp. Paratype 2 NHMW 2010/0036/0014 (NMMW; ex. BL coll.), length 12.2 mm. lower shell

bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ Peninsula, Falcôn State,

Venezuela; 88-91. Propustulariaparaguanensis n. sp. Holotype NHMW 2010/0036/0015 (NMMW; ex. BL

coll.), length 17.2 mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo,

Paraguanâ Peninsula, Falcôn State, Venezuela.

92-93. Propustularia paraguanensis n. sp. Paratype 1 NHMW 2010/0036/0016 (NMMW; ex. BL coll.), length

1 7.8 mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ

Peninsula, Falcôn State, Venezuela; 94-95. Propustularia paraguanensis n. sp. Paratype 2 NHMW

-010/0036/0017 (NMMW; ex. BL coll.), length 16.1 mm. lower shell bed, 1 km Southwest of Casa Cantaure,

about 10 km west of Pueblo Nuevo, Paraguanâ Peninsula, Falcôn State, Venezuela.

B. M. Landau & L. T. Groves

Novapex 12(1-2): 1-38, 10 mars 2011

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

Although ascribed by various authors to the genus

Macrocypraea, Cypraea trinitatensis belongs within

the genus Trôna (see generic discussion above).

Unfortunately the fossula is covered by matrix in the

Schilder (1939) and Jung (1965) specimen from

Carriacou, however, the shape of the dilated

hemispherical expansion on anterior third ot the

aperture and the absence of spines on the siphonal

canal indicate they are also Trôna. Trôna trinitatensis

differs from T. ingratni by its smallcr shell (at least in

Cantaure, although the Trinidad and Carriacou shells

are larger), more elongated cylindrical shape, much

less sinuous and narrower aperture, and somewhat

more crowded teeth on both labral and columellar

sides. The re-illustration of the holotype of Cypraea

fossula Ingram, 1947 (Figs 33-36) clearly show it to

belong within the genus Trôna and not Luria as

suggested by Dolin (1991) and Dolin & Lozouet

(2004). The shell is slightly broader than our other two

specimens of T. trinitatensis , but less inflated than T.

ingrami, also the columellar density is slightly less

than typical T. trinitatensis, but probably within the

variability of this species. We therefore consider C.

fossula a junior subjective synonym of T. trinitatensis.

Geological and geographical distribution.

Lower Miocene: Cantaure Formation, Venezuela;

Tamana Formation, Trinidad.

Middle Miocene: Grand Bay Formation of Carriacou.

Subfamily EROSARIINAE Schilder, 1924

Genus Propustularia Schilder, 1927

Type species: Cypraea surinamensis Perry, 1811, by

original désignation.

Discussion. Schilder (1939) and Schilder & Schilder

(1971) assigned a small group of tropical American

cypraeids to the genus Pustularia Swainson, 1840 (ie.,

P. mejasensis, P. americana , P. caribaea ail Schilder,

1939, and C. gurabonis Ingram, 1939). However, their

shells do not conform to the generic characteristics of

the Recent Indo-Pacific Pustularia species. The fossil

taxa ail hâve shells with far less pronounced rostration

of the extremities which do not carry the spines so

characteristic of the Recent species. Lorenz (1999)

also observed that the posterior extremity composed

of a callus-accumulation and two marginal spines

formed by the posterior terminal ridges, seen in the

Recent species, is not found in any of the Caribbean

fossil species assigned by Schilder & Schilder (1971)

to Pustularia. The fossula is narrow and smooth in ail

Recent Pustularia species, whereas it is denticulate in

the Neogene Caribbean species. Most of this group of

species share more shell characteristics with the genus

Propustularia Schilder, 1927, which is today

represented by a single taxon Propustularia

surinamensis (Perry, 1811) found in the Caribbean.

Propustularia is characterised by pyriform-inflated

shells, with somewhat rostrate extremities, a weakly

produced spire and fine teeth. Pustularia americana

(holotype; Figs 147-150), on the other hand, shows the

shell characteristics of the genus Erosaria Schilder,

1924. Lorenz (1999) and Lorenz & Hubert (2000)

synonymised Propustularia with Pi oadusta Sacco

1894, and suggested Pustularia might hâve evolved

from Proadusta- like forms. Propustularia

longidentata n. sp. (herein) also shares some shell

characteristics with the deep water Indo-Pacific and

southeast African genus Nesiocypraea Azuma &

Kurohara, 1967, particularly the species N. lisetae

Kilbum, 1972. These characteristics include globular

shape, distinctly curved aperture in the posterior third,

denticulate fossula, and a callus bridge, in

Nesiocypraea connecting both, labral and columellar

teeth. In Nesiocypraea , how'ever, the teeth are short

and do not extend onto the ventrum or outer lip.

Meyer's (2004) molecular work on cypraeids placed

the clade consisting of Propustularia, Nesiocypraea

and Ipsa as basal to ail cowries.

Propustularia longidentata n. sp.

Text-figure 6; Figs 78-87

Dimensions and type material. Holotype; NHMW

2010/0036/0012, height 15.2 mm (Figs 78-81);

paratype 1; NHMW 2010/0036/0013, height 14.5 mm

(Figs 82-85); paratype 2; NHMW 2010/0036/0014,

height 12.2 mm (Figs 86-87).

Type locality. lower shell bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Falcôn State, Venezuela (=locality GS12PGNA of

Gibson-Smith & Gibson-Smith, 1979).

Type stratum. Cantaure Formation (early Miocene:

Burdigalian).

Description. Shell small for genus, fragile, globular,

posteriorly swollen, spire weakly projecting, covered

by adapical callus, extremities moderately projecting;

sides rounded, moderately callused, lip marginal

callus sharply delimited and slightly raised; ventrum

flattened, convex in profile; aperture narrow, edges

parallel, conspicuously curved to the left in the

posterior third; siphonal canal moderately long,

externally barely margined; exhalant channel

moderately produced, strongly bent adaxially,

externally barely margined; terminal ridge well-

developed, tooth-like at its inner extremity; fossula

narrow, hardly delimited from the rest of the

columella, bearing narrow elongated extensions of the

labral teeth; teeth on both sides well developed, 21-22

columellar teeth, anterior columellar teeth fused at the

angulation, forming an elevated vertical callus bridge,

mid-height the teeth weaken or subobsolete, again

strongly developed in the posterior portion, teeth

extending onto the ventrum especially in the anterior

and posterior portions, inwards the teeth hardly extend

onto the columella, columella smooth; 20-23 labral

B. M. Landau & L. T. Groves

Novapex 12(1-2): 1-38, 10 mars 2011

teeth extending just over half ot the labral width. No

dorsal sulcus, pustules or pitting présent. Colour

pattern not preserved, under UV light suggestion of

large spots on the dorsum.

Shell Formula. 14.0 (66.4 - 55.7) 26: 26

Specimen

collection number

length

width

height

Holotype (Figs 78-81 )

NHMW 2010/0036/0012

15.2

10.8

8.7

Paratype 1 (Figs 82-85)

NHMW 2010/0036/0013

14.5

9.4 (distorted)

8.3

Paratype 2 (Figs 86-87)

NHMW 2010/0036/0014

12.2

7.6

6.5

Table 7. Dimensions and number of teeth; Propustularia longidentala n. sp.

Text-figure 6. Propustularia longidentata n. sp.

Holotype NHMW 2010/0036/0012 (NMMW coll.; ex.

BL coll.), length 15.2 mm. lower shell bed, 1 km

Southwest of Casa Cantaure, about 10 km west of

Pueblo Nuevo, Paraguanâ Peninsula, Falcôn State,

Venezuela.

Discussion. Several Neogene Caribbean forms hâve

been described. Pustularia mejasensis Schilder, 1939

from Trinidad is represented by a single internai

mould. However, because it is impossible to

détermine what taxon this represents it should be

considered nomen dubiurn. Pustularia americana

Schilder, 1939 from the early Miocene of Cuba has a

more solid shell, with less produced, extremities,

wider siphonal channel, fewer columellar and labral

teeth, which on the columellar side to not extend onto

the ventrum and pustules on the dorsum adjacent to

the labral marginal callus, and is better placed in the

genus Erosaria. Propustularia caribaea (Schilder,

1939) (holotype; Figs 151-154) from the early Middle

Miocene Grand Bay Formation of Carriacou is

extremely similar to P. longidentata [shell formula

13.5 (64 - 50) 28: 23], but no bridge is developed

between the anterior columellar teeth.

Geological and geographical distribution.

Lower Miocene: Cantaure Formation, Venezuela

Propustularia paraguanensis n. sp.

Text-figure 7; Figs 88-95

Dimensions and type material. Holotype; NHMW

2010/0036/0015, lieight 17.2 mm (Figs 88-91);

paratype 1; NHMW 2010/0036/0016, height 17.8 mm

(Figs 92-93); paratype 2; NHMW 2010/0036/0017,

height 16.1 mm (Figs 94-95).

Type Iocality. lower shell bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Falcôn State, Venezuela (=locality GS12PGNA of

Gibson-Smith & Gibson-Smith, 1979).

Type stratum. Cantaure Formation (early Miocene:

Burdigalian).

Description. Shell small for genus, solid, globular,

weakly swollen posteriorly, spire involute, covered by

adapical callus, extremities weakly projecting; sides

rounded, nioderately callused, lip marginal callus

sharply delimited and slightly thickened; ventrum

flattened, weakly convex in profile; aperture narrow,

edges parai lel, conspicuously curved to the left in the

posterior third; siphonal canal moderately long,

externally barely margined; exhalant channel

moderately produced, strongly bent adaxially,

externally barely margined; terminal ridge well-

developed, tooth-like at its inner extremity; fossula

narrow, weakly delimited from the rest of the

columella, labral teeth extend across the fossula as

elevated ridges and form teeth again at the inner edge

of the fossula; 19-20 short columellar teeth, not

extending past the angulation onto the ventrum; 20-22

labral teeth extending less than half of the labral

width. No dorsal sulcus, pustules or pitting présent.

Colour pattern not preserved, under UV light

suggestion large irregular spots on the posterior

portion of the dorsum, the spots much wider than the

distance between them.

Etymology. named for the long teeth, seen especially Shell Formula. 17.0 (67.1 - 51.8) 25: 22

in the holotype.

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

specimen

collection number

length

width

height

Holotype (Figs 88-91 )

NHMW 2010/0036/0015

17.2

1 1.7

9.1

Paratype 1 (Figs 92-93)

NHMW 2010/0036/0016

17.8

1 1.5

9.0

Paratype 2 (Figs 94-95)

NHMW 2010/0036/0017

16.1

11.0

8.3

Table 8. Dimensions and number of teeth; Propustularia paraguanensis n. sp.

Text-fïgure 7. Propustularia paraguanensis n. sp.

Holotype NHMW 2010/0036/0015 (NMMW coll.; ex.

BL coll.), length 17.2 mm. lower shell bed, 1 km

Southwest of Casa Cantaure, about 10 km west of

Pueblo Nuevo, Paraguanâ Peninsula, Falcôn State,

Venezuela.

Discussion. Propustularia paraguanensis n. sp.

differs from P. longidentata n. sp. in its less

posteriorly swollen shell shape, more strongly

sculptured fossula and shorter teeth on both labral and

columellar sides. There is no trace of the callus bridge

so prominently developed in P. longidentata.

Propustularia bartschi (Ingram, 1939) (holotype; Figs

155-156) from the early Pleistocene of Costa Rica is

rnost similar to paratype 2 of P. paraguanensis (Figs

94-95) in overall shape but the widest portion of the

shell is more posterior in P. bartschi. The dentition

seems to be the same in ail of the figured type

specimens but the aperture is most similar to paratype

1 (Figs 92-93). In the latéral views P. bartschi is much

more tapered towards the anterior than P.

paraguanensis.

Etymology. named after the Paraguanâ Peninsula,

location of the Cantaure deposits.

Geological and geographical distribution.

Lower Miocene: Cantaure Formation, Venezuela

Subfamily ZONARIINAE Schilder, 1932

Genus Zonaria Jousseaume, 1884

Type species: Cypraea zonaria Gmelin, 1791, by

original désignation.

Zonaria is tranditonally assigned to the subfamily

Erroneinae Schilder, 1927. However, based on

mitochondrial molecular data Meyer (2004)

demonstrated that the genus belongs within a separate

group; Zonariinae Schilder, 1932. Zonaria is

characterized by species with oval to pyriform, small

to moderate-sized shells, the teeth are fine and widely

spaced, the fossula is much reduced, the columellar

area is smooth, the callus collar surrounding the

siphonal canal is well-developed and the colour

pattern consists of marginal spotting and a dorsum

with irregular or interrupted mottled banding. Zonaria

has been known in Europe since the early Miocene

and is distributed today in the Mediterranean and West

Africa (Lorenz & Hubert, 2000). Groves (1997)

reported the genus in the Pacific tropical American

Neogene assemblages with the description of Z.

pittorum Groves, 1997 (holotype; Figs 157-158) from

the early Pliocène Esmeraldas beds of the Onzole

Formation of Ecuador. Later Groves & Nielsen

(2003) described Z. frassinetti (Figs 159-160) from the

early late Miocene, Tortonian of Chile which is here

reassigned to the genus Pseudozonaria. There is an

additional species of Zonaria from the Neogene of the

Dominican Republic, which awaits publication (BL

coll.; unpublished data). The genus has also been

recorded from the middle Miocene of the Madeira

Archipelago (Lorenz & Groh, 1998).

Figures 96-111

96-99. Zonariapingata n. sp. Holotype NHMW 2010/0036/0018 (NMMW; ex. BL coll.), length 45.8 mm. lower

s e bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ Peninsula, Falcôn

State, Venezuela; 100-103. Pseudozonariapraeaequinoctialis n. sp. Holotype NHMW 2010/0036/0019

(NMMW, ex. BL coll.), length j 2.3 mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west

u i UË ff 0 ’ ^ ara ë uan:l Peninsula, Falcôn State, Venezuela; 104-107. Pseudozonaria primarobertsi n. sp.

^ * 8364 (NMB coll.), length 22.7 mm. lower shell bed, I km Southwest of Casa Cantaure, about

km west of Pueblo Nuevo, Paraguanâ Peninsula, Falcôn State, Venezuela; 108-111. Pseudozonaria

I l imaio ieitsi n. sp. Paratype 1 NMB H18365 (NMB coll.), length 21.3 mm. lower shell bed, 1 km Southwest of

uasa Lantaure, about 10 km west of Pueblo Nuevo, Paraguanâ Peninsula, Falcôn State, Venezuela.

B. M. Landau & L. T. Groves

NOVAPEX 12(1-2): 1-38, 10 mars 2011

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

The “ Pseudozonaria " group, which is

characterised by its elongate-oval shells with well

developed, often sharp eut teeth, densely spotted

margins, freckled dorsum and prominent extremities,

is closely related. Groves (1997) considered

Pseudozonaria a subgenus of Zonaria ; Lorenz &

Hubert (2000) used “ Pseudozonaria ” as an informai

group within Zonaria; Landau & Silva (2010)

considered the two separate at full generic rank.

Pseudozonaria has a fossil record in the tropical

American Neogene and is today restricted to the

Pacific side of its original wider distribution (i.e.

western side of Central America), and is an example

of a Paciphile taxon (Woodring, 1966; Landau et al.,

2009).

Zonaria pingata n. sp.

Text-figure 8; Figs 96-99

Dimensions and type material. Holotype NHMW

2010/0036/0018, height 45.8 mm (Figs 96-99).

Type locality. lower shell bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Falcôn State, Venezuela (=locality GS12PGNA of

Gibson-Smith & Gibson-Smith, 1979).

Type stratum. Cantaure Formation (early Miocene:

Burdigalian).

Description. Shell medium-sized for genus, solid,

ovate-pyriform, posteriorly swollen, in latéral profile

dorsal curvature regularly attenuated abapically, spire

weakly projecting, covered by adapical callus; sides

rounded, strongly callused; callus ascending above

mid-height, its highest point at the dorsal hump;

ventrum flattened, slightly convex in profile; aperture

weakly sigmoidal; siphonal canal long, abaxially

asymmetrical and recurved, extemally surrounded by

thick callus collar; exhalant channel weakly bent

adaxially, externally surrounded by thick adapical

callus; terminal ridge well-developed. tooth-like at its

inner extremity; fossula smooth, rnarked by small

dépréssion; three anterior columellar teeth, most

anterior thinnest, adapically 14 short, stout, parallel,

well-spaced inner teeth, ending abruptly at the basal

and columellar planes, not extending onto the ventrum

nor the aperture; columella smooth; 20 heavy,

regularly-spaced labral teeth; Ventral and marginal

zones spotted, marginal callus lighter coloured,

dorsum golden-brown with orange watermark-like

stains placed in irregular bands.

Shell Formula. 45.8 (67.9-58.0) 17: 14

Specimen

collection number

length

width

height

Holotype (Figs 96-99)

NHMW 2010/0036/0018

45.8

31.1

26.1

Table 9. Dimensions and number of teeth; Zonaria pingata n. sp.

Text-figure 8. Zonaria pingata n. sp. Holotype

NHMW 2010/0036/0018 (NHMW coll.; ex. BL coll.),

length 45.8 mm. lower shell bed, 1 km Southwest of

C asa Cantaure, about 10 km west of Pueblo Nuevo,

Paraguanâ Peninsula, Falcôn State, Venezuela.

Discussion. Zonaria pingata n. sp. is described based

on a single beautifully preserved specimen and is

extremely rare in the Cantaure Formation. The

Cantaure Formation shell differs from the early

Pliocène Esmeraldas Formation Z. pittorum in being

more globose rather than pyriform in shape, the

posterior portion of the aperture is more twisted to the

left in Z. pittorum and the teeth are longer on both

sides in Z. pittorum. The shell formula of the two is

very similar (Z. pittorum : 40.1 (68.0 -52.1) 17:15).

Zonaria pingata is most similar to Z. zonaria

(Gmelin, 1791), the type-species of the genus, but has

a more ovate shell, the inner lip teeth are shorter and

stouter and the siphonal canal is more recurved. The

Recent Mediterranean and West African Zonaria

pyrum (Gmelin, 1791) is similar in shape, but the

anterior third of the aperture is expanded, the inner lip

teeth are again longer and the margins are unifonu

orange without spots. The Mediterranean Pliocène

Zonaria porcellus (Brocchi, 1814) is easily separated

by its more elongate-pyriform shape and much finer

teeth, which on the inner lip become obsolète

adapically.

Etymology. From Latin for stained Zonaria.

Geological and geographical distribution.

Lower Miocene: Cantaure Formation, Venezuela.

B. M. Landau & L. T. Groves

Novapex 12(1-2): 1-38, 10 mars 2011

Zonaria pseudotumulus n. sp.

Text-figure 9; Figs 132-139

Dimensions and type material. Holotype; NMB

H18360, height 27.2 mm (Figs 132-135); paratype 1

NHMW 2010/0036/0022, height, 26.4 mm (Figs 136-

137); paratype 2 NHMW 2010/0036/0023, height,

22.2 mm (Figs 138-139).

Type locality. lower shell bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Falcôn State, Venezuela (=locality GS12PGNA of

Gibson-Smith & Gibson-Smith, 1979).

Type stratum. Cantaure Formation (early Miocene:

Burdigalian).

Description. Shell medium-sized, solid, pyriform,

posteriorly swollen, in latéral profile dorsal curvature

regularly attenuated abapically, spire involute, covered

by adapical callus; sides rounded, moderately

callused, callus ascending to 'A-'A shell height;

ventrum weakly convex in profile; margination (sensu

Dolin & Lozouet, 2004) prominent, giving the left

anterior terminal a pinched appearance; aperture of

medium width, anterior portion almost straight,

posteriorly curving to the left, anterior portion very

slightly wider; siphonal canal relatively long, narrow,

abaxially asymmetrical; exhalant channel moderately

produced, linrited by parallel lips; columella

denticulate along entire length, bearing 22-23 short

denticles, which extend as folds a short distance onto

the columella peristome and a short distance across

the angulation onto the ventrum; anterior portion of

angulation swollen and ridge like, overhanging the

fossula; anterior columellar teeth more prominent;

inner % of columella peristome smooth; terminal fold

oblique (sensu Schilder & Schilder, 1938), separated

from the columellar teeth by a deep groove,

moderately developed, bordering the siphonal canal,

small internai lamina (sensu Dolin & Lozouet, 2004);

fossula relatively well developed but poorly delimited,

concave, very steep, ridges originating from the

anterior columellar teeth run onto the fossula, become

obsolète mid-fossula, which is smooth, strengthening

again at the inner edge to forrn inner teeth; 20-27 short

labral teeth developed at the lip edge, hardly running

onto the labrum. Traces of spotted colour pattern

preserved on the dorsum, enhanced under UV light.

Shell Formula. 25.3 (65.1 - 50.1) 23: 22

specimen

collection number

length

Width

height

Holotype (Figs 132-135)

NMB H18360

27.2

17.5

14.1

Paratype 1 (Figs 136-137)

NHMW 2010/0036/0022

26.4

17.4

12.3

Paratype 2 (Figs 138-139)

NHMW 2010/0036/0023

22.2

14.5

11.6

Table 10. Dimensions and number of teeth; Zonaria pseudotumulus n. sp.

Text-figure 9. Zonaria pseudotumulus n. sp. Holotype

NMB H18360 (NMB coll.), length 27.2 mm. lower

shell bed, 1 km Southwest of Casa Cantaure, about 10

km west of Pueblo Nuevo, Paraguanâ Peninsula,

Falcôn State, Venezuela.

Discussion. Zonaria pseudotumulus n. sp. is closely

similar to the early Miocene Z. tumulus (Heilprin,

1886) from the Chipola Formation of Florida, but

differs in having a less globose shell, the anterior

portion of the aperture is less expanded, the outer lip

is not sinuous and the fossula is not as wide nor as

strongly ridged as in Z. tumulus. Zonaria spurcoides

(Gabb, 1873) (lectotype; Figs 161-164) from the early

Miocene Baitoa Formation (Figs 140-143), early

Pliocène Gurabo Formation of the Dominican

Republic, and early Miocene Chipola Formation of

Florida (Dolin, 1991) is even more similar, but differs

most notably in the character of the anterior portion of

the columella and anterior left terminal; in Z.

pseudotumulus the anterior portion of the angulation is

markedly swollen, overhanging the fossula, the strong

anterior teeth extend a short distance over the fossula

and are obsolète mid-fossula in the paratypes, in the

holotype the anterior two ridges run right across the

fossula, although weakened. In Z. spurcoides the

anterior portion of the angulation is not as swollen, the

columellar teeth are wider spaced and continuous

across the fossula as narrow ridges joining the

columellar teeth at the angulation with the teeth at the

inner edge of the fossula. In Z. pseudotumulus the

anterior left terminal has a pinched and somewhat

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

twisted appearance, constricted between the groove

between the anterior terminal and first columellar

tooth on the ventral side and the indented margination

on the dorsum. This feature is not seen in Z.

spurcoides. The fossula is also flatter in Z.

pseudotumulus.

Etymology. Named reflecting similarity to Cypraea

tumulus from the early Miocene Chipola Formation of

Florida.

Geological and geographical distribution.

Lower Miocene: Cantaure Formation, Venezuela.

Subfamily PSEUDOZONARIINAE

Genus Pseudozonaria Schilder, 1927.

Type species Cypraea arabicu/a Lamarck, 1810,

original désignation.

The “ Pseudozonaria ” group is characterised by its

elongate-oval shells with well developed, often sharp

eut teeth, densely spotted margins and freckled

dorsum and prominent extremities. Today the group is

represented by five tropical American Pacific species;

P. aequinoctialis (Schilder, 1931), P. annettae (Dali,

1909), P. arabicula (Lamarck, 1810), P. nigropuctata

(Gray, 1828), and P. robertsi (Hidalgo, 1906) (Lorenz

& Hubert, 2000; Meyer, 2003, 2004). Several fossil

species belonging to Zonaria Jousseaume, 1884 and

Pseudozonaria Schilder, 1927 hâve been described

from the Gatunian Neogene. Zonaria and

Pseudozonaria are closely related groups, and not ail

cypraeid specialists are in agreement as to their rank

or which species belong in which of the two groups.

Groves (1997) considered Pseudozonaria a subgenus

of Zonaria ; Lorenz & Hubert (2000) use

“Pseudozonaria ” as an informai group within

Zonaria , and Landau & Silva (2010) considered both

valid généra.

Lôpez Soriano (2006) justified the séparation of a

new subfamily Pseudozonariinae from the Zonariinae

Schilder, 1932 based on molecular data (Meyer,

2004), anatomical différences of the mande and

papillae, and some small différences in shell

morphology. They also hâve distinct geographical

distributions; Pseudozonariinae are tropical American,

today restricted to the western side of Central

America; Zonariinae are known since the Miocene in

Europe and Recent of West Africa (Lorenz & Hubert,

2000 ).

Pseudozonaria praeaequinoctialis n. sp.

Text-figure 10; Figs 100-103

Dimensions and type material. Holotype; NHMW

2010/0036/0019, height 32.3 mm (Figs 100-103).

Type locality. lower shell bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Falcôn State, Venezuela (=locality GS12PGNA of

Gibson-Smith & Gibson-Smith, 1979).

Type stratum. Cantaure Formation (early Miocene:

Burdigalian).

Description. Shell of medium size and thickness for

genus, inflated pyriform, posteriorly swollen, in latéral

profile dorsal curvature regularly attenuated

abapically, spire weakly projecting, covered by

adapical callus; sides rounded, moderately callused;

ventrum flattened, convex in profile; aperture

relatively wide and straight, posteriorly curving to the

left, anterior portion much wider; siphonal canal

narrow, abaxially asymmetrical; exhalant channel

hardly produced, limited by parallel lips; Columella

denticulate along entire length, bearing 19 sharp,

narrow denticles, which extend as folds onto the

columella peristome within the aperture, but not onto

the ventrum with the exception of the three adapical

denticles, which extend a short distance over the

angulation onto the ventrum; terminal fold marginal,

bordering the siphonal canal, where it is strengthened

and keel-like, separated from the columellar teeth by a

deep groove; fossula small, weakly concave, very

steep, poorly delimited from the rest of the columella,

bearing ridges which do not extend to the smooth

inner edge; 20 very short, sharp labial teeth,

developed at the inner edge, but not extending onto

the labrum; labrurn moderately thickened in the

médial portion. A suggestion of a spotted marginal

colour pattern and dorsal banding is preserved, this

pattern enhanced under UV light.

Shell Formula. 32.3 (65.9 - 51.3) 18: 18

Figures 112-127

112-115. Pseudozonariaprimarobertsi n. sp. Paratype 3 NMB H18359 (NMB coll.), length 28.9 mm. lower

shell bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ Peninsula, Falcôn

State, Venezuela; 116-119. Pseudozonaria primarobertsi n. sp. Paratype 6 NMB H18366 (NMB coll.), length

-4.8 mm. lowei shell bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguanâ

Peninsula, Falcôn State, Venezuela; 120-123. Pseudozonaria primarobertsi n. sp. Paratype 5 NMB H18363

(NMB coll.), length 25.3 mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo

mP eninsula, Falcôn State, Venezuela; 124-127. Pseudozonariafalconensis n. sp. Holotype

^8 (NMB coll.), length 16.5 mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km

west ot Pueblo Nuevo, Paraguanâ Peninsula, Falcôn, State Venezuela.

B. M. Landau & L. T. Groves

Novapex 12(1-2): 1-38, 10 mars 2011

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

Specimen

collection number

length

width

height

Holotype (Figs 100-103)

NHMW 2010/0036/0019

32.3

21.3

16.6

Table 11. Dimensions and number of teeth; Pseudozonaria praeaequinoctialis n. sp.

Text-fîgure 10 . Pseudozonaria praeaequinoctialis n.

sp. Holotype NHMW 2010/0036/0019 (NHMW coll.;

ex. BL coll.), length 32.3 mm. lower shell bed, 1 km

Southwest of Casa Cantaure, about 10 km west of

Pueblo Nuevo, Paraguanâ Peninsula, Falcôn State,

Venezuela.

the widest aperture of any of the group and the least

number of teeth, especially on the columellar side

where they are obsolète on the posterior portion of the

columella (shell formula for Z. jfrassinetti; 28.1 (62.2-

44.5) 13: 4).

Etymology. Combination of before = ‘prae’ and

equinoctial = ‘aequinoctialis’[Latin],

Geological and geographical distribution.

Lower Miocene: Cantaure Formation, Venezuela.

Pseudozonariaprimarobertsi n. sp.

Text-llgure 11; Figs 104-123

Dimensions and type material. Holotype; NMB

H18364, length 22.7 mm (Figs 104-107); paratype 1

NMB H18365, length, 21.3 mm (Figs 108-111);

paratype 2 NMB H18367, length, 24.7 mm; paratype

3 NMB H18359, length, 28.9 mm (Figs 112-115);

paratype 4 NMB H18361, length, 24.4 mm; paratype

5 NMB H18363, length, 25.3 mm (Figs 120-123);

paratype 6 NMB H18366, length, 24.8 mm (Figs 116-

119).

Discussion. Pseudozonaria praeaequinoctialis n. sp.

is clearly the predecessor of the Pseudozonaria

annettae- group now living in the tropical American

Pacific. This group is characterised by their very short

teeth and wide apertures. In the Recent faunas two

species/subspecies are recognised, which we prefer to

recognise at full species level, a northern form, P.

annettae (Dali, 1909) found along the Pacific coast of

Mexico, and a Southern form P. aequinoctialis

(Schilder, 1933) from Panama to Ecuador. The

northem form is easily separated from P.

praeaequinoctialis having a much more elongated-

pyri form shell shape. The Southern form differs from

the northem form in having a heavier, broader shell

with slightly coarser teeth, and therefore is doser to

the tossil species. Pseudozonaria praeaequinoctialis

ditters from P. aequinoctialis in having an even

broader shell (width/height 59.1 vs 65.9 in P.

praeaequinoctialis ) and in having considerably fewer

columellar denticles (CT = 14 vs. 18 in P

praeaequinoctialis).

Zonaria frassinetti Groves & Nielsen, 2003 fi-,

the early late Miocene, Tortonian of Chile is h

reassigned to the Pseudozonaria annettae-grc

characterised by its very wide aperture and wid

spaced and sharp denticles. In Pseudozona

. rassinetti these features are exaggerated, so that it 1

Other material. 20 specimens BL coll.

Type locality. lower shell bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Falcôn State, Venezuela (=locality GS12PGNA of

Gibson-Smith & Gibson-Smith, 1979).

Type stratum. Cantaure Formation (early Miocene:

Burdigalian).

Desci iption. Shell small, solid, oval-depressed to

cylindrical-pyriform, posteriorly swollen, in latéral

Profile dorsal curvature regularly attenuated

abapically, spire involute, covered by adapical callus;

sides rounded, strongly to moderately callused, callus

ascending to 'A-'A shell height; ventrum flattened, very

weakly convex at the extremities; aperture of medium

width, parasigmoida), posteriorly strongly curving to

the left, anterior portion very slightly wider; siphonal

canal narrow, abaxially asymmetrical; exhalant

channel weakly produced, deep, limited by parallel

lips, columella denticulate along entire length, bearing

16-22 coarse, elevated denticles, which extend as folds

onto the columella peristome and a very short distance

over the angulation onto the ventrum, anterior two

columellar teeth larger and often fused or partially so;

terminal fold strongly developed and elevated,

B. M. Landau & L. T. Groves

Novapex 12(1-2): 1-38, 10 mars 2011

bordering the siphonal canal, where it is strengthened

and keel-like with a prominent, elevated internai

lamina (sensu Dolin & Lozouet, 2004), separated from

the columellar teeth by a deep groove; fossula hardly

developed and poorly delimited from the rest of the

columella, very steep, bearing ridges which extend to

the inner edge, ending in a denticle; 20-24 coarse

labral teeth, outer lip bevelled inwards, with teeth

extending across the bevelled inner portion and a short

distance onto the labrum. A suggestion of a fine,

messy spotted marginal colour pattern is preserved in

sorne specimen, enhanced under UV light.

Shell Formula, ‘cylindrical-pyriform morphotype’;

23.0 (65.8-51.7) 22: 18

Specimen

collection number

length

width

height

holotype (Figs 104-107)

NMB H18364

22,7

15,3

11,7

paratype 1 (Figs 108-111)

NMB H18365

21,3

13,9

11,1

paratype 2

NMB H18367

24,7

15,8

12,6

paratype 3 (Figs 112-115)

NMB H18359

28,9

17,7

13,1

BL coll. 1

29,2

18,5

14,3

BL coll. 2

20,7

14,4

11,4

BL coll. 3

25,1

16,3

13,1

BL coll. 4

18,8

12,3

9,8

BL coll. 5

21,4

14,1

11,2

BL coll. 6

21,7

13,9

10,9

BL coll. 7

22,6

15,2

11,9

BL coll. 8

20,1

14,5

11,5

BL coll. 9

21,9

15,6

12,1

BL coll. 10

22,7

14,4

11,8

BL coll. 11

23,3

15,2

11,9

Table 12a. Dimensions and number of teeth; Pseudozonariaprimarobertsi n. sp., ‘cylindrical-pyriform

morphotype’.

Shell Formula, ‘oval-depressed morphotype’; 23.8 (71.1 -54.0) 22: 20

Specimen

collection number

length

width

height

paratype 4

NMB H18361

24,4

13,4

paratype 5 (Figs 120-123)

NMB H18363

25,3

14,2

paratype 6 (Figs 116-119)

NMB H18366

24,8

17,1

12,6

BL coll. 1

27,3

20,7

15,2

BL coll. 2

23,1

16,3

11,7

BL coll. 3

22,2

15,3

11,4

BL coll. 4

22,9

16,6

12,1

BL coll. 5

24,8

17,3

BL coll. 6

18,8

14,4

BL coll. 7

15,9

12,5

BL coll. 8

19,8

14,3

10,8

BL coll. 9

21,4

14,8

Table 12b. Dimensions and number of teeth; Pseudozonaria primarobertsi n. sp., ‘oval-depressed morphotype’.

Text-figure 11 (opposite). Pseudozonaria

primarobertsi n. sp.. Paratype 1 NMB Fil8365 (NMB

coll.), length 21.3 mm. lower shell bed, 1 km

Southwest of Casa Cantaure, about 10 km west of

Pueblo Nuevo, Paraguanâ Peninsula, Falcôn State,

Venezuela.

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

Discussion. Pseudozonaria primarobertsi n. sp. is the

most common cypraeid in the Cantaure Formation.

Two morphotypes occur; a ‘cylindrical-pyritoim

morphotype’ and an ‘oval-dcpressed morphotype .

Apart from the différence in shell shape the ‘oval-

depressed morphotype' tends to hâve one or two more

columellar teeth than the ‘cylindrical-pyriform

morphotype’ and the teeth in general tend to be

somewhat less coarse and elevated in the ‘oval-

depressed morphotype’, however, the character of the

strong terminal ridge, very weakly developed fossula

and ridged columellar peristome is the same in both

morphotypes. Moreover, a fair number of specimens

could not be clearly ascribed to one or other

morphotypes. The différences seen in the two

morphotypes could indicate sexual dimorphism.

height

1,70

Pseudozonaria sp.

o cylindrical-pyriform

O O O °

morphotype

| 1,50

O % é> °

■ oval-depressed

<>■■■ 1 ■

morphotype

■§) 1,40

■> O." "

■

1,30

1,20

■

Text-fîgure 12. Height against height/width; Pseudozonaria primarobertsi n. sp.

Pseudozonaria primarobertsi n. sp. is clearly closely

similar in size, shape and character of the teeth to the

Recent tropical American P. robertsi (Hidalgo, 1906).

As seen in our fossil taxon, the development of the

marginal callus in P. robertsi is also somewhat

variable The main différence between the two taxa is

that the Recent species has fewer columellar teeth (CT

= 12 vs 18-20 in P. primarobertsi).

Figures 128-143

128-129 . Pseudozonaria falconensis n. sp. Paratype 1 NMB H18369 (NMB coll.), length 14.4 mm. lower shell

bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguand Peninsula, Falcôn State,

Venezuela; 130-131. Pseudozonaria falconensis n. sp. Paratype 2 NHMW 2010/0036/0020 (NMMW; ex. BL

coll.), length 16.7 mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo,

Paraguand Peninsula, Falcôn State, Venezuela; 132-135. Zonariapseudotumulus n. sp. Holotype NMB H18360

(NMB coll.), length 27.2 mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo

Nuevo, Paraguand Peninsula, Falcôn State, Venezuela;

136-137. Zonaria pseudotumulus n. sp. Paratype 1 NHMW 2010/0036/0022 (NMMW; ex. BL coll.), length 26.4

mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Paraguand

Peninsula, Falcôn State, Venezuela. 138-139. Zonaria pseudotumulus n. sp. Paratype 2 NHMW 2010/0036/0023

(NMMW; ex. BL coll.), length 22.2 mm. lower shell bed, 1 km Southwest of Casa Cantaure, about 10 km west

of Pueblo Nuevo, Paraguand Peninsula, Falcôn State, Venezuela;

140-143. Zonaria spurcoides (Gabb, 1873) NHMW 2010/0036/0031 (NMMW; ex. BL coll.), length 28.3 mm.

Baitoa Formation, early Miocene, Arroyo Hondo, Yaque del Norte River, Dominican Republic.

itfl36û

B. M. Landau & L. T. Groves

NOVAPEX 12(1-2): 1-38, 10 mars 2011

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

In the Neogene several species hâve been

described with shells closely similar to those of P.

robertsi. Olsson ( 1964) described P. telembiensis from

the Pacific portion of the Neogene Gatunian Province,

from a single specimen from the late Miocene

Angostura Formation of Ecuador. Groves (1997) re-

illustrated the holotype. This Ecuadorian species is

extremely similar in shape to P. primarobertsi,

differing only in the less crowded nature of the teeth

(shell formula for P. telembiensis ; 19.7 (67-52) 16:

15). Groves (1997) described Zonaria cathyae from

the early Pliocène Esmeraldas beds of the Onzole

Formation of Ecuador. Now considered a

Pseudozonaria, it is extremely similar to the ‘oval-

depressed morphotype’ of P. primarobertsi, and

indeed, apart from slightly fewer teeth in P. cathyae,

their shell formulae are very similar (shell formula for

P. telembiensis-, 22.3 (71-57) 19: 17). The main

différence between the two taxa is in the apertural

characteristics of the anterior portion. In ail members

of the P. robertsi- group, including P. primarobertsi,

P. porteili Petuch, 1990, P, raymondrobertsi (Pilsbry,

1922) (holotype; Figs 169-172), P. telembiensis and P.

fehsei Landau & Silva, 2010 (holotype; Figs 165-168)

the terminal ridge is strong and keel-like followed by

a deep groove separating the most anterior columellar

tooth, which is usually more strongly developed.

Pseudozonaria cathyae does not belong to this group,

having the anterior third of the aperture dilated by a

small hemispherical expansion, and there is no deep

groove separating a weaker terminal ridge from the

relatively fine columellar teeth.

Pseudozonaria raymondrobertsi, from the early

Pliocène Gurabo Formation of the Dominican

Republic and early Pliocène Bowden Formation of

Jamaica, can be easily separated by the consistently

well developed and angular marginal callus giving the

shell a very broad, depressed shape. Pseudozonaria

raymondrobertsi differs from the rest of the P.

robertsi -group in usually having the terminal ridge

less strongly developed, the groove between the

terminal ridge and first columellar tooth shallower

(although it is quite prominent in the holotype Figs

104-107) and in having fewer, finer, longer teeth

(shell formula for P. raymondrobertsi', 28.2 (70.5-

51.1) 16: 14: ten specimens from Arroyo Zamba,

Dominican Republic, BL coll.).

Pseudozonaria fehsei from the early Pliocène

Cubagua Formation of Cubagua Island, Venezuela

also has a similar shell shape, but differs in the nature

of the aperture, which is a little wider in the anterior

portion in P. fehsei and the teeth are a little longer and,

especially on the labial side, are a little more widely

spaced (shell formula for P. fehsei-, 24.7 (66-50.6) 18:

16). Pseudozonaria porteili from the Pleistocene

Bermont Formation of Florida has very coarse

denticles (shell formula for P. porteili', 25 (63.9-51.5)

19: 15), similar in number to P. fehsei and P.

telembiensis, but differs in that the columellar

denticles do not extend over the columellar peristome

(see Petuch, 1990, fig. 9).

Etymology. Reflecting the earliest species in the P.

robertsi species-group known.

Geological and geographical distribution.

Lower Miocene: Cantaure Formation, Venezuela.

Figures 144-172 (type specimens of some tropical American Neogene Cypraeidae).

144-145. Muracypraea henekeni (G. B. Sowerby I, 1850) Lectotype BMNH G83940, length, 58 mm, Miocene,

^ que R ' v n e 1 r ’ St Domingo ' Colld - CoL T S - 1 lencken. (‘J.S. Heniker’), c. 1848; 146. Luriapatrespatriae

(Maury, 1917) Holotype PR1 28780 (ex Gabb collection), length 27.9 mm. Bluff 1, Cercado de Mao, Santo

omingo (specimen figured by Maury, 1917, pl. 19, fig. 10; adapical portion of outer lip seems to hâve been

damaged since the original figure was taken). Image courtesy of the Paleontological Research Institution; 147-

150 Pustularm (Pustulana) americana Schilder, 1939 Holotype NMB H 11259, length 11.2 mm, width 7.8 mm,

height 6.^ mm, Lower Miocene?: Cauto Fluss, road from Alto Cedro to Palma Soriano Cuba- 151-154

Propustularia caribaea (Schilder. 1939) Holotype NMB H11260, length 13.6 mm, width 9.0 mm heVht 6 8

mm early Middle Miocene Grand Bay Formation, Carriacou; 155-156. Propustularia bartschi (Ingram. 1939)

Holotype USNM 559684, length 25.8 mm, width, 17 mm, height 12.8 mm, Lower Pleistocene, Moin Formation,

Limon, Costa Rica; 157-158. Zonariapittorum Groves, 1997 Holotype LACMIP 12432, length 40.1 mm, width

-.0.9 mm height 12.8 mm, Lower Pliocène, TU locality 1399, roadcut on west side of village of Camarones, 20

km east ot bridge over Rio Esmeraldas, Esmeraldas Beds, Onzole Formation, Ecuador; 159-160. Pseudozonaria

frassinetti Groves & Nielsen, 2003) Hypotype LACMIP 13720, length 23.1 mm, width, 14.8 mm, height 11 2

mm early Upper Miocene, Torton.an, Navidad Formation, tidal platform approximately I km north of Matanzas,

auenal Caro Province, central Chile; 161-164. Zonariaspurcoides (Gabb, 1873) Lectotype ANSP 2999, length

<51 ~ mm ’ Miocene, Santo Domingo; 165-168. Pseudozonaria fehsei Landau &

Tinn ~i i r 2010/0038/0013 (NHMW coll., ex B L coll.), Lower Pliocène, Araya Formation,

Canon de las Calderas, Cubagua Island. Length 26.6 mm, width 16.9 mm; 169-172. Pseudozonaria

Dom/ngt' Sl ( y ' l922) HOl ° tyPe ANSP 3 " 5 ' kngth 275 mm ’ 19 mm ’ heighl - 15 "™, Miocene, Santo

B. M. Landau & L. T. Groves

NOVAPEX 12(1-2): 1-38, 10 mars 2011

B. M. Landau & L. T. Groves

Cypraeidae from the early Miocene of northem Venezuela

Pseudozonaria falconemis n. sp.

Text-figure 13; Figs 124-131

Dimensions and type material. Holotype; NMB

H18368, height 16.5 mm (Figs 124-127); paratype 1

NMB H18369, height 14.4 mm (Figs 128-129);

paratype 2 NHMW 2010/0036/0020, height, 16.7 mm

(Figs 130-131); paratype 3 NHMW 2010/0036/0021,

height, 14.9 mm.

Type locality. lower shell bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Falcôn State, Venezuela (=locality GS12PGNA of

Gibson-Smith & Gibson-Smith, 1979).

Type stratum. Cantaure Formation (early Miocene:

Burdigalian).

Description. Shell very small, solid, cylindrical-

pyriform, posteriorly swollen, in latéral profile dorsal

curvature regularly attenuated abapically, spire

involute to very weakly projecting, covered by

adapical callus; sides rounded, moderately callused,

callus ascending to 'A-'A shell height; ventrum weakly

convex in profile; aperture of medium width, almost

straight, posteriorly weakly curving to the left.

Text-figure 13. Pseudozonaria falconensis n. sp.

Holotype NMB H18368 (NMB coll,), length 16.5

mm. lower shell bed, 1 km Southwest of Casa

Cantaure, about 10 km west of Pueblo Nuevo,

Paraguanâ Peninsula, Falcôn State, Venezuela.

Discussion. This is the smallest Pseudozonaria

species in the Cantaure Formation, and easily

anterior portion very slightly wider; siphonal canal

natTow, abaxially asymmetrical; exhalant channel

moderately produced, widening slightly adapically;

columella denticulate along entire length, bearing 15-

21 short, coarse denticles, which extend as folds a

short distance onto the columella peristonre, but not

across the angulation onto the ventrum, anterior

columellar teeth slightly more prominent; inner A of

columella peristome smooth; terminal fold strongly

developed and elevated, bordering the siphonal canal,

where it is strengthened and keel-like with a

prominent, elevated internai lamina (sensu Dolin &

Lozouet, 2004), separated from the columellar teeth

by a groove; fossula relatively well developed,

concave, but poorly delimited from the rest of the

columella, very steep, bearing five ridges of variable

strength extending from the anterior columellar teeth

to the inner edge of the fossula, where they form

relatively long, extremely large, rounded inner teeth;

19-24 coarse labral teeth, anterior third of labral teeth

stretched out on slightly constricted, depressed,

hemicircular area; outer lip bevelled inwards, with

teeth extending across the bevelled inner portion and a

variable distance onto the labrum, at most just short of

mid-labrum. No colour pattern preserved.

Shell Formula. 14.3 (64.5 - 52.2) 26; 20

recognised by its rather produced posterior extremity

and relatively well developed fossula, which is smooth

in the middle, but with enormous teeth developed at

the inner edge of the fossula. We hâve not found any

congener with this feature similarly strongly

developed.

Etymology. Named for Falcôn State, Venezuela.

Geological and geographical distribution.

Lower Miocene: Cantaure Formation, Venezuela.

BIOGEOGRAPHJC IMPLICATIONS

The southem Caribbean early Miocene had a rich and

varied cypraeid assemblage, with thirteen species here

recorded Irom the early Miocene, Burdigalian

assemblage of Cantaure, ten of which are described as

new. The presence of an equally rich cypraeid

assemblage in the early Miocene of the Chipola

Formation, northem Florida, was described by Dolin

(1991). As mentioned in the introduction, cypraeids

are a difficult group to work with, and both the

spécifie and supraspecific classification varies

specimen

collection number

length

width

height

holotype (Figs 124-127)

NMB H18368

16,5

10,5

8,6

paratype 1 (Figs 128-129)

NMB H18369

14,4

8,9

7,3

paratype 2 (Figs 130-131)

NHMW 2010/0036/0020

16,7

10,7

8,5

paratype 3

NHMW 2010/0036/0021

14,9

9,7

7,9

BL coll. 3

11,2

7,6

6,2

Table 13. Dimensions and number of teeth; Pseudozonaria falconemis n. sp.

B. M. Landau & L. T. Groves

Novapex 12(1-2): 1-38, 10 mars 2011

enormously between researchers. In order for us to

compare the faunas on either side of the Caribbean

early Neogene we must first review the classification

offered by Dolin (1991). Table 14 présents a generic

révision of Dolin’s (1991) taxa. This is not a species-

level révision, which is beyond the scope of this work,

and some of the taxa may well be junior subjective

synonyms (see Fehse, 2004).

Classification fide Dolin (1991)

Current taxonomy

Cypraeorbis emilyae n.sp.

Cypraeorbis emilyae (Dolin, 1991)

Cypraeorbis hertleini (Ingram, 1948)

Cypraeorbis willcoxi willcoxi (Dali, 1890)

Cypraeorbis bal lis ta (Dali, 1915)

Cypraeorbis willcoxi ballista (Dali, 1915)

Zoila willcoxi (Dali, 1890)

Cypraeorbis willcoxi willcoxi (DM, 1890)

Zoila arlettae n.sp.

Cypraeorbis arlettae (Dolin, 1991)

Siphocypraea chilona (Dali, 1900)

Cypraeorbis chilona (Dali, 1900)

Trôna leporina calhounensis n.ssp.

Trôna calhounensis Dolin, 1991

Talparia dominiciensis (Gabb, 1873) [of Dolin,

1991]

Luria dominiciensis (Gabb, 1873) [of Dolin, 1991]

= L. dockervi (Dolin & Lozouet, 2004)

Talparia mariaelisabethae n.sp

Luria mariaelisabethae (Dolin, 1991)

Mauritia campbelliana (Pilsbry, 1922)

Luria campbelliana (Pilsbry, 1922)

Lyncina theresae n.sp.

Zonaria theresae (Dolin, 1991) [?= Z.

mariaelisabethae (Dolin, 1991)]

Erronea (Adusta) tumulus (Lleilprin, 1886)

Zonaria tumulus (Heilprin, 1886)

Erronée/ (Adusta) heilprinii (Dali, 1890)

Zonaria heilprinii (Dali, 1890)

Erronea (Adusta) spurcoides (Gabb, 1873)

Zonaria spurcoides (Gabb, 1873)

Erronea (Adusta) shirleyae n.sp.

Zonaria shirleyae (Dolin, 1991)

| Bistolida praelatior n.sp.

Zonaria praelatior (Dolin, 1991)

Table 14. Cypraeid assemblage présent in the early Miocene Chipola Formation of northern Florida;

classification of Dolin (1991) vs. current taxonomy.

From a palaeobiogeographic perspective the

Cantaure Formation assemblage is chronologically the

oldest Neogene deposit, situated geographically on the

southemmost part of the Gatunian

palaeobiogeographic province (Vermeij & Petuch,

1986; Vermeij, 2005; Landau et al., 2008). The

Chipola Formation assemblage is situated

geographically within the neighbouring northern

Caloosahatchian Province, which was already a

distinct palaeobiogeographic province by the early

Miocene (Vermeij, 2005). In the présent work we

identify two généra, Muracypraea and Propustularia ,

which seem to be endemic to the Gatunian Province.

Pseudozonaria has not been found in the

Caloosahatchian Province during the Miocene or

Pliocène, but is represented by a single, extremely rare

species in the Pleistocene, P. portelli (Petuch, 1990).

Conversely, Cypraeorbis Conrad, 1865 has not so far

been recorded in the Neogene Gatunian Province.

Acknowledgments

Our thanks to Walter Etter and Olivier Schmidt of the

Naturhistorisches Muséum Basel (NMB), Switzerland,

for access to the Gibson-Smith collection and the loan

of type specimens from the Naturhistorisches Muséum

Basel collection. To Paul Callomon, Amanda Lawless

(ANSP), Mary Stecheson and Harry Filkorn

(LACMIP), Mark Goodwin (UCMP), and Jann

Thompson, Tom Waller (USNM) for the loan of type

specimens in their respective institutions. Dr. Alan G.

Beu, GNS Science, Lower Flutt, New Zealand

reviewed the MS and made many useful suggestions

for improvement.

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R. Houart&S.Gori

Novapex 12(1-2): 39-45, 10 mars 2011

Description of two new Favartia species

(Gastropoda: Muricidae: Muricopsinae)

from Masirah Island, Oman, Arabian Peninsula

Roland HOUART

Research Associate

Institut royal des Sciences naturelles de Belgique

Rue Vautier, 29, B-1000 Bruxelles, Belgium

roland.houart@skynet.be

Sandro GOR1

Via Sernesi, 7

57123 Livorno, Italy

sandrogori@fastwebnet.it

KEYWORDS. Eastern Indian Océan, Arabian Peninsula, Oman, Gastropoda, Muricidae, Favartia

n. sp.

ABSTRACT. Two new species of Muricidae are described and compared with similar looking

species from the western Indian Océan. The Favartia species from the studied area are ail

illustrated.

INTRODUCTION

There are currently 64 known Recent species of

Favartia s.s. and 22 of Favartia (Pygmaepterys)

(sensu Houart, in litt.). Twenty-five of these species

occur in the Indian Océan of which 12 are also

recorded in the western Indian Océan and 8 in the

studied area, more precisely, Oman and nearby areas:

Favartia (Favartia) cecalupoi Bozzetti, 1993, F. (F.)

cyclostoma (Sowerby, 1841), F. (F.) deynzeri Houart,

1998, F. (F.) flexirostris (Melvill, 1898), F. (F.)

marjoriae (Melvill & Standen, 1903), F.

(Pygmaepterys) adenensis (Houart & Wranik, 1989),

F. (P.) paulboschi Smythe & Houart, 1984 and F. (P.)

yemenensis (Houart & Wranik, 1989).

The Mollusca of this part of the world were

recently commented on and illustrated in several

publications and books. The first publication, by

Biggs (1973), includes Mollusca of the Trucial Coast,

at the southem end of the Persian Gulf (United Arabs

Emirates), but only a few muricids were listed and one

was illustrated: Hexaplex kusterianus (Tapparone-

Canefri, 1875). Doreen Sharabati (1981) followed

with a fully illustrated volume with over 100 species

of mollusks depicting their habitats, their uses by man

throughout the years and illustrating rnany species in

their habitat. In 1982, Kathleen Smythe published a

small book about the seashells of the Arabian Gulf,

but only a few species were illustrated. Still in 1982

and then in 1989, Donald and Eloise Bosch published

two very useful books devoted to the seashells of

Oman and of Southern Arabia with numerous color

illustrations of the shells. Then more recently, a team

of specialists, well known for their many contributions

to the study of mollusks, Donald T. Bosch, S. Peter

Dance, Robert G. Moolenbeek and P. Graham Oliver

(1995) published a huge and impressive book where

more than a thousand species were described and

illustrated.

Three Favartia species were illustrated by these

authors: F. (F.) cyclostoma (Sowerby, 1841) (Bosch et

al., 1995: 119, fig. 470), F. marjoriae (Melvill &

Standen, 1903) (Bosch et al., 1995: 119, fig. 472), F.

(P.) paulboschi Smythe & Houart, 1984 (Bosch &

Bosch, 1989: 58; Bosch et al., 1995: 119, fig. 472) and

F. (P.) colombi n. sp. [under the name F. (P.)

yemenensis] (Bosch et ah, 1995: 120, fig. 478).

Favartia marjoriae (Melvill & Standen, 1903)

(Figs 23-25) is probably conspecific with F. (F.)

maculata (Reeve, 1845) (Fig. 26), a species living

throughout the Indo-West Pacific and known to hâve

very variable shell characters. The protoconch of F.

marjoriae lias not yet been examined, but this and

eventually DNA studies of F. marjoriae and of

different forms of F. maculata will be necessary to

confirm or réfuté their conspecificity.

R. Houart & S. Gori

Two new Favartia species from Masirah Island

Table 1. Terminology used to describe the spiral cords and apertural denticles (after Merle, 1999 and 2001)

Terminology in parenthèses: erratic feature.

Primary cord

s :

secondary cord

IP:

Infrasutural primary cord (primary cord on subsutural ramp)

adis :

adapical infrasutural secondary cord (on subsutural ramp)

abis :

abapical infrasutural secondary cord (on subsutural ramp)

PI :

Shoulder cord

P2-P6 :

Primary cords of the convex part of the teleoconch whorl

sl-s6 :

secondary cords of the convex part of the teleoconch whorl

example: si = secondary cord between PI and P2; s2 = secondary cord between P2 and P3, etc.

ADP :

adapertural primary cord on the siphonal canal

MP:

médian primary cord on the siphonal canal

ABP :

abapertural primary cord on the siphonal canal

ads :

adapertural secondary cord on the siphonal canal

ms :

médian secondary cord on the siphonal canal

abs :

abapertural secondary cord on the siphonal canal

eabs :

extreme abapertural secondary cord on the siphonal canal

Example: eabsl = secondary cord between EABP and EABP1

APERTURE

ID:

Infrasutural denticle

DI to D6:

Abapical denticles

Abbreviations

BMNH: Natural History Muséum, London, U,K.

IRSNB: Institut royal des Sciences naturelles de

Belgique, Bruxelles, Belgium.

MNHN: Muséum national d'Histoire naturelle, Paris,

France.

JC: Collection of Jacques Colomb, Marseille, France.

RH: Collection of Roland Houart.

SG: Collection of Sando Gori.

Figure 1 . Terminology of the spiral sculpture

morphology in Favartia colombi n. sp.

SYSTEMATICS

Family MURICIDAE Rafinesque, 1815

Subfamily MURICOPSINAE Radwin & D'Attilio,

1971

Genus Favartia Jousseaume, 1880

Type species by original désignation: Murex

breviculus Sowerby, 1834, Recent, Indo-West Pacific.

Favartia (Favartia) colombi n. sp.

Figs 1,2-6, 27

Pygmaepterys yemenensis - Bosch et al. 1995: 120,

fig. 478.

Type material. Oman, Masirah Island, Ras Al Ya,

20°29’42" N, 58°57'029" E, 21 m, on coarse sand,

holotype IRSNB IG 31676/ MT 2316.

Paratypes: Oman, Masirah Island, Ras Al Ya,

20°29’42" N, 58°57'029" E, 21 m, on coarse sand, 1

MNHN 23206; Sur Masirah, east of Masirah Island, 1

m, low tide, under stones, 1 BMNH 20100521 ; Oman,

Masirah Island, Ras Al Ya, 20°39’504" N, 58°52’138"

E, 21 m, on sand 4 RH, 7 SG; Masirah Island, 2 m, 1

JC, 1 RH.

Other material examined. Oman, Masirah Island,

Ras Al Ya, 20°32’895" N, 58°57’420" E, 13 m, on

sand (8 SG, 2 RH); Masirah Island (3 JC); Masirah

Island, 2 m (2 JC); Sur Masirah, east of Masirah, 1 m,

low tide, under stones (15 JC); North of Muscat,

Mina al Fahl, 24-30 m, dredged, sand and mud (4

RH).

R. Houart & S. Gori

NOVAPEX 12(1-2): 39-45, 10 mars 2011

Distribution. Oman, north of Muscat and Masirah

Island.

Description. Shell medium-sized for the genus, up to

16.5 mm in height at maturity. Height/width ratio

1.63-1.93:1. Slender, lanceolate, shoulder weakly

sloping, weakly concave.

Greyish-white or tan with lighter colored primary

spiral cords. Aperture light brown within.

Spire high with 1.6 protoconch whorls and teleoconch

up to 5 weakly angular, strongly shouldered whorls.

Suture impressed, partially obscured by third primary

spiral cord (P3) of previous whorl. Protoconch small.

Whorls with a single strong but narrow adapical keel.

Terminal lip thin, erect, weakly curved.

Axial sculpture of teleoconch whorls consisting of

prominent, narrow varices, each with short, scabrous,

open spinelets decreasing in strength and length

abapically, connected to each other with scabrous

flange. Nine varices from first to third teleoconch

whorl, decreasing to 6-8 on fourth whorl and 4 or 5 on

last whorl. Spiral sculpture of high, strong, narrow,

smooth primary cords and occasionally very narrow

secondary cords. First and second whorls with visible

PI and P2 or P1-P3. Third and fourth whorls of

juvénile specimens with PI, P2, s2, P3, P4, P5, P6,

ADP and MP. Fifth whorl with PI. (si), P2, s2, P3,

(s3), P4, P5, s5, P6, (s6). ADP, MP, ABP. Shoulder

ramp smooth except for axial varices. P1-P4 almost

similar in size and strength, P4 and P5 close to each

other, P5 weakly narrower, P6 narrower spiral cord,

almost half the size of P1-P4; ADP and MP broad.

Primary cords more obvious on adapertural part of

varices. Secondary cords very narrow, only obvious

on varices.

Aperture moderately small, ovate. Columellar lip

narrow, smooth, rim partially erect at abapical

extremity, otherwise adhèrent. Anal notch shallow,

broad. Outer lip weakly erect, weakly crenulated, with

weak, elongate denticles within: ID, DI or DI split,

D2 split, D3 split, D4 and D5. 1D very low, DI low,

D2-D5 increasing in strength abapically. Siphonal

canal short, 23-29% of total shell height, narrow,

weakly dorsally recurved at tip, narrowly open, with 2

or 3 short scabrous spines originating from ADP, MP

and ABP. ADP broadest.

Operculum and radula unknown.

Remarks. Other Favartia s.s. species occur in the

western Indian Océan, mostly along the East African

coast, up to Somalia and the Red Sea. The species

most similar to F. colombi n. sp. are Favartia

(Favartia) cecalupoi Bozzetti, 1993, F. cyclostoma

(Sowerby, 1841), F. nucula (Reeve, 1845), F. peasei

(Tryon, 1880), F. rosamiae D'Attilio & Myers, 1985

and F. sykesi (Preston, 1904).

Favartia cecalupoi (Fig. 13) from Somalia differs

in many respects i.e. the protoconch which consists of

2 smooth bulbous whorls. The shell is more rounded

with a broader last teleoconch whorl, the varices are

lower and the spiral cords are more numerous. F.

cyclostoma is larger relative to the number of

teleoconch whorls, the shell is also much broader with

a rounded paucispiral protoconch of 1.5 whorls. F.

nucula, F. rosamiae , F. peasei and F. sykesi differ in

many respects such as breadth and height of the shell,

axial and spiral sculpture and mainly by having

different protoconch whorls, small, rounded and

paucispiral in F. nucula and F. peasei, conical and

multispiral in F. rosamiae (Figs 14, 32) and F. sykesi.

Favartia (Pygmaepterys) paulboschi Smythe &

Flouart, 1984 (Figs 15-16, 28), another Favartia

species from Oman differs in having a relatively

narrower shell vs its height, broader and fiat primary

spiral cords, fewer or absence of secondary cords,

narrower varices with more adapically recurved

shoulder spine, and relatively shorter and broader

siphonal canal.

Favartia (F.) colombi was also confused with, and

wrongly identified as F. (P.) yemenensis Houart &

Wranik, 1989 (Figs 17-18, 29) (Houart, in litt.) which

also has a keeled protoconch, and therefore was also

illustrated as Pygmaepterys yemenensis in Bosch et al.

(1995: 120). However, F. yemenensis differs from F.

colombi n. sp. in having a more angular and broader

shell with higher and narrower varices, in having a

comparatively broader aperture, a longer siphonal

canal and more numerous secondary spiral cords.

Favartia (Pygmaepterys) adenensis (Fig. 19) also

occur in this area but is very different and does not

need to be compared here.

Etymology. colombi'. Named after Jacques Colomb

(Marseille, France) who collected several specimens

of the new species in Masirah, together with the junior

author.

Favartia (Favartia) roseotincta n.sp.

Figs 7-12, 30

Type material. Oman, Masirah, Ras Al Ya,

20°39’504" N, 58°52’138" E, 21 m, on sand, holotype

MNHN 23205

Paratypes: 3 RH, 3 SG.

Distribution. Oman, Masirah Island.

Description. Shell small for the genus, up to 8.3 mm

in length at maturity. Height/width ratio 1.66 - 1.86:1,

biconical, narrow, weakly spinose. Shoulder strongly

sloping, weakly convex.

Pinkish white or light pink, occasionally with white

varices and white siphonal canal. Aperture pink

within, surrounded with white on columellar lip outer

rim and outer apertural lip.

Spire high with 1.5 protoconch whorls and teleoconch

up to 5 weakly convex, angular, strongly shouldered

whorls. Suture impressed. Protoconch small, elongate,

whorls rounded, last whorl strongly elongate.

Terminal lip eroded.

R. Houart & S. Gori

Two new Favartia species from Masirah Island

Axial sculpture of teleoconch whorls consisting of

high, strong, narrow varices, each witli short, broad,

blunt, open primary spines. First whorl with 8 varices,

second to fourth with 8 or 9 varices, last whorl with 4

varices. Varices of last whorl broader and more distant

from each other than in other whorls, with a large gap

between third and fourth (apertural) varices, almost

twice the distance between second and third varices.

Spiral cords from First to penultimate teleoconch

whorl of high, narrow, smooth primary cords. First

and second whorls with visible PI and P2, or P1-P3,

P3 partially covered with next whorl, third and fourth

whorl with visible P1-P3, P3 still partially covered by

next whorl. Last whorl with IP, P1-P5, (s5), P6, (s6),

ADP, MP. IP only weakly visible on axial varices,

P1-P3 almost of same strength, P3 occasionally

narrower, P4-P6 decreasing in strength abapically, P4

and P5 high, broad, P6 very small, almost obsolète,

occasionally preceded by almost obsolète s5.

Aperture small, ovate, columellar lip narrow, smooth,

lip partially erect, adhèrent on shell at 30-40% of

adapical extremity. Anal notch moderately broad.

Outer lip erect, with weak elongate, narrow denticles

within: ID (split), DI split, D2 (split), D3, D4, D5.

Siphonal canal short, narrow, strongly dorsal ly

recurved at tip, narrowly open, with ADP and MP

giving rise to short, open, blunt spines.

Operculum and radula unknown.

Remarks. A few species of Favartia hâve an elongate

and narrow shell such as observed in F. roseotincta n.

sp., namely F. flexirostris (Melvill, 1898), F.

guamensis Emerson & D’Attilio, 1979, F. iredalei

Ponder, 1972, F. jeanae Bertsch & D’Attilio, 1980

and F. peregrina (Olivera, 1980).

Favartia flexirostris (Figs 20, 31) which also

occurs in Oman, differs from F. roseotincta n. sp. by

many shell characters, namely in having a rounded,

globose protoconch (Fig. 31), more rounded

teleoconch whorls with a globose last whorl and a

rounded aperture, numerous, broad, secondary spiral

cords on the whole shell, and a longer, more strongly

tapered siphonal canal.

Favartia guamensis differs also in many ways,

namely by having a very different, conical protoconch

of sinusigeral type, consisting of 3-3.5 whorls,

denoting planktotrophic larval development. Its

geographical distribution is therefore also much more

expanded, ranging from off Mozambique (East Africa)

to the Tuamotus (Central Pacific). Other différences

are numerous such as the lower and broader spire

whorls, the narrower and straighter shoulder ramp, the

broader and more squamous primary spiral cords, the

small, rounded aperture with a characteristic narrow,

deep anal notch and the single spine (ADP) on the

narrower siphonal canal.

Favartia iredalei (Fig. 21) from Lord Howe

Island, NSW, Australia and West Sumatra also has a

different protoconch of 1.5 rounded whorls. It differs

also in having a less sloping and narrower shoulder

ramp with IP, or adis and IP, and in having more

squamous primary cords and obvious secondary cords

on the convex part of the last teleoconch whorl.

Both F. jeanae and F. peregrina from the

Philippines also differ in having a different larval

development, denoted by their conical, multispiral

protoconch of 3-3.5 whorls and a terminal lip of

sinusigeral type. The shell morphology of F. jeanae is

close to F. roseotincta n.sp but besides the protoconch

différences, the shell differs in having a comparatively

higher spire, narrower primary spiral cords, obvious

adis and IP on the shoulder ramp, and a narrower,

tapered siphonal canal.

Favartia peregrina differs in many ways and does

not need to be compared further here.

Another species from Elat (Red Sea), was named

Favartia elatensis Emerson & D'Attilio, 1979 but it

actually tumed out to be a Mitrexsid species (Houart,

1994, Merle & Houart, 2003) and differs from F.

roseotincta in many respects.

Figures 2-22

2-6. Favartia (Favartia) colombi n. sp.

2-3. Oman, Masirah Island, Ras Al Ya, 20°29’42" N, 58°57'029" E, 21 m, holotype IRSNB IG 31676/MT 2316,

15.6 mm; 4-5. Oman, Masirah Island, Ras Al Ya, 20°39’504" N, 58°52’138" E, 21 m, paratype RH 12 mm; 6.

Oman, Masirah Island, Ras Al Ya, 20°29’42" N, 58°57'029" E, 21 m, paratype MNHN 23206

7-12. Favartia (Favartia) roseotincta n. sp., Oman, Masirah, Ras Al Ya, 20°39’504" N, 58 0 52’138” E, 21 m.

7-8. Holotype MNHN 23205, 8.3 mm; 9-10. Paratype SG, 8.2 mm; 11-12. Paratype RH, 8.1 mm.

13. Favartia (F.) cecalupoi Bozzetti, 1993, Ras Hafun, Somalia, SG, 12.4 mm; 14. Favartia (F.) rosamiae

D Attiho & Myers, 1985, Seychelles, RH, 15.6 mm; 15-16. Favartia (F.) paulboschi Smythe & Houart, 1984,

iuw\\a Masirah, Oman, paiatype RH, 17 mm; 17-18. Favartia (Pygmaepterys) yemenensis (Houart &

ïamk, 1989); 17. Gulf ot Aden, RH, 23 mm; 18. Oman, Masirah Island, Ras Al Ya SG 16 7 mm - 19

Favartia (P.) adenensis (Houart & Wranik, 1989), Gulf Of. Aden, PDR Yemen, paratypeRH. 15.4 mm; 20.

!" (F)flexirostris (Melvill, 1898), Oman, N Of. Muscat, RH, 11.3 mm; 21. Favartia (F.) iredalei Ponder,

. f Australla - NSW, Lord Howe Island, RH, 6.1 mm; 22. Favartia (F.) jeanae Bertsch & D'Attilio. 1980,

Philippines, Cebu, RH, 8.5mm.

R. Houart & S. Gori

NOVAPEX 12(1-2): 39-45, 10 mars 2011

R. Houart & S. Gori

Two new Favartia species from Masirah Island

Favartia colombi n.sp. described above as

occurring in the same area differs in being larger with

a similar number of teleoconch whorls, in having a

different protoconch morphology (Figs 27 vs 30) and

in having more numerous secondary spiral cords.

The variable spacing between penultimate and last

(apertural) varix is an unusual feature observed in F.

roseotincta. Such unusual cases of a distinct larger

gap between these two varices are seen in two other

muricopsine species: Favartia rosamiae (Fig. 14) and

F. varimutabilis Houart, 1991 from the Lesser Antilles

and southeastern Brazil.

Etymology. roseotincta (L): Named for its color

which is reminiscent of a beautiful pink and white

rose.

Figures 23-32

23-25. Favartia (Favartia) marjoriae (Melvill & Standen, 1903)

23. Persian Gulf, Sheikh Shuaib Island, 15 fms (27 m), syntype BMNH 1903.11.5.18.19, 27.5 mm.; 24-25.

Persian Gulf, IRSNB IG 10591, 29 mm.

26. Favartia maculata (Reeve, 1845), Thailand, Phuket Island, RH, 18.7 mm.

27-32. Protoconchs (scale bars 0.5 mm)

27. Favartia (Favartia) colombi n.sp (RH); 28. Favartia (Pygmaepterys) paulboschi (RH); 29. Favartia (P.)

yemenensis (SG); 30. Favartia (F.) roseotincta n. sp. (SG); 31. Favartia (F.) flexirostris (RH); 32. Favartia (F.)

rosamiae (RH).

R. HOU ART & S. GORI

NOVAPEX 12(1-2): 39-45, 10 mars 2011

ACKNOWLEDGEMENTS. We are very thankful to

Jacques Colomb, Marseille, France, for the loan of his

material and the gift of two specimens, to Greg

Herbert, University of South Florida, Tampa, U.S.A.

for his useful comments on the ms and to John Wolff,

Lancaster, Pennsylvania, U.S.A., for checking the

English text.

REFERENCES

Biggs, H. E. J. 1973. The marine Mollusca of the

Trucial coast, Persian Gulf. Bulletin of the British

Muséum (Natural History), ZoologyV ol. 24(8):

343-421,6 pis.

Bosch, D. & Bosch, E. 1982. Seashells of Oman.

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Bosch, D. & Bosch, E. 1989. Seashells of Southern

Arabia. Motivate Publishing, U.A.E. pp. 1 -95.

Bosch, D.T., Dance, S.P, Moolenbeek, R.G & Oliver,

PG. 1995. Seashells of Eastem Arabia. Ed. P.

Dance, Motivate Publishing. pp. 1-296.

Houart, R. 1994. Illustrated catalogue of Recent

species ofMuricidae named since 1971.

Wiesbaden: 1-179.

Merle, D. 1999. La radiation des Muricidae

(Gastropoda : Neogastropoda) au Paléogène:

approche phylogénétique et évolutive. Paris.

Unpublished thesis, Muséum national d'Histoire

naturelle : i-vi, 499 pp.

Merle, D. 2001. The spiral cords and the internai

denticles of the outer lip in the Muricidae:

terminology and methodological comments.

Novapex 2 (3): 69-91.

Merle, D. & Flouart, R. 2003. Ontogenetic changes of

the spiral cords as keys innovation of the muricid

sculptural patterns: the example of the Muricopsis-

Murexsul lineages (Gastropoda: Muricidae:

Muricopsinae). C.R. Palevol. 2: 547-561.

Sharabati, D. 1981. Saudi Arabian seashells. Selected

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G. T. Watters

Novapex 12(1-2): 47-48, 10 mars 2011

Redescription and range extension of Antillophos bahamasensis Petuch, 2002

(Gastropoda: Buccinidae)

G. Thomas WATTERS

Department of Evolution, Ecology and Organismal Biology

Ohio State University

Columbus, OH USA 43212

Watters. 1 @osu.edu

KEYWORDS. Caribbean Sea, Buccinidae, Antillophos

ABSTRACT. Antillophos bahamasensis Petuch, 2002, previously known only from the type

locality, is recorded from Guadeloupe. The species is redescribed based on this additional material.

Watters (2009) reviewed the western Atlantic

Océan species of Antillophos. At the time the species

Antillophos bahamasensis Petuch, 2002, was known

only from the faded holotype and a single paratype

collected off Victory Cay, Bimini Chain, Bahamas,

apparently from 35 m (Figures 1, 2). Because of the

lack of material a description of the variability of the

species was not possible. It was suggested that the

species might even prove to be synonymous with the

widespread Antillophos chazaliei (Dautzenberg,

1900).

After publication of the review I was fortunate to

receive numerous lots of Antillophos from

Guadeloupe sent by Mr. Dominique Lamy. Five

species were represented, some in abundance: A.

chazaliei (Dautzenberg, 1900), A. candeanus

(d’Orbigny, 1842), A. smithi (Watson, 1885), A.

beauii (Fischer and Bemardi, 1857), and surprisingly,

A. bahamasensis. Specimens of A. bahamasensis were

recorded from Saint-Françoise, Grande-Terre (250 m)

(Figure 3, 4), Basse-Terre (350 m) (Figure 5), and

Port-Louis, Grande-Terre (150 m). At least one

specimen was live-taken and the species was found

either alone or in association with A. smithi and A.

beauii. Antillophos bahamasensis appears to live in

deeper water than most western Atlantic Antillophos

and is obviously very rare in collections. These are

also the fi rst published records of A. candeanus and A.

chazaliei from Guadeloupe, although well within the

overall range of both species.

These new specimens not only increase the range

of the species significantly, but indicate that the

species is distinct from A. chazaliei, as well as

allowing a more comprehensive description of the

species beyond the type lot. Although quite similar, A.

bahamasensis differs from A. chazaliei (Figure 6) in

having more numerous axial ribs on the penultimate

whorl (11 — 17 in A. bahamasensis, 8 - 12 in A.

chazaliei), in having less pronounced sculpture, and in

having the lirae in the outer lip broken into pustules

(entire in A. chazaliei). The description given by

Watters (2009) is herein modified and additional

specimens are figured.

Description. Shell 12.8 - 18.0 mm in length.

Fusiform; spire 50% - 60% of total length.

Protoconch conical, of 2.25 - 2.5 smooth whorls with

sharp keel at periphery near suture. Teleoconch of 6.5

whorls. Teleoconch whorls sculptured with narrow,

widely spaced, fiat, spiral cords separated by wide

intervals, 14 - 17 on last whorl. Some interspaces with

a single, fine, 2° spiral thread. Axial sculpture of

widely-spaced, low, rounded ribs, 11 - 19 on last

whorl (excluding varices) and 11 - 17 on penultimate

whorl (excluding varices). Varices well-developed,

about one varix every 1/3 - 1/4 tum including final

last whorl (not apparent on holotype); occasionally

varices may be adjacent to each other. Terminal varix

low, wide, crossed by numerous axial ribs.

Intersections of axial and spiral sculpture form

pustulose, rachet-like sculpture. Aperture elongate-

oval, with one plication anteriorly; anal canal set off

by one or two denticles. Outer lip with 12-17 lirae

deep within mouth, with oceasional intercalated 2°

ones; the primary lirae are broken up into peculiar

linear pustules. Columella continuous; pariétal lip

adhèrent to previous whorl. Siphonal canal short,

open. “Stromboid notch” shallow to of medium depth.

Base color grayish-white, most specimens hâve some

evidence of three brown spiral bands at suture,

periphery, and siphonal canal; terminal varix always

white without spots. Aperture white. Operculum leaf-

shaped, tan, with anterior terminal nucléus. Radula

and anatomy unknown.

Acknowledgements

I thank Mr. Dominique Lamy for the opportunity to

study these important specimens.

G. T. Watters

Redescription of Antillophos bahamensis Petuch, 2002

Figures 1-6

1-5. Antillophos bahamasensis Petuch, 2002.

1-2. Holotype UF 277198, off Victory Cay, Bimini Chain, Bahamas, 18.0 mm length; 3-4. Watters coll. 14435a,

250 m, Saint-Françoise, Grande-Terre, 17.3 mm length; 5. BMSM 42999, 350 m, Basse-Terre, 12.9 mm length.

6 . Antillophos chazaliei (Dautzenberg, 1900). Watters coll. 13749d, 150 m, Guadeloupe, 15.7 mm length.

REFERENCES

Petuch, E. J. 2002. New deep water gastropods from

the Bimini Shelf, Bimini Chain, Bahamas.

Ruthenica 12: 59-65, figs. 1,2.

Watters, G.T. 2009. A révision of the western Atlantic

Océan généra Anna, Antillophos, Bailyci,

Caducifer , Monostiolum, and Parviphos , with

description of a new genus, Dianthiphos, and notes

on Engina and Hesperistemia (Gastropoda:

Buccinidae: Pisaniinae) and Cumia

(Colubrariidae). Nantiliis 123: 225-275.

C. VlLVENS, F. SWINNEN & F. DENIZ GUERRA

Novapex 12(1-2): 49-55, 10 mars 2011

A new species of Clelandella

(Gastropoda: Trochoidea: Trochidae: Cantharidinae)

from Western Sahara

Claude VlLVENS

Rue de Hermalle, 113 - B-4680 Oupeye, Belgium

Scientific Collaborator, Muséum national d'Histoire naturelle, Paris

vilvens.claude@skynet.be

Frank SWINNEN

Lutlommel, 10 - B-3920 Lommel, Belgium

Scientific Collaborator, Museu Municipal do Funchal, Madeira

f.swinnen@skynet.be

Francisco DENIZ GUERRA

Avda de la Democracia 47 - CP 35018 Las Palmas de Gran Canada, Spain

fdeniz@telefonica.net

KEYWORDS. Gastropoda, Trochoidea, Trochidae, Cantharidinae, Western Sahara, Clelandella,

new species.

ABSTRACT. Clelandella artilesi n. sp., a new species from Western Sahara is described and

compared with similar species from the north-eastem Atlantic, especially off West Africa.

RESUME. Une nouvelle espèce de Clelandella provenant du Sahara Occidental, Clelandella

artilesi n. sp., est décrite et comparée avec d'autres espèces similaires de l'Atlantique Nord-est,

plus particulièrement au large de l'Afrique Occidentale.

INTRODUCTION

Until the next last years, there was only a few

literature about the malacofauna from off former

French West Africa and the species from this area

were still rather poorly known (see Vilvens &

Swinnen, 2007 for an abbreviated historical account of

the main expéditions in this area). By now, some

valuable Works hâve highlighted the malacofauna not

only from West Africa but also from adjacent Atlantic

islands such as Canary Is., Madeira Is. and Cape

Verde Is. (e.g. Curini-Galletti, 1985; Segers, 2002;

Ardovini & Cossignani, 2004; Gofas, 2005; Rolan.

2005; Rolan & Swinnen. 2009; Segers, Swinnen & De

Prins, 2009).

Arnong the species reported or newly described in

ail these papers and books, the Cantharidinae

(accepted subfamily of the Trochidae family) species

are rather numerous. This paper describes a new

species that belongs to this subfamily.

Material and methods

The material studied was found by some dredging

performed by dragging boats working in the Canary -

Saharan fishing bank. Ail the shells where collected

between 1993 and 2001, in an area off Western Sahara

that goes from Cap Boujdour (Cabo Bojador: 22°18'N,

16°40'W) to Cap Barbas (Cabo Barbas: 26°07N,

14°29'W), at between 50-70 m deep on sandy bottoms

(see text figure 2 below).

Regarding the description methodology, the main

conchological features used are (see text figure 1

below) :

♦ general shape of the shell;

♦ shape of the whorls;

♦ spiral cords and axial threads of the whorls;

♦ spiral cords on the base;

♦ shape of the aperture, the outer and the inner lip;

♦ colour pattern.

C. VlLVENS, F. SWINNEN & F. DENIZ GUERRA

A new species of Clelandella

Text figure 1 : Features of Cantharidinae shells; FI :

height; W : width; HA : height ofthe aperture; PI. P2,

P3,... : primary cords; SI, S2, S3, ... : secondary

cords (shell : Clelandella miliaris (Brocchi, 1814),

Brittany, Roscoff, 9.4 x 8.0 mm).

Statistical calculations hâve been performed with

Microsoft Excel 2003 and Tanagra 1.4, a free

statistical software from Lyon II University.

Abbreviations

Repository

IRSNB : Institut royal des Sciences naturelles de

Belgique, Brussels, Belgium.

MNHN : Muséum national d'Histoire naturelle, Paris,

France.

Other abbreviations

H : height

W : width

TW : number of teleoconch whorls

PI, P2, P3,...: primary cords (PI is the most adapical)

SI. S2, S3,...: secondary cords (SI is the most

adapical)

lv : live-taken specimen(s) présent in sample

dd : only dead specimen(s) présent in sample

sub : subadult specimen(s)

juv : juvénile specimen(s)

SYSTEMATICS

We follow here the new classification of Williams et

al. (2010) who has elevated the Cantharidini tribe (as

it was considered by Hickman & McLean, 1990 and

still by Bouchet & Rocroi (2005)) at a subfamily level

of the Trochidae.

Text figure 2 : Prospecting area in this paper

Superfamily : TROCHOIDEA Rafinesque, 1815

Family : TROCHIDAE Rafinesque, 1815

Subfamily : CANTHARIDINAE Gray, 1857

Genus : Clelandella Winckworth, 1932

Type species : Trochus clelandi W.Wood, 1828

(=Tmchus miliaris Brocchi, 1814) (by o.d.)-

Pliocene, northern Italy.

Distinctive features. Shell rather small (height up

to 14 mm for the known species), elevated spire

almost as high as wide, conical shape with fiat whorls.

Smooth protoconch with a thin terminal varix. Rather

strong spiral sculpture of beaded spiral cords. Angular

to subangular periphery. interrupted peristome,

aperture prosocline, rhomboidal. Base from almost fiat

to moderately convex. No umbilicus or umbilicus very

narrow. Variable colour, with various patterns.

Remarks. Gofas (2005) described recently 4 new

species from north-western Atlantic and 1 new species

from eastern Mediterranean.

C. VlLVENS, F. SWINNEN & F. DENIZ GUERRA

Novapex 12(1-2): 49-55, 10 mars 2011

Clelandella miliaris (Brocchi, 1814)

Figs 7-14, Text Figs 1, 3

Trochus miliaris Brocchi, 1814: 353, pl. 6, fig. 1.

Type locality: northern Italy, no

specified locality; lectotype from qualification as

holotype.

Trochus clelandi — W. Wood, 1828: 16, pl. 5, fig. 15.

Trochus millegranus - Philippi, 1836: 183, pl. 10, fig.

25.

Calliostoma clelandi - Kaicher, 1979: card#2094.

Calliostoma miliaris - Poppe & Goto. 1991: 74, pl.6,

fig. 9.

Clelandella miliaris - Giannuzzi-Savelli et al., 1994:

88, figs.279-281.

Clelandella miliaris - Cretella et al., 1999 : 12-1A,

figs. 13-15, 17-18, 25,28,33-34.

Calliostoma miliaris - Ardovini & Cossignani, 2004:

71.

Clelandella miliaris - Gofas, 2005: 134-136, figs. 1—

3, 12A-B, 13A-B.

Clelandella miliaris - WoRMS, 2011 :

p=taxdetails&id= 141774.

Material examined. France. Britanny, Finistère, near

Ouessant, +/- 100 m, 40 dd. - Roscoff, +/- 3 m, 25 lv.

Italy. Tuscany Archipelago, 2 lv. Spain. Off Malaga,

2 lv. West Africa. Off Mauritania, 17°45'N, 16°25'W,

3 lv, 1 juv lv.

Distribution. North-eastern Atlantic, from Norway to

West Africa and Mediterranean Sea, 35-800 m.

Remarks. The main characteristics of this species

are :

- height up to 14 mm, width up to 13.8 mm;

- a high spire, a conical shape, with up to 7 fiat

whorls;

- strongly prosocline axial threads and up to 8 spiral

cords, granular by intersection with axial threads; on

First whorl, 4 cords Pl, P2, P3 and P4 appearing

immediately, smooth; Pl slightly weaker than other

cords; cords granular by intersection with axial

threads as soon as second whorl; secondary cords S2

and S3 appearing by intercalation; P4 thicker on third

whorl, dividing into 2 cords on fourth whorl, up to 10

cords on last whorl of big specimens, producing a

thick rim making carina;

- columella nearly straight, oblique, with a weak

médian swelling;

- base almost fiat, with 8 to 13 granular spiral cords;

- no umbilicus or very narrow umbilicus reduced to a

small slit.

Nicklès (1950) reviewed the known marine species

of Western and Equatorial Africa but curiously didn't

mention this species. Kaicher (1979) still used the

synonym name. At last, Cretella et al. (1999)

produced an accurate study of this species.

This is a variable species (Gofas, 2005) regarding size

and colour, also sometimes with a slightly

cyrtoconoidal shape instead of a conical one and a

subangular instead of angular periphery (some

specimens from western Africa). Gofas mention even

(without illustrations) some specimens from Ivory

Coast with a narrow umbilicus.

Clelandella artilesi n. sp.

Figs 1-4, Table 1, Text Fig. 2, 3

Type material. Holotype (6.9 x 7.2 mm) 1RSNB

1G.31784/MT2330 for database DaRWIN. Paratypes:

1 MNHN 23351, 2 coll. F. Deniz Guerra, 2 coll.

F.Swinnen, 1 coll. C.Vilvens.

Type locality. West Africa. Off Western Sahara, 50-

60 m.

Material examined. West Africa. Off Western

Sahara, 12/1999, 50-60 m, 2 dd sub, 3 dd juv. - Off

Western Sahara, 7/2001, 50-60 m, 3 dd (with holotype

and 2 paratypes). - Off Western Sahara, 1993-1999,

50-70 m, 15 dd, 7 dd sub, 2 dd juv (with 4 paratypes).

Distribution. West Africa, off Western Sahara, 50-60

Diagnosis. A typical cantharid species with an

elevated, conical spire, up to 6 even granular spiral

cords on the whorls, an angular periphery, up to 6 or 7

smooth spiral cords on the base and a very narrow or

closed umbilicus.

Description. Shell of moderate size for the genus

(height up to 7.2 mm, width up to 5.1 mm),

higher than wide, conical; spire elevated, height 1,2x

to 1.4x width; angulate periphery; umbilicus closed or

very narrow.

Protoconch about 180 pm wide, of 1 to 1.25 whorls,

rounded, smooth with a thin, straight terminal lip.

Teleoconch of up to 6.9 whorls; two first whorls

convex, other whorls almost straight, with spiral cords

first smooth, granular later and axial threads much

more stronger on abapical whorls than on adapical

whorls. Suture visible, not canaliculated. First whorl

convex, without axial sculpture and 4 smooth cords Pi

(i=l,2,3,4) appearing almost immediately, Pl slightly

later and weaker; interspace between cords about 1.5

larger than cords; ail cord brown except Pl lighter;

axial threads very weak, hard to detect; suture

impressed, not canaliculated. On second whorl, S2

appearing at first midwhorl, S3 at second midwhorl,

both quickly similar to Pi; ail cords still smooth. On

third whorl, ail cords more or less similar in size,

except P4 slightly stronger; distance between cords

similar to cords; S4 may appear, partly covered by

next whorl; weakly prosocline axial threads clearly

visible on abapical part, especially between S3, P4 and

S4. Shape of whorl weakly convex, almost straight

near end. On fourth whorl, axial thread stronger

between spiral cords; threads still weakly prosocline;

C. VlLVENS, F. SWINNEN & F. DENIZ GUERRA

A new species of Clelandella

cords subgranular. On next whorls, spiral cords

granular; interspaces almost as wide as cords, except

interspace between S3 and P4 slightly greater. On last

whorl, beads of P4 (sometimes also of S3) sharp; S4

appearing, thin, almost smooth to weakly subgranular,

close to P4.

Aperture roundly rhomboidal; outer lip thin; inside

nacreous.

Columella nearly straight, only slightly oblique, with a

weak basal swelling forming a kind of blunt tooth.

Base weakly convex, with thin axial threads on whole

surface and with 6 or 7 rather low, spiral cords; cords

similar in size to the cords on whorls, weakly

subgranular by intersection with axial threads;

interspaces between cords grater than or equal to

cords; axial threads much thinner and more crowded

than axial threads on whorls.

No umbilicus or very narrow umbilicus reduced to a

small sût.

Colour of teleoconch pinkish white, sometimes with

brownish fiâmes on First whorls only and with

brownish dashes on peripheral cords, or sometimes

with brownish fiâmes on the whole whorls;

protoconch off-white.

H/W

TW/H

holotype IRSNB

6.9

7.2

5.1

1.41

0.96

paratype MNHN

6.7

5.9

4.9

1.20

1.14

paratype FDG-1

6.5

5.8

4.6

1.26

1.12

paratype FDG-2

6.4

6.8

1.36

0.94

paratype FS-1

6.6

6.4

1.28

1.03

paratype FS-2

6.5

6.2

4.9

1.27

1.05

paratype CV

6.5

6.2

4.7

1.32

1.05

mecrns

6.59

6.36

4.89

1.30

1.04

minima

6.40

5.80

4.60

1.20

0.94

maxima

6.90

7.20

5.10

1.41

1.14

standard déviation

0.17

0.50

0.18

0.07

Table 1. - Clelandella artilesi n. sp.: Shells measurements in mm for types (FDG : F. Deniz Guerra coll.; FS : F.

Swinnen coll.; CV : C. Vilvens coll.).

Discussion. The new species is close to Clelandella

miliaris (Brocchi, 1814) (Figs 7-14) but differs from

it mainly by a greater height/width ratio (about 1.30

instead of about 1.15 for C. miliaris — see scatter plot

there under showing the different slope for the

régression lines), a spiral cord P4 similar in size to the

other cords (instead of being much stronger,

producing usually a prominent peripheral rim) never

divided in or covered by thinner spiral cords (3-10

such cords are présent on C. miliaris), axial threads

weakly prosocline with an angle of about 30° with

vertical (instead of 45°).

Plate 1. Figures 1-12. Scale bar: 1 mm.

1-4. Clelandella artilesi n. sp., West Africa. Off Western Sahara, 50-60 m. 1-2. Holotype IRSNB (IG.31784),

7.2 x 5.1 mm. 3-4. Paratype MNHN (23351), 5.9x4.9 mm.

C- madeirensis Gofas, 2005, holotype MNF1N, Madeira archipelago, off Porto Santo, 430 m, 8.3 x 7.0 mm

7-14. C. miliaris (Brocchi, 1814). 7-8. Off Mauritania, 6.2 x 5.7 mm. 9-10. France, Britanny, Finistère, near

Ouessant, 8.7 x 7.7 mm. 11-12. Italy, Tuscany Archipelago, 9.1 x 7.1 mm. 13-14. Spain, off Malaga, 15.2 x 12.8

mm.

C. VlLVENS, F. SWINNEN & F. DEN1Z GUERRA

NOVAPEX 12(1-2): 49-55, 10 mars 2011

C. VlLVENS, F. SWINNEN & F. DENIZ GUERRA

A new species of Clelandella

Text Figure 3 : Scatter plot for height (H) and width (W)

examined specimens of C. miliaris.

Clelandella artilesi n. sp. is also rather close to

Jujubinus catenatus Ardovini, 2006 from Sicily, but

this similar in size species has a more elevated spire

(with a greater H/W ratio of about 1.50), a slightly

cyrtoconoidal shape, a transversely elongated flared

aperture, thinner spiral cords on the base and a

different coloration with reddish brown cords on a

light green background.

The new species may also be compared to

Clelandella madeirensis Gofas, 2005 (Figs 5-6) from

Madeira, but this species is slightly greater (height up

to 8.3 mm for a similar number of whorls), has a

different ontogeny of the cords (S3 appearing first, S2

later and 2 tertiary cords appearing and growing to

reach the same size as the other cords, giving 9 spiral

cords on the last whorl ol the holotype), lacks the

axial threads between the spiral cords and has more

numerous (up to 12 or 13), uneven and thinner spiral

cords on the base.

Etymology. After Miguel Artiles, collector from

Arinaga, South East of the Island of Gran Canaria,

interested in the marine malacofauna of the entire

world and occasional collaborator of the third author

for the study of land shells from Canarias.

Acknowledgements

We would like to thank P. Bouchet (Muséum national

d Histoire naturelle, Paris) for access to the

malacological resources of the MNHN and V. Héros

(MNHN) for lier help in borrowing types and finding

various scientific papers.

Also, we are grateful to T. Backeljau (Institut royal

des Sciences naturelles de Belgique, Bruxelles) for his

constant help in borrowing types.

of types Clelandella artilesi n. sp. of and some

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Bouchet, P. & Rocroi, J.P. 2005. Classification and

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Gofas, S. 2005. Geographical différentiation in

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In: taxonomie database of the European Marine

Mollusca of the Department of Systematics &

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http://www.somali.asso.fr/clemam/biotaxis.php on

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In: Bouchet, P.; Gofas, S.; Rosenberg, G. (2010)

World Marine Mollusca database. Accessed

through: World Register of Marine Species at

http://www.marinespecies.org/aphia.php?p=taxdet

ails&id=141 774 on 2011-01-13

C. VlLVENS, F. SWINNEN & F. DENIZ GUERRA

Novapex 12(1-2): 49-55, 10 mars 2011

Hickman, C.S. & Mc Lean, J.H. 1990. Systematic

révision and suprageneric classification of

trochacean gasteropods. Natural History Muséum

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côte occidentale d'Afrique, Ed. P. Lechevalier. 269

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Rolan, E. & Swinnen, F. 2009. Two new species of

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Segers, W. 2002. On some shallow-water marine

molluscs of the Azores. Gloria Maris 41(4-5): 84-

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Segers, W., Swinnen, F. & De Prins, R. 2009. Marine

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Vilvens, C. & Swinnen, F. 2007. Description of a new

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La liste des références, en ordre alphabétique, respectera la forme suivante (les titres des publications ne devraient pas être abrégés):

Keen, A.M. & Campbell, G.B. 1964. Ten new species of Typhinae (Gastropoda : Muricidae). TheVeliger 7(1): 46-57.

Powell, A.W.B. 1979. New Zealand Mollusca. Marine, land and freshwater shells. William Collins Publishers Ltd: xiv + 500 pp.

Mayr, E. 1989. Attaching names to objects, In: What the philosophy of biology is : essays for David Hull (M. Ruse, ed.), Klumer Academie, Dordrecht: 235-243.

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Keen, A.M. & Campbell, G.B. 1964. Ten new species of Typhinae (Gastropoda : Muricidae). TheVeliger 7(I): 46-57.

Powell, A.W.B. 1979. New Zealand Mollusca. Marine, land and freshwater shells. William Collins Publishers Ltd: xiv + 500 pp.

Mayr, E. 1989. Attaching names to objects. In: What the philosophy of biology is : essays for David Hull (M. Ruse, ed.), Klumer Academie, Dordrecht- 235-243

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VAPEX

Trimestriel de la Société Belge de Malacologie

association sans but lucratif

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VOL 12 (3-4)

2011

10 OCTOBRE

SOMMAIRE

Articles originaux - Original articles

E. F. Garcia

A new species of Mitra (Fusimitra ) (Gastropoda: Mitridae) 57

from the northwestem Gulf of Mexico

A new species, a lost type and its forgotten name and more 63

terebrid discoveries in the Caribbean (Gastropoda: Terebridae)

The Cantharus group (Gastropoda: Buccinidae) on Almirante 73

Leite Bank (Mozambique) with description of two new

species and one new genus

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Gastropoda: Terebridae) du Pléistocène de Niau, Tuamotu

(Polynésie Française)

New record of Typhinellus labiatus (Cristofori & Jan, 1832) 91

(Gastropoda: Muricidae) from Sào Tomé and Principe and

discussion about its classification and geographical

distribution

E. F. Garcia

Two new species of Epitonium (Gastropoda: Epitoniidae)

from the western Atlantic

D. Massemin, S. Clavier

& J.-P. Pointier

First record of Pisidium punctiferum (Guppy, 1867) and

Eupera viridans (Prime, 1865) (Mollusca: Sphaeriidae) from

French Guiana

109

B. M. Landau &

G. J. Vermeij

New Lyriinae (Mollusca: Volutidae) from the Lower Miocene

Cantaure Formation of Venezuela

119

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SOCIETE BELGE DE MALACOLOGIE

E. F. Garcia

Novapex 12(3-4): 57-62,10 octobre 2011

A new species of Mitra ( Fusimitra ) (Gastropoda: Mitridae) from the

northwestern Gulf of Mexico

Ernilio F. GARCIA

115 Oak Crest Dr.

Lafayette, LA 70503

Efg21 12@louisiana.edu

KEYWORDS. Mitridae, Mitra s.s., new species, Gulf of Mexico

ABSTRACT. A new species of Mitra (Fusimitra ) inhabiting the pinnacles off the Louisiana coast

is described and compared with Mitra (F.) antillensis Dali, 1889, its most similar congener.

INTRODUCTION

On the continental shelf off the Louisiana and

Texas coasts there is a sériés of banks or “pinnacles”

peculiar to the northwestern Gulf of Mexico. Their

geological history indicates that at one time, when the

sea- level was much lower, these were shallow

intertidal areas where coral grew. Nowadays these

banks, which arise abruptly from a soft, muddy

bottom to 20 to 90 m from the surface, hâve a summit

composed of calcareous rubble, a product of their

former coral fauna. These formations hâve been

discussed by Williams (1951), Stetson (1951),

Goedicke (1955) and others; however, their molluscan

fauna was not seriously addressed until Parker &

Curray (1956), who listed a large number of species

and noted their similarity to the shallow-water

Caribbean fauna ( Idem : 2428). The list of the rich

molluscan fauna of the pinnacles was later expanded

by a number of expéditions supported by grants,

mostly from the National Science Foundation, to the

Biology Department at the University of Louisiana at

Lafayette. They hâve been reported elsewhere (Garcia,

2000, 2002, 2007, 2008, 2010; Garcia & Lee, 2002,

2003, 2004).

The new species of Mitra (Fusimitra ) described

herein seems to be confmed to the top of the offshore

pinnacles discussed above. Although there hâve been

dredging cruises in the northeastern quadrant of the

Gulf of Mexico south to Dry Toitugas, and in the

southeastern quadrant at Bahia de Campeche, Mexico,

no specimens of the new species hâve been collected

in those areas. The 10 specimens that comprise the

type material were collected during seven expéditions

expanding a period of 11 years, from 2000 to 2010.

Only two specimens hâve been collected alive.

Ail cruises were done on the R/V Pélican, a

research vessel administered by the Louisiana

Universities Marine Consortium (LUMCON) using a

box dredge roughly 3 ft. by 3 ft. by 1 ft.

Abbreviations

BMSM: Bailey- Matthews Shell Muséum, Sanibel,

Florida, USA.

CMT: Charlotte M. Thorpe collection, Jacksonville

Beach, Florida, USA.

EFG: author’s collection

FF: Frank Frumar collection, Kirkwood, Missouri,

USA.

SBMNH: Santa Barbara Muséum of Natural History,

Santa Barbara, California, USA.

TCWC: The Texas Cooperative Wildiife Collection,

Texas A & M University, College Station, Texas,

USA.

USNM: United States National Muséum, Washington,

D.C., USA

SYSTEMATICS

Family MITRIDAE Swainson, 1829

Genus Mitra Lamarck, 1798

Type species: Voluta episcopalis Linnaeus, 1758 (= V.

mitra Linnaeus, 1758 [ICZN, 1969]) by subséquent

désignation of Montf'ort (1810: 543).

Mitra ulala n. sp.

Figs. 1-15

Type material. Holotype (Figs. 1-3) USNM 1155049,

28°05.95'N, 91°01.34'W, 69-68 m, length 30.5mm.

width 10.3 mm; paratype 1 (Fig. 4) USNM 1155050,

28°06.217N, 91°070'W, 75-65 m, length 21.9 mm,

width, 8.75 mm; paratype 2 USNM 1155051,

28 U 06.71'N, 91°02.50'W, 57-71 m, length, 22.o mm,

width 8.7 mm; paratype 3. TCWC 4-5247,

28°05.76'N, 91°01.15'W -, 64.7-62.1 m, length 25.6

mm, width 9.2 mm; paratype 4 (Fig. 7) USNM

1155052, 27°58.01'N 92° 35.67'W, 75-85 m, length

27.1 mm, width 8.4 mm; paratype 5 (Fig. 8) BMSM

17952, 28° 6.2 UN. 91° 2.23’W, 99.3 m, length 21.9

mm, width 8.0 mm; paratype 6 (Figs. 5-6) SBMNH

149759. 28°38.16'N, 89°33.19'W, 60-70 m, length

18.8 mm, width 7.6 mm; paratype 7 (Fig. 9) EFG

24308, 28°5.85’N, 91°1.28’W, 68.3 m, length 32.2

mm, width 10.8 mm; paratype 8 (Figs. 10-11) EFG

23177, 27° 59.141 'N; 91° 38.832’W, 91-65 m, length

23.2 mm, width 7.8 mm; paratype 9 (Figs. 12-15)

CMT, 28°05.552'N, 91°00.82'W, 63-64 m, length 23.7

mm, width 8.7 mm.

E. F. Garcia

A new species of Mitra

Type locality. Louisiana, off Isles Demieres,

28“05.95'N, 91°01,34'W, 69-68 m

Distribution. Offshore Louisiana pinnacles, 57- 99 m.

Description. Shell up to 32.2 mm in length, elongate-

ovate, average with/ length ratio 0.37. Protoconch

damaged, slightly tilted from axis (Fig. 14), of at least

3 smooth, conical, tan whorls. Teleoconch of 8

whorls; whorls very narrowly shouldered, straight-

sided. Suture channeled. Axial sculpture of numerous

narrow, strong cords on first 5 or 6 whorls; cords

narrower than interspaces, diminishing in strength on

later whorls, obsolète on last whorl. Spiral sculpture of

4 strong, wide cords on early whorls, 5 on later

whorls; interspaces pitted; cords becoming strongly

nodulous as they cross over axial éléments, creating a

strong clathrate pattern of nodes and pits on first 5

whorls (Figs. 3, 6, 11), diminishing in strength on later

whorls, obsolète on last whorl of mature specimens

except for two cords next to suture and at base of

whorl, where numerous weak, slightly nodulous spiral

cords still show. Apeiture approximately half the

length of shell, narrowly pointed posteriorly, with

relatively wide siphonal canal anteriorly; outer lip

simple, only slightly thickened; pariétal wall of adult

specimens with well-delineated, thin callus

posteriorly, thickened and slightly raised anteriorly,

producing 4 columellar folds; posterior fold largest,

prominent; next two folds proportionately smaller; last

fold almost obsolète. Shell rusty-brown in fresh

specimens, with white amorphous white blotches at

suture; blotches tending to form small to large axial

flammules that may cross entire whorl; a well-

delineated to nebulous white band appearing at mid-

section of last whorl; aperture in fresh specimens

mauve. Animal translucent-white, with opaque, white

to pale- yellow blotches; foot and siphon pale

yellowish-cream (Fig. 15).

Remarks. Cernohorsky (1976: 383) differentiates

Fusimitra Conrad, 1855 from other subgenera in

Mitridae by its early whorls showing a clathrate

sculpture similar to Concilia but adult whorls

becoming smoother, not unlike those of Mitra s.s. As

these characters apply to Mitra ulala , I hâve placed

this new species in Fusimitra.

The type sériés of Mitra (Fusimitra) ulala is

consistent in most characters. The two variables are

the basic color, which ranges from rusty-brown in live

collected specimens to orange and yellow in older

specimens, and the quantity and shape of the white

blotching. Only one, a live-collected specimen, had an

almost intact protoconch, which is tilted (Fig. 14); the

second live-collected specimen has a partial

protoconch that also seems to be tilted. However, at

this point one cannot be sure if this is a true character

of the species.

Mitra (Fusimitra) ulala n. sp. is very different

from most other western Atlantic Mitra s.s. such as

Mitra barbadensis (Gmelin, 1791), Mitra

damasomonteiroi Cossignani & Cossignani, 2007,

Mitra espinosai Sarasüa, 1978, Mitra lenhilli Petuch,

1988, Mitra leonardi Petuch, 1990, Mitra nodulosa

(Gmelin, 1791), Mitra pallida Usticke, 1959, and

Mitra semiferruginea Reeve, 1845. It can only be

confused with its congener, Mitra (F.) antillensis Dali,

1889.

Mitra antillensis (Figs. 16-20) is a widespread

species, ranging from North Carolina to Brazil. It is

very rare in the Gulf of Mexico, most of the

specimens having been found in the southeastern

quadrant of the Gulf. To the author’s knowledge, only

three specimens hâve been collected in the

northwestern Gulf of Mexico, one off Texas, at

Alaminos Station 72-A4 (Fig. 19) and two off

Louisiana: at Diaphus Bank, 28°5’N, 90“42’W, in 92

m (length 52,5 mm, width 15.6 mm); and at one of the

pinnacles, 27° 49 N 92° 53.5 W, 75-85 m (Figs. 17-

18). These three specimens are consistent with USNM

62103 (Fig. 16), the specimen designated by

Cernohorsky (1976: 387- 388; pi. 326, fïg. 4) as the

"selected holotype" (lectotype) for this taxon and

preclude the possibility that Mitra ulala is an

ecomorph of M. antillensis. Mitra antillensis differs

from the new species in the following characters:

1. It grows to a larger size. The lectotype has 8

teleoconch whorls and measures 81 mm. The holotype

as well as paratypes 7 and 8 of M. ulala also hâve 8

teleoconch whorls and measure respectively 30.5mm,

32.2 mm and 23.2 mm; moreover, ail other paratypes,

which measure 18.8 mm to 27.1 mm, hâve at least 7

whorls.

Figures 1-11. Mitra ulala n. sp.

1-3. Holotype USNM 1155049, 28°05.95’N, 91°01.34'W, 69-68 m, length 30.5mm. width 10.3 mm. 4. Paratype

1, USNM 1155050, 28°06.217N, 91°070'W, 75-65 m, length 21.9 mm, width, 8.75 mm. 5-6. Paratype 6 SBMNH

149759, 28°38.16TM89°33.19'W, 60-70 m, length 18.8 mm, width 7.6 mm 7. Paratype 4 USNM 1155052,

27°58.01'N 92° 35.67'W, 75-85 m, length 27.1 mm, width 8.4 mm. 8. Paratype 5, BMSM 17952, 28° 6.2l’N, 91°

2.23’W, 99.3 m, length 21.9 mm, width 8.0 mm. 9. Paratype 7, EFG 24308, 28°5.85’N, 91°1.28’W, 68.3 m,

length 32.2 mm, width 10.8 mm. 10-11. Paratype 8, EFG 23177, 27° 59.14l’N; 91°38.832'W, 91-65 m, length

23.2 mm, width 7.8 mm.

E. F. Garcia

Novapex 12(3-4): 57-62, 10 octobre 2011

E. F. Garcia

A new species of Mitra

2. It is proportionately narrower than the new species.

The lectotype, and four specimens from the Gulf of

Mexico (1 from Dry Tortugas, 2 from Louisiana, and

1 from Texas) hâve respectively a width/length ratio

(W/L) of 0.29, 0.30, 0.30 and 0.29. The type material

of M. ulala ranges from 0.34 to 0.40, with an average

of 0.37 W/L.

3. Its surface sculpture is different. Mitra antillensis

has a sculpture dominated by spiral éléments, which

become unusually strong below the suture, 3 or 4 in

earlier whorls, 6 or 7 on last whorl. In M. ulala the

spiral sculpture near the shoulder is of equal strength

on ail teleoconch whorls, and only two cords show by

the body whorl suture of two adult specimens.

Moreover, the First 5 whorls of M. ulala hâve strongly

developed axial and spiral éléments (Figs. 3, 6, and

11). Although some specimens of M antillensis from

Barbados are more heavily sculptured on early whorls

than those from more northern latitudes, they don’t

show the heavily clathrate pattern of the new species.

Finally, the number of spiral cords on Mitra

antillensis continue to increase with each whorl, the

lectotype showing about 10 on the penultimate whorl.

Mitra ulala has 4 on early whorls and no more than 5

on later whorls (Compare Figs. 3 and 18).

4. The coloration of Mitra antillensis is rather

subdued, from grayish-white to tan, sometimes with a

suffused white band by the suture; it also has an

olivaceous periostracum. Mitra ulala has a thin,

transparent periostracum and a pattern of coloration

not found in M. antillensis.

Etymology. Named for the University of Louisiana at

Lafayette, located in Acadia Parish, a French-speaking

area of Louisiana. Although its official acronym is

ULL, the jovial acronym “ULALA” has been used to

refer to this institution.

Acknowledgements

I am very much indebted to Drs. Darryl Felder and

Suzanne Fredericq, faculty members of the Biology

Department at the University of Louisiana at Lafayette

for inviting me to join them in their expéditions on the

R/V Pélican , on which most of the specimens used for

this article were collected. I am also very grateful to

Mrs. Charlotte Thorpe, of Jacksonville Beach, Florida,

tor allowing me to use her photograph of the animal of

Mitra ulala, as well as for lending me the specimen

for further photography. Dr. Harry G. Lee, of

Jacksonville, Florida, has reviewed this study and

considerably improved on its quality, I also would like

to thank the following individuals for providing

information and/or images of Mitra antillensis for this

study: Linda Ward and Mignonette Doley Johnson,

United States National Muséum; Dr. Fabio

Moretzsohn, Harte Research Institute for Gulf of

Mexico Studies, and Texas A&M University-Corpus

Christi, and Dr. Mary Wicksten, Texas A&M

University-College Station; and Mr. Frank Frumar,

Kirkwood, Missouri, and Mr. Steve Kern, Key West,

Florida. Most of the material for this study is based

upon work supported by the National Science

Foundation under Grant No. 0315995 and the RAP1D

grant.

REFERENCES

Cernohorsky, W.O., 1976. The Mitridae of the world

Part I. The subfamily Mitrinae. Indo-Pacific

Mollusca 5(17): 273-528. September 28.

Garcia, E. F. 2000. Surprising new molluscan records

from Louisiana and the northwestern Gulf of

Mexico. American Conchologist 28(3):5-6.

Garcia, E. F. 2002. More discoveries from a collecting

expédition off the Louisiana coast. American

Conchologist 30(l):6-8.

Garcia, E. F. 2007. A new species of Cosmioconcha

(Gastropoda: Columbellidae) from the northern

Gulf of Mexico. Novapex 8(2): 43-46.

Garcia, E. F. 2008. An extension of the genus

Spinosipella (Bivalvia: Verticordidae) in the Gulf

of Mexico. American Conchologist 36(3): 8-9.

Garcia, E. F. 2010. A géographie extension for two

species of Favartia (Muricidae: Muricopsinae)

from the western Atlantic. American Conchologist

38(3): 10-11.

Garcia, E. F. & Fl. G. Lee. 2002. Report on molluscan

species found in the offshore waters of Louisiana,

including many extensions of known range and un-

named species. American Conchologist 30(4): 10-

13.

Garcia, E. F. & H. G. Lee. 2003. Report on molluscan

species found in the offshore waters of Louisiana,

including many extensions of known range and un-

named species. II. American Conchologist

31(1 ):26-29.

Figures 12-20

12-15. Mitra ulala n. sp. Paratype 9 CMT, 28°05.552’N, 91°00.82'W, i63-64 m, length 23.7 mm, width 8.7 mm.

(Photograph of Fig. 15 by Charlotte Thorpe). 16- 20. Mitra antillensis Dali, 1889. 16. Lectotype, USNM 62103,

36 mi SE oi Cape Lokout, North Carolina, 336 m, length 81.0 mm, width 40.0 mm (Photo crédit: Mignonette

Doley Johnson). 17-18. EFG 22261, Louisiana, 27° 49 N 92° 53.5 W, 75-85 m, length (approx.) 48 mm, width

14.5 mm. 19. TCWC 4- 2600, Texas, Alaminos Station 72-A4, 28°34.7’N, 92° 05.5'W,39 m, length 59.9 mm,

width 17,1 mm. 20. FF, 16 mi. SW of Key West, Florida, 167 m (Photo crédit: Steve Kern).

E. F. Garcia

NOVAPEX 12(3-4): 57-62, 10 octobre 2011

E. F. Garcia

A new species of Mitra

Garcia, E. F. & H. G. Lee. 2004. Report on lhe

malacofauna of offshore Louisiana Waters -

including many range extensions and un-named

species. III .American Conchologist 32(3): 21-24.

Goedicke, T. R. 1955. Origin of the pinnacles of the

continental shelf and slope of the Gulf of Mexico.

Texas Journal of Science 7 (2): 149-159.

ICZN (International Commission for Zoological

Nomenclature). 1969. Opinion 885. Voluta mitra

Linnaeus, 1758 (Mollusca: Gastropoda) added to

the Official List. Bulletin of Zoological

Nomenclature 26: 125-127. 24 October.

MontfortD. de. 1810. Conchyliologie systématique, et

classification méthodique des coquilles; offrant

leurs figures, leur arrangement générique, leurs

descriptions caractéristiques, leurs noms; ainsi

que leur synonymie en plusieurs langues. Ouvrage

destiné à faciliter l'étude des coquilles, ainsi que

leur disposition dans les cabinets d'histoire

naturelle. Coquilles univalves, non cloisonnées.

Tome second. Schoell, Paris. Pp. [1-3], 1-676.

<http://www.biodiversitylibrary.Org/item/4 1566#p

age/553/mode/l up>

Parker, R. H. and J. R. Curray. 1956. Fauna and

bathymetry of banks on continental

shelf, northwest Gulf of Mexico. Bulletin of the

American Association of Petroleum

Geologists 40: 2428- 2439.

Stetson, Fl. C. 1953. The sédiments of the western

Gulf of Mexico. Part I - The

continental terrace of the western Gulf of Mexico.

Its surface sédiments, origin and

development, Papers in Physical Oceanography

and meteorology of Massachusetts Institute of

Technology and Woods Ploie Océanographie

Institute 12(4)1-45.

Williams, H. F. 1951. The Gulf of Mexico adjacent to

Texas. Texas Journal of Science 3(2): 237-250.

Y.Terryn

NOVAPEX 12(3-4): 63-72, 10 octobre 201

A new species, a lost type and its forgotten name and more terebrid

discoveries in the Caribbean (Gastropoda: Terebridae)

Yves TERRYN

Scientific Associate MNHN & RB1NS

Kapiteinstraat 27, 9000 Gent, Belgium

yves@naturalart.be

KEY WORDS. Terebridae, Caribbean, Western Atlantic, Guadeloupe, Terebra (s. 1.) lamyi sp.

nov., Terebra limatula, shell morphology, biodiversity.

ABSTRACT. A small collection of terebrid specimens from Guadeloupe is discussed, with the

description of a new species Terebra (s. I.) lamyi sp. nov. and the history of the taxon Terebra

limatula Dali, 1889 is extensively researched.

INTRODUCTION

While examining specimens of the new species

hereafter described, several terebrid experts were

asked for an opinion as to their identity.

One responded that the typical reticulated/beaded

sculpture is a feature typically seen in deeper water

species from the Caribbean and adjacent areas.

Although it bore almost no resemblance to any known

species, he advised the author to study also the

“forgotten species” Terebra limatula Dali, 1889a as

the new shell could well match its very short

description. As mentioned by Bratcher & Cemohorsky

(1987), the type was thought to be lost and in fact the

original description offers little discriminative help, so

this investigative lead was easily rejected. It wasn’t

until a quick search in the online type database of the

USNM was performed that specimens were found

logged as Terebra limatula. This contradicted earlier

report and prompted further investigation. Not only

was there more than one syntype but the shells were

actually présent in the type collection and fitted the

description and each were identical with one another.

The présent paper tries to clarify the situation, and its

effects.

Material and methods

Ail specimens from Guadeloupe were collected by Mr

Dominique Lamy and résidé in his private collection

unless otherwise mentioned. Ail pictures of these

specimens were taken by the collecter, with the

exception of the new species Terebra (s. 1.) lamyi sp.

nov. which were taken by the author.

Ail pictures of type material held in the USNM were

provided by Ellen Strong.

Ail pictures of type material held in the ANSP were

provided by Paul Callomon.

Abbreviations

ANSP: Academy of Natural Sciences, Philadelphia,

PA, USA

1NV: Instituto de Investigaciones Marinas, Santa

Maria, Colombia

MCZ: Muséum of Comparative Zoology, Harvard,

MA, USA

MNHN: Muséum national d'Histoire naturelle, Paris,

France

MZUSP: Museu Zoologia da Universidade de Sao

Paulo, Sao Paulo, Brazil

RBINS: Royal Belgian Institute of Natural Sciences,

Bmssels, Belgium

USNM: United States Nation Muséum - The

Smithsonian Institute, Washington, DC, USA

DL: Private collection Dominique Lamy, Guadeloupe

YT: Private collection of Yves Terryn, Belgium

Ancient history

W. H. Dali (1889a: 66) described the species

Terebra ( Acus ) limatula in the reports of the

Albatross-ex pedition to the Caribbean. The type

locality was not actually given but he enumerated a

list of localities as ‘habitat’: “ Barbados, 100fins; Gulf

of Mexico at Station 36, in 84 fins.; Bahamas, west of

North Bemini (sic!), in 200 fins (Dr. Rush); US Fish

Commission Station 2402, in the Gulf of Mexico,

between the delta of the Mississippi and Cedar Keys,

Fia., in 111 fms., mud; and Station 2610, 24 miles S.

E. from Cape Lookout on the Carolina coast, in 22

fms., sand, bottom température 79°.0F

Dali mentioned no total number of specimens he

had before him and only lists the size of a specimen as

being 18.0 mm long, 3.5 mm wide and with 14

whorls. The actual description of its discriminative

features is a bit scattered throughout the manuscript:

in a key and in the actual short description, and can be

summarized as follows: Shell small, acute, elongate,

columella not keeled, whorls fiat. Strongly cancellate

sculpture and nodulose at the interstices, about 18-20

costae on the body whorl. Subsutural band nodulose

and 2-4 spiral rows of nodules on the remainder of the

whorl. Color white to pale buff.

Dali furthermore mentioned the possible need to

distinguish the specimens coming from ‘more North’

Y. Terryn

A new terebrid species from the Caribbean

i.e. South Carolina, USA as var. acrior, defined as

having only 2-3 spiral rows that are not as strongly

sculptured.

After its description the taxon was rarely used

again, often in faunistie lists, only twice crudely

figured e.g. Dali, 1889b: 94; Dali & Simpson, 1901:

pl. 29, fig. N; de Jong & Kristensen, 1965: 47;

Warmke & Abbott, 1962: 133 (as var. acrior)-, Abbott,

1968: 164 (as a subspecies of Terebra protexta );

Kaicher, 1981: card 2710 (“holotype” of Terebra

acrior)-, de Jong & Coomans, 1988: 104. In ail these

cases the identity of the taxon could not be well

defrned by description, drawing or picture.

Recent history

The name Terebra limatula was mentioned also in

the Terebridae révision by Bratcher and Cemohorsky

(1987: 30), who listed the species as dubious because

the type material was thought to be iost and the

description was considered too general to be of any

help.

While the présent author was revising ail relevant

taxa for a possible comparison with Terebra (s. 1.)

lamyi sp. nov., the naine Terebra limatula remained

enigmatic because of its description and ‘lost types’.

A quick search by Dr Ellen Strong (Curator of

Mollusca, USNM, USA) revealed that possible ‘types’

are présent at the USNM. Upon a more in-depth look

through the type collection and records of the USNM,

she was able to trace back four (+1 possible) records

labeled as syntypes of Terebra limatula and var.

acrior.

USNM 103436: Unconfirmed type, not listed in the

online database because of its unconfirmed status.

United States Fish Commission, Station 2120, at 134

m, Grenada.

The specimen was catalogued Jan. 31, 1890, simply as

“Terebra ”, is conspecific with the hereafter-

mentioned specimens but is clearly does not belong to

the original type sériés as noted by Dali (see above).

USNM 92870: Catalogued February 13, 1888 with no

identification. 1 specimen in dry condition. Collected

19 October 1885, RV Albatross, US Fish Commission

Station 2610, North Carolina Cape Lookout at 40 m.

USNM 93971: Catalogued March 20, 1888 with no

identification. 1 specimen in dry condition. Collected

14 March 1885, RV Albatross, US Fish Commission

Station 2402 Florida, between Mississippi Delta and

Cedar Keys at 203 m, 34.33N - 76.2W.

USNM 61229: 2 specimens in dry condition;

collected by W. Rush, Bahamas, NW Bimini Island at

366 m. Additional notes accompanying this sample in

the USNM mention: “see also USNM 92870, 93971;

ANSP 33723 (syntypes)”.

The specimen was catalogued Dec. 19, 1885, and

labeled “ Terebra ”, Specimens are currently out on

loan.

USNM 87294: Terebra ( Acus) limatula var. acrior

Dali, 1889. 1 specimen in dry condition. RV Blake.

US Coast Survey Station 100, Barbados, 13.167N -

59.533W at 183 m. Leg. A. Agassiz. The specimen

was catalogued June 17, 1888 and labeled as “Acus

limatulus Dali” Specimen is currently out on loan.

The database notes field in the type record USNM

61229 mentions additional specimens or types in the

ANSP. An investigation by Paul Callomon (Curator of

Mollusca, ANSP, USA) revealed the following:

ANSP 33723: 3 ‘syntypes’ with the following

information on the labels:

Original label: L. protexta Cow / Bemini, Bahamas

[sic! = Bimini] / Dr. Wm. H. Rush!

Subséquent label 1 : Terebra limatula form acrior Dali

/ det. T. Bratcher / 17 Oct 1981.

Subséquent label 2: Probable syntypes of Terebra

limatula Dali, 1889 / A. R. Kabat. 17 Feb 1995.

One specimen (9.3 mm) seems conspecific with ail the

‘syntypes' of Terebra limatula in the USNM, but the

other two (6.0 mm & 12.2 mm) clearly belong to a

different species (and genus/group). Why these three

were kept together as one species and regarded as such

by Bratcher is a complété mystery as the différence is

quite évident. A bit of research revealed that the two

belong in fact to a probably undescribed species.

The total amount of ‘syntypes’ is thus actually six,

five for limatula and one for limatula var. acrior.

They are deposited in two muséums: five (of which

one is of limatula var. acrior) in USNM and one in

ANSP.

As quoted above, the var. acrior is merely based upon

small différences in the strength of sculpture and

number of spiral rows of beads. As it concems a

smaller specimen than the specimens of the

nominative type, we can conclude that this represents

a mere juvénile of the latter. Thus Terebra limatula

var. acrior is here interpreted as a junior synonym of

Terebra (s. 1.) limatula.

Figures 1-7

DI ' p S>e ^ ,a VV! * * * * S * 7 * P' nov ' MNHN 23204, Flolotype, 10.8 mm, Guadeloupe, off Saint François, 250 m; 2.

Saint p- Ven mm ’ Guadelou P e > offSaint François, 250 m; 3. YT, Paratype, 3 10.7 mm, Guadeloupe, off

Saint François, 250 m; 4. DL, Paratype, 2 9.1 mm, Guadeloupe, off Saint François, 250 m.

mm nlh >Ü SP VV S 5 ' ANSP 425025 > 12 - 2 mm ’ Bahamas, N Bimini Island, 366 m; 6. ANSP 425025, 5.9

mm, Bahamas, N Bimmi Island, 366 m.

7. Terebra limatula ANSP 33723, paralectotype, 9.3 mm, Bahamas, N Bimini Island 366

Y.Terryn

Novapex 12(3-4): 63-72, 10 octobre 2011

Y. Terryn

A new terebrid species from the Caribbean

As mentioned above, Bratcher & Cernohorsky ( 1987)

considered the type of Terebra limatula and var.

acrior to be lost. Subséquent researchers amateurs

studying terebrids from the Caribbean and adjacent

areas hâve often ignored the taxon.

Since then, however quite a nurnber of West

Atlantic/Caribbean ‘cancellate’ and/or ‘beaded’

terebrids hâve been described that resemble Terebra

limatula'.

Terebra (s. 1.) crassireticulata Simone, 1999 = nom.

nov. pro Terebra reticulata Simone & Verrisimo,

1995;

Terebra (s. 1.) colombiensis Simone & Gracia, 2006;

Terebra (s. 1.) simonei Lima, Tenorio & Barros, 2007;

And to a lesser extent, we can add the following to the

list:

Terebra (s. 1.) leptapsis Simone, 1999 = Terebra

doellojuradoi Carcelles, 1953 fide Faber, 2007,

synonymy agreed;

Terebra (s. 1.) intumescyra Lima, Tenorio & Barros,

2007;

Terebra (s. 1.) alagoensis Lima, Tenorio & Barros,

2007.

The discovery of the types of Terebra (s. I.) limatula

might affect the validity of the species here

mentioned, and this is discussed below.

Further discoveries

The fishing and collecting operations around the

island of Guadeloupe by Mr Dominique Lamy hâve

yielded over the last décades a large nurnber of rare

discoveries in malacology thanks to his personal

investment of time and vigorous effort and because of

his particular fishing methods (dredgings, baited bottle

traps etc... ). Besides the above-mentioned newly

described species newly, a nurnber of noteworthy

species/specimens are here briefly commented on and

figured. Ail the taxa will need further investigation to

better understand the intraspecific variability of these

hard to obtain deeper water species, but this must

await the arrivai of further specimens. The présent

énumération serves merely as a photographie

iconography to illustrate the difficulty of research, the

huge variety and biodiversity in the Terebridae of the

Caribbean, and our Iack of knowledge of the fauna.

No attempt has been made at this stage to assign taxa

or even give detailed descriptions as they would be

based on incomplète, dead collected or too few

specimens.

SYSTEMATICS

Family TEREBRIDAE Môrch, 1852

Ail species here discussed and described will be

placed in the informai grouping Terebra (s. 1.) (thus

sensu Bratcher & Cernohorsky, 1987) as opposed to

Terebra s. s. ( sensu Terryn, 2007), unless otherwise

mentioned. The actual generic status of the species is

the subject of continuous research and will be updated

as information cornes available.

Terebra lamyi sp. nov.

Figs 1-4

Type material. Holotype MNHN 23204, dredged off

Saint François, Guadeloupe at 250 m, 10.8 mm.

Paratypes. Ail dredged off Saint François, Guadeloupe

at 250 m - Paratype 1: DL, 10.6 mm; Paratype 2: DL,

9.1 mm; Paratype 3: YT, 10.7 mm.

Type Locality. Off Saint François, Guadeloupe.

Dredged on muddy bottom at 250 m.

Additional material. DL, 1 specimen dredged at 150

m off Port Louis.

Distribution. Known only from the type locality and

off Port Louis.

Habitat. Ail specimens were dredged on a muddy

substrate. Presumed bathymetrical range for living

specimens between 150 and 250 m.

Figures 8-12

8. Terebra colombiensis Simone & Gracia, 2006. INVMOL 1963, paratype, 15.5 mm, off Bocas de Ceniza,

Lolombia, 31_-326 m. (Courtesy of Luiz Ricardo L. Simone, MZUSP) (x 0.5).

’ , t ' c h la ^' ,hl,ll l a Da\\, 1889. USNM 93971, lectotype, 17.9 mm, USA, Florida, between Mississippi Delta

and Cedar Keys, 203 m. (Courtesy of Ellen Strong, USNM).

pü" I e 'n ’ la -^' mone ^ Verissimo, 1995. MZUSP 27930, holotype, 22.2 mm, slope off Ubatuba, Sao

Pau o, Brazil, 320 m. (Picture courtesy of Luiz Ricardo L. Simone, MZUSP ).

Nnat r^V^ Hmatula Dali, 1889. 11. USNM 92870, paralectotype, 14.7 mm (reconstructed estimate), USA,

r , al ,° ! ' i na ’ Cape Lookout ’ 40 m. (Photo courtesy of Ellen Strong, USNM). 12. USNM 103436, 5 9 mm,

Grenada, 134 m. (Photo courtesy of Ellen Strong, USNM).

Y. Terryn

NOVAPEX 12(3-4): 63-72, 10 octobre 2011

Y. Terryn

A new terebrid species from the Caribbean

Description. Shell srnall to 11 mm. Shell color

maroon to fawn; subsutural band often giving a

somewhat lighter appearance. Outline of whorls

slightly concave to alrnost straight. Protoconch of

about 2.0 whorls; shiny brown. Subsutural band

decorated with numerous axial ribs, as numerous as on

the remainder of the whorl. Subsutural band delimited

by a deeper groove compared to those on the

remainder of the whorl. Axial sculpture of numerous

slightly arcuate to rnostly straight ribs, wider than the

interspace. Spiral sculpture of 5-6 grooves, somewhat

cutting the axial ribs; ail giving the shell a somewhat

elongate-beaded appearance. Columella curved with a

shiny brown callous. Animal and operculum

unknown.

Etymology. The species is named in honor of the

Guadeloupe conchologist Mr Dominique Lamy in

gratitude for providing the conchological community

with molluscan material from Guadeloupe, including

the here-described species.

Terebra species 1

Figs 5-6

Material: ANSP 425025, previously included in

ANSP 33723, 2 sps.

Locality data: Bimini, Bahamas.

Commentary: Protoconch with about 2 whorls,

mamillate. Subsutural band and remainder of whorl of

identical sculpture consisting of numerous low axial

ribs. Radial sculpture consisting of numerous shallow

grooves. Subsutural marcation consisting of a shallow

groove, yet deeper incised than the radial scupture: 1-2

radial grooves on the subsutural band there, about 5-6

on the remainder of the whorl (measured on the

largest specimens penultimate whorl).

Discussion and comparison: Both specimens are

small, heavily damaged and badly eroded, probably

representing dead juvéniles or subadult forms. Clear

description of the features therefore is not possible.

However it is very évident that these two specimens

are not conspecific with the ‘ limatula ’ contained in the

original type lot ANSP33723. Both the différence in

protoconch and sculpture is very évident. Hence the

two specimens should be separated out of the type

sample, leaving only 1 paralectotype of Terebra (s. 1.)

limatula in ANSP33723 (see further).

Terebra limatula Dali, 1889

Figs 7, 9-12

Terebra (Aeus) limatula Dali, 1889a: 66.

Terebra ( Acus ) limatula var. acrior Dali, 1889a: 66.

Terebra crassireticulata Simone, 1999 (= nom. nov.

pro Terebra reticulata Simone & Verissimo, 1995):

222 .

Type material

Terebra limatula: USNM 92870: 1 specimen (dry).

Collected 19 October 1885, RV Albatross, US Fish

Commission Station 2610, North Carolina Cape

Lookout at 40 m; USNM 93971: 1 specimen (dry).

Collected 14 March 1885, RV Albatross, US Fish

Commission Station 2402 Florida, between

Mississippi Delta and Cedar Keys at 203 m, 34.33N -

76.2W; USNM 61229: 2 specimens (dry); collected by

W. Rush, Bahamas, N Bimini Island at 366 m; ANSP

33723: 1 specimen (dry); collected by W. Rush,

Bahamas, N Bimini Island at 366 m, 9.3 mm (see

above).

The specimen registered as USNM 93971 (Fig. 9) is

selected as the lectotype, in order to enhance the

stability of the nomenclature in accordance with ICZN

art. 74.7.3. Ail other type material specimens of

Terebra limatula (USNM 92870, USNM 61229) and

ANSP (33723) are paralectotypes.

Terebra limatula var. acrior: USNM 87294: Terebra

{Acus) limatula var. acrior Dali, 1889. 1 specimen in

dry condition. RV Blake. US Coast Survey, Barbados,

13.167N - 59.533W at 183 m.

Terebra reticulata: MZUSP 27930: 1 specimen in dry

condition. Station 5361, 24°42’0”S-44 o 30’5”W, slope

off Ubatuba, Sao Paulo, Brazil, 320 m. Numerous

paratypes at MZUSP.

Terebra crassireticulata : Although Terebra

crassireticulata was a re-naming of Terebra

reticulata, a heading was made in the manuscript

entitled ‘type material’ which lists a large number of

specimens from the MNHN but without formally

designating types. Assumed is that this was a mere

éditorial lapsus.

Figures 13-23 (Ail from coll. Dominique Lamy and from Guadeloupe).

lt]f: Tere À bn ‘ evel y nae Clench & Aguayo, 1939. 13. 51.1 mm, off Saint-François, 250 m. 14. 69.7 mm, off

ord Grande-Terre, 300 m. 15. 62.5 mm, off Nord Grande-terre, 300 m.

16. Terebra glossema Schwengel, 1940. 23.3 mm, off Port-Louis, 150 m.

17. Terebra species 2. 29.8 mm, off Port-Louis, 100 m.

ll'l 9 ' T ' erebra species 3. 18. 9.2 mm, off Saint-François, 250 m. 19. 7.4 mm, off Anse La Gourde, 250 m.

q ' eK çç™ species 4. 20. 6.2 mm, off Anse La Gourde, 250 m. 21. 9.1 mm, off Anse La Goude, 250 m. 22.

9.5 mm, off Vieux Habitants, 300 m.

23. Terebra species 5. 14.7 mm, off Saint-François, 250 m.

Y. Terryn

NOVAPEX 12(3-4): 63-72, 10 octobre 2011

Y.Terryn

A new terebrid species from the Caribbean

Additional matcrial studied. USNM 103436 (see

remarks above).

Type locality. Lectotype: US Fish Commission

Station 2402, Florida, between Mississippi Delta and

Cedar Keys at 203 m, 34.33N - 76.2W

Description. See Simone & Verissimo, 1995; Simone,

1999 for an extensive description of both the shell

morphology and anatomy. The holotype of Terebra

crassireticulata shows exactly the sanie protoconch

and shell morphology as Terebra limatula. The

specimens of Terebra limatula studied were ail shells

with damaged body whorl (columellar plicae visible),

compared to the fully adult and complété holotype of

Terebra crassireticulata with thickened columellar

callus.

Habitat. Unknown.

Distribution. From North Carolina, Florida and the

Bahamas (Dali, 1889a), probably South along the

chain of islands bordering the Atlantic to Brazil,

where it has been confirmed from Rio de Janeiro to

Sao Paulo (Simone, 1999).

Terebra evelynae Clench & Aguayo, 1939

Figs 13-15

The species is easily identifiable among the West

Atlantic and Caribbean terebrids but has been rarely

recorded. Only a handful of dead collected specimens

are known besides the live collected holotype (MCZ

135077: 118.3 mm). As far as the author could

détermine, the species had only been recorded from

off Cuba (type locality: off N Santa Clara Province,

225 fms) till the présent discovery of one live and

several dead (crabbed) collected specimens off

Guadeloupe.

Terebra glossema Schwengel, 1940

Fig. 16

The species is fairly identifiable amongst the West

Atlantic and Caribbean terebrids but has been rarely

recorded. The range of the species is restricted to the

northern Caribbean (Florida, Bahamas and Cuba) but

Faber (2007: Aruba) and the présent finding

(Guadeloupe) confirm a much wider range southwards

and it probably lives throughout the eastem rim of the

Caribbean islands.

Terebra species 2

Fig. 17

The species bears some resemblance to T. glossema in

the characteristics of the sculpture, but the density of

the spiral and axial sculpture is rather different. The

sculpture is much coarser and the general outline is

much more slender than in T. glossema. At présent

this species is only represented by a single specimen.

Thus nothing is known concerning intraspecific

variability. The species can not be compared to any

other known taxon in the Caribbean and is most

probably new to science. Due to the lack of additional

specimens at the moment, the species remains under

study awaiting more finds.

Terebra species 3

Figs 18-19

The following three species can obviously be

accredited as belonging to the limatula- group because

of the shell surface morphology, which is reticulated

and beaded; but because of constant différences in

protoconch and variations in the sculpture, they must

be assigned under different taxa.

The spiral sculpture of this species is sharp: deep

grooves separated by almost protruding beads, yet

close-set and dense.

Terebra species 4

Figs 20-22

The présent species was also dredged at the type

locality of Terebra lamyi. It was preliminary stored as

T. cf. colombiensis pending further research but the

présent paper proves that it is does not belong to that

species nor with any of the related species. The main

différence is the conical, paucispiral protoconch of

about 2.5-3.0 whorls, T. limatula and T. colombiensis

both hâve a near-mammillate protoconch. Furthermore

the présent species is quite distinguishable because the

fine beading and deep and wide spiral grooves gave

rise to a somewhat step-like or turreted outline of the

whorls. Because of the close relationship in sculpture,

this species is without doubt related to the

aforementioned. It is here figured for the first time

awaiting more specimens for further study.

Figures 24-33 (Ail from coll. Dominique Lamy and from Guadeloupe).

24. Terebra species 6. 14.5 mm, off Port-Louis, 130 m.

25-27. Terebra species 7. 25. 16.3 mm, off Port-louis, 130 m. 26. 14.3 mm, off Port-Louis, 100 m. 27. 14.8 mm,

off Port-Louis, 150 m.

28. Terebra species 8. 1 1.5 mm, off Port-Louis, 160 m.

29-33. Terebra species 9. 29. 9.8 mm, off Port-Louis, 100 m. 30. 12.5 mm, off Port-louis 100 m. 31.9.2 mm, off

Port-Louis, 120 m. 32. 6.1 mm, off Port-Louis, 100 m. (x 1.5). 33. 9.1 mm, off Port-Louis, 120 m.

Y.Terryn

Novapex 12(3-4): 63-72,10 octobre 2011

Y. Terryn

A new terebrid species from the Caribbean

Terebra species 5

Fig. 23

The species shows a strong resemblance to T.

limcitula but again the protoconch and the even coarser

general sculpture leaves no doubt that it represents a

different and perhaps undescribed taxon.

Terebra species 6

Fig. 24

The species has no comparison in the Caribbean but is

somewhat similar in sculpture to a species occuring in

the Indo-Pacific (coll. MNHN, Marquesas Islands)

currently under study and description. The species is

only known from one dead collected specimen which

would make description prématuré at this stage.

Terebra species 7, 8 & 9

Figs 25-33

The mentioned 3 morpho-species are without doubt

related and close in resemblance to T. alba.

Although T. alba is reported throughout the

Caribbean, findings are usually restricted to the North

(Gulf of Mexico, Bahamas, Florida etc... ).

Species 6 & 8 are closely related to one another and to

T. alba because of similar sculpture, yet differ

amongst each other in protoconch size and shape.

Species 7 has a coarser sculpture and a different

protoconch than the aforementioned.

Acknowledgements

I wish to express my gratitude to Mr Dominique

Lamy for bringing a large number of terebrids from

Guadeloupe and especially the species here named

after him, to my attention. Also a great word of

appréciation for ail their efforts goes to Dr Ellen

Strong (USNM) and Paul Callomon (ANSP).

Furthermore, I wish to thank Dr Philippe Bouchet and

Virginie Héros (MNHN) for their ongoing assistance

in the study of Tercbridae. 1 sincerely appreciate the

assistance I had from Luiz Ricardo L. Simone

(MZUSP), in providing pictures of types and valuable

comments in the matter. 1 also sincerely thank Willem

Faber (The Netherlands) and Koen Fraussen

(Belgium) for critically appraising the text and Gavin

Malcolm (UK) for assistance and comments during

the research.

REFERENCES

Abbott, R. T. 1968. Seashells of North America.

Golden Press, New York, USA. 280 pp.

Bratcher, T. & W. O. Cernohorsky 1987. Living

Terebras of the World. Madison Publishing

Associates, New York, NY, USA. 240 pp.

Dali, W. H. 1889a. Report on the Results of

Dredgings, under the supervision of A. Agassiz, in

the Gulf of Mexico (1877-78) and in the Caribbean

Sea (1879-80), by the U. S. Coast Survey Steamer

“Blake 29. Report on the Mollusca. 2.

Gastropoda and Scaphopoda. Bulletin of the

Muséum of Comparative Zoology, Harvard , 18: 1 -

492.

Dali, W. H. 1889b. A preliminary catalogue of the

shell-bearing marine mollusks and brachiopods of

the southeastem coast of the United States, with

illustrations of many of the species. Bulletin of the

United States National Muséum, 37: 94.

Dali, W. H. & Simpson, C. T. 1901. The Mollusca of

Porto Rico. Bulletin of the United States Fish

Commission, 20: 351-524, pis 53-58.

de Jong, K. M. & Coomans, H. E. 1988. Marine

gastropods from Curaçao, Aruba and Bonaire. E.

J. Brill, Leiden, The Netherlands. 261 pp.

de Jong, K. M. & Kristensen, I. 1965. Gegevens over

Mariene Gastropoden van Curaçao.

Correspondentie Blad van de Nederlandse

Malacologische Vereniging, suppl.: 1-56.

Faber, M. J. 2007. Marine gastropods from the ABC-

islands and other localities. 14. The family

Terebridae (Gastropoda, Neogastropoda) with

description of a new species from Aruba.

Miscellanea MaUacologica , 2 (3): 49-55.

Kaicher, S. D. 1981. Card Catalogue of World- Wide

Shells, 27: Terebridae II, no. 2710.

Simone, L. R. L. & Verrissimo, P. 1995. Terebra

reticulata, new species of Terebridae (Gastropoda,

Prosobranchia, Conoidae) from Southeastem

Brazil. Bulletin of Marine Sciences, 57 (2): 460-

466.

Simone, L. R. L. 1999. Comparative morphology and

systematics of Brazilian Terebridae (Mollusca,

Gastropoda, Conoidae), with description of three

new species. Zoosytema, 21 (2): 199-248.

Terryn, Y. 2007. Terebridae, a Collectors Guide.

Conchbooks, Hackenheim, Germany &

NaturalArt, Gent, Belgium. 57 pp. + 65 colour pis.

Wannke, G. L. & Abbott, R. T. 1962. Caribbean

Seashells. Livingstone Publ., Wynnewood,

Pensylvania, USA. i-x, 348 pp.

K. Fraussen & J. Rosado

NOVAPEX 12(3-4): 73-79, 10 octobre 2011

The Cantharus group (Gastropoda: Buccinidae)

on Almirante Leite Bank (Mozambique)

with description of two new species and one new genus

Koen FRAUSSEN

Leuvensestraat 25, B-3200 Aarschot, Belgium

koen.fraussen@skynet.be

José ROSADO

Av. Friedrich Engels, n. 373, le,

Maputo, Moçambique

joserosadoi@hotmail.com

ABSTRACT. A new genus and two new species of deep water Buccinidae collected during

MAINBAZA are described: Pollia imprimelata sp. nov. and Micrologus mochatinctus gen. & sp.

nov., both front Almirante Leite Bank. Pollia sowerbyana (Melvill & Standen, 1903) is recorded

from Madagascar and the variability of this species is discussed.

KEYWORDS. Mollusca, Gastropoda, Buccinidae, Pisaniinae, Pollia, Mozambique,

MAINBAZA, new taxa.

INTRODUCTION. Ignored in the past, the

Mozambique fauna has become the subject of

intensive collecting and studies during the latest

décades. It is obvious that Mozambique is blessed

with a unique fauna, with components from the South

African fauna as well as from the East African fauna,

due to its geographical situation. Ridden up fragments

from the deep water faunas are présent, from the Natal

Bassin (in the south) and the Somalian Bassin (in the

north). In addition, a large number of species which

are most probably endemic for the regio are known.

The second author participated in the recent

MAINBAZA cruise in the Mozambique Channel. Part

of the Admirante Leite Bank was investigated during

this expédition, an area with strong currents resulting

in difficult dredging and trawling conditions. Among

rnany animais two particular species belonging to the

Cantharus group attracted attention. One is

recognized as belonging to an undescribed species of

the genus Pollia Gray in Sowerby, 1834 and named

Pollia imprimelata sp. nov. For the second species,

which is distinct in spiral sculpture and columellar

characteristics from ail known Cantharus group

species, we had to décidé to describe a new genus

because it seems out of place in any of the other

known généra. This taxon is added to the East African

fauna under the name Micrologus mochatinctus gen.

& sp. nov.

Material is, unless otherwise stated, deposited in

MNHN. The material is, unless being a type (which

are allocated to catalogue numbers), unambiguously

designated and retrievable by the combination of

expédition acronym and station number.

ABBREVIATIONS

KF: collection Koen Fraussen, Belgium.

MHNM: Museu de Historia Natural de Maputo,

Mozambique.

MNHN: Muséum national d'Histoire naturelle, Paris,

France.

JR: collection José Rosado, Portugal,

dd: empty shell, dead collected.

juv: juvénile or subadult shell.

DW: (drague Warén) Warén dredge.

SYSTEMATICS

BUCCINIDAE Rafinesque, 1815

Subfamily Pisaniinae Gray, 1857

Genus Pollia Gray in Sowerby, 1834

Pollia, Gray in Sowerby, 1834: pl. 237, fig. 12. Type

species by monotypy: “ Triton undosus Lamk.” =

Buccinum undosus Linnaeus, 1758 from the Indo-

West Pacific, Recent.

For a discussion on the taxonomie position of Pollia

we refer to Vermeij & Bouchet (1998: 472-473) and

Vermeij, (2006: 85-86, 89-91).

Pollia imprimelata sp. nov.

Figs 7-11

Type material. Holotype, 19.9 mm, Mozambique

Channel, Almirante Leite Bank, MAINBAZA stn.

DW3167, 26°12’S, 35°02’E, 228-230 m deep,

MNHN-23770.

K. Fraussen & J. Rosado

The Cantharus group on Almirante Leite bank

Paratypes 1-7, 19.1-21.6 mm, saine locality, MNHN-

23771, MFTNM, JR, KF-6129.

Paratype 8, 12.6 mm, juvénile (protoconch), saine

locality, MNHN-23771.

Type locality. Mozambique Channel, Almirante Leite

Bank, MA1NBAZA stn. DW3167, 26°12’S, 35°02’E,

228-230 m.

Material examined. Mozambique: Mozambique

Channel, Almirante Leite Bank, MAINBAZA stn.

DW3167, 26°12’S, 35°02’E, 228-230 m, 28 dd (9 jv.).

- Stn DW3169, 26°11 ’S, 35°01’E, 450 m, 4 dd.

Range and habitat. Only known from the Almirante

Leite Bank. Bathymétrie range, ail empty shells,

between 228 and 450 m.

Syntopic with Micrologie mochatinctus sp. nov. on

Almirante Leite Bank (MAINBAZA stn. DW3167

and DW3169).

Almirante Leite Bank is an area with strong currents

overflowing a bottom of hard rocks. The habitat is

characterized by the presence of rock, coral, sponges

and gorgonians (Text Fig. 1).

Text Figure 1. Habitat of Pollia imprimelata sp. nov.

Description. Shell of medium size for genus (up to

21.6 mm in height). Shape semi-oval, spire rather

conical. Teleoconch whorls 5 Zi in number, laterally

flattened, suprasuturally with narrow. but clearly

constricted area. Colour white to yellowish brown

with brown spiral lines on top of axial sculpture.

Protoconch consisting of slightly more than 1 3 / 4

whorl, 2.2 mm high, 1.1 mm in diameter. Colour

yellowish with brown spot on tip. Whorls rapidly

increasing in size, last whorl big, convex. Surface

smooth, rather glossy, but covered with numerous

minute shallow holes. Transition to teleoconch

marked by 3 or 4 fine incrémental lines.

First whorl with 3 well developed primary spiral

cords; a fine, fourth cord présent in subsutural

interspace. A single, fine secondary spiral cord

appears along second whorl, situated in the iniddle of

each interspace. Subsutural spiral cord increasing in

stiength, on third whorl almost as strong as other

primary spiral cords; a fifth primary spiral cord partly

concealed under suture with subséquent whorl. One

subsutural secondary spiral cord becoming as strong

as primary spiral cords along penultimate whorl,

resulting in the presence of 2 spiral cords on

subsutural slope which render a bilirate appearance

and torm a deep suture above axial ribs. Penultimate

K. Fraussen & J. Rosado

NOVAPEX 12(3-4): 73-79, 10 octobre 2011

whorl with 5 primary spiral cords, two subsutural ones

slightly weaker, separated by a narrower interspace; a

sixth primary spiral cord partly concealed under suture

of subséquent whorl. Body whorl with about 40 spiral

cords of different strength, 20 of them primary and

secondary spiral cords of about sanie strength,

altemating with about 20 tlner tertiary spiral threads.

First teleoconch whorl with 13 fine axial ribs;

abapically, on the suprasutural adpression, becoming

weaker, resulting in a suture with constricted

appearance; interspaces of equal size or slightly

broader. Subséquent whorls with 12 axial ribs. Ribs

becoming gradually weaker, but slightly broader.

Body whorl with 8 axial ribs, about 3 additional ribs,

or traces of them, on prelabral varix. Last 1/4 part of

body whorl covered with a broad, but weak prelabral

varix. Whole surface covered with minute, sharp

incrémental lamellae.

Aperture oval, adapical part pinched. Columella

strongly concave; callus thin, white, rather adhèrent to

preceding whorl, covered with big lirae according to

sculpture of preceding whorl, with a single adapical

columellar lira and 3 strong, abapical columellar folds

near transition to siphonal canal. Outer lip thin;

omamented with about 16 internai lirae of different

strength, not according to sculpture of outer surface.

Siphonal canal short, broad, open. Aperture and

siphonal canal together slightly more than 1 14 of total

shell length.

Animal and operculum unknown.

Comparision. Pollia imprimelata sp. nov. is

characterized by the laterally slightly flattened whorls

which give the spire a more conical appearance and,

as a conséquence, a more pinched adapical part of the

aperture. The maximum diameter of each whorl is

situated below the periphery. This shape is rather

atypical for the genus.

The colour ranges front snow-white and off-white

with sonie pale brownish bands to the presence of

reddish brown spiral Unes on top of the spiral

sculpture, especially on the periphery and subsutural

slope. Snow-white shells and coloured ones were

found in the sanie haul (MAINBAZA stn. DW3167

and DW3169).

AU known Pollia species differ by having a

convex subsutural area and the slightly more rounded

adapical part of the outer lip. (See below.)

Pollia sowerbyana (Melvill & Standen, 1903)

(type locality: “Gulf of Oman and Mekran Coast,

especially between Gwadûr and Jask, frorn 25-30

fathoms”) (Fig. 13) is characterized by the convex,

slightly shouldered whorls, the big axial ribs, the

round rather than oval aperture with a weak adapical

tabulation. The suture is distinct, suprasuturally

slightly appressed. Specimens with a broad shape, or

short spire, niay hâve the appressed band partly

concealed under the next whorl.

Pollia sowerbyana is similar in spiral sculpture,

but differs by the convex teleoconch whorls which

reach their maximum diameter near the shoulder

(instead of below the suturai line), consequently the

adapical part of the outer lip is broader and more

tabulate where connected with the preceding whorl;

the prelabral varix is stronger but narrower; the

suprasutural constricted area is usually broader; the

subsutural spiral cords are straight (rather than waved

or twisted where Crossing the axial ribs) and finer with

a broader interspace; the number of axial ribs is

smaller on the upper teleoconch whorls (about 10) but

higher on the body whorl (10 including the prelabral

varix); the columellar callus is broader, especially the

adapical part which has a gently curved edge; the

pattern usually consists of 2 broad bands (instead of

fine fines) and it seerns that the hairs on the

periostracum are longer.

It may be possible that the more eastern species

(Philippines: P. vicdani’, Coral Sea and New

Caledonia: P. pellita) hâve to be regarded as

conspecific with P. sowerbyana. Further study may

throw a light on this question but in the meantime we

regard these taxa as distinct (see below).

Pollia vicdani (Kosuge, 1984) (type locality:

Philippines, Bohol, off Panglao, 120 fms.) is similar,

but differs front P. imprimelata sp. nov. by the bigger

number of axial ribs on the body whorl (the number of

axial ribs on the upper spire whorls is the same) and

the slightly more convex subsutural area and the

slightly more rounded adapical part of the outer lip.

The number of axial ribs on penultimate and body

whorls seems to vary considerably in many Pollia

species, but it is quite constant in Philippine

populations of P. vicdani. We doubt this is a solid

characteristic to distinguish these species or fonns.

Pollia pellita Vermeij & Bouchet, 1998 (type

locality: New Caledonia, Loyalty Ridge, 20°42’S,

167°00’E, 270 ni) is similar to P. imprimelata sp. nov.

in shape and pattern, especially the fonn front Coral

Sea, but differs by the slightly more convex subsutural

area and the slightly more rounded adapical part of the

outer lip.

Pollia delicata (E. A. Smith, 1899) (type locality:

Investigator “Station 237, lat. 13° 17’ N., long. 93° 7’

E., off Andaman Islands, 90 fath.”) is similar, but

differs front P. imprimelata sp. nov. by the smaller

number of protoconch whorls (about 1), the clearly

more convex whorls, the spiral sculpture which is fine

and running straight near the subsutural slope (instead

of being twisted when Crossing the axial ribs).

Etymology. Pollia imprimelata sp. nov. is derived

from imprimere (Latin, verb) meaning "press upon" in

combination with latus (Latin) meaning "side", which

refers to the slightly flattened latéral sides. The last

letters of imprimere are contracted to avoid deviating

pronunciation.

K. FRAUSSEN & J. ROSADO

The Cantharus group on Almirante Leite bank

Pallia sp.

Fig. 12

Material. Mozambique Channel, Almirante Leite

Bank, MAINBAZA stn. DW3168, 26°12’S, 35°03’E,

87-90 m, 1 dd.

Remarks. The single dead collected specimen has a

slightly damaged columellar part of the siphonal canal

and is characterized by the protoconch which consist

of a single whorl with a rather broad, but slightly

flattened tip.

The shell is similar to Pollia imprimelata sp. nov.

in spiral sculpture, but differs by the protoconch

which consist of a single whorl (instead of 2 whorls)

with a flattened tip, the more convex whorls resulting

in a more convex adapical part of the aperture and a

shape typical of the genus, the adapical part of the

axial ribs which is weaker and the smaller adult size.

Pollia subcoslata (Krauss, 1848) (type locality “In

litore natalensi”: South Africa, Natal, littoral), a

species which also lives along the coast of

Madagascar and Mozambique in shallow to fairly

deep water, is easily distinguishable by the bigger

spiral cords and by the body whorl with weak or

absent axial sculpture. P. subcoslata also has a thicker

shell, a darker colour and a thick periostracum, but

these three features may vary in the genus, according

to bathymétrie occurrence.

Micrologus gen. nov.

Type species. Micrologus mochatinctus sp. nov. (type

locality: Mozambique Channel, Almirante Leite Bank,

MAINBAZA stn. DW3169, 26 0 U’S, 35 o 01’E, 450 m

deep).

Diagnosis. Shell rather small. Shape semi-oval; spire

moderately high, fusiform; base slightly stretched.

Protoconch paucispiral, consisting of about 1 smooth

whorl; surface rather rough, covered with minute

shallow holes. Transition to teleoconch indistinct,

marked by the start of the sculpture of the teleoconch.

Spiral sculpture fine, consisting of thin primary spiral

cords (4 on first whorl, 5 on other spire whorls, 11 on

body whorl). Spiral interspaces broad on periphery,

with 1 (on upper spire whorls) to 3 (on fourth whorl to

body whorl) fine secondary spiral threads of equal

strength, occasionally 4.

Axial sculpture consisting of broad ribs with broad

interspaces, running from suture to suture on spire

whorls, from suture to just below midwhorl on body

whorl.

Aperture large, ovate, adapically slightly pinched with

a single columellar knob and a labral knob bordering

the anal notch. Columella strongly concave, smooth

and glossy, with an adapical columellar knob

(bordering the anal notch) and a strong abapical

columellar knob on transition to siphonal canal. Callus

thin, smooth, glossy, broad; abapical part more

developed, forming a broad layer well adhèrent to

columella. Outer lip rounded, lip thick, edge sharp,

with broad, but low internai knobs, not situated

according to primary spiral cords on outer side;

adapical knob slightly bigger, occasionally split;

abapical knob slightly bigger, on transition to siphonal

canal. This abapical knob not situated in front of

abapical columellar knob but strongly diagonally

orientated to each other. Labral varix low but broad,

slightly prosocline. Siphonal canal short, broad, open.

Comparison. The diagnosis is based on the single

known species, described below. Therefore we do not

know the magnitude of variation within the genus.

Based on known variability in other généra of the

Cantharus group we may assume with some certainty

that the way the spiral cords are arranged is a solid

diagnostic feature, as well as the convex shape of the

teleoconch whorls and the narrow siphonal area

without constricted base. Contrary to the strength of

the primary spiral cords, the width/length index and

the size are features variable in most other related

généra.

Hesperisternia Gardner, 1944 (type species:

Hersperisternia waltonia Gardner, 1944: 445-447,

from the Miocene of Florida), has a similar sculpture

consisting of fine spiral cords and a columellar fold on

the transition to the siphonal canal. The strongly

sculptured species of that genus look much different at

first glance, but the more subtly sculptured species

like H. multangula grandanus (Abbott, 1986) and H.

shaskyi (Berry, 1959) hâve a similar spiral sculpture

consisting of a few primary spiral cords with many

fine secondary spiral threads of equal strength in the

interspaces. Hesperisternia species differ from the

new genus by the presence of 3 dominant spiral cords

on the periphery (while in Micrologus gen. nov. ail

primary spiral cords, also on the base, are of a rather

equal strength); the secondary spiral cords of unequal

strength, the decreasing number of axial ribs towards

the body whorl; the presence of well developed

sculpture in the aperture and the shorter, slightly

twisted siphonal canal with a slightly more

pronounced umbilical fissure and a slightly more

constricted area between base and siphonal canal.

Hesperisternia waltonia Gardner, 1944 (type

locality: “No. 3742, Shell Bluff, Shoal River, Walton

County, Florida”), the type species of the genus, is

described with 3 protoconch whorls while Recent

species of that genus hâve 1 to 1 3 A protoconch whorls.

Hesperisternia shaskyi (Berry, 1959) (type

locality: south olï Guaymas, Sonora), which is in fact

not a typical species of its genus, is ornamented with a

spiral sculpture most similar to Micrologus gen. nov.

(evenly spaced primary spiral cords, secondary spiral

cords of more or less equal strength) and a weakly

sculptured aperture, but differs (apart from the broad

shape and distinct range) by the more constricted

shape of the base, the shorter siphonal canal and by

the broad spiral cords on the siphonal canal.

K. Fraussen & J. Rosado

Novapex 12(3-4): 73-79, 10 octobre 2011

The range of Hesperisternia is restricted to East

Pacific and West Atlantic warm waters (at présent no

Indo-West Pacific species are known) while

Micrologie gen. nov. is known from eastern Africa

which is the other side of the planet).

For a detailed account on the fossil and recent

Hesperisternia species we refer to Vermeij (2006: 81-

82, 89-91).

Pollia Gray in Sowerby, 1834 (type species:

Buccimm undosum Linnaeus, 1758: 740) may look

similar in shape and size, but differs by the spiral

sculpture consisting of cords of a rather irregular

strength, the more oval shape with shorter siphonal

canal and the presence of sculpture on the columella.

Cancellopollia Vermeij & Bouchet, 1998 (type

species: Cancellopollia gracilis Vermeij & Bouchet,

1998: 480-483) differs by the more oval shape and the

bigger spiral cords of unequal strength.

Species with a similar columellar callus, for

example Pollia shepstonensis Tomlin, 1926,

Buccimm cinis Reeve, 1846 and Triton egregia

Reeve, 1844, ail of uncertain generic placement (often

placed in Engina J. E. Gray, 1839 or Prodotia Dali,

1924), may look similar in size, shape and colour, but

differ by the broader spiral cords, especially on the

base and by the presence of knobs or lirae on the

columella.

Etymology. Micrologus gen. nov. is named after the

book “Micrologus”, written in 1026 by Guido

d’Arezzo and one of the two most popular books

about music in médiéval times. The musical notation

(staff notation) introduced in that work is still used

nowadays. The name refers to the fine spiral fines of

the shell, which are 5 in nurnber when counting the

secondary spiral threads and adjacent primary spiral

cords, which is similar to, and the same nurnber of the

staff notation. This sculpture is also présent on the

base of the shell, contrary to other known généra in

the group, but similar to the ensemble of staffs on a

score.

Micrologus mochatinctus sp. nov.

Figs 1-6

Type material. Hoiotype, 15.0 mm, Mozambique

Channel, Almirante Leite Bank, MAINBAZA stn.

DW3169, 26°H’S, 35°01'E, 450 deep, MNHN-

23772.

Paratype 1, 9.5 mm, juvénile (protoconch), same

locality, MNHN-23773.

Paratypes 2-5, 13.4-13.8 mm, same locality, MNHN-

23773, MHNM, JR, KF-6130.

Type locality. Mozambique Channel, Almirante Leite

Bank, MAINBAZA stn. DW3169, 26° 11 ’S, 35°01 ’E,

450 m deep.

Material examined. Mozambique Channel,

Almirante Leite Bank, MAINBAZA stn. DW3167,

26°12’S, 35°02'E, 228-230 m, 5 dd (1 fragment). -

Stn DW3169, 26°11’S, 35 o 01’E, 450 m, 8 dd (1 juv.,

4 fragments).

Range and habitat. Only known from Almirante

Leite Bank. Bathymétrie range, ail empty shells,

between 228 and 450 m.

Syntopic with Pollia imprimelatus sp. nov. on both

DW3167 and DW3169 stations.

Almirante Leite Bank is an area with strong currents

overflowing a bottom of hard rocks. The habitat is

characterized by the presence of rock, coral, sponges

and gorgonians (Text Fig. 1).

Description. Shell small (up to 15.0 mm). Shape

semi-oval, spire fusiform, base slightly stretched.

Axial sculpture dominant in combination with

accentuated spiral pattern. Protoconch paucispiral,

consisting of about I smooth whorl with rather rough

surface. Transition to teleoconch indistinct, marked by

a fine incrémental fine and the start of the sculpture of

the teleoconch. Teleoconch with 5 14 whorls. Colour

white with brown spiral bands; upper spire whorls

white, with small brown dots on axial ribs in between

spiral cords; spiral interspaces of lower spire whorls

and body whorl omamented with broad, chocolaté

brown spiral bands, usually darker on axial ribs,

secondary spiral threads usually brown, primary spiral

cords snow-white. Narrow band along subsutural

slope and tip of siphonal canal snow-white.

First teleoconch whorl with 4 fine spiral cords, 2

subsutural ones slightly finer, interspaces of equal

size. Second whorl with 5 primary spiral cords,

abapical one rather big, gradually decreasing in

strength towards upper suture, adapical spiral cord

finer. Third whorl with a single fine secondary spiral

thread (hoiotype) or cord (paratype 1 ) in the middle of

each spiral interspace. Spiral interspaces gradually

becoming broader, at First on periphery; nurnber of

secondary spiral threads increasing to 3 on fourth

whorl and body whorl. Spiral sculpture fine, of equal

strength along the whorl, occasionally slightly higher

when Crossing axial ribs; primary spiral cords evenly

spaced also on base, slightly narrower on subsutural

slope; secondary spiral threads of equal strength.

First teleoconch whorl with 11 narrow axial ribs;

abapically slightly weaker or ended, resulting in a

rather constricted suture; interspaces broad. Axial ribs

gradually becoming broader, their nurnber slightly

increasing to 10 on second whorl, 9 on third whorl.

Penultimate whorl with 11 axial ribs, body whorl with

13 ribs, including prelabral varix. Axial ribs running

from suture to just below periphery on body whorl.

Aperture large, ovate, adapically slightly pinched with

a single columellar knob and a labral knob bordering

the anal notch. Columella concave, smooth and

glossy, with an adapical columellar knob (bordering

the anal notch) and a strong abapical columellar knob

on transition to siphonal canal. Callus thin, smooth,

glossy, broad; abapical part more developed, forming

a broad layer well adhèrent to columella. Outer lip

K. Fraussen & J. Rosado

The Cantharus group on Almirante Leite bank

rounded, lip thick, edge sharp, wilh broad but low

internai knobs, not situated according to primary spiral

cords on outer side; adapical knob slightly bigger,

occasionaily split; abapical knob slightly bigger, on

transition to siphonal canal. This abapical knob is not

situated in front of the abapical columellar knob, but

strongly diagonally orientated abapically of it. Labral

varix low but broad, slightly prosocline. Siphonal

canal short, broad. open.

Animal and operculum unknown.

Comparison. Micrologus mochatinctus sp. nov. is

characterized by the fine spiral sculpture consisting of

equally spaced primary spiral ribs of equal strength

with the interspaces omamented with 3 (occasionaily

4) fine secondary spiral threads of equal strength, in

combination with a smooth columella.

Ail possibly similar species of other groups differ

by the presence of stronger spiral cords on the base

and by the presence of knobs and lirae on the

columella.

Etymology. Micrologus mochatinctus sp. nov. is

derived from “mocha”, originally Mocha in Yemen

where coffee has been exported since ancient times,

now used for a mixture of coffee and chocolaté, and

for expressing the brown colour; in combination with

“tinctus” (Latin), meaning “a dye”.

ACKNO WLEDG M ENTS

We are grateful to Philippe Bouchet, Philippe

Maestrati and Virginie Héros (MNHN, France) for

making the material avai labié for study. Guido T.

Poppe (Belgium, Philippines) for support and sharing

his knowledge. Philippe Poppe (Philippines) for

procuring material for comparison. Alain Fraussen

(Belgium) for etymological help. David Monsecour

(Belgium) for correcting the English text.

REFERENCES

Bouchet, P. & Warén, A. 1986. Mollusca Gastropoda:

Taxonomical notes on tropical deep water

Buccinidae with descriptions of new taxa. In:

Résultats des Campagnes MUSORSTOM. I & II.

Philippines (1976-1980). Tome 2. Mémoires de

Muséum national d'Histoire naturelle, série A,

Zoology, 133:455-499, text figs. 1-4, pis. 1-18.

Gardner, J., 1944. The molluscan fauna of the Alum

Bluff Group of Florida. VII. Stenoglossa (in part).

United States Geological Survey. Professional

Papers, 142-G: 437-491.

Linnaeus, C., 1758. Systema naturae per régna tria

naturae. Editio décima, reformata. Vol. 1, Regnum

animale. Stockholm, 824 pp.

Sowerby, G. B., 1834. The Généra of Recent and

Fossil Shells, vol. II. London, pis. 127-262.

Vermeij, G. J. & Bouchet, Ph., 1998. New Pisaniinae

(Mollusca, Gastropoda, Buccinidae) from New

Caledonia, with remarks on Cantharus and related

généra. Zoosystema, 20(3): 471-485.

Vermeij, G. J., 2006. The Cantharus Group of

Pisaniine Buccinid Gaspropods: Review of the

Oligocène tp Recent Généra and Description of

Some New Species of Gemophos and

Hesperisternia. Cainozoic Research, 4(1-2): 71-

96.

Figures 1-13

1-6. Micrologus mochatinctus gen. & sp. nov., holotype MNHN-23772, 15.0 mm, Mozambique Channel

Almirante Leite Bank, MAINBAZA stn. DW3169, 26°1LS, 35°0LE, 450 m deep.

7-11. Pollia imprimelata sp. nov.

7-8. Holotype MNHN-23770, 19.9 mm, Mozambique Channel, Almirante Leite Bank, MAINBAZA stn.

DW3 |6 7, 26°12’S, 35°02’E, 228-230 m deep; 9-10. Paratype 1, MNHN-23771, 21.3 mm, same locality;

1. Piotoconch of paratype 8, MNHN-23771, diameter 1.1 mm, same locality.

12. Pollia sp., 17.6 mm, Mozambique Channel, Almirante Leite Bank, MAINBAZA stn. DW3168

-6° 12 S, 35°03’E, 87-90 m deep, MNHN.

P' VWVl 61 ^’ ana (Melvill & Standen, 1903), northwestern Madagascar, between Majunga and Cape

St. André, MIRIKY stn CP3260, 15°35’S, 45°45’E, 179-193 m deep, MNHN.

K. Fraussen & J. Rosado

Novapex 12(3-4): 73-79, 10 octobre 2011

L. G. Brown & B. D. Neville

Novapex 12(3-4): 81-86, 10 octobre 2011

Nomenclatural notes on Amaea arabica (Nyst, 1871) comb. nov.

and Cirsotrema fimbriolatum (Melvill, 1897) (Gastropoda: Epitoniidae),

two similar species from the Indo-Pacifîc faunal province

Leonard G. BROWN

5 Vumbaco Drive, Wallingford, CT 06492, USA

E-mail: Epmanshell@AOL.com

Bruce D. NEVILLE

2700 Sandy Circle, College Station, TX 77845-5309, USA

E-mail: Bneville@tamu.edu

KEYWORDS. Gastropoda, Epitoniidae, Amaea, Cirsotrema, nomenclature.

ABSTRACT. Seal aria decussata ‘ Lamarck’ Kiener, 1838, and S. decussata ‘Lamarck’ Sowerby

II, 1844, are shown to refer to a Recent species different from the true S. decussata Lamarck,

1804, a fossil. The illustration of the holotype of Scalaria arabica Nyst, 1871, is compared to

photographs of the probable holotype of Cirsotrema kieneri Tapparone-Canefri, 1876, to

document that the two species are conspecific, that Nyst’s name has precedence, and that S.

arabica belongs to the genus Amaea. Scalaria fimbriolata Melvill, 1897, a similar species from

the Indo-Pacific faunal province that has been confused with A. arabica, is shown to be

specifically and generically distinct from A. arabica.

INTRODUCTION

A review of the literature on the Epitoniidae, e. g.

Nakayama (2003: 21) illustrâtes that questions exist

regarding the relationship between Scalaria decussata

Lamarck, 1804, an Eocene fossil species collected

near Paris, France, and the Recent species Scalaria

arabica Nyst, 1871 [= Scalaria decussata Lamarck

sensu Sowerby II (1844: 103, pi. 35, fig. 140)] and

Cirsotrema kieneri Tapparone-Canefri, 1876

[=Scalaria decussata Lamarck sensu Kiener (1838:

21, pl- 7, fig. 23)].

To résolve these questions, we set out to track

down photographs of the type specimens of S.

decussata, S. arabica, and C. kieneri and also

conducted a thorough review of the literature that

included référencés to these species names. In the

course of our research, we discovered that the species

listed above hâve also been confused with the species

described under the name Scalaria fimbriolata

Melvill. 1897, a similar Recent species referable to the

genus Cirsotrema. Therefore, in addition to

documenting our conclusion that the correct name for

the species illustrated by Kiener and Sowerby is

Amaea arabica (Nyst, 1871), we hâve included a

discussion of Melvill’s species to document that it is

not a synonym of A. arabica.

Abbreviations

NHMUK: Natural History Muséum, London.

MHNG: Muséum d’EIistoire Naturelle de Genève,

dd: specimen(s) collected dead.

Iv: specimen(s) collected alive.

SYSTEMATICS

Family EPITONIIDAE S. S. Berry, 1910

Genus Amaea H. & A. Adams, 1853: 223

Type species: Scalaria magnifica G. B. Sowerby II,

1844, by subséquent désignation (Boury, 1909: 258)

Amaea decussata (Lamarck, 1804)

Fig 1-2

Scalaria decussata Lamarck, 1804: 213; Lamarck,

1806: pl. 10, fig. 3; Lamarck, 1822: 229; MHNG,

1918: pl. 6, figs. 74 a, b, 75 a, b; non Scalaria

decussata Pease, 1867: 289 [=Epitonium

sandwichense (Nyst, 1871)].

Distribution. France. Fossil, Eocene.

Reniarks. The original engravings of Lamarck’s S.

decussata (Lamarck, 1806) show both spiral and axial

sculpture, though the axial sculpture overlays the

spiral. Photographs of the syntypes of Lamarck’s S.

decussata published by the MHNG (1918) show that

this fossil species has convex teleoconch whorls that

are not angular below the suture and the axial costae

are not raised where they cross the spiral cords (Fig.

1-2). The sculpture indicates that the species is

correctly placed in the genus Amaea.

Pease (1867: 289) also named a Scalaria

decussata. Pease’s shell, however, is an Epitonium

and the name was also replaced by Nyst (1871: 132)

with Scalaria sandwichensis.

L. G. Brown & B. D. Neville

Nomenclatural notes on Amaea arabica and Cirsotremafimbriolatum

Amaea arabica (Nyst, 1871)

Figs 3-7, 11

Scalaria decussata Kiener, 1838: 21, pl. 7, fig. 23;

Sowerby II, 1844: 103, pl. 35, Fig. 140; Sowerby II,

1874: species 114, pl. 15, fig. 114a; Clessin, 1897: 39,

pl. 12, fig. 2.; non Scalaria decussata Latnarck, 1804.

Scalaria arabica Nyst, 1871: 105; Boury, n.d., pl. 35,

fig. 1, 3. Nom. nov. for Scalaria decussata Sow. (non

Lam.). Type locality: coast of Arabia.

Cirsotrema Kieneri Tapparone-Canefri, 1876: 155;

Kaicher, 1983: 3584. Nom. nov. for Scalaria

decussata Kiener (non. Lam.). Type locality: none

given.

Amaea Sowerbyi Dunker, 1882: 69; nom. nov. for

Scalaria decussata Sow. (non Lam.)

Scalaria kieneri : Tryon, 1887: 81, pl. 17, fig. 21, 22,

26.

Amaea decussata : Cleevely, 1980: 240, fig. 2, 7.

Amaea {Amaea) decussata'. Weil, et. al., 1999: 82, fig.

234.

Material Examined. Urangan, south Queensland,

Australia, 2 dd. Palandra Beach, Townsville,

Queensland, Australia, ldd. Swan Reefs, Queensland,

Australia, trawled, 1 dd.

Distribution. Red Sea, south to Mauritius, east to

Queensland, Australia. Intertidal to 15 m.

Remarks. In their respective monographs of the

genus Scalaria, Kiener (1838) and Sowerby II (1844)

each illustrated species labeled Scalaria decussata

Lamarck. Nyst, in his Tableau Synoptique et

Synonymique (the “Tableau") presented at the

December 3, 1871, meeting of the Société

Malacologique de Belgique, erected the replacement

name Scalaria arabica for the species illustrated by

Sowerby. Apparently, Nyst concluded that the species

illustrated by Sowerby was not the same as Lamarck’s

fossil species. Tapparone-Canefri (1876) erected the

replacement name Cirsotrema kieneri for the species

illustrated by Kiener because it was his opinion that

Kiener’s species was distinguishable from both S.

decussata Lamarck, 1804, and S. decussata sensu

Sowerby II, 1844. Dunker ( 1882) also recognized that

Sowerby’s species differed from Lamarck’s and

erected the replacement name Amaea Sowerbyi, which

is a junior objective synonym of Scalaria arabica

Nyst, 1871. While Tryon (1887: 81) agreed that the

species illustrated by Kiener was distinct from

Lamarck’s fossil species, he went on to say that in his

opinion, Kiener and Sowerby illustrated the same

Recent species. Tryon, who apparently overlooked

the replacement name erected by Nyst, used the name

Scalaria kieneri for this species. Cleevely (1980: 240)

commented that the species labeled Amaea decussata

in his paper is virtually indistinguishable from

‘ Cirsotrema ’ kieneri, documenting that there were not

only questions regarding a possible synonymy at the

species level, but also questions regarding the correct

generic assignment of this species.

Because Nyst erected the replacement name S.

arabica based on the specimen figured by Sowerby

under the name S. decussata, Sowerby’s figured

specimen is the holotype of S. arabica (ICZN, 1999:

Art. 72.7). Kathie Way at NHMUK has informed us

that the specimen from Arabia figured by Sowerby

under the name Scalaria decussata is supposed to be

in the collection at her institution. She was, however,

unable to locate it. We hâve there fore reproduced an

enlarged illustration of Sowerby’s figure (Fig. 7).

Similarly, because Tapparone-Canefri erected the

replacement name C. kieneri for the specimen figured

by Kiener under the name S. decussata, Kiener’s

specimen is the holotype of C. kieneri. We hâve

reproduced Kiener’s figures (Fig. 5-6). Kiener noted

that the figured specimen is in “the Muséum

collection.” Kiener’s material, which presumably

included this specimen, was acquired by Delessert and

is now in the collection at the MHNG (Y. Finet pers.

comm.). Yves Finet provided photographs of the sole

specimen of Scalaria decussata in the Delessert

collection (Fig. 3-4). Because it matches well, and is

consistent in size with, the specimen figured by

Kiener, we consider this specimen to be the probable

holotype of Scalaria decussata Kiener, 1838, and, by

extension, Cirsotrema kieneri.

Comparison of these figures confions that the

species figured by Kiener is conspecific with the

species figured by Sowerby. Both specimens hâve the

same angular teleoconch whorls with cancellate

sculpture and the basal ridge and the sculpture on the

base of the shell appears to be the same. Because of

the combination of the cancellate sculpture on the

convex teleoconch whorls, the occasional varices, and

the strong basal ridge, we consider this taxon to be

referable to the genus Amaea.

Figures 1-12

1-2. Scalaria decussata Lamarck, 1804, MHNG, syntypes, photo of Fig 74 & 75 in Catalogue illustré de la

Collection Lamarck. Mollusques Trachélipodes Fossiles, with permission. 3-4,11. Scalaria decussata Lamarck,

sensu Kiener, 1838, length 40 mm, no locality data, MNHG 67919, probable holotype of Cirsotrema kieneri

lapparone-Canefri, 1876. 5-6. Scalaria decussata Lamarck, sensu Kiener, 1838 (Kiener’s figure). 7. Scalaria

decussata Lamarck, sensu Sowerby II, 1844 (Sowerby’s figure), holotype of Scalaria arabica Nyst, 1871.

8-9. Scalaria fimbriolata Melvill, 1897, length 19 mm, width 5 mm, Karachi, NHMUK 1897.7.30.90.

(holotype). 10, 12. Cirsotrema fimbriolatum (Melvill, 1897), length 59.6 mm, width 14.5 mm, Broome, Western

Australia, 15 m, B. Neville coll. no. 1256.

L. G. Brown & B. D. Neville

Novapex 12(3-4): 81-86, 10 octobre 2011

L. G. Brown & B. D. Neville

Nomenclatural notes on Amaea arabica and Cirsotrema fimbriolatum

Therefore, Tryon and Tapparone-Canefri were

correct in concluding that the Recent species figured

by Kiener and Sowerby is not synonymous with the

fossil species S. decussata Lamarck, 1804, because S.

decussata has a different sculpture that is évident in

the photographs of the syntypes.

Finally, in the course of preparing this manuscript,

we discovered there is a question regarding the

publication date of Nyst’s Tableau, which included his

replacement name S. arabica, as well as replacement

names for a number of other epitoniid taxa. While it

has generally been given as 1871, the actual

publication date could be as late as 1874. The World

the date of Elegantiscala arabica as 1872, though the

source of the date is “not documented.” Nyst’s other

names from the Tableau are cited in WoRMS as 1871.

Nakayama (2003: 21) gives the date of Scalaria

arabica as 1873, citing Bouiy (1913). This is

evidently a typographical error on Nakayama’s part,

however, as Boury (1913: 103) clearly gives 1871 as

the date for the Tableau ; Nakayama (2003: 94) in his

own bibliography gives the date of the Tableau as

1871. Because the Tableau was presented at a

December 3, 1871, meeting, we question whether it

would hâve been published that same year. When we

checked the Zoological Record , we discovered that

Nyst’s Tableau appeared for the first time in the 1874

volume, although it was listed with an 1871 date.

While not definitive, it is an indication that the

Tableau was published after 1871 and certainly no

later than 1874. Buckhuys (1985) prepared a

bibliographie note on the journals published by the

Société Malacologique de Belgique. However, he

began with the volume for 1872 and did not provide

information on individual publication dates. Because

we were unable to résolve this publication date

question, we are following earlier authors in using an

1871 publication date for the Tableau.

Notwithstanding the uncertainty regarding the

publication date, since it is clear that Nyst’s Tableau

was published prior to Tapparone-Canefri’s 1876

paper proposing the name C. kieneri for this Recent

species, there is no question that the oldest available,

and valid, name for the Recent taxon is therefore

Nyst’s replacement name, Amaea arabica (Nyst,

1871).

Genus Cirsotrema Môrch, 1852: 49

Type species: Scalaria varicosa Lamarck, 1822, by

monotypy.

Cirsotrema Jimbriolatum (Melvill, 1897)

Figs 8-10, 12

Scalaria fimbriolata Melvill, 1897: 11, pl. 6, fig. 10;

Melvill, 1898: 2, pl. I, fig. 12. Type locality: Karachi.

Scala ( Cirsotrema ) fimbriolata : Melvill and Standen,

1903: 349.

Cirsotrema kieneri: Wilson, 1993: 274, pl. 44, fig. 3;

Weil, et. al. 1999: 128, fig. 401 (non Tapparone-

Canefri, 1876).

Epitonium fimbriolatum: Bosch and Bosch, 1982: 52.

Amaea fimbriolata: Bosch, Dance, Moolenbeek, et.

al., 1995: species no. 400.

Amaea ( Scalina ) kieneri: Nakayama, 2003: 21, pl. 20,

fig. 1-3 (non Tapparone-Canefri, 1876).

Material Examined. Off Hervey Bay, Queensland,

Australia, 61-79 m, 1 dd. Urangan, south Queensland,

Australia, 1 dd. Broome, Western Australia, 15 m, 1

lv. 80 Mile Beach, Western Australia, 1 Iv.

Distribution. Gulf of Oman and Persian Gulf, east to

Japan and Queensland, Australia. Intertidal to 79 m.

Remarks. Cirsotrema fimbriolatum has also been

confused with A. arabica, but differs from it markedly

in structure, indeed belonging to a different genus.

The original figures of Scalaria fimbriolata (Melvill,

1897, and particularly Melvill, 1898) illustrate

particularly well the characters of this species.

Unfortunately the quality of the original image is not

suitable for enlargement and reproduction here.

While it is superficially similar to A. arabica, this

species can be distinguished by the characteristic

Cirsotrema costae and the base of the shell. The base

of C. fimbriolatum is ringed by a sériés of triangular

projections resembling teeth on a cog, whereas A.

arabica has a simple basal ridge lacking these

triangular projections (Fig. 1 I, 12).

Scalaria fimbriolata is referable to the genus

Cirsotrema because of the combination of the strong

basal disk, spiral lirae, and the strongly crispate costae

that consist of numerous plates that are fused together.

This combination of teleoconch characters is présent

in Cirsotrema varicosum (Lamarck, 1822) and a

number of other species that hâve been referred to the

genus Cirsotrema s.s. Species in the genus Amaea

can hâve similar teleoconch scupture, but lack the

crispate costae consisting of numerous fused plates.

Acknowledgments

We want to thank Yves Finet at the MHNG,

Geneva, Switzerland, who furnished the photographs

of the specimen of S. decussata (sensu Kiener) in the

Delessert collection and authorized us to reproduce the

figured specimens of S. decussata Lamarck in this

manuscript. We also want to thank Kathie Way at

NHMUK, London, England, who provided

information on the holotype of S. arabica and

furnished the photographs of the holotype of S.

fimbriolata. Henry Domke photographed the

i 11 ustrated specimen in the Bruce Neville collection.

Alan J. Kohn ot University of Washington Biology

provided excerpts of the catalog of Lamarck’s types.

Emilio Garcia called our attention to the question of

the dates of Nyst’s Tableau , and Paul Callomon at the

L. G. Brown & B. D. Neville

NOVAPEX 12(3-4): 81-86, 10 octobre 2011

Academy of Natural Sciences, Philadelphia and

Richard E. Petit provided helpful comments on this

question. Lawrence Gall of the Peabody Muséum of

Natural History, Yale University, prepared the plate.

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J. Trôndle & J. Letourneux

Novapex 12(3-4): 87-90, 10 octobre 2011

Description de Terebra niauensis n. sp.

(Mollusca: Gastropoda: Terebridae) du Pléistocène de Niau,

Tuamotu (Polynésie Française)

Jean TRÔNDLÉ

Attaché au Muséum national d'Histoire naturelle

Département Systématique et Evolution

55, rue de Buffon, 75005 Paris, France

j .trondle@orange.fr

Jean LETOURNEUX

Mahina - Tahiti - Polynésie Française

natualeyla@mail.pf

MOTS-CLEFS. Mollusca, Gastropoda, Terebridae, Polynésie Française, Pléistocène

KEYWORDS. Mollusca, Gastropoda, Terebridae, French Polynesia, Pleistocene

RÉSUMÉ. Une nouvelle espèce Terebra niauensis n.sp. du Pléistocène est décrite de Niau,

Archipel des Tuamotu (Polynésie Française) et est comparée à Terebra gouldi Deshayes, 1857,

espèce proche actuelle et endémique des Iles Flawaii.

ABSTRACT. Terebra niauensis n. sp. is described from Niau, Tuamotu Archipelago (French

Polynesia) dated Pleistocene and is compared with Terebra gouldi Deshayes, 1857, a quite similar

présent species endemic to the Hawaiian Islands.

INTRODUCTION

Les tests de Terebra niauensis n. sp. proviennent de

sables coralliens et coquilliers récoltés sur l’atoll de

Niau. L’atoll présente actuellement une altitude de 7,5

m et son lagon saumâtre est sans communication

directe avec l'océan et d’une profondeur moyenne de 2

m. Les sables détritiques extraits en bordure du lagon

correspondraient à un niveau marin du dernier

interglaciaire (Pléistocène, environ 125 000 ans),

niveau plus élevé qu’actuellement (de 6 à 10 m) alors

que le lagon communiquait avec l’océan. Dans ces

sédiments détritiques une autre espèce de mollusque

gastropode a été récemment décrite par Trôndlé et

Salvat (2010).

Abréviations

BMNH: The Natural History Muséum, London, U.K.

CRIOBE: Centre de Recherches Insulaires et

Observatoire de l’Environnement, Moorea, Polynésie

française.

EPHE: École Pratique des Hautes Études, Perpignan,

France.

1RSNB: Institut royal des Sciences naturelles de

Belgique.

MNHN: Muséum national d'Histoire naturelle, Paris,

France.

AW: Anders Warén, Swedish Muséum of Natural

History, Stockholm, Sweden.

GK: Gijs Kronenberg, Netherlands.

GP: Gustav Paulay, Florida Muséum of Natural

History, University of Florida, Gainesville, Florida,

USA.

JL: Collection Jean Letourneux.

JT: Collection Jean Trôndlé.

MB: Collection Michel Boutet.

PF: Polynésie Française.

SYSTÉMATIQUE

Famille TEREBRIDAE Môrch, 1852

Genre Terebra Bruguière, 1789

Espèce type: Buccinum subulatum Linné, 1767, par

monotypie, Lamarck, 1799

Terebra niauensis n. sp.

Figs 1-5, 8, 9

Matériel type. 18 individus; holotype: 96,5 mm,

MNHN 23685 (Figs. 1, 2); paratype: 89,5 mm,

CRIOBE (Fig. 3, 8, 9); 9 paratypes MNHN 23686: 78

mm (Fig. 4), 81,6 mm, 74 mm, 62,7 mm, 62 mm, 60,4

mm, 55 mm. 50,7 mm et 49,8 mm; paratype 61,2 mm,

JT (Fig. 5); paratype 86,1 mm, JL; paratype 84 mm,

GP; paratype 79,1 mm, AW; paratype 87,5 mm,

(Mairie de Niau); paratype 87,8 mm, MB; paratype

60,6 mm, GK.

Localité type. Atoll de Niau, Tuamotu, Polynésie

Française, 16°08’S, 146°20’W.

J. Trôndle & J. Letourneux

Terebra niauensis n. sp. de Polynésie Française

Description de l'holotype. Coquille d'une hauteur de

96,5 mm, d'aspect brillant et de couleur uniformément

blanche; sous UV le test apparaît de teinte ivoire et

présente quelques taches brunes sous la suture. La

protoconque est absente et la téléoconque compte 14

tours. Les tours sont légèrement convexes. La bande

sous-suturale est délimitée par une suture profonde et

un sillon sous-sutural bien marqué; elle est ornée de

petits nodules, 12 sur le dernier tour, en forme

d'accent grave occupant toute la bande sur les

premiers tours puis plus espacés et se détachant

sensiblement de la suture sur les tours suivants.

L'espace entre ces nodules est orné de fines stries

axiales occupant toute la hauteur de la bande sous-

suturale, Les tours sont sculptés de très nombreuses

côtes axiales, anguleuses sur les premiers tours et

d'épaisseur variable; ces côtes sont dans le

prolongement des stries et des nodules de la bande

sous-suturale. La sculpture axiale est obsolescente en

avant des derniers tours et sur le dernier tour. Absence

de sculpture spirale. L'avant dernier tour présente un

orifice circulaire de 5 mm de diamètre, empreinte d'un

prédateur. Le dernier tour et l'ouverture sont allongés.

Le diamètre du dernier tour est de 20 mm et la hauteur

de l'ouverture de 24 mm. La lèvre est fine. La

columelle est légèrement convexe, oblique et présente

un pli médian.

Distribution. Uniquement connue de la localité type.

Remarques. La taille maximum observée est celle de

l'holotype. La protoconque est absente chez tous les

exemplaires examinés. L'espace entre les nodules de la

bande sous-suturale et leur nombre (de 12 à 20) sur le

dernier tour sont variables. Quatorze des dix-huit

exemplaires examinés présentent une perforation

circulaire, témoin de la présence de prédateurs dans le

même habitat. Ces perforations régulières, plus larges

à l'entrée qu’au fond sont typiques de celles

occasionnées par les Naticidae (Cemohorsky, 1971).

Notocochlis gualteriana (Récluz, 1844) et Polinices

tumidus (Swainson, 1840), présents en grand nombre

dans les sédiments étudiés de la thanatocénose, sont

probablement ces prédateurs.

Terebra niauensis n. sp. est proche de Terebra

gouldi Deshayes, 1857 (Figs 6, 7) par sa taille et son

aspect général. Elle en diffère par sa sculpture axiale

plus irrégulière au niveau de la bande sous-suturale et

plus fine au niveau des tours. Cette sculpture tend à

être obsolescente sur les derniers tours alors qu'elle

reste constante et bien marquée chez T. gouldi. Le

dernier tour et l'ouverture de T. niauensis sont

nettement plus allongés. L'observation sous UV n'a

mis en évidence que quelques taches sous la suture

alors que le test de T. gouldi est orné de maculations

régulières sur la bande sous-suturale et de deux bandes

brunes sur le dernier tour.

Le récent inventaire des mollusques de Polynésie

Française (Trôndlé & Boutet, 2009) fait état de la

présence de 66 espèces de Terebridae dont 59

clairement identifiées. Parmi les grandes espèces

Terebra dimidiata (Linné, 1758), espèces à large

répartition indo-pacifique, est la seule a posséder un

aspect général relativement proche de Terebra

niauensis. Cependant la sculpture de Terebra

dimidiata est constante (Bratcher & Cernohorsky,

1987) et son test entièrement lisse, à l'exception des

tout premiers tours, à la différence de Terebra

niauensis. Une autre espèce Terebra caledonica

Sowerby, 1909, récemment redécouverte (Conde &

Terryn, 2003), endémique de l'Ile des Pins,

extrêmement variable dans sa sculpture, possède

certaines formes également proches de Terebra

niauensis mais en diffère par sa taille plus petite et son

aspect général plus trapu.

Plusieurs études ont été réalisées sur les faunes

malacologiques du pléistocène des îles avoisinantes de

la Polynésie Françaises où peu ou pas de Terebridae

ont été signalés: Tongatabu [Ostergaard , 1935 (T.

cerithina, T. subulata)], Hawaii [Ostergaard, 1939 (T.

gouldi)], Henderson [Spencer & Paulay, 1989 (aucun

Terebridae)]. En revanche 9 espèces sont citées par

Ladd (1982, p. 81-85, pl. 30) du pléistocène des

Nouvelles Hébrides (Vanuatu). Parmi elles, deux

espèces sont proches de T. niauensis: T. dimidiata

[déterminée de façon erronée comme T. cf. felina (

pl.30, fig. 10-12)] et Terebra (Oxymeris) interlineata

Deshayes (pl. 30, fig. 9). La détermination de cette

dernière est faite par Ladd sur un seul exemplaire. T.

interlineata, considérée comme synonyme de la très

variable Terebra crenulata (Linné, 1758), possède une

sculpture axiale de ses premiers tours différente; par

ailleurs seules des lignes brunes ornent la bande sous

suturale et non des taches. L'exemplaire de Ladd est

difficile à séparer des exemplaires juvéniles, de taille

identique, de T. niauensis.

Etymologie. L'espèce est nommée d'après la localité

type.

Figures 1-9

1-5, 8, 9. Terebra niauensis n. sp. 1, 2. Holotype MNHN 23685, 96,5 mm; 3, 8, 9. Paratype CRIOBE, 89,5 mm:

4. Paratype MNHN 23686, 78 mm; 5. Paratype Coll. JT, 61,2 mm.

6, 7. Terebra gouldi Deshayes, Lectotype 197962/1 BMNH, 61,6 mm.

J. Trôndle & J. Letourneux

Novapex 12(3-4): 87-90, 10 octobre 2011

J. Trôndle & J. Letourneux

Terebra niauensis n. sp. de Polynésie Française

Remerciements

Nous remercions Ludwig Blanc (PF) qui le premier a

attiré notre attention sur la présence de coquilles

fossiles sur l'atoll de Niau, Philippe Bacchet (PF) et

Robert Gourguet (PF) qui ont activement participé aux

récoltes, Michel Boutet (PF), connaisseur éclairé de la

faune malacologique actuelle de Polynésie française,

Pierre Lozouet (MNHN) et Yves Terryn (IRSNB) qui

nous ont prodigué leurs conseils afin d'améliorer le

manuscrit, Amelia MacLellan (BMNH) qui a mis à

notre disposition les photos des types de Terebra

gouldi Deshayes, Philippe Maestrati (MNFIN) pour les

photos et la réalisation de la planche et Didier Merle

(MNFIN) qui a permis l'observation sous lumière UV

du matériel étudié.

REFERENCES

Bratcher, T. & Cemohorsky, W.O., 1987. Living

Terebras of the World. A monograph of the Recent

Terebridae of the World. American Malacologists,

Melbourne, Fia. 236p.

Cernohorsky, W.O., 1971. The Family Naticidae

(Mollusca: gastropoda) in the Fiji Islands. Records

of the Auckland Institute and Muséum, 8: 169-208.

Condé, J. & Terryn, Y., 2003. Additional descriptive

notes on Terebra caledonica Sowerby, 1909.

Gloria Maris, 43(2-3): 31-39.

Ladd, H.S., 1982. Cenozoic fossil mollusks from

western Pacific islands; Gastropods (Eulimidae

and Volutidae through Terebridae). Geological

Survey Professional Paper, 1171: 100p.

Ostergaard, M.J., 1935. Recent and fossil marine

Mollusca of Tongatabu. Bernice P. Bishop

Muséum , Bulletin 131: 3-59.

Ostergaard, M.J., 1939. Report on fossil Mollusca of

Molokai and Maui. Occasional Papers of Bernice

P. Bishop Muséum , Honolulu, Ffawaii, 15(6): 57-

77.

Spencer, T. & Paulay, G., 1989. - Geology and

geomorphology of Flenderson Island. Atoll

Research Bulletin, 323: 18p.

Trôndlé, J. & Boutet, M., 2009. Inventory of marine

molluscs of French Polynesia. Atoll Research

Bulletin, 570: 87p.

Trôndlé, J. & Salvat, B, 2010. La thanatocènose du

lagon de l’atoll de Niau (Polynésie française) avec

la description d’une nouvelle espèce de Strombus

(Mollusca, Gastropoda, Strombidae). Zoosystema,

32(4): 613-623.

R. Houart, S. Gori & P. Ryall

NOVAPEX 12(3-4): 91-97, 10 octobre 2011

New record of Typhinellus labiatus (Cristofori & Jan, 1832) (Gastropoda:

Muricidae) from Sâo Tomé and Principe and discussion about its

classification and geographical distribution

Roland HOUART

Research Associate

Institut royal des Sciences naturelles de Belgique

Rue Vautier, 29, B-1000 Bruxelles, Belgium

roland.houart@skynet.be

Sandro GORI

Via Semesi, 7

57123 Livomo, Italy

sandrogori@fastwebnet.it

Peter RYALL

St Ulrich 16

A-9161 Maria Rain, Austria

peterryalll @hotmail.com

KEY WORDS. Gulf of Guinea, Sâo Tomé and Principe, biogeography, Muricidae, Typhinae.

ABSTRACT. The natural history of the islands of Sào Tomé and Principe is briefly discussed and a

new record of a large typhine from Sao Tomé and Principe is commented. The examined specimens

belongs to Typhinellus and are compared with the type species T. Icihiütus (Cristotori & Jan, 1832),

occasionally better known as T sowerbii (Sowerby, 1833), a junior synonym. A list of the Muricidae

collected in these islands is given in the appendix.

INTRODUCTION

Sào Tomé and Principe Islands are a group of small

islands, relict of an ancient volcanic mountain range

situated otfthe west African coast. They consists of two

principal islands, Sào Tomé and Principe. They lie

along a magmatic geological feature known as the

Guinea line which is a flaw in the African tectonic

plate more or less 1500 kms long that lias served as a

channel for magmas for million ot years. The Guinea

line extends across the océan continent and magmatic

extrusions up through it hâve given lise to majoi

oceanic and continental relief extending from

Southwest to northeast, including the islands ot

Annobon, Sào Tomé, Principe and Bioko and the

mainland features of Mount Cameroon, the Cameroon

highlands and the Jos plateau of Nigeria (Drewes &

Wilkinson, 2004).

The islands of Annobon, Sào Tomé and Principe

are long known for the high level of endemism in theii

biota and some taxa are shared in the latter two. The

three islands are separated from each other and from

the West African coast by océan depth up to 3000 m.

Principe islands is geologically the oldest. Sao Tome

being the largest with 850 square kilometers is situated

on the equator line, south of Principe island and with

other small islets (Ilheu das Cabras, Ilheu Santana,

Sete Pedras, Ilheu das Rolas, Uheus Gabado/San

Miguel and Ilheu Coco near Sâo Tomé; Ilheu Tmhosa

grande, Ilheu Tinhosa pequena, Ilheu Bone de Joquei,

Pedra Gale and Ilheu dos Mosteiros near Principe).

Sâo Tomé, Principe and the small islets fonn the

Republic of Sâo Tomé and Principe.

One of the junior authors (SG) has travelled several

times to both Islands, collecting eight times in Sào

Tomé and twice in Principe. Collecting has been in

various habitats and with different methods (scuba

diving, using tangle nets or washing dead corals and

small stones) both by day and night producing a

sampling from 5 to 40 m depth.

Previous research has been conducted by a number

of earlier collectors as summarized by Fernandes &

Rolân (1993) who themselves collected extensively in

the islands primarily by snorkelling. Other recent

works concerning muricids from the islands include

Fernandes & Rolân (1990), Rolân & Fernandes (1991),

Houart & Rolân (2001). Thanks to the introduction of

scuba facilities in the area it was also possible more

recently to explore the rocky coastlines more

intimately and other new species of Muricidae hâve

been described : Houart (2005), Rolân & Gori (2007),

Houart & Gori (2008).

Among the various muricids collected in both

Islands (see appendix), a large typhine has led to a

careful and refined examination.

Typhinae species hâve a small size, rarely exceeding

40 mm in height, and hâve the particularity to grow

hollow tubes (Fig. 1) situated between each pair of

varices. These anal tubes are gradually closed and

broken during the growth of the shell, only the last tube

R. Houart, S. Gori & P. Ryall

New record of Typhinellus labiatus

remains open and can be long to very long when intact

(Fig. 17).

The situation of these tubes compared to the axial

varices is used among other things as a tool for their

generic classification. These tubes may be either

situated half way between the varices or nearest to

preceeding or succeeding varix, or even originate from

the varix itself. The shell of Typhinae also has a sealed

siphonal canal, four or very rarely five varices per

whorl and, in some généra, a partition (Fig. 1), which is

a lamellar outgrowth, or erect plate, joining the last

varix of the whorl with the previous one. The edge of

the aperture is erect and forms an entire peristome.

The examined protoconchs of ail the Recent species are

paucispiral, consisting of 1.5 to 2 whorls.

Abbreviations

JLD: Coll. Jean-Louis Delemarre - P: Principe - PR:

Coll. Peter Ryall - RFI: Coll. Roland Houart - SG: Coll.

Sandro Gori - ST: Sâo Tomé - lv: live-taken specimen

- dd: empty shell.

Descriptions of spiral sculpture use the terminology

introduced by Merle (1999, 2001):

P - Primary cord; PI - Shoulder cord; P2-P6 - Primary

cords of the convex part of the teleoconch whorl.

SYSTEMATICS

Family TYPHINAE Cossmann, 1903

Genus Typhinellus Jousseaume, 1880

Type species by original désignation: Typhis sowerbii

Broderip, 1833 = Murex labiatus Cristofori & Jan,

1832, Mediterranean, East Atlantic

The shells belonging to Typhinellus are characterized in

having a partition. There are four flange-like, frilled

varices, constricted above the aperture and flaring at its

abapical end; the varical flange of the last teleoconch

whorl extends to almost the tip of the siphonal canal.

The anal tube does not originate from the varix but is

situated near the preceeding varix, and adpressed to the

preceeding partition.

Typhinellus labiatus (Cristofori & Jan, 1832)

Figs 1-29

Murex fistulatus Risso, 1826 (not Muricites fistulatus

Schlotheim, 1820).

Murex labiatus Cristofori & Jan, 1832

Typhis sowerbii Broderip, 1833

Murex tetrapterus Bronn, 1838

Murex syphonellus Bonelli in Bellardi & Michelloti,

1841

Typhis (Typhinellus) tetrapterus var. protetrapterus

Sacco, 1890

Typhis sowerbyi var. fulva Pallary, 1906

Typhis sowerbyi var. minor Pallary, 1906

Typhis (Cephonochelus) recens Nordsieck, 1972

Typhis sowerbyi elongatus Settepassi, 1977

Material examined from Sâo Tomé and Principe

Sâo Tome. In fine sand and silt, 6 m, 1 lv (PR) (Figs

11-14); Ubabudu Reef, NE Sào Tomé, 00°15'804" N,

06°45'569" E, 20 m, 1 dd (PR); Lagoa Azul, NW Sào

Tomé, 1 lv (SG) (Figs 6-7); Lagoa Azul, NW Sào

Tomé, 10-14 m, 1 lv (SG); Lagoa Azul, NW Sào Tomé,

15 m, 1 lv (SG)

Principe. Bahia das Agulhas, 01°36'06" N, 07°20'55"

E, 15 m, 1 lv (SG) (Figs 9-10); Praia Evora, San

Antonio, 01°38’24" N, 07°26'34" E, 7 m, on muddy

sand, 1 dd (SG).

DISCUSSION

Keen (1944: 56) regarded Murex labiatus as a

possible species dubium (sic) because Bellardi (1873)

had earlier placed Murex labiatus in the synonymy of

M. fistulosus Brocchi, 1814. Her decision was tentative

because she had not seen a copy of their work and later

Gertman (1969: 156) considered it was best to follow

Keen's suggestion and consider Murex labiatus as a

nomen dubium. However, the lectotype of Murex

labiatus was figured afterwards by Pinna ( 1971 : pl. 76,

fig. 12) and by Pinna & Spezia (1978: pl. 35, fig. 1).

Typhis labiatus was described from the Pliocène of

Castell'Arquato and is undoubtedly conspecific with the

Recent Typhinellus sowerbii. A Pliocène specimen

from Castell'Arquato is here illustrated for comparison

(Figs 23-24). Typhinellus labiatus was also commented

and illustrated by Houart (2001 ).

There are no stable différences observed between

the Mediterranean shells and the specimens collected in

Sào Tomé and Principe. The specimens from West

Africa are larger, occasionally almost twice as big as the

Mediterranean shell but also with at least one additional

teleoconch whorl. The spire looks also to be lower but

we could not yet examine a specimen from Sao Tomé

or Principe with an intact apex, ail of them having the

first whorls eroded, so that :

1. The spire could be as high than in T. labiatus from

the Mediterranean if we add the protoconch and

teleoconch whorls 1 and 2.

2. We don't know exactly the number of teleoconch

whorls in the specimens from Sâo Tomé and Principe,

although the larger ones probably has 6 or 6.5

teleoconch whorls vs 4.5 or 5.5 in the examined

Mediterranean specimens; as a reminder also, one of

the syntypes of Typhis sowerbii is a large specimen of

24.5 mm (Fig. 2).

The axial sculpture consists of 4 rounded varices

with a shaip lamellae, ending with a short, strongly

recurved spine at shoulder. The anal tube is strongly

backward recurved, forming an angle of approximately

65-85° with the axis of the shell. The last anal tube

(when intact) is long and hollow, the others are

gradually shorter and closed. They are strongly

adpressed to the preceding partition. The spiral

sculpture consists of low cords, more apparent on the

R. Houart, S. Gori & P. Ryall

Novapex 12(3-4): 91-97, 10 octobre 2011

3 4

Figures 1-4 - Typhinellus labiatus (Cristofori & Jan, 1832)

1. Nomenclature. Livomo, Italy, 13.6 mm, (RH).

2. Syntype of Typhis sowerbii Broderip, 1833, Mediterranean, 24.5 mm, BMNH 197461.

3-4. Protoconch. SEM J. Cillis, IRSNB, scale bars 1000 pm.

varices, ending as more or less recurved, short spines on

varices. There are 6 primary cords (Fig. 1), P1-P6,

occasionally shallow or almost obsolète in some

specimens, sometimes with one to three secondary

cords between P2 and P3 and/or P3 and P4, or P5 and

P6.

The protoconch has not yet been examined. The

protoconch of Mediterranean specimens consists ot 1.5-

1.75 whorls (Figs 3-4).

Other specimens of T. labiatus were reported lrom

the western Atlantic. Gertman (1969: pl. 1, fig 5a-5c)

illustrated a typical specimen of 17 mm from the

Leeward lslands, near Nevis and reported the species

also from Portobelo, Panama, from off Texas and from

Egmont Key, Florida. Other specimens from the

western Atlantic are occasionally larger and broader

(Figs 25-29) but like Gertman (1969: 156), we also feel

that these différences are within the range of variation

of T. labiatus.

A specimen collected in Principe by SG was eating

a small bivalve.

Fernandes & Rolân (1993: 38) reported they had

collected in Sào Tomé a species they recorded as

Typhis cf clarki Keen & Campbell, 1964. We suspect

these authors were referring to the species which is the

basis of this paper as one of the current authors (PR)

R. Houart, S. Gori & P. Ryall

New record of Typhinellus labiatus

has material from the late Francisco Fernandes herein

illustrated (fig. 11-14).

Houart (1997: 84-85, figs. 234, 235 ) illustrâtes a

specimen of Typhina expansa (Sowerby, 1873) as

Typhis (Talityphis) expansus from the geographically

close area of lie Banié, Gabon. However this species

lives in a different substrata of very fine and thick

sédiment washed down the nearby rivers. The animal

lives submerged in détr itus and only the tip of the canal

exits the surface (PR personal observation). In the

islands, however, T. labiatus live on silty or muddy

sand (SG). Typhina and T. expansa can be distinguished

from Typhinellus by having a varical Range broadly

expanded adapically, extending to midway of the

siphonal canal; by having a broader last teleoconch

whorl relative to the shell height, and by having the

anal tube near the preceeding varix, but not adpressed

as in Typhinellus.

The examination of the syntypes of T. belcheri proved

Typhina to be congeneric with Talityphis (Houart,

2002 ).

Acknowledgements

We are grateful to Emilio Rolân (Museo de Historia

Natural, Santiago de Compostela, Spain) and Jean-

Louis Delemarre (St. Nazaire, France) for checking the

list of species collected in Sao Tomé and Principe and

for their comments,

Sandro Gori also thanks Jean-Louis Testori, owner

of Club Maxel diving Center of Sao Tome and ail his

team, composed by Edmilson Augusto, Lucas Tabares,

Helder Brito, Apolo Pires, Esmael Cristovao, Eslander

Campos and Armando Pires, that with patience and skill

help him during the researches

REFERENCES

Bellardi, L. 1873.1 Molluschi dei terreni terziarii del

Piemonte e delle Liguria. Part 1 : Cephalopoda,

Pteropoda, Heteropoda, Gasteropoda (Muricidae e

Tritonidae). Memorie délia Reale Accademia delle

Scienze di Torino, ser. 2, vol. 27: 33-324.

Figures 6-29 - Typhinellus labiatus (Cristofori & Jan, 1832)

6-8. Sào Tomé, Lagoa Azul, 31.2 mm, (SG); 9-10. Bahia das Agulhas, Principe, 15 m, 24.1 mm, (SG);

11-14. OU Sao Tomé, 6 m, in line sand/silt, 21.9 mm, (PR); 15-17. Kerkennah, Tunisia, 20.1 mm, (RH);

18-20. Kerkennah, Tunisia, 15.2 mm, (RH); 21-22. Mallorca, 16.6 mm, (RH); 23-24. Castell'Arquato, Piacenza,

Italie, Pliocène, 16.8 mm, (RH); 25-26. Portobelo, East Panama, 60 m, 28.7 mm, (RH); 27-29. Portobelo, East

Panama, 60 m, 26.5 mm, (RH).

• •

R. Houart, S. Gori & P. Ryall

Novapex 12(3-4): 91-97, 10 octobre 2011

R. Houart, S. Gori & P. Ryall

New record of Typhinellus labiatus

Drewes, R.C. & Wilkinson, J. A. 2004. The California

Academy of Sciences Gulf of Guinea Expédition

(2001). I. The taxonomie Status of the Genus

Nesionilaxus Perret, 1976 (Anura: Hyperoliidae),

Treefrogs of Sao Tome and Principe, with

comments on the Genus Hyperolius. Proocedings of

the California Academv of Sciences 55(20): 395-

407.

Femandes, F. & Rolân, E. 1990. Nuevo genero y

nuevas especies de la familia Buccinidae

Rafinesque, 1815 (Mollusca, Neogastropoda) de la

Isla de Sao Tomé. Bollettino Malacologico 25(9-

12): 341-348.

Femandes, F. & Rolàn, E. 1993. Marine molluscs of

Sâo Tomé and Principe: bibliographie actualization

and new contributions. Iberus, 11(1): 31 -47.

Gertman, R.L. 1969. Cenozoic Typhinae (Mollusca:

Gastropoda) of the Western Atlantic région. Tulane

Stndies in Geology andPaleontology 7(4): 143-191.

Houart, R. 1997. Les Muricidae d'Afrique occidentale

II. Ocenebrinae, Ergalataxinae, Tripterotyphinae,

Typhinae, Trophoninae & Rapaninae. Apex 12(2-3):

44-91.

Houart, R. 2001. A review of the Recent Mediterranean

and Northeastem A tlantic species of Muricidae.

Evolver: 1-227 (5 May 2001).

Houart, R. 2002. Description of a new typhine

(Gastropoda: Muricidae) from New Caledonia

with comments on sonie generic classifications

within the subfamily. Venus 61(3-4): 147-159.

Houart, R. 2005. Description of a new species of

Muricopsis (Gastropoda: Muricidae:

Muricopsinae) frorn Sào Tome, West Africa.

Novapex 6(4): 119-122.

Houart, R. & Gori, S. 2008. Description of a new

Muricopsis species (Muricidae: Muricopsinae)

from Northwest Sào Tomé. Novapex 9(4): 149-

153.

Houart, R. & Rolân, E. 2001. A new Muricopsis

(Gastroposa, Muricidae) from Annobôn Island,

Eastern Atlantic. Novapex 2(2): 61-66.

Keen, A.M. 1944. Catalogue and révision of the

gastropod subfamily Typhinae. Journal of

Paleontology 18 (1): 50-72.

Merle D. 1999. La radiation des Muricidae

(Gastropoda: Neogastropoda) au Paléogène:

approche phylogénétique et évolutive. Paris.

Unpublished thesis. Muséum national d'Histoire

naturelle : i-vi, 499 pp.

Merle D. 2001. The spiral cords and the internai

denticles of the outer lip in the Muricidae:

terminology and methodological comments.

Novapex 2(3): 69-91.

Pinna, G. 1971.1 Tipi delle specie di Gasteropodi

terziari istituite de Giuseppe De Cristofori e Giorgio

Jan nel 1832 conservati nelle collezioni del Museo

Civico di Storia Naturale di Milano. Atti Società

Italiana Scienza Naturale et Museo Civico di Storia

Naturale di Milano 112: 421 -440.

Pinna, G. & Spezia, L. 1978. Catalogo dei Tipi del

Museo Civico di Storia Naturale di Milano V. I Tipi

dei gasteropodi fossili. Atti Società Italiana Scienza

Naturale et Museo Civico di Storia Naturale di

Milano 119(2): 125-180.

Rolân, E. & Femandes, F. 1991. Muricopsis

(Risomurex) (Gastropoda, Muricidae) de las islas de

Sao Tomé y Principe (Golfo de Guinea, Africa

Occidental). Apex 6(1-2): 11 -20.

Rolân, E. & Gori, S. 2007. A new species of Muricopsis

(Muricidae: Muricopsinae) from Sâo Tome Island.

Novapex 8(1): 23-26.

R. Houart, S. Gori & P. Ryall

Novapex 12(3-4): 91-97, 10 octobre 2011

Appendix

List of Muricidae (excluding Coralliophilinae) collected in Sâo Tome (ST) and Principe (P)

(*) = endemic

Bolinus cornutus (Linnaeus, 1758) (ST)

Fernandes & Rolân, 1993

Hexaplexrosarium (Rôding, 1798) (ST & P)

JLD. PR, SG Fernandes & Rolân, 1993

Favartia (Favartia) emersoni (Radwin & D'Attilio, 1976 (ST)

SG new record

Favartia (Murexiella) bojadorensis (Locard, 1897) (ST)

SG new record

Homalocantha melanamathos (Gmelin, 1791) (ST & P)

SG new record

Muricopsis delemarrei Houart, 2005 (ST & P) (*)

JLD, PR, RH, SG

Muricopsis hernandezi Rolân & Gori, 2007 (ST) (*)

PR, RH, SG

Muricopsis matildeae Rolân & Fernandes, 1991 (ST) (*)

JLD, PR, RH, SG Fernandes & Rolân, 1993

Muricopsis principensis Rolân & Fernandes, 1991 (P)(*)

Muricopsis rutilus mariangelae Rolân & Fernandes, 1991 (ST & P) (*)

JLD, PR, RH, SG Fernandes & Rolân, 1993

Muricopsis testorii Houart & Gori, 2008 (ST) (*)

PR, RH, SG

Pradoxa confirmata Fernandes & Rolân, 1989 (ST & P) (*)

JLD, PR, RH, SG Fernandes & Rolân, 1993

Pradoxa thomensis Fernandes & Rolân, 1989 (ST& P) (*)

PR, RH, SG Fernandes & Rolân, 1993

lnermicosta inermicosta ( Vokes, 1871 ) (ST & P)

JLD, Fernandes & Rolân, 1993

Monda nodulosa (C.B. Adams, 1845) (ST & P)

JLD, PR, SG, Fernandes & Rolân, 1993

Trachypolia turricula (Maltzan, 1884) (ST & P)

PR (maybe reported in Fernandes & Rolân,

1993 as Ocinebrina suga )

Stramonita haemastoma (Linnaeus, 1767) (ST & P)

JLD, Fernandes & Rolân, 1993

Thais nodosa (Linnaeus, 1767) (ST & P)

JLD, Fernandes & Rolân, 1993

Typhinellus labiatus (Cristofori & Jan, 1832) (ST & P)

PR, SG (maybe reported in Fernandes & Rolân,

1993 as Typhis cf clarcki)

A few species mentioned by Fernandes & Rolân (1993) are questionable or/and actually junior synonyms. They

were mostly quoted from older records:

Murex turbinatus Lamarck, 1822. Is a synonym of Hexaplex duplex (Rôding, 1798)

Murex hoplites Fischer, 1876. Is a synonym of Hexaplex duplex (Rôding, 1798)

Murex tumulosus Sowerby. 1841. Is a synonym of Bolinus cornutus (Linnaeus, 1758) (see above)

Muricopsis blainvillei Payraudeau, 1826. Is a synonym of Muricopsis cristatus (Brocchi, 1814)

As mentioned by Fernandes & Rolân (1993: 45 [C9]): "we hâve not found this species and the record is

probably due to the confusion with some of the recently describcd species and mentioned in the list", this makes

référencé to Muricopsis mariangelae and M. matildae (E. Rolân in litt.).

Ocinebrina suga (Fischer-Piette, 1942). See above under Trachypollia turricula

Typhis cf. clarki Keen & Campbell, 1964. See above under Typhinellus labiatus

Thais ascensionis (Quoy & Gaimard, 1832). Is a synonym of Thais nodosa (see above)

Thais neritoidea (Linné. 1767). Is a synonym ot Thais nodosa (see above)

E. F. Garcia

Novapex 12(3-4): 99-107, 10 octobre 2011

Two new species of Epitonium (Gastropoda: Epitoniidae)

from the western Atlantic

Emilio F. GARCIA

115 Oak Crest Dr.

Lafayette, LA 70503

Efg21 12@louisiana.edu

KEYWORDS. Gulf of Mexico, Florida, Puerto Rico, taxonomy, Epitoniidae, Epitonium n. spp.

ABSTRACT. Two new Epitonium species from eastem Florida, the Gulf of Mexico and Puerto

Rico are described and compared with similar congeners. Spécial attention is given to E.

championi Clench & Turner, 1952, the most similar to the two proposed new taxa.

INTRODUCTION

In June, 2005, faculty members and graduate

student in the Biology Department at the University of

Louisiana, Lafayette went on a research expédition to

Bahia de Campeche. Southern Gulf of Mexico, on

board the R/V Pélican, a research vessel operated by

the Louisiana Universities Marine Consortium

(LUMCON). Results of this expédition hâve been

reported elsewhere (Garcia, 2006, 2007, 2008a,

2008b, 2008c). This area of the Gulf, located in the

Southwest quadrant, lias been poorly sampled. The

single campaign in 2005 increased the known

diversity of the area from 575 to 674, that is by 17%

(Rosenberg et al., 2009: 584).

Among the interesting material dredged in 120

hauls in Bahia de Campeche, there was an epitoniid

species, collected at only two consecutive stations.

Although there hâve been 56 species of Epitoniidae

recorded from the Gulf of Mexico (Rosenberg et al,

2009: 583), the Campeche specimens defied

identification. The species looked similar to

Epitonium championi Clench & Turner, 1952, a taxon

that had been erroneously reported from the Gull of

Mexico (Garcia & Lee, 2002:11; Rosenberg et al.,

2010: 641); however, upon close inspection of the

holotype of E. championi and other type material,

important différences came to light. Moreover, after

studying other epitoniids from the Gulf and elsewhere,

rnostly from the Harry G. Lee collection, a second

undescribed species was revealed that had been

hitherto identified as E. championi and E. turritellula.

This was a very surprising fmd, as the species inhabits

a large area, at least from Texas and Louisiana to

Sanibel Island, northeast Florida and Puerto Rico.

This study proposes two new Epitonium taxa and

scrutinizes them against their most similar congener,

E. championi , as well as other less- likely congeneric

taxa.

Abbreviations

BMSM: Bailey- Matthews Shell Muséum, Sanibel,

Florida, USA.

EFG: author’s collection.

HGL: Harry G. Lee collection, Jacksonville, Florida,

USA.

HMNS: Houston Muséum of Natural Science,

Houston, Texas, USA.

LACM: Los Angeles County Natural History

Muséum, Los Angeles, California, USA.

MCZ: Muséum of Comparative Zoology, Cambridge,

Massachusetts, USA.

SBMNH: Santa Barbara Muséum of Natural History,

Santa Barbara, California, USA.

UNAM: Universidad Nacional Autônoma de México,

Ciudad México,México.

USNM: United States National Muséum, Washington,

D.C., USA.

spec: specimen(s).

SYSTEMATICS

Family EPITONIIDAE S. S. Berry, 1910

Genus Epitonium Rôding, 1798

Type species: Turbo scalaris Linnaeus, 1758 by

subséquent désignation by Suter (1913).

Epitonium championi Clench & Turner, 1952

Figs 1-12, Table I

Material examined. Holotype MCZ 182900 length

11.5 mm, width 4.6 mm, Massachusetts, Cape Cod,

Hyannis, Lewis Bay (Figs 1-5); 3 paratypes, MCZ

162585 Massachusetts, Martha’s Vineyard, Gay Head

(see Figs 6 and 7) ; 6 spec.; Emerald I., North

Carolina, 34°40T’N 77°0’49”W (HMNS 47485);

Stone Harbor, New Jersey, 39°02’8”N 74°46’03”W

(HMNS 47484) (Fig. 9), 1 spec.; Florida, Duval

County, off Big Talbot I., 10-20 m (see Figs 10 to 12)

(HGL); 2 spec., Florida, Duval County, S.Jacksonville

Beach (see Fig. 8) (HGL); 1 spec., Florida, Duval

County, Fort George(HGL).

Distribution. South coast of Cape Cod, Massachusetts

to Duval County, NE Florida.

Original description. “ Shell reaching about 14 m

(1/2 inch in length, attenuate, imperforate, rather solid

E. F. GarcIa

Two new species of Epitonium

and strongly sculptured. Whorls 10 to 11, convex and

attached. Color a liât white to a light cream. Aperture

subcircular, with both the palatal and pariétal margins

thickened. The palatal or outer lip being greatly

thickened in older specimens. Columella short and

arched. Spire extended and produced at an angle of

20°. Suture moderately impressed. Axial sculpture

consisting of 8 or 9 flattened cord-like, slightly

impressed costae which are rather variable as to width.

Spiral sculpture consisting of 19 to 20 flattened ridges,

those nearest the umbilical area being a little narrower.

Basal ridge absent. Operculum thin, paucispiral and

brown in color. Nuclear whorls 2 1/2 to 3,smooth and

opaque” (Clench and Turner, 1952:318).

Discussion. In their original description Clench and

Turner (1952:318) described E. championi as having

flattened, cord-like axial costae and flattened spiral

ridges. Later on, in their “Remarks,” they elaborate on

this distinctive sculpture, stating that it “appears much

like a basket weave in which the upright and outer

struts (the axial costae) arc woven tightly, causing the

horizontal weave (the spiral ridges)to bulge outwardly

between the struts” (p. 319). This peculiar character of

the species is clearly seen on Fig. 4, a close-up of the

holotype, and Fig. 12, a specimen dredged off Duval

County, Florida.

The authors subsequently State that E. championi

“is perhaps a divergent element of E. candeanum, in

which both axial and spiral sculpture hâve had an

excess of development” (p. 319); however, E.

candeanum has blade-like axial costae and a

microsculpture of axial and spiral threads. Clench &

Turner did not observe the presence of a

microsculpture in the type material of E. championi,

nor did 1 on examining them. This character was also

lacking from ail the specimens examined, including

those from northeast Florida (see Figs 4 & 12). The

juxtaposition of E. candeanum and E. championi, i.e.,

blade-like vs. cord-like axial costae, has lead to the

misidentification of the species described herein as E.

leali, which does show the cord-like (although not

very flattened) costae of E. championi but also a

microsculpture between the axial éléments lacking in

the latter. Other différences will be shown in the

discussion of the former.

The maximum reported size of E. championi is

13.7 mm (Rosenberg, 2009); however, I hâve

examined a specimen from the Houston Muséum of

Natural Science ( No. 47484) that measures 17mm

(Fig. 9).

Although Dr. Lee’s collection shows that E.

championi is well-represented in NE Florida, the

specimens of Epitonium ”championi ” from the Gulf of

Mexico that I hâve examined, as weli as the specimen

shown in the Encyclopedia of Texas Seashells

(Tunnell et al, 2010:190) are misidentifications.

Epitonium leali n. sp.

Figs 13-26, Table II

Epitonium championi Clench & Turner, 1952- Garcia

& Lee, 2002:11.

Epitonium championi Clench & Turner, 1952- Lee,

2009: 95, fig. 454.

Epitonium championi Clench & Turner, 1952-

Rosenberg et al., 2010:641.

Epitonium tumtellula (Môrch, 1875)- Tunnell et al.,

2010: 193.

Type material. Florida, Gulf County, St. Joe Bay,

Palm Point, 29°48’29”"N 85°17’52”W. Holotype

USNM 1 150471 length 9.7 mm, width 3.6 mm (Figs

13-17) ;1 paratype SBMNH 149690 (Fig. 18); 1

paratype BMSM 17956 1 paratype EFG 29966; 7

paratypes HGL ail from the type locality.

Type locality. Florida, Gulf County, St. Joe Bay,

Palm Point, 29°48’29”"N 85°17’52”W.

Other material examined. Puerto Rico: San Juan,

Tsla Verde, 18° 25’ 39” N, 66° 0’ 36” W (Figs 24-

26)(1 spec., HGL). Florida: Duval County, off Big

Talbot I., 10-20 m (7 spec. HGL)(see Figs 19 and 20) ;

Lee County, Sanibel lsland ,26 o 27’N 82 o 0r W (3

spec.; BMSM 5687, BMSM 25698, BMSM

25708)(see Fig. 21); west end of St, Vincent I., 29.66°

N 85.13° W (HMNS 47413). Louisiana: Terrebonne

Parish, Isles Derniers, 29°3’41.44”N, 90°57’1.5r’W

(1 spec., HGL; 2 spec. EFG I0350)(see Fig. 23).

Texas: Heald Bank 29°12.5’N, 92°10.8’W, 10-13 m

(1 spec., EFG 12861); Nueces Co, Port Aransas, 27°

50’ 1” N, 97° 3’ 39” W (2 spec., HGL)(see Fig. 22);

San Luis Pass, Galveston, 29°5’2”N 95°7’12”W

(HMNS 39743);

Distribution. Puerto Rico, east Florida, west Florida,

Louisiana, Texas.

Figures 1-12. Epitonium) championi Clench & Turner

1-5. Massachusetts, Cape Cod, Hayannis, Lewis Bay. Holotype MSZ 182900 length 11.5 mm,width 4.6 mm. 6.

Massachusetts, Martha’sVineyard, off Gay head. Paratype, 5.9 mm. 7. Massachusetts, Martha’sVineyard, off '

Gay head. Paratype, 9.2 mm 8. Florida, Duval County, S.Jacksonville Beach, 4.6 mm (HGL) 9 New Jersev

Stone Harbor, 39°02’8”N 74°46’03”W, 17 mm (HMNS 47484), 10. Florida, Duval Co., off Big Talbot lsland

10-20 m, 8.4 mm (HGL). 11-12. Florida, Duval Co., off Big Talbot lsland, 10-20 m, 7.7mm (HGL)

100

E. F. Garcia

Novapex 12(3-4): 99-107, 10 octobre 2011

101

E. F. Garcia

Two new species of Epitonium

Description. Holotype (Figs 13-17) 11.5 mm in

length, attenuate (width/ length ratio 0.37).

Protoconch damaged, remaining whorl smooth, white.

Teleoconch of 8 moderately convex, joined whorls.

Suture relatively deep. Axial ornamentation on early

whorls of 16 or 17 narrow, well- defined, rounded

costae, some becoming varicoid starting on fourth

whorl; at least one varix per whorl after fourth; costae

diminishing in number on later whorls, 10 on Iast

whorl; posterior terminal embedding into suture,

joining earlier whorl (Fig. 15); axial interspaces

approximately 3 to 4 times as wide as costae, except

where varical formations occur; 4 strong varices on

last whorl. Spiral sculpture of 14 or 15 cords between

sutures; cords uneven in strength, narrower than

interspaces (Fig. 16); interspaces ornamented with

numerous microscopie axial threads, which wrinkle

top of spiral cords as they cross over (Fig. 16); base of

last whorl convex, developing a shallow chink at

umbilical area (Fig. 17). Aperture elongate-ovate,

complété, with strong, wide labral varix, narrower on

pariétal side. Operculum unknown. Shell pale pinkish-

tan with white axial costae.

Discussion. There is very little variation in ail the

material studied, other than number and position of

varices and slight variation in number of axial costae.

Compare holotype Figs 13, 15 and 16 with Figs 24, 25

and 26, the specimen from Puerto Rico; also, see

Table II. Some specimens from deeper water hâve a

tendency to develop a more capacious last whorl,

which also causes them to hâve a stronger umbilical

dépréssion (Figs 19 and 20). Fresh specimens, such as

those from the type locality and the specimen from

Puerto Rico (Figs 24 to 26), hâve a pale pinkish- tan

shell with white axial costae; ail other specimens are

white.

This new species has been confused with

Epitonium championi, from which it differs by having

more rounded axial costae which ernbed into a deeper

suture (Compare Fig. 3 with Figs 15 and 26), in

having narrower spiral cords that do not “bulge”, in

having a secondary microsculpture in the axial

interspaces (Compare Figs 4 and 12 with Figs 16 and

25), and in generally having a narrower shell (width/

length ratios 0.36 vs o.40). Although some specimens

of E. leali do approach the same ratio as E. championi,

this is due to the strong development of the last whorl.

Epitonium leali may also be confused with

Epitonium pigrum n. sp. The différences between the

two taxa will be treated in the discussion of the latter.

Epitonium tiburonense Clench & Turner, 1952 is

superfïcially similar to E. leali, but the former is

smaller; the holotype having 9 whorls at only 6.8 mm

in length. Moreover, E. tiburonense has evenly

distributed non- varicoid axial costae which “form a

thickened pad” (Clench & Turner, 1952: 305) in the

umbilical area, and lacks the secondary microsculpture

of E. leali.

Although Epitonium turritellula Morch, 1874 has

been confused with the new species (Tunnell et al.,

2010: 93), the former only grows to 6.5 mm

(Rosenberg, 2009), producing 10 whorls at only 6.4

mm in length (Clench & Turner, 1952: 298). Also, E.

turritellula has blade- like, more numerous costae

(about 20 on last whorl) that tend to “peak” at the

suture, giving the appearance of narrowly coronated

whorls.

Although ail of the specimens of Epitonium leali

in this study were collected empty, it seems that the

species inhabits very shallow water in the Gulf of

Mexico, as most specimens hâve been collected at

beaches from southwestem Florida to Texas.

Etymology. Named for Dr. José H. Leal, Director of

the Bailey- Matthews Shell Muséum, Sanibel, Florida

and Editor of the prestigious malacological journal

The Nautilus. This taxon honors him for his

accomplishments in the field and for his willingness to

help both personally and in his capacity as the

Director of the Bailey-Matthews Shell Muséum.

Epitonium pigrum n. sp.

Figs 27- 40, Table III

Type material. Mexico: Balu'a de Campeche,

20°51.49N, 92 Ü 21.44’W, in 63-65 m. Holotype

USNM1150470 length 11.5 mm, width 4.3 mm; 1

paratype MCZ 373765 (Fig. 32); 1 paratype UNAM; 1

paratype EFG 26207 (Figs 33-34) 1 paratype BMSM

17957 (Fig. 35); 1 paratype SBMNH 149689 (Fig 36);

1 paratype HGL (Fig. 37). Bahia de Campeche,

20 ü 52.40 N, 92 U 24.83’W, in 77-81 m; 1 paratype

LACM 3189 (Fig. 38), 1 paratypes EFG 26273 (Figs

39-40).

Figures 13-26. Epitonium leali n. sp.

13-17. Florida, Gulf County, St. Joe Bay. Palm Point, 29°48’29”"N 85° 17’52”W. Holotype USNM 1150471

length 9.7 mm, width 3.6 mm. 18. Florida, Gulf County, St. Joe Bay, Palm Point, 29°48’29”"N 85°17’52”W

Paratype SBMNH 149690 11mm. 19. Florida, Duval County, off Big Talbot F, 10-20 m, 14 mm (HGL) 20.

Florida, Duval County, off Big Talbot I., 10-20 m, 13.5 mm (HGL). 21. Florida, Lee County, Sanibel Island,

26°27’N 82°01’ W, 18.5 mm (BMSM 25708). 22. Texas, Nueces Co, Port Aransas, 27°50T" N, 97°3'9" W. 8.5

mm (HGL) 23. Louisiana, Terrebonne Parish, Isles Derniers, 29°3’41,44”N, 90°57’1.51”W, 11.5 mm (EFG

10350). 24- 26. Puerto Rico, San Juan, Isla Verde, 18°25’39" N, 66° 0' 36" W, 9 mm (HGL).

102

E. F. Garcia

NOVAPEX 12(3-4): 99-107, 10 octobre 2011

103

E. F. Garcia

Two new species of Epitonium

Type locality. Bahia de Campeche, 20°51.49’N,

92°21.44’W, in 63-65 m.

Other material examined. Louisiana: 28°05’N,

90°59’W, in 114 m (HMNS 39750); 28° 37.920’N, 90°

36.550’W, in 22 m (EFG 23531).Texas: SSE of

Freeport, 28°15’N, 95°0’W (HMNS 39734); Heald

Bank, SSE of Galveston, 29°08’N, 94°01’W(HMNS

39735); S. of Galveston, 29 Ü 55’N, 94°41’W, in 20 m

(HMNS 39736); SE of Freeport, 28 l, 19’N, 94°29’W, in

50 m (HMNS 39737); off Galveston, 28°52’N,

94°42’W, in 22 m (HMNS 39738); off Galveston,

29°16’N, 94°29’W, in 16 m (HMNS 39740); off

Galveston, 28°21 ’N, 94°53’W (HMNS 39744); 30 mi.

off Port Isabel, in 22-28 m (HMNS 39745); off

Galveston, 28°18’N, 94°28’W, in 50 m (HMNS

39746); off Matagorda peninsula, 28°35’41”N,

95°58’59’W, in 15 m (HMNS 39747); Padre L, in 4 m

(HMNS 39748); off Padre I., in 22 m (HMNS 39749)

Distribution. Western Gulf of Mexico: Bahia de

Campeche, Mexico; Texas and Louisiana, USA

Description. Holotype (Figs 27-31 ) 11.5 mm in

length, imperforate, attenuate (width/ length ration

0.37). Protoconch conical, smooth, glassy, with a

brownish band on shoulder of whorls, of

approximately 4 whorls; transition to teleoconch

delineated by growth scar and change in opacity.

Teleoconch of 9.5 convex whorls; whorls rapidly

increasing in width. Suture relatively shallow, crossed

by axial ornamentation. Axial sculpture of

approximately 22 thin, erect costae on early whorls;

costae curving adaperturally, slightly flattening, as

they cross over suture, pasting on to the earlier whorl,

some forming inconspicuous “peaks’ (Fig. 29),

decreasing in number, becoming more cord- like on

later whorls; 17 costae on last whorl; varices forming

randomly starting at the end of fifth whorl; 3 varices

on last whorl; microscopie axial wrinkles showing on

interspaces between axial costae. Spiral sculpture of

primary cords of uneven strength; microscopie spiral

wrinkles appearing between them, creating pustulose

surface as they cross axial éléments (Fig. 30); spiral

ornamentation not Crossing over axial éléments;

approximately 12 primary cords on early whorls,

increasing in number on later whorls. Base of shell

solid; umbilical area covered by pariétal thickness

(Fig. 31). Aperture oval; pariétal and labral margins

thickened; labral rnargin becoming patulose anteriorly,

pariétal rnargin narrowing at posterior end. Shell

milky-white with slight satin luster.

Discussion. The main characters of the paratypes are

consistent with those of the holotype; there is some

expected variation in the number of axial costae (see

Table III), and in the number and placement of

varices.

Epitonium pigrum is most similar to E. leali n.sp.

They both hâve the same general proportions, a

number of varicoid axial costae and similar

microsculpture on interspaces. However, E. pigrum

lias more numerous axial costae on early whorls (22

vs. 15) and on the last whorl (15 vs. 10), a shallower

suture, and a solid umbilical area without a chink

(compare Figs 7 and 35). More importantly, the

production of the terminais of the axial costae of the

two species as they cross the suture are quite different:

E. pigrum produces flattened costae which curve

adaperturally and adhéré to the previous whorl; E.

leali maintains the rounded costae which embed into

the deeper suture and do not turn adaperturally

(compare Figs 15 and 26 with Figs 29 and 40).

Although both species inhabit the Coastal areas of

Louisiana and Texas, ffesh specimens of Epitonium

leali can be found on beaches while E. pigrum

inhabits offshore banks.

Epitonium championi is also similar to E. pigrum in

general shape, and the production of the terminais of

the axial varices is similar, particularly when the

specimens are not fresh (compare Fig. 3 with Figs 29

and 40) ; however, E. championi has fewer axial

costae on early whorls (16 vs. 22) and on the last

whorl (10 vs. 15), has more flattened axial costae,

wider spiral cords that “bulge” out, and lacks a

microsculpture (compare Figs 4 and 12 with Fig. 30).

Etymology. From the Latin pigrum (adjective

meaning lazy), referring to the relatively large number

of prominent varices, presumably periods of rest.

Figures 27-40. Epitonium pigrum n. sp.

27-37. Mexico, Bahia de Campeche, 20°51.49'N, 92°21.44'W, 63-65 m. 27-31. Holotype USNM 1150470 length

11.5 mm, width 4.3 mm. 32. Paratype MCZ 373765 33-34. Paratype BMSM 17957 35. Paratype SBMNH &

149689 36. Paratype LACM 3189 37 Paratype EFG 26207 38-40. Mexico, Bahia de Campeche 20°52 40'N

92°24.83'W, 77-81 m 38. Paratype HGL col. 39-40.Paratype EFG 26273.

104

E. F. Garcia

Novapex 12(3-4): 99-107, 10 octobre 2011

105

E. F. Garcia

Two new species of Epitonium

Acknowledgements

My spécial thanks to Dr. Harry G. Lee, of

Jacksonville Florida, for the loan of much of the

material used for this study, as well as for the donation

of the holotype and several paratypes of Epitonium

leali. The specimens of E. leali were eollected by the

late Barbara Barfield, Southport, Florida. I am very

grateful to Dr. José H. Leal, Director of the Bailey-

Matthews Shell Muséum and Editor of The Nautilus,

for giving me access to the type material of E.

championi and for the loan of specimens front the

Muséum; and very much appreciate the efforts of Tina

Petway, Lucy Clampit and Eydie Rojas for rnaking

available specimens housed at the Houston Muséum

of Natural Science. Part of the material for this study

is based upon work supported by the National Science

Foundation under Grant No. 0315995.

REFERENCES

Clench, W. J. and R. D. Turner. 1952. The généra

Epitonium (part II), Depressiscala, Cylindriscala,

Nystiella and Solutiscala in the Western Atlantic.

Johnsonia 2: 289-356

Garcia, E. F. 2006. Six new species of mollusks

(Gastropoda: Cerithioidea, Buccinoidea,

Muricoidea) front Bahia de Cantpeche,

southwestern Gulf of Mexico. Novapex 7(4): 77-

89.

Garcia, E. F. 2007. Report on ntollusks eollected in a

dredging expédition to Bahia de Campeche,

southwestern Gulf of Mexico. American

Conchologist 35(2)4-11.

Garcia, E. F. 2008a. Eight new molluscan species

(Gastropoda: Turridae) from the western Atlantic,

with the description of two new généra. Novapex

9(1): 1-15.

Garcia, E. F. 2008b. Eight new molluscan species

(Gastropoda: Turridae) front the western Atlantic,

with the description of two new généra. Novapex

9(1): 1-15.

Garcia, E. F. 2008c An extension of the genus

Spinosipella (Bivalvia: Verticordidae) in the Gulf

of Mexico. American Conchologist 36(3): 8-9.

Garcia, E. F. & H. G. Lee. 2002. Report on molluscan

species found in the offshore waters of Louisiana,

including many extensions of known range and un-

named species. American Conchologist 30(4): 10-

13.

Lee, H. G. 2009. Marine shells of northeast Florida.

Jacksonville Shell Club: Jacksonville. 204 pp

including numerous txt figures.

Rosenberg, G. 2009. Malacolog 4.1.1 : A Database of

Western Atlantic Marine Mollusca. [WWW

database (version 4.1.1 )] URL

http://www.malacolog.org/.

Rosenberg, G., F. Moretzsohn & E. Garcia. 2009.

Gastropoda (Mollusca) of the Gulf of Mexico. /«:

Gulf of Mexico: Ils Origins, Waters, and Biota.1,1.

Biodiversity. D. L. Felder & D. K. Camp, eds,

Texas A & M University Press, pp. 579-699.

Suter, H. 1913. Manual of the New Zealand Mollusca.

Wellington: Government Printer, pp. 1-1120.

Tunnell, J.W, J. Andrews, N. Barrera, & F.

Moretzshon. 2010. Encyclopedia o f Texas

Seashells. Texas A & M University Press: College

Town, 512 pp, including many color photos.

TABLES

Abbreviations: H: Holotype; P: Paratype; BT. Big Talbot L, NE Florida; Ft.G.: Fort Geroge, NE Florida; HB:

Healds Bank, Texas; ID: Isles Derniers, Louisiana; JX: Jacksonville Beach, NE Florida; NJ: Stone Harbor,

New Jersey; PA: Port Aransas, Texas; PR: Isla Verde, N. Puerto Rico; SI: Sanibel Island, W Florida; SJB: St.

Joe Bay, NW Florida; SL Pass: San Luis Pass, Galveston, Texas.

- Juvénile specimens were not included in the tables below.

TABLE I. Epitonium championi Clench & Turner, 1952

Specimen

Number of

whorls

Length in

Width in

W/L

ratio

Number of

early axial

costae

Number of axial

costae last

whorl

H Figs 1-5

11.5

4.6

0.40

P Fig, 7

9.2

3.6

0.39

7.5

3.8

0.38

BT Fig. 11

7.7

3.2

0.42

7.5

3.5

0.47

7.1

3.1

0.44

BT Fig. 10

8.4

3.5

0.42

7.5

8.8

3.8

0.43

Ft.G.

5.7

2.2

0.39

N,J. Fig. 9

0.29

Average

0.40

106

E. F. Garcia

Novapex 12(3-4): 99-107, 10 octobre 2011

TABLE II. Epitonium leali n. sp

Specimen

Number

whorls

Length in

Width in

W/1L

ratio

Number

of early

axial

costae

Number of axial

costae last whorl

H Figs 13-17

9.7

3.6

0.37

PSJB

9.9

3.5

0.35

PSJB Fig. 18

3.8

0.35

PSJB

10.7

4.2

0.39

PSJB

8.5

10.7

4.0

0.37

PSJB

9.4

3.6

0.38

PSJB

6.5

9.8

3.6

0.37

PSJB

8.5

3.2

0.38

PSJB

3.8

0.38

PSJB

9.0

3.3

0.37

PSJB

6.5

7.7

2.8

0.36

BT Fig. 19

8.5

14.0

5.3

0.38

BT Fig. 20

8.5

15.8

6.3

0.40

8.5

13.5

5.0

0.37

12.1

4.9

0.40

8.0

3.0

0.38

10.2

3.6

0.35

7.2

2.7

0.38

6.5

4.0

1.7

0.42

PA Fig. 22

7.5

8.5

3.5

0.41

7.5

7.4

2.7

0.37

ID Fig. 23

11.5

4.3

0.37

7.5

9.8

3.8

0.39

7.5

10.7

4.5

0.42

PR Figs 24-26

3.2

0,36

7.5

16.2

5.8

0.36

8.5

18.6

6.3

0.34

SI Fig. 21

8.5

18.5

6.3

0.34

SL Pass

9.2

3.6

0.39

Average

0.36

TABLE III. Epitoniumpigrum n. sp.

Specimen

Number

whorls

Length in

Width in

W/1L

ratio

Number

of early

axial

costae

Number of axial

costae last whorl

H Figs 27-31

9.5

11.5

4.3

0.37

P Fig. 32

0.36

9.5

12.6

4.3

0.34

P Figs 33-34

8.5

10.9

4.1

0.38

P Fig. 35

0.36

P Fig. 36

7.5

9.6

3.7

0.39

P Fig. 37

8.5

11.6

4.3

0.37

Average

0.36

107

«aW’PW^ . WHBEI B l

D. Massemin, S. Clavier & J.-P. Pointier

Novapex 12(3-4): 109-118, 10 octobre 2011

First record of Pisidium punctiferum (Guppy, 1867) and Eupera viridans

(Prime, 1865) (Mollusca: Sphaeriidae) from French Guiana

David MASSEMIN

Angle des rues Du Four et Des Remparts / 34380 Notre-Dame de Londres, France

yoda.massemin@hotmail.fr yoda.massemin@wanadoo.fr

Simon CLAVIER

HYDRECO, Laboratoire Environnement de Petit Saut / B.P 823 / 97388 Kourou cedex, Guyane,

France

simon.clavier@hydrecolab.com

Jean-Pierre POINTIER

USR 3278 CNRS-EPHE CRIOBE, Université de Perpignan / 66860 Perpignan cedex, France

pointier@univ-perp.fr

KEYWORDS. French Guiana, continental Bivalvia, Pisidium punctiferum, Eupera viridans,

identification key.

MOTS CLÉS. Guyane, bivalves continentaux, Pisidium punctiferum, Eupera viridans, clé

d’identification.

ABSTRACT. Two freshwater Bivalvia species identified as Pisidium punctiferum (Guppy, 1867)

and Eupera viridans (Prime, 1865) were collected at severai places from French Guiana. Those

new records bring the number of freshwater-molluscs species from this French overseas territory

to 27 and extend the distribution area of the mentioned species to the Guiana Shield. An

identification key to continental Bivalvia from French Guiana is theretore provided.

RESUME. A l'occasion d'une étude hydrobiologique, Pisidium punctiferum (Guppy, 1867) et

Eupera viridans (Prime, 1865) ont été collectés en Guyane. Ces deux bivalves, nouveaux pour

l’inventaire, portent à 27 le nombre de mollusques dulcicoles connus de ce territoire ; leur aire de

distribution naturelle est étendue en direction du Plateau des Guyanes. Une clé d identification des

bivalves continentaux de Guyane est proposée.

INTRODUCTION

The Guiana Shield is an old Precambrian

geological formation of more than 1,600 km long,

located in the north-eastern part of South America,

including Guyana, Suriname, French Guiana and a

part of Brazil (Amazonas, Roraima, Para and Amapâ

States, west of the Amazona River). French Guiana

(2°- 6° North latitude) stretches over almost 84,000

square kilométrés (Figure 1); it is a hot spot of

biodiversity because 80% of the territory is covered by

a tropical rain forest. considered as one of the 15 last

worldwide remaining clumps only partially affected

by human activities (Gargominy, 2003 & Hammond,

2005). An abundant and complex hydrographical

System feeds this territory where acids and poorly

mineralized waters are unfavourable habitats to

freshwater molluscs.

The difficulty to reach the upstream areas explains

why this group was so poorly studied despite of some

conséquent but old works (Drouet, 1859 & Tillier,

1980), until the recent publication of a molluscs

identification guide (Massemin et al., 2009) in which

24 species of freshwater molluscs are listed.

Nevertheless, thanks to hydrobiological investigations

carried out in French Guiana under the auspices of the

Water Framework Directive and EDF, one species

was recently added to that list (Clavier et al, 2010).

The aim of the présent article is to report for the First

finie the presence of two freshwater Bivalvia species

in French Guiana (Figure 2). This discovery carries

the number of freshwater molluscs species known

from French Guiana to 27 (Gastropoda & Bivalvia),

belonging to 7 families; it gives the opportunity to the

authors to présent a general illustrated identification

key of the known species of continental Bivalvia

inhabiting this territory (Figure 3; appendix 1 ).

Material and Methods

In order to assess the benthic French Guiana

macroinvertebrate fauna [following the définition of

Rosenberg and Resh (1993), the term “benthic” nieans

“bottom-living” and the prefix “macro” indicates that

109

D. Massemin, S. Clavier & J.-P. Pointier

Pisidium punctiferum and Eupera viridans from French Guiana

Figure 1. Geographical localization of French Guiana and mentioned localities from French Guiana.

these organisais are retained by mesh size of 200-500

pm], an important hydrobiological study was carried

out by HYDRECO laboratory during the dry season

2009 (from September to December), under the

auspices of the local représentation of the French

Ministry in charge of environmental questions

(DIREN Guyane).

Samplings were conducted by one of us (SC) on the

entire territory in different habitats (large rivers, mid-

order streams, small streams named locally “criques”

and réservoir lakes) using a D-framed hand net (200

pm mesh size). Samples were collected by sweeping

the net along the banks. The first centimètres of the

substratum (organic and/or inorganic) were stirred to

dislodge attached or buried macroinvertebrates.

RESULTS

A total of 125 live specimens of small freshwater

Bivalvia peaclams species belonging to the

Sphaeriidae Family and to two different généra, were

collected in many watersheds (Figure 1, tables 1 & 2)

and preserved in éthanol 70°. Registered specimens

were deposited both in public (EPHE, Peipignan,

France) and private collections (HYDRECO, Kourou,

French Guiana). The material collected included a

total of 81 Pisidium punctiferum (Guppy, 1867) and

44 Eupera viridans (Prime, 1865).

Identilications of the species were made according

to spécifie conchologically characters (Kuiper, 1983)

and original descriptions of Guppy (1867) for P.

punctiferum and Prime (1865) for E. viridans.

Diagnose of the striate peaclam P. punctiferum

was established according to the following criteria: 1/

The average size when adult is around 3 mm in length

and 2.5 mm in height; 2/ The shell is oval and thin, the

anterior valve is shorter and angulated in its posterior

part; 3/ The test is diaphanous, finely striate

concentrically and covered with numerous granular

points, which are finer and more crowded on the

110

D. Massemin, S. Clavier & J.-P. Pointier

Novapex 12(3-4): 109-118, 10 octobre 2011

Figure 2. A. Eupera viridans (Prime, 1865), Crique Leblond, Sinnamary 4.1 mm; B. Eupera viridans Crique

Leblond, Sinnamary 3.8 mm; C. Pisidiumpunctiferum (Guppy, 1867), Crique Arataï, Régina 2.9 mm,

C. Pisidium punctiferum Saut Fracas 3.1 mm.

umbones, where the concentric striation is less

évident; 4/ A short periostracum may be présent and

5/ The hinge is well-developed with latéral teeth in

both valves (1-1/1-1 ) and small cardinal teeth (2/2).

Diagnose of the mottled fmgernail clam Eupera

viridans, was established according to the following

criteria: 1/ The size can be of about 8 mm when adult

(Pointier, 2008); 2/ The shell is oblong, compressed

and inequilateral, the anterior side is shorter and

rounded, the posterior one is sub abrupt and the basal

margin is slightly rounded; 3/ The test lias very

regular and délicate striation, a cream colour with

irregular dark brown spots, 4/ There is no mention of

periostracum and 5/ It présents a sub central umbo and

a cardinal tooth on each valve (Pointier, 2008).

discussion

The small freshwater mussels belonging to the

Sphaeriidae Family are distributed ail over the world

and the main généra, Pisidium , is cosmopolitan

(Kuiper, 1983). Pisidium punctiferum was described

from the Caribbean Island of Trinidad in 1867, under

the name Cyclas punctifera ; the type locality is a pond

at Saint Ann's River, near Port of Spain (10°39'00" N /

61°31’01" W): specimens were collected by the

government botanist Prestoe and by Guppy [The first

example was found by Mr. Prestoe, the colonial

botanist, when we were examining the weeds in a

pond at St. Ann, near Port of Spain, for Mollusca ].

Authors took a spécial care to compare the

collected specimens to the peaclam Pisidium

sterkianum Pilsbry, 1897 because this species is

widely distributed from the Guiana Shield [from

Venezuela (specimens collected by one of us, JPP, in

1987 and identified by Kuiper) and Brazil (Agudo-

Padrôn, 2009; Mansur & Pereira, 2006)] to Argentina

(Darrigran & Pastorino, 1995; Ituarte, 2007),

including the Pemvian Amazon (Ituarte, 2004),

Paraguay and Uruguay (Scarabino & Mansur, 2007).

Pisidium sterkianum occupies the same type of

D. Massemin, S. Clavier & J.-P. Pointier

Pisidium punctiferum and Eupera viridans from French Guiana

Figure 3. A. Ostreidae Crassostrea rhizophorae (Guilding, 1828), specimen collected on rocks, Montsinéry

French Guiana, 93 mm, B. Ostreidae Crassostrea rhizophorae (Guilding, 1828), specimen collected on a root in

a mangrove, Montsinéry French Guiana, 78 mm; C. Dreissenidae Mytilopsis leucophaeata (Conrad, 1831),

Montsinéry French Guiana, 20 mm; D. Corbiculidae Polymesoda ch aequilatera (Deshayes, 1855). specimen

without periostracum, Montsinéry French Guyana, 24 mm; E. Corbiculidae Polymesoda cf. aequilatera

(Deshayes, 1855), specimen with periostracum, Montsinéry French Guyana, 17 mm; F. Hyriidae Castalia

sulcata (Krauss, 1849), Tampok French Guiana, 47 mm; G. Hyriidae Diplodon granosus (Bruguière. 1792),

French Guiana, 19 mm; H: Hyriidae Diplodon voltzi Vemhout, 1914, MNHN, Litany French Guiana, 64 mm;

H. Hyriidae Diplodon granosus (Bruguière, 1792), Holotype MNHN, Cayenne French Guiana, 38 mm.

habitats than P. punctiferum , i.e. small streams, lakes,

etc. Consequently, P. sterkianum could probably be

présent in the study area. This species may be

distinguished from P. punctiferum according to the

following characters (Pilsbry, 1897): 1. The average

size when adult is around 6 mm in length and 5 mm in

height; 2. The shell is équilatéral with the anterior end

broadly truncated and beaks full but rather small; 3.

The test, glossy, light yellowish outside and greyish-

white inside, is very finely striated concentrically

without granular points; 4. There is no mention in

literature on the presence of periostracum and 5/ The

cardinal teeth are parailel and the latéral tooth is lower

and longer in the left valve.

Pisidium punctiferum is mentioned in literature

(Figure 4) from the United States of America

[Kentucky, Virginia, Illinois, Texas and northeast

Florida (Heard, 1979; Turgeon et al., 1998)] to

Southern Brazil [Rio Grande do Sul State (Agudo-

Padrôn, 2009; Martello et al., 2006; Simone, 2006)]

and Uruguay (Henry, 1897), including the West Indies

[Dominica (Starmühlner, 1988), Guadeloupe (Pointier

1974; 1976; 2008), Martinique (Guyard & Pointier,

1979; Pointier 2008), Saint-Lucia (Jordan. 1985 ; Mc

Killop & Harrison, 1981; 1982) and Saint-Vincent

(Harrison & Rankin, 1978)].

The species is mentioned in literature as living in

Cuba (Starmühlner, 1988). It is probably due to

misidenti lication, because even after recent

investigations it has not been found there (Pointier et

al., 2005). It is also signalled by the same author as

D. Massemin, S. Clavier & J.-P. Pointier

Novapex 12(3-4): 109-118, 10 octobre 2011

Figure 4. Distribution area of Pisidium punctiferum.

inhabiting Mexico, Panama, Jamaica, Venezuela and

Paraguay but without any précisé mention nor author.

The présent discovery expands the distribution of P.

punctiferum to French Guiana and, consequently, to

the Guiana Shield. Otherwise, the presence in

neighbouring Suriname, where the species is not yet

mentioned, is highly probable because 20 specimens

were found in the Maroni River, frontier between

Suriname to the west and French Guiana to the east

[Apsik Icholi (05°09'06" N / 54 o 20’21" W), Crique

Apatou (03°48’20" N / 54°08'36" W) and Papaïchton

(3°05'34" N / 52°20'28" W)]. So, if the species has

already been recorded in southem Brazil, its

distribution area is potentially also extended to the

northeast Brazil, because one piece was collected in

the Oyapock River, frontier between French Guiana to

the west and the Amapa State to the east.

In literature, most authors mentioned to hâve

observed P. punctiferum in lentic habitat: on the Saint

Vincent Island, Harrison & Rankin (1978) found the

maximum population densities in swamps; In

Dominica, it was observed in exposed sunlit ponds

(Starmühlner, 1988); In Guadeloupe, the species is

inhabiting swamps, canals and flooded meadows

(Pointier, 1974, 1976). In French Guiana, P.

punctiferum has a large distribution area ail over the

territory, from east to west, near the coastal line

[Crique Apatou (03°48'20" N / 54°08’36'' W) and

Crique Eau Claire (05°08'45" N / 52° 52'09" W)] to

the head watershed [Apsik Icholi (02°59'10" N /

54°10'53" W)]. The species has a large ecological

plasticity because it occupies a numerous variety of

habitats as it was recorded from small streams with

clear water (Crique Apatou and Crique Eau Claire) to

muddy large rivers [Papaïchton (3°05'34" N /

52°20'28" W)]. The site where the most important

number of specimens was collected (33) is a natural

rocky ri file environment without signifïcant entropie

pressures, [Saut Dalles, Sinnamary Watershed

(04°33'21 " N / 52°54'03" W)]. A single individual was

also found into the réservoir lake of Petit-Saut [Vata

(04°51'58" N / 52°57'4r' W)]. The species was mainly

found in water without entropie pressure. It would be

interesting to focus on the sampling effort in these

habitats in the rainy season, when the freshwater level

grows, creating a lot of standing water habitats.

Eupera viridans was described from the Caribbean

Island of Guadeloupe in 1865, under the name of

Sphaerium viridanfe by Prime (1865). The type

locality is the city of Pointe-à-Pitre (16°14'00" N /

61°31'00" W). Little is known about its natural

distribution (Figure 5). The species is only mentioned

in literature from the West-Indian Islands of

Guadeloupe (Pointier, 2008), Martinique (Delannoye,

Personal communication, 2010), Saint-Lucia (Pointier,

2001) and Saint-Martin (Coonans, 1967). Eupera

cubensis (Prime, 1865) is probably a synonym of E.

viridans. Consequently, the distribution of the species

113

D. Massemin, S. Clavier & J.-P. Pointier

Pisidium punctiferum and Eupera viridans from French Guiana

Figure 5. Distribution area of Eupera viridans.

could be enlarged to Cuba, Costa Rica (Volkmer-

Ribeiro & Machado, 2009), Texas and Florida (Heard,

1979) and to the Upper Mississippi River Basin too

(Illinois), but as an introduced species (Sneen et al.,

2009). The discovery of E. viridans in French Guiana

would be the first record of this species for the South

American continent. However, this information must

be taken with caution, because the systematic of this

genus remains confusing and misidentifïcations may

occur in the literature.

In the Lesser Antilles, Eupera viridans is common

in marshes, canals and ponds, attached to the aquatic

végétation or to wood débris (Pointier, 2008). In

French Guiana, the 44 observed specimens were

collected in a single spot at Leblond [Sinnamary

watershed (04°46'37" N / 53°07’08" W)], an altered

stream under entropie pressures such as illégal gold-

mining or domestic sewages (Comité de Bassin de

Guyane, 2006); this Bivalvia appears like a not

sensitive taxa in terms of bio indication. Punctually

distributed and first mentioned on the South America

continent, the species lias a patrimonial value for

French Guiana.

In conclusion, we hâve to mention that the

détection

of minute clams is difficult in the coloured and

charged in wood débris of the Guiana freshwaters.

Lack of studies on this group together with these

characteristics i.e. small size, difficult visual détection,

explains the late discovery ofrelatively abundanttaxa

on this territory. Sampling method is also an

explanatory factor. Without a standardized collect

protocol, knowledge on freshwater molluscs remains

uncompleted (Massemin et al., 2009). The use of a

hand net with very small mesh size (200 pm) would

be a good alternative to collect minute species. In

2010, three new species of freshwater molluscs (one

Gastropoda and two Bivalvia) from French Guiana

were found using this technique, increasing the

number of freshwater species mentioned from this

overseas department. Why couldn’t the discovery of

other species be expected in the future?

Acknowledgements

Simon Clavier is grateful to DIREN 973, to EDF

GUYANE for finding the sampling in the Sinnamary

watershed, and to the teams of HYDRECO and NT2

Aquatic Environmental Lab (AEL).

114

D. Massemin, S. Clavier & J.-P. Pointier

Novapex 12(3-4): 109-118, 10 octobre 2011

Study site

Watershed

Habitat type

GPS

Number of

specimens

Crique Eau Claire

Kourou

little stream

05°08'45" N / 52° 52'09" W

Takari Tanté

Sinnamary

large river

04°37'18" N / 52°55'38" W

Vata

Sinnamary

réservoir lake

04°51 ’58" N / 52°57'41 " W

Saut Fracas

Mana

large river

04°45'30" N / 53°40'49" W

Arataï

Approuague

large river

04°01'36" N / 52°41'35" W

Saut Dalles

Sinnamary

large river

04°33'21" N / 52°54'03" W

Athanase

Approuague

large river

04° 10'50" N / 52°21 ' 14" W

Apsik Icholi

Maroni

large river

02°59’10" N / 54°10’53" W

Crique Apatou

Maroni

little stream

05°09'06" N / 54°20'21" W

Papaïchton

Maroni

large river

03°48'20" N / 54° 08'36" W

Para Itou

Oyapock

large river

03°05'34" N / 52°20'28" W

Total 81

Table 1

Study site

Watershed

Habitat type

GPS

Number of

specimens

Leblond

Sinnamary

mid-order stream

04°46'37" N / 53°07'08" W

Total 44

Table 2

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D. Massemin, S. Clavier & J.-P. Pointier

Novapex 12(3-4): 109-118, 10 octobre 2011

Appendix 1

In French Guiana, Bivalvia of brackish waters, as oysters, can be observed up to more than 50km inside lands

and are therefore considered by the authors to be continental species; size is expressed for adult shells.

Drawing by the authors (DM).

1 1.1

Hinge dysodont

(absence of tooth) 2

1.2

Hinge heterodont

(cardinal and latéral teeth présents and dissimilar) 3

2 2.1

Shell extended, widened and rounded

behind, end not sharp (mytiliform); internai

surface entirely pearly; in brackish water;

shell attached by a byssus to hard substratum

size of 10 to 20 mm Dreissenidae

Mytilopsis leucophaeata ( Fig. 1)

2.2

Shell oval to variable-shaped,

kept closely; internai surface

entirely pearly; in brackish water

size more than 9 mm Ostreidae

Crassostrea rhizophorae (Fig. 2)

3 3.1

Shell circular to suboval;

in treshwater: size less than 8mm 4 (Sphaeriidae)

3.2

Shell rectangular to subtrigonal;

in freshwater; size up to 40 mm 5 (Hyriidae)

3.3

Shell subtriangular clam-shaped;

in hrackish water: size up to 10mm 7 (Corbiculidae)

4 4.1

4.2

The iirnbo is central 8 (Pisidium)

The umbo is subcentral 9 ( Eupera )

5 5.1

Shell subtrigonal;

cardinal teeth well developed;

test concentrically sculptured;

up to 45 mm Castci 1 ici sulccita (Fig. 3)

5.2

Shell rectangular;

cardinal teeth not well developed__6

6 6.1

Tessellata sculpture;

muscle scar rounded at the posterior end;

size np to 40mm Diplodon yronosus (Fig. 4)

6.2

Concentric sculpture;

muscle scar angulated at the posterior end;

size up to 60mm Diplodon voltzi (fig.5)

Palliai sinus well developed;

size more than 20 mm__ Polymesoda aequilatera (Fig. 6)

8 8.1

Shell oval and compressed;

test matt and diaphanous; whitish, slightly tinted

with fuscous patches;

surface finely concentrically striated and covered with

granular points;

latéral teeth similar in both valves;

minute species (3-5mm)__ Pisidium punctiferum (Fig. 7)

8.2

Shell inequilateral and rounded;

117

D. Massemin, S. Clavier & J.-P. Pointier

Pisidium punctiferum and Eupera viridans from French Guiana

test glossy light yellowish;

surface very finely concentrically striated and uncovered

with granular points;

latéral teeth lower and shorter in left valve;

size from 6 to 8 mm_ Pisidium sterkianum

(Species probably présent)

9 The test is cream colour ornamented with

dark brown sports;

one cardinal tooth on each valve;

size of about 8 mm_ Eupera viridans (Fig. 8)

I 18

B. Landau & G. J. Vermeij

Novapex 12(3-4): 119-123, 10 octobre 2011

New Lyriinae (Mollusca: Volutidae) from the Lower Miocene Cantaure

Formation of Venezuela

Bernard LANDAU *

Centra de Geologia da Universidade de Lisboa. Campo Grande, 1749-016 Lisboa, Portugal and

International Health Centres, Av. Infante de Henrique 7, Areias Sào Joào, P-8200 Albufeira, Portugal

bemielandau@sapo.pt

Geerat J. VERMEIJ

Department of Geology, University of California at Davis, One Shields Avenue, Davis, CA 95616,

USA. vermeij@geology.ucdavis.edu

* Corresponding author

ABSTRACT. The subfamily Lyriinae (Volutidae) is recorded from the early Miocene Cantaure

Formation of Venezuela for the first time. Two species are recorded: Lyria gabbi Vokes, 1998,

which is also found in the Lower Miocene Baitoa Formation ot the Dominican Republic, and

Enaeta inornata n. sp.

KEYWORDS. Lyriinae, Mollusca, Lyria, Enaeta ,

record, new species

INTRODUCTION

No species of Lyriinae Pilsbry & Olsson, 1954 were

mentioned in Jung’s (1965) systematic description of

the Lower Miocene Cantaure Formation fossil

molluscan assemblage on the Paraguanâ Peninsula,

Venezuela. This is hardly surprising, as both the

species recorded here are extremely rare in these

deposits. We discuss one new record of Lyria Gray,

1847 and describe a new species of Enaeta Adams &

Adams, 1853 from the Cantaure Formation.

Material and Methods

The material described here is from the Gibson-Smith

collection housed in the Naturhistorisches Muséum

Basel (NMB coll.), Switzerland and the Bernard

Landau collection (BL coll.), now deposited in the

Naturhistorisches Muséum Wien (NHMW coll.),

Vienna. We hâve examined other collections from this

locality, and none contains any species of Lyriinae.

SYSTEMATIC PALAEONTOLOGY

CLASS GASTROPODA

SUPERFAMILY VOLUTOIDEA Rafinesque, 1815

Family VOLUTIDAE Rafinesque, 1815

Subfamily LYRIINAE Pilsbry & Olsson, 1954

Genus Lyria Gray, 1847

Type species (by original désignation): Voluta nucléus

Lamarck, 1811.

Lyria gabbi Vokes, 1998

Figs 1, 2

Lyria pulchella (Sowerby). Gabb 1873, p. 219 (in

part; non G. B. Sowerby I, 1850).

Miocene, Cantaure Formation, Venezuela, new

Lyria soror (Sowerby). Pilsbry 1922, p. 338, pl. 24,

figs 11-12 (non G. B. Sowerby 1, 1850).

Lvria (Lyria) gabbi E. H. Vokes 1998, p. 11, pl. 1, figs

7-8.

Material and dimensions. Two specimens; larger,

NHMW 2009z0079/0006 (ex BL coll.), height 33.7

mm; lower shell bed, 1 km Southwest of Casa

Cantaure, about 10 km west of Pueblo Nuevo, Falcôn,

Venezuela (= locality GSI2PGNA of Gibson-Smith &

Gibson-Smith 1979) (1 other in BL coll).

Discussion. Lyria gabbi Vokes, 1998 was originally

described from the coeval Baitoa Formation of the

northern Dominican Republic. Vokes (1998) singled

out the presence of sharp barbs (prolongations of

denticles présent within the outer lip) along the margin

of the outer lip as the most characteristic feature of

this species. Neither of the shells from Cantaure are

perfectly preserved; both hâve the apex missing and

the barbs on the outer lip are worn, but the denticles

within the outer lip are clearly visible (Fig.l), and in a

similar position to those seen in a specimen of Lyria

gabbi from the type locality (Fig. 3). Vokes (1998)

discussed the presence of a very faint colour pattern

on the Dominican shells, consisting of fine spiral fines

about 1 mm apart. A similar pattern can be seen on the

Cantaure specimens (Fig.l). The position and

character of the axial omament and denticles on the

columellar lip are also identical. Lyria limata Hoerle

& Vokes, 1978 (Figs 5-6) from the contemporaneous

Chipola Formation of NW Florida also has these barbs

on the outer lip, but is otherwise not especially

similar, with fewer, broader axial ribs and a taller

spire.

Later Tropical American Lyria species seent to

hâve lost the barbs on the edge of the outer lip. Lyria

B. Landau & G. J. Vermeij

New Lyriinae from the Lower Miocene of Venezuela

pulchella (Sowerby, 1850) (Figs 7-8) from the Upper

Miocene Cercado, Lower Pliocène Gurabo and Mao

formations of the Dominican Republic lias quite a

different shell shape, with a far more inflated last

whorl, less numerous, broader ribs and a thickened,

abapically flared outer lip. This is also the only

common Lyria species in the tropical American

deposits. Lyria incomperta Hoerle & Vokes, 1978

(Figs 9-10), also from the Cercado and Gurabo

Formations of the Dominican Republic, but almost

totally confmed to coralline faciès (Vokes 1998), has

finer axial sculpture more similar to that of L. gabbi,

but has a proportionately higher spire. Both L.

pulchella and L. incomperta hâve smooth outer lips,

without the barbs seen in L. gabbi. For further

comparisons see Vokes (1998).

To our knowledge, the two early Miocene Lyria

species with barbed outer lips are the only tropical

American gastropods with this character. In the

Recent fauna, barbed lips are confmed to the Indo-

West Pacific, or to species that hâve only very

recently colonized the eastern Pacific (e.g. Casniaria

vibexmexicana Stearns, 1894). Barbed lips occur in

the strombid genus Tridentarius (Kronenberg &

Vermeij 2002), the cassid généra Casmaria and

Phalium (see Beu 2008), the mitrid genus Mitra

(sensu stricto) (Cernohorsky 1976), some species of

Harpa (see Rehder 1973), and in several nassariids

including the genus A/ectrion (Cernohorsky 1984).

The function of these barbs remains unclear, but

defence is a distinct possibility, especially in those

cassids, strombids, and nassariids in which some of

the barbs point anteriorly.

From a paleobiogeographical point of view, the

Dominican deposits belong in the West Indian

Subprovince, whereas the Cautaure deposits are

within the Columbian-Venezuelan-Trinidad

Subprovince of the Neogene Gatunian Province (see

Woodring 1974; Vermeij & Petuch 1986; Landau et

al. 2008). We can add L. gabbi to the relatively short

list of species that occur in both of these

paleobiogeographical subprovinces, together with the

following species of Enaeta, and Vasum tuberculatum

Gabb, 1873 (Vokes, 1998) and V. haitense (G. B.

Sowerby I, 1850) (BL coll.).

Genus Enaeta H. Adams & A. Adams, 1853

Type species (by subséquent désignation, Cossmann

1899): Voluta harpa Bames, 1824 (junior primary

homonym of V. harpa Mawe, 1823) = Voluta barnesii

Gray, 1825.

Enaeta inornata n. sp.

Figs 11-16

Type material and dimensions. Holotype NMB

H18371, height 30.8 mm, width 15.9 mm (Figs 11-

13); paratype 1 NHMW 2009z0078/0001, height 25.0

mm, width 13.2 mm (Figs 14-16); paratype 2 NHMW

2009z0078/0002, height 29.9 mm, width 15.8 mm.

Type locality. Cantaure Formation (early Miocene:

Burdigalian), Lower shell bed, 1 km Southwest of

Casa Cantaure, about 10 km west of Pueblo Nuevo,

Falcôn, Venezuela (= locality GS12PGNA of Gibson-

Smith & Gibson-Smith 1979).

Diagnosis. An Enaeta species with a medium-sized,

solid, biconic shell, inflated last whorl and subdued

sculpture consisting of weak spiral threads on the spire

and six to eight axial ribs, developed only on the mid-

portion of the last whorl.

Description. Shell medium-sized for genus, solid,

biconic, with inflated last whorl. Protoconch missing.

Teleoconch of five whorls with superficial linear

suture. Spire short, conical, with periphery at abapical

suture. Sculpture on spire of very faint spiral threads,

visible only under magnification. Last whorl large,

approximately 80% of total height, convex, weakly

constricted at base, bearing six to eight relatively

broad axial ribs developed only on mid-portion of

whorl. Aperture elongate, approximately 55% of total

height. Outer lip weakly convex, slightly flared

abapically, thickened by labral varix, smooth within,

except for one heavy denticle about two-thirds of

distance from anterior end to posterior end of lip. Anal

canal very narrow, shallow; siphonal canal short,

open, narrow, posteriorly recurved. Pariétal callus

weakly developed immediately adjacent to anal canal,

a thin callus wash below; columellar callus very

slightly thickened. Columella bearing four strong

oblique folds on abapical portion. Siphonal fasciole

flattened, separated from base by small ridge of shell.

Discussion. Hoerle & Vokes (1978) and Vokes (1998)

considered Enaeta to be a subgenus of Lyria.

However, we follow Poppe & Goto ( 1992) and Bail &

Poppe (2002) and consider the différences between the

two sufficiently important to separate them at generic

level within the Lyriinae (for discussion on radular

characteristics of the group and taxonomie position

see Poppe & Goto 1992, p. 19).

Two well-defined Enaeta species groups are

présent in Tropical American Recent and Neogene

assemblages. The first is represented in the fossils

assemblages by Enaeta perturbatrix (Maury, 1917)

from the Lower Pliocène Gurabo Formation of the

Dominican Republic. It is characterized by an

elongate fusiform shell with a high spire and sculpture

of numerous, well-developed axial ribs, although they

are only well-developed at the periphery on the last

whorl. This species is an early représentative of the

Caribbean group including E. guildingi (G. B.

Sowerby 1, 1844), E. reevei Dali, 1907 and E.

leonardhilli Petuch, 1988 (probably a synonym of E.

guildingi ; Poppe & Goto 1992), which ail hâve the

same elongate fusiform shell shape and differ in

details of their sculpture. Ail these species seem to

hâve two stronger anterior folds on the columella and

several weaker ones above. Enaeta trechmanni Jung,

120

B. Landau & G. J. Vermeij

NOVAPEX 12(3-4): 119-123, 10 octobre 2011

1971 from the early Middle Miocene Grand Bay

Formation (Robinson & Jung 1972; Donovan et al.

2003) probably also belongs in this group. Therefore,

this group is recorded earliest in the Atlantic.

The second group also has fusiform shells,

although stockier and more solid, with stronger axial

sculpture that forms nodules at the shoulder. This is

represented in the Recent Caribbean fauna by Enaeta

cylleniformis (G. B. Sowerby I, 1844) and in the

Recent tropical eastem Pacific fauna by E. cumingii

(Broderip, 1832). According to Pitt & Pitt (1995), the

predecessor of E. cumingii is E. propecumingii Pitt &

Pitt, 1995 from the Upper Miocene Esmeraldas Beds

of Ecuador. Ail these species hâve three strong

abapical folds on the columella and a few wealcer

folds above. Therefore, this second group is recorded

earliest in the Pacific.

Enaeta inornata n. sp. falls into a third group of

less well-defined Tropical American species with

more ovate shells, having a much larger last whorl and

a proportionately shorter spire and rather discrète

sculpture. Two fossil species bear some similarity to

E. inornata: E. isabellae (Maury, 1910) [= Strigatella

americana Dali, 1915, see Gardner 1937] from the

Lower Miocene Chipola Formation of northwestern

Florida (Figs 17-18) is the most similar in shape,

height of spire and size of last whorl. It differs in the

character of its ornament, which consists of narrow,

weak, close-set axial ribs. Enaeta ecnomia Woodring,

1964 from the middle Middle Miocene Lower Gatun

Fonnation of Panama is also similar to our

Venezuelan species in shell shape, having a broad,

solid last whorl, but is immediately distinguished by

its sculpture consisting of about 18 axial ribs on the

last whorl and close-set spiral threads covering the

entire shell surface. Enaeta ecnomia and E. isabellae

are more similar to each other than they are to E.

inornata, but E. ecnomia has an even broader shell

shape, a somewhat coeloconoid spire and fewer,

slightly stronger axial ridges than E. isabellae. Enaeta

olssoni Hoerle & Vokes, 1978 also from the Late

Miocene middle-upper Gatun Fonnation of Cativa,

Panama, and also known from a single individual, is

similar to E. isabellae, but has no axial sculpture at

ail, moreover it has placations on the posterior portion

ofthe columella.

In the Recent faunas there are no représentatives of

this group in the Caribbean, although Enaeta barnesii

(Gray, 1825) from the Recent tropical eastem Pacific

is similar in having almost no surface sculpture and an

inflated last whorl, even more so than any ot the tossil

forms. Ail of these species, except E. olssoni , hâve

four strongly developed folds on the columella. It is

likely that E. inornata is more closely related to this

Recent tropical eastem Pacific species than to the

extant Caribbean forms. However, we stop short ol

calling them a paciphi 1 ic clade (sensu Vermeij &

Petuch 1986) as this phylogeny is not indisputable.

From a paleobiogeographical angle, both Lyria and

Enaeta were présent in both the Gatunian and

Caloosahatchian Paleobiogeographical provinces

during the early Miocene, but subsequently suffered a

range contraction so that by the late Miocene-early

Pliocène they were only présent in the more Southern

Gatunian Province (see Vermeij & Petuch 1986;

Landau et al. 2008).

Etymology. The name reflects the very weak

ornament.

Acknowledgements

We would like to thank Walter Etter and Olivier

Schmidt of the Naturhistorisches Muséum Basel

(NMB), Switzerland, for access to the Gibson-Smith

collection and the loan of type specimens from the

Naturhistorisches Muséum Basel collection. Dr. Alan

G. Beu, GNS Science, Lower Flutt, New Zealand

reviewed the MS and made many useful suggestions

for improvement.

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New Lyriinae from the Lower Miocene of Venezuela

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revisited. New data from the Pliocène of Cubagua,

Venezuela. Palaeogeography, Palaeoclimatology,

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Figures 1-18

1-2. Lyria gabbi Vokes, 1998, NHMW 2009z0079/0006 (ex BL coll.), height 33.7 mm, Cantaure Formation

(early Miocene: Burdigalian), lower shell bed, Casa Cantaure, Paraguana Peninsula, Falcôn, Venezuela. 3-4.

Lyria gabbi Vokes, 1998, NHMW 2009z0079/0001 (ex BL coll.), height 44.3 mm, Baitoa Formation (early

Miocene: Burdigalian), E side Rio Yaqué del Norte, just above mouth of Arroyo Hondo, Dominican Republic (=

NMB 17290; see Saunders et al. 1986). 5-6. Lyria limata Hoerle & Vokes, 1978, NHMW 2009z0079/0002 (ex

BL coll.), height 33.1 mm, Chipola Formation (early Miocene: Burdigalian), Tenmile Creek, 2.8 km W of

Chipola River, Calhoun County, Florida (= TU 546; see Hoerle & Vokes 1978). 7-8. Lyria pulchella (Sowerby,

1850), NHMW 2009z0079/0003 (ex BL coll.), height 28.8 mm, Gurabo Formation (early Pliocène: Zanclean),

Gurabo River, E bank, firsl bluff downstream from ford on Los Quemados-Sabaneta road, Dominican Republic

(= TU 1210; see Saunders et al. 1986). 9-10. Lyria incomperta Hoerle & Vokes, 1978, NHMW 2009z0079/0004

(ex BL coll.), height 47.0 mm, Cercado Formation (late Miocene: Tortonian), unnamed formation, Lôpez area,

Rio Yaqué del Norte, Dominican Republic (= NMB 17273; see Saunders et al. 1986). 11-13. Enaeta inornata n.

sp., holotype, NMB H18371, height 30.8 mm, Cantaure Formation (early Miocene: Burdigalian), lower shell

bed, Casa Cantaure, Paraguana Peninsula, Falcôn, Venezuela. 14-16. inornata n. sp., paratype, NHMW

2009z0078/000l (ex BL coll.). height 25.0 mm, Cantaure Formation (early Miocene: Burdigalian), lower shell

bed. Casa Cantaure, Paraguana Peninsula, Falcôn, Venezuela. 17-18. Enaeta isabellae (Maury, 1910), NHMW

2009z0079/0005 (ex BL coll.), height 33.1 mm, Chipola Formation (early Miocene: Burdigalian), Tenmile

Creek, 2.8 km W of Chipola River, Calhoun County, Florida (= TU 546; see Hoerle & Vokes 1978).

122

B. Landau & G. J. Vermeij

Novapex 12(3-4): 119-123, 10 octobre 2011

123

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