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Vita Malacologica, 1: 49-54 12 December 2002
VITA MALACOLOGICA, 1: 49
(Kronenberg, 1993, 1998; Petuch, 1994; Yokogawa, 1999;
Jung and Heitz, 2001; Simone, in prep.) and from highly pre-
liminary phylogenetic analyses (Stone, 2001).
As was true of many authors of his generation and of ear-
lier workers, Abbott (1960, 1961) was typically vague about
the characters that distinguish higher taxa, with the result that
the lines among his subgenera of Strombus are blurred and
arbitrary. This is notably the case with Canarium Schuma-
cher, 1817 (type species: Canarium ustulatum Schumacher,
1817, = Strombus urceus Linnaeus, 1758), which Abbott
(1960) regarded as a subgenus of Strombus but which we treat
as a distinct genus (see below). In an attempt to identify
supraspecific taxa that correspond to true clades, we have sur-
veyed the diversity of species currently recognized in
Canarium, and here name two new genera, Terestrombus and
Tridentarius, for small, aberrant, late Neogene to Recent
strombids from the tropical Indo-Pacific region. In recogniz-
ing these new taxa, we specify a more precise characterization
of Canarium. We also take the opportunity to comment on the
status and geographic distributions of two of the three species
included in Terestrombus.
Finally, we discuss several hitherto poorly understood
shell characters in Strombidae, characters we believe may
help to delineate phylogenetic relationships in this group.
SYSTEMATICS
Strombidae Rafinesque, 1815
Terestrombus gen. nov.
Type species: Lambis fragilis Röding, 1798
Fusicanarium Romagna Manoja, 1980a: 13 [nomen nudum]
Diagnosis. — Small, thin-shelled, spirally and axially
almost smooth strombids with rounded, basally unconstricted
whorls, thin, determinate, unglazed outer lip, indistinct strom-
boid notch, and thin columellar callus.
ABSTRACT
Terestrombus (type species: Lambis fragilis Röding,
1798) and Tridentarius (type species: Strombus dentatus
Linnaeus, 1758) are two new Indo-Pacific genera of Strombi-
dae based on a comparative study of shell characters. We
extend the geographic ranges of Terestrombus afrobellatus
(Abbott, 1960) and T. terebellatus (Linnaeus, 1758), taxa
from the Indian and Pacific Oceans respectively, which we
consider full species instead of subspecies as previous authors
have done. We discuss a previously unrecognized character,
nature of edge of outer lip, as a potentially informative feature
from strombid phylogeny. All American and some Indo-
Pacific strombids, including Terestrombus, have an unglazed
adult outer-lip edge; whereas glazed lip edges are today found
only in Indo-Pacific strombids, including Tridentarius.
Projections on the outer lip have evolved multiple times in
Stromboidea, including Tridentarius. Among Neogene
Strombidae, these features evolved only in Indo-Pacific
clades.
INTRODUCTION
Abbott's (1960, 1961) monographs on the broadly defined
genera Strombus Linnaeus, 1758 (type species: Strombus
pugilis Linnaeus, 1758) and Lambis Röding, 1798 (type
species: Strombus lambis Linnaeus, 1758) have served as the
point of departure for all subsequent work on these groups
(see e.g. Romagna Manoja, 1973, 1980a, b; Walls, 1980;
Kronenberg & Berkhout, 1984, 1986; Wilson, 1993; Drivas &
Jay, 1998; De Turck et al., 1999; Stone, 2001). Species that
have been described or revised since Abbott's monographic
studies have generally been assigned to supraspecific taxa in
Abbott's classification (see e.g. Mienis, 1969, 1971; Man in ’t
Veld & Visser, 1993; Man in ’t Veld & De Turck, 1998;
Willan, 2000; reviewed by De Turck et al., 1999). Challenges
to Abbott's taxa Strombus and Lambis have come from
detailed morphological and paleontological studies
Terestrombus and Tridentarius, new genera of Indo-
Pacific Strombidae (Gastropoda), with comments on
included taxa and on shell characters in Strombidae
Gijs C. KRONENBERG
Milieu Educatie Centrum, P.O. Box 435, NL-5600 AK Eindhoven, the Netherlands
e-mail: gijsckro@worldonline.nl
Geerat J. VERMEIJ
Department of Geology, University of California at Davis, One Shields Avenue, Davis, CA 95616, U.S.A.
e-mail: vermeij@geology.ucdavis.edu
Key words: Gastropoda, Strombidae, Terestrombus, Tridentarius, taxonomy, new genus, Indo-Pacific
Kronenberg, G.C. & G.J. Vermeij - Terestrombus and Tridentarius
VITA MALACOLOGICA, 1: 50
Description. — Shell small for family (maximum height
about 49 mm), fusiform, basally unconstricted on left side
opposite outer lip, very weakly constricted at outer lip of
adult. Protoconch of about 2.5 smooth whorls. First one to
two teleoconch whorls with about eight fine spiral incised
lines, other whorls of spire smooth, rounded, little inflated,
with or without weak rounded varices. Last whorl smooth
except for fine spiral grooves on base. Terminal adult varix
present but low, defined by weak axial ridge behind sharp,
unglazed edge of lip. Inner (adaxial) surface of outer lip with
or without very fine, short spiral striae. Columellar callus
thin, narrow, smooth. Stromboid notch on adult outer lip shal-
low, poorly defined; projection between stromboid notch and
anterior canal narrow, pointed. Adapical canal present or
absent. Operculum stromboid, elongate, arched, serrated on
convex edge.
Derivation. — The prefix "tere-" has two meanings here.
First, it is a shortened version of Linnaeus' specific name tere-
bellatus, one of the species we assign to Terestrombus.
Second, the Dutch word tere (or teer) means fragile, an apt
characterization of the type species, Terestrombus fragilis.
Geographic and stratigraphic distribution. — Indo-
Pacific, late Miocene to Recent.
Other included species. — Strombus (Canarium) afrobel-
latus Abbott, 1960 (thus here considered a full species);
Strombus terebellatus Linnaeus, 1758.
Remarks. — Romagna Manoja (1980a, b) introduced the
name Fusicanarium for four species of Strombus: S. terebel-
latus Linnaeus, 1758, S. fragilis (Röding, 1798), S. dentatus
Linnaeus, 1758, and S. fusiformis Sowerby, 1842. He neither
designated a type species nor gave a description or diagnosis,
stating only that Fusicanarium differs from Canarium
Schumacher, 1817. The name Fusicanarium is therefore a
nomen nudum under ICZN Article 13. As we shall discuss
further below, the four species assigned by Romagna Manoja
(1980a, b) to Fusicanarium belong, in our view, to three gen-
era: Canarium (S. fusiformis), Terestrombus (S. terebellatus
and S. fragilis), and Tridentarius gen. nov. (S. dentatus).
Abbott (1960) included the species of Terestrombus in his
concept of Canarium, which he treated as a subgenus of
Strombus. Canarium as understood here is a large, morpho-
logically well-defined group of small Indo-Pacific Strombidae
characterized by a fusiform, basally constricted shell with an
externally (abaxially) thickened and flaring outer lip whose
edge is unglazed. The surface is generally spirally ridged. The
columellar callus in the adult is narrow and well demarcated.
A deep and distinct adult stromboid notch is separated from
the anterior canal by a rounded projection. Axial sculpture is
typically present. These features, though variable, are typical-
ly well expressed in such species as Canarium urceum
(Linnaeus, 1758), C. erythrinum (Sowerby, 1842), C. macula-
tum (Sowerby, 1842), C. labiatum (Röding, 1798), C.
microurceum (Kira, 1959), and C. mutabile (Swainson, 1821).
The species of Canarium most similar to Terestrombus is C.
fusiforme (Sowerby, 1842).
This species is typical of Canarium except that the projec-
tion between the stromboid notch and the anterior canal is less
well developed and therefore more like that of Terestrombus.
C. fusiforme differs from Terestrombus in lacking spiral lines
on early teleoconch whorls, a more distinct stromboid notch
and a well demarcated columellar callus, which bears some
plicae on the adapical side.
Species of Terestrombus show a remarkably juvenilized
appearance relative to Canarium. Adult features of
Terestrombus that resemble those of juvenile Canarium
include an indistinct, shallow stromboid notch, an outer lip
without strongly flaring or abaxially thickened form, and a
thin, poorly demarcated, smooth, narrow columellar callus.
Adult Terestrombus tend to be basally little or not at all con-
stricted, a condition again reminiscent of that in many young
Canarium. Accordingly, we propose that Terestrombus is
derived from relatively unsculptured species of Canarium
similar to the Recent C. fusiforme, C. maculatum, and some
forms of C. mutabile.
The fossil record is consistent with this hypothesis. The
oldest record of Terestrombus is that of Strombus cf. S. fra-
gilis of Ladd (1972) from the late Miocene of Bikini Atoll,
Marshall Islands. Terestrombus fragilis is also known from
the Pliocene of Vanuatu (as Strombus terebellatus of Abrard,
1946; see Abbott, 1960). Fossil specimens of Recent species
of Canarium are known from the Pliocene of Vanuatu and
Fiji (Abbott, 1960; Ladd, 1972). The earliest species of
Canarium are the extinct early and late Miocene Strombus
unifasicatus Martin, 1884, and S. spolongensis Martin, 1916,
from Java (see Abbott, 1960). There is an as yet unidentified
specimen of Canarium present in the private collection of Mr.
Robert Marquet, Antwerp, Belgium from the early Miocene
(Aquitanien) of France.
Some readers may object to our naming a taxon Tere-
strombus as a derived clade within Canarium, for such an
action would make Canarium at least paraphyletic. We take
this taxonomic action to emphasize the difference between
typical (and presumably ancestral) species of Canarium and
the more derived, more paedomorphic trio of species of
Terestrombus. Members of the latter group seem to occupy
sandy pockets and other soft sediment settings. Most typical
Canarium species occur on vegetated sand flats and grass-
beds, or, in the case of C. mutabile and related species, on the
hard substrata of reef flats and benches (Vermeij, pers. obs.).
Terestrombus has several characters in common with two
other Indo-Pacific strombid genera, Conomurex and Gibberu-
lus. In Conomurex P. Fischer, 1884 (type species: Strombus
luhuanus Linnaeus, 1758), the base is entirely unconstricted
(a condition more extreme than in Terestrombus, where the
left side is unconstricted and the right side is very weakly
concave at the outer lip), and the columellar callus is
extremely thin and indistinctly set off from the outer surface
of the body whorl. Conomurex
differs from Terestrombus by
the generally very low spire, and by the presence of a distinct,
broad, adapical notch on the outer lip, a stronger developed
stromboid notch and a distinct, broad flange between the
stromboid notch and the anterior canal. The spire whorls of
Conomurex have more or less well developed axial ribs. The
development of axial sculpture on the shoulder of the body
Kronenberg, G.C. & G.J.Vermeij - Terestrombus and Tridentarius
VITA MALACOLOGICA, 1: 51
whorl of Conomurex varies. In C. persicus (Swainson, 1821),
axial sculpture is absent, whereas in C. fasciatus (Born, 1778)
it is expressed as distinct knobs (see Moolenbeek & Dekker,
1993, for assigning this species to Conomurex). In the other
three species of Conomurex, viz. C. luhuanus, C. decorus
(Röding, 1798) and C. masirensis (Moolenbeek & Dekker,
1993) [=? C. coniformis (Sowerby, 1842)], the expression of
shoulder axials varies among individuals and among popula-
tions. Shoulder axials are always absent in Terestrombus.
The genus Gibberulus Jousseaume, 1888 (type species:
Strombus gibberulus Linnaeus, 1758) resembles Terestrom-
bus in having a very thin, indistinctly demarcated columellar
callus; and in some immature specimens, an almost uncon-
stricted base. Like Conomurex, however, Gibberulus differs
from Terestrombus in having a distinct adapical notch, at least
in the flaring adult outer lip. Moreover, Gibberulus has a
glazed outer-lip edge in the adult, whereas in Terestrombus
the edge remains unglazed. Species of Gibberulus usually
have broad varices. Spiral sculpture is variable in expression.
In G. gibberulus from the Indian Ocean and G. albus (Mörch,
1850) from the Red Sea, the abaxial surface of the outer lip
has distinct spiral sculpture, a feature variably developed in
the one or more species comprising the Pacific taxon G. gib-
bosus (Röding, 1798). [Abbott, 1960, and Coomans & van
Amsterdam (1970) considered Gibberulus to be a subgenus of
Strombus, and recognized a single species with three geo-
graphic subspecies. We prefer to treat the three included taxa
as three species pending a thoroughly needed revision.]
There is a striking but superficial resemblance between
Terestrombus and the Recent Indo-Pacific seraphsid Terebel-
lum terebellum (Linnaeus, 1758). The latter species differs
from Terestrombus in lacking a stromboid notch, in the
absence of any sculpture on the spire whorls, in having the
edge of the adult outer lip glazed, and in having the base as
the unconstricted widest part of the shell. As Jung (1974) has
pointed out, T. terebellum is the only Recent member of the
stromboidean family Seraphsidae, which is known back to at
least the Paleocene. The resemblance between Terebellum
Röding, 1798, and Terestrombus is thus entirely the result of
convergence. Both taxa appear to be rapid burrowers in sand.
Whether species of Terestrombus can swim for short distan-
ces, as Terebellum can (Jung & Abbott, 1967), is unknown.
Tridentarius gen. nov.
Type species: Strombus dentatus Linnaeus, 1758
Fusicanarium Romagna Manoja, 1980a: 13 [nomen nudum]
Diagnosis. — Small, high-spired strombids with strongly
reduced spiral sculpture, determinate outer lip with glazed
adult edge and three sharp basal projections. An indistinct
stromboid notch is separating the two abapicalmost projec-
tions. A thick, narrow columellar callus is present, and a long
adapical apertural channel.
Description. — Shell small for family (maximum height
56.5 mm), fusiform, basally very weakly or not constricted.
Protoconch of about 2.5 smooth whorls. First one to two
teleoconch whorls with eight very fine incised spiral lines and
fine growth lines. First three to four teleoconch whorls with
varices. Later spire whorls with gradually appearing axial
ribs, most strongly expressed at periphery, abapically fading
out above midpoint of whorl. Determinate adult outer lip lit-
tle expanded, with glazed edge, adapically extending as nar-
row apertural channel with the adapical extension of the
thickened, narrow columellar callus. Edge of adult outer lip
dentate, the adapical two teeth axially broad and rounded,
lobe-like, the abapical three teeth sharp, triangular, spinelike,
directed ventrally and abapically. Stromboid notch shallow
but distinct, situated between the two abapicalmost teeth.
Abapical end of inner lip pointed, extending beyond abapi-
calmost tooth on outer lip. Columella abapically and adapi-
cally with a few indistinct thin plicae. Outer lip very little
expanded abaxially, finely spirally striate on inner (adaxial)
surface. Aperture narrow, widening abapically. Operculum
stromboid, elongate, arched, serrated on convex edge.
Derivation. — The name refers to the three spine-like
abapical projections or teeth on the lip. The name is a tribute
to Gmelin (1791), who so aptly (if posteriorly) named the
type species Strombus tridentatus. Linnaeus' S. dentatus, a
less evocative name, takes nomenclatorial precedence.
Gender, masculine.
Geographic and stratigraphic distribution. — Indo-
Pacific, Pleistocene to Recent.
Remarks. — As pointed out in the remarks under Tere-
strombus, Romagna Manoja (1980a, b) included Strombus
dentatus as one of four species in his new genus
Fusicanarium. The latter taxon was not described or com-
pared to other taxa, and no type species was designated. It is
therefore a nomen nudum.
Abbott (1960) included Strombus dentatus in Canarium
Schumacher, 1817. This assignment was evidently based on
general shape and small size. The species differs in so many
ways from Canarium that a close phylogenetic relationship
seems unlikely. We therefore propose the new genus Triden-
tarius for it.
Tridentarius differs from Canarium in having an adapical
apertural channel, a glazed rather than unglazed adult outer
lip edge, a nearly or entirely unconstricted base instead of a
distinctly concave abapical sector on the body whorl, and
sharp projections on the abapical sector of the outer lip,
whose entire length is wavy or dentate. Despite their general-
ly similar fusiform shells, Tridentarius and Terestrombus
also differ in many characters. Tridentarius has, and Tere-
strombus lacks, axial ribs on later teleoconch whorls. The
outer lip edge of Tridentarius adults is glazed and abapically
adorned with spine-like projections, whereas that of
Terestrombus is unglazed and unornamented. Finally, the
columellar callus is thick and distinctly demarcated from the
body whorl in Tridentarius, whereas it is thin and indistinctly
set off in Terestrombus.
There is a striking superficial resemblance between
Tridentarius and the middle Eocene Rimella-like strombid
genus Cowlitzia Clark & Palmer, 1923, from western North
America. Both taxa have axial ribs on the spire whorls and
body whorl, and sculpture at the edge of the abapical sector
Kronenberg, G.C. & G.J. Vermeij -Terestrombus and Tridentarius
VITA MALACOLOGICA, 1: 52
extend the known geographic range of T. afrobellatus. We
now have material from Mozambique (Fernal Veloso Beach,
Nacala Bay, three specimens, GCK 2823); southwestern
Madagascar (Tulear, shallow water on sand, one specimen
taken February, 2001); Maldives (Raabandhiuraa Island,
Malaku Atoll, one living and one dead specimen taken 7
November, 1998 by Sandro Gori, GCK 5908). There are two
specimens of Terestrombus in the Kronenberg collection from
the Red Sea, Gulf of Aqaba, which look very much like T.
terebellatus. The paucity of specimens from the Red Sea
available at present does not permit a definite conclusion
about their identiy. For the time being we follow Mienis
(1970, 1975, pers. comm. Gijs C. Kronenberg 2001) who con-
siders the Red Sea specimens conspecific with T. terebellatus.
The Kronenberg collection also contains a specimen of T.
terebellatus, which extends the known geographic range of
that mostly Pacific species westward into the eastern Indian
Ocean. The specimen, taken by Bunjamin Dharma in "sand
fill" at Serangan, Bali (GCK 5934), is the only Indian Ocean
record of this taxon.
REMARKS ON STROMBID CHARACTERS AND
PHYLOGENY
Although Stone (2001) has made a good beginning in his
phylogenetic study of the strombid genus Lambis Röding,
1798, much work remains to be done on the family as a
whole, including a more refined approach to the recognition
and description of shell characters than is currently available
in the literature. Lambis, Strombus s.l., and probably other
supraspecific taxa in Strombidae are polyphyletic or para-
phyletic (Stone, 2001; Yokogawa, 1999; Simone, unpublished
data).
In our examination of material for this paper, we have
identified a character of the adult outer lip that has not previ-
ously been mentioned in the literature. In some strombids, the
edge of the outer lip remains unglazed. That is, the smooth
adaxial shell layer (the one on the "inner" side of the outer lip)
does not extend across the edge to the outer (abaxial) side of
the outer lip. This is the condition in the Indo-Pacific genera
Canarium Schumacher, 1817; Conomurex Fischer, 1884;
Doxander Iredale, 1936; Harpago Gill 1870 and Terestrom-
bus gen. nov., as well as in several species currently assigned
to other supraspecific taxa in Abbott's (1960) scheme. By
contrast, other strombids have the abaxial shell layer extended
across the edge to form a narrow band on the abaxial side,
with the result that the edge of the outer lip is glazed. This
condition occurs in the American and eastern Atlantic clade
of Strombus Linnaeus, 1758, and allied taxa (see below), and
in the Indo-Pacific genera Dolomena Iredale, 1931 (here
taken narrowly to include the type species Strombus pulchel-
lus Reeve, 1851), Euprotomus Gill, 1870, Gibberulus Jous-
seaume, 1888, Labiostrombus Oostingh, 1925, Laevistrombus
Abbott, 1960 (not Kira, 1955, who introduced the name as
nomen nudum; see Bieler & Petit, 1996), Lambis Röding,
1798, Mirabilistrombus Kronenberg, 1998, Tricornis Jous-
of the outer lip. Species of Cowlitzia, however, have distinct
spiral sculpture, unlike Tridentarius. The serrations on the
outer lip of Cowlitzia are situated in a more abapical position,
and continue through the shallow stromboid notch, whereas in
Tridentarius the stromboid notch separates the two abapical-
most denticles, which are larger and more spinelike than the
serrations of Cowlitzia. The adapical canal of Cowlitzia dif-
fers from that of Tridentarius in being Rimella-like, extending
from the main part of the aperture in an adapical direction
along the spire, then recurving toward the spire's dorsum (see
also Wrigley, 1938, for a discussion of the adapical canal in
Rimella-like strombids). In Tridentarius, the canal extends
straight back adapically and does not recurve. Although we
suspect that the resemblance between Cowlitzia and
Tridentarius is due to convergence, a careful study is needed
in which these two taxa are compared to the Rimella-like gen-
era Ectinochilus Cossmann, 1889, and Dientomochilus Coss-
mann, 1904, and to several Canarium-like strombids.
A NOTE ON TERESTROMBUS AFROBELLATUS
Abbott (1960) named Strombus (Canarium) terebella-
tus afrobellatus from the Indian Ocean as a subspecies of S.
terebellatus Linnaeus, 1758. He differentiated it from the
nominal subspecies by noting that "the apertural wall of the
body whorl lacks the small spiral brown color streaks,
although the colors of the outer shell may show through.
Spire only one third the length of the entire shell" (p. 88). In
his description of S. t. terebellatus, Abbott (1960: 87) noted
"spire height almost one half the length of the entire shell.".
Abbott (1960) examined specimens of S. t. afrobellatus from
Kenya, Tanzania, and Mozambique, and noted that specimens
from Nossi-Bé (northwestern Madagascar) and Jubal Island
(Red Sea) are "probably this subspecies" (p. 88). In examin-
ing additional material at the Zoologisch Museum Amsterdam
(ZMA), Mienis (1969) noted that S. t. afrobellatus has a more
or less well developed posterior (adapical) canal, a feature
more or less lacking in T. t. terebellatus. We do not find this
to be a consistent difference. Mienis (1969) questioned the
occurrence of S. t. afrobellatus in the Red Sea, and reported
typical S. t. terebellatus from the Red Sea locality of Obhur,
Saudi Arabia. He further reported S. t. afrobellatus from the
Andaman Islands, but questioned the reliability of this record
because the material is from an old collection. Mienis (1975)
later mentioned "many additional specimens from several
other Red Sea localities" but explicitly listed only Eilat,
Israel. Pickery & Wellens (1998) reported both subspecies
from Eilat. Mienis (pers. comm. to Gijs C. Kronenberg, 2001)
stated that to his knowledge there are no reliable records of T.
afrobellatus from the Red Sea.
Examination of material in Kronenberg's collection
(GCK, 20 specimens) leads us to the conclusion that the two
taxa previously considered subspecies of S. terebellatus are
specifically distinct members of Terestrombus. We observed
no specimens with intermediate characters.
Specimens in the Kronenberg collection enable us to
Kronenberg, G.C. & G.J. Vermeij - Terestrombus and Tridentarius
VITA MALACOLOGICA, 1: 53
ple times independently. Among Neogene Strombidae, these
projections have evolved only in Indo-Pacific clades.
ACKNOWLEDGEMENTS
Gijs C. Kronenberg wants to thank: Mr. Sandro Gori
(Livorno, Italy) for the donation of two specimens of
Terestrombus afrobellatus; Mr. Bunjamin Dharma (Djakarta,
Indonesia) for the donation of a specimen of Terestrombus
terebellatus; Mr. Henk K. Mienis (Hebrew University of
Jerusalem) for information concerning the occurrance of
Terestrombus in the Red Sea; Dr. Luis Ricardo L. Simone
(Museo de Zoologia da Universidade de São Paulo, Brazil)
for information about his research on Strombidae based on
anatomical characters; Mr. Robert Marquet (Antwerp,
Belgium) for showing his fossil specimen of Canarium; Mr.
Adri Burger (Heerhugowaard, the Netherlands) for stimulat-
ing discussions; Marianne Matthijssen for her support and
encouragement.
REFERENCES
ABBOTT, R.T., 1960. The Genus Strombus in the Indo-
Pacific. — Indo-Pacific Mollusca 1(2): 33-146.
ABBOTT, R.T., 1961. The Genus Lambis in the Indo-Pacific.
— Indo-Pacific Mollusca 1(3): 147-174.
BIELER, R. & R.E. PETIT, 1996. Additional notes on nomi-
na first introduced by Tetsuaki Kira in "Coloured illustra-
tions of the shells of Japan". — Malacologia 38(1-2): 33-
34.
CLARK, B.L. & D.K. PALMER, 1923. Revision of the
Rimella-like gastropods from the west coast of North
America. — Bulletin of the department of Geological sci-
ences of the University of California 14(7): 277-288, pl. 51.
COOMANS, H.E. & M.L.M. VAN AMSTERDAM, 1970.
Distribution of the Strombus gibberulus complex in Indo-
nesia (Gastropoda, Strombidae). — Beaufortia 18(232):
113-118.
DE TURCK, K., K. KREIPL, L. MAN IN ’T VELD & G.T.
POPPE, 1999. The family Strombidae. In: G.T. POPPE &
K. GROH, dir., A conchological Iconography: 1-60, pls 1-
130. Hackenheim.
DRIVAS, J. & M. JAY, 1998. The Strombidae of Réunion.
— La Conchiglia 30(suppl. 289): 10-15.
INTERNATIONAL COMMISSION ON ZOOLOGICAL
NOMENCLATURE, 1999. International code of zoological
nomenclature. Ed. 4: i-xxix, 1-306. London.
JUNG, P. & A. HEITZ, 2001. The subgenus Lentigo (Gastro-
poda: Strombidae) in tropical America, fossil and living. —
The Veliger 44(1): 20-53.
KRONENBERG, G.C., 1993. On the identity of Lambis
wheelwrighti Greene, 1978 and L. arachnoides Shikama,
1971. — Vita Marina 42(2): 41-56.
KRONENBERG, G.C., 1998. Revision of Euprotomus Gill,
1870. 1. The systematic position of Strombus listeri Gray,
seaume, 1886, Lentigo Jousseaume, 1886, and Tridentarius
gen. nov.
Jung & Heitz (2001) assigned a series of late Miocene to
Recent strombids from the western Atlantic and eastern
Pacific to the genus Lentigo. The American group, typified by
Strombus raninus Gmelin, 1791, superficially resembles the
Indo-Pacific Lentigo by having broad, nodose cords and a
wide, thick, columellar callus. The two groups differ, how-
ever, in several characters of the outer lip. In the S. raninus
group, the adapical end of the outer lip is a rounded lobe sepa-
rated from the spire by a wide space, corresponding to a broad
sinus. In Lentigo, the adapical end of the outer lip is peculiar-
ly angled and situated close to the spire, forming a long adapi-
cal channel along the spire and terminating in a distinct adapi-
cal notch. The adapical part of the columellar callus of the S.
raninus group is smooth, whereas that in Lentigo covers
nodes that were on the shell exterior before the columellar
callus spread over them. In Lentigo there are small triangular
projections on the outer lip between the stromboid notch and
the anterior canal. These projections are lacking in the S. ran-
inus group. Lentigo is an Indo-Pacific group consisting of
only two species, L. lentiginosus and L. pipus (Röding, 1798).
Species of the S. raninus group are convergent with Lentigo,
but evidently belong to an American, probably monophyletic
clade, group that includes Strombus s.s. (based on Strombus
pugilis Linnaeus, 1758, the type species, the Recent Floridian
S. alatus Gmelin, 1791, and the eastern Pacific S. gracilior
Sowerby, 1825, together with many fossil taxa) and the taxa
Aliger Thiele, 1929, Eustrombus Wenz, 1940, Macrostrom-
bus Petuch, 1994, and Titanostrombus Petuch, 1994. This
mainly American group also contains the Recent West
African S. latus Gmelin, 1791, and its forerunner S. coronatus
Defrance, 1827. The entire Atlantic-eastern Pacific clade can
be traced back to such species as S. radix (Brongniart, 1823)
and S. bonellii Brongniart, 1823 in Europe, which extend
back to the late Oligocene (see also Lozouet & Maestrati,
1986). For S. bonellii an older name, S. nodosus Borson,
1820, might be available (pers. comm. Adri Burger to Gijs C.
Kronenberg).
Another character of interest is the presence of denticles
along the edge of the outer lip. Projections of the outer lip are
the rule in the stromboidean family Aporrhaidae, and are
widespread in Strombidae. In the strombid genera Lambis and
Harpago, these projections are oriented in the plane of the
aperture or are curved dorsally. In the Indo-Pacific rostellarid
genera Tibia Röding, 1798, Rostellariella Thiele, 1929, and
Rimellopsis Lambiotte, 1979 the shells are fusiform, have
wide-spaced projections situated just behind the glazed edge
of the outer lip, and have a ventralward orientation, as in
Tridentarius. The latter genus is the only Recent strombid
genus in which the projections have a downward and abapical
orientation and are situated at the edge of the outer lip. The
more closely spaced serrations of the Eocene Rimella-like
strombid Cowlitzia also lie along the lip's edge. Lip projec-
tions in Strombidae have thus formed at different locations
and have different orientations, and evidently evolved multi-
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Strombidae). — Basteria 33(5-6): 109-114.
MIENIS, H.K., 1971. Revision of Strombus (Canarium) mu-
tabilis ochroglottis Abbott (Gastropoda: Strombidae). —
Archiv für Molluskenkunde 101 (5-6): 301-304.
MIENIS, H.K., 1975. Strombus terebellatus from the Red
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of Strombus decorus and Strombus persicus, with the de-
scription of Strombus decorus masirensis n.ssp. and a note
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VITA MALACOLOGICA, 1: 54
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