On the identity of Strombus coniformis Sowerby II, 1842 (Gastropoda, Strombidae), with additional notes on its distribution.

Abstract

Based on shell characters, Strombus (Conomurex) coniformis Sowerby II, 1842 can be distinguished from S.
(C.) decorus (Röding, 1798). The distribution range of both forms is partly overlapping, without intermediates being known. Therefore both have to be considered valid species. As the type specimens of S. (C.) coniformis are considered lost, a neotype for this species is designated. S. (C.) decorus masirensis Moolenbeek & Dekker, 1993 is a junior objective synonym of S. (C.) coniformis. Conomurex is considered a genus, rather then a subgenus of Strombus. New locality data for Conomurex coniformis are provided, forming a range extension for this species from Iran southwards to Eastern Africa.

Full text

659
INTRODUCTION
The genus Conomurex P. Fischer, 1884 (type spe-
cies Strombus luhuanus Linnaeus, 1758), here con-
sidered to be a genus in the family Strombidae, is
characterised by an overall conical shape, absence
of spiral sculpture, a sharp edged outer lip which
barely flares, and a columella which is hardly or
not thickened by a callus. Moolenbeek & Dekker
(1993) have discussed some of the species allo-
cated to Conomurex (as a subgenus of Strombus).
Summarised, their conclusions were: 1) Strombus
persicus Swainson, 1821 is a species rather then
a subspecies; 2) S. fasciatus Born, 1778 should be
allocated to Conomurex; 3) the description of a
new subspecies, viz. S. decorus masirensis.
In their paper, Moolenbeek & Dekker (1993: fig.
15) also illustrated a shell that they identified as
S. (C.) decorus forma coniformis, thereby follow-
ing the opinion of Abbott (1960: 138). They did
not, however, refer to Sowerby’s (1842) original
description and figures.
During the 1990’s and early 2000’s some
Conomurex specimens from further south on
the Arabian peninsula (mainland of Yemen and
Socotra) and localities along the East African coast
(Somalia, Brava (= Baraawe), Kisimajo and Bagiuni
islands (both close to the border with Kenya);
Zanzibar; Madagascar, Mahajanga and Tulear)
were found, which fitted the description of S. (C.) d.
masirensis. These new, more southern, localities for
this taxon as well as the unclear status of S. decorus
f. coniformis urged us to re-evaluate the status of the
taxon described by Moolenbeek & Dekker, as well
as that of Strombus coniformis Sowerby II.
We also examined a number of specimens of S.
decorus to verify the differences between this spe-
cies and C. decorus forma coniformis sensu Abbott
(1960) and Moolenbeek & Dekker (1993). From
the observations made, it appears that in the
eastern part of the Indian Ocean the form of S.
decorus with plicae and/or knobs on the shoulder
of the teleoconch whorls is absent.
Based on conchological characters, it has
been advocated to consider the strombid taxa
employed as subgenera by Abbott (1960) as full
genera (Kronenberg & Vermeij, 2002 and refer-
ences therein). More recently, this approach was
confirmed by Simone (2005) based on anatomi-
cal data, and Latiolais (2003) and Latiolais et al.
(2006) based on molecular data. Simone, Latiolais
and Latiolais et al. conclude that Strombus sensu
Abbott is paraphyletic, and based on the clado-
grams presented by Latiolais and Latiolais et al.,
the name Strombus should only be employed to
denote a group of species within a clade that
is now restricted to the (sub)tropical Americas
and to West Africa; in the latter this clade is
represented by S. latus (see Kronenberg &
Vermeij, 2002). This species is now allocated to
Persististrombus Kronenberg & Lee, 2007
Fossil representatives of this clade are known
from the Miocene in Europe. This is discussed
by Kronenberg & Lee (2007) and will be further
discussed by Harzhauser & Kronenberg (2008;
in prep.). Conomurex is considered here to be
a monophyletic group of Indo-Pacific strombs,
ON THE IDENTITY OF STROMBUS CONIFORMIS SOWERBY
II, 1842 (GASTROPODA, STROMBIDAE), WITH ADDITIONAL
NOTES ON ITS DISTRIBUTION
GIJS C. KRONENBERG
1
, VIRGILIO LIVERANI
2
& HENK DEKKER
3
1
c/o Milieu Educatie Centrum, P.O. Box 435, NL-5600 AK Eindhoven, The Netherlands
2
Via Batticuccolo 45, I-48018 Faenza Ra., Italy,
3
Zoological Museum, Universiteit van Amsterdam, Dept of Malacology, P.O.Box 94766, NL-1090 GT Amsterdam, The Netherlands
Abstract Based on shell characters, Strombus (Conomurex) coniformis Sowerby II, 1842 can be distinguished from S.
(C.) decorus (Röding, 1798). The distribution range of both forms is partly overlapping, without intermediates being known.
Therefore both have to be considered valid species. As the type specimens of S. (C.) coniformis are considered lost, a neotype
for this species is designated. S. (C.) decorus masirensis Moolenbeek & Dekker, 1993 is a junior objective synonym of S.
(C.) coniformis. Conomurex is considered a genus, rather then a subgenus of Strombus. New locality data for Conomurex
coniformis are provided, forming a range extension for this species from Iran southwards to Eastern Africa.
Key words Strombidae, Conomurex, neotype, range extension, Indian Ocean.
Contact author : gijs.kronenberg@tiscali.nl
JOURNAL OF CONCHOLOGY (2009), VOL.39, NO.6

GC KRONENBERG, V LIVERANI & H DEKKER
660
closely related to Gibberulus fide Latiolais et al.
(2006).
ABBREVIATIONS
GB Gianluigi Bini, Città di Castello, Italy,
private collection
GCK Gijs C. Kronenberg, Eindhoven, The
Netherlands, private collection
HD Henk Dekker, Winkel, The Netherlands,
private collection
JW Joop Wiersma, Kerkrade, The
Netherlands, private collection
MF Maurizio Forli, Prato, Italy, private col-
lection
MNHN Museum National d’Histoire Naturelle,
Paris, France
RMNH Nationaal Natuurhistorisch Museum,
Leiden, The Netherlands
spec./specs (in material examined) specimen(s)
VL Virgilio Liverani, Faenza, Italy, private
collection
unreg. unregistered
ZMA Zoölogisch Museum, Amsterdam, The
Netherlands
SYSTEMATICS
Family STROMBIDAE Rafinesque, 1815
Genus Conomurex P. Fischer, 1884
Type species Strombus luhuanus Linnaeus, 1758
by monotypy
Conomurex decorus (Röding, 1798)
(Figs 1-8, 13)
Lambis decora Röding, 1798: 62, sp. 777 [referring
to Chemnitz, 1788: pl. 157 figs 1499 and 1500]
For principal synonyms (except Strombus coni-
formis) see Abbott (1960). Records of Conomurex
decorus can be found in literature, but we do not
include a full list, since Abbott synonymised C.
coniformis with C. decorus, which was followed
by subsequent authors, and therefore makes
their true identity questionable, except when
specimens are illustrated. Good illustrations of
this species can be found in Walls (1980: 142 top
figs); Kronenberg & Berkhout (1984: pl. 8 fig 9);
Moolenbeek & Dekker (1993: figs 14, 15); DeTurck
et al. (1999: pl. 112 figs 1-5, pl. 113 figs 1-5, pl. 114
fig. 1); Heiman et al. (2007: figs 2, 4).
Type material Chemnitz (1788: figs 1499 and
1500) illustrated two views of one specimen (here
reproduced Figs 1, 2). Röding (1798) referred to
Chemnitz’s illustrations for his new species, this
figured specimen should be considered a syn-
type of Lambis decora. To unequivocally clarify the
identity of this species, and distinguish it from
Strombus coniformis Sowerby, 1842 (see further
below) we designate the specimen figured by
Chemnitz as lectotype of Lambis decora Röding,
1798. The present whereabouts of this specimen
are unknown; it might be in Copenhagen, or still
in the Chemnitz collection, now in St. Petersburg
(Martynov, 2002). The type locality is Mauritius
(designated by Abbott, 1960).
Brief diagnosis Species of Strombidae with coni-
cal shape, low spire, distinct stromboid notch and
posterior notch. Outer lip hardly flaring, thin,
unglazed. Columellar callus very thin. Colour
very variable, white background with irregular
narrow or broad mottling, nettings or zigzag
streaks, whether or not set in a pattern of more
or less distinct spiral bands. Adcolumellar side of
outer lip white, deeper within the aperture pink
to orange red, base of columella white.
Conomurex decorus is a well-known Indian
Ocean species, and can only be confused with
C. persicus (Swainson, 1821) and C. coniformis.
For differences from C. persicus see Moolenbeek
& Dekker (1993), although it should be noted
that the differences in the radula as noted by
Moolenbeek & Dekker should be regarded with
some reservations, as Mutlu (2004) clearly dem-
onstrated sexual dimorphism in the radula of C.
persicus. For differences with C. coniformis see
further below.
Material examined Unknown: ex Jousseaume,
1916, MNHN unreg., 1 (knobbed form, in sam-
ple with 1 C. coniformis); ZMA unreg., 3 specs;
ex Hoogeveen, RMNH unreg., 1 spec. “Indian
Ocean”, ex Reinwardt, RMNH unreg., 6 specs
(2 juvenile, 1 knobby form). “East Africa”,
1962, RMNH unreg. 1 spec. South Africa: Port
Elizabeth, Schoenmakerskop, at low tide in rock
pool, GCK 3698, 1 spec.. Mozambique: ex coll.
Jan Berkhout, GCK 5696, 1 spec.; VL 900-01, 3

IDENTITY OF STROMBUS CONIFORMIS
661
specs (knobby form); VL 900-19, 4 specs; Ilha
Magaruque, on sand at low spring tide, ix 1971,
ex C.A. v.d. Peijl, RMNH unreg., 3 specs; District
Inhambane, Ilha Santa Carolina, offshore between
Inhassoro and Chuambo, leg. M. de Lanoy Meijer
– de Geer, RMNH unreg., 6 specs; (as Portuguese
East Africa), ex J.Mulder, RMNH unreg., 10 specs;
(as Portuguese East Africa), “Porto Amelia”,
ex Mulder, RMNH unreg., 4; Nacala Bay, JW
03.540c(2), 1 spec.; Nacala Bay, JW 03.540c(3), 1
spec.; Maputo Bay, Ihaca Isl., JW 03.540(6), 1spec.;
Maputo Bay, Ihaca Isl., JW 03.540(7), 1 spec.;
Bazaruta Isl., JW 03.540(1), 1 spec.; Tanzania:
Zanzibar, x 1926, leg. G. Van Roon, RMNH unreg.,
1 spec.; Zanzibar, Uroa, leg. H.H. Kool, 1994, ZMA
unreg., 1; Zanzibar, E side Uroa, near Tamarind
Beach Hotel, coral bank at low tide, leg. H.H.
Kool, 29 ix 1994 – 11 x 1994, RMNH unreg., 1
spec.; Zanzibar, Uroa, leg. H.H. Kool, 1995, ZMA
unreg., 2 specs; Zanzibar, N point Nungwi, at
low tide, leg. H.H. Kool, ix 1995, RMNH unreg.,
1 spec.; Zanzibar, viii.1999, leg. V.Liverani, GCK
5998, 4 specs; Zanzibar, Bweju, ix.1997, beached,
GCK 5822, 1 spec.; Zanzibar, Ras Nungwi, at low
tide xii 2000 VL 900-14, 30 specs (some knobby
form); Zanzibar, Kiwengwa beach, ix 2002, by
snorkelling, VL 900-15, 12 specs (some knobby
form); Zanzibar, Jambiani, 1998, HD 547, 6 specs;
Zanzibar, Uroa, 1995, HD 1482, 2 specs. Kenya:
no further data JW 03.540(8), 4 specs; Mombasa,
GCK 438, 1 spec.; Mombassa, in shallow water,
1986, GCK 906, 1 spec.; near Mombasa in shallow
water, 1986, ex Jan Berkhout, GCK5691, 1 spec.;
Mombasa, leg. P.F. Fontei, ZMA unreg., 2 specs;
Ukunda, leg. S. Hillaar, 1956, ZMA unreg., 1 spec.;
Diana Beach, leg. T. Bruins, 1973, ZMA unreg., 2
specs; Diana Beach, leg. R.G. Moolenbeek, 1981,
ZMA unreg., 2 specs; Seychelles: St. Anne Island,
ex coll. N. Koekoek, 1990, ZMA unreg., 1 spec.;
NIOP-E, Tyro Seychelles 1992/1993, Sta. 758 St.
Joseph atoll, N rim 5°24’S 53°20’E, reef flat 27 xii
1992 RMNH unreg., 1 subadult; NIOP-E, Tyro
Seychelles 1992/1993, Sta. 780 Poivre atoll, W rim
5°46’S 53°18’E, reef slope at 10m 1 i 1993 RMNH
unreg., 1 spec.. Somalia: Brava, 1984, by snorkel-
ling, VL 900-06, 1 spec.; Merca, beached, GB 11578,
1 spec.. Madagascar: no further data JW 03.540(9),
3 specs; Nosy Be isl., VL 900-13, 2 specs; Mahajanga
bay, beached, GB 459, 2 specs; Tulear, at 1m on
rocky bottom covered with algae, ex Jan Berkhout,
GCK 5695, 1 spec. (knobby form); Tulear, from
shell market, 1985, ex M. Bloecher, ZMA unreg.,
2 specs; Tulear VL 900-10, 8 specs;Itafy, beach ven-
dors, 1985, ex M. Bloecher, ZMA unreg., 2 specs;
Itafy lagoon, 1985, leg. M. Bloecher, ZMA unreg.,
1juv., 1 subadult, 3 adult; Ifaty Lagoon, at 3 m on
sand, 1987, GCK 990, 1 spec.; Ifaty, leg. Bozzetti,
VL 900-09, 6 specs; Anakao, 1985, by snorkel-
ling, VL 900-11, 18 specs (some knobby form);
Fort Dauphin, JW 03.540(1), 1 spec. Comores Isl:
Majotte, VL 900-24, 6 specs; Europa Isl.: in lagoon
in shallow water, GCK 2562, 1 spec.. Réunion: St.
Gilles, dived at night at 35m on algae, GCK 3068,
1 spec.; Cap la Houssaye, dived at 10m on algae,
GCK 3069, 1 spec.; Cap la Houssaye, at 12m on
sand, 1987, ex coll. Jan Berkhout, GCK 5690, 1
spec.; Cap la Houssaye, at 11m on sand, 1988, ex
coll. Jan Berkhout, GCK 5690, 2 specs. Mauritius:
ex coll. Frank, 1897, RMNH unreg., 2 specs; ex
coll. C.A. v.d. Peijl, RMNH unreg., 2 specs; leg.
W.Gyngell, ex coll. J. Mulder, RMNH unreg., 1
spec.; don. P.L. van Pel, ZMA unreg., 1 spec; ex
coll. M.M. Schepman, ZMA unreg., 1 spec.; ex coll.
L. de Priester, ZMA unreg., 1 spec.; Ile Marianne,
leg. W. Regter & B. Gras, 2002, ZMA unreg., 1
spec.; ex coll. Schepman, ZMA unreg., 3 specs;
ex coll. Ph. Dautzenberg, ex coll. L. de Priester,
1934, ZMA unreg., 1 spec; idem, 4 specs; ex coll.
G. Lieftinck, 1940, ZMA unreg., 1 spec.; Ile Plate,
GCK 101, 1 spec.; Le Morne, ex. Jan Berkhout,
GCK 5694, 1 juvenile; Trou, near shipwreck, leg
Jan Paul Buys, 16.viii.1996, GCK unnumbered, 1
spec.. Rodriguez: Birds islet, 1983 by snorkelling,
VL 900-03, 1 spec.. Chagos Isls: Diego Garcia, VL
900-07, 2 specs. Oman: Persian Gulf (doubtful), ex
coll. Jan Berkhout, GCK 5688, 1 spec.. Maldives:
VL 900-02, 2 specs. India: SW coast, don. E. Wils,
1977, ZMA unreg., 1 spec.. Sri Lanka: Trincomalee,
China Bay, leg. P.L. van Pel, 1981, ZMA unreg., 2
specs; Trincomalee, 1988, ex coll. Jan Berkhout,
GCK 5692, 1 spec.; Trincomalee, 1988, GCK 1398,
1 spec.; Trincomalee, 1990, in weedy sand, leg.
Kalika K. Perera, don. Richard J. Kelly, GCK 5132,
1 spec.. Thailand, Andaman Sea: Phuket area,
local shops, VL 900-16, 8 specs; Koh Pee-Pee, by
fisherman VL 900-17, 6 specs; Krabi province, Koh
Lipe, beached, VL 900-18, 1 spec.; Racha Island,
1995, HD 11809, 1 spec.; Phuket, ex coll. S. Martin,
1990, ZMA unreg., 1 spec.; Phuket, Rawai beach,
1995, HD 11810, 16 specs; Phuket, Maiton Isl., JW
03.540(2), 1 spec.; Phuket, near Rawai, JW 03.540(4),
1 spec.; Phuket, near Rawai, JW 03.540(5), 1 spec.;.

GC KRONENBERG, V LIVERANI & H DEKKER
662
Figs 1-8, 13 Conomurex decorus (Röding, 1798). 9-12, 14-18. Conomurex coniformis (Sowerby II, 1842). 1-2 Original
figures of Lambis decora Röding from Chemnitz, 1788, Conchylien Cabinet: pl. 157 figs 1499-1500. 3-4 Madagascar,
Tuléar, Anakao, 57 mm, VL 900-09. 5-7,13 Madagascar, Tuléar, Anakao, 54 mm, VL 900-10. 8 Juvenile, Tanzania,
Zanzibar, Nungwi, beached, 40 mm, VL 900-15. 9-10 Original figures of Strombus coniformis from Sowerby, Thesaurus
Conchyliorum: pl. 7 figs 55,61. 11-12 Jemen, Socotra, 32 mm, VL 905-05. 14-17 Somalia, Cape Guardafui, shallow
water, 49 mm, VL 905-04. 18 Juvenile, Somalia, Brava, shallow water among seagrass, 37 mm, VL 905-03.

IDENTITY OF STROMBUS CONIFORMIS
663
Malaysia, west coast: Pulau Pangkor, 1971, ex coll.
Jan Berkhout, GCK 5689, 1 spec.; Indonesia: ZMA
unreg., 1 spec.; Sumatera, Sabang, leg. C.H.J. van
Benthem, 1920, ZMA unreg., 4 specs.
Conomurex coniformis (Sowerby II, 1842)
(Figs 9-12, 14-18)
Strombus coniformis Sowerby II, 1842: 29, pl. 7 figs
55, 61
Strombus coniformis Kiener, 1843: 36, pl. 19 fig. 2
Strombus coniformis [in part] Tryon, 1885: 122
(as synonym of S. decorus), pl. 8 fig 90 (a copy of
Sowerby’s fig. 55)
? Strombus cylindricus Swainson & S. mauritiana
Lamarck Shopland, 1896: 225
? Strombus mauritianus var. coniformis Melvill &
Standen, 1901: 381
? Strombus cylindricus Swainson & S. mauritianus
Lamarck Shopland, 1902: 175
Strombus coniformis [in part] Abbott, 1960: 138
(listed in synonymy of S. decorus as knobbed
form)
Strombus coniformis [in part] Walls, 1980: 188 (in
synonymy index as synonym of S. decorus)
Strombus conformis [sic; in part] Kronenberg &
Berkhout, 1984: 295 (as synonym of S. decorus)
Strombus decorus Wranik & Saad, 1992: 60
Strombus decorus masirensis Moolenbeek &
Dekker, 1993: 6-7, 9 figs 9-13
Strombus decorus masirensis Samaie, 1997: 107-
108, colour photo
Strombus decorus Wranik, 1998: 179
Strombus decorus masirensis De Turck et al., 1999:
53, pl. 114 figs 2-4 (not pl. 114 fig. 1= C. decorus
knobbed form)
Strombus decorus Wranik, 2004: pl. 204 fig. c
Conomurex decorus masirensis Heiman et al., 2007:
figs 5, 6
Not “form coniformis” Heiman et al. 2007: table
1, figs 24, 25 = Conomurex persicus
Type material Syntypes of Strombus coniformis
are considered lost (see below), Sowerby (1842)
did not indicate a locality for Strombus coniformis.
Holotype of Strombus (Conomurex) decorus masi-
rensis is ZMA Moll. 3.92.029, the type locality is
Oman, Masirah Island (Moolenbeek & Dekker
1993).
Sowerby’s original description (1842: 29) reads
(translated): “Shell cone-shaped, smooth, palely
yellowish-brown, brown bands interrupted
encircled, strongly angled; extremely short spire;
whorls plicate at shoulder; narrowed aperture,
external lip barely expanded, with small sinus.”.
This description could refer to a number of
species of Conomurex. Some confusion may
arise as Sowerby, in the additional remarks
(in English), wrote down the phrase: “(…)
absence of the posterior sinus (…)”, which
contradicts the “small sinus” in the descrip-
tion.
Sowerby (1842: pl. 7 figs 55, 61) illustrated two
specimens from the collections of F.J. Stainforth
and of T. Norris. These figured shells are to
be considered syntypes of Strombus coniformis.
Both collections were sold at auctions, the first
about 1850, the second in 1873 (Dance, 1986:
166, 169). None of these syntypes are present in
the Natural History Museum, London (Kathie
Way, pers. comm. to GCK, Feb. 2003), nor in the
National Museum of Wales, Cardiff; National
Museum of Scotland, Edinburgh; nor the muse-
ums of Manchester or Leeds (Harriet Wood,
pers. comm. to GCK, Feb.-March 2003) and are
presumed lost. No type locality was given by
Sowerby (1842: 29).
Sowerby’s (1842) illustrations are here repro-
duced (Figs 9, 10). Figure 9, (Sowerby’s fig. 55)
shows a dorsal view, Fig. 10 (his fig. 61) shows
an apertural view. Both specimens show (mark-
edly less conspicuous in fig. 55) the so-called
strombid notch. The presence or absence of a
posterior sinus cannot be established in fig. 55,
while in fig. 61 there is no posterior sinus vis-
ible, but an adapical extension of the outer lip,
forming a gutter-like posterior canal against the
spire, is present.
It is evident that Sowerby distinguished S. coni-
formis from S. decorus (as S. cylindricus Swainson)
since he mentions this latter species (Sowerby,
1842: 29) and illustrates it on the same plate (pl.
7 figs 50, 57, 59). The specimen illustrated on pl.
7 fig. 50 is particularly interesting, as it repre-
sents a white specimen of the knobbed form of
Conomurex decorus, see also further below.
Kiener (1843) considered S. coniformis a spe-
cies different from S. decorus, judging from his
comment (1843: 36) “... des caractères distinctifs,
nous pensons qu’elle doit être conservée.”.
Moolenbeek & Dekker (1993) made an
extensive description of S. decorus masirensis,
based especially on the holotype of that spe-

GC KRONENBERG, V LIVERANI & H DEKKER
664
cies. Sowerby’s fig. 61 shows a specimen that
accommodates the differences as mentioned by
Moolenbeek & Dekker (1993: 7), viz. it shows the
tongue-like flap (forming the gutter-like posterior
canal), and a reddish colour on the columella.
Comparison of the figures of Sowerby (1842)
with the description by Moolenbeek & Dekker
(1993) and comparison of the material available
have convinced us that these two taxa are syn-
onymous. Strombus decorus masirensis is a junior
synonym of S. coniformis. The holotype of S.
decorus masirensis shows a strong resemblance in
colour pattern to one of Sowerby’s (1842: pl. 7 fig.
55) figured specimens. To establish the identity of
S. coniformis unequivocally, we hereby designate
the holotype of Strombus decorus masirensis as
neotype of Strombus coniformis. Besides the holo-
type and 36 paratypes present in ZMA and the
paratype present in HD (Moolenbeek & Dekker,
1993) of Strombus decorus masirensis, we studied
the following material of Conomurex coniformis:
Material examined Unknown: ex Jousseaume,
1916, MNHN unreg., 1 spec. (in sample with 1
C. decorus knobbed form). Oman: Mirbat, Jazirat
Hino, leg. R.G. Moolenbeek & P.L. van Pel, 1996,
ZMA unreg., 2 specs; Masirah, leg. J. Bryan, 1998,
ZMA unreg., 14 specs; Masirah, leg. D.T. Bosch,
1992, ZMA unreg., 7 specs; Masirah, Ras Radum,
leg. M. Day, 1997, ZMA unreg., 1 spec.; Masirah,
2 km S of Haql, leg. R.G. Moolenbeek & P.L. van
Pel, 1995, ZMA unreg., 4 specs; Masirah, behind
RAF station, leg. R.G. Moolenbeek & P.L. van Pel,
1997, ZMA unreg., 1 spec.; Masirah, Ras ad Dan,
leg. R.G. Moolenbeek & P.L. van Pel, 1995, ZMA
unreg., 1 spec.; Masirah, E coast, leg. M. Day, 1995,
ZMA unreg., 2 specs; Masirah, BERS station, leg.
R.G. Moolenbeek & P.L. van Pel, ZMA unreg.,
3 specs; Masirah JW 03.542(1), 1 spec.; Gulf of
Oman, SE Arabia, 1983 GCK 706, 2 specs; Gulf of
Oman, SE Arabia, GCK 779, 1 spec.; Masirah Isl.,
ex Jan Berkhout, GCK 5697, 1 spec.. Yemen: 120
km W of Mukallah, leg. C.A. Wagenvoort, 1993,
ZMA inreg., 1 spec.; al-Mahrah, Ra’s Darjah, W.
side of cape. 15°27’31”N-051°48’35”E, beached,
HD 9314, 1 spec.; al-Mahrah, Tabut, 5 km S of vil-
lage, 15°54’37”N-052°09’49”E, beached, HD 9313, 1
spec.; al-Mahrah, Musayna’ah, 12 km E of village,
15°05’32”N-050°48’43”E, beached, HD 9315, 1
spec.; Aden area, HD 8825, 3 specs; Socotra Island,
coll. Petruccioli, Rome, 3 specs; Socotra Island,
ex Petruccioli, coll. Pirazzini, Lugo, Italy, 1 spec.;
Socotra Island, leg. Lavranos, 1967, MNHN unreg.,
15 specs; Socotra, Abd el Kuri isl., legit Lavranos,
1967, MNHN, unreg., 6 specs. Somalia: near
Mugdiisho (Mogadiscio) JW 03.542(2), 1 spec.; near
Mugdiisho JW 03.542(3), 1 spec.; near Mugdiisho
JW 03.542(4), 2 specs; Brava, 1984, by snorkelling
VL 905-03, 1 juvenile; Somalia?, from Somali fisher-
men, deep water?, VL 905-04, 14 specs; from Somali
fishermen, shallow water, VL 905-05, 1 spec.; HD
8921, 2 specs; Kisimaio, collected in sand at 12 m.
depth, HD 8881, 2 specs ; Bagiuni islands, beach,
1970, MF unreg., 1 spec.. Tanzania: Zanzibar, Ras
Nungwi, 1998, VL 905-01, 1 spec.; Kiwengwa
beach, Bravo Club hotel, by snorkelling, ix.2002,
character
C. persicus C. decorus C. coniformis
shoulder knobs on
last whorl
absent sometimes present always present
posterior sinus deep deep shallow
adapical lip extension absent absent present
columellar callus
thin to moderately
thick
very thin very thin
colour columellar
callus
white
pattern of outer shell
shining through, base
white
pattern of outer shell
shining through, base
orange
colour aperture white to pale pink
white border, pink to
orange red inside
rosy border, white
inside
Table 1 Differences in shell characters between Conomurex persicus, C. decorus and C. coniformis

IDENTITY OF STROMBUS CONIFORMIS
665
VL 905-02, 1 spec.. Madagascar: Mahajanga bay,
beached, GB 459, 1 spec.; Anakao (Tulear), live in
shallow water, GB unreg., 1 spec..
DISCUSSION
Conomurex coniformis was synonymised with
Conomurex decorus (within the genus Strombus)
by Abbott (1960), in spite of some outstanding
characters of C. coniformis shown by the original
illustrations, which distinguish this species from
C. decorus.
Conomurex decorus masirensis was differenti-
ated by Moolenbeek & Dekker (1993: 7) from
C. decorus by two constant characters, viz. the
presence of the outer lip posterior extension,
the “tongue-like flap” as Moolenbeek & Dekker
called it, which is absent in C. decorus, and the
colouring of the columella, red to orange in C.
masirensis (Figs 10, 12, 16-18), but no statement
was made about the colouring of the colume-
lla in C. decorus. The columella of C. decorus is
always white (Figs 2, 4, 6-8). The holotype of C. d.
masirensis (Moolenbeek & Dekker, 1993: 5 fig. 2),
when viewed from the apex, clearly shows a pos-
terior sinus. This posterior sinus is often much
less distinct in other specimens, and is generally
Fig 19 Map of the Indian Ocean with the distribution of Conomurex species. + = C. coniformis, • = C. decorus. Note
the exclusive presence of C. coniformis along southern Arabia.

GC KRONENBERG, V LIVERANI & H DEKKER
666
(although not always) deeper and u-shaped in
C. decorus (Figs 13, 14). As already mentioned by
Moolenbeek & Dekker (1993: 7), shoulder knobs,
especially on the last half of the body whorl, are
present in C. coniformis, but may also be present
in specimens of C. decorus, and this feature is
therefore of little use in distinguishing both spe-
cies (Figs 5-7, 13). To the differences mentioned
above we may add that the colouring of the aper-
ture is reversed in the two species: in C. decorus it
is pinkish/red deep within, with a white border,
while in fresh and adult C. coniformis it is white
inside with light pink-orange towards the rim of
the outer lip. This difference is even present in
juvenile specimens (Figs 8, 18). Differences we
observed are summarised in Table 1.
Based on the constant differences, the sympatric
occurrence in a part of the areas of distribution of both
taxa, without specimens with intermediate characters,
we conclude that these taxa should be separated as
distinct species in the genus Conomurex.
REMARKS
The ranges of both taxa are distinct, but over-
lap in Eastern Africa (see material examined,
and Fig. 19). The finding of C. decorus at Brava,
Somalia confirms the occurrence of this species
in Somalia, which was reported from Merca by
Borri et al. (2002: 21).
The Persian Gulf record of C. decorus in the
GCK-collection (see specimens examined) is
doubtful and needs confirmation, as no speci-
mens of C. decorus are reported with certainty
from that area (Moolenbeek & Dekker, 1993).
However, Melvill & Standen (1901) recorded
Strombus mauritianus var. coniformis from the
Persian Gulf, which they regard as a nodular
form of S. mauritianus (= C. decorus). The identity
of these noduled specimens, as well as the typical
specimens, should be re-examined.
Samaie (1997) recorded specimens of C. coni-
formis from Ramin, Iran (just east of Cah Bahar),
close to the Pakistani border. Mr. Moolenbeek
identified these specimens. This record is the
most eastern locality we know of.
The Aden record in the HD collection is of
shells bought from a local shell shop in Aden.
Shopland (1896, 1902) recorded both Strombus
cylindricus Swainson and S. mauritianus Lamarck
from Aden; both names are now considered syn-
onyms of C. decorus (Abbott, 1960: 138). Although
Shopland used two names to denote his samples,
it seems likely that Shopland had in fact only
specimens of C. coniformis before him.
Conomurex coniformis is the only species of
Conomurex recorded from southern Oman
and Yemen. The finding of a good number of
specimens at Socotra (MNHN and Petruccioli)
indicates that viable populations exist there,
confirmed by the records of Wranik (2004) and
F.G. de Ceuninck van Capelle, Leiden, The
Netherlands (pers. comm., HD). The finding of
C. coniformis in southern Somalia, Zanzibar, and
Madagascar, however, is no evidence that viable
populations of that species actually live there.
But it is at least an indication that larvae can
be dispersed by ocean currents in a southward
direction (although such dispersal would be
opposed by the prevailing East African Coastal
Current), and, therefore that C. coniformis may
establish itself along the East African coast.
The name Strombus decorus fa. conospira in
Moolenbeek & Dekker (1993: caption fig. 15) is an
error for Strombus decorus fa. coniformis. The name
Strombus conospira Moolenbeek & Dekker, 1993 is
not available as it appears as both an infrasub-
specific name (ICZN art. 10.2) and without a
description or diagnosis (ICZN art. 13).
NOTE ADDED IN PROOF
Bandel (2007: 148) described the subgenus
Decoristrombus (Type species by Original des-
ignation Strombus fasciatus Born, 1778, endemic
to the Red Sea and possibly the adjacent part of
the Indian Ocean), differentiating Decoristrombus
from Conomurex only by differences between the
two type species, viz. that that “Strombus fasciatus
resembles Strombus (Conomurex) luhuanus but has
the corner of its body whorl ornamented by short
ribs or spikes” (Bandel, 2007: 149). Bandel (2007)
also allocated S. decorus to his new genus, retain-
ing S. persicus in Conomurex.
In Conomurex fasciatus (Born, 1778) the shoulder
knobs may vary from distinct, pointed knobs to
almost indiscernible (the so called forma elegans).
As shown in this paper, both C. decorus and C.
coniformis may have distinct shoulder knobs and
C. decorus may have a completely smooth shoul-
der. The type species of Conomurex, Strombus
luhuanus, usually has a smooth shoulder on the
last whorl, but specimens that have low knobs on
the shoulder of the last whorl are known.

IDENTITY OF STROMBUS CONIFORMIS
667
As there are no other differences in overall
shape, nor in structure of the outer lip, see
Kronenberg & Vermeij (2002) for a preliminary
discussion on characters of the outer lip, and no
biogeographical separation we see no reason to
separate Decoristrombus from Conomurex, and
therefore consider Decoristrombus a junior syno-
nym of Conomurex.
ACKNOWLEDGEMENTS
We would like to thank Mr. Pasquale Fazzini
(Forlì, Italy) for the donation of specimens to the
second author, and photography; Mr. Luciano
Petruccioli (Rome, Italy) for information regard-
ing the occurrence of Conomurex coniformis at
Socotra; Dr. Theo C.M. Kemperman (Amsterdam,
The Netherlands) for help with the translation of
the original description of Strombus coniformis;
Dr. Philippe Bouchet (MNHN) for admittance
to the collection and making valuable remarks
on an earlier draft of this paper; Ms. Katie Way
(The Natural History Museum, London, United
Kingdom) and Ms. Harriet Wood (National
Museum of Wales, Cardiff, United Kingdom) for
checking the collections under their custody and
making further inquiries; Mr. Robert Moolenbeek
(ZMA) for calling our attention to the work of
Samaie and his kind hospitality; Mr. Jeroen Goud
(RMNH) for kind hospitality; Mr. Gianluigi Bini
and Mr. Maurizio Forli for allowing the second
author to examine their collections; Mr. Joop
Wiersma, Kerkrade, The Netherlands, for show-
ing his collection to the first author; Dr. David
Reid, The Natural History Museum for reading
the final draft. GCK wants to thank Ms. Marianne
Matthijssen for her lasting support.
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