Description of eight new species and one new genus of Muricidae (Gastropoda) from the Indo-West Pacific

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Description of eight new species and one new genus of
Muricidae (Gastropoda) from the Indo-West Pacific
Roland HOUART
Research associate
Institut royal des Sciences naturelles de Belgique
and
Muséum national d'Histoire naturelle, Paris, France
UMR7205 ISyEB
roland.houart@skynet.be
KEYWORDS. Gastropoda, Muricidae, new genus, new species, neotype, lectotype, Indo-West
Pacific.
ABSTRACT. Eight new species of Muricidae are described from Mozambique, South Africa and
Papua New Guinea. A new genus, Tylothais n. gen. is described for the species formerly included
in Thalessa H. & A. Adams, 1853, a subjective junior synonym of Volema Röding, 1798, a genus
of Melongenidae. A neotype is designated for both Murex plicatus Gmelin, 1791 and Murex
virgatus Dillwyn, 1817 and a lectotype is designated for Purpura distinguenda Dunker, 1866.
Five subfamilies are represented in this paper: Muricinae (Aspella aclydis n. sp., from Papua New
Guinea), Ergalataxinae (Spinidrupa aethes n. sp., from Papua New Guinea), Trophoninae
(Leptotrophon fusiformis n. sp., from the Solomon Sea), Typhinae (Laevityphis libos n. sp., from
Papua New Guinea) and Rapaninae (Tylothais n. gen. akidotos n. sp., T. funilata n. sp., T.
inhacaensis n. sp. and T. ovata n. sp., from Mozambique and South Africa).
RESUME. Huit nouvelles espèces de Muricidae sont décrites du Mozambique, d'Afrique du Sud
et de Papouasie-Nouvelle-Guinée. Un nouveau genre, Tylothais n. gen., est décrit pour les espèces
auparavant incluses dans Thalessa H. & A. Adams, 1853, synonyme subjectif plus récent de
Volema Röding, 1798, un genre appartenant aux Melongenidae. Un néotype est désigné à la fois
pour Murex plicatus Gmelin, 1791 et Murex virgatus Dillwyn, 1817 et un lectotype est désigné
pour Purpura distinguenda Dunker, 1866.
Cinq sous-familles sont représentées dans cet article: Muricinae (Aspella aclydis n. sp. de
Papouasie-Nouvelle-Guinée), Ergalataxinae (Spinidrupa aethes n. sp. de Papouasie-Nouvelle-
Guinée), Trophoninae (Leptotrophon fusiformis n. sp. de la mer des Salomon), Typhinae
(Laevityphis libos n. sp. de Papouasie-Nouvelle-Guinée) et Rapaninae (Tylothais n. gen. akidotos
n. sp., T. funilata n. sp., T. inhacaensis n. sp. et T. ovata n. sp. du Mozambique et d'Afrique du
Sud).
INTRODUCTION
New species of Trophoninae from Papua New
Guinea have been recently described from deep water
by Houart & Héros (2016) from the BIOPAPUA and
NIUGINI campaigns. Two additional species of
Muricidae are here described from those same
expeditions. One additional shallow water species
from Papua New Guinea is described from the
KAVIENG campaign and a deep water trophonine is
described from the Solomon Sea, from the MADEEP
campaign. Species of Muricidae from the western
Indian Ocean were also described by Houart & Héros
(2013 and 2015). The species described in this paper
from the western Indian Ocean all belong to
Rapaninae, originate from Mozambique and South
Africa and are shallow water or coastal species,
mainly found in sand and on rocks at low tide.
Several studies pertaining to the new classification of
Muricidae, based on genetic results were published
recently. Six of those cover Indo-West Pacific species
or genera that are included here: Claremont et al.,
2008 (Ergalataxinae and Rapaninae); Barco et al.,
2010 (overall classification); Barco et al., 2012
(Pagodulinae and Trophoninae); Claremont et al.,
2013a (Ergalataxinae); Claremont et al., 2013b
(Rapaninae) and Barco et al., 2015 (Haustrinae and
Pagodulinae).
Material and methods
Material
The material studied here originates from expeditions
in the West Pacific, in the vicinity of Papua New
Guinea and the Solomon Islands and in the Western
Indian Ocean, from Inhaca Island, Mozambique. Other
material originates from the Natal Museum,
Pietermaritzburg, South Africa and from the author's
personal collection.
The following expeditions of MNHN/IRD are
involved:

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BIOPAPUA: from August 22 to October 18, 2010.
This expedition explored parts of the Bismarck Sea
and of the Solomon Sea (see also Houart & Héros,
2016).
INHACA: from November 23 to December 12, 2011.
The island of Inhaca (26°S, 33°E) is about thirty
kilometers off the continent. It extends from the
peninsula of Machangulo towards the north and is
separated from it by the strait of Ponta Torres. The
island measures 12 km in its greatest length from
north to south and less than 7 km from east to west. It
consists mainly of an extremely fragile sand dune. The
few underlying rocks encountered around the island
are made up of beach-rock sand dating from the late
Pleistocene glaciation. At low tide, the exposed areas
cover an area larger than that of the island (P.
Maestrati, unpublished report).
PAPUA NIUGINI: from October 25 to December 26,
2012. Intensive study of the Madang region, with
MADANG, a deep-sea cruise off the Bismarck Sea
coast, from Vitiaz Strait to the border with West
Papua (Irian Jaya) (see also Houart & Héros, 2016).
MADEEP: from April 6 to May 8, 2014 in Papua New
Guinea. This deep-sea cruise was dedicated to the
study of the deep sea biodiversity in the Bismarck and
Solomon seas, in continuation of the PAPUA
NUIGINI project.
KAVIENG: from June 1 to 30 (Kavieng Lagoon
Biodiversity Survey) and from August 27 to
September 7, 2014 (deep water). During the lagoon
survey, a total of 404 stations were surveyed covering
the entire Kavieng lagoon between 0 and 45 meters.
The PAPUA NIUGINI, MADEEP and KAVIENG
expeditions were part of the Our Planet Reviewed
program, conducted by the National Museum of
Natural History (MNHN), Pro-Natura Internation
(PNI), the Institute for Research for Development
(IRD), and their in-country scientific partners, the
University of Papua New Guinea (UPNG) and the
National Fisheries Authority.
Methods
The characters used to describe the shell morphology
are the general aspect of the shell, its shape and size,
colour, shape of the spire and number of protoconch
and teleoconch whorls, features of the protoconch,
shape of the teleoconch whorls and features or form of
the suture and of the subsutural ramp, of axial and
spiral sculpture, the aperture and siphonal canal. When
known, the characters of the operculum and radula are
also used. Unless otherwise mentioned, the species
descriptions are based on the holotype and the
paratypes.
The morphology of the radula is described starting
from the rachidian tooth followed by the lateral teeth.
The method for determining diameter, height and
counting the number of protoconch whorls is
explained in Fig. 1.
The bathymetric range given here is provided using
the inner values of the recorded depth: the largest
value of the minimum values and the lowest value of
the maximum values of all the recorded ranges.
Fig. 1. Method for determining diameter, height and counting the number of protoconch whorls (scale bars: 500
µm).
Abbreviations
Repository
IRSNB: Institut royal des Sciences naturelles de
Belgique, Bruxelles, Belgium.
MNHN: Muséum national d'Histoire naturelle, Paris,
France.
NMSA: The KwaZulu-Natal Museum,
Pietermaritzburg, South Africa.
RH: collection of Roland Houart.
NHMD: Natural History Museum of Denmark
(Zoology), Copenhagen.
ZMB: Museum für Naturkunde der Humboldt
Universität zu Berlin, Zoologisches Museum,
Germany.
Other abbreviations
CP: chalut à perche (beam trawl)

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DW: drague Warén (Warén dredge)
lv: live collected specimen(s)
dd: empty shell(s).
Terminology used to describe the radula (Fig. 2)
cc: central cusp
ild: inner lateral denticle
lc: lateral cusp
LT: lateral teeth
mc: marginal cusp
md: marginal denticles.
Terminology used to describe the spiral cords and
the apertural denticles (after Merle 2001 and 2005)
(Fig. 3). Terminology in parentheses: variable
feature.
Convex part of teleoconch whorl and siphonal canal
ab: abapical (or abapertural);
abis: abapical infrasutural secondary cord (on
subsutural ramp);
ABP: abapertural primary cord on the siphonal canal;
ad: adapical (or adapertural);
adis: adapical infrasutural secondary cord (on
subsutural ramp);
ADP: adapertural primary cord on the siphonal canal;
ads: adapertural secondary cord on the siphonal canal;
IP: infrasutural primary cord (primary cord on
subsutural ramp);
MP: median primary cord on the siphonal canal;
ms: median secondary cord on the siphonal canal;
P: primary cord;
P1: shoulder cord;
P2-P6: primary cords of the convex part of the
teleoconch whorl;
s: secondary cord;
s1-s6: secondary cords of the convex part of the
teleoconch whorl (example: s1 = secondary cord
between P1 and P2; s2 = secondary cord between P2
and P3, etc.);
SP: subsutural cord.
Aperture
D1 to D5: abapical denticles;
ID: Infrasutural denticle.
Fig. 2. Radula of Tylothais akidotos n. sp.
Family MURICIDAE Rafinesque, 1815
Subfamily MURICINAE Rafinesque, 1815
Genus Aspella Morch, 1857
Type species by monotypy: Ranella anceps Lamarck,
1822, Mediterranean.
Aspella aclydis n. sp.
Figs 3A, 5A-G, L, O
Type material. Holotype MNHN IM-2013-54542; 1
paratype MNHN IM-2013-53620; 3 paratypes Reg.
No. MSF mx214 (a, b, c), Molluscan Science
Foundation, Inc., Owings Mills, Maryland, USA, 1
paratype RH (as listed below).
Material examined. KAVIENG, stn KB48, type
locality, 18 June 2014, 1 lv (holotype); stn KZ02,
Kavieng Lagoon, Edamago Id, 02°37' S, 150°44' E, 9
m, soft corals, rocks, sand, 1 lv (paratype MNHN IM-
2013-53620); Albatross Passage east of Manne Island,
Kavieng area, 02°45' S 150°43' E, with grit at 30-40 m
(3 paratypes Molluscan Science Foundation, Inc.).
Type locality. Papua New Guinea, Kavieng Lagoon,
west side of Tsoilaunung Island, 02°33' S, 150°31' E,
6 m, coral block in middle of sand.
Distribution. Papua New Guinea, Kavieng, living at
6-30 m.
Description. Shell large for the genus, 21.1 mm in
length (paratype MSF mx214, Molluscan Science
Foundation, Inc.). Length/width ratio 2.3-2.5. Slender,

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lanceolate, flattened, smooth. Subsutural ramp very
narrow, almost horizontal, strongly concave. Shell
entirely covered by a thick, dirty white intritacalx.
Aperture white.
Spire very high with 2.15 protoconch whorls and 8
elongate, flattened, weakly shouldered teleoconch
whorls. Suture strongly impressed, ventrally and
dorsally partially obscured by broad buttress
connecting preceding whorl. Protoconch (Fig. 5O)
small, conical. Whorls smooth, glossy, penultimate
whorl with narrow keel abapically. Maximum width
500 µm, height 550 µm. Terminal lip erect, of
sinusigera type.
Axial sculpture of teleoconch whorls consisting of
high and low, narrow, lightly nodose varices and low
intervariceal ribs. First and second whorl with 6
narrow, low ribs; third whorl starting two high,
expanded, lateral varices, one additional low varix
between lateral varices and a low intervariceal rib;
fourth to last whorl with two high, expanded, narrow,
lateral varices, two lower varices and two broad, low
buttresses connecting preceding whorl. Spiral
sculpture almost indistinct, consisting of P1-P6, ADP,
MP and ABP (Fig. 3A), only distinct on lateral varices
covered by thick, very weakly axially and spirally
striate, intritacalx (Fig. 5L).
Aperture narrow, ovate. Columellar lip narrow,
smooth, adherent. Anal notch indistinct. Outer lip
weakly erect, smooth with 5 obvious denticles within:
D1-D5. Siphonal canal short, straight, narrowly open.
Operculum (Fig. 5G) dark brown, ovate with apical
nucleus. Radula unknown.
Remarks. There are 12 extant species of Aspella
known from the Indo-West Pacific but only two with a
conical, multispiral protoconch, attesting to a
planktotrophic larval development, Aspella hildrunae
Houart & Tröndlé, 2008 from French Polynesia and A.
producta (Pease, 1861), living throughout the Indo-
West Pacific. All the other species have a rounded,
paucispiral protoconch of 1.5 whorls.
Aspella hildrunae (Figs 5H-J, M, P) differs in having
a reticulate, strongly spirally and axially striate
intritacalx (Fig. 5M), a broader shell with a lower
spire, a comparatively broader aperture, narrower but
more obvious buttresses and a strongly dorsally bent
siphonal canal.
Aspella producta (Fig. 5K, N) differs in many ways,
having a comparatively broader and larger, strongly
nodose shell, reaching 23 mm in length. It also differs
in having a thick, strongly cancellate intritacalx (Fig.
N).
In total three species of Aspella were collected in
Papua New Guinea during the KAVIENG expedition:
A. producta, A. media Houart, 1987 (Fig. 5Q) and
Aspella aclydis n. sp.
Etymology. Aclydis (L): small javelin or spear, refers
to the lanceolate form of the shell.
Subfamily Ergalataxinae Kuroda, Habe & Oyama,
1971
Genus Spinidrupa Habe & Kosuge, 1966
Type species by original designation: Murex
euracanthus A. Adams, 1853, Indo-West Pacific.
Spinidrupa aethes n. sp.
Figs 3B-C, 6A-D
Type material. Holotype MNHN IM-2000-32835 and
1 paratype MNHN IM-2000-32836 (as listed below).
Material examined. PAPUA NIUGINI, stn PD54,
type locality, 2 dd (holotype and paratype MNHN).
Type locality. Papua New Guinea, Madang Lagoon,
West of Panab Island, 05°11' S, 145°48' E, 15 m.
Distribution. Only known from the type material.
Figure 3. Spiral cords and apertural denticles morphology
A. Aspella aclydis n. sp. Papua New Guinea, Kavieng Lagoon, west side of Tsoilaunung Island, 02°33' S,
150°31' E, 6 m, holotype MNHN IM-2013-54542.
B-C. Spinidrupa aethes n. sp. Papua New Guinea, West of Panab Island, 05°11' S, 145°48' E, 15 m. B. Paratype
MNHN IM-2000-32836; C. Holotype MNHN IM-2013-32835.
D. Leptotrophon fusiformis n. sp. Solomon Sea, Budibudi Island, north of Archipel Laughlan Islands, 09°11' S,
153°55' E, 380-411 m, holotype MNHN IM-2013-45607.
E. Tylothais akidotos n. sp. Inhaca Island, Mozambique, Maputo Bay, Xixuana, 25°60' S, 32°56' E, 0-1 m,
holotype MNHN IM-2013-43791.
F-G. Tylothais funilata n. sp.
F. Mozambique, south of Vilanculos, mangroves, paratype NM G1300/T4222; G. Holotype NM L2726/T4221.
H. Tylothais inhacaensis n. sp. Inhaca Island, Mozambique, Ponta do Farol, 25°58 S, 32°59' E, holotype MNHN
IM-2000-32843.
I-J. Tylothais ovata n. sp.
I. Paratype RH; J. Holotype NM P0992/T4224.

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Description. Shell medium-sized for the genus,
holotype 19.8 mm in length. Length/width ratio 1.6.
Biconical, broadly ovate, heavy, spinose and
squamous. Subsutural ramp broad, strongly sloping,
strongly convex. Shell light tan with partially dark and
light brown coloured spines, primary cords and
siphonal canal. Aperture dirty white. Spire high.
Teleoconch of 7 angulate, strongly shouldered,
spinose whorls. Suture strongly adpressed. Protoconch
unknown, eroded in both specimens.
Axial sculpture of teleoconch whorls consisting of
moderately high, strong, broad ribs and varices. Other
axial sculpture of numerous, weak, growth lamellae.
First and second teleoconch whorls with 10 ribs, third
and fourth with 9, fourth and fifth whorls with 8
varices, last with 7 varices. Spiral sculpture of high,
strong, narrow, squamous, primary, secondary and
tertiary cords (Fig. 3B). Last teleoconch whorl with
high, narrow P1, P3, P5 and ADP; P2, P4 and P6 low,
narrow. Tertiary cords lower and narrower, of equal
strength. P1 and P3 highest, P6 narrow and low.
Previous teleoconch whorls with visible P1, s1; P2
visible from antepenultimate whorl. Subsutural ramp
with 3 or 4 low, narrow, squamous threads of same
strength.
Aperture large, narrow, ovate. Columellar lip narrow,
smooth in juvenile paratype, Columellar lip of
holotype with 3 elongate, weak knobs, decreasing in
strength abapically and weak, low parietal knob at
adapical extremity. Rim adherent. Anal notch deep,
broad. Outer lip with 6 strong, moderately high,
narrow denticles within: ID, D1-D5 (Fig. 3C),
extended as narrow lirae into aperture. Siphonal canal
short, broad, very weakly dorsally recurved, broadly
open, with ADP and MP.
Operculum and radula unknown.
Figure 4
A-B. Laevityphis coronarius Deshayes, 1865. Barisis-au-Bois, Paris Basin, Lower Eocene, France, RH, 6 mm.
C-D. Siphonochelus arcuatus (Hinds, 1843). Algoa Bay, South Africa, live in crayfish trap, 100 m, RH, 12.8
mm (type species of Siphonochelus).
(SV: succeeding varix; AT: anal tube; PV: preceding varix)
Figure 5 (scale bars 500 µm)
A-G, L, O. Aspella aclydis n. sp.
A-C, G, L, O. Papua New Guinea, Kavieng Lagoon, west side of Tsoilaunung Island, 02°33' S, 150°31' E, 6 m,
coral block in middle of sand (all holotype); A-C. Holotype MNHN IM-2013-54542; G. Operculum; L. Detail of
intritacalx; O. Protoconch.
D-F. Papua New Guinea, Kavieng, Albatross Passage east of Manne Island, Kavieng area, 2°45'S 150°43'E, with
grit at 30-40 m, paratype MSF mx214, 21.1 mm.
H-J, M, P. Aspella hildrunae Houart & Tröndlé, 2008
H-J. French Polynesia, Society Archipelago, Tahiti, Punaauia, Amiral reef, coral fragments, holotype MNHN
IM-2000-20165, 15.1 mm; M. French Polynesia, Society Archipelago, Huahine Island (RH), detail of intritacalx;
P. Protoconch (RH).
K, N. Aspella producta (Pease, 1868)
K. Kavieng Lagoon, Papua New Guinea, Northwest tip of Big Nusa Island, 02°34'S, 150°46,7'E, 6-20 m,
MNHN IM-2013-54683, 12.5 mm; N. Touho, New Caledonia (RH), detail of intritacalx.
Q. Aspella media Houart, 1988. Protoconch (RH).

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Remarks. The shell morphology of Spinidrupa aethes
n. sp. is particular in its spiny, narrow, high primary
cords and broad, elongate columellar denticles, close
to S. euracanthus (A. Adams, 1853) (Fig. 6E-F).
However, S. euracanthus has a comparatively broader
and a more rounded last teleoconch whorl, usually
with broader, more rounded primary cords, a quite
shorter, narrower siphonal canal and 2 or 3 small
nodes at abapical extremity of the columellar lip
instead of 3 broad elongate nodes almost all along the
columella in S. aethes n. sp. Spinidrupa infans (E. A.
Smith, 1884) (Fig. 6G) is a smaller species with the
same differences as between S. euracanthus and S.
aethes n. sp.
The genus Orania Pallary, 1900 was another
possibility for this new species. While the new species
seems closer to Spinidrupa it is instructive to compare
it with O. rosadoi Houart, 1998, O. fischeriana
(Tapparone Canefri, 1882) and O. rosea Houart, 1996.
Orania rosadoi also has a spiny shell but more
rounded and broader primary cords, a narrower
siphonal canal, a smooth columellar lip and a uniform
colour. Orania fischeriana also bears 3 elongate
denticles on the columellar lip but they are more
obvious and narrower, while the shell morphology
strongly differs from Spinidrupa aethes n. sp. Orania
rosea is a spiny shell but differs in many ways from
O. aethes n. sp. in having a uniform colour, a higher
spire, a broadly convex last teleoconch whorl, a
shorter and narrower siphonal canal and much
narrower elongate denticles on the abapical part of the
columellar lip. Lastly, due to the particular
morphology of the aperture the new species was also
compared with some buccinids, in particular with
some Clivipollia species. However some features such
as a strongly spiny shell and a straighter columellar
lip, less concave adapically, separate it from those
species.
Etymology. Aethes (G): unusual, strange, refers to its
particular morphology, unusual in Spinidrupa.
Subfamily Trophoninae Cossmann, 1903
Genus Leptotrophon Houart, 1995
Type species, by original designation: Leptotrophon
caroae Houart, 1995, New Caledonia.
Leptotrophon fusiformis n. sp.
Figs 3D, 6H-L
Type material. Holotype MNHN IM-2013-45607, 6
paratypes MNHN IM-2013-43633, IM-2013-43634,
IM-2013-43635, IM-2013-45608 and MNHN IM-
2013-45610, MNHN IM-2000-32837 and 1 paratype
coll. RH (as listed below).
Material examined. MADEEP, stn DW4285,
Solomon Sea, Budibudi Island, north of Laughlan
Archipelago , 09°11'S, 153°55'E, 380-411 m, 7 lv, 7
dd (holotype MNHN IM-2013-45607, 6 paratypes
MNHN IM-2013-436333, MNHN IM-2013-43634,
MNHN IM-2013-43635, MNHN IM-2013-45608,
MNHN IM-2013-45610, MNHN IM-2000-32837), 1
lv, 6 dd, (1 paratype RH, dd); stn DW4286, Budibudi
Island, north of Laughlan Archipelago, 09°12'S,
153°55'E, 306-365 m, MNHN IM-2013-4616, 1 lv;
stn DW4287, Budibudi Island, north of Laughlan
Archipelago, 09°12'S, 153°56'E, 340-375 m, MNHN
IM-2013-45623, 1 lv.
Type locality. Solomon Sea, Budibudi Island, north of
Laughlan Archipelago, 09°11'S, 153°55'E, 380-411 m.
Distribution. Solomon Sea, Budibudi Island, north of
Laughlan Archipelago, living at 365-380 m.
Description. Shell large for the genus, up to 16.2 mm
in length (holotype). Length/width ratio 2.2.
Lanceolate, broadly ovate, smooth, lightly built.
Subsutural ramp moderately broad, strongly sloping,
straight or weakly convex. Shell entirely white.
Spire high with 1.5 protoconch whorls and teleoconch
up to 5 broad, convex, weakly shouldered whorls.
Suture weakly adpressed. Protoconch large, broad,
whorls rounded, smooth, glossy. Maximum width and
height 1000 µm. Terminal lip shallow, delicate, thin,
almost straight.
Figure 6 (scale bar 500 µm)
A-D. Spinidrupa aethes n. sp. Papua New Guinea, West of Panab Island, 05°11' S, 145°48' E, 15 m.
A-B. Holotype MNHN IM-2000-32835, 19.8 mm; C-D. Paratype MNHN IM-2000-32836, 16.4 mm.
E-F. Spinidrupa euracantha (A. Adams, 1853)
E. Gulf of Eilat, Red Sea, RH, 19.6 mm; F. Guam, Hapra Harbour, RH, 18.8 mm.
G. Spinidrupa infans (E. A. Smith, 1884). Pemba, Mozambique, RH, 15.8 mm.
H-L. Leptotrophon fusiformis n. sp. Solomon Sea, Budibudi Island, north of Archipel Laughlan Islands, 09°11'
S, 153°55' E, 380-411 m.
H-J. Holotype MNHN IM-2013-45607, 16.2 mm; J. Protoconch; K-L. Paratype MNHN IM-2013-43634, 14.6
mm.
M-N. Leptotrophon marshalli Houart, 1995. New Caledonia, Grand-Passage, 18°55' S 163°24' E, 460 m,
holotype MNHN IM-2000-199, photo M. Caballer 2014, 9 mm.
O-P. Leptotrophon surprisensis Houart, 1995. North of New Caledonia, 18°56' S, 163°22' E, 444-452 m, RH,
12.1 mm.

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Axial sculpture of teleoconch whorls consisting of
high, strong, narrow, rounded ribs and numerous,
minute, growth striae. First teleoconch whorl with 9
ribs, second and third with 8, penultimate with 8 or 9
ribs, last whorl with 7 or 8. Spiral sculpture of low,
rounded, narrow, barely visible primary and secondary
cords, more obvious on early whorls. Last teleoconch
whorl with adis, IP, abis, P1, s1, P2, s2, P3, s3, P4, s4,
P5, s5, P6, s6 and 3-5 cords on siphonal canal,
probably ADP, (ads), MP, (ms), (ABP).
Aperture large, broadly ovate. Columellar lip broad,
strongly flaring, smooth or with minute node
abapically. Rim partially erect, adapically adherent.
Anal notch shallow, broad. Outer lip smooth with very
low denticles or smooth within. Holotype with only
D1 visible. Siphonal canal short, narrow, very weakly
dorsally recurved, narrowly open.
Operculum light brown, thin, fragile, ovate, with
apical nucleus and numerous concentric ridges.
Radula unknown.
Remarks. Only two species, both from north New
Caledonia can be reasonably compared with
Leptotrophon fusiformis n. sp., all others are strongly
dissimilar.
Leptotrophon marshalli Houart, 1995 (Fig. 6M-N) is a
smaller species with less rounded whorls, narrower,
less numerous axial ribs and without spiral sculpture
except numerous, minute lirae all over the shell.
Leptotrophon surprisensis Houart, 1995 (Fig. 6O-P) is
also smaller, with an obviously smaller protoconch, a
broader last teleoconch whorl with a smaller aperture,
shell without obvious spiral sculpture except P1 with
small spinelets formed at the intersection with the
axial ribs.
Etymology. Fusiformis (L): Spindle-shaped, named
for its fusiform shape.
Subfamily Typhinae Cossmann, 1903
Genus Laevityphis Cossmann, 1903
Type species by original designation: Typhis
coronarius Deshayes, 1865 (= Typhis muticus J.
Sowerby, 1834). Lower Eocene, France and England.
DISCUSSION
The genus Laevityphis is morphologically close to
Siphonochelus Jousseaume, 1880 but differs
constantly from it in having the anal tube clearly
separated from the varix, nearer to the preceding one,
while in Siphonochelus the anal tube originates
directly from the axial varix or is directly connected to
it (Fig. 4A-D).
There are currently three species classified in
Laevityphis: L. bullisi (Gertman, 1969) from the
western Atlantic (Colombia), L. tillierae (Houart,
1986), from New Caledonia, and L. tubuliger (Thiele,
1925) from the Zanzibar Channel.
Typhis transcurrens Martens, 1902 was first included
in Laevityphis by Keen (1944) followed by Radwin &
D'Attilio (1976), by D'Attilio & Hertz (1988) and then
by Houart et al. (2011) but I do not agree any longer
because the anal tubes in T. transcurrens, strongly
expanded and flattened at the base, are directly
connected to the axial varices, typical for
Siphonochelus s.s.
Laevityphis libos n. sp.
Fig. 7A-G
Type material. Holotype MNHN IM-2013-58289, 7
paratypes MNHN IM-2000-32838 - IM-2000-32841
and 1 paratype RH (as listed below).
Material examined. BIOPAPUA, stn CP3709, off
Madang, 04°58'S, 145°52'E, 640-675 m, 1 lv
(paratype MNHN IM-2000-32838).
PAPUA NIUGINI, stn CP4033, Cape Croisiles,
04°52'S, 145°53'E, 780-780 m, 2 lv (paratypes MNHN
IM-2000-32839); stn CP4038, east Kotakot, 04°27'S,
145°34'E, 800-840 m, 4 lv (3 paratypes MNHN IM-
2000-32840, 1 paratype coll. RH).
KAVIENG, stn CP4436, New Ireland, 02°16'S,
150°45'E, 1128-1135 m, 1 lv (holotype MNHN IM-
2013-58289).
MADEEP, stn DW4318, Solomon Sea, off Marshall
Bennett Island, west Woodlark Island, 08°37'S,
151°47'E, 705-817 m, 1 lv (paratype MNHN-IM-
2000-32841).
Figure 7
A-G. Laevityphis libos n. sp.
A-C. Papua New Guinea, New Ireland, 02°16' S, 150°45' E, 1128-1135 m, holotype MNHN IM-2013-58289,
8.2 mm; D-G. Papua New Guinea, off Madang, 04°58' S, 145°52' E, 640-675 m, paratype MNHN IM-2000-
32838, 9.8 mm.
H-J. Laevityphis tillierae (Houart, 1985). South of New Caledonia, 825-860 m, RH, 7.3 mm.
K-M. Laevitiphis tubuliger (Thiele, 1925). Zanzibar Channel, 465 m, syntype ZMB 109314, 5.1 mm.
N-Q. Tylothais akidotos n. sp.
N-O. Inhaca Island, Mozambique, Maputo Bay, Xixuana, 25°60' S, 32°56' E, 0-1 m, holotype MNHN IM-2013-
43791, 33.7 mm; P. Maputo Bay, Xixuana, 25°60' S, 32°56' E, Inhaca Island, paratype MNHN IM-2013-43792,
34.3 mm; Q. Mozambique, Inhaca Island, Delagoa Bay, paratype NM 7346/T4220, 40.3 mm.

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Type locality. Papua New Guinea, New Ireland,
02°16'S, 150°45'E, 1128-1135 m.
Distribution. 02°16' - 08°37' S, 145°34' E - 150°45'
E, living at 780-1128 m.
Description. Shell small for the genus, up to 11 mm
in length (paratype MNHN IM-2000-32839).
Length/width ratio 2.1-2.2. Slender, lanceolate,
narrowly ovate. Subsutural ramp narrow, weakly
sloping, weakly concave. Shell entirely white. Spire
high with 1.5 protoconch whorls and teleoconch up to
5 narrow, weakly convex, strongly shouldered whorls.
Suture impressed. Protoconch large, broad, whorls
rounded, smooth. Maximum width 1000 µm, height
1100 µm. Terminal lip broken.
Axial sculpture of teleoconch whorls consisting of 4
low varices with rounded, narrow, oblique, raised
fold, from first to last teleoconch whorl. Other axial
sculpture of low, narrow, rounded intervariceal ribs
connecting base of anal tubes at abapertural part.
Spiral sculpture of very shallow, almost invisible P1
with anal tubes and P2 (shoulder). Anal tubes broad at
base, weakly tapered at extremity, ovate, separated
from axial varices and closer to preceding varix.
Aperture small, ovate, forming a continuous
peristome. Columellar lip narrow, flaring, smooth.
Outer lip erect, smooth within. Siphonal canal short,
broad, straight, ventrally sealed, weakly tapered
abapically.
Operculum light brown, ovate, with subapical nucleus
in lower right, with numerous concentric ridges.
Radula unknown.
Remarks. The shell of Laevityphis tillierae from New
Caledonia (Fig. 7H-J) is smaller and narrower, being
half the size of L. libos n. sp. for the same number of
teleoconch whorls, with a narrower siphonal canal and
with the anal tubes being less flattened and narrower,
less tapered at extremity.
Laevityphis tubuliger (Fig. 7K-M) is also smaller but
has a comparatively broader shell with a less elongate
spire, a narrower, more strongly dorsally recurved
siphonal canal, narrower anal tubes and a rounded
protoconch.
Etymology. Libos (G): drop, tear, in reference to the
drop-shaped shell.
Subfamily Rapaninae Gray, 1853
Tylothais n. gen.
Type species, here designated: Purpura savignyi
Deshayes, 1844, western Indian Ocean.
Description.
Shell broad, spire high or moderately high, nodose, with
7-11 axial ribs, crossed by broad, low, primary cords
and several secondary and tertiary cords. Primary cords
forming low to high knobs at their intersection with
axial ribs; subsutural cord broad, occasionally high.
Aperture broad, with rim of columellar lip adherent to
shell and low, elongate parietal node at adapical
extremity; anal notch narrow, moderately deep. Outer
apertural lip crenulated with low to moderately high,
narrow denticles within. Siphonal canal broad, very
short, broadly open.
Radula consisting of a rachidian bearing a broad, long,
central cusp, a narrower, somewhat shorter lateral cusp
on each side with a short lateral inner denticle. Marginal
area with 2-4 strong folds, ending as small denticles,
and of a short bifid marginal cusp. Lateral teeth narrow
and sickle shaped.
Included species
Tylothais savignyi (Deshayes, 1844) new. comb.
T. aculeata (Deshayes, 1844) new comb.
T. virgata (Dillwyn, 1817) new comb.
T. akidotos n. sp.
T. funilata n. sp.
T. inhacaensis n. sp.
T. ovata n. sp.
Etymology: from Tylos (G), meaning knob, for the
knobby morphology of the shell and Thais, the type
genus of the former Thaidinae, now synonym of
Rapaninae. The genus Thais was named after Thaïs, a
famous Athenian courtesan who lived during the time
of Alexander the Great.
Note: The species included here in Tylothais were
previously included by Claremont et al. (2013b) in
Thalessa H. & A. Adams, 1853.
F. C. Baker (1895: 183) designated Murex
hippocastanum Linnaeus, 1758 as type species of
Thalessa, and this designation was confirmed by ICZN,
Opinion 911 (1970: 20). Dodge (1957: 137-139)
designated as lectotype of Murex hippocastanum
Linnaeus, 1758, a specimen inscribed with the serial
number of Murex hippocastanum in Linnaeus' hand,
and concluded that Murex hippocastanum as described
by Linnaeus in the "Systema" was conspecific with
Pyrula galeodes Lamarck, 1822 (= Volema myristica
Röding, 1798). This conclusion was also followed by
Cernohorsky (1969: 295) and Tröndlé & Houart (1992:
98).
Thalessa is thus a subjective junior synonym of Volema
Röding, 1798 [family Melongenidae] and cannot be
used further to classify the muricids considered here (G.
Rosenberg, in litt.).
Claremont et al. (2013b) recognized four species of
Tylothais (as species of Thalessa):
Thalessa aculeata (Deshayes, 1844) (Figs 10H-I, 11G),
T. savignyi (Deshayes, 1844) (Figs 8A-D, 11E), T.
distinguenda (Dunker, 1866) = T. virgata (Dillwyn,
1817) (Fig. 10A-F, 11F) and a fourth unnamed species
close to T. savignyi. The name Purpura distinguenda
Dunker, 1866 has been considered (Houart,

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unpublished, as indexed in WoRMS) a junior subjective
synonym of Murex virgatus Dillwyn, 1817, a substitute
name for Murex plicatus Gmelin, 1791, non Lightfoot,
1786. When he established the name Murex plicatus,
Gmelin (1791: 3551) referred to Lister (Conchology t.
939, fig. 34.a), Klein (t. 3, fig. 56), Seba (t. 49, fig. 70)
and Chemnitz (t. 23, fig. 1141-1142). Of these
illustrations, Lister's and Klein's represent the same
shell, probably an ergalataxine; Seba's figure could be
anything, even a ranellid. Only Chemnitz illustrated a
shell similar to what is now known as "Thalessa"
virgata. Dillwyn (1817: 732) referred only to these last
figures but mis-citing them as "1441 and 1442" rather
than 1141 and 1142. However as pointed out by D. Reid
(in litt.) a doubt exists as to what the shell illustrated by
Chemnitz really is; it could also be Tylothais aculeata,
of which some forms are close to T. virgata.
The illustrations from Martini & Chemnitz (figs 1141
and 1142) are reproduced here (Fig. 10A) and
compared to a similar shell of T. virgata from
Southwest Thailand (Fig. 10B).
The name Murex virgatus Dillwyn, 1817 is the oldest
available name and has priority over Purpura
distinguenda Dunker, 1866. Moreover, the conditions
of ICZN Article 23.9 (reversal of precedence) are not
met, as Thais virgata has been used as valid by a
number of authors in recent years (see below).
In order to preserve the name Murex virgatus, I
designate the specimen illustrated by Chemnitz (fig.
1141-1142) as the lectotype for both names, Murex
plicatus Gmelin, 1791 and Murex virgatus Dillwyn,
1817. Chemnitz figured a shell from his own
collection: “In museo nostro”.
The shell collection of Johann H. Chemnitz is housed in
part at the Zoological Museum in Copenhagen.
Some of his muricids there were illustrated by
Cernohorsky (1974). Other specimens of the Chemnitz
collection were purchased for the Russian Imperial
Academy of Sciences at a public auction in
Copenhagen in 1802 (Martinov, 2002) and are now
housed at the Zoological Institute of Russian Academy
of Sciences, St. Petersburg.
The shell illustrated by Chemnitz could not be located
in either of those institutions, therefore I here designate
the specimen illustrated in Fig. 10B as the neotype for
both names.
Chemnitz's material originated from the East Indies and
the type locality of Murex plicatus is India so that the
locality of the neotype (Phuket, SW Thailand) fulfills
the ICZN art. 75.3.6 which states that the neotype
should come as nearly as practicable from the original
type locality. "East Indies" is here referred to in its
broadest context as defined by Encyclopaedia
Britannica: https://www.britannica.com/place/East-
Indies
The neotype is deposited in the Zoological Museum,
Copenhagen because a few rapanines from the
Chemnitz collection are already housed there as
illustrated by Cernohorsky (1974).
Tylothais virgata (Dillwyn, 1817)
Figs 10A-F, 11F
Murex plicatus Gmelin, 1791: 3551, ref. to Chemnitz,
figs. 1141, 1142 (not M. plicatus Lightfoot, 1786)
Murex virgatus Dillwyn, 1817: 732 (new name for
Murex plicatus Gmelin, 1791, p. 3551). Neotype
NHMD-189923, here designated.
Purpura distinguenda Dunker, in Dunker & Zelebor,
1866: 910. Two syntypes ZMB /Moll 9542, here
selected as lectotype ZMB/Moll 9542a and
paralectotype ZMB/Moll 9542b.
Purpura pseudohippocastanum Dautzenberg, 1929:
427 (new name for Purpura hippocastanum KIENER,
1836 (non Linné, nec Lamarck), Icon. coq. viv., p. 52,
pl.12, fig. 33; TRYON, 1880 (pars, non Lin. nec
Lam.), Manual, II, p. 162, pl. 45, fig. 42, 43 (both =
Murex virgatus Dillwyn, 1817).
Additional references
Thais aculeata ― Cernohorsky, 1967: 130, pl. 28, fig.
172; 1969: 295, pl. 47, fig. 1; Wilson & Gillett, 1971:
90, fig. 5; Hinton, 1972: 40, pl. 20, fig. 13; Wells &
Bryce, 1985: 90, fig. 303; Lai, 1987: 68, pl. 32, fig. 7;
Wilson, 1994: 238, pl. 4, fig. 12 (not Tylothais
aculeata).
Thais hippocastanum ― Abbott & Dance, 1982: 147,
text fig.; Subba Rao, 2003: 243, pl. 57, fig. 4 (not
Murex hippocastanum Linnaeus, 1758 = Volema
myristica Röding, 1798).
Thais distinguenda ― Tan, 2000: 499; Tan &
Kastoro, 2004: 50.
Thais (Mancinella) savignyi ― Tsuchiya, 2000: 397,
pl. 197, fig. 172 (not Tylothais savignyi).
Thais virgatus ― Hylleberg & Kilburn, 2003: 77;
Thach, 2005: 129, pl. 35, fig. 15; Dharma, 2005: 170,
pl. 60, fig. 7; Houart, 2008: 218, pl. 404, fig. 2; Houart
& Héros, 2008: 478.
Mancinella aculeate (sic) ― Zang, 2008: 182, text fig.
(not Tylothais aculeata).
Thais (Thalessa) virgatus ― Houart et al., 2010: 264,
text fig.
Thalessa distinguenda ― Claremont et al., 2013b: 94.
Thalessa virgata ― Tsuchiya, 2017 (I): 298, pl. 254,
fig. 1, (II): 961.
Type locality. East Indies.
Description of the neotype. Shell 26.6 mm in length,
19.4 mm in width, biconical, broad, heavy, nodose.
Subsutural ramp broad, strongly sloping, concave.
Spire moderately high.
Shell blackish brown with white spots or lines
between axial ribs and spiral cords. Aperture bluish
white within with 5 dark brown, narrow spiral lines;
inner edge of outer apertural lip blackish brown.
Columellar lip light brown with whitish band in
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Axial sculpture of last teleoconch whorl consisting of
8 broad, low, rounded ribs, each with high nodes at
intersection with spiral cords. Spiral sculpture of 5
strong, broad, low primary cords (P1-P5), decreasing
in strength and width abapically and 2 or 3 narrow,
secondary cords between each pair of primary cords.
Aperture broadly ovate. Columellar lip smooth,
adherent to shell. Outer lip crenulated with 5 narrow
denticles within (ID, D1-D4). Siphonal canal very
short, broadly open.
Remarks. Tylothais aculeata and T. virgata live
throughout the Indo-West Pacific while T. savignyi is
confined to the western Indian Ocean, from Zululand in
South Africa to Somalia and then throughout the Red
Sea, Yemen, Oman and the Persian Gulf.
Another species, Purpura tumulosa Reeve, 1846,
assigned to "Thalessa" in WoRMS was not included in
the phylogeny by Claremont et al. (2013b). Its
classification in "Thalessa" in WoRMS was based only
on the shell characters. However, the radula (Fig. 11I) is
much closer to the dagger type radula (Herbert et al.
2007) of Thaisella (Fig. 11H) than to Tylothais (Fig.
11A-G). That species is therefore better included in
Thaisella pending further research.
Tylothais akidotos n. sp.
Figs 2, 3E, 7N-Q, 11A-B
Thais distinguenda (Dunker) ― Kensley, 1973: 146,
fig. 509 (not Purpura distinguenda Dunker, 1866).
Type material. Holotype MNHN IM-2013-43791 and
2 paratypes MNHN IM-2013-43790 and MNHN IM-
2013-43792; 4 paratypes NM 7346/T4220; 1 paratype
RH (as listed below).
Material examined. INHACA 2011, stn MM11,
Inhaca Island, Mozambique, Maputo Bay, Xixuana,
25°60' S, 32°56' E, 0-1 m, 3 and 9 December 2011, 3
lv (holotype MNHN IM-2013-43791 and 2 paratypes
MNHN IM-2013-43790 and MNHN IM-2013-43792);
Inhaca Island, Mozambique, Delagoa Bay, 14 lv (10
ex NM7346 and 4 paratypes NM 7346/T4220, 1
paratype RH); Mozambique, south of Vilanculos,
mangrove, June 1971, 1 lv, NM L2726; 3 miles south
of Vilanculos, mangrove, 2 lv, NM G1300.
Type locality. Inhaca Island, Mozambique, Maputo
Bay, Xixuana, 25°60' S, 32°56' E, 0-1 m.
Distribution. Inhaca Island and in the vicinity of
Vilanculos, Mozambique.
Description. Shell medium-sized for the genus, up to
40.3 mm in length (paratype NM 7346/T4220).
Length/width ratio 1.4-1.6. Biconical, broadly ovate,
heavy, nodose and weakly squamous. Subsutural ramp
broad, weakly sloping, lightly concave. Shell dirty
white with brown or blackish brown spiral cords, dirty
white between cords and on nodes. Aperture bluish
white with dark brown axial band within outer
apertural lip rim; columellar lip bluish white, dark
brown abaperturally and adaperturally near of anal
notch.
Spire high, acute, up to 6 broad, convex, strongly
shouldered, nodose teleoconch whorls. Suture
impressed, obscured by broad subsutural cord.
Protoconch unknown, eroded in all examined
specimens.
Axial sculpture of teleoconch whorls consisting of
low, narrow, nodose ribs with high, narrow, pointed
nodes at intersection with primary spiral cords. Other
axial sculpture of numerous growth lamellae. Axial
sculpture of two first teleoconch whorls eroded, third
with 11 ribs, fourth with 11-13, fifth with 10 or 11,
last whorl with 10 ribs. Spiral sculpture of moderately
high, rounded, nodose, primary cords. Cords
longitudinally grooved, forming 3-5 narrow, weakly
squamous, low threads on crest. Secondary and
tertiary spiral sculpture of low, narrow cords of
different magnitude. All whorls with broad, nodose
SP. Early whorls with visible P1, s1 and tertiary cord;
P2 almost always covered by next whorl. Last
teleoconch whorl with broad, nodose SP followed by
adis, IP, P1-P5, ADP, with s1 or s1 and tertiary cord
between each pair of primary cords.
Aperture moderately large, ovate. Columellar lip
narrow, smooth, rim completely adherent with low
parietal tooth at adapical extremity. Anal notch
moderately deep, narrow. Outer lip erect, crenulated,
with 5 low, narrow denticles within: ID, D1-D4; D2
occasionally split. Siphonal canal very short, broad,
straight, broadly open, with broad P5 and occasionally
narrow ADP.
Figure 8
A-D. Tylothais savignyi (Deshayes, 1844)
A. Sinai, Gulf of Suez, Egypt, RH, 41.6 mm; B-C. Kenya, RH, 31.2 mm; D. Mozambique (no other data), RH,
28.2 mm.
E-L. Tylothais funilata n. sp.
E-F. Mozambique, south of Vilanculos, mangroves, holotype NM L2726/T4221, 34.9 mm; G-H. Mozambique,
south of Macoque, mangrove, paratype NM F9936/T4223, 31.3 mm; I-J. Mozambique, south of Vilanculos,
mangrove, June 1973, paratype NM G1300/T4222, 29.2 mm; K. Protoconch. INHACA, stn MM13, Inhaca
Island, 26°04' S, 32°54' E, sand, mud and rocks, paratype MNHN IM-2000-32842 (scale 500 µm); L. Operculum
(holotype) (scale 2 mm).

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Operculum dark brown D-shaped with lateral nucleus
in upper right.
Radula (Fig. 11A-B ) consisting of a rachidian with
broad, long, central cusp, a narrower lateral cusp on
each side flanked by a short lateral inner denticle.
Marginal area with 2 or 3 strong folds, ending as small
denticles, and a short marginal cusp. Lateral teeth
narrow and sickle shaped.
Remarks. Tylothais akidotos n. sp. resembles T.
savignyi (Fig. 8A-D) but differs in having a narrower
and higher spire, a slightly narrower, lower, last
teleoconch whorl, a narrower, more horizontal
subsutural ramp, the suture of the last whorl in T.
savignyi reach the P1 spiral cord of preceding whorl
while only covering P2 in the new species, s1 and one
or two threads remaining visible. Tylothais akidotos n.
sp. also has shorter and narrower nodes at intersection
of spiral and axial sculpture.
The holotype and 2 paratypes were preserved in
ethanol (95%) for future genetic studies.
Etymology. Akidotos (G): pointed, named in reference
of its pointed, acute spire.
Tylothais funilata n. sp.
Figs 3F-G, 8E-L
Type material. Holotype NM L2726/T4221, 3
paratypes NM G1300/T4222 and NM F9936/T4223, 1
paratype MNHN IM-2000-32842, 1 paratype RH (as
listed below).
Material examined. Mozambique, south of
Vilanculos, Mangroves, June 1971, lv (holotype NM
L2726/T4221); south of Macoque, mangrove, lv (1
paratype NM F9936/T4223); 3 miles south of
Vilanculos, mangrove, lv (2 paratypes NM
G1300/T4222, 1 paratype RH); INHACA 2011, stn
MM13, 26°04' S, 32°54' E, Inhaca Island, west of
Ponta Punduine, intertidal, sand, mud and rocks, 05
December 2011, dd (1 paratype MNHN IM-2000-
32842).
Type locality. Mozambique, south of Vilanculos,
mangrove.
Distribution. From Inhaca Island to south of
Vilanculos, Mozambique.
Description. Shell medium sized for the genus, up to
34.9 mm in length (holotype). Length/width ratio 1.4-
1.5. Biconical, broad, heavy, nodose and squamous.
Subsutural ramp broad, weakly sloping, weakly
concave. Shell creamy white with broad, orange
brown axial bands, more obvious on nodes of spiral
cords. Aperture white or light orange with orange-
brown narrow band within outer lip rim. Columellar
lip white.
Spire high with 2.5 protoconch whorls and teleoconch
up to 6 broad, strongly shouldered, nodose whorls.
Suture impressed, obscured by broad subsutural cord.
Protoconch (Fig. 8K) small, conical, elongate, whorls
smooth but weakly eroded in examined specimen.
Maximum width 900 µm, length 1000 µm.
Axial sculpture of teleoconch whorls consisting of
low, narrow, nodose ribs, each rib with low, narrow
nodes. Other axial sculpture of few growth lamellae.
First whorl eroded, second with 12 ribs, third and
fourth with 9-12, penultimate whorl with 9 or 10 ribs,
last with 8-10. Spiral sculpture of low, rounded, broad,
squamous and nodose primary cords, strongly
longitudinally grooved, forming 3 narrow, squamous
threads on crest, and squamous, narrow, secondary
cords and threads between primary cords. First
teleoconch whorl of paratype with visible, narrow P1
and P2, second with narrow SP, adis, IP, abis, P1, s1,
t, P2. Primary cords becoming broader from second or
third to last whorl. Adapical shoulder cords partially
covered by SP. Last whorl with broad SP, narrow IP,
broad P1-P5 with narrow secondary cords and threads
between primary cords.
Aperture large, ovate. Columellar lip narrow, smooth,
rim completely adherent with weak, broad, parietal
tooth at adapical extremity. Anal notch moderately
deep, narrow. Outer lip erect, crenulated, with 5 low,
narrow denticles within: ID, D1-D4. Siphonal canal
very short, broad, straight, broadly open, with broad
P5 and occasionally narrow ADP.
Operculum (Fig. 8L) dark brown, D-shaped with
lateral nucleus in upper right. Attached surface with
broad, callused rim.
Radula unknown.
Figure 9 (scale bar 2 mm)
A-E. Tylothais inhacaensis n. sp. Inhaca Island, Mozambique, Ponta do Farol, 25°58 S, 32°59' E, rocks, sand
and algae.
A-B. Holotype MNHN IM-2000-32843, 24.4 mm; C-D. Paratype MNHN IM-2000-32844, 20.6 mm; E.
Operculum (holotype).
F. Neothais intermedia (Kiener, 1836). Japan, Onna Point, Okinawa, 1963, RH, 33.4 mm.
G-J. Tylothais ovata n. sp. South Africa, Mission Rocks, Zululand, on rocks at low tide.
G-H. Holotype NM P0992/T4224, 32 mm; I-J. Paratype RH, 32 mm.
K-L. Thaisella coronata (Lamarck, 1816). Lagos, Nigeria, RH, 33.7 mm.

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Remarks. Tylothais akidotos n. sp. is close but
Tylothais funilata n. sp. differs in having a broader
last teleoconch whorl and a comparatively lower and
less pointed spire in adult specimens; it also has
broader primary spiral cords with lower, blunter nodes
except SP which is broader and more conspicuous in
Tylothais akidotos n. sp. The colour also strongly
differs, orange-brown, broad axial bands, a whitish
aperture with a white columellar lip and orange-brown
narrow band within the outer apertural lip in Tylothais
funilata n. sp., compared to brown or blackish brown
spiral cords in Tylothais akidotos n. sp., with a bluish
white aperture and dark brown band within the outer
apertural lip rim and a bluish white columellar lip,
dark brown abaperturally and near the anal notch.
Both species are sympatric and the holotype of
Tylothais funilata n. sp. was syntopic with Tylothais
akidotos n. sp.
Etymology. Funis (L): cord and lata (L): broad, in
reference to the broad spiral cords.
Tylothais inhacaensis n. sp.
Figs 3H, 9A-E
Type material. Holotype MNHN IM-2000-32843 and
1 paratype MNHN IM-2000-32844 (as listed below).
Material examined. INHACA 2011, stn MM6, type
locality, 27 November, 8 and 10 December 2011, 2 dd
(holotype MNHN IM-2000-32843 and paratype
MNHN IM-2000-32844).
Type locality. Inhaca Island, Mozambique, Ponta do
Farol, 25°58’S, 32°59'E, rocks, sand and algae.
Distribution. Known only from the type locality.
Description. Shell small for the genus, 24.4 mm in
length for holotype. Length/width ratio 1.5. Broadly
ovate, nodose, lightly built. Subsutural ramp broad,
weakly sloping, straight. Shell dirty white with dark
brown axial coloured spiral cords interrupted by dirty
white blotches or striae. Aperture dark bluish white
within with dark blackish brown broad axial band
within outer apertural rim. Columellar lip suffused
with different shades of brown, darker on abapertural
tip.
Spire moderately high with probably 5 broad, convex,
weakly angulate, shouldered, nodose teleoconch
whorls (early whorls partly eroded). Suture impressed,
obscured by broad subsutural cord. Protoconch
unknown, first whorls eroded in both examined
specimens.
Axial sculpture of teleoconch whorls consisting of
very low, narrow, nodose ribs with low, moderately
broad and high nodes at intersection with primary
spiral cords. Last whorl with 10 unequidistantly
spaced ribs. Previous whorls partly eroded. Spiral
sculpture of high, rounded, nodose primary cords and
low, narrow secondary and tertiary cords. Primary
cords longitudinally grooved, forming 2 or 3 low
threads on crest, partly eroded. Last whorl with (adis),
broad, high SP, IP, broad, high P1-P4; P5 narrower
and lower. Three secondary and tertiary cords between
P1 and P2, two between P2 and P3, one (paratype) or
three (holotype) between P3 and P4, one between P4
and P5.
Aperture large, broad, ovate. Columellar lip broad,
smooth, adherent, with very weak, low parietal tooth
at adapical extremity. Anal notch moderately deep,
narrow. Outer lip crenulated, smooth within. Siphonal
canal very short, broad, straight, broadly open.
Operculum (Fig. 9E) dark brown, D-shaped, with
lateral nucleus in center right. Attached surface with
broad callused rim.
Radula unknown.
Remarks. Tylothais savignyi differs from T.
inhacaensis n. sp. in having a larger, more solid and
heavier shell with a smaller aperture and heavier,
broader primary spiral cords with higher and broader
nodes.
Tylothais inhacaensis n. sp. could be also confused
with young specimens of Menathais intermedia
(Kiener, 1836) (Fig. 9F), but the shell morphology in
M. intermedia is different in being thicker and more
solid with broader primary spiral cords and nodes,
higher, more strongly sloping subsutural ramp with
absent or less obvious subsutural cord (SP), a usually
higher spire, a smaller aperture, white within instead
of dark bluish white in T. inhacaensis n. sp.
Etymology. Named after Inhaca Island where the
species was collected.
Figure 10
A-G. Tylothais virgata (Dillwyn, 1817)
A. Murex plicatus Gmelin, 1791. Original figures 1141 and 1142 from Chemnitz; B. Surin Beach, west coast of
Phuket, SW Thailand, neotype NHMD-189923, 26.6 mm; C-D. Nha Trang, Vietnam, RH, 40 mm; E-F. Nicobar
Islands, lectotype and paralectotype of Purpura distinguenda (Dunker, 1866) ― E. Lectotype ZMB/Moll 9542a,
35.5 mm; F. Paralectotype ZMB/Moll 9542b, 22.5 mm; G. New Caledonia, Bay of Touho, RH, 36.7 mm.
H-I. Tylothais aculeata (Deshayes, 1844). Masbate Island, Philippines, RH, 45.5 mm.

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Tylothais ovata n. sp.
Figs 3I-J, 9G-J, 11C-D
Thais distinguenda (Dunker, 1852) ― Richards, 1981:
56, pl. 29, fig. 233 (not Purpura distinguenda Dunker,
1866)
Thais (Thalessa) sp. ― Houart et al. 2010: 269, Text
figs (holotype NM and paratype RH).
Type material. Holotype NM P0992/T4224.
Paratypes: 1 MNHN IM-2000-32845; 1 IRSNB I.G.
33404/MT. 3465; 4 RH (as listed below).
Material examined. South Africa, Mission Rocks,
Zululand, on rocks at low tide, February 1989, lv
(holotype NM P0992/T4224 and 2 paratypes RH);
Mission Rocks, Zululand, in shallow water, 1986, lv
(1 IRSNB I.G. 33404/MT. 3465; 1 RH); St Lucia
Light House, Zululand, January 1974, 1 lv (paratype
MNHN IM-2000-32845); Park Rynie, Natal, on rock
in crevices, low tide, 1986, 1 lv (paratype RH);
Durban, Natal, on reef at 1-4 m, 1 lv, RH; Mission
Rocks, Natal, live on rocks at low tide, 1 lv, RH.
Type locality. South Africa, Mission Rocks,
Zululand, on rocks at low tide.
Distribution. South Africa, KwaZulu-Natal, from
Park Rynie to Mission Rocks.
Description. Shell medium sized for the genus, up to
32 mm in length (holotype). Length/width ratio 1.3-
1.4. Broadly ovate, heavy, nodose. Subsutural ramp
broad, weakly sloping, straight. Shell greyish brown.
Aperture bluish white within; columellar lip light tan,
darker coloured on outer border and abapertural
extremity; inner side of outer apertural lip with broad
brown axial band and narrow whitish axial band near
edge.
Spire high, up to 5 broad, convex, strongly
shouldered, nodose teleoconch whorls. Suture
obscured by broad subsutural ramp cord. Protoconch
unknown, eroded in all specimens, partly broken in
holotype.
Axial sculpture of teleoconch whorls consisting of
very low, narrow ribs, more obvious on early whorls,
with low, narrow nodes at intersection with primary
cords. Other axial sculpture of numerous growth
lamellae and striae. Holotype with partly eroded
sculpture on first and second whorl, third and fourth
whorls with 14 ribs, last with 12. Paratypes with 10 or
11 axial ribs on last whorl; other whorls with eroded
sculpture. Spiral sculpture of moderately broad,
nodose, primary cords and weakly squamous
secondary and tertiary cords. Last teleoconch whorl of
holotype with SP, adis, IP, P1, t, s1, t, P2, t, s2, t, P3,
t, s3, t, P4, s4, P5, ADP. Primary cords longitudinally
grooved, forming 3 or 4 lightly squamous, rounded
threads on crest. All whorls with broad, nodose SP,
P1-P4 of similar width and height, P5 and ADP
narrower and lower.
Aperture large, ovate. Columellar lip moderately
broad, smooth; rim adherent, very weakly partially
erect on a small portion abapically, with low, elongate
parietal tooth at adapical extremity. Anal notch
moderately deep, broad. Outer lip crenulated, with 5
weak, low denticles within: ID, D1-D4; ID
occasionally obsolete. Siphonal canal very short,
broad, straight, broadly open, with ADP.
Operculum dark brown, D-shaped with lateral nucleus
in center right. Attached surface with large callused
rim.
Radula (Fig. 11C-D) with a rachidian tooth bearing a
long, moderately broad central cusp, a broad lateral
cusp on each side, flanked with inner lateral denticle.
Marginal area with 3 or 4 small, obvious denticles,
and marginal cusp at extremity.
Remarks. The shell characters of Tylothais ovata n.
sp. are very close to Thaisella coronata Lamarck,
1816 (Fig. 9K-L), the type species of Thaisella, an
amphi-Atlantic species, living along the West African
coast, from south Mauritania to Angola (Houart, 1997:
62), but also in several localities in the Western
Atlantic. However, the morphology of the radula in
the new species differs from the typical Thaisella
species in having a rachidian with a long, narrow
central cusp compared to the dagger-type morphology
of Thaisella (Fig. 11H) in which the rachidian bears a
very long, broad, triangular cusp. The lateral cusps in
the new species differ also from Thaisella in being
slenderer and shorter in opposition to the long,
triangular cusps turned outwardly at their distal ends
in Thaisella. The radula of T. ovata n. sp. is closer to
Tylothais, the species is therefore here included in that
genus.
Figure 11. Radulae
A-B. Tylothais akidotos n. sp., Mozambique, Inhaca Island, RH (scale bars: A. 100 µm; B. 20 µm); C-D.
Tylothais ovata n. sp., South Africa, Zululand, RH (scale bars: C: 200 µm; D: 50 µm); E. Tylothais savignyi
(Deshayes, 1844), Mozambique, Inhaca Island, RH (scale bar: 100 µm); F. Tylothais virgata (Dillwyn, 1817),
New Caledonia, RH (scale bar: 50 µm); G. Tylothais aculeata (Deshayes, 1844), New Caledonia, RH (scale bar:
50 µm); H. Thaisella coronata (Lamarck, 1816), Benin, West Africa, RH (scale bar: 50 µm); I. Thaisella
tumulosa (Reeve, 1846), South Korea, RH (scale bar: 50 µm).
A-E: SEM Y. Kantor; F-I: SEM A. Warén.

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Other differences in the shell morphology are also
obvious, Thaisella coronata is usually broader, the
anal notch is deeper, the inner outer apertural lip is
sculptured by numerous, low lirae extending into the
aperture, instead of 5 low denticles in Tylothais ovata
n. sp. The colour of the aperture is also different.
No other related or similar species have been found in
Tylothais or in any of the other rapanine genera.
Etymology. Ovata (L): egg-shaped, named after the
broadly ovate form of the shell.
A
CKNOWLEDGEMENTS
I am very grateful to Philippe Bouchet (Muséum
national d'Histoire naturelle, Paris, France) for giving
me the opportunity for many years to study the
material collected during the Tropical Deep Sea
Benthos campaigns in the Indo-West Pacific, for
useful information and for his help in many ways
during the finalizing of this paper.
Many thanks also to Dai Herbert and the late Richard
(Dick) Kilburn (KwaZulu-Natal Museum, South
Africa) for the loan of material, to the late Rudolf
Kilias (Humboldt-Universität zu Berlin, Germany)
and Matthias Glaubrecht, (then Humboldt-Universität
zu Berlin, Germany, now in Centrum für Naturkunde
an der Universität Hamburg, Germany), for the loan of
the syntypes of Typhis tubuliger Thiele, 1925 and
Purpura distinguenda Dunker, 1866, to Christine
Zorn, Collection Assistent of Mollusca, Museum für
Naturkunde Berlin, for providing the registration
numbers of Purpura distinguenda, to Tom Schiøtte
(Natural History Museum of Denmark, Copenhagen)
and Boris I. Sirenko (Zoological Institute of the
Russian Academy of Sciences, St. Petersburg) for
searching in the Chemnitz collection in their
respective institution, to Virginie Héros and Philippe
Maestrati (MNHN) for helpful information about the
MNHN/IRD campaigns, to Anders Warén (Natural
History Museum, Stockholm, Sweden) and Yuri
Kantor (A. N. Severtsov Institute of Ecology and
Evolution, Russian Academy of Sciences, Moscow,
Russia) for preparation of radulae and for SEM work.
Manuel Caballer (MNHN) provided the images of the
MNHN holotype of Leptotrophon marshalli, E-
Recolnat Project: ANR-11-INBS-0004. Thanks also to
Gary Rosenberg (Academy of Natural Sciences of
Drexel University, Philadelphia, USA) and Philippe
Bouchet for their advice and comments about the new
genus described in this paper, to John Wolff
(Lancaster, Pennsylvania, USA), for checking the
English text and for other comments and to Felix
Lorenz (Buseck, Germany) and Dietmar Amon
(Kavieng, New Ireland) for the additional specimens,
now paratypes, of Aspella aclydis n. sp.. Koen
Fraussen (Aarschot, Belgium) is acknowledged for his
advice about Spinidrupa aethes n. sp. in connection
with the genus Clivipollia.
I am also very thankful to Gary Rosenberg for a
thorough review of the manuscript and for his helpful
suggestions.
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