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SOCIETE ROYALE BELGE DE MALACOLOGIE

R. HOUART, D. ZUCCON & N. PUILLANDRE NOVAPEX 20(HS 12): 1-52, 10 mars 2019

Description of new genera and new species of Ergalataxinae

(Gastropoda: Muricidae)

Roland HOUART*

roland.houart @ skynet.be

Dario ZUCCON**

dario.zuccon@ mnhn.fr

Nicolas PUILLANDRE**

puillandre @ mnhn.fr

* Research associate

Institut royal des Sciences naturelles de Belgique, rue Vautier, 29, 1000 Bruxelles

Institut Systématique Evolution Biodiversité (ISYEB), Muséum national d'Histoire naturelle, CNRS, Sorbonne

Université, EPHE, 57 rue Cuvier, CP 26, 75005 Paris, France.

** [Institut Systématique Evolution Biodiversité (ISYEB), Muséum national d'Histoire naturelle, CNRS,

Sorbonne Université, EPHE, 57 rue Cuvier, CP 26, 75005 Paris, France.

KEY WORDS. New taxa, Gastropoda, Muricidae, Ergalataxinae, Indo-Pacific, South Atlantic.

ABSTRACT. The recent genetic analysis of the muricid subfamily Ergalataxinae has led to a

better understanding of this subfamily, but some species were left without appropriate generic

assignments and the classification of others required revision. This knowledge gap is partially

filled herein, with new combinations and the description of three new genera. The examination of

new material, along with a careful re-examination of and comparison to existing material, resulted

also in the identification of nine new species. These new genera and new species are described

herein, lectotypes are designated and new combinations are given. The geographical range of all

the new species is provided on maps. AIl new species are compared with related or similar species.

The radula of Morula palmeri Powell, 1967 is 1llustrated for the first time.

New genera: Claremontiella n. gen., type species Purpura nodulosa C.B. Adams, 1845;

Murichorda n. gen., type species Purpura fiscellum Gmelin, 1791; Lauta n. gen., type species

Ricinula parva Reeve, 1846.

New species: Cytharomorula arta n. sp. from Reunion Island; C. absidata n. sp. from New

Caledonia; C. elegantula n. sp. from the western Pacific Ocean; C. fatuhivaensis n. sp. from the

Marquesas; C. manusuduirauti n. sp. from the Philippines; Orania pseudopacifica n. sp. from the

Marquesas; Tenguella chinoi n. sp. from the eastern Indian and western Pacific Oceans: T. ericius

n. sp. and T. chinoi from the western Pacific Ocean; and Claremontiella adiakritos n. sp. from

western Mexico. Claremontiella adiakritos n. sp. was previously confused with Ricinula

ferruginosa Reeve, 1846, a species which has been combined with Morula, Evokesia and Pascula,

and which is a synonym of C. nodulosa (C.B. Adams, 1845).

Lectotypes are designated for Fusus pachyrhaphe E.A. Smith, 1879 (= Orania pachyrhaphe) and

Purpura granulata Duclos, 1832 (= Tenguella granulata).

New combinations: Morula nodulosa (C.B. Adams, 1845) and Morula consanguinea (E.A. Smith,

1891) are transferred to Claremontiella n. gen.; Orania ornamentata Houart, 1995 ïs transferred to

Cytharomorula; Morula parva (Reeve, 1846) is transferred to Lauta n. gen.; Muricodrupa

fiscellum (Gmelin, 1791), Morula rumphiusi Houart, 1996 and Muricodrupa jacobsini Emerson &

D'Attilio, 1985 are transferred to Murichorda n. gen; Morula anaxares (Kiener, 1835) is

transferred to Muricodrupa; Muricopsis carnicolor Bozzetti, 2009 and Morula taiwana Lai &

Jung, 2012 are transferred to Orania; and Morula palmeri Powell, 1967 is transferred to Pascula.

Cantharus albozonatus E.A. Smith, 1890 is removed from Orania, to which it was assigned in

WoRMS (MolluscaBase 2018), and transferred to the genus Enginella Monterosato, 1917

(Buccinoidea, Pisantidae). | | | |

New synonymy: Séstrum ventricosulum G. & H. Nevill, 1875 is considered synonym of Morula

echinata (Reeve, 1846).

INTRODUCTION contracta (Reeve, 1846), E. tokugawai Kuroda &

Habe, 1971 and Cytharomorula vexillum Kuroda,

Four species were originally assigned to the 1953. The subfamily diagnosis was based on

subfamily Ergalataxinae (Gastropoda, Muricidae): morphological characters of the shell, radula and egg

Bedevina birileffi (Lischke, 1871), Ergalatax capsule (Kuroda et al. 1971). Later on, many genera

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

A _—_—_—_]————"—— — ———————]——

and species were added to the subfamily, counting

now 20 genera and 146 species recognized as valid

(WORMS). Recent molecular genetic analyses

confirmed the validity of this subfamily within the

Muricidae (Barco et al. 2010; Claremont et al. 2008;

Claremont et al. 2013).

Claremont et al. (2013) analysed 52 ergalataxine

species in 18 genera, representing 36% of the

currently accepted species and 90% of the genera, and

showed that many of the currently accepted genera

were polyphyletic, thus requiring further analysis. In

addition, some of the species included by Claremont

et al. (2013) were unidentified, and potentially

represented new species. To address some of these

issues, we herein discuss and clarify the taxonomy of

the genera Muricodrupa Iredale, 1918, Cytharomorula

Kuroda, 1953, Orania Pallary, 1900, Pascula Dall,

1908, and Tenguella Arakawa, 1965. In addition, three

new genera are described, together with several new

species in Cyfharomorula, Orania, Pascula, Tenguella

and Claremontiella n. gen. Our conclusions are based

on the results from Claremont et al 2013, completed

by the examination of additional material, including

newly sequenced samples.

Material and methods

Material

The material studied here primarily includes

specimens collected on various cruises conducted by

the MNHN/IRD in the Indo-West Pacific. Other

specimens were from the collections of the Natural

History Museum, United Kingdom; the KwaZulu-

Natal Museum, South Africa; the Houston Museum of

Natural History, Houston, Texas, U.S.A; and the

personal collection of the first author.

Specimens from the following expeditions of the

MNAN/IRD were examined: BENTHEDI: Mayotte,

northern Madagascar, Mozambique Channel, 1977;

MD32: Reunion Island, 1982; BIOCAL: New

Caledonia, 1985; CHALCAL 2: New Caledonia,

1986; SMIB 3: New Caledonia, 1987; SMIB 4: New

Caledonia, 1989; SMIB 5: New Caledonia, 1989;

BERYX 11: New Caledonia, 1992; MUSORSTOM 7:

SW Pacific, Wallis & Futuna, 1992; SMIB 8: New

Caledonia, 1993; BATHUS 2: New Caledonia, 1993;

MUSORSTOM 8: Vanuatu, 1994; MUSORSTOM 9:

Marquesas, 1997; LIFOU 2000: New Caledonia,

2000; BORDAU 2: Tonga, 2000; NORFOLK 1: New

Caledonia, 2001; NORFOLK 2: New Caledonia,

2003; PANGLAO 2004 and 2005: Philippines;

EBISCO: New Caledonia, 2005; SANTO 2006:

Vanuatu, 2006; AURORA 2007: Philippines;

TERRASSES: New Caledonia, 2008; MAINBAZA:

Mozambique, 2009; MIRIKY: Madagascar, 2009;

ATIMO VATAE: Madagascar, 2010; EXBODI: New

Caledonia, 2011; PAPUA NIUGINI: Papua New

Guinea, 2012; PAKAIHI I TE MOANA: Marquesas,

2012; KANACONO: New Caledonia, 2016;

KANADEEP: New Caledonia, 2017; MD208: Walters

Shoal, south of Madagascar, 2017 and BIOMAGLO:

Mayotte, Glorieuses and Comoros islands, 2017. |

The material used for the phylogenetic tree 1S

listed in Table I

Morphological analyses

The characters used to describe shell morphology

address the general aspect of the shell, its shape, size,

and colour, the shape of the spire including the

number and features of the protoconch and teleoconch

whorls, details of the suture and of the subsutural

ramp, details of axial and spiral sculpture, the

aperture, the siphonal canal, and when available, the

characters of the operculum and radula.

The method used to determine diameter and

height, and to count the number of protoconch whorls,

follows Bouchet & Kantor (2004) as shown in Fig. 1.

The morphology of the radula is described starting

from the rachidian tooth, followed by the lateral teeth

(Fig. 2). Unless otherwise indicated, species

descriptions are based on the holotype and the

paratypes. The bathymetric ranges given herein are the

inner values of the recorded depths: the deepest

minimum and the shallowest maximum of each

recorded depth range.

Molecular analyses

The protocols used for DNA extraction and the

PCR and sequencing of the Barcode fragment of the

Cytochrome oxidase I (COI) gene are detailed in

Puillandre et al. (2017). PCR products were purified

and sequenced by the Eurofins sequencing facility.

Sequences were deposited in BOLD (Barcode of Life

Datasystem) and GenBank (Table 1). Newly obtained

sequences were combined in a single dataset together

with the sequences from Claremont et al. (2013) that

- corresponds to the same genera, 1.e. Cytharomorula,

Orania and Tenguella and two outgroups

(Concholepas concholepas and Thais nodosa) (Table

1).

All the sequences were aligned manually (no indel

was detected), and the dataset was analysed using a

Bayesian approach as implemented in Mr.Bayes v. 3.2

(Huelsenbeck et al. 2001), with two runs consisting of

four Markov chains of 10,000,000 generations each,

with 8 chains, 5 swaps, and a sampling frequency of

one tree every 2,000 generations. Each codon position

of the COI gene was treated as an unlinked partition,

each following a general time reversible (GTR)

model, with a gamma-distributed rate variation across

sites approximated in four discrete categories and a

proportion of invariable sites. Convergence of each

analysis was evaluated using Tracer v. 1.6 (Rambaut.

Drummond, 2014) to check that all ESS values were

greater than 200. The trees were then calculated after

omitting the first 25% trees as burnin. Statistical

support Was evaluated as Bayesian posterior

probability (PP).

R. HOUART, D. ZUCCON & N. PUILLANDRE

NOVAPEX 20(HS 12): 1-52, 10 mars 2019

Table 1. Species used in the phylogeny

Cytharomorula vexillum MNHN-IM-2007-18174 EBISCO

Tenguella chinoi n. sp. NHMUK 20080772 Guam

ABBREVIATIONS

Repository

AMS: The Australian Museum, Sydney, Australia.

HMNS: Houston Museum of Natural History,

Houston, Texas, U.S.A.

IRSNB: Institut royal des Sciences naturelles de

Belgique, Bruxelles, Belgium.

MCZ: Museum of Comparative Zoology, Harvard

University, Cambridge, Massachusetts, U.S.A.

MHNG: Muséum d'Histoire Naturelle, Genève,

Switzerland.

MNHN: Muséum national d'Histoire naturelle, Paris,

France.

NHMUK: Natural History Museum, London, United

Kingdom.

NMSA: The KwaZulu-Natal Museum,

Pietermaritzburg, South Africa.

RH: collection of Roland Houart.

MK216548

Cytharomorula grayi (Dall, 1889) MK216549

MK216546

MK216544

MK216554

MK216556

MK216558

DW4647 :

HE584055.1

HES84052.1

Orania carnicolor (Bozzetti, 2009) MK216552

DW3608

K216557

[Orania castaneg _" [NHMUK 20100166 7 Jsouthafrics [7 |

K216555

MK216551

MK216550

MK216541

HES84053.1

HES84054.1

Malaysia

akaihi | te Moana

Tenguella ericius n. sp. MNHN-IM-2013-67609 Pakaihi | te Moana MQ7-M holotype MNHN |MUBA788-18 |MK216559

Tenguella granulata NHMUXK 2007645 Seychelles ar)

Tenguella marginalba NHMUXK 20090088 Queensland, Australid |

Concholepas concholepas NHMUXK 19990303 ÉTN

NHMUK 20070652 CN

MK216553

MNHN MUBA773-18 |MK216543

DW2496 MNHN MUBA771-18 |HE584062.1

MNHN MUBA312-15 |MK216545

z|z|z

[MK216550 |

IMK216541 |

HE5840531 |

HES840541 |

[______ [NHmUK U (HES840151 |

| ___ Jholotype NHMUK] [FNe77a8 |

IMK216559 |

INHMUK | JFN6774181 |

INHMUK | |HES84016.1

INHMUK | JFN6774171 |

INHMUK | __ JEu391581 |

INHMUK | _ |EU391579

Other

IRD: Institut de Recherche pour le Développement.

WoRMS: World Register of Marine Species.

Station number prefixes

CH: Chalut de loutre (otter trawl)

DR: Drague à roches (rocks dredge)

CP: Chalut à perche (beam trawl)

DC: Drague Charcot (Charcot dredge)

DW: Drague Warén (Warén dredge)

PM: M = marées (tides)

PR: R = récoltes à vue (handpicking)

Specimens

dd: empty shell(s).

lv: live collected specimen(s)

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

Number of Whoris (here 4)

E«

Maximum width

Figure 1. Method for determining diameter, height and

counting the number of protoconch whorls.

Terminology used to describe the spiral cords and

the apertural denticles (after Merle 2001, 2005)

(Figs 10-12). Variable features are given in

parentheses.

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 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: si = secondary cord

between P1 and P2; s2 = secondary cord between P2

and P3;etc.);

SP: subsutural cord.

t: tertiary cord

Aperture

D1 to DS: abapical denticles;

ID: infrasutural denticle.

Figure 2. Terminology used to describe the radula, here Pascula darrosensis (E.A. Smith, 1884)

cc: central cusp; Id: lateral denticle; le: lateral cusp; ma: marginal area; LT: lateral teeth.

Results and Discussion

Molecular analyses

In the phylogenetic tree (Fig. 3), the genera Orania

and Tenguella are recovered monophyletic.

Cytharomorula 1s recovered paraphyletic, contrary to

the results obtained by Claremont et al. (2013), but

this is probably due to the fact that we used only one

gene. Each included species, and in particular, the

newly described species, Cytharomorula absidata n.

sp., Cytharomorula elegantula n. sp., Tenguella chinoi

n. sp. and Tenguella ericius n. Sp., corresponds to an

independent lineage, and when several specimens per

species are included, they are grouped in a single

supported clade (Posterior Probabilities > 0.95).

Taxonomic implementation

Given the results obtained in Claremont et al.

(2013), the phylogenetic tree presented here (Fig. 3)

and the morphological analyses of additional material

(see below), we propose to revise several genera of

Ergalataxinae and to describe new genera and new

species. The different taxa are discussed below in the

order they appear in the fig. 1 of Claremont et al.

(2013), following their clade naming (A, B, C, V, W.

X, Y and Z); it should be noted that the clades for

which we do not propose any taxonomic change or

novelty are not discussed.

Purpura nodulosa (C.B. Adams, 1845 has

previously been assigned to Morula Schumacher,

1817 or Trachypollia Woodring, 1928 by a few

authors: to Morula by Rios (1970, 1985), Abbott

(1974), Bernard (1984) and Houart (1997); and to

Trachypollia by Radwin & D'Attilio (1976), De Jone

& Coomans (1988), Leal (1991) and Diaz Merlano &

Puyana Hegedus (1994). A sister relationship between

“Morula” nodulosa and all other species in major

clade À was well supported in Claremont et al. (2013).

who suggested to create a new genus for the western

and eastern Atlantic representatives of this clade.

R. HOUART, D. ZUCCON & N. PUILLANDRE

NOVAPEX 20(HS 12): 1-52, 10 mars 2019

EE _———_ _ _ _—_

“Morula" nodulosa was previously included in

Morula because its shell morphology was similar to

that of species of Tenguella, previously considered a

Synonym Of Morula. However, molecular genetic

studies have shown this species to be separated from

both Tenguella and Morula, as well as from

Trachypollia. À new genus is here described to

include "Morula" nodulosa and two other species.

Muricodrupa has long been considered to contain

three species: the type species M. fenestrata

(Blainville, 1832) (Fig. 29A-B), M. fiscellum (Gmelin,

1791) (Fig. 28F-H) and M. jacobsini Emerson &

D'Attilio, 1985 (Fig. 281) However, based on

Claremont et al. (2013), M. fiscellum should be

excluded from Muricodrupa, probably together with

M. jacobsini, a different although morphologically

related species. Muricodrupa fenestrata would then

remain as the sole representative of this genus.

However, Claremont et al. (2013) noted that "Morula"

anaxares (Kiener, 1835) formed a marginally

significant clade with M. fenestrata, which was not

contradicted in any other analyses. Indeed, the shell

morphology of "Morula" anaxares is very similar to

M. fenestrata (Fig. 29C-D). As it is clear that

"Morula" anaxares is excluded from Morula and

Tenguella, we here include this species in

Muricodrupa.

In the molecular analysis of Claremont et al.

(2013), M. fiscellum formed a well-supported clade

with "Morula" rumphiusi Houart; 1996, which was

clearly excluded from Morula Schumacher, 1817.

Both species, together with M. jacobsini, are

morphologically distinct from the other species

included in the clade B (Claremont et al. 2013), and

we here assigned them to a new genus (see Table 2).

Eleven species typically assigned to Morula

formed a monophyletic group within major clade B

(subclade V) of Claremont et al. (2013). Since this

subclade included M. uva, the type species of Morula,

Claremont et al (2013: 27) proposed that this generic

name should be restricted to this subclade. This

subclade also contained M. biconica, the type species

of Habromorula Houart, 1995, but the Habromorula

species analysed by Claremont et al (2013) and

assigned to this group [as a subgenus of Morula: M.

(H.) biconica, M. (H.) coronata, M. (H.) japonica, M.

(H.) spinosa and M. (H.) striata] did not form a clade

in any analysis. However, the authors also noted that

the monophyly of this morphologically distinctive

group was not strongly contradicted, so further

sampling and analysis could yet support Habromorula

as a monophyletic subgenus of Morula.

Three species currently assigned to Orania, ©.

gaskelli (Melvill, 1891) (Fig. 21L), ©. serotina (A.

Adams, 1853) (Fig. 2IM-N) and O. bimucronata

(Reeve, 1846) (Fig. 21Q) formed the subclade W in

the molecular phylogeny of Claremont et al. (2013),

clustering with Lataxiena fimbriata (Hinds, 1843)

(Fig. 22J) and Usilla avenacea (Lesson, 1842) (Fig.

210-P), and not with the other species of Orania

(clade Z, see below). Based on shell morphology, it is

certainly possible to group ©. gaskelli, O. serotina and

possibly U. avenacea in the same genus. However, it

would be difficult to justify the inclusion of ©.

bimucronata and particularly L. fimbriata in that

morphological group, and additional studies are

required before grouping them in the same genus. We

suggest Continuing to retain these species in the genera

to which they are currently assigned: Orania (0.

gaskelli, O. serotina and O. bimucronata), Lataxiena

(L. fimbriata) and Usilla (U. avenacea) (see Table 2).

The type species of Oppomorus Iredale, 1937, O.

nodulifera (Menke, 1829), formed a well-supported

clade with O. funiculata (Reeve, 1846) and O.

purpureocincta (Preston, 1909), and Claremont et al.

(2013) therefore proposed that Oppomorus should be

accorded full generic rank and not considered a

subgenus of Morula, as had been previously proposed

by Houart (2004). Three species formerly assigned to

Morula [M. nodulosa (C.B. Adams, 1845), M. parva

(Reeve, 1845) and M. rumphiusi Houart, 1996] were

excluded from the genus.

In major clade C of Claremont et al. (2013),

"Morula” parva (Reeve, 1846) was sister to all other

members of subclade Z, all morphologically clearly

distinct from “Morula” parva, indicating that a new

genus was also required for this species.

The genus Pascula Was also recovered as

polyphyletic by Claremont et al. (2013), and :1s

included in their Clade C, subclade Z, together with

Cytharomorula and Orania. The type species of

Pascula, P. citrica (Dall, 1908) (Fig. 22K-L) was not

analysed in that study. However, the study did include

a related species, P. darrosensis (E.A. Smith, 1884),

with a similar radula morphology (Figs 7F and 22M-

N). Pending further genetic research all species

previously included in Pascula are here retained 1n

this genus (see Table 2).

Based on the results of the present analysis, and

also because the shell morphology of the type species

of Orania [(Orania fusulus (Brocchi, 1814)], as well

as that of other Orania Species, 1s similar to the shell

morphology of Cyfharomorula, the latter could be

considered a possible junior synonym of Orania,

pending future research. Indeed, Claremont et al.

(2013: subclade Z) considered the validity of

Cytharomorula uncertain, as this genus was not

monophyletic in any analysis. However, this non-

monophyly was due to the presence of Orania

ornamentata Houart, 1995 (Fig. 13D-E) in the

Cytharomorula clade. The shell morphology of ©.

ornamentata is Strongly similar to that of the type

species of Cytharomorula (C. vexillum Kuroda, 1953)

(Fig. 13A-C), and we thus re-assign ©. ornamentata

in Cytharomorula. This renders the Cytharomorula of

Claremont et al. (2013: subclade Z) monophyletic.

Unfortunately, as long as Orania fusulus (Brocchi,

1814) (Fig. 21A-C), the type species of Orania, has

not been analysed genetically, it will remain unclear

whether Cytharomorula should be considered a junior

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

D] ——————————Z—

synonym of Orania or not. Here, we still consider that

Orania and Cytharomorula are two distinct genera.

Four ergalataxine genera were not included in

Claremont et al. (2013) because of a lack of

appropriate material, and the type species were not

EU391581 Concholepas concholepas

EU391579 Thais nodosa

0.99

0.2 subst/site

HE584015.1 Tenguella ceylonica

analysed for two additional genera (Table 2). The

inclusion of these four genera in the Ergalataxinae,

and the composition of Orania and Pascula, therefore

remains doubtful. Their classification is primarily

based on morphological shell and/or radula characters.

FN677417.1Tenguella musiva

HE584016.1 Tenguella marginalba

FN677414.1 Tenguella granulata

NHMUK20080772 Tenguella chinoi n. sp.

17 MNHN-M-2013-67608 LE Tenguella ericius n. sp.

MNHN-IM-2013-67609

MNHN-IM-2007-18225 Cytharomorula absidata n.sp.

HE584062.1 Cytharomorula vexillum

HE584055.1 Cytharomorula ornamentata

MNHN-IM-2013-60932

MNHN-IM-2013-61561

MNHN-IM-2013-69797

MNHN-IM-2013-69802

MNHN-IM-2013-69809

MNHN-IM-2013-66421

MNHN-IM-2013-66419

MNHN-IM-2013-69808

MNHN-IM-2013-69796

MNHN-IM-2013-69798

Cytharomorula grayi

MNHN-IM-2013-63326 Cytharomorula elegantula n.sp.

HE584052.1 Cytharomorula springsteeni

HE584054.1 Orania mixta

HE584056.1 Orania pacifica

HE584053.1 Orania fischeriana

1 MNHN-IM-2009-22449

MNHN-IM-2009-14366

MNHN-IM-2009-22448

HE584061.1

MNHN-IM-2013-66424

MNHN-IM-2013-66434

MNHN-IM-2013-66431

MNHN-IM-2013-66432

Orania camicolor

Orania castanea

Figure 3. Bayesian phylogeny of Ergalataxinae included in this paper and rapanine outgroups

SYSTEMATIC ACCOUNT

Family MURICIDAE Rafinesque, 1815

Subfamily ERGALATAXINAE Kuroda, Habe &

Oyama, 1971

Genus Cytharomorula Kuroda, 1953

Type species by monotypy: Cyfharomorula vexillum

Kuroda, 1953, Japan (Fig. 13A-C)

Diagnosis. Shell with high spire, elongate, ovate or

broadly ovate, weakly nodose, not or rarely exceeding

20 mm in length at maturity. Axial sculpture of last

teleoconch whorl consisting of 8 or 9 narrow, high,

rounded ribs. Spiral sculpture of low or moderately

high, narrow, rounded, low, primary and secondary

cords and few threads.

Aperture ovate. Columellar lip weakly concave,

smooth or with weak, narrow knobs abapically, rim

almost entirely adherent to shell. Anal notch broad,

deep. Outer lip with narrow, obvious denticles within.

Siphonal canal short, broadly open ventrally.

Radula of the type species (Fig. 6A) with three

dimensional rachidian tooth bearing a long, narrow,

projecting central cusp, a small, very narrow lateral

denticle, a moderately long, narrow, lateral cusp and

weak marginal folds. No marginal cusp. Lateral teeth

sickle shaped, with broad base.

Remarks. The genus Cyfharomorula includes 10

Recent species in WoRMS (MolluscaBase 2018):

Cytharomorula ambonensis (Houart, 1996) (Indo-

West Pacific); C. benedicta (Melvill & Standen, 1895)

(Pacific Ocean); C. danigoi Houart, 1995 (Pacific

Ocean); C. dollfusi (Lamy, 1938) (Indo-West Pacific);

C. grayi (Dall, 1889) (western and eastern Atlantic,

southwestern Indian Ocean); C. lefevreiana

(Tapparone Canefri, 1880) (western Indian Ocean); C.

paucimaculata (Sowerby, 1903) (Japan); C. pinguis

Houart, 1995 (New Caledonia, New Hebrides Arc and

Tonga); C. springsteeni Houart, 1995 (Indo-West

Pacific); C. vexillum Kuroda, 1953 (Pacific Ocean).

Only one of these species occurs in the Atlantic, all

others are found in the Indo-West Pacific region. A

revision of a few species (Cyfharomorula ambonensis.

C. benedicta, C. dollfusi, C. lefevreiana and C.

paucimaculata) was recently published (Houart.

2013a). Five additional new species from the Indo-

West Pacific are described below, and a new

combination 1s given for C. ornamentata (Houart.

1995), which gives us a total of 16 species for this

genus.

R. HOUART, D. ZUCCON & N. PUILLANDRE

NOVAPEX 20(HS 12): 1-52, 10 mars 2019

a] _—_—_

In several species the primary spiral cord P4 is

generally broader and larger, with broader, more

Conspicuous knobs at the intersection with the axial

ribs.

Cytharomorula absidata n. sp.

Figs 4; 11F; 13F-L

Type material. Holotype MNHN-IM-2000-34201,

New Caledonia, NORFOLK 2, stn DW2123, 23°185,

168°15'E, 187-197 m.

Paratypes: New Caledonia, NORFOLK 2, stn

ENW2TS, 2S008)S, LOS, 87-197 mn, 1 ‘A,

paratype MNHN-IM-2007-18225 (BOLD MUBA765-

18; GenBank HES584051.1); SMIB 5, stn DW93,

22°20'S, 168°43'E, 240-255 m, 1 dd, MNHN-IM-

2010-23374.

Type locality. New Caledonia, 23°18'S, 168°15'E,

187-197 m.

Distribution. Southern New Caledonia, living at 187-

197 m.

Description. Shell medium sized for the genus, up to

13 mm in length at maturity (paratype MNHN).

Length/width ratio 2.0 (paratype). Lanceolate,

biconical, broadly ovate. Subsutural ramp broad,

strongly sloping, concave.

Shell creamy white, axial ribs light orange coloured

near suture. Light tan on crest of some primary spiral

cords and at tip of siphonal canal. Aperture white.

Spire high, acute, with 3+ protoconch whorls (tip

broken). Teleoconch up to 5 broad, strongly convex,

weakly shouldered whorls. Suture strongly adpressed.

Protoconch moderately small, conical, acute. Whorls

smooth, glossy. Maximum width 850 um. Terminal

lip of sinusigera type.

Axial sculpture consisting of high, broad, rounded,

weakly nodose ribs. Ribs of last whorl connected to

preceding whorl with strong buttresses. Other axial

sculpture consisting of numerous, very narrow, small,

growth lamellae. Spiral sculpture of low, rounded,

narrow, smooth primary, secondary and tertiary cords,

and additional, few smooth threads. Primary and

secondary cords of same strength, tertiary cords

narrow. Last whorl with SP, adis, IP, abis, PI, s1, P2,

s2, P3, s3, P4, sd, P5, s5, P6, s6, ADP, (ads), MP, ms,

ABP, (abs).

Aperture moderately small, ovate. Columellar lip

narrow, smooth except 2 or 3 weak knobs abapically.

Rim adherent, weakly erect on a small portion

abapically. Anal notch deep, broad. Outer lip smooth,

with low. narrow denticles within: ID, DI and D2

fused, D3, D4 and D5. Siphonal canal short, narrow,

straight, narrowly open.

Operculum and radula unknown.

Remarks. This is the species identified as

Cytharomorula cf. grayi from New Caledonia

(MNHN-IM-2007-18225) and analyzed in Claremont

et al. (2013).

Cytharomorula elegantula n. sp. differs from C.

absidata n. Sp. in having a higher, more elongate

spire, less shouldered teleoconch whorls, a less

concave subsutural ramp and a relatively narrower last

teleoconch whorl. The spiral cord morphology differs

also, consisting of narrower primary, secondary and

tertiary cords and more widely spaced primary cords

in C. elegantula n . sp.

Cytharomorula pinguis Houart, 1995 (Fig. 18M-N)

differs greatly from C. absidata n. sp. in having a

higher spire, strong columellar folds and heavy outer

apertural lip denticles, a broader outer apertural lip

and much broader, flat, spiral cords.

Cytharomorula springsteeni (Figs 10F; 13M-O) also

recorded in New Caledonia differs in its shell

morphology. C. springsteeni has also a strongly

concave subsutural ramp and adpressed suture, but the

shell is lighter, more fragile, with a slightly narrower

last teleoconch whorl, narrower axial ribs, less

expanded on the preceding whorl, and narrower spiral

cords.

Cytharomorula manusuduirauti n. Sp. a species here

described from the Philippines (Fig. 18G-L), is a

smaller shell, also with broad axial ribs but which are

obviously less expanded on the penultimate whorl.

The numerous, narrower, primary, secondary and

tertiary cords are of approximately similar strength in

C. manusuduirauti and more crowded than im C

absidata n. sp.

Etymology. absidatus (a) (L): arched, named for the

strongly arched form of the last teleoconch whorl.

150°

90° 105° 120° 135° 165°

Figure 4. Distribution of Cyfharomorula absidata n.

Sp.

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

A —— "—…"—" —"

Cytharomorula arta n. sp.

Figs 5; 1I0A-B; ISA-E

Type material. Holotype MNHN-IM-2000-34203,

Reunion, MD32 (REUNION), stn CP57, 21°05S,

SSAUE 210-227mdd.

Paratypes: Reunion, MD32, stn DC2, 21°12'S,

55°49'E, 160-190 m, 2 dd, MNHN-IM-2000-34204;

siDCS6 21P05$S, 552125, 1982 170-22Stmr ad

MNAN-IM-2000-34205; stn CP57, 21°05'S, 5S°11'E,

210-227 m, dd, 1 MNAHN-IM-2000-34206; stn

DC176, 21°02'S, SS°11'E, 165-195 m, 1 dd, MNHN-

IM-2000-34207, 1 dd, RH.

Type locality. Western Indian Ocean, Reunion,

21805 SSUE 210-2271.

Distribution. Western Indian Ocean, East Reunion

Island, 190-210 m (dd) (Fig. 5).

Description. Shell small for the genus, up to 14.3 mm

in length at maturity (holotype). Length/width ratio

2.0-2.2. Slender, lanceolate, narrow, nodose, lightly

built. Subsutural ramp broad, strongly sloping, weakly

concave.

Creamy-white or light tan with numerous spiral cords

topped with brown. Tip of siphonal canal light brown.

Aperture white.

Spire very high, acute, with 3.5-4 protoconch whorls

and teleoconch up to 5 weakly convex, elongate,

weakly angulate, weakly shouldered, nodose whorls.

Suture of whorls adpressed. Protoconch small,

conical, whorls smooth, glossy, height and width 900

um. Terminal lip of sinusigera type.

Axial sculpture of teleoconch whorls consisting of

moderately high, broad, nodose ribs. First whorl with

9 ribs, second with 9 or 10, third with 10 or 11, fourth

with 10-12, last whorl with 8 or 9 ribs. Occasional

presence of a single varix on penultimate and last

whorls. Spiral sculpture of high, rounded, nodose,

primary, secondary and tertiary cords. First whorl with

visible IP, PI and P2 or P1-P3; second with SP, adis,

IP, abis, P1, si, P2, s2, P3 with additional tertiary

cords on third and fourth whorls. Last teleoconch

whorl with SP, adis, IP, abis, SP, P1, s1, P2, s2, P3,

s3, P4, s4, PS, s5, P6, ADP, MP, ms and ABP with

additional tertiary cords (Fig. 10A). Cords of

approximately similar size except broader and higher

P4, usually lighter coloured. Crossing of axial ribs and

primary cords giving rise to low nodes, more obvious

on P4.

Aperture narrow, ovate. Columellar lip narrow,

smooth, rim partially weakly erect, adherent at a small

portion at adapical extremity. Anal notch moderately

deep, broad. Outer lip erect, smooth, with 6 low,

elongate denticles within, consisting of IP, DI-D5,

occasionally obsolete (Fig. 10B). Siphonal canal short,

17% of total shell length in holotype, narrow, weakly

dorsally recurved, broadly open.

Operculum and radula unknown.

Remarks. Cytharomorula springsteeni (Figs 10F;

13M-O), a species currently known from several

localities in the Pacific Ocean (New Caledonia,

Vanuatu, Philippines and the Marquesas), differs from

C. arta n. sp. in having a comparatively broader shell

with a slightly steeper shoulder ramp, more

shouldered teleoconch whorls, a shorter, less elongate

spire, and a weakly narrower, longer siphonal canal.

Cytharomorula elegantula n. sp. described herein

(Figs 10C; 16H-N) differs in having a comparatively

larger and broader shell reaching 17.3 mm in length, a

broader anal notch, a small but obvious parietal tooth

and a low, narrow, weak, elongate knob on the

adapical part of the columellar lip, which is smooth in

C. arta n. sp. Young specimens of C. arta n. sp. and

C. elegantula n. sp. are oddly similar (Fig. 10 À & C),

but C. elegantula still differs in having a broader,

larger shell for a same number of teleoconch whorls, a

low, broad SP, a comparatively narrower siphonal

canal and already in young specimens a very low

parietal tooth and a low knob on the columellar lip.

Cytharomorula lefevreiana (Fig. 15F-G), another

species occurring off Mauritius and Reunion that was

collected during the MD32 cruise (Houart, 2013a),

differs in many aspects, including having a

comparatively smaller and narrower shell, from 7 to

11 mm in length, more strongly shouldered whorls

with very obvious P1-P3 spiral cords, and a relatively

smaller aperture.

Etymology. Arfa (L): meaning narrow, named for its

narrow shell.

15° 30° 45° 60° 15° 90° 105° 120°

Figure 5. Distribution of Cytharomorula arta n. sp.

CYTHAROMORULA GRAYI GROUP

À very careful comparative study of shell and genetic

characters, including additional newly available

material, indicates that specimens identified or

doubtfully identified as C. grayi (Dall, 1889),

R. HOUART, D. ZUCCON & N. PUILLANDRE

NOVAPEX 20(HS 12): 1-52, 10 mars 2019

—_—_…— _—

distributed in five distinct areas, represent in fact three

distinct species. C. grayi s.s. lives in the tropical

western and eastern Atlantic with a range extension

which includes south-eastern South Africa and the

south-western Indian Ocean; a second species, C.

elegantula n. sp., is limited to the southwest Pacific, in

New Caledonia, Vanuatu and Tonga; and a third, C.

fatuhivaensis n. sp., is restricted to the Marquesas

Archipelago. In the present state of our knowledge,

none of these species appear to have overlapping

ranges.

Cytharomorula grayi (Dall, 1889)

Figs 10D; 11A-B; 14; 15H-P

Nassaria (Nassarina) grayi Dall, 1889: 183, pl. 32,

fig. 12a.

Cantharus (Tritonidea) laevis E.A. Smith, 1890: 261,

pl2k he 1.

Trophon lowei Watson, 1897: 244, pl. 19, fig. 12.

Cytharomorula (?) grayi — Houart, Kilburn &

Marais, 2010: 223.

Not Cytharomorula grayi — Houart, 1995: 254, figs

9-10, 67-68; Houart & Héros, 2008: 459 (= C.

elegantula n. Sp.)

Not Cytharomorula sp. cf. C. grayi — Tründlé &

Houart, 1992: 88, fig. 48 (= C. fatuhivaensis n. sp.).

Not Cytharomorula grayi — Houart & Trôndlé, 2008:

70, 92 (= C. fatuhivaensis n. sp.).

Not Cyfharomorula grayi — Tsuchiya, 2000: 381, pl.

16% 00e 61; Isuchiya 2017: 954, pl. 247, fie: 3 (=

Cytharomorula sp.).

Type material. Nassaria (Nassarina) grayi Dall,

1889: The specimen illustrated by Dall (MCZ 7256)

was "removed" from its box in the MCZ, and the

specimen presently in the box labelled C. grayi 1s not

C. grayi but another species, not even a muricid

(Houart & Tründlé, 1992; Vokes, 1996), but rather a

buccinid. Nevertheless, the original figure in Dall

(1889: pl. 32, fig. 12a) is excellent and its identity is

undoubtable. The specimen illustrated by Dall was

designated as the lectotype by Vokes (1996: 29);

Cantharus (Tritonidea) laevis E.A. Smith, 1890:

lectotype NHMUK 1889.10.1.2362 (Fig. 150O-P),

designated by Vokes (1996: 29) (see remarks) and 4

paralectotypes NHMUK 1889.10.1.2363-2366;

Trophon lowei Watson, 1897: lectotype NHMUK

1911.7.17.2 (Fig. I15N), designated by Houart &

Abreu (1994: 122) (see remarks).

Type localities. Nassaria (Nassarina) grayi Dall,

1889: off Barbados, in 73 fathoms (133.5 m), Blake

Station 290: Cantharus (Tritonidea) laevis E.A.

Smith, 1890: Saint Helena; Trophon lowei Watson,

1897: Madeira.

Other material examined. Western Atlantic: East

of Guadeloupe (south of Désirade), 250 m, 2 Iv, RH;

west coast of Barbados, 165 m, 1 Iv, RH; southeastern

Brazil, MDS55 (BRESIL), stn DCS55, 20°32'S,

28°52'W, 780-795m, 12 dd, MNHN-IM-2012-33133;:

MDSS, stn CB103, 23°36'S, 42°02'W, 200-217 m, 1

Iv, MNHN-IM-2012-33127.

Eastern Atlantic: Canary Islands, Gran Canaria,

Sardina, dredged, 2 Iv, RH; Canary Islands, Tenerife,

Palma, 150-200 m, 52 1v, RH; Madeira, Porto Santo, 1

IV, RH; Madeira, Funchal Bay, 100 m, 1 1v, RH;

Madeira (no other data), 1 1v, RH.

South Africa: Durban, off Umlaas Canal, 128 m, 1 Iv,

NM B6282; South Africa, Transkei, off Mtamvuna

River, 31°08'S, 30°17'E, 160 m, 1 dd, NM C9646;

southern Natal, 75-95m, in grit samples, 3 Iv, coll.

Roy Aiken; ! dd, RH.

Walters Shoal: MD208, stn DW4877, 33°105,

43°49'E, 217-256 m, 1 Iv, MNHN-IM-2013-66418, 1

IV, MNHN-IM-2013-66419 (BOLD MUBA781-18;

GenBank MK216540); stn DW4880, 33°175,

43°5l'E, 275-318 m, 2 Ivy, MNHN-IM-2013-66421

(BOLD MUBA782-18; GenBank MK216553); stn

DW4885, 33°175S, 43°55E, 272-380 mm, 2 I, 1

MNEN, 1 RH; stn DW4894, 33°09'S, 43°50'E, 199-

261 m, 2 Iv, MNAN; stn DW4896, 33°07'S, 43°5l'E,

325-357 m, 1 dd, MNHN; stn DW4897, 33°09'S,

43°59'E, 490-584 m, 1 1v, MNEN.

Mozambique Channel: MAINBAZA, stn DW3167,

Almirante Leite Bank, 26°12'S, 35°02'E, 228-230 m, 3

dd, MNEN.

Reunion: MD32 (REUNION), stn DC2, 21°125S,

55°49'E, 160-190 m, 3 dd, MNHN; stn DCI128,

20°S1'S, 55°36'E, 280-340 m, 1 dd, MNAHN; stn

CP129, 20°5L'S, 55°36'E, 290-300 m, 1 dd, MNHN.

Mayotte, (Comoros and Glorieuses Islands:

BIOMAGLO, stn DW4789, 12°22'S, 46°25'E, 340-

342 m, 4 Iv, MNHN-IM-2013-69797 (BOLD

MUBA774-18; GenBank MK216549), MNHN-IM-

2013-69802, (BOLD MUBA775-18; GenBank

MK216544), MNAN-IM-2013-69808, (BOLD

MUBA776-18; GenBank MK216554), MNHN-IM-

2013-69809, (BOLD MUBA777-18; GenBank

MK216556); stn DW4790, 12°22'S, 46°25'E, 360-375

m, 1 Ivy, MNHN-IM-2013-69798 (BOLD MUBA778-

18; GenBank MK216546); stn DW4841, 12°23S,

43°33'E, 154-333m, 1lv, MNHN-IM-2013-69796

(BOLD MUBA779-18; GenBank MK216548).

Glorieuses Islands: SW Grande Glorieuse,

BENTHEDI, stn 93DS, 11°32'S, 47°16'E, 480-550 m,

1 dd, MNHN.

Distribution. Cyrharomorula grayi occurs from the

Lesser Antilles in 100-450 m (Garrigues & Lamy

2018) to southern Brazil in the western Atlantic, and

in the eastern Atlantic in the Canary Islands, Madeira

and the banks off Portugal, but not on the mainland

European continental shelf (SEAMOUNT I! cruise

material in MNHN). The species also occurs off Saint

Helena and extends 1ts range to South Africa, Walters

Shoal, Mozambique, Reunion, Mayotte, Comoros and

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

D nd EEE ————]__—

Glorieuses Islands, Indian Ocean, there living in 128-

275 m (Fig. 14).

Remarks. Vokes (1996: 29) mentioned and illustrated

the originally figured syntype of Cantharus laevis as

the holotype. Smith (1890) never designated any shell

as the holotype. The type material in the NHMUK

consists of 5 specimens, mentioned as syntypes by

Houart (1997: 56). Vokes (1996) only mentions what

she then called the "holotype", but having examined

the material when visiting the NHMUXK she certainly

was aware of the existence of the other 4 specimens.

The mention of Vokes is here considered as an

indirect lectotype designation by inference as

holotype, following ICZN Art. 74.5: "In a lectotype

designation made before 2000, either the term

"lectotype", or an exact translation or equivalent

expression (e.g. "the type”), must have been used or

the author must have unambiguously selected a

particular syntype to act as the unique name-bearing

type of the taxon. When the original work reveals that

the taxon had been based on more than one specimen,

a subsequent use of the term "“holotype" does not

constitute a valid lectotype designation unless the

author, when wrongly using that term, explicitly

indicated that he or she was selecting from the type

series that particular specimen to serve as the name-

bearing type".

Houart & Abreu (1994: 122, fig. 5) illustrated the

syntype and only existing specimen of Trophon lowei

as the holotype. This is here also considered as a valid

lectotype designation following ICZN Art. 74.6:

"When it has been accepted that a nominal species-

group taxon was based on a single specimen and the

original description neither implies nor requires that

there were syntypes, and if it is considered

subsequently that the original description was based

on more than one specimen, the first author to have

published before 2000 the assumption that the

species-group taxon was based upon a single type

specimen is deemed to have designated that specimen

as the lectotype".

Discussion. Material consisting of numerous live and

dead collected specimens from the southwestern

Indian Ocean was examined and initially considered a

separate species. Adult specimens are bigger than the

typical Atlantic Cyfharomorula, wider and more

shouldered, have a slightly wider aperture and a

narrower Siphonal canal (Figs 11A-B; 16A-G).

Nevertheless, genetic analysis performed on

specimens from Guadeloupe (KARUBENTHOS 2),

Walters Shoal (MD 208) and other specimens

collected off Mayotte, Comoros and Glorieuses

islands confirm the conspecifity of the populations of

the Atlantic Ocean and the Indian Ocean.

The type material of Trophon lowei (Madeira) (Fig.

ISN) and Cantharus laevis (Saint Helena) (Fig. 150-

P) seems to be composed of intermediate forms

between the populations of the western Atlantic and

the Canary Islands (Fig. 15H-L) and those of the

Indian Ocean.

Cytharomorula elegantula n. sp.

Figs 6B-C; 10C; 14; I6H-N

Cytharomorula grayi — Houart, 1995: 254, figs 9-10,

67-68; Houart & Héros, 2008: 459 [(not C. grayi

(Dall, 1889)].

Type material. Holotype MNHN-IM-2013-63326

(BOLD MUBA780-18; GenBank MK216542), New

Caledonia, KANACONO, stn DW4778, 23°035,

168°18'E, 170-248 m.

Paratypes: New Caledonia, KANACONO, stn

DW4741,-2252/S 674 210-2100 101,

MNAN-IM-2013-63337;

KANADEEP, stn DW4945, 25°22'S, 159°43'E, 108-

130 m, 1 Iv, MNHN-IM-2013-48638;

BIOCAL, stn DW66, 24°55'S, 168°22'E, 505-515 m,

2 1v, MNAN-IM-2010-23359 (radula illustrated in

Houart, 1995: figs 9-10 and reproduced here Fig. 6B-

C);

SMIB 4, stn DW46, 24°47'S, 168°09'E, 260 m, 1 Iv,

MNAN-IM-2010-23369;

SMIB 8, sin DWI154, 24°46'S, 168°08'E, 235-252 m,

13 Iv & dd, 11 MNHN-IM-2000-34208, 2 RH; stn

DWI159, 24°46'S, 168°08'E, 241-245 m, 9 Iv & dd,

MNEN-IM-2000-34209; stn DWI167, 23°38S5,

168°43'E, 430-452 m, 2 1v, MNHN-IM-2010-23357.

Type locality. South of New Caledonia, 24°48'S,

168°09'E, living at 245 m.

Other material examined. New Caledonia.

BIOCAL: stn DW65, 24°48'S, 168°10'E, 245-275 m,

1 dd, MNAN-IM-2010-23360;

CHALCAL 2: sin CH8, 23°13'S, 168°03'E, 300/'m, 3

dd, MNHN-IM-2010-23366; stn DW69, 24°44'E.

168°08'E, 260 m, 1 dd, MNHN-IM-2010-23365: stn

DW70, 24°46S, 168°09'E, 232 m, 1 1v, MNHN-IM-

2010-23364; stn DW71, 24°42'S, 168°10'E, 230 m. 8

Iv & dd, MNHN-IM-2010-23363;

SMIB 3: stn DWB8, 24°45'S, 168°08'E, 233 m, 2 dd,

MNAHN-IM-2010-23368; DW14, 23°40'S, 167°60'E.

246 m, 4 1v & dd, MNHN-IM-2010-23367;

SMIB 4: stn DW44, 24°46S, 168°08'E, 300 m., 1 dd.

MNEAN-IM-2010-23371; stn DW49, 24°465S.

168°09'E, 300 m, 1 dd, MNHN-IM-2010-23370:

SMIBB 5: stn DW95, 22°60'S, 168°20'E, 200 m, 1 dd.

MNEAN-IM-2010-23372; stn DW96. 23 00,

168°19'E, 245 m, 1 dd, MNHN-IM-2010-23373:

SMIB 8: stn DW159, 24°46'S, 168°08'E, 241-245 m.

1 1v, MNHN-IM-2010-23356;

BERYX 11: sin DW11+CP23, 24°44'S, 168°10'E.

320-350 m, 1 dd, MNHN-IM-2010-23361: DWI18.

24°48'S, 168°09'E, 250-270 m, 3 dd, MNHN-IM-

R. HOUART, D. ZUCCON & N Ù

. PUILLANDRE NOVAPEX 20(HS 12): 1-52, 10 mars 2019

Figure 6 — Radulae

À. Cytharomorula vexill

B-C. Cytharomorula ele

D. Cytharomorula ornamentata

E-F. Orania carnicolor Bozzetti

G. Orania fusulus (Brocchi, 1814), Angola. Scale bar: 50 um (SEM A. Warén)

(Küster, 1858), East London. South Africa. Scale bar: 50 um (SEM A. Warén)

um Kuroda, 1953, New Caledonia. Scale bar: 50 um (SEM A. Warén)

gantula n. Sp. New Caledonia. Scale bars: B. 100 um; C. 20 um (SEM P. Bouchet)

(Houart, 1995), Transkeiï, Sourth Africa. Scale bar: 50 um (SEM A. Warén)

i, 2009, Madagascar. Scale bars: E: 50 um; F. 20 um (SEM A. Warén)

H. Orania castanea

R. HOUART, D. ZUCCON & N. PUILLANDRE New genera and new species of Ergalataxinae

Figure 7 — Radulae

A-B. Orania pseudopacifica n. sp. Marquesas, MUSORSTOM 9, stn CP1239, paratype MNHN-IM-2000-34213.

Scale bar: A. 100 um, B. 50 um (SEM Y. Kantor).

C. Orania pacifica (Nakayama, 1988), Philippines. Scale bar: 20 um (SEM P. Bouchet)

D-E. Orania pleurotomoides (Reeve, 1845), Papua New Guinea. Scale bar 10 um (SEM P. Bouchet)

F. Pascula darrosensis (E.A. Smith, 1884), Philippines. Scale bar: 50 um (SEM A. Warén)

G-H. Pascula palmeri (Powell, 1967), New Zealand, W of Maroro Point, Aorangi Island, Poor Knights Islands.

35°28.57 S - 174°44.18 E, 5-20 m. NMNZ M.164595. Scale bar: 200 um (SEM Y. Kantor) È

R. HOUART, D. ZUCCON DRE

& N. PUILLANDRE NOVAPEX 20(HS 12): 1-52, 10 mars 2019

Figure 8 — Radulae

A-B. Tenguella chinoi n. sp., Vanuatu, SANTO, stn RAP15, paratype MNHN-IM-2000-34215. Scale bar: A. 50

um, B. 30 um (SEM Y. Kantor)

C. Tenguella granulata (Duclos, 1832), Gulf of Aden. Scale bar: 50 um (SEM A. Warén)

D. Claremontiella adiakritos n. sp., San Felipe, Baja California. Scale bar: 50 um (SEM A. Warén)

E-F. Claremontiella consanguinea (E.A. Smith, 1891), Säo Tomé. Scale bar: G: 50 um, H: 20 um (SEM P.

Bouchet)

G-H. Murichorda rumphiusi (Houart, 1995), Ambon. Scale bars: C: 100 um; D: 50 um (SEM A. Warén)

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

Figure 9 — Radulae

À. Murichorda fiscellum (Gmelin, 1791), Ambon. Scale bar: 50 um (SEM A. Warén)

B. Muricodrupa anaxares (Kiener, 1835), Ambon. Scale bar: 50 um (SEM A. Warén)

20100-23362; stn CP25, 24°44'S, 168°09'E, 230-235 m,

[NIV

BATAUS 2: DW789/22"SSISM66227E, 405-525

ni, Add:

LIFOU 2000: stn 1462, 20°47'S, 167°03'E, 70-120 m,

1 dd, MNAN-IM-2010-21108;

NORFOLK 1: stn 1674, 23°40'S, 168°00'E, 245-253

m, 2 dd, MNHN-IM-2010-23354; stn DW1675,

24°45'S, 168°09'E, 231-233 m, 9 dd, MNHN-IM-

2010-23350; stn CP1677, 24°44'S, 168°09'E, 233-259

m, 1 1v, MNAN-IM-2010-23352; stn DW1724,

23°17'S, 168°14'E, 200-291 m, 1 dd, MNAHN-IM-

2010-2551: sn /DW 1726, 25%418)S, LEOSASIE 185

207 m, 2 dd, MNHN-IM-2010-23353; stn DW1727,

23°17'S, 168°14'E, 190-212 m, 4 dd, MNHN-IM-

2010-23349;

NORFOLK 2: stn DC2089, 24°44'S, 168°09'E, 227-

230 m, 3 dd; stn DW2093, 24°44'S, 168°09'E, 230 m,

1 1v (MNAN-IM-2007-18205), 5 dd;

EBISCO: stn DW2639, 20°47'S, 161°0O1'E, 289-294

m, 1 dd, MNHN-IM-2010-5135;

TERRASSES: stn DW3103, 23°02'S, 168°21'E, 180-

220 m, 2 dd;

EXBODI: stn DW3866, 22°52'S, 169°26'E, 100 m, 1

Iv, MNHN-IM-2009-22839; stn DW3867, 22°52%S,

169°26'"E, 146-610 m, 1 dd, MNHN-IM-2014-2394;

KANACONO: stn DW4763, 23°17'S, 168°15'E, 192-

260 m, 3 dd, MNHN;

KANADEEP: stn DW4945, 25°22'S, 159°43'E, 108-

130 m, 3 dd, MNHN.

Vanuatu. MUSORSTOM 8: stn DW964, 20°20'S,

169°49'E, 360-408 m, 1 dd; stn DWI1101, 15°04S,

(6086 205210 01 dd sm CPIMISS 559

167°03'E, 174-210 m, 1 dd;

SANTO: Pleistocene of Santo, Kere River, near

Finmele (site 1), 15°33'S, 166°57'E, coarse detrital

level, 5 specimens.

Wallis & Futuna. MUSORSTOM 7: Futuna, stn

DW496, 14°20'S, 178°04'W, 250-330 m, 1 dd; Field

Bank, stn DW594, 12°31'S, 174°20'W, 495-505 m, 1

dd.

Tonga. BORDAU 2: stn DW1508, 21°02S ,

175°19°W, 555-581 m, 1 dd, IM-2008-965; stn

DW1512, 21°19'S , 175°01°W, 183-184 m, 1 dd, IM-

2008-9635: SMmDWIS21021M9S"-175010N0225233

m, 1 dd, MNHN-IM-2008-966; stn DW1567, 21°02'S,

175°19°W, 351-356 m, 1 dd, IM-2008-964: stn

DW1605,,2212S 7S 20 1692106 5 dd,

MNAN-IM-2008-967.

Distribution. New Caledonia, Vanuatu, Futuna and

Tonga, living at 100-505 m, but most living specimens

were collected between 230 and 265 m (Fig. 14).

Description. Shell medium sized for the genus, up to

17.3 mm in length at maturity (paratype MNHN-IM-

2000-34208). Length/width ratio 2.0-2.1. Slender,

lanceolate, narrowly ovate, heavy, nodose. Subsutural

ramp broad, strongly sloping, weakly concave.

White or light tan, usually with spiral cords topped

with brown; occasionally pure white. Siphonal canal

brown or light brown and darker on tip. Aperture

white.

Spire high with 3.5 protoconch whorls and teleoconch

up to 5 broadly ovate, convex, weakly shouldered,

nodose whorls. Suture adpressed. Protoconch small.

conical with a very narrow keel abapically. Whorls

smooth, glossy, height and width 900 um. Terminal

lip erect, of sinusigera type.

Axial sculpture of teleoconch whorls consisting of

high, strong, broad, rounded, nodose ribs. First whorl

with 9 ribs, second and third with 9 or 10, fourth with

9-11, last whorl with 8 or 9 ribs. Spiral sculpture of

low, rounded, narrow, smooth, primary, secondary

R. HOUART, D. ZUCCON & N. PUILLANDRE |

NOVAPEX 2O0(HS 12): 1-52, 10 mars 2019

Figure 10 — Spiral sculpture and apertural morpholog)y

. SD. Reunion, MD32, stn DC56, 21°05'S, 55°12'E, 1982, 170-225 m. A. Paratype

2 Di

3 mm: B. Holotype MNHN-IM-2000-34203, 14.3 mm.

Kaimon Maru Bank. SMIB 8. stn DW159, 28/01/1993,

A-B. Cytharomorula arta n

MNHN-IM-2000-34205, 12

C. Cytharomorula elegantula n. sp., New Caledonia,

24°46'S. 168°08' E, 241-245 m, paratype MNHN-IM 20000-34209, 15.7 mm.

D. Cytharomorula grayi (Dall, 1889), Canary Islands, Tenerife, Palma, in shrimp nets, 150-200 m, 1985, RH,

19.6 mm.

E. Cytharomorula fatuhivaensis n. Sp. French Polynesia, Marquesas Islands. Fatu Hiva. 03/09/1990, 10°31'S,.

138°39' W, 210 m, holotype MNHN-IM-2012-18829, 14.7 mm.

F. Cytharomorula springsteeni Houart, 1995, Philippines, tangle nets, 146 m, paratype RE, 12.7 im:

R. HOUART. D. ZUCCON & N. PUILLANDRE New genera and new species Of Ergalataxinae

Figure 11 — Spiral sculpture and apertural morphology

A-B. Cytharomorula grayi (Dall, 1889), MD208, stn DW4877, Walters Shoal, 33°10'S, 43°49'E, 217-256 m,

MNHN-IM-2013-66419, 23.2 mm.

C-D. Orania pseudopacifica n. sp., French Polynesia, Marquesas Islands, Nuku Hiva, 25/08/1997, 8°45'S,

140°13' W, 104-109 m, holotype MNHN-IM-2000-34211, 14.5 mm; D. Detail of the columella. Scale bar: 1

mm.

E. Orania pacifica (Nakayama, 1988), Philippines, Panglao, 55-81 m, detail of the columella, RH. Scale bar: 1

mm.

F. Cytharomorula absidata n. sp., New Caledonia, NORFOLK 2, stn DW2123, 23°18'S, 168°15'E, 187-197 m.

Holotype MNHN-IM-2000-34201, 9.6 mm.

G. Claremontiella adiakritos n. sp., West Mexico, Mazatlan, holotype NHMUK 20180545, 23.3 mm.

R. HOUART, D. ZUCCON & N. PUILLANDRE

NOVAPEX 20(HS 12): 1-52, 10 mars 2019

Figure 12 — Spiral sculpture and apertural morphology

A-B. Tenguella chinoi n. sp. Papua New Guinea, Papua Niugini, stn PR214, 08/12/2012, 1-8 m (no other

information), MNHN, 12.5 mm.

C-D. Tenguella ericius n. sp., Vanuatu, SANTO, stn FM36, 15°22,4'S 167°13'E 27/09/2006 0O-I m, paratype

MNHN-IM-2000-34218, 13.9 mm.

and tertiary cords. First teleoconch whorl with visible

SP. IP. PI and P2: second whorl with SP, adis, IP, PT,

si. P2. s2: third and fourth with additional abis and

tertiary cords. Last whorl with SP, adis, IP, abis, PI,

s1. P2. s2, P3, s3, P4, s4, P5, s5, P6, s6, ADP, ads, MP

and ABP: P4 occasionally broader and higher with

high knobs at intersection with axial ribs, then usually

lighter or white coloured.

Aperture narrow, ovate. Columellar lip narrow with

weak, low parietal tooth at adapical extremity and 2 or

3 elongate, low, narrow folds abapically. Rim weakly

partially erect, adherent at small portion at adapical

extremity. Anal notch deep, broad. Outer lip erect,

smooth with 6 weak, low, elongate denticles within,

consisting of ID, DI-DS. Siphonal canal short, 16-

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species Of Ergalataxinae

A …——"…—"….———." —_

18% of total shell length, narrow, dorsally recurved,

broadly open.

Operculum unknown. Radula with three dimensional

rachidian tooth bearing a long, narrow, projecting

central cusp, a small, narrow lateral denticle on each

side, a medium sized lateral cusp, as broad as central

cusp but shorter, and a few marginal folds. No

marginal cusps. Lateral teeth sickle shaped, relatively

broad.

Remarks. Cytharomorula grayi, with which C

elegantula n. sp. has often been confused, differs in

having a more strongly shouldered, less fusiform shell

with a narrower, more concave and less strongly

sloped subsutural ramp, and broader, stronger, higher

axial ribs, broader and more obvious on shoulder.

These axial ribs are also less numerous, 6 or 7 on the

last teleoconch whorl in C. grayi as opposed to 8 or 9

in C. elegantula n. sp. C. grayi also has a broader, less

fusiform aperture with more obvious denticles.

For differences with Cytharomorula absidata n. sp.,

C. arta n. sp. and C. fatuhivaensis n. sp. see the

descriptions of these species herein.

The Pleistocene specimens from Vanuatu were

collected during an exploration in February 2006 of

fossil sites as part of the SANTO 2006 Expedition

(Lozouet et al. 2011). The fossil specimens come from

the Kere 1 deposit (= Kere Shellbed of Mallik and

Ladd). A well-preserved coral (Flabellum sp.)

collected in this outcrop gave an age of 133 000 yr.

(Lozouet et al., 2011) (see also Houart, 2013b).

Etymology. Elegantula (L): diminutive of elegans,

meaning small and elegant.

Cytharomorula fatuhivaensis n. sp.

Figs 10E; 14; 18A-F

Cytharomorula sp. cf. C. grayi — Trôndlé & Houart,

1992: 88, fig. 48 [(not C. grayi (Dall, 1889)].

Figure 13 (scale bar: 500 um)

A-C. Cytharomorula vexillum Kuroda, 1953.

Cytharomorula grayi — Houart & Tründlé, 2008: 70,

92 [(not C. grayi (Dall, 1889)].

Type material. Holotype MNHN-IM-2012-18829,

French Polynesia, Marquesas, Fatu Hiva, SMCB: stn

CAS303, 10°31'S, 138°39'W, 210 m, lv.

Paratypes: French Polynesia, Marquesas, Fatu Hiva,

10°31'S, 138°39'W, 210 m, 16 lv, MNHN-IM-2000-

34210, 1 1v, RH.

Type locality. French Polynesia, Marquesas, Fatu

Hiva, 10°31'S, 138°39'W, living at 210 m.

Other material examined. MUSORSTOM 9, stn

DW1145, Marquesas, Ua Pou, 9°19'S, 140°06'W, 150-

180 m, MNHN-IM-2008-2929, 7 Iv & dd; stn

DW1146, 9°19'S, 140°06'W, 200 m, MNHN-IM-

2008-2928, 3 dd; stn DW1148, 9°19'S, 140°06'W, 300

m, MNHN-IM-2008-2927, 2 dd; stn DW1197, Hiva

Oa, 9°57'S, 140°02'W, 277-372 m, MNHN-IM2008-

2926, 1 dd; stn DR1247, Fatu Hiva, 10°34S,

138°42'W, 1150-1250 m, MNHN-IM-2008-2925, 1!

dd.

Distribution. French Polynesia, Fatu Hiva, Ua Pou

and Hiva Oa, living at 180-210 m (Fig. 14).

Description. Shell medium sized for the genus, up to

16.5 mm in length at maturity (paratype MNHN).

Length width ratio 1.8-2.0. Biconical, broadly ovate,

heavy, nodose. Subsutural ramp broad, strongly

sloping, weakly concave.

Light tan or whitish with dark brown spiral cords.

Aperture white.

Spire high with 3.5 protoconch whorls and teleoconch

up to 5 broad, strongly convex, weakly shouldered,

nodose whorls. Suture strongly adpressed. Protoconch

small, conical. Whorls smooth, glossy, with a very

narrow keel abapically. Height and width 1000 um.

Terminal lip erect, of sinusigera type.

A-B. Japan, Tosa, 128 m, MNEN, 21.1 mm; C. Protoconch, New Caledonia.

D-E. Cyfharomorula ornamentata (Houart, 1995), South Africa, Durban, RH, 16.8 mm.

E-L. Cytharomorula absidata n. sp.

F-H. New Caledonia, NORFOLK 2, stn DW2123, 23°18'S, 168°15'E, 187-197 m, holotype MNHN-IM-2000-

34201, 9.6 mm; I. New Caledonia, NORFOLK 2, stn DW2123, 23°18'S, 168°15'E, 187-197 m, paratype

MNHN-IM-2007-18225 (BOLD MUBA765-18; GenBank HE584051.1) (fragment of analysed specimen, 8

mm); J-L.New Caledonia, SMIB 5, stn DW93, 22°20'S, 168°43'E, 240-255 m, paratype MNHN-IM-2000-

34202, 13.7 mm.

M-0O. Cytharomorula springsteeni Houart, 1995

M. Philippines, tangle nets, 146 m, paratype RH, 12.7 mm; N-O. New Caledonia, Kaimon Maru Bank, SMIB 8.

stn DW159, 28/01/1993, 24°46'S, 168°08'E, 241-245 m, 13.2 mm, MNHN.

R. HOUART, D. ZUCCON & N. PUILI ANDRI

NOVAPEX 20(HS 12): I

52. 10 mars

2019

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

D ee EN RE

Axial sculpture of teleoconch whorls consisting of

high, strong, broad, rounded, nodose ribs. First and

second teleoconch whorls with 9 ribs, third with 8-10,

fourth with 9 or 10 and last whorl with 6 or 7 ribs.

Spiral sculpture of low, strong, narrow, nodose,

primary, secondary and tertiary cords. First whorl with

visible IP, P1, P2, starting si; second whorl with SP;

IP, PI, si, P2, s2; third and fourth whorls with

additional adis and occasionally with abis and tertiary

cords. Last whorl with SP, adis, IP, abis, PI, s1, P2,

tertiary cords (Fig. 10Ë). Primary and secondary cords

of approximately similar size.

Aperture moderately small, narrow, ovate. Columellar

lip moderately broad with weak, low parietal tooth at

adapical extremity. Rim adherent or partially erect on

a small portion abapically. Anal notch deep, broad.

Outer lip erect, smooth, with 6 strong, elongate

denticles within, decreasing in strength abapically,

consisting of ID, DI-DS$. Siphonal canal very short,

12-14% of total shell length, strongly dorsally bent,

s2, P3, P4, PS5, P6, s6, ADP, MP and ABP and few broadly open.

Operculum and radula unknown.

ke are

3 sn m.

as Lot

TRE Sal 4

Tan Fe

165° 150° 135° 120° 105° 90° ” 39° 45° 60° 75°

Figure 14. Distribution of the species in the Cytharomorula grayi group.

Cytharomorula grayi (@western Atlantic Ocean; Meastern Atlantic Ocean; À western Indian Ocean); + Ce

elegantula n. sp: X C. fatuhivaensis n. sp.

Figure 15 (scale bar: 500 um)

A-E. Cytharomorula arta n. sp.

A-B. Reunion, MD32, stn CP57, 21°05'S, SS°11'E, 1982, 210-227 m, holotype MNHN-IM-2000-34203, 14.3

mm; C-D. Reunion, MD32, stn CP57, 21°05'S, SS°11'E, 1982, 210-227 m, paratype MNHN-IM-2000-34206,

11.9 mm; E. Protoconch, Reunion, MD372, stn DC2, 1982, 21°12'S, 55°49'E, 160-190 m, paratype MNHN-IM-

2000-34204.

F-G. Cytharomorula lefevreiana (Tapparone Canefri, 1880), MD32, Stn DCS85, Reunion, 20°59'S, 55°15'E, 58-

70 m, MNAN, 6.6 mm.

H-P. Cyrharomorula grayi (Dall, 1889)

H-IL. Canary Islands, Tenerife, Palma, in shrimp nets, 150-200 m, RH, 19.6 mm; J. East of Guadeloupe, south of

Desirade, 250 m, RH, 13.8 mm; K. West coast of Barbados, dredged 165 m, RH, 13.2 mm; L South-Eastern

Brazil, Martin Vaz Id, 20°32"S, 28°52' W, 780-795 m (dd), MNHN-IM-2012-33133, 14.1 mm; M. Madeira, RH,

17.8 mm; N. Trophon lowei Watson, 1897, Madeira, lectotype NHMUK 1911.7.17.2, 19.3 mm, photo Harry

Taylor, NHMUK Photographic Unit, © Natural History Museum of London; O-P. Cantharus laevis E.A. Smith.

1890, St Helena, lectotype NHMUK 1889.10.1.2362, 22.2 mm, photo Harry Taylor, NHMUK Photographic

Unit, © Natural History Museum of London.

R. HOUART, D. ZUCCON & N. PUILLANDRE NOVAPEX 20(HS 12): 1-52. 10 mars 2019

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

A ———"—.— ——————

Remarks. Cytharomorula elegantula n. sp. from New

Caledonia (Fig. 16H-N), also described herein, differs

from C. fatuhivaensis n. Sp. in having a usually

lighter, narrower shell, with a length/width ratio of

2.0-2.1 as opposed to the 1.8-2.0 of C. fatuhivaensis

(C. fatuhivaensis is stockier and broader); in having a

narrower and longer siphonal canal, 16-18% of total

shell length, vs 12-14 % in C. fatuhivaensis; a higher,

more elongate spire; narrower, lower, usually more

numerous axial ribs; a comparatively more elongate

aperture; less apparent, usually narrower spiral cords,

except for P4 which is very often higher and more

apparent in C. elegantula than has been observed in

other Cytharomorula species, but which 1s of equal

height to other cords in C. fatuhivaensis.

Etymology. Named after Fatu Hiva Island, the type

locality.

Cytharomorula manusuduirauti n. sp.

Figs 17; 18G-L

Cytharomorula pinguis — Tsuchiya, 2000: 381, pl.

189, fig. 88; Houart, 2008: 23, 198, pl. 394, fig. 6 [not

C. pinguis Houart, 1995)]

Cytharomorula springsteeni — Tsuchiya, 2017: 954,

pl. 247, fig. 4 [not C. springsteeni Houart, 1995)].

Type material. Holotype HMNS 2017.2223

Po.406866, Philippines, Aliguay Island, near 8°44" N,

123°12'E, sand, sandy gravel & sandy mud bottoms,

50-150 m, lv.

Paratypes: Philippines, Balicasag Island, near 9°31"N,

120°41'E, tangle or lumun-lumun net, 100-150 m, Iv,

1 HMNS 2017.2223 Po.524523; Philippines,

Balicasag Island, near 9°31' N, 120°4l'E, tangle or

lumun-lumun net, 50-80 m, dd, 1 HMNS 2017.2223

Po.571785; Philippines, Aliguay Island, rocky mud,

160 m, 1v, 1 RH.

Type locality. Philippines, Aliguay Island, near 8°44"

N, 123°1240'E, sand, sandy gravel & sandy mud

bottoms, 50-150 m.

Distribution. Philippines Islands, living at 150-160 m

and Japan, Ogasawara Islands (Tsuchiya, 2017) (Fig.

17).

Description. Shell small, up to 11.6 mm in length

(holotype). Length/width ratio 1.9-2.0. Biconical,

broadly ovate, heavy, lightly nodose.

Light tan, extreme abapical part of last teleoconch

whorl and siphonal canal dark brown or blackish

brown. Occasionally with weak brown blotches

between axial ribs of last teleoconch whorl. Aperture

white, columellar lip occasionally with brown blotch

abapically.

Spire high with 3+ protoconch whorls (first whorl

broken) and teleoconch of 4 or 4.5 broad, weakly

shouldered, nodose whorls, spire whorls more strongly

shouldered. Suture adpressed.

Protoconch large, high, conical, glossy. Maximum

width 800 um, height 1000 um. Terminal lip of

sinusigera type.

Axial sculpture of teleoconch whorls consisting of

moderately high, broad, rounded ribs, with 8 or 9 ribs

on all whorls, ribs obviously broader on last whorl.

Spiral sculpture of numerous, narrow, rounded cords,

more than 35 on last whorl, equally distant and of

same strength on whole length of shell.

Aperture small, ovate. Columellar lip narrow, smooth

Or With two weak, narrow Kknobs abapically. Rim

weakly erect abapically, adherent adapically. Anal

notch deep, broad. Outer lip smooth, with 6 strong,

elongate denticles within, decreasing in strength

abapically, consisting of ID, D1-D5. Siphonal canal

very Short, 8-10% of total shell length, narrow, weakly

dorsally recurved, broadly open.

Operculum and radula unknown.

Figure 16 (scale bars: 500 um)

A-G. Cytharomorula grayi (Dall, 1889)

A-D. MD208, stn DW4877, Walters Shoal, 33°10'S, 43°49'E, 217-256 m; A-C. MNHN-IM-2013-66419 (BOLD

MUBA781-18; GenBank MK216540), 23.2 mm; D. MNHN-IM-2013-66418, 22.0 mm; E. Protoconch. Walters

Shoal, MD208, stn DW4894, 33°09'S, 43°50'E, 199-261 m, MNHN; F. South Africa, Durban, off Umlaas Canal.

coarse rubble, 14/12/1983, 128 m, NM B6282, 19.2 mm; G. SW Grande Glorieuse Id, BENTHEDI, stn DS93. |

07/04/1977, 11°33'S, 47°16'E, 480-550 m, MNEN, 17.2 mm.

H-N. Cyrharomorula elegantula n. sp.

H-J. New Caledonia, KANACONDO, stn DW4778, 23°03'S, 168°18'E, 170-248 m, holotype MNHN-IM-2013-

63326, (BOLD MUBA780-18; GenBank MK216542), 10.2 mm; K-L. New Caledonia, N.O. "Jean Charcot"

BIOCAL, stn DW66, 03/09/1985, 24°56S, 168°22 E, 505-515 m, paratype MNHN-IM-2010-23359, 16.3 mm:

M. New Caledonia, N.0. "Jean Charcot" BIOCAL, stn DW66, 03/09/1985, 24°56S, 168°22 E, 505-515 m.

paratype MNHN-IM-2010-23359, 16.3 mm; N. Protoconch, New Caledonia, Kaimon Maru Bank. SMIB 8. stn

DW159, 28/01/1993, 24°46'S, 168°08'E, 241-245 m, paratype MNHN-IM-2000-34209.

? ] ? : OC | ) * pe

R. HOUART, D. ZUCCON & N. PUILLANDRE NOVAPEX 20(HS 12): 1-52, 10 mars 2019

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

150°

90° 105° 120° 135? 165°

Figure 17. Distribution of Cytharomorula

manusuduirauti n. Sp.

Remarks. Cytharomorula manusuduirauti n. sp. has

been confused with C. pinguis Houart, 1995 and C.

springsteeni Houart, 1995. However both species

differ from C. manusuduirauti.

Cytharomorula pinguis (Fig. 18M-N) differs in having

a comparatively narrower shell with broader, flatter

and less numerous spiral cords. C. pinguis also has a

comparatively narrower aperture with strong DI

denticle and strong, heavy knobs on the columellar lip,

as opposed to the smooth columellar lip with very

weak adapical knobs in C. manusuduirati n. sp. Both

species have primary, secondary and tertiary cords of

approximately similar strength, which are difficult to

separate. However, C. pinguis has 20-22 broad, flat

cords on the last teleoconch whorl as opposed to 37-

40 narrow, rounded cords in €. manusuduirauti n. sp.

Figure 18 (scale bars: 500 um)

Cytharomorula springsteeni (Fig. 13M-O) also differs

from C. manusuduirauti n. sp. in many ways. The

shell of C. springsteeni is narrower and more elongate,

with a larger, higher aperture and a higher spire. The

spiral sculpture also differs in being more irregular

with higher, weakly broader primary and secondary

cords compared to the narrower tertiary cords (Fig.

10F), rather than the similar sized, more numerous and

crowded cords in C. manusuduirauti n. Sp.

Etymology. Named for the late Emmanuel (Manu)

Guillot de Suduiraut, friend and well-known shell

enthusiast who sent a specimen (now paratype) of this

new species to one of the authors (RH) several years

ago.

Genus Orania Pallary, 1900

Type species by original designation: Pseudomurex

spadae Libassi, 1859 (= Murex fusulus Brocchi,

1814), Mediterranean (Fig. 21A-C).

Diagnosis. Shell with high spire, elongate, ovate,

nodose, not or rarely exceeding 25 mm in length at

maturity. Axial sculpture of last teleoconch whorl

consisting of 7-9 narrow, low, sharp or weakly

rounded ribs. Spiral sculpture of low, narrow, primary,

secondary and tertiary cords.

Aperture ovate, columellar lip weakly concave,

smooth or with weak folds abapically, rim almost

entirely adherent to shell. Anal notch broad, deep.

Outer lip with strong, elongate denticles within.

Siphonal canal short, broadly open ventrally.

Radula of the type species with three dimensional

rachidian tooth bearing a long, moderately broad,

projecting central cusp, a small, narrow lateral

denticle, a moderately long, broad, lateral cusp and

strong marginal folds giving rise to small, short

marginal denticles, and a bifid marginal cusp. Lateral

teeth sickle shaped, with broad base.

A-F. Cytharomorula fatuhivaensis n. sp., French Polynesia, Marquesas Islands, Fatu Hiva, 03/09/1990, 10°31'S,

138°39' W, 210 m.

A-B. Holotype MNEN-IM-2012-18829, 14.7 mm; C. Protoconch, paratype MNHN-IM-2000-34210; D.

Paratype MNHN-IM-2000-34210, 14.8 mm; E-F. Paratype MNHN-IM-2000-34210, 15.5 mm.

G-L. Cytharomorula manusuduirauti n. sp.

G-H. Philippines, Aliguay Island, near 8°44" N, 123°12'40'E, sand, sandy gravel & sandy mud bottoms, 50-150

m, holotype HMNS 2017.2223 Po.406866, 11.6 mm, photo Gary Kidder, The Houston Museum of Natural

Science, Houston, Texas, U.S.A.; L Philippines, Balicasag Island, near 9°31'N, 120°41'E, tangle or lumun-

lumun net, 100-150 m, paratype HMNS 2017.2223 Po.524523, 10.1 mm, photo Gary Kidder, The Houston

Museum of Natural Science, Houston, Texas, U.S.A.; J-L. Philippines, Aliguay Island, rocky mud, 160 m,

paratype RH, 10.2 mm.

M-N. Cytharomorula pinguis Houart, 1995, New Caledonia, Loyalty Ridge, 20°41'S 167°07'E, 360 m, holotype

MNAN-IM-2000-952, 16.7 mm, photo Manuel Caballer (MNHN) E-Recolnat Project: ANR-1 1-INBS-0004.

R. HOUART, D. ZUCCON & N. PUILI ANDRE

NOVAPEX 20(HS 12): 1-52, 10 mars 2019

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

a. —_ —_ —]_—]—]—] ——]— ——"——

Remarks. At present, 32 Recent species and one

subspecies are assigned to Orania in WoRMS

(MolluscaBase 2018). Of these, ©. ornamentata

Houart, 1995 is here transferred to Cyfharomorula,

and ©. albozonata (E.A. Smith, 1890) is removed

from Muricidae to the Buccinoidea (Pisaniidae).

One of the syntypes of Cantharus albozonatus E.A.

Smith, 1890 in the NHMUK, most probably the shell

figured by Smith (1890: 260, pl. 21 fig. 9), is here

illustrated (Fig. 291-J). That species is not a muricid

but rather a buccinid close to Enginella leucozona

(Philippi, 1844) (K. Fraussen, in litt.). However, based

on the images we received from A. Salvador at the

NHMUK, other buccinids are included in the 35

syntypes of C. albozonatus (see Fig. 29K-L), which is

here tentatively assigned to Enginella Monterosato,

joe

One new species of Orania is described herein, while

Muricopsis carnicolor Bozzetti, 2009 (Figs 6E-F;

21D-H) and Morula taiwana are transferred to

Orania. This leaves 33 species and one subspecies in

this genus (Table 2).

The radula of Orania fusulus (Fig. 6G), the type

species, 1s typical for the Orania type described in

Houart (1995: 246, fig. 3), as it has broader cusps and

denticles, small, short marginal cusps and a short

marginal cusp. See also the radula of Orania

carnicolor (Fig. 6E-F).

Orania pseudopacifica n. sp.

Figs 7A-B; 11C-D; 19; 20A-F

?Morula (?Morula) pacifica — Trôndlé & Houart,

1992: 102, fig. 81 [not Orania pacifica (Nakayama,

1988)].

Orania pacifica — Houart & Trôndlé, 2008: 92 [not

Orania pacifica (Nakayama, 1988)].

Type material. Holotype MNHN-IM-2000-34211,

MUSORSTOM 9, stn DW1170, French Polynesia,

Marquesas Archipelago, Nuku Hiva, 8°45'S, 140°

10'W, 104-109 m, lv.

Paratypes: French Polynesia, Marquesas Archipelago,

Nuku Hiva, MUSORSTOM 9, stn DW1170, 8°45'S,

140°, 10'W, 104-109 m, 10 lv, MNHN-IM-2000-

34212, 2 [v, RH; stn CP1239, 09°42'S, 139°04'W, 89-

95 m, 7 1v, MNHN-IM-2000-34213.

Type locality. French Polynesia, Marquesas

Archipelago, Nuku Hiva, 8°45'S, 140°, 10'W, living at

104-109 m.

Other material examined. Marquesas Archipelago.

MUSORSTOM 9: stn DW1144, 09°19'S, 140°04'W,

85-95 m, 6 1v & dd; stn DW1146, 09°19'S, 140°06'W,

200 m, 1 dd; sin DW1148, 09°19'S, 140°06'W, 300 m.

hdd; sin DRIIS1, 0919, 140204, 70-77 1. 10

dd; stn DW1152, 07°59'S, 140°44'W, 85-150 m, 4 dd:

stn CP1158, 07°59'S, 140°44'W, 109-110 m, 1 Iv:; stn

CP1160, 07°58'S, 140°42'W, 49-55 m, 1 Iv; stn

DW1163, 08°57'S, 140°06'W, 78-85 m, 3 dd; stn

DW1164, 08°58'S, 140°06'W, 170-180 m, ! dd; stn

DW1170, 08°45'S, 140°13'W, 104-109 m, 94 Iv & dd:

stn CP1177, 08°45'S, 140°14'W, 108-112 m, 3 dd; stn

CP1178, 08°46'S, 140°15'W, 74-75 m, 2 dd; stn

DRI181, numerous, specimens, 08°465, 140°03'W,

102-130 m, dd; stn DR1182, 08°46'S, 140°04'W, 90-

120 m, 3 dd; stn 1183, 08°46'S, 140°04'W, 86-120 m,

2 dd; stn DR1200, 09°50'S, 139°09'W, 96-100 m, 6

dd; stn DW1208, 09°49'S, 139°10'W, 117 m, 2 Iv &

dd; stn DW1210, numerous specimens, 09°505,

139°01'W, 98-100 m, dd; stn DWI1217, numerous

specimens, 09°44,5'S, 138°50'W, 85-87 m, lv & dd;

stn DR1223, 09°45'S, 138°51'W, 90-150 m, 7 dd; stn

DW1224, numerous specimens, 09°45'S, 138°51"W,

115-120 m, dd; stn CP1227, numerous specimens,

09°44'S, 138°53'W, 84-85 m, lv & dd; stn CP1228,

09°45'S, 138°52'W, 107-108 m, 19 dd; stn DW1230,

09°44'S, 139°07'W, 95-100 m, 18 dd; stn DW1235,

09°42'S, 139°04'W, 105-285 m, 7 dd; stn CP1237,

09°42'S, 139°04'W, 95-305 m, 17 Iv & dd; stn

CP1239, 09°42'S, 139°04'W, 89-95 m, 26 1v & dd; stn

DR1240, 10°25'S, 138°41'W, 70-90 m, 1 dd; stn

DW1241, 10°28'S, 138°41'W, 85-130 m, 1 dd; stn

DR1245, 10°29'S, 138°36'W, 85-130 m, 14 dd; stn

DW1256, 09°25'S, 140°08'W, 70-72 m, 1 dd; stn

DR1257, 09°26'S, 140°08:W, 85-127 1m, 13 dd; ‘sin

DW1260, 09°25'S, 140°07'W, 49-100 m, 9 dd; stn

CP1265, 09°20'S, 140°07'W, 90-92 m, 19 Iv; stn

DW1274, 07°55'S, 140°40'W, 100-120 m, 2 dd; stn

DW1280, 07°59'S, 140°43'W, 87-98 m, 3 dd; stn

DW1281, 07°48'S, 140°21'W, 450-455 m, 1 dd; stn

DW1288, 08°54'S, 139°38"W, 200-220 m, 1 dd; stn

DR1292, 08°54'S, 139°37'W, 95-100 m, 43 dd; stn

DW1293, 08°54'S, 139°38'W, 50 m, 4 1v & dd; stn

CP1294, 08°54S, 139°38"W, 100 m, 4 Iv: sim

DR1297, 08°54S, 139°37"W, 90-150 m, 1 [v: stn

DR1298, 08°49'S, 140°17'W, 305 m, 3 dd; stn

CP1304, 08°54'S, 140°14'W, 50-58 m, 2 dd; stn

DRI1305, numerous specimens, 08°54'S, 140°15'W,

90-155 m, dd.

Distribution. French Polynesia,

Archipelago, 7°48'S — 10°29'S,

140°44'W, living at 50-117m (Fig. 19).

Marquesas

138°56W. —

Description. Shell medium sized for the genus, up to

16.8 mm in length at maturity (paratype MNHN).

Length/width ratio 1.7-1.8. Biconical, broadly-ovate,

heavy, weakly nodose, finely squamous. Subsutural

ramp broad, strongly sloping, weakly concave.

White, light tan or light peach. Aperture white.

occasionally tinged with light pink at edge of

columellar lip.

Spire high with 3.5-4 protoconch whorls and

teleoconch of up to 5 broad, weakly shouldered

whorls. Suture of whorls adpressed. Protoconch small.

conical, acute. Whorls smooth, glossy, with a very

narrow, single keel abapically. Height 900 um, width

750-800 um. Terminal lip of sinusigera type.

R. HOUART, D. ZUCCON & N. PUILLANDRE

NOVAPEX 20(HS 12): 1-52, 10 mars 2019

a] ——]—

Axial sculpture of teleoconch whorls consisting of

low, broad, rounded ribs. First and second teleoconch

Whorls with 7-9 ribs, third with 8 or 9, fourth with 7 or

8, last whorl with 6 or 7 ribs. Spiral sculpture of high,

rounded, narrow, finely squamous, primary, secondary

and tertiary cords and few additional narrow threads.

First whorl with SP, IP, P1. second whorl with SP, IP,

Starting abis, P1, sl, third with SP, starting adis, IP,

abis, PI, t, sl, fourth with SP, adis, IP, abis, PI, &, si.

last teleoconch whorl usually with SP, adis, IP, abis.

PCs 2,52, 1 P5,53, P4, sd, t, PS5, 85, ADP, MP

and one or two additional threads. SP sometimes split,

broad, strongly squamous, occasionally with high,

projecting scales.

Aperture moderately large, narrow, ovate. Columellar

Hp narrow with 2-4 strong knobs on whole length,

deeply extended within aperture (Fig. 11D). Rim

partially erect, adherent at small portion adapically.

Occasionally with weak, low parietal tooth at adapical

extremity. Anal notch deep, broad, with expanded lip.

Outer lip erect, crenulated, with 6 strong, elongate

denticles within, decreasing in strength abapically,

consisting Of ID, DI-DS. Siphonal canal short, broad,

dorsally recurved, broadly open.

Radula of three-dimensional type consisting of a

rachidian with à narrow, long central cusp, a short

lateral denticle and a fairly broad, moderately long

lateral cusp on each side, few, obvious, marginal folds

and a very short, broad, marginal cusp. Lateral teeth

sickle shaped with broad base and narrow extremity.

Remarks. Orania pseudopacifica n. sp. may be

compared with only three species: ©. pacifica

(Nakayama, 1988), with which it was confused in

recent literature, ©. rosea Houart, 1996 and ©.

pleurotomoides (Reeve, 1845).

Orania pacifica (Figs 7C; 11E; 20G-H) differs in

having a more strongly squamous shell with a less

obvious subsutural primary cord (SP); a narrower

subsutural ramp; and differently-arranged, narrower

and weaker columellar folds, which are less deeply

extended within the aperture (Fig. 11E). The anal

notch is also narrower in ©. pacifica, With a less

expanded lip. Orania pacifica is widely distributed in

the Indo-West Pacific, from Japan (type locality),

Taiwan, the Philippines, and several other localities in

the Indo-West Pacific, to Mozambique and South

Africa in the Indian Ocean.

Orania rosea (Fig. 20K-N), described from Reunion

Island in the Indian Ocean but also found in the

Philippines grows much larger, up to 23-25 mm in

length, is more fusiform with a more squamous shell.

The aperture is obviously larger and comparatively

more strongly ovate, with a relatively smooth

columellar lip except for the low abapical folds. It also

has a less obvious subsutural cord.

Orania pleurotomoides (Fig. 201-J) has à more

strongly shouldered shell with a less bent, narrower

subsutural ramp; a narrower subsutural cord: and

much narrower or absent secondary cords. In this

species, the last teleoconch whorl is more sharply bent

anteriorly while the siphonal canal is obviously

narrower. The aperture is more triangular with a series

Of low abapical folds decreasing in strength

abapically; a narrow, more obvious parietal tooth and

a narrower anal notch with a less expanded lip.

To my knowledge, none of these three species occur

in French Polynesia.

Etymology. Pseudo (G) = false. Named

pseudopacifica because it was misidentified as Orania

pacifica in recent literature.

30°

45°

l ji > } l H

90° 105° 120° 135° 150° 165° 180° 165° 150°

Figure 19. Distribution of Orania pseudopacifica n.

Sp.

Orania carnicolor (Bozzetti, 2009) comb. nov.

Figs 6E-F; 21D-H

Muricopsis carnicolor Bozzetti, 2009: 19, text figs.

Type material. Holotype MNHN-IM-2000-22900.

Type locality. South Madagascar, Lavanono, 280 kms

southwest of Tolagnaro.

Other Material examined. Madagascar. ATIMO

VATAE, stn TS02, 25°01.3'S, 47°00.5'E, 18 m, 1dd;

stn BB03, 25°26.4'S, 44°56.1'E, 14-18 m, 3 lv,

(MNHN-IM-2009-22465, MNHN-IM-2009-22485), 1

dd; stn BB04, 25°26.9'S, 44°55.9'E, 14-18 m, 5 1v &

dd; stn BS04, 25°26.9'S, 44°55.9'E, 14-18m, 1 dd; stn

TS04, 25°02.3'S, 47°00.3'E, 22-24 m, 2 dd; stn TBOS,

25°02.2'S, 47°00.4'E, 23 m, 3 1v, 1 dd (MNHN-IM-

2009-14481, MNHN-IM-2009-22445); BP06,

25°25.4'S, 44°54.5-T'E, 19-20 m, 6 Iv (MNAN-IM-

2009-22501-05, MNHN-IM-2009-22511-12); stn

BP07, 25°27.2-6'S, 44°55.6-9'E, 18-22 m, 4 dd; stn

BPO8, 25°27.3-6'S, 44°55.2"E, 25-26 m, 2 dd, stn

BP10, 25°25.5-8'S, 44°54.4-6'E, 23-25 m, 4 1v & dd;

stn TA10, 25°28.3'S, 44°55.6'E, 23 m, 1 dd; stn BS11,

15°28:65, AA°S0SE, 8-11 M, 1 0d; sin BP2i,

25°23.1-2'S, 44°51.4-6'E, 20-23 m (MNHN-IM-2009-

22505); stn BP22, 25°23.4'S, 44°51.7'E, 20-22 m, 5 Iv

& dd; stn BP31, 25°23.6-7'S, 44°53.3-5S'E, 10-12 m, 1

dd; stn BP33, 25°25.80-8'S, 44°55.7-8'E, 11-13 m, 4

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

D op

lv & dd; stn TA35, 24°45.6'S, 47°12.4'E, 5-6 m, 1 Iv,

1 dd, (MNHN-IM-2009-22441); stn BP36, 25°21.9',

AAS502E, 10-17:0m, 1 Iv: sin BP357, 2502274275;

44°50.2-T'E, 19-20 m 1 1v, 1 dd; stn BP41, 25°22.9-

23.2'S, 44°51.0-6'E, 19-21 m, 3 1v; stn BP42, 25°22.8-

23.1S, AS NE, 18-21rm 10e dd sm CPES 10;

25°14.6'S, 47°09.1'E, 79-80, 1 Iv (MNEHN-IM-2009-

22439); stn DW3519, 24°51.9'S, 47°28.0'E, 80-83 m,

6 lv & dd; stn CP3545, 25°29'S, 46°42'E, 108-110 m,

1 dd; stn CP3546, 25°22.7'S, 46°42.5'E, 84-85 m, 5 [v

& dd; stn CP3547, 25°18.0'S, 46°40.3'E, 69-70 m, 5

lv; stn DW3550, 26°03.2'S, 45°32.1'E, 98 m, 2 dd; stn

CRSS72 2SM0TS, M SE 75 77m Sr dde tn

CP3579, 25°54.5'S, 45°33.2'E, 65-66 m, 1 1v, 1 dd; stn

DW3606, 25°48.4'S, 44°5S1.1'E, 44-46 m, 17 dd; stn

DW3608, 25°39.4'S, 44°53.0'E, 37-38 m, 16 lv & dd

[MNHAN-IM-2009-14366 (BOLD MUBA290-I5;

GenBank MK216552), MNHN-IM-2009-22448

(BOLD MUBA312-15; GenBank MK216545),

MNEAN-IM-2009-22449 (BOLD MUBA313-I5;

GenBank MK216557)]; stn DW3609, 25°343S,

MÉSS2E 52 Um 3 dd sin CP5624, 258810S;

45°57.0'E, 63 m, 11 1v & dd; stn DW3626, 25°30.2%S,

45°46.3'E, 41-42 m, 1 dd.

Distribution. South Madagascar, living at 13-84 m.

Remarks. Orania carnicolor Was described as a

species of Muricopsis, a genus unknown in the Indo-

West Pacific, based on three dead collected

specimens, but the general shape of these shells are

reminiscent of several species of Orania, including ©.

fusulus, the type species. Its assignment to

Ergalataxinae is confirmed by the morphology of the

radula, here 1llustrated for the first time (Fig. 6E-F).

Orania castanea (Küster, 1858)

Figs 6H; 211-K

Purpura castanea Küster, 1858: 170, pl. 28, figs 8, 9.

Cominella fasciata Sowerby II, 1886: 3.

Cominella unifasciata Var. concolor Sowerby I,

1897: 4.

Cominella unifasciata nigronodulosa Turton, 1932:

SSD ie 02

Thais castanea (Krauss) — Barnard, 1974: 691.

Thais castanea — Kensley, 1973:146, fig. 507.

Thais castanea (Küster, 1886) — Barnard, 1959: 224;

Richards, 1981: 56, pl. 29, fig. 230; Kilburn &

Rippey, 1982: 89, pl. 20, fig. 7; Steyn & Lussi, 1998:

96, fig. 371.

Nucella castanea (Küster, 1886) — Houart et al.

2010: 197.

Type material. Purpura castanea: "There is no

indication. The whereabouts of this type (even if from

Kuster's coll.) are unknown. P. castanea Was founded

on material coming from F. Krauss, probably

destroyed during the war in Stuttgart" (R. Janssen, in

litt.); Cominella fasciata: three syntypes NHMUK

86.4.2.10; Cominella unifasciata var. nigronodulosa:

one syntype Oxford Museum.

Type localities. Purpura castanea: Cape Agulhas;

Cominella fasciata: Port Elizabeth; Cominella

unifasciata var. concolor: Natal; Cominella

unifasciata nigronodulosa: Port Alfred.

Distribution. South Africa, from False Bay to the

south coast of Natal, and Walters Shoals, south of

Madagascar, approximately 33°12'S, 43°50'E (new

locality).

Other material examined. South Africa: W Cape

Province, Witsand, 2 Iv (RH); East London, 3 Iv (RH)

(radula illustrated); East Cape Province, WSW of East

London, 6 1v (RH); Gonubie, East London, 1 1v (RH);

Haga Haga, 3 1v & dd (RH); South Africa (no other

data), 10 dd (RH).

Walters Shoal seamount: MD208: stn WSO8,

33° 14'S, 43°56'E, 30-33 m, MNHN-IM-2013-66431, 1

Iv (BOLD MUBA784-18; GenBank MK216551); stn

WS08, 33°14'S, 43°56'E, 30-33 m, MNHN-IM-2013-

66432, (BOLD MUBA785-18; GenBank MK216550)

1 1v; stn WB09, 33°14'S, 43°56'E, 27-30 m, MNHN-

IM-2013-66434, (BOLD MUBA786-18: GenBank

MK216541), 1 Iv; stn WB10, 33°09'S, 43°52'E, 30 m.

MNEN-IM-2013-66424, (BOLD MUBA783-18:

GenBank MK216555), 1 Iv; stn WB10, 33°095,

43°52'E, 30 m, 2 1v.

Figure 20 (scale bar: 500 um)

A-F. Orania pseudopacifica n. sp. French Polynesia, Marquesas Islands, Nuku Hiva, 8°45'S, 140°13' W. 104-

109 m.

A-D. Holotype MNHN-IM-2000-34211, 14.5 mm; E-F. Paratype MNHN-IM-2000-34212, 14.0 mm.

G-H. Orania pacifica (Nakayama, 1988), Philippines, Balut Island, RH, 17.5 mm.

I-J. Orania pleurotomoides (Reeve, 1845), Philippines, Mactan Island, 60 m, RH, 16.1 mm.

K-N. Orania rosea Houart, 1996

K-L. Reunion Island, off Saint Pierre, 21°21'S 55°27'E, 73-77 m, holotype MNHN-IM-2000-910, 18.6 mm.

photo Manuel Caballer (MNHN) E-Recolnat Project: ANR-11-INBS-0004; M-N. Philippines Mactan Id. Cebu.

120 m, sand and broken coral, RH, 20.9 mm.

R. HOUART, D. ZUCCON & N. PUILLANDRE

NOVAPEX 20(HS 12): 1-52. 10 mars 2019

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new Species of Ergalataxinae

A —_——.—— —— _— _—_—

Remarks. In Claremont et al (2013: 23, fig. 1; 24, fig.

2), subclade Z contains "Thais" castanea, Which was

well supported in a cluster with Orania fischeriana

(Tapparone Canefri, 1882), Orania pacifica and O.

mixta Houart, 1995, and we here confirm this result

(Fig 3). "Thais" castanea is clearly not a member of

any rapanine or ocenebrine genus, and should be

reassigned to Ergalataxinae. This assignment 1s further

confirmed by its radula morphology (Fig. 6H), which

is Similar to Orania.

The date of Küster's publication of the original

description of Orania castanea is wrongly indicated

as 1886 by numerous authors, including one of the us

(RH) and in WoRMS (MolluscaBase 2018). This is

probably due to Barnard (1959: 224), who was

probably the first to make this mistake.

Six live specimens were collected during the recent

MD 208 expedition to the Walters Shoal Mountain

(Fig. 21K). This range extension, even if not

unexpected in this area, remains very interesting.

Orania pachyrhaphe (E.A. Smith, 1879)

Fig. 22A-E

Fusus pachyrhaphe E.A. Smith, 1879: 205, pl. 20, figs

91, SA

Bedeva bireleffi — Tsuchiya, 2000: 381, pl. 189, fig.

90 [not Bedevina birileffi (Lischke, 1871)].

Type material. Lectotype NHMUK 1878.10.16.2/1,

here designated; 1 paralectotype (©. pachyrhaphe)

NHMUK 1878.10.16.2/2 and 3 paralectotypes

(actually B. birileffi) 1878.11.7.28.

Type locality. Japan, Ukushima, Goto Islands,

33°15.5' N, 129°5'E, 11 fathoms (20 m) (station 8 in

E.A. Smith, 1879: 182) (here selected).

Figure 21 (scale bar: 500 um)

Material examined. Lectotype NHM UK and 4

paralectotypes; Japan, Okinawa, Haneji Inland Sea, on

rocks, 0.3-1 m, 1v, 13 RH.

Distribution. Japan, Okinawa and Goto Islands.

Remarks. Smith (1879: 205) mentioned two

specimens in his description, one of 21 mm

(illustrated in his plate 20, fig. 37) and one of 16 mm

(his plate 20, fig. 37a). However, further in his text he

wrote: "The largest specimen from the latter locality

differs from the rest..." and "I feel convinced that they

all belong to one and the same species". Thus we may

conclude that more than two specimens were

involved, and there are indeed five syntypes in

NHMUK, of which two are the ones originally

illustrated. However, the specimens illustrated by E.A.

Smith belong to two different species. His fig. 37 is

clearly the largest specimen deposited in NHMUXK,

here designated lectotype of ©. pachyrhaphe (Fig.

22A-B), but his fig. 37a and two other paralectotypes

are what is now known as Bedevina birileffi (Lischke,

1871), the intricate history of which was analysed by

Houart et al. (2013). The smallest of the four

paralectotypes is a young specimen of O. pachyrhaphe

(Fig. 22C). À specimen of B. birileffi is here

illustrated (Fig. 22H-1) together with the paralectotype

figured by E.A. Smith (1879: fig. 37a) (Fig. 22F-G).

Bedevina was analysed in Claremont et al (2013) and

its validity in the FErgalataxinae was confirmed.

However, the boundaries of this genus remain unclear

because the same clade contained some species

included in Spinidrupa (but not the type species). The

definition of Bedevina and Spinidrupa thus remain to

be delimited exactly, but, as noted in Claremont et al.

(2013: 27), the conservative retention of both genera

is recommended until further analyses are undertaken.

A-C. Orania fusulus (Brocchi, 1814), A. Spain, Fuengirola, RH, 22.6 mm; B. Italy, Piacenza, Piacenziano,

Castell'Arquato, Pliocene, RH, 23.2 mm, C. West Africa, Mauritania, RH, 17.4 mm.

D-H. Orania carnicolor (Bozzetti, 2009)

D-E. Madagascar, Lavanono, holotype MNHN-IM-2000-22900, 9.15 mm, photo Manuel Caballer (MNHN) E-

Recolnat Project: ANR-11-INBS-0004; E-G. ATIMO VATAE, stn BP22, Madagascar secteur Ouest de

Lavanono, 25°23.4'S, 44°51]1.7'E, 20-22 m, MNEN, 11.3 mm; H. Protoconch, ATIMO VATAE, stn CP3545.

25°29'S, 46°42'E, 108-110 m, MNEN.

I-K. Orania castanea (Küster, 1858)

I-J. South Africa, Haga Haga, RH, 13.5 mm; K. MD208, stn WB10, Walters Shoal, 33°09,1'S, 43°51.8'E. 30 m.

MNHN-IM-2013-66424, (BOLD MUBA783-18; GenBank MK216555), 12.4 mm. | |

L. Orania gaskelli (Melvill, 1891), Papua New Guinea, Hansa Bay, RH, 16.6 mm.

M:-N. Orania serotina (A. Adams, 1853), Philippines, Cebu, Mactan Id, Punta Engaño, RH, 14.3 mm.

O-P. Usilla avenacea (Lesson, 1842), Marquesas, Ua Huka, RH, 11.1 mm.

Q. Orania bimucronata (Reeve, 1846), Malaya, Tioman, RH, 14.6 mm.

R. HOUART, D. ZUCCON & N. PuII LANDRI NOVAPEX 20(HS 12): 1-52, 10 mars 2019

R. HOUART, D. ZUCCON & N. PUILLANDRE

« c 2] d a] »1 » Ç 4 c ec 1 Le £

New genera and new species of Ergalataxina

A EEE

Orania pachyrhaphe differs from B. birileffi in

reaching a larger size relative to the number of

teleoconch whorls; in having a lower spire; in having

a spiral sculpture consisting of narrow primary cords

together with weakly narrower secondary cords and

small tertiary cords versus rounded, more crowded

primary and secondary cords of approximately similar

size; and in that adults specimens of B. birileffi lack

tertiary cords. In addition, the siphonal canal of ©.

pachyrhaphe is relatively broader and the shell is

more squamous. Its inclusion in Orania is based only

on its shell morphology, which is close to ©. fusulus,

the type species of Orania.

Genus Pascula Dall, 1918

Type species by original designation: Trophon citricus

Dall, 1908, Easter Island.

Evokesia Radwin & D'Attilio, 1972

Type species by original designation:

rufonotatum Carpenter, 1864.

Sistrum

Diagnosis. Shell with high spire, broadly elongate,

ovate, nodose, not or rarely exceeding 20 mm in

length at maturity. Axial sculpture of last teleoconch

whorls consisting of moderately broad, high, rounded

ribs. Spiral sculpture of moderately high and broad

primary cords, narrow secondary cords and few

threads.

Aperture ovate. Columellar lip weakly concave,

smooth or with weak folds abapically, rim weakly

erect abapically, adherent at small portion adapically.

Outer lip with narrow, elongate denticles within.

Radula of the type species illustrated by Rehder

[1980: 135, pl. 3 (3-4)] with three dimensional

rachidian tooth bearing a long, narrow, projecting

central cusp, a very small, narrow lateral denticle, a

moderately long, moderately broad, lateral cusp and

very weak marginal folds. No marginal cusp. Lateral

teeth sickle shaped, with broad base.

Remarks. Eight Recent species are assigned to

Pascula in WoRMS (MolluscaBase 2018): Pascula

citrica (Dall, 1908) (Easter Island and French

Polynesia); P. darrosensis (E.A. Smith, 1884) (Indo-

West Pacific); P. muricata (Reeve, 1846) (Indo-West

Pacific); P. ochrostoma (Blainville, 1832) (Indo-West

Pacific); P. ozenneana (Crosse, 1861) (Indo-West

Pacific); P. philpoppei Houart, 2018 (Philippines and

Japan); P. rufonotata (Carpenter, 1864) (Baja

California, Mexico and Galapagos Islands); and P.

submissus (E. A. Smith, 1903) (Indo-West Pacific).

One species, P. palmeri (Powell, 1967) (Fig. 220-P)

from New Zealand, is here added as a new

combination. In the original description by Powell

(1967: 193), P. palmeri was assigned to Morula.

Although molecular genetic analyses are not yet

available, the shell and radula morphology of P.

palmeri (Figs 7G-H; 220-P) is very close to P. citrica

(Fig. 22K-L), the type species of Pascula. Thus, while

waiting for a more complete genetic analysis of this

genus, we suggest assigning P. palmeri to Pascula

rather than to Morula, as the type species, Morula uva

(Rüding, 1798), is quite different from P. palmert.

Genus Tenguella Arakawa, 1965

Type species by original designation:

granulata Duclos, 1832, Indo-West Pacific.

Purpura

Diagnosis. Shell with high spire, broadly ovate,

nodose, rarely exceeding 30 mm in length at maturity.

Axial sculpture of last teleoconch whorl consisting of

8-10 broad, rounded, nodose ribs. Spiral sculpture of

strong, high, broad primary cords and numerous

threads, forming relatively high nodes at intersection

with axial ribs.

Aperture ovate. Columellar lip sinuous, smooth or

with 1 or 2 narrow knobs abapically, rim very weakly

erect abapically, otherwise adherent. Outer lip with

strong denticles within.

Radula of the type species (Fig. 8C) with three

dimensional rachidian tooth bearing a long,

moderately broad, projecting central cusp, a very

small, narrow lateral denticle, a moderately long,

broad, lateral cusp and strong marginal folds giving

rise to small, short marginal denticles, and a bifid

marginal cusp. Lateral teeth sickle shaped, with broad

base.

Figure 22

A-E,. Orania pachyrhaphe (E.A. Smith, 1879)

A-B. Japan, Ukushima, Goto Islands, 33°15.5'N, 129°5'E, 11 fathoms (20 m), lectotype (here designated)

NHMUK 1878.10.16.2/1, 20.9 mm; C. Paralectotype NHMUK 1878.10.16.2/2, 11 mm; D-E. Okinawa. Haneji

Inland Sea, muddy areas on rocks, 0.3 — 1 m, RH (D. 26.4 mm; E. 23.7 mm).

FE-L. Bedevina birileffi Lischke, 1871)

F-G. Paralectotype of Fusus pachyrhaphe E.A. Smith, 1879, NHMUXK 1878.11.7.28, 15.8 mm: H-L. Indonesia.

near Belintung Is, Carima Straits, 15-20 m, RH, 14.8 mm.

J. Lataxiena fimbriata (Hinds, 1843), West Sumatra, RH, 23.7 mm.

K-L. Pascula citrica (Dall, 1908), Easter Id, Hanga Piko, RH, 13.8 mm.

M-N. Pascula darrosensis (E.A. Smith, 1884), Philippines, Sulu Sea, RH, 10.3 mm.

O-P. Pascula palmeri (Powell, 1967), New Zealand, Poor Knights Islands, on rock wall, 20 m, RH. 20.6 mm.

R. HOUART, D. ZUCCON & N. PUILLANDRE NOVAPEX 20(HS 12): 1-52. 10 mars 2019

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

A do oo 000087 Ve pm ZZ———_——_—

Remarks. The genus Tenguella currently includes

five Recent species: Tengualla ceylonica (Dall, 1923),

T. granulata (Duclos, 1832), T. hoffmani Houart,

2017, T. marginalba (Blainville, 1832) and 7. musiva

(Kiener, 1835). Two new species are described below

giving a new total of 7 species in this genus (Table 2).

Tenguella granulata (Duclos, 1832)

Figs 8C; 23; 25Q-S

Purpura granulata Duclos, 1832: 111, pl.2, fig. 9.

Purpura tuberculata Blainville, 1832: 204, pl.9, fig. 3.

Purpura tuberculata var. cingulifera Kiener, 1835:

pl.S, fig. 10°.

Type material. Purpura granulata: not located (not in

MNEIN). The specimen 1llustrated by Duclos 1s here

designated as lectotype (ICZN Art. 74.4). The original

drawing 1s excellent (Fig. 23) and easily recognizable

while the length given by Duclos (1 pouce =

approximately 27 mm) fits perfectly for this species;

Purpura tuberculata: 4 syntypes MNHN-IM-2000-

777; Purpura tuberculata var. cingulifera: Not located

in MENG nor in MNEN.

Type localities. Purpura granulata: Australia

("Nouvelle-Hollande"); Purpura tuberculata: Red Sea

and Madagascar, here restricted to the Red Sea;

Purpura tuberculata var. cingulifera: unknown.

Distribution. Tenguella granulata is a very common

species living throughout the Indo-West Pacific.

Remarks. Tenguella granulata is here compared with

two species, T. chinoi n. Sp. and T: ericius n. sp.,

collected during the MNHN/IRD expeditions in The

Marquesas, Vanuatu and Papua New Guinea and

known from a few other localities (see below).

Fig. 23. Purpura granulata. Original illustration from

Duclos, 1832.

Tenguella chinoiï n. sp.

Figs 8A-B; 12A-B; 24; 25A-H

Morula mutica — Wilson, 1994: 44, pl. 5, figs 4 a-b

[not Azumamorula mutica (Lamarck, 1816)].

Azumamorula sp. — Dharma, 2005: 168, pl. 59, figs

17a, b.

Morula (Habromorula) sp. — Tsuchiya, 2017: 958,

be 2S Es 1

Type material. Holotype NHMUK 20080772/1

(reconstructed aperture and two pieces of tissue

labelled "MORMUD.GM") (GenBank: FN677418),

Guam, Mangalao, Pago Bay, near shore of eroded

limestone cliffs, in open bay with near-shore fringing

reef (syntopic with T. ericius n. sp).

Paratypes: Guam, Mangalao, Pago Bay, near shore of

eroded limestone cliffs, in open bay with near-shore

fringing reef, 3 1v, NHMUK 20080772/2 (fragments)

(ex NHMUK 20080772).

Vanuatu, SANTO 2006, stn VMI2, Palikulo

Peninsula, 15°39'S, 167°16'E, 0-1 m, intertidal, soft

bottom, 1 1v, MNHN-IM-2000-34214 (syntopic with

Tenguella granulata), stn RAPIS, North Tuvana

Island, 15°37'S, 167°0l'E, c. 2 m, 6 1v, 4 MNHN-IM-

2000-34215; 2 RH (syntopic with Tenguella

granulata); stn FM36, Vaucluse Passage, 15°22,4 ,

167°13'E, 0-1 m, intertidal, 1 1v, MNHN-IM-2000-

34216 (syntopic with Tenguella granulata and T.

ericlus n. SP.).

Papua New Guinea, PAPUA NIUGINI, stn PM41,

Wonad Island, 05°08'S, 145°49'E, sandy beach and

intertidal rocks, 0-1 m, 6 Ivy, MNHN-IM-2000-34217

(syntopic with Tenguella granulata and T. ericius n.

Sp.);

Type locality. Guam, Mangalao, Pago Bay, near

shore of eroded limestone cliffs, in open bay with

near-shore fringing reef.

Other material examined. Papua New Guinea:

PAPUA NIUGINI, stn PFMO3, Bilbil Island, exposed

steep, fringing reef, under rocks, exposed, 05°15'S,

145°4TE, 2 m, 2 Iv (syntopic with Tenguella

granulata), stn PM25, Barag Island, O5°01S,

145°48'E, fringing reef, on narrow barrier island, 3 lv:

stn PM33, Rempi area, bay on south side, coral reef,

O5S°02'S, 145°48'E, coral reef, 1 1v; stn PM43, Wonad

Island, 05°08'S, 145°49'E, night tide, sandy beach and

intertidal rocks, 0-1 m, 1 1v (syntopic with Tenguella

granulata), sn PR76, Rempi area, SW Hargun Island,

05°02'S, 145°48'E, 2-15 m, 1 Iv (syntopic with

Tenguella granulata); stn PR192, Kranket Island.

Cape Jantzen, 05°12'S, 145°49'E, 3-60 m, 2 1v: stn

PR204, BilBil L., southern tip, no coordinates, 3-15 m.

2 Iv (syntopic with Tenguella granulata); stn PR214.

Tab Island, no coordinates, 1-8 m, 4 Iv & 1 dd

(syntopic with Tenguella granulata and T. ericius n.

Sp.); Hansa Bay, Madang Province, Laing Island, 2 1v.

juv, (RH).

R. HOUART, D. ZUCCON & N. PUILLANDRE

NOVAPEX 20(HS 12): 1-52, 10 mars 2019

——__—.———————————————.

China Sea: (no other data), 11 1v (ex Staadt coll.

1969) (MNHN).

Western Australia: Exmouth, 1 1v (RH).

Distribution. Western Australia, Exmouth (1 RH),

East Java (Dharma, 2005), Japan, Izu Islands and

southwards (Tsuchiya, 2017), Papua New Guinea and

Vanuatu; perhaps also the China Sea (MNHN).

Intertidal to 3 m depth, sandy beach and intertidal

rocks (Fig. 24).

Description. Shell small for the genus, up to 13.6 mm

in length at maturity (paratype MNHN-IM-2000-

34216) (Fig. 25B-C). Length/width ratio 1.3-1.4,.

Broad, heavy, nodose. Subsutural ramp broad, weakly

sloping, concave.

Blackish-brown or black. Aperture white within with

narrow black coloured narrow band at edge and dark

bluish broad blotch at adapical part of strong curve of

columellar lip.

Spire high, low. Teleoconch of up to 5 or 6 strongly

convex, shouldered, nodose whorls. Suture adpressed.

Protoconch and early teleoconch whorls strongly

eroded in type material.

Axial sculpture of teleoconch whorls consisting of

low, broad, nodose ribs with strong, low, broad nodes

when crossing primary spiral cords. Last teleoconch

whorl with 8 or 9 low, occasionally indistinct ribs,

eroded on earlier whorls. Spiral sculpture of low,

strong, rounded and flat, broad, nodose, primary cords

and low, narrow threads. Last teleoconch whorl with

SP, P1-P5 with 2-4 flat, narrow, smooth threads

between each pair of cords. SP very broad, obvious,

usually broader than other spiral cords. P1-P2 similar

in size, P3-PS decreasing in strength abapically. PS

very narroW.

Aperture large, very narrow, strongly ovate, sinuous.

Columellar lip broad, strongly curved, with one or two

very narrow knobs abapically and low parietal tooth at

adapical extremity. Rim weakly partially erect

abapically, adherent adapically. Anal notch narrow,

moderately deep. Outer lip weakly erect, smooth, edge

broad with 4 strong, high denticles within, consisting

of D1-D4. DI very broad and high, D2 more than half

the size of D1, D3 and D4 lower and narrower, more

elongate within aperture. Siphonal canal very short, 9-

10% of total shell length, narrow, straight, broadly

open.

Radula consisting of a rachidian with a moderately

large, long, central cusp, a short, narrow, lateral

denticle and a moderately broad, long, lateral cusp on

each side, 2 or 3 obvious, short, marginal denticles

and a short marginal cusp. Lateral teeth sickle shaped

with broad base and narrow extremity.

Remarks. Based on molecular data, Tenguella chinoiï

n. sp. was separated from Tenguella granulata, T.

ceylonica, T. musiva and T. marginalba by Claremont

et al (2013, as "Tenguella n. sp."). These five species

formed a well-supported subclade. We here confirm

this result (Fig 3).

The material studied in Claremont et al. (2013) (as

Tenguella n. sp.) consists of six specimens, four of

them were crashed to obtain the soft parts, two

remained intact, one of those is here illustrated (Fig.

25P). These two intact specimens are obviously what

is here described as T. ericius n. sp. and $o, the

NHMUK material was first identified as this species.

However, in the meantime, two specimens of T7.

ericius n. Sp. from the PAKAIHI I TE MOANA

expedition in the Marquesas Archipelago were also

analyzed in MNHN and proved to be different with

around 10% of genetic distance (COI gene) from the

specimen analyzed in Claremont et al. (2013).

This unexpected result could mean two things: either

the Marquesas species is a sibling species but is

genetically different, or the material in Claremont et

al. (2013) was composed of different species and the

specimen analyzed by these authors was not T. ericius

n. sp. After a very careful examination of the crushed

shells, the reconstruction of the aperture of one

studied specimen was possible (Fig. 25A) and it is

clearly this second hypothesis which is the correct

one. The species studied in Claremont et al. (2013) is

clearly the species described here as T. chinoi n. sp.

while the two intact, not studied specimens are

undoubtedly T: ericius n. sp.

These two species, T. ericius n. sp. and T. chinoi n. sp.

are syntopic in some localities and thus also in Guam,

where the specimens analyzed in Claremont et al.

(2013) were collected. Thanks now to the results

obtained by the analysis of T. ericius n. sp. from the

Marquesas it 1s also proved that the two species are

genetically different. For other differences between

these two species see under T. ericius n. sp.

The protoconch and the first teleoconch whorls were

preserved in two juvenile specimens from Papua New

Guinea (RH). The protoconch is conical and glossy

brown, consisting of 3.5 whorls and a terminal lip of

the sinusigera type (Fig. 25G). Some specimens

collected in the China Sea (no other locality data)

before 1960 and deposited in MNHN are bigger and

reach a length of 16 mm (Fig. 25H).

The shell morphology of this little species confirms its

separation from the other Tenguella species and, more

particularly, from the similar looking 7. granulata.

Tenguella granulata is a species with a quite variable

shell morphology (Fig. 25Q-S), but it differs

consistently from 7. chinoiï n. Sp. in being less stocky,

with more acute nodes, in having a comparatively

larger shell, often reaching a length of more than 25

mm, with an average size of 15-20 mm, compared to

the 12-16 mm of 7: chinoï. The spire in 7. granulata is

higher and more acute: the aperture is larger, broader,

and less sinuous, with relatively smaller, less obvious

denticles and a narrower outer apertural lip.

Tenguella chinoi n. Sp. was illustrated by Dharma

(2005) as Azumamorula sp. This monotypic genus still

needs to be analysed carefully, but A. mutica

R. HOUART, D. ZUCCON & N. PUILLANDRE New genera and new species of Ergalataxinae

(Lamarck, 1816), the type species of Azumamorula Paratypes: Marquesas: PAKAIHI I TE MOANA, stn

(Fig. 27A-B), differs consistently from Tenguella in MQ7-M, Baie des Controleurs, 8°54'S, 140°03'W,

having a broadly convex, smoother shell with obsolete intertidal, 1 Ivy, MNHN-IM-2013-67608 (BOLD

axial ribs and very low spiral sculpture. It also has a MUBA787-18: GenBank MK216547).

broader aperture with comparatively weaker denticles. Vanuatu: SANTO 2006, stn VM06, Vanuatu, Maloka

Island, 15°35 S, 166°59'E, intertidal, rock bottom, 1 1v

Etymology. Named after Mitsuo Chino (Japan), (paratype MNHN-IM-2007-18187); stn FM36,

known for his numerous contributions to malacology Vaucluse Passage, 1572248 , 167°13E, 0-1 m,

and author of several new species, who sent me some intertidal, 3 Iv, MNHN-IM-2000-34218 (Fig. 25N-O)

of the shells included in the studies of Ergalataxinae. (syntopic with Tenguella granulata and T. chinoi n.

Sp.).

Papua New Guinea: PAPUA NIUGINI, sin PMA41,

Wonad Island, 05°08'S, 145°49'E, sandy beach and

" intertidal rocks, 0-1 m, 14 Iv (13 MNHN-IM-2000-

34219, 1 RH) (syntopic with Tenguella granulata and

° T. chinoi n. sp.); stn PR214, Tab Island, 05°15S,

145°47'E, 1-8 m, 1 dd, MNHN-IM-2000-34220

+ (syntopic with Tenguella granulata and T. chinoi n.

Sp.).

+ Guam: Mangalao, Pago Bay, near shore of eroded

limestone cliffs, in open bay with near-shore fringing

. reef, 2 Iv, NHMUK 20180544 (ex NHMUK

20080772) (syntopic with T. chinoi n. sp.) (see under

that species).

90° 105 120° 4135° 150° 165° 180° 165 150° Type locality. Marquesas, Pakaïhi I Te Moana, stn

Figure 24. Distribution of Tenguella chinoi n. sp. MQ7-M, Baie des Controleurs, 8°54S, 140°03"W,

intertidal.

Tenguella ericius n. sp.

Figs 12C-D; 251-P; 26 Other material examined. Marquesas: Taiohae,

Nuku Hiva Island, 1 1v, 1 dd (RH).

Morula granulata — Wells et al., 1990: 44, pl. 21, fig. China Sea: (no other data), 9 Iv, 2 dd (ex Staadt coll,

141 (not Tenguella granulata). 1969) (MNEN).

Type material. holotype MNHN-IM-2013-67609 Distribution. China Sea (no other data), Christmas

(BOLD MUBA788-18; GenBank MK216559), Island (Indian Ocean) (Wells et al. 1990), Guam,

Marquesas, PAKAIHI I TE MOANA, stn MQ7-M, Papua New Guinea, Vanuatu and Marquesas (Fig. 26).

Baie des Controleurs, 8°54'S, 140°03'W, intertidal.

Figure 25 (scale bar: 500 um)

A-H. Tenguella chinoi n. sp.

À. Guam, Mangalao, Pago Bay, near shore of eroded limestone cliffs, in open bay with near-shore fringing reef,

holotype NHMUK 20080772/1, GenBank: FEN677418, reconstructed aperture, 10 mm; B-C. Vanuatu, SANTO),

stn FM36, 15°22,45, 167°13'E, 0-1 m, paratype MNHN-IM-2000-34216, 13.6 mm; D-F. Papua New Guinea, |

Papua Niugini, stn PM41, 05°08,l'S, 145°49, 0-1 m, paratype MNHN-IM-2000-34217, 12.1 mm; G. Protoconch,

Papua New Guinea, Hansa Bay, Laing Island, RH; H. China Sea (no other data), MNEHN, 15.3 mm.

I-P. Tenguella ericius n. sp.

I-J. Marquesas, Pakaïhi I Te Moana, stn MQ7-M, Baie des Controleurs, 8°54'S, 140°03' W, intertidal, holotype

MNHN-IM-2013-67609, (BOLD MUBA788-18; GenBank MK216559), 13.1 mm; K-M. Vanuatu, SANTO stn

VMO6, Vanuatu, Maloka Id,15°35', 166°59'E, intertidal, paratype MNHN IM-2007-18187, 13.9 mm; N-O.

Vanuatu, SANTO, stn FM36, 15°22,4S, 167°13'E, 0-1 m, paratype MNHN-IM-2000-34218, 13.9 mm: P. Guam.

Mangalao, Pago Bay, near shore of eroded limestone cliffs, in open bay with near-shore fringing reef, paratype

NHMUK 20180544 (ex NHMUK 20080772), 14.8 mm.

Q-S. Tenguella granulata (Duclos, 1832)

Q-R. Papua New Guinea, Papua Niugini, stn PR214, 08/12/2012, 1-8 m (no other information), MNHN. 172

mm; S. Vanuatu, SANTO), stn RAP15,15°36,6'S, 167°0l'E, 2 m, MNEHN, 19.9 mm. ner

4 rs ds )19

N VA X

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R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new Species of Ergalataxinae

Figure 26. Distribution of Tenguella ericius n. sp.

Description. Shell small for the genus, up to 15.3 mm

in length at maturity. Length/width ratio 1.4-1.5.

Biconical, broad, heavy, weakly spinose and nodose,

tuberculate. Subsutural ramp strongly sloping,

concave.

Blackish-brown or black. Aperture white within with

narrow black coloured narrow band at edge and dark

bluish broad blotch at adapical weak curve of

columellar lip.

Spire moderately high. Teleoconch of up to 5 broad,

shouldered, nodose whorls. Suture adpressed.

Protoconch eroded 1n all specimens.

Axial sculpture of teleoconch whorls consisting of

narrow, moderately high, nodose ribs. Early spire

whorls with 12 or 13 ribs; last whorl with 9 ribs with

strong, relatively high, broad nodes at intersection

with primary spiral cords. Nodes higher and more

apparent on apertural rib. Spiral sculpture of low,

strong, rounded, broad, nodose, primary cords with 2

or 3 very narrow, rounded threads between each pair

of cords. Last teleoconch whorl with broad SP, similar

to P1-P3, P4 narrower, P5 narrowest primary cord.

Aperture large, ovate. Columellar lip broad, smooth or

with one or two very small denticles abapically and

low parietal tooth at adapical extremity. Anal notch

moderately deep, broad, with an open channel shortly

extending in SP spine. Outer lip weakly erect, smooth,

with 4 strong, broad denticles within consisting of DI-

* D4: D1 broadest and strongest, D2-D4 decreasing in

strength abapically. Siphonal canal very short, 3.5-

5.5% of total shell length, narrow, straight, broadly

open.

* Operculum and radula unknown.

. Remarks. Tenguella chinoi n. sp. differs consistently

from T. ericius in having a slightly broader, more

globose, stocky and solid shell, with broader and

* lower axial ribs; and in having blunt, broad nodes

compared to the more spiny shell of T. ericius n. sp. T.

chinoi also has a more sinuous columellar lip and

broader, more apparent apertural denticles within the

outer lip, resulting in a much more narrower aperture

as opposed to a relatively broad aperture in T. ericius

n. sp. In T. chinoi the spire is also lower and less

acute, the outer apertural lip is relatively broader, and

the siphonal canal, although short in T. chinoi n. sp. is

comparatively longer (9-10% of the total shell length)

than in T. ericius (3.5-5.5 % of the shell length).

Tenguella granulata differs from T. ericius n. sp. in

having an obviously larger shell and a comparatively

narrower aperture due to a strongly folded columellar

lip. The outer apertural lip of T. granulata is almost

similar than that of T. ericius n. sp., but it is obviously

less expanded abaperturally at the level of P4, between

D3 and D4 (Fig. 12D).

Etymology. Ericius: named after its

spinose shell.

hedgehog,

Discussion. Tenguella granulata, T. chinoi n. sp. and

T. ericius n. Sp. are syntopic in PAPUA NIUGINI, stn

PMA41 and PR214, and in SANTO 2006, stn FM36:; T.

chinoi n. Sp. and T. ericius are also syntopic in Guam

(see under T. chinoi n. Sp.). Tenguella granulata and

T. chinoi n. Sp. are syntopic in PAPUA NIUGINI stn

PEMO03, PM43, PR76 and PR204, and in SANTO stn

VM12 and RAP15.

Figure 27 (scale bar: 500 um)

A-B. Azumamorula mutica (Lamarck, 1816), Reunion Id, harbour, 2-3 m, RH, 18.2 mm.

C-L. Claremontiella nodulosa (C.B. Adams, 1845)

C-D. Jamaica, lectotype MCZ 177045, 14.9 mm (photo Jennifer Trimble, curatorial assistant, MCZ); E-G.

Syntypes of Ricinula ferruginosa Reeve, 1846, “W. Indies added to board”, H. Cuming collection, NHMUK

1968462. E-F. 18.9 mm; G. 17.3 mm (photo Harry Taylor, NHMUK Photographic Unit, © Natural History

Museum of London); H. Cape Verde, Sal Id, RH, 14.6 mm; I. Tobago, Rocky Point, Mt Irvine Bay, RH, 15.4

mm; J. Protoconch, Cuba, RH; K-L. West Africa, Angola, Cape Esterias, RH, 15.2 mm.

M-P. Claremontiella consanguinea (E.A. Smith, 1891)

M-N. St. Helena, lectotype NHMUK 1889.10.1.2368, 17.4 mm (photo Harry Taylor, NHMUK Photographic

Unit, © Natural History Museum of London); O-P. Sao Tome, Praia Emilia, RH, 13.8 mm.

R. HOUART, D. ZUCCON & N. PUILLANDRE

NOVAPEX 20(HS 12): 1-52, 10 mars 2019

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species Of Ergalataxinae

Un ue ep SSSR

Tenguella ericius n. sp. was also collected in the

China Sea prior to 1960, where it is probably

sympatric with 7. granulata and T. chinoiï n. sp., but

specific locality data is missing (MNHN). As for T.

chinoi n. Sp., the specimens of T. ericius n. sp.

collected in the China Sea are larger, reaching a length

of 17.5 mm.

Another shell, misidentified as M. granulata, was

illustrated by Wells et al. (1990). It is undoubtedly T.

ericius. The localities given (Christmas Island and

Cocos/Keelings Islands, Indian Ocean) are not

unexpected, but the stated size of the shell (2 cm) 1s

far larger than the other known specimens.

Claremontiella new genus

Figs 8D-F; 11G; 27C-P; 28A-E

Type species: Purpura nodulosa C.B. Adams, 1845:

2, eastern and western Atlantic (Fig. 27C-L).

Other included species. Claremontiella adiakritos n.

sp. and C. consanguinea (E.A. Smith, 1891) (new

combination).

Description. Shell moderately broad with high spire

and 7 or 8 rounded, nodose axial ribs, crossed by a

subsutural spiral cord and 5 primary spiral cords on

convex part of teleoconch whorl, with additional, low,

narrow, secondary and tertiary cords. Protoconch

conical with 3.5 whorls and sinusigeral terminal

notch.

Aperture narrow, ovate. Columellar lip smooth or with

low denticles abapically and low parietal tooth at

adapical extremity, weakly concave, rim adherent to

shell. Anal notch broad, moderately deep. Outer

apertural lip smooth with low denticles within.

Siphonal canal short, 12-14% of total shell length,

narrow, open.

Radula of Claremontiella of three-dimensional type

consisting of a rachidian bearing a narrow, long,

central cusp, a narrow, very short, lateral denticle on

each side, a broad, short lateral cusp, a few marginal

folds and a short marginal cusp. Lateral teeth sickle

shaped, narrow with a slightly broader base (Fig. 8D-

IE),

Distribution. Eastern and western South Atlantic,

Eastern Pacific, Gulf of California and Baja

California, Mexico.

Remarks. Houart (1997: 64) considered

Claremontiella consanguinea (E. À. Smith, 1891)

(Figs 8E-F; 27M-P) a valid species and assigned it to

Morula, together with C. nodulosa. The shell of C:

consanguinea Was separated from C. nodulosa by

having a narrower shell with lower nodes, more

broadly spaced spiral cords, a relatively longer

siphonal canal and a less denticulate or smooth

aperture. Since then, we have had the opportunity to

examine more specimens of Claremontiella species

from off West Africa, especially a population living in

Säo Tomé (Fig. 270-P). In these specimens, the shell

looks strongly similar to the original illustration of

Cantharus consanguineus.

Having sent typical specimens of C. nodulosa from

Virgin Islands, close to Jamaica, the type locality of C.

nodulosa and a species identified as C. consanguinea

from Säo Tomé for genetic analyses, we received an

opinion from Andrea Barco (in litt, 17 June 2013) who

said, "There is little genetic distance between the two

groups, but it is really low. I would expect higher

values between different species."

There is little doubt that some genetic differences

could exist between populations from Säo Tomé and

Saint Helena, so the names Claremontiella

consanguinea and C. nodulosa could be subjective

synonyms. Further analyses are needed before making

a final decision. In the meantime C. consanguinea is

here provisionally retained as a valid species.

Claremontiella adiakritos n. sp. is assigned to

Claremontiella because of its similar shell and radula

morphology. The shell of C. adiakritos is close to the

type species (Figs 27H and 28E) with which it has

often been confused, as Morula, Evokesia, or Pascula

ferruginosa (see under C. adiakritos n. sp.).

Etymology. The new genus is named for Martine

Claremont whose contribution to the molecular

phylogeny of Muricidae, in particular here for the

Ergalataxinae, while working at the Natural History

Museum, London, was extremely useful and

appreciated.

—————..—.—… —.—…—.…—.———….…——…—…——…—…—…—…—…—.—.—…—…—…——————.—.—.—.—_.._.._.._._._._

Figure 28

A-E. Claremontiella adiakritos n. sp.

A-D. West Mexico, Mazatlan. A-B. Holotype NHMUK 20180545, 23.3 mm; C-D. Paratype RH, 24.7 mm: E.

West Mexico, Baja California, near San Felipe, Alecia Playa, RH, 15.2 mm.

F-H. Murichorda fiscellum (Gmelin, 1791)

E-G. South of west Java, Sancang, low tide, under rocks, RH, 23.2 mm; H. Aden, Gulf of Aden. RH, 26.2 mm.

L. Murichorda jacobsini (Emerson & D'Attilio, 1985), Solomon Islands, Guadalcanal, Marau Sound, RH, 37.3

mm.

J-L. Murichorda rumphiusi (Houart, 1996).

J. Ambon, Hito, east side of Laha, paratype RH, 19.7 mm; K-L. Singapore, east coast of Park Beach. intertidally,

muddy rocks, RH, 19.6 mm.

)

NOVAPEX 20(HS I!

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

A — ]——]]—

Claremontiella adiakritos n. sp.

Figs 8D; 11G:; 28A-E; 30

Morula ferruginosa — Keen, 1958: 376, sp. 411;

Abbott, 1974: 178, fig. 1875 (not Ricinula ferruginosa

Reeve, 1846).

Morula (Morunella) ferruginosa — Keen, 1971: 554,

fig. 1092; Victor Alamo & Violeta Valdivieso, 1997:

54 (not Ricinula ferruginosa Reeve, 1846).

Evokesia ferruginosa — Radwin & D'Attulio, 1972:

338, fig. 1A; Radwin & D'Attilio, 1976: 143, pl. 3, fig.

5; Kaicher, 1979, card 2038 (not Ricinula ferruginosa

Reeve, 1846).

Pascula ferruginosa — Vokes, 1984: 2014, 215 (not

Ricinula ferruginosa Reeve, 1846).

Morula (Morula) ferruginosa — Skoglund, 2002: 116

(not Ricinula ferruginosa Reeve, 1846).

Type material. Holotype NHMUK 20180545, West

Mexico, Mazatlan, lv.

Paratypes: West Mexico, Mazatlan, 1v, 2 RH; Mexico,

Baja California Sur, Playa Santispac, about 30 km

south of Mulegce, 2 IRSNB I.G. 33893; MT. 3744, 2

MNAN-IM-2000-34221

Type locality. West Mexico, Mazatlan.

Other Material examined. Baja California, Mexico,

Alecia Playa, near San Felipe, 24 Iv & dd, RH;

Guaymas, 3 1v & dd, RH (radula illustrated); Mexico,

Adair Bay, 2 1v, RH; Gulf of California (no other

data), 3 Iv, RH; Mexico, Mulege, 2 dd, RH; Baja

California, Mexico, Puertocitos, 1 dd, RH; Baja

California, Mexico, La Paz, 2 1v, RH.

Distribution. Magdalena Bay, Baja California,

through the Gulf of California and south along the

Sonoran coast of Mexico at least as far as Guaymas,

intertidally, under rocks (Keen, 1971). Its distribution

south to Bocapän, El Rubio and Tumbes, Peru (Alamo

& Valdivieso, 1987, 1997 and Skoglund, 2002) is

doubtful (see remarks). Finet (1994: 50) mentions

records from the Galapagos Islands but without any

illustration, which is also here considered doubtful

(Fig. 30).

Description. Shell large for the genus, up to 24.7 mm

in length at maturity (paratype RH). Length/width

ratio 2.1-2.3. Lanceolate, narrow, heavy, nodose.

Subsutural ramp broad, strongly sloping, concave.

Dark brown or blackish-brown, white between nodes

of first primary cord (P1), also on other cords but less

obvious. Aperture bluish-white within with brown

band within outer lip up to white apertural denticles.

Columellar lip bluish-white with a darker area

abapically.

Spire very high with teleoconch up to 6 weakly

convex, narrow, angulate, shouldered, nodose whorls.

Suture adpressed. Protoconch in examined material

unknown, eroded in all specimens.

Axial sculpture of teleoconch whorls consisting of

low, weak, narrow, rounded ribs with moderately

high, sharp nodes at intersection of primary spiral

cords. First and second whorls eroded in all

specimens, third whorl with 10 ribs, fourth with 9,

fifth with 9 or 10, last whorl with 7 ribs. Spiral

sculpture of low, rounded, broad, nodose primary

cords and numerous threads. Last teleoconch whorl

with SP, PI-PS and additional threads between each

pair of cords. P1 broadest, P2-P3 weakly narrower,

similar in strength, P4 narrower and lower, PS

smallest on adapical part of siphonal canal.

Aperture small, ovate. Columellar lip narrow, smooth,

rim adherent, with very weak low parietal tooth at

adapical extremity. Anal notch shallow, broad. Outer

lip smooth with low, small denticles within, consisting

of DI-D4, strongly decreasing in strength abapically;

very low ID occasionally observed. Denticles

sometimes obsolete. Siphonal canal very short, 10-

12% of total shell length, narrow, weakly dorsally

recurved, broadly open.

Operculum dark brown, ovate with subapical nucleus

in lower right.

Radula of three-dimensional type consisting of a

rachidian bearing a narrow, long, central cusp, a

narroWwW, very short, lateral denticle on each side, a

broad, shorter lateral cusp and few marginal folds.

Lateral teeth sickle shaped, narrow with a slightly

broader base.

Figure 29

A-B. Muricodrupa fenestrata (Blainville, 1832), New Caledonia, Ouvea, RH, 30.3 mm.

C-D. Muricodrupa anaxares (Kiener, 1835), South Africa, Durban, Reunion rocks, low tide, in rock crevices,

RH, 14.6 mm.

E-F. Lauta parva (Reeve, 1845), Philippines, Cebu, RH, 9.4 mm.

G-H. Engina alveolata (Kiener, 1836), Fiji, Wayasawa group, Wasayama Id, under coral slab, low tide. RH.

12.9 mm.

I-L. Cantharus (Tritonidea) albozonatus Smith, 1890, St. Helena, W.H. Turton collection.

I-J. Syntype NHMUK 1889.10.1.2356-2361, 15.8 mm; K-L. Syntype NHMUK 1889.10.1.152-161. 10.9 mm.

R. HOUART, D. ZUCCON & N. Put LANDRI NOVAPEX 20(HS 12): 1-52, 10 mars 2019

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

Remarks. Vokes (1984) figured Reeve's 1llustrated

syntype of Ricinula ferruginosa (NHMUK 1968462)

and noted that the syntypes were specimens of the

Atlantic Trachypollia nodulosa (C.B. Adams), leaving

the Pacific species without a name. Vokes (1984) also

illustrated the lectotype of T. nodulosa.

Claremontiella adiakritos n. sp. is assigned to

Claremontiella because the shell and radula

morphology are similar to those of C. nodulosa, type

species of the genus, and to C. consanguinea.

The species has a very high spire with a low

subsutural spiral cord; low, narrow cords on

subsutural ramp; five low, broad, spiral cords on

convex part of teleoconch whorl; numerous spiral

threads; and a very short siphonal canal. The outer

apertural lip bears low denticles within, and 1s

occasionally smooth.

The ontogeny of the spiral cords is unknown since all

examined specimens were badly corroded from the

first to the second or third teleoconch whorls. For this

reason, the protoconch could also not be examined.

However, Radwin & D'Attilio (1972: 338) described

and illustrated the protoconch of C. adiakritos n. sp.

(as Evokesia ferruginosa) with 3.5 papillose whorls,

similar to the protoconch morphology of the type

species of Claremontiella (Fig. 27J).

The radula of this species, 1llustrated in Radwin &

D'Attilio (1972: 340, fig. 16), 1s strangely different

from that illustrated here (Fig. 8D), extracted from a

specimen collected near San Felipe. The drawing

shows a broader and shorter central cusp, and broader

lateral cusps as well as broad lateral teeth, while the

SEM here 1llustrated shows a typical ergalataxine

radula, very close to that of C. consanguinea, With a

narrow, long central cusp, a very short lateral denticle

on each side, short, moderately broad lateral cusps and

narrow, slender, lateral teeth.

The assignation of this species to Claremontiella n.

gen. 1s tentative and requires genetic analyses for

confirmation.

Alamo & Valdivieso (1987, 1997) reported this

species (without any figure) from Bocapän and El

Rubio, Tumbes, Peru, but Claremontiella adiakritos n.

Sp. has been frequently confused with Trachypollia

lugubris and no voucher material was referenced (V.

Mogollon, in litt.). Therefore, the presence of C.

adiakritos in Peru and the Galapagos Islands remains

doubtful.

Claremontiella nodulosa differs from C. adiakritos n.

sp. in having a shell with shightly larger primary spiral

cords with rounded nodes instead of more sharp

knobs, and in having a generally less high spire, a

broader, obvious and nodose subsutural cord, and a

darker coloured aperture with thicker outer apertural

lip and more obvious denticles within.

Etymology. Adiakritos (G) meaning mixed,

undistinguishable, named for its ressemblance to some

forms of Claremontiella nodulosa (C.B. Adams,

1845), with which it has often been confused in

literature.

Figure 30. Distribution of Claremontiella adiakritos

n. SP.

Murichorda new genus

Figs 8G-H; 9A; 28F-L

Type species: Purpura fiscellum Gmelin, 1791: 3552,

Indo-West Pacific.

Other included species. Murichorda rumphiusi

(Houart, 1996) (new combination) and Murichorda

jacobsini (Emerson & D'Attilio, 1985) (new

combination).

Description. Shell large, up to 44.5 mm in length,

broadly ovate, weakly angulate, shouldered, with 6-8

broad, high, rounded axial ribs, crossed by 5 high,

longitudinally grooved primary spiral cords,

occasionally deeply excavated between each pair of

cords. Intersection of axial ribs and primary spiral

cords forming low, broad knobs. Aperture broadly

ovate with low denticles within outer lip. Siphonal

canal short to moderate in length, 13-29% of total

shell length, broadly open.

Radula ergalataxine (Figs 8G-H; 9A), of three-

dimensional type, consisting of a narrow, long central

Cusp, a very Short, narrow, lateral denticle on each

side, a broad, moderately long lateral cusp and

occasionally à few low marginal folds. Lateral teeth

sickle shaped, broad.

Distribution. Throughout the Indo-Pacific.

Remarks. Murichorda fiscellum was previously

assigned to Cronia H. & A. Adams, 1853 by

Cernohorsky (1969: 311; 1982: 113), but Cronia. type

species C. amygdala (Kiener, 1835), was included in a

different clade by Claremont et al. (2013), together

R. HOUART, D. ZUCCON & N. PUILLANDRE

NOVAPEX 20(HS 12): 1-52, 10 mars 2019

————_—.

with Ergalatax Iredale, 1931 and Maculotriton Dall,

. which could be potential Synonyms (see Table

Because its shell morphology is close to M. fenestrata

(Blainville, 1832), the type species of Muricodrupa,

Murichorda fiscellum has been assigned to

Muricodrupa by numerous authors, Starting with Kay

(1979). However, in Claremont et al. (2013) "Morula"

fiscella and "Morula" rumphiusi formed a well-

supported clade in all analyses but were excluded

from Muricodrupa. Both are therefore here assigned

to Murichorda new genus, along with Murichorda

Jacobsini.

Etymology. Prefix Muri, from Muricidae + suffix

chorda (L): rope, twine. Named for the strong, broad

Spiral cords, a characteristic shared by the shells

assigned to this new genus.

Lauta new genus

Fig. 29E-F

Type species: Ricinula parva Reeve, 1846: pl. 6, fig.

43.

Description. Shell small, up to 10 mm in length,

weakly ovate and shouldered, nodose, with 7 or 8

axial ribs crossed by narrow, rounded, primary cords

consisting Of SP near suture, P1-PS on convex part of

teleoconch whorl and ADP on siphonal canal, with

rounded, high nodes at intersection of axial ribs and

spiral cords.

Spire high with conical protoconch of 3.5 whorls.

Aperture strongly ovate, narrow, with strong denticles

within outer lip. Columellar lip narrow, almost

straight, smooth, rim almost entirely adherent to shell.

Anal notch broad, moderately deep. Siphonal canal

short, 10-13% of total shell length. Radula unknown.

Distribution. Philippine Islands.

Remarks. Lauta parva is sister to all other members

of subclade Z in Claremont et al (2013: fig. 1-3B),

suggesting that a new genus is required for this

species. The genus Morula Schumacher, 1817 in

which Lauta parva was previously included consisted

of 19 species. One of them, M. anaxares, is now

assigned to Muricodrupa Tredale, 1918; two species,

M. nodulosa and C. consanguinea, are now assigned

to Claremontiella n. gen.; one species, M. rumphiust,

is now assigned to Murichorda n. gen; and one

species, M. parva, is here assigned to Lauta n. gen. Of

the 14 remaining species of Morula Ss.s., six were

analysed by Claremont et al (2013): M. aspera

(Lamarck, 1816), M. chrysostoma (Deshayes, 1844),

M. echinata (Reeve. 1846) (= Sistrum ventricosulum

G. & H. Nevill, 1875, new synonymy) (as M.

benedicta in Claremont et al., 2013), M. nodicostata

(Pease, 1868), M. uva (Rôüding, 1798) and M. zebrina

Houart, 2004. The eight remaining species are here

provisionally retained in Morula s.s. (Table 2),

pending future genetic analyses.

Etymology. The generic name Lauta (L: neat, elegant)

is derived from Ricinula lauta Reeve, 1846, junior

Synonym Of Engina alveolata (Kiener, 1836) (Fig.

29G-H), a species of Pisaniidae (Buccinoidea) whose

shell morphology and (particularly) colouration is

Strangely reminiscent of the type species, L. parva.

ACKNOWLEDGEMENTS

Most of the material in this paper originates from

numerous shore-based expeditions and deep water

cruises, conducted between 1977 and 2017 by the

MNHN and Pro-Natura International (PNI) as part of

the Our Planet Reviewed programme (SANTO 2006,

ATIMO VATAE, MIRIKY, MAINBAZA, PAPUA

NIUGINI, PI Philippe Bouchet; KANACONO, PI

Nicolas Puillandre; KANADEEP, PI Sarah Samadi),

and/or by MNHN and Institut de Recherche pour le

Développement (IRD) as part of the Tropical Deep-

Sea Benthos programme in New Caledonia (BIOCAL.

CHALCAL 2, SMIB 3, 4, 5, 8, BERYX 11, BATHUS

2, NORFOLK 1, NORFOLK 2, EBISCO,

TERRASSES, EXBODI), Wallis & Futuna

(MUSORSTOM 7), Vanuatu (MUSORSTOM 8), the

Marquesas (MUSORSTOM 9), Tonga (BORDAU 2),

the Philippines (PANGLAO 2005, AURORA 2007),

the Mozambique Channel (BIDMAGLO) and Walters

Shoal (PIs Claude Lévi, Bertrand Richer de Forges,

Patrick Lehodey, Cécile Debitus, Philippe Bouchet,

Sarah Samadi, Laure Corbari). Additional sources are

BENTHEDI (PI Bernard Thomassin), the Marion-

Dufresne MD32 cruise off Réunion (PI Alain Guille),

the LIFOU 2000 expedition to the Loyalty Islands, the

PANGLAO 2004 expedition to the Philippines, and

the PAKAIHI I TE MOANA expedition to the

Marquesas Is. Scientific partners included the

University of Papua New Guinea (UPNG), University

San Carlos (Cebu City) and the Bureau of Fisheries

and Aquatic Resources (BFAR, Manila), Institut

d'Halieutique et des Sciences Marines, University of

Toliara (IH.SM) and the Madagascar bureau of

Wildlife Conservation Society (WCS), Instituto

Español de Oceanografia, the Marine and Polar

Programme of the International Union for

Conservation of Nature (IUCN International), and

Agence des Aires Marines Protégées (AAMP), with

access to Ship time through the Flotte

Océanographique Française, and funding from the

Total Foundation, Prince Albert IT of Monaco

Foundation, Stavros Niarchos Foundation, Vinci

Entrepose Contracting, Fondation EDF, the French

Global Environment Fund (FFEM), Fonds Pacifique

and the Government of New Caledonia. For station

lists and context of the expeditions, see

https://expeditions.mnhn.fr/ and Bouchet et al. (2008).

We are very grateful to Philippe Bouchet (Muséum

national d'Histoire naturelle, Paris, France) for the

opportunities he has given us to study the material

collected during the Tropical Deep Sea Benthos

campaigns in the Indo-West Pacific, for information

about the various expeditions and for constructive

R. HOUART, D. ZUCCON & N. PUILLANDRE

New genera and new species of Ergalataxinae

An pp aa—

comments. We are also very thankful to Barbara

Buge, Virginie Héros and Philippe Maestrati (MNHN)

for their help in searching for and sending specimens

in loan, and for relevant information. Manuel Caballer

(MNHN) provided the images of the MNEN

holotypes, E-Recolnat Project: ANR-11-INBS-0004.

Thanks also to everyone who contributed to this work

by sending photos and loans, and for other

information. Without their help, it would have been

difficult to complete this article. Our sincerest thanks

to Adam Baldinger and Marcia Kazmierczak

(Museum of Comparative Zoology, Harvard

University, Cambridge, U.S.A) for the image of the

lectotype of Purpura nodulosa C.B. Adams, 1845; to

Andrea Barco, GEOMAR, Helmholtz Centre for

Ocean Research, Kiel, Germany for information about

genetic analyses; to Philippe Bouchet (MNHN),

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. to Martine Claremont (Boston,

Massachusetts, U.S.A) for reading, commenting on

and correcting the first draft of this ms; to Jacques

Colomb (Marseille, France) for providing me with

specimens of C. nodulosa from Virgin Islands and C.

consanguinea from Säo Tomé for genetic analysis; to

Dai Herbert, Linda Davis and the late Richard (Dick)

Kilburn (KwaZulu-Natal Museum, South Africa) for

the Iloan of material; to Roland Janssen

(Forschungsinstitut Senckenberg, Frankfurt, Germany)

for information about the type material of Purpura

castanea Küster, 1858; to Bruce Marshall (Museum of

New Zealand Te Papa Tongarewa, Wellington, New

Zealand) for sending us soft parts of Pascula palmeri

for study and SEM of the radula; to Shawn Miller

(Okinawa, Japan) for providing me with some

specimens Of Orania pachyrhaphe a few years ago; to

Valentin Mogollén Avila (Universidad Nacional

Federico Villarreal, Lima, Peru) for information about

the possible presence of Claremontiella adiakritos n.

Sp. in Peru; to Tina Petway and Gary Kidder (The

Houston Museum of Natural Science, Houston, Texas,

U.S.A.) for information and photos of the holotype

and paratypes of Cytharomorula manusuduirauti n.

Sp.; to Jane Pickering (then assistant curator, zoological

collections at the Oxford University Museum, United

Kingdom), for the loan of the syntype of Cominella

unifasciata nigronodulosa Turton, 1932 in 1995; to

Andreia Salvador and Harry Taylor (Natural History

Museum, United Kingdom), for providing information

and photographs of the type material deposited in

NHMUK; and to Emmanuel Tardy (Muséum

d'histoire naturelle, Genève, Switzerland) for

information about a type from Kiener.

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R. HOUART, D. ZUCCON & N. PUILLANDRE NOVAPEX 20(HS 12): 1-52, 10 mars 2019

a" —_—_

Table 2, List of the valid extant taxa in ERGALATAXINAE

Bold: new taxa described in this paper.

Asterisk: taxa sequenced in Claremont et al. (2013) or in this paper.

Question mark: doubtful classification after sequencing, but a témporary placement in the genera in which they

are Currently assigned is suggested.

NC: New combination.

Genus Type species Species

Azumamorula Emerson, 1968 Ricinula mutica Lamarck, A. mutica (Lamarck, 1816)

1816

*Bedevina Habe, 1946 Trophon birileffi Lischke, *B. birileffi (Lischke, 1871)

1871

“Claremontiella n. gen. Purpura nodulosa C.B. C. adiakritos n. sp.

Adams, 1845 C. consanguinea (E.A. Smith, 1891) NC

*C. nodulosa (C.B. Adams, 1845) NC

*Cronia H. & A. Adams, 1835 Purpura amygdala Kiener, *C. amygdala (Kiener, 1835)

1835 *C. aurantiaca (Hombron & Jacquinot,

1853) (see note)

C. avellana (Reeve, 1846)

*Cytharomorula Kuroda, 1953 Cytharomorula vexillum *Cytharomorula absidata n. sp. [(as

Kuroda, 1953 Cytharomorula cf. grayi in Claremont et al.

(2013)]

C. ambonensis Houart, 1996

C. arta n. sp.

C. benedicta (Melvill & Standen, 1895)

C. danigoi Houart, 1995

C. dollfusi (Lamy, 1938)

*C. elegantula n. sp.

C. fatuhivaensis n. sp.

*C. grayi (Dall, 1889)

C. lefevreiana (Tapparone Canefri, 1880)

C. manusuduirauti n. sp.

*C. ornamentata (Houart, 1995) NC

*C. paucimaculata (Sowerby, 1903)

C. pinguis Houart, 1995

*C. springsteeni Houart, 1995

*C. vexillum Kuroda, 1953

Daphnellopsis Schepman, 1913 Daphnellopsis lamellosa D. fimbriata (Hinds, 1843)

Schepman, 1913 D. hypselos Houart, 1995

D. lamellosa Schepman, 1913

D. lochi Houart, 2013

D. lozoueti Houart, 2013

D. pinedai Houart, 2013

*Drupella Thiele, 1925 Drupa cornus Rôding, *D. cornus (Rôüding, 1798)

1798 *D). eburnea (Küster, 1862)

*D. fragum (Blainville, 1832)

*D. margariticola (Broderip, 1833)

D. minuta Fujioka, 1984

*D. rugosa (Born, 1778)

*Ergalatax Iredale, 1931 Ergalatax recurrens *E, contracta (Reeve, 1846)

Iredale, 1931 = Murex E. crassulnata (Hedley, 1915)

pauper Watson, 1883 E. dattilioi Houart, 1998

E. heptagonalis (Reeve, 1846)

*E. junionae Houart, 2008

E. martensi (Schepman, 1892)

E. pauper (Watson, 1883)

E. tokugawai Kuroda & Habe, 1971

E. zebra Houart, 1995

R. HOUART, D. ZUCCON & N. PUILLANDRE New genera and new species of Ergalataxinae

D de qe EE —

Table 2 (continued)

RE ne ee in mener: (de

Genus Type species Species

*Lataxiena Jousseaume, 1883 Lataxiena lataxiena L. blosvillei (Deshayes, 1832)

Jousseaume, 1883 = L. bombayana (Melvill, 1893)

Trophon fimbriatus Hinds, L. cumella (Jousseaume, 1898)

1844 L. desserti Houart, 1995

*L. fimbriata (Hinds,1844)

L. habropenos Houart, 1995

L. lutescena Zhang & Zhang, 2015

L. solenosteiroides Houart, Fraussen &

Barbier, 2013

*Lauta n. gen. Ricinula parva Reeve, 1846 *L. parva (Reeve, 1846) NC

Lindapterys Petuch, 1987 Lindapterys vokesae L. domlamyi Garrigues & Merle, 2014

Petuch, 1987 (early L. murex (Hedley, 1922)

Miocene, Florida) L. sanderi Petuch, 1987

L. soderiae Callea, Volpi, Martignoni &

Borri, 2001

*Maculotriton Dall, 1904 Triton bracteatus Hinds, *M. serriale (Deshayes, 1834)

1844 = Buccinum serriale

Deshayes, 1834

*Morula s.s. Schumacher, 1817 Drupa uva Rôding, 1798 M. (M.) albanigra Houart, 2002

M. (M.) angulata (Sowerby, 1893)

*M. (M.) aspera (Lamarck, 1816) (see note)

M. (M.) cernohorskyi Houart & Trôndlé,

1997

*M. (M.) chrysostoma (Deshayes, 1844)

*M. (M.) echinata (Reeve, 1846)

*M. (M.) nodicostata (Pease, 1868)

M. (M.) oparensis (Melvill, 1912)

M. (M.) peasei Houart, 2002

M. (M.) praecipua Rehder, 1980

M. (M.) rodgersi Houart, 2000

*M. (M.) uva (Rôding, 1798)

M. (M.) variabilis (Pease, 1868)

*M. (M.) zebrina Houart, 2004

*Morula (Habromorula) Houart, Purpura biconica Blainville, M. (H.) aglaos (Houart, 1995)

1995 1832 M. (H.) ambrosia (Houart, 1995)

M. (Æ.) bicatenata (Reeve, 1846)

*M. (1) biconica (Blainville, 1832)

*M. (H.) coronata (H. Adams, 1869)

M. (H.) dichrous (Tapparone Canefri, 1880)

M. (Æ.) euryspira (Houart, 1995)

M. (H.) fuscoimbricata (Sowerby, 1915)

*M. (H.) japonica (Sowerby, 1903)

M. (H.) lepida (Houart, 1995)

M. (H.) porphyrostoma (Reeve, 1846)

*M. (H.) spinosa (H.& A. Adams, 1853)

*M. (H.) striata (Pease, 1868)

M. (H.) whiteheadae Houart, 2004

*Murichorda n. gen. Murex fiscellum Gmelin, *M. fiscella (Gmelin, 1791) NC

1791 M. jacobsoni Emerson & D'Attilio, 1981 NC

*M. rumphiusi Houart, 1996 NC

*Muricodrupa Iredale, 1918 Purpura fenestrata *M. anaxares (Kiener, 1835) NC

Blainvville, 1832) *M. fenestrata (Blainville, 1832)

*Oppomorus Iredale, 1937 Purpura nodulifera Menke, *O. funiculatus (Reeve. 1846)

1829 *O. noduliferus (Menke, 1829)

“O. purpureocinctus (Preston, 1909)

R. HOUART, D. ZUCCON & N. PUILLANDRE NOVAPEX 20(HS 12): 1-52, 10 mars 2019

Table 2 (continued)

ee ——_—_…—————— "cp

*Orania Pallary, 1900 Murex spadae Libassi, O. adiastolos Houart, 1995

Note: Type species not analysed in 1859 = Murex fusulus O. archaea archaea Houart, 1995

Claremont et al. (2013) Brocchi, 1814 O. archaea hitomiae Houart & Moe, 2011

O. atea Houart & Trôndlé, 2008

O. badia (Reeve, 1845)

* 70. bimucronata (Reeve, 1846)

*O. carnicolor (Bozzetti, 2009) NC

*O. castanea (Küster, 1886)

O. corallina (Melvill & Standen, 1903)

O. dharmai Houart, 1995

O. ficula (Reeve 1848)

*O. fischeriana (Tapparone-Canefri, 1882)

O. fusulus (Brocchi, 1814)

* 70. gaskelli (Melvill, 1891)

O. livida (Reeve, 1846)

*O. mixta Houart, 1995

O. nodosa (Hombron & Jacquinot, 1841)

O. nodulosa (Pease, 1869)

O. pachyrhaphe (Smith, 1879)

*O. pacifica (Nakayama, 1988)

O. pholidata Watson, 1883

O. pleurotomoides (Reeve, 1845)

O. pseudopacifica n. sp.

O. rosadoi Houart, 1998

*O. rosea Houart, 1996

* 20. serotina (A. Adams, 1853)

O. simonetae Houart, 1995

O. subnodulosa (Melvill, 1893)

O. taeniata Houart, 1995

O. taiwana (Lai & Jung, 2012) NC

O. walkeri (Sowerby, 1908)

O. xuthedra (Melvill, 1893)

*Pascula Dall, 1908 Trophon citricus Dall, 1908 P. citrica (Dall, 1908)

Note: Type species not analysed in *P. darrosensis (E.A. Smith, 1884)

Claremont et al. (2013) *P. muricata (Reeve, 1846)

*P. ochrostoma (Blainville, 1832)

P. ozenneana (Crosse, 1861)

P. palmeri (Powell, 1967) NC

P. philpoppei Houart, 2018

P. rufonotata (Carpenter, 1864)

*P. submissa (E. A. Smith, 1903)

*Phrygiomurex Dall, 1904 Triton sculptilis Reeve, *P. sculptilis (Reeve, 1844)

| 1844

*Spinidrupa Habe & Kosuge, 1966 Murex euracanthus A. S. aethes Houart, 2017

Adams, 1853 * S. euracantha (A. Adams, 1853)

* 95. infans (E.A. Smith, 1884)

*Tenguella Arakawa, 1965 Purpura granulata Duclos, *Tenguella ceylonica (Dall, 1923) |

1832 *T. chinoi n. sp. [as Tenguella n. sp. in

Claremont et al (2013)]

*T. ericius n. sp.

*T. granulata (Duclos, 1832)

T. hoffmani Houart, 2017

*T. marginalba (Blainville, 1832)

*T. musiva (Kiener, 1835)

R. HOUART, D. ZUCCON & N. PUILLANDRE New genera and new species of Ergalataxinae

D ———__ ————— — — —

Table 2 (continued)

A _ _ —_ — — .,.

Genus Type species Species 14

*Trachypollia Woodring, 1928 Trachypollia sclera T. didyma (Schwengel, 1943)

Woodring, 1928 *T. lugubris (C.B. Adams, 1852)

T. sclera (Woodring, 1928)

T. turricula (Maltzan, 1884)

*Usilla H. Adams, 1861 Vexilla nigro-fusca Pease, *T. avenacea (Lesson, 1842)

1860 = Vexilla fusconigra

Pease, 1860 = Purpura

avenacea Lesson, 1842

Uttleya Marwick, 1934 Uttleya arcana Marwick, U. ahiparana (Powell, 1927)

1934 (lower Pleistocene, U. marwicki Powell, 1952

New Zealand) U. williamsi Powell, 1952

Note: Small genetic distances are suggestive of possible synonymy of species; for example, there was < 1% divergence

among COI sequences for the pair Morula uva and M. aspera, and similarly for the pair Cronia amygdala and C. aurantiaca

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