European Journal of Taxonomy 134: 1-23

http://dx.doi.org/10.5852/ejt.2015.134

www. europeanj ournaloftaxonomy. eu

2015 • Anseeuw P. et al.

ISSN 2118-9773

This work is licensed under a Creative Commons Attribution 3,0 License.

Research article

um:lsid:zoobank.org:pub:B623BClC-96CD-410B-97E6-4D03864D29EC

Perotrochus caledonicus (Gastropoda: Pleurotomariidae) revisited:

descriptions of new species from the South-West Pacific

Patrick ANSEEUW 1 , Nicolas PUILLANDRE 2 *, Jose UTGE 3 , Philippe BOUCHET 4

1 Mispelstraat 18, 9820 Merelbeke, Belgium.

2 Institut de Systematique, Evolution, Biodiversite ISYEB -UMR7205 - CNRS, MNHN, UPMC,

EPHE, Museum National d’Histoire Naturelle, Sorbonne Universites,

43 Rue Cuvier, F-75231 Paris, France.

3 UMS 2700, Museum National d’Histoire Naturelle, Departement Systematique et Evolution,

43 Rue Cuvier, F-75231 Paris, France.

4 Institut de Systematique, Evolution, Biodiversite ISYEB - UMR7205 - CNRS, MNHN, UPMC,

EPHE, Museum National d’Histoire Naturelle, Sorbonne Universites,

55 Rue Buffon, F-75231 Paris, France.

* Corresponding author: puillandre@rrmhn fr

1 urn:lsid:zoobank.org: author: 89470910-9E6E-4B84-91E4-9C10019DD7DD

2 urn:lsid:zoobank.org:author:00565F2A-C170-48Al-AAD9-16559C536E4F

3 urn:lsid: zoobanlc. org: author: 3 888EAFD-B518-4979-A2DF-4A7A1D4B7F 89

4 urn:lsid:zoobank.org:author:FC9098A4-8374-4A9A-AD34-475E3AAF963A

Abstract. Morphological (shell) and molecular examination of a large suite of specimens of

pleurotomariids from around New Caledonia and the Coral Sea reveals the existence of four species in

the complex of Perotrochus caledonicus'. Perotrochus deforgesi Metivier, 1990 and P. pseudogranulosus

sp. nov. live allopatrically on the plateaus and guyots of the Coral Sea; Perotrochus caledonicus Bouchet

& Metivier, 1982 and Perotrochus wareni sp. nov. live sympatrically - but essentially not syntopically -

on the slopes of New Caledonia, Norfolk Ridge and the Loyalty Ridge. All species live in the 300-500

m interval, and together form a significant component of the mollusc fauna living on hard bottoms in the

SW Pacific, with individual dredge hauls containing up to 25 specimens of Perotrochus.

Keywords. Perotrochus wareni sp. nov., Perotrochus pseudogranulosus sp. nov., Coral Sea, New

Caledonia, COI Barcode gene.

Anseeuw R, Puillandre N., Utge J. & Bouchet P. 2015. Perotrochus caledonicus (Gastropoda: Pleurotomariidae)

revisited: descriptions of new species from the South-West Pacific. European Journal of Taxonomy 134: 1-23.

http://dx.doi.org/10.5852/ejt.2015.134

Introduction

Perotrochus caledonicus Bouchet & Metivier, 1982 and P. tangaroanus Bouchet & Metivier, 1982 were

the first pleurotomariids described from the South Pacific, from New Caledonia and from the Lau Ridge

between New Zealand and Fiji (Bouchet & Metivier 1982), respectively. Since then, additional species

have been documented from New Caledonia (. Perotrochus deforgesi Metivier, 1990 and Bayerotrochus

1

European Journal of Taxonomy 134: 1-23 (2015)

boucheti (Anseeuw & Poppe, 2001)) and Tonga (Bayerotrochuspoppei Anseeuw, 2003). Moreover, many

populations at first sight referable to Perotrochus caledonicus were sampled around New Caledonia and

in the Coral Sea.

The type material of P. caledonicus from off southern New Caledonia consists of specimens with a

macroscopically non-pustulose shell, with rather smooth to weakly beaded spiral cords. Soon after

its description, it appeared that a more granulose or pustulose morph also occurred around New

Caledonia. However, as such material appeared very similar in terms of size, color and general profile,

it was considered to represent just a form of P. caledonicus. The two forms were reported in print

(Anseeuw 1990; Anseeuw & Goto 1996) and referred to as the “smooth form” and “pustulose form” of

P. caledonicus , respectively. Since then, dealers and shell collectors have maintained this distinction,

using the expression “caledonicus pustulose form” or “caledonicus granulose form”, with some even

suggesting that the latter could warrant full recognition as a species (e.g., www.conchology.be T

New material from New Caledonia and the Coral Sea has now allowed revisiting this issue, using

an integrative taxonomic approach combining DNA sequencing and shell characters of both juveniles

and adults. DNA sequences provide independent characters to test whether the different forms of P.

caledonicus correspond to divergent lineages or not. The congruence of the different sets of characters

analyzed revealed that three species were hidden under the name P. caledonicus , two of which are now

described as new. In addition, Perotrochus deforgesi was found to be molecularly closely related to this

complex - which was not suspected from shell characters alone - and in this paper we will designate

these small Perotrochus from the SW Pacific together as the “Perotrochus caledonicus complex”.

Fig. 1. Distribution maps of the four species. A. Perotrochus wareni sp. nov. B. Perotrochus deforgesi

Metivier, 1990. C. P erotrochus caledonicus Bouchet& Metivier, 1982. D. Perotrochus pseudogranulosus

sp. nov. Stations with sequenced specimens in black.

2

Material and methods

Material

ANSEEUW P. et al ., New species of Pleurotomariidae

A total of284 lots and 879 specimens, including 59 sequenced specimens, attributable to the Perotrochus

caledonicus complex were collected in the New Caledonia region, including the Coral Sea (Table 1,

Fig. 1) during research cruises of the Tropical Deep-Sea Benthos program (Bouchet, Heros, Lozouet &

Maestrati 2008). After 2002, live-taken specimens were specifically processed on board for molecular

analyses. Apiece of the foot was cut and placed in 95° ethanol; the remaining body and the shells were

also preserved for further examination. All specimens are preserved at the Museum National d’Histoire

Naturelle (MNHN); each sequenced specimen is linked to a unique MNHN collection number. These

specimens are registered in BOLD, the Barcode of Life database, and the corresponding sequences are

also registered in GenBank (Table 1, Fig 1).

DNA sequencing

Total DNA was extracted from the piece of foot using the 6100 Nucleic Acid Prepstation system (Applied

Biosystem) or the Epmotion 5075 robot (Eppendorf) following the manufacturer’s recommendations.

The barcode fragment of the COI gene (658 bp) was tentatively amplified using the universal primers

LCO1490 and HC02198 (Folmer, Black, Hoeh, Lutz & Vrijenhoek 1994). However, the rate of success

was very low (less than 10%), and several other protocols and primer pairs were tested. Finally, we were

able to amplify most of the specimens, including several for each morphological species hypothesis,

using the primers FISHR2 (Steinke & Hanner 2011) and 140F (Ketmaier, Giusti & Caccone 2006), and

the following protocol: PCR reactions were performed in 20 pi containing 3 ng of DNA, 10x reaction

buffer containing 15 m M MgCf, 0.26 m M dNTP, 0.3 pM of each primer, 5% DMSO, 1 mg/ml BSA,

and 1 unit of QBiotaq (MPBiomedicals). Amplification consisted of an initial denaturation step at 95°C

for 5’, followed by 38 cycles of denaturation at 95°C for 40”, annealing at 50°C for 40”, followed by

extension at 72°C for 50’. The final extension was at 72°C for 3’. PCR products were purified and

sequenced by the Eurofins sequencing facility. Both directions were sequenced to confirm accuracy of

each sequence.

Phylogenetic analyses

In addition to the specimens of Perotrochus sequenced by us, COI sequences of Pleurotomariidae from

GenBank were included in the dataset, as well as a sequence of Haliotis tuberculata (Haliotidae), used

as outgroup (Table 1). Phylogenetic analyses were performed using MrBayes (Huelsenbeck, Ronquist

& Hall 2001), running two parallel analyses, consisting each of five Markov chains of 20,000,000

generations with a sampling frequency of one tree each 2,500 generations. The number of swaps was set

to 3, and the chain temperature at 0.02. Parameters of the substitution model were estimated during the

analysis (6 substitution categories, a gamma-distributed rate variation across sites approximated in four

discrete categories and a proportion of invariable sites). A different model of substitution was applied

for each codon position of the COI gene. Convergence of each analysis was evaluated using Tracer 1.4.1

(Rambaut & Drummond 2007) to check that ESS values were all greater than 200 (default burning).

K2P genetic distances were calculated with MEGA 5 (Tamura et al. 2011).

Results

Morphological analyses

With the exception of some juveniles, we could separate all the specimens of the Perotrochus caledonicus

complex into four different morphotypes (Table 2): one group presents shell characters consistent with

the name-bearing holotype of P. caledonicus as described by Bouchet & Metivier (1982); the second

group corresponds to the “rugose” or “pustulose” form, and is described below as P. wareni sp. nov.;

a third group, with a geographically limited distribution within the Coral Sea, shows a microgranular

3

European Journal of Taxonomy 134: 1-23 (2015)

sculpture and is described as P. pseudogranulosus sp. nov.; the fourth group is readily assignable to P.

deforgesi.

Phylogenetic analyses

The phylogenetic analyses show that the P. caledonicus group is a well-supported clade (Posterior

Probability PP = 1) within the Pleurotomariidae (Fig. 2). Although based on a single gene, the tree would

also suggest that the genus Perotrochus is not monophyletic. Furthermore, some GenBank sequences

appear to be either misidentified or contaminated; for example, the sequence L78912.1, identified as

Bayerotrochus teramachii (Kuroda, 1955), is almost identical to three sequences identified as B. midas

(Bayer, 1965). Within P. caledonicus s.l., the analysis of the COI gene diversity revealed the presence

of four groups, each corresponding to a well supported clade in the phylogenetic tree (PP = 1 for each

of them), and totally congruent with the morphological analysis. P. deforgesi is the sister-clade of a

group that includes the specimens attributed to P. caledonicus and P. wareni sp. nov. The mean K2P

genetic distance between P. wareni sp. nov. and P. caledonicus is 3.8%, which is similar to the distance

Haliotis tuberculata JQ894846.1

- Entemnotrochus adansonianus L78910.1

- Entemnotrochus rumphii L78911.1

0.99r Perotrochus amabilis AY923929.1

•- Perotrochus maureri L78914.1

r Perotrochus quoyanus L78915.1

"I- Perotrochus lucaya L78916.1

0.99 r Mikadotrochus beyrichii AM049331.1

L Mikadotrochus beyrichii EU530109.1

Bayerotrochus africanus AMQ49330.1

Bayerotrochus midas L78913.1

Bayerotrochus teramachii L78912.1

Bayerotrochus midas AY296820.1

Bayerotrochus midas AY923930.1

0.93

T

- IVII

r M

r^fr 1

MNHN-IM-2007-32067

M N H N-1M-2007-32054

MNHN-IM-2007-32058

MNHN-IM-2009-7492

MNHN-IM-2009-7491

MNHN-IM-2009-7495

r MNHN-iM-2009-7487

f MNHN-IM-2007-32066

Hr MNHN-IM-2009-7489

P MN H N-1M-2007-32056

V MNHN-IM-2009-7488

¥ MNHN-IM-2007-32064

t IMNHN-IM-2009-7485

' MNHN-IM-2009-7493

MNHN-1M-2009-7486

MNHN-IM-2007-32060

MNHN-IM-2007-32063

MNHN-IM-2007-32088

MNHN-IM-2007-32087

MNHN-IM-2007-32090

MNHN-IM-2007-32083

- MNHN-IM-2007-32091

r MNHN-IM-2009-7494

f- MN HN-1M-2007-32062

t M N M N-1M-2007-32061

1 MNHN-IM-2007-32085

Perotrochus pseudogranulosus sp. nov.

fit

0.1 subst./site

£

Perotrochus deforgesi

r MNHN-IM-2007-36456

MNHN-IM-2007-36461

MNHN-IM-2007-34685

M N H N-l M-2007-36460

M N HN-1 M-2007-34686

M N HN-1 M-2007-34688

r M.NHN-IM-2007-34690

Ur MNHN-IM-2007-34677

f M NHN-l M-2007-34684

\ MNHN-IM-2007-34680

1 MN HN-I M-2007-34682

r MNHN-IM-2007-36541

fir MNHN-IM-2009-7476

\ MNHN-IM-2007-36302

1 t- MNHN-IM-2007-36596

— r MNHN-IM-2007-36519

H- MNHN-IM-2007-36557

r MNHN-IM-2009-7483

\s IMNHN-IM-2009-7480

MN HN-1M-2007-36301

MNHN-IM-2009-7478

MNHN-IM-2007-36303

M N H N-l M-2007-36592

MNHN-IM-2007-36595

r MNHN-IM-2007-36458

|L MNHN-IM-2007-36459

I MNHN-IM-2007-36542

' MNHN-IM-2007-36822

r MNHN-IM-2007-36463

f MNHN-IM-2007-36505

r MNHN-IM-2007-36455

V MNHN-IM-2007-36300

" MNHN-IM-2007-36546

Perotrochus wareni sp. nov.

Perotrochus caledonicus

Fig. 2. Bayesian phylogenetic tree. Posterior probabilities (> 0.90) are shown for each node.

4

ANSEEUW P. et al ., New species of Pleurotomariidae

between P. deforgesi and P. caledonicus (3.7%) and greater than the distance between P. deforgesi and

P. wareni sp. nov. (2.1%). This would tend to confirm that if P deforgesi is considered a valid species

- which is morphologically indisputable - then the two morphs (P. caledonicus s.s. and P. wareni sp.

nov.), included until now in P caledonicus , should be ranked as species as well. More surprisingly,

P. pseudogranulosus sp. nov. is the sister-group to a clade that includes the other three species, with a

mean genetic distance of 9.9% to the other three species, confirming its validity as a separate species.

Systematics

Phylum Mollusca Cuvier, 1795

Class Gastropoda Cuvier, 1795

Subclass Vetigastropoda Salvini-Plawen, 1980

Superfamily Pleurotomarioidea Swainson, 1840

Family Pleurotomariidae Swainson, 1840

Genus Perotrochus P. Fischer, 1885

Comparative material (all in MNHN)

Perotrochus caledonicus. 135 lots comprising 560 specimens (Table 1).

Perotrochus deforgesi. 36 lots comprising 59 specimens (Table 1).

Perotrochus wareni sp. nov.

um:lsid:zoobank.org:act:51E7B304-66D2-4AE6-84C0-3A4427203065

Figs 1A, 3A-F, 4A-J

Etymology

This new species is named in honor of Dr Anders Waren of Naturhistoriska Riksmuseet, Stockho lm , in

recognition of his lifetime interest in deep-sea exploration and his participation in expeditions around

New Caledonia and elsewhere in the South Pacific, many of which yielded specimens used in this paper.

Material examined

73 lots comprising 176 specimens (Table 1).

Type material

Holotype

NEW CAFEDONIA: a sequenced specimen, MNHN-IM-2007-36460.

Paratypes

NEW CALEDONIA: MNHN-IM-2007-34680 (Fig. 4A-B); MNHN-IM-2007-34684 (Fig. 4C-D);

MNHN-IM-2007-34685 (Fig. 4E-F); MNHN-IM-2007-36456 (Fig. 4G-H); MNHN-IM-2007-36461

(Fig. 41—J).

Type locality

Norfolk Ridge, MunidaBank, 22°59’ S, 168°2FE, 320-390 m (TERRASSES sta. DW3101).

Description (holotype)

Shell of medium size, solid, thick, general profile rather conical, with wealdy convex, rather straight¬

sided whorls, with a diameter a little smaller than its height (H/D ratio =1.08), numbering 10.5 teleoconch

whorls, with a mean spire angle of 65° with weakly impressed suture, whorl surface dull. Protoconch

glassy, rather obtusely depressed. Teleoconch with heavily beaded spiral cords very early on. Dominant

5

European Journal of Taxonomy 134: 1-23 (2015)

Fig. 3. Perotrochus wareni sp. nov., MNHN-IM-2007-36460, holotype (H = 50.3 mm). A-D. Teleoconch.

E. Slit. F. Protoconch.

6

s p

ANSEEUW P. et al.. New species of Pleurotomariidae

Fig. 4. Perotrochus wareni sp. nov., paratypes. A— B. MNHN-IM-2007-34680 (H = 28.2 mm )

MNHN-IM-2007-34684 (H = 43.8 mm). E-F. MNHN-IM-2007-34685 (H = 21.5 m m)

MNHN-IM-2007-36456 (H = 55.1 mm). I-J. MNHN-IM-2007-36461 (H = 48.8 mm )

7

European Journal of Taxonomy 134: 1-23 (2015)

teleoconch sculpture consisting of finely but strongly beaded spiral cords, intersecting less marked axial

riblets. Periphery of basal disc crenulated due to a strongly marked spiral cord running at edge of disc.

On last whorl, 13 spiral cords above selenizone, 4 spiral cords below, and 2 major cords in the selenizone

itself Slit short, about Vz* the circumference of last whorl, situated below midwhorl, and very narrow.

Aperture rectangularly depressed. Basal disc rather flat, depressed in its center and sharply edged at its

periphery, with a very wide (extending over 40% of base diameter) callus pad which is finely ridged

radially. Inside aperture, inner slit lips only partially covered by nacre, leaving a V-shaped area uncovered

(approximately 25% of surface of inner slit lip extremity in aperture uncovered by nacre, showing only

porcellaneous layer). Nacre coverage thick, no surface sculpture showing through it. Background color

yellowish beige, with some faint axial orange-red flammulations, not really arranged into a distinct

checker-board pattern; basal disc of same colour, with some faint orange axial flammulation reaching

only to basal disc edge. Operculum small, multispiral, circular, light yellowish.

Measurements

Ma xim um basal diameter (D) 46.75 mm, minimum diameter 43.9 mm Height (H) 50.27 mm H/D =

1.08. Length of slit at upper margin 26.7 mm, at lower margin 16.2 mm Slit width: 1.6 mm ; slit length:

V6.5 i th of circumference of last whorl. Weight of empty shell 26.3 g.

Discussion

One of the distinctive shell characters separating Perotrochus wareni sp. nov. from P. caledonicus is

the well-marked beading on the teleoconch spiral cords, visible also on the earlier whorls, giving it at

first glance its typical pustulose or granular appearance. The intensity of the beading varies between

specimens, leading to “heavily beaded”/“very pustulose” specimens and to “weakly beaded” / “light

pustulose” specimens, with all different intergrades. This variability may reflect environmental

conditions, as the beading intensity is generally stable within one haul / lot and varies between hauls /

lots. Leaving aside beading intensity, shell characters are quite stable in P. wareni sp. nov. The general

profile of the teleoconch, the outline of the aperture, the selenizone and slit width, and the extension of

the callus on the basal disc, all show consistent differences between P. wareni sp. nov., P. caledonicus

and P. pseudogranulosus sp. nov. (Table 2). Perotrochus wareni sp. nov. also bears some resemblance to

P. gotoi Anseeuw, 1990, and, in fact, a specimen of the granulated P. “ cfr. caledonicus ” ([i.e., P. wareni

sp. nov.) had been used for comparison at the time of its original description (Anseeuw 1990).

Perotrochus wareni sp. nov. can be separated from P. gotoi by its somewhat shorter slit length (A. 51 th of

basal diameter in wareni sp. nov. vs VcP in gotoi), the larger number of spiral cords on adult specimens,

the more irregularly banded checkerboard colour pattern in the area below the selenizone, the heavier

and thicker shell (around 60-70% heavier at comparable shell sizes), the less extensive area uncovered

by the nacreous layer inside the apertural inner slit lip extremities, and the umbilical callus occupying

a much larger surface on the basal disc (45% in P. wareni sp. nov. vs 28% in P. gotoi). The two species,

however, share (also with Mikadotrochus salmianus (Rolle, 1899) the nacreous coverage of the inner

slit lips in the aperture, a feature that separates them from Perotrochus caledonicus s.s. and P. pseudo¬

granulosus sp. nov. (Table 2, Figs 3-4, 7). Other features, like a deeper, more intense colour pattern on

the teleoconch and basal disc, fine microgranulosity on the spiral cords, a thin, light shell, and a more

lustrous shell surface, further separate Perotrochus pseudogranulosus sp. nov. from P. wareni sp. nov. (Table

2, Figs 3-6).

8

ANSEEUW P. et al ., New species of Pleurotomariidae

Perotrochus pseudogranulosus sp. nov.

um:lsid:zoobank.org:act:CE972B51-FB50-4C06-BlB2-DD01A6BC737D

Figs ID, 5A-F, 6A-J

Etymology

The specific epithet emphasizes the beaded spiral sculpture of the species.

Material examined

40 lots comprising 84 specimens (Table 1).

Type material

Holotype

NEW CALEDONIA: a sequenced specimen, MNHN-IM-2009-7495.

Paratypes

NEW CALEDONIA: MNHN-IM-2007-32058 (Fig. 6A-B); MNHN-IM-2007-32066 (Fig. 6C-D);

MNHN-IM-2009-7485 (Fig. 6E-F); MNHN-IM-2009-7491 (Fig. 6G-H); MNHN-IM-2009-7493

(Fig. 6I-J).

Type locality

Coral Sea, Capel Bank, 24°45’ S, 159 0 42’ E, 348-354 m (EBISCO sta. CP2494).

Description (holotype)

Shell of medium size, light, thin, general profile rather conical, with weakly convex to straight-sided

whorls with weakly impressed suture, diameter significantly exceeding height (H/D = 0.78), numbering

8 teleoconch whorls, with a mean spire angle of 80°. Protoconch ivory white, depressed. Dominant

sculpture of teleoconch consisting of numerous lightly beaded spiral cords, with microsculptural pattern

of fine radiating threads, giving the entire whorl surface a shiny metallic luster. On last whorl, 11 spiral

cords above selenizone, 7 below and 3 major cords in the selenizone itself. Slit long, about A 1 ' 1 the

circumference of the last whorl, situated below midwhorl, and rather narrow. Aperture depressed. Basal

disc rather flattened, with angular edge, with a relatively narrow (extending over 30% of base diameter)

light nacreous callus pad which is finely ridged radially and ends in a raised porcellaneous edge. Inside

the aperture inner slit lips nearly completely covered by nacre, leaving a narrow area (approximately 15%

of the surface) parallel to the inner slit lips uncovered, showing only porcellaneous layer. Background

colour yellowish beige, with intense reddish crimson colour markings arranged in very regular checker¬

board pattern, overall reinforcing color intensity; basal disc showing some contrasting reddish crimson

flammulations, particularly visible at its periphery, and more yellowish tan towards the center. Operculum

small, multispiral, circular, light yellowish (fallen off/missing in holotype).

Measurements

Ma xim um basal diameter (D) 64.9 mm, minimum diameter 59.1 mm Height (H) 50.7 mm H/D = 0.78.

Depth of slit at upper margin 42.1 mm, depth of slit at lower margin 26.9 mm Slit width 3.1 mm Slit

length: A.64 th of circumference of last whorl. Weight of empty shell 42.6 g.

Discussion

Perotrochus pseudogranulosus sp. nov. most closely resembles P. caledonicus (Fig. 7) at first glance, but

is distinguished by its more conical and higher shell, a more flattened basal disc profile and a more intense

and regular checkerboard colour pattern and more lustrous shell surface. It differs from P. wareni sp.

nov. by its weakly beaded spiral cords with a microsculpture of fine radiating threads, its more intensely

9

European Journal of Taxonomy 134: 1-23 (2015)

Fig. 5. Perotrochus pseudogranulosus sp. nov., MNHN-IM-2009-7495, holotype (H = 54.3 mm).

A-D. Teleoconch. E. Slit. F. Protoconch.

10

ANSEEUW P. et al.. New species of Pleurotomariidae

Fig. 6. Perotrochus pseudogranulosus sp. nov., paratypes. A—B. MNHN-IM-2007-32058 (H = 53 mm ).

C-D. MNHN-IM-2007-32066 (H = 29.8 mm). E-F. MNHN-IM-2009-7485 (H = 42.1 mm)

G-H. MNHN-IM-2009-7491 (H = 36 mm). I-J. MNHN-IM-2009-7493 (H = 32.1 m m)

11

European Journal of Taxonomy 134: 1-23 (2015)

marked checkerboard colour pattern, with metallic luster, a much longer slit and a much smaller callus

pad area on the basal disc. Finally, it differs from P. deforgesi (Fig. 7), the only other species occurring

in the Coral Sea, by its general outline which is distinctly higher conical, its thin shell, its more intense

checkerboard colour markings, its less granular spiral cords and smaller callus pad coverage on the basal

disc.

Some specimens of P. pseudogranulosus sp. nov. have over the years turned up in the shell trade as

“P. cfr. caledonicus ”, supposedly originating from NW Australia or even from the South China Sea

(Anseeuw & Goto 1996). However, based on the lack of precise and trustworthy locality data, the

lack of more recent confirmation of those alleged findings and, most of all, the general unavailability

of such material for study, we reject these localities as intentionally or unintentionally unproven and

unverifiable.

Discussion

The geographical distribution within the New Caledonia region (Fig. 1) is different for the four species.

Many samples of Perotrochus caledonicus s.s. originate from the SW off He des Pins, southern New

Caledonia, with scattered specimens from Norfolk Ridge, the Loyalty Ridge and the Far North of New

Caledonia (Grand Passage); the average depth of occurrence, based on 135 lots, is 407 meters. The

distribution of Perotrochus wareni sp. nov. includes Norfolk Ridge, the Loyalty Ridge, the Far North

of New Caledonia (Grand Passage) and the Lansdowne Plateau in the Coral Sea; the average depth

of occurrence, based on 73 lots, is 344 meters. Perotrochus caledonicus and P. wareni sp. nov. thus

co-occur locally in the south of New Caledonia, Norfolk Ridge and Grand Passage, where they have

different bathymetric preferences, although they were sometimes found at the same stations (LITHIST

sta. CP14 and sta. CP16; NORFOLK 1 sta. DW1658 and sta. DW1709; MUSORSTOM4 sta. DW222;

TERRASSES sta. DW3101 and sta. DW3110). Perotrochus pseudogranulosus sp. nov. has a distribution

restricted to the Chesterfield Plateau, with a mean depth of 360 meters based on 40 lots. Perotrochus

deforgesi is also restricted to the Coral Sea, but occurs only on the Bellona Plateau and Capel Bank,

thus allopatrically with regard to P. pseudogranulosus sp. nov., but at similar depths (mean depth of 380

meters based on 36 lots).

The morphological, molecular and micro distribution data, thus all converge to support the conclusion

that there are four small Perotrochus in the New Caledonia region, of which two have been described

as new in the present paper.

At genus level, although Anseeuw & Poppe (2005) and Aktipis & Giribet (2010) had used the

combinations Mikadotrochus caledonicus and M. deforgesi , the molecular phylogeny shows that

Perotrochus caledonicus and P. deforgesi are not congeneric with Pleurotomaria beyrichii Hilgendorf,

1877, the type species of Mikadotrochus Lindholm, 1927. However, the tree also shows that the four

small New Caledonia Perotrochus do not form a monophyletic group with Pleurotomaria quoyana P.

Fischer & Bernardi, 1856, the type species of Perotrochus P. Fischer, 1885. Perotrochus quoyanus is a

species from the western Atlantic, and the species (P. amabilis (Bayer, 1963), P. maureri Harasewych &

Askew, 1993, P. lucaya Bayer, 1965) that cluster with it in the tree are also from the western Atlantic.

It thus appears that, although they are traditionally placed in Perotrochus (e.g., Anseeuw & Goto 1996;

Harasewych 2002), the small pleurotomariids from the Pacific are not congeneric with true Perotrochus

from the western Atlantic, and a new genus will have to be established to classify them.

Acknowledgments

The material on which this paper is based has been accumulated during many expeditions of the Tropical

Deep-Sea Benthos programme in the New Caledonia region. We refer to Bouchet et al. (2008) for an

12

ANSEEUW P. et al ., New species of Pleurotomariidae

Fig. 7. A-F. Perotrochus caledonicus Bouchet & Metivier, 1982. A-B. MNHN-IM-2000-1262, holotype

(H = 31.2 mm). C-D. MNHN-IM-2007-36300 (H = 51.2 mm). E-F. MNHN-IM-2007-36301 (H = 47.5

mm). — G-L. Perotrochus deforgesi Metivier, 1990. G-H. MNHN-IM-2000-1391, holotype (H = 33.9

m m ) I-J. MNHN-IM-2007-32062 (H = 33.7 mm). K-L. MNHN-IM-2007-32085 (H = 25.1 mm).

13

European Journal of Taxonomy 134: 1-23 (2015)

overview of the programme and the acknowledgements to the captains, principal scientists and crew

involved are applicable here. We thank Yoshihiro Goto and Guido Poppe, with whom we discussed

the taxonomic hypotheses presented in this paper. Manuel Tenorio and Javier Conde attempted a

morphometric analysis (results not shown). Barbara Buge, Virginie Heros, Julien Brisset and Philippe

Maestrati curated the material, Philippe Maestrati and Manuel Caballer-Gutierrez provided the photos.

This work was supported by the Service de Systematique Moleculaire (UMS 2700 CNRS-MNHN), and

we acknowledge the advice of M.G. Harasewych on DNA amplification in pleurotomariids.

References

Aktipis S.W. & Giribet G. 2010. A phylogeny of Vetigastropoda and other “archaeogastropods”: re¬

organizing old gastropod clades. Invertebrate Biology 129 (3): 220-240. http://dx.doi.org/10.llll/

i. 1744-7410.2010.00198.x

Anseeuw P. 1990. Anew species of pleurotomariid gastropod from the Philippines. La Conchiglia 22:

10-15.

Anseeuw P. & Goto Y. 1996. The Living Pleurotomariidae: 1. Elle Scientific Publications, Osaka.

Anseeuw P. & Poppe G. 2005. Pleurotomariidae: An iconographic visit anno 2005. Visaya supplement

1, Cebu, Philippines.

Bouchet P, Heros V., Lozouet P. & Maestrati P. 2008. A quarter-century of deep-sea malacological

exploration in the South and West Pacific: Where do we stand? How far to go? In: Heros V., Cowie R.H.

& Bouchet P. (eds) Tropical Deep-Sea Benthos 25. Memoires du Museum national d’Histoire naturelle

196: 9—40. Museum national d’Histoire naturelle, Paris.

Bouchet P. & MetivierB. 1982. Living Pleurotomariidae (Mollusca: Gastropoda) from the South Pacific.

New Zealand Journal of Zoology 9: 309-317.

Folmer O., Black M., Hoeh W., Lutz R. & Vrijenhoek R. 1994. DNA primers for amplification of

mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine

Biology and Biotechnology 3: 294-299.

Harasewych M.G. 2002. Pleurotomarioidean gastropods. Advances in Marine Biology 42: 235-292.

Huelsenbeck J.P, Ronquist F. & Hall B. 2001. MrBayes: Bayesian inference of phylogeny. Bioinformatics

17: 754-755. http://dx.doi.Org/10.1093/bioinfonnatics/17.8.754

Ketmaier V., Giusti F. & Caccone A. 2006. Molecular phylogeny and historical biogeography of the

land snail genus Solatopupa (Pulmonata) in the peri-Tyrrhenian area. Molecular Phylogenetics and

Evolution 39 (2): 439—451. http://dx.doi.Org/10.1016/i.vmpev.2005.12.008

Metivier B. 1990 Description of a new Perotrochus from the Coral Sea, Southwest Pacific (Gastropoda:

Pleurotomariidae). Venus 49(1): 1-7.

Rambaut A. & Drummond A.J. 2007. Tracer v. 1.4. Available from http://beast.bio.ed.ac.uk/Tracer

[accessed on 25 Jun. 2014],

Steinke D. & Hanner R. 2011. The FISH-BOL collaborators’ protocol. Mitochondrial DNA 22 (SI):

10-14. http://dx.doi.org/10.3109/19401736.201Q.536538

Tamura K., Peterson D., Peterson N., Stecher G., Nei M. & Kumar S. 2011. MEGA5: Molecular

Evolutionary Genetics Analysis using Maximum Likelihood, Evolutionary Distance, and Maximum

Parsimony methods. Molecular Biology and Evolution 28: 2731-2739. http://dx.doi.org/10.1093/

molbev/msrl21

14

ANSEEUW P. et al ., New species of Pleurotomariidae

Manuscript received: 1 April 2015

Manuscript accepted: 11 June 2015

Published on: 24 August 2015

Topic editor: Rudy Jocque

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; Botanic Garden Meise,

Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London,

United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History

Museum of Denmark, Copenhagen, Denmark.

15

European Journal of Taxonomy 134: 1-23 (2015)

Table 1. List of specimens.

MNHN ID

N

SPECIES

MNHN EXPEDITION

STATION

GENERALAREA

COORDINATES; DEPTH (in)

BOLD ID

GENBANK #

IM-2000-1262

1

P. caledonicus

VAUBAN

DR15

22°49’S, 167°12’E; 390-395

IM-2000-1324

2

P. deforgesi

CHALCAL1

DC32

Chesterfield

19°43’S, 158°33’E; 350

IM-2000-1361

1

P. deforgesi

MUSORSTOM 5

DW337

Chesterfield

19°54’S, 158°38’E; 412-430

IM-2000-1387

2

P. caledonicus

VAUBAN

DR15

22°49’S, 167°12’E; 390-395

IM-2000-1389

1

P. deforgesi

MUSORSTOM 5

DC361

Chesterfield

19°52’S ! 158°38’E; 400

IM-2007-32054

1

P. pseiidograniilosus

EBISCO

DW2530

S Nova Bank

22°48’S, 159°23’E; 338-343

PLEROOlO-15

KR087206

IM-2007-32055

1

P. pseudogranulosus

EBISCO

DW2530

S Nova Bank

22°48’S, 159°23’E; 338-343

IM-2007-32056

1

P. pseiidograniilosus

EBISCO

DW2530

S Nova Bank

22°48’S, 159°23’E; 338-343

PLEROOll-15

KR087205

IM-2007-32057

1

P. pseiidograniilosus

EBISCO

DW2530

S Nova Bank

22°48’S, 159°23’E; 338-343

IM-2007-32058

1

P. pseiidograniilosus

EBISCO

DW2501

Capel Bank

24°50’S, 159°51’E; 325-520

PLERO058-15

KR087194

IM-2007-32059

1

P. wareni

EBISCO

DW2631

S Lansdowne

21°03’S, 160°44’E; 372-404

IM-2007-32060

1

P. deforgesi

EBISCO

DW2577

N Bellona

20°20’S, 158°39’E; 399-602

PLERO012-15

KR087218

IM-2007-32061

1

P. deforgesi

EBISCO

DW2577

N Bellona

20°20’S, 158°39’E; 399-602

PLER0002-15

KR087208

IM-2007-32062

1

P. deforgesi

EBISCO

CP2567

NW Bellona

20°20’S, 158°42’E; 395-400

PLERO013-15

KR087217

IM-2007-32063

1

P. deforgesi

EBISCO

CP2593

Chesterfield

19°43’S, 158°32’E; 300-323

PLERO014-15

KR087216

IM-2007-32064

1

P. pseiidograniilosus

EBISCO

DW2526

S Nova Bank

22°46’S, 159°23’E; 330-340

PLERO015-15

KR087193

IM-2007-32065

1

P. pseiidograniilosus

EBISCO

DW2526

S Nova Bank

22°46’S, 159°23’E; 330-340

IM-2007-32066

1

P. pseiidograniilosus

EBISCO

DW2526

S Nova Bank

22°46’S, 159°23’E; 330-340

PLERO016-15

KR087204

IM-2007-32067

1

P. pseiidograniilosus

EBISCO

DW2525

S Nova Bank

22°48’S, 159°23’E; 408-410

PLERO017-15

KR087203

IM-2007-32068

1

P. wareni

EBISCO

DW2634

S Lansdowne

21°05’S, 160°46’E; 342-347

IM-2007-32069

1

P. wareni

EBISCO

DW2634

S Lansdowne

21°05’S, 160°46’E; 342-347

IM-2007-32070

1

P. pseiidograniilosus

EBISCO

DW2525

S Nova Bank

22°48’S, 159°23’E; 408-410

IM-2007-32071

1

P. pseiidograniilosus

EBISCO

DW2525

S Nova Bank

22°48’S, 159°23’E; 408-410

IM-2007-32072

1

P. pseiidograniilosus

EBISCO

DW2528

S Nova Bank

22°49’S, 159°23’E; 320-345

IM-2007-32073

1

P. pseiidograniilosus

EBISCO

DW2528

S Nova Bank

22°49’S, 159°23’E; 320-345

IM-2007-32074

1

P. pseiidograniilosus

EBISCO

DW2528

S Nova Bank

22°49’S, 159°23’E; 320-345

IM-2007-32075

1

P. pseiidograniilosus

EBISCO

DW2528

S Nova Bank

22°49’S, 159°23’E; 320-345

IM-2007-32076

1

P. pseudogi'anulosus

EBISCO

CP2529

S Nova Bank

22°47’S, 159°23’E; 330-340

IM-2007-32077

1

P. pseiidograniilosus

EBISCO

CP2529

S Nova Bank

22°47’S, 159°23’E; 330-340

IM-2007-32078

1

P. pseiidograniilosus

EBISCO

CP2529

S Nova Bank

22°47’S, 159°23’E; 330-340

IM-2007-32079

1

P. pseiidograniilosus

EBISCO

CP2529

S Nova Bank

22°47’S, 159°23’E; 330-340

IM-2007-32080

1

P. pseiidograniilosus

EBISCO

CP2529

S Nova Bank

22°47’S, 159°23’E; 330-340

IM-2007-32081

1

P. pseiidograniilosus

EBISCO

CP2529

S Nova Bank

22°47’S, 159°23’E; 330-340

IM-2007-32082

1

P. pseiidograniilosus

EBISCO

CP2529

S Nova Bank

22°47’S, 159°23’E; 330-340

IM-2007-32083

1

P. deforgesi

EBISCO

CP2593

Chesterfield

19°43’S, 158°32’E; 300-323

PLER0004-15

KR087207

IM-2007-32084

1

P. deforgesi

EBISCO

CP2593

Chesterfield

19°43’S, 158°32’E; 300-323

IM-2007-32085

1

P. deforgesi

EBISCO

CP2593

Chesterfield

19°43’S, 158°32’E; 300-323

PLER0005-15

KR087215

IM-2007-32086

1

P. deforgesi

EBISCO

CP2593

Chesterfield

19°43’S, 158°32’E; 300-323

IM-2007-32087

1

P. deforgesi

EBISCO

CP2593

Chesterfield

19°43’S, 158°32’E; 300-323

PLER0006-15

KR087214

IM-2007-32088

1

P. deforgesi

EBISCO

CP2593

Chesterfield

19°43’S, 158°32’E; 300-323

PLER0007-15

RR087213

IM-2007-32089

1

P. deforgesi

EBISCO

CP2593

Chesterfield

19°43’S, 158°32’E; 300-323

[M-2007-32090

1

P. deforgesi

EBISCO

CP2593

Chesterfield

19°43’S, 158°32’E; 300-323

PLER0008-15

RR087212

IM-2007-32091

1

P. deforgesi

EBISCO

CP2593

Chesterfield

19°43’S, 158°32’E; 300-323

PLER0009-15

KR087211

16

N

1

ANSEEUW P. et al.. New species of Pleurotomariidae

p.

p

p.

SPECIES

MNHN EXPEDITION

STATION

GENERALAREA

COORDINATES; DEPTH (in)

BOLD ID

GENBANK#

P. wareni

CONCALIS

DW2946

Grand Passage

19°02’S, 163°27’E; 276-277

PLERO024-15

KR087219

P. wareni

CONCALIS

DW2946

Grand Passage

19°02’S, 163°27’E; 276-277

P. wareni

CONCALIS

DW3025

Grand Passage

18°57’S, 163°23’E; 396-400

P. wareni

CONCALIS

DW2944

Grand Passage

18°59’S, 163°24’E; 320-335

PLERO025-15

KR087228

P. wareni

CONCALIS

DW2949

Grand Passage

19°00’S, 163°28’E; 261-272

P. wareni

CONCALIS

DW3024

Grand Passage

18°57’S, 163°22’E; 349-370

PLERO018-15

KR087224

P. wareni

CONCALIS

DW2941

Grand Passage

19°3’S, 163°27’E; 271-272

P. wareni

CONCALIS

DW2973

Grand Passage

18°14’S, 163°06’E; 275-288

PLERO019-15

KR087223

P. wareni

CONCALIS

DW2943

Grand Passage

18°57’S, 163°23’E; 380-430

PLER0020-15

KR087222

P. wareni

CONCALIS

DW2973

Grand Passage

18°14’S, 163°06’E; 275-288

PLERO021-15

KR087221

P. wareni

CONCALIS

DW2974

Grand Passage

18°15’S, 163°06’E; 283-326

P. wareni

CONCALIS

CP2975

Grand Passage

18°15’S, 163°06’E; 297-316

PLERO022-15

KR087220

P. wareni

CONCALIS

DW2977

Grand Passage

18°15’S, 163°07’E; 326-368

P. wareni

CONCALIS

DW2977

Grand Passage

18°15’S, 163°07’E; 326-368

PLERO023-15

KR087229

P. wareni

CONCALIS

DW2977

Grand Passage

18°15’S, 163°07’E; 326-368

caledonicus

TERRASSES

DW3101

Norfolk Ridge

22°59’S, 168°21’E; 320-390

PLERO044-15

KR087186

caledonicus

TERRASSES

DW3075

Norfolk Ridge

23°17’S, 168°14’E; 270

PLER0040-15

KR087190

caledonicus

TERRASSES

DW3075

Norfolk Ridge

23°17’S, 168°14’E; 270

PLERO041-15

KR087189

caledonicus

TERRASSES

DW3035

Loyalty Ridge

22°41’S, 168°56’E; 790-800

PLERO038-15

KR087192

caledonicus

TERRASSES

DW3035

Loyalty Ridge

22°41’S, 168°56’E; 790-800

caledonicus

TERRASSES

DW3075

Norfolk Ridge

23°17’S, 168°14’E; 270

PLERO042-15

KR087188

P. wareni

TERRASSES

DW3101

Norfolk Ridge

22°59’S, 168°21’E; 320-390

PLERO045-15

KR087227

caledonicus

TERRASSES

DW3075

Norfolk Ridge

23°17’S, 168°14’E; 270

caledonicus

TERRASSES

DW3073

Norfolk Ridge

23°17’S, 168°14’E; 250-270

PLERO039-15

KR087191

caledonicus

TERRASSES

DW3076

Norfolk Ridge

23°14’S, 168°13’E; 390-570

PLERO043-15

KR087187

P. wareni

TERRASSES

DW3101

Norfolk Ridge

22°59’S, 168°21’E; 320-390

PLERO046-15

KR087226

P. wareni

TERRASSES

DW3101

Norfolk Ridge

22°59’S, 168°21’E; 320-390

PLERO047-15

KR087225

P. wareni

TERRASSES

DW3101

Norfolk Ridge

22°59’S, 168°21’E; 320-390

caledonicus

TERRASSES

DW3110

Norfolk Ridge

23°02’S, 168°16’E; 270-310

PLERO048-15

KR087182

caledonicus

TERRASSES

DW3127

lie des Pins

23°00’S, 167°16’E; 400-420

PLERO052-15

KR087178

caledonicus

TERRASSES

DW3127

lie des Pins

23°00’S, 167° 16’E; 400-420

PLERO053-15

KR087177

caledonicus

TERRASSES

DW3123

He des Pins

22°53’S, 167°13’E; 420-450

caledonicus

TERRASSES

DW3122

He des Pins

22°47’S, 167°12’E; 390-410

PLERO034-15

KR087185

caledonicus

TERRASSES

DW3122

He des Pins

22°47’S, 167°12’E; 390-410

PLERO049-15

KR087181

caledonicus

TERRASSES

DW3124

He des Pins

22°54’S, 167°15’E; 460

caledonicus

TERRASSES

DW3123

He des Pins

22°53’S, 167°13’E; 420-450

PLERO051-15

KR087179

caledonicus

TERRASSES

DW3122

He des Pins

22°47’S, 167°12’E; 390-410

PLER0050-15

KR087180

caledonicus

TERRASSES

DW3127

He des Pins

23°00’S, 167°16’E; 400-420

caledonicus

TERRASSES

CP3048

Norfolk Ridge

23°44’S, 168°16’E; 380-400

PLERO035-15

KR087173

caledonicus

TERRASSES

DW3127

lie des Pins

23°00’S, 167°16’E; 400-420

PLERO054-15

KR087176

caledonicus

TERRASSES

CP3052

Norfolk Ridge

23°49’S, 168°16’E; 500-800

PLERO037-15

KR087171

caledonicus

TERRASSES

CP3049

Norfolk Ridge

23°42’S, 168°16’E; 380-410

PLERO036-15

KR087172

caledonicus

NORFOLK 2

DW2156

lie des Pins

22°54’S, 167°15’E; 468-500

PLER0003-15

KR087183

caledonicus

NORFOLK 2

DW2156

He des Pins

22°54’S, 167°15’E; 468-500

caledonicus

NORFOLK 2

DW2156

He des Pins

22°54’S, 167°15’E; 468-500

PLEROOOl-15

KR087174

17

European Journal of Taxonomy 134: 1-23 (2015)

MNHNID

N

SPECIES

MNHN EXPEDITION

STATION

GENERALAREA

COORDINATES; DEPTH (in)

BOLD ID

GENBANK #

IM-2009-7479

1

P. caledonicus

NORFOLK 2

DW2156

lie des Pins

22°54’S, 167°15’E; 468-500

IM-2009-7480

1

P. caledonicus

NORFOLK 2

DW2156

lie des Pins

22°54’S, 167°15’E; 468-500

PLERO059-15

KR087184

IM-2009-7481

1

P. caledonicus

NORFOLK 2

DW2156

lie des Pins

22°54’S, 167°15’E; 468-500

IM-2009-7482

1

P. caledonicus

NORFOLK 2

DW2156

He des Pins

22°54’S, 167°15’E; 468-500

IM-2009-7483

1

P. caledonicus

NORFOLK 2

DW2156

He des Pins

22°54’S, 167°15’E; 468-500

PLERO055-15

KR087175

IM-2009-7485

1

P. pseudogranulosus

EBISCO

DW2528

S Nova Bank

22°49’S, 159°23’E; 320-345

PLERO026-15

KR087202

IM-2009-7486

2

P. deforgesi

EBISCO

CP2593

Chesterfield

19°43’S, 158°32’E; 300-323

PLERO056-15

KR087210

IM-2009-7487

1

P. pseudogranulosus

EBISCO

CP2529

S Nova Bank

22°47’S, 159°23’E; 330-340

PLERO027-15

KR087201

IM-2009-7488

1

P. pseudogranulosus

EBISCO

CP2529

S Nova Bank

22°47’S, 159°23’E; 330-340

PLERO028-15

KR087200

IM-2009-7489

1

P. pseudogranulosus

EBISCO

DW2526

S Nova Bank

22°46’S, 159°23’E; 330-340

PLERO029-15

KR087199

IM-2009-7490

1

P. deforgesi

EBISCO

CP2567

NW Bellona

20°20’S, 158°42’E; 395-400

IM-2009-7491

1

P. pseudogranulosus

EBISCO

DW2525

S Nova Bank

22°48’S, 159°23’E; 408-410

PLER0030-15

KR087198

IM-2009-7492

1

P. pseudogranulosus

EBISCO

DW2526

S Nova Bank

22°46’S, 159°23’E; 330-340

PLERO031-15

KR087197

IM-2009-7493

1

P. pseudogranulosus

EBISCO

DW2530

S Nova Bank

22°48’S, 159°23’E; 338-343

PLERO057-15

KR087195

IM-2009-7494

1

P. deforgesi

EBISCO

DW2608

Chesterfield

19°33’S, 158°40’E; 393-396

PLERO032-15

KR087209

IM-2009-7495

1

P. pseudogranulosus

EBISCO

CP2494

Capel Bank

24°45’S, 159°42’E; 348-354

PLERO033-15

KR087196

IM-2009-7496

1

P. pseudogranulosus

EBISCO

DW2501

Capel Bank

24°50’S, 159°51’E; 325-520

IM-2009-7497

1

P. caledonicus

NORFOLK 2

DW2031

Stylaster Bank

23°39’S, 167°44’E; 440

IM-2010-11117

7

P. caledonicus

SMIB 8

DW197-199

lie des Pins

22°51’—22°52’S,

168°12’-168°13’E; 408-436

IM-2010-11118

1

P. caledonicus

SMIB 8

DW185

Antigonia Bank

23°15’S, 168°04’E;311-355

IM-2010-11119

27

P. caledonicus

LITHIST

CP16

Jumeau Ouest Bank

23°43’S, 168°16’E; 379-391

IM-2010-11120

1

P. caledonicus

LITHIST

DW13

Jumeau Est Bank

23°45’S, 168°17’E; 400

IM-2010-11121

1

P. pseudogranulosus

EBISCO

DW2528

S Nova Bank

22°49’S, 159°23’E; 320-345

IM-2010-11122

1

P. caledonicus

SMIB 2

DW05

S He des Pins

22°56’S, 167°15’E; 398-410

IM-2010-11123

1

P. caledonicus

BIOCAL

DW38

S He des Pins

23°00’S, 167°15’E; 360

IM-2010-11124

2

P. caledonicus

BATHUS 3

DW829

Norfolk Ridge

23°21’S, 168°02’E; 386-390

IM-2010-11125

13

P. caledonicus

BATHUS 3

CP811

Norfolk Ridge

23°41’S, 168°16’E; 383-408

IM-2010-11126

9

P. caledonicus

BATHUS 3

DW817

Norfolk Ridge

23°42’S, 168°16’E; 405-410

IM-2010-11127

3

P. caledonicus

BATHUS 3

DW818

Norfolk Ridge

23°44’S, 168°16’E; 394-401

IM-2010-11128

1

P. caledonicus

BATHUS 3

DW829

Norfolk Ridge

23°21’S, 168°02’E; 386-390

IM-2010-11129

5

P. caledonicus

NORFOLK 2

CP2048

Jumeau Est Bank

23°44’S, 168°16’E; 380-389

IM-2010-11130

4

P. caledonicus

NORFOLK 2

DW2049

Jumeau Est Bank

23°43’S, 168°15’E; 470-621

IM-2010-11131

7

P. caledonicus

NORFOLK 2

CP2050

Jumeau Est Bank

23°42’S, 168°16’E; 377

IM-2010-11132

2

P. caledonicus

NORFOLK 2

DW2052

Jumeau Est Bank

23°42’S, 168°15’E; 473-525

IM-2010-11133

1

P. caledonicus

NORFOLK 2

DW2108

Jumeau Est Bank

23°47’S, 168°17’E; 403-440

IM-2010-11134

1

P. caledonicus

NORFOLK 2

CP2114

Jumeau Est Bank

23°45’S, 168°17’E; 390-398

IM-2010-11135

8

P. caledonicus

NORFOLK 2

CH2115

Jumeau Est Bank

23°45’S, 168°17’E; 377-401

IM-2010-11136

2

P. caledonicus

NORFOLK 2

CP2118

Antigonia Bank

23°23’S, 168°01’E; 383-393

IM-2010-11137

3

P. caledonicus

NORFOLK 2

DW2126

Crypthelia Bank

23°16’S, 168°14’E; 398-550

IM-2010-11138

6

P. caledonicus

NORFOLK 2

DW2155

lie des Pins

22°52’S, 167°13’E;453-455

IM-2010-11139

4

P. caledonicus

NORFOLK 2

CP2153

lie des Pins

22°47’S, 167°12’E; 395-400

IM-2010-11140

4

P. caledonicus

NORFOLK 2

DW2156

He des Pins

22°54’S, 167°15’E; 468-500

IM-2010-11141

1

P. caledonicus

NORFOLK 2

DW2023

Brachiopode Bank

23°27’S, 167°5 HE; 282-297

IM-2010-11142

1

P. caledonicus

NORFOLK 2

DW2027

Brachiopode Bank

23°26’S, 167 0 51’E; 465-650

IM-2010-11143

2

P. wareni

BATHUS 4

DW926

Grand Passage

18°57’S, 163°25’E; 325-330

18

N

1

2

14

8

2

8

10

36

8

13

5

9

4

13

11

6

23

14

3

5

1

15

21

1

2

4

3

2

17

6

1

13

7

ANSEEUW P. e? al New species of Pleurotomariidae

p

p.

SPECIES

MNHN EXPEDITION

STATION

P wareni

BATHUS 4

DW925

caledonicus

SMIB 8

DW185

caledonicus

SMIB 8

DW179

caledonicus

SMIB 8

DW199

caledonicus

SMIB 8

DW197

caledonicus

SMIB 8

DW198

caledonicus

NORFOLK 1

DW1704

caledonicus

NORFOLK 1

DW1657

caledonicus

NORFOLK 1

DW1658

caledonicus

NORFOLK 1

CP 1705

caledonicus

NORFOLK 1

DW1733

caledonicus

NORFOLK 1

CP1708

caledonicus

NORFOLK 1

DW1709

caledonicus

NORFOLK 1

DW1710

caledonicus

NORFOLK 1

DW1707

caledonicus

NORFOLK 1

DW1738

caledonicus

NORFOLK 1

DW1734

caledonicus

NORFOLK 1

DW1737

caledonicus

NORFOLK 1

DW1736

caledonicus

NORFOLK 1

DW1735

caledonicus

NORFOLK 1

DW1739

caledonicus

NORFOLK 1

CP 1669

caledonicus

NORFOLK 1

CP1715

caledonicus

NORFOLK 1

DW1653

caledonicus

NORFOLK 1

CPI 711

caledonicus

NORFOLK 1

DW1712

caledonicus

NORFOLK 1

CP1671

caledonicus

NORFOLK 1

DW1652

caledonicus

LITHIST

CPI 7

caledonicus

LITHIST

CPI 5

caledonicus

LITHIST

DW13

caledonicus

LITHIST

CP 14

P. wareni

LITHIST

CP 16

caledonicus

TERRASSES

DW3127

caledonicus

TERRASSES

DW3053

caledonicus

TERRASSES

CP3049

caledonicus

TERRASSES

DW3122

caledonicus

TERRASSES

DW3123

caledonicus

TERRASSES

DW3124

caledonicus

TERRASSES

CP3048

caledonicus

MUSORSTOM 4

DW222

ACC.

386350

caledonicus

SMIB 2

DW05

caledonicus

BATHUS 3

DW817

caledonicus

BATHUS 2

DW729

caledonicus

SMIB 1

DW02

GENERAL AREA COORDINATES; DEPTH (in)

Grand Passage

18°55’S, 163°24’E; 370-405

Antigonia Bank

23°15’S, 168°04’E;311-355

Jumeau Est Bank

23°46’S, 168°17’E; 400-405

lie des Pins

22°52’S, 167°12’E; 408-410

lie des Pins

22°52’S, 167°13’E; 414-436

He des Pins

22°52’S, 167°13’E; 414-430

Norfolk Ridge

23°47’S, 168°17’E; 400-420

Norfolk Ridge

23°26’S, 167°50’E; 305-332

Norfolk Ridge

23°27’S, 167°50’E; 320-336

Norfolk Ridge

23°45’S, 168°16’E; 400-463

Norfolk Ridge

22°57’S, 167°16’E; 427-433

Norfolk Ridge

23°41’S, 168°15’E; 381-384

Norfolk Ridge

23°43’S, 168°16’E; 380-389

Norfolk Ridge

23°45’S, 168°17’E; 386-426

Norfolk Ridge

23°41’S, 168°15’E; 381-493

Norfolk Ridge

22°51’S, 167°12’E; 340-381

Norfolk Ridge

22°54’S, 167°13’E; 403-429

Norfolk Ridge

22°51’S, 167°10’E; 400

Norfolk Ridge

22°52’S, 167°12’E; 383-407

Norfolk Ridge

22°53’S, 167°13’E;415-445

Norfolk Ridge

22°51’S, 167°14’E; 404-448

Norfolk Ridge

23°41’S, 168 o 01’E; 302-325

Norfolk Ridge

23°22’S, 168°03’E; 270-312

Norfolk Ridge

23°26’S, 167°51 ’E; 328-340

Norfolk Ridge

23°48’S, 168°17’E; 409-439

Norfolk Ridge

23°22’S, 168°03’E; 180-250

Norfolk Ridge

23°42’S, 168 o 01’E; 320-397

Norfolk Ridge

23°27’S, 167 0 51’E; 290-378

Jumeau Ouest Bank

23°41’S, 168 o 01’E; 247-281

Jumeau Ouest Bank

23°40’S, 168°15’E; 389-404

Jumeau Est Bank

23°45’S, 168°17’E; 400

Jumeau Ouest Bank

23°42’S, 168° 16’E; 378-402

Jumeau Ouest Bank

23°43’S, 168°16’E; 379-391

He des Pins

23°00’S, 167°16’E; 400-420

Norfolk Ridge

23°45’S, 168°16’E; 410-440

Norfolk Ridge

23°42’S, 168°16’E; 380-410

He des Pins

22°47’S, 167°12’E; 390-410

He des Pins

22°53’S, 167°13’E; 420-450

lie des Pins

22°54’S, 167°15’E; 460

Norfolk Ridge

23°44’S, 168°16’E; 380-400

S Grande-Terre

22°58’S, 167°33’E; 410-440

S lie des Pins

22°56’S, 167°15’E; 398-410

Norfolk Ridge

23°42’S, 168°16’E; 405-410

SW Passe de Dumbea

22°52’S, 167°12’E; 400

S He des Pins

22°52’S, 167°13’E; 415

BOLD ID GENBANK #

19

European Journal of Taxonomy 134: 1-23 (2015)

MNHNID N SPECIES MNHN EXPEDITION STATION GENERAL AREA COORDINATES; DEPTH (m) BOLD ID GENBANK#

IM-2010-11190

3

P. caledoniciis

SMIB 1

DW07

S lie des Pins

22°55’S, 167°16’E; 500

IM-2010-11191

6

P. caledoniciis

SMIB 1

DW09

S lie des Pins

22°55’S, 167°15’E; 450

IM-2010-11192

1

P. caledoniciis

BIOCAL

CP42

S He des Pins

22°46’S, 167°14’E; 380

IM-2010-11193

1

P. caledoniciis

LAGON

DW1147

Belep

19°08’S, 163°30’E; 210

IM-2010-11194

1

P. caledoniciis

VAUBAN

DR15

22°49’S, 167°12’E; 390-395

IM-2010-11195

24

P. caledoniciis

SMIB 2

DW03

S He des Pins

22°54’S, 167°14’E; 412-428

IM-2010-11196

3

P. caledoniciis

SMIB 2

DW01

S He des Pins

22°53’S, 167°13’E; 438-444

IM-2010-11197

3

P. caledoniciis

SMIB 2

DW17

S He des Pins

22°56’S, 167°16’E; 428-448

IM-2010-11198

6

P. caledoniciis

SMIB 2

DW14

S He des Pins

22°53’S, 167°13’E; 405-444

I M-2010-11199

4

P. caledoniciis

SMIB 2

DW08

S He des Pins

22°53’S, 167°14’E; 435-447

IM-2010-11200

3

P. caledoniciis

SMIB 2

DW16

S He des Pins

22°52’S, 167°12’E; 390

IM-2010-11201

2

P. caledoniciis

SMIB 2

DW07

S He des Pins

22°56’S, 167°14’E; 428

IM-2010-11202

2

P. caledoniciis

SMIB 2

DW04

S He des Pins

22°53’S, 167°14’E; 410-417

IM-2010-11203

1

P. caledoniciis

SMIB 2

DW15

S He des Pins

22°53’S, 167°12’E; 375-402

IM-2010-11204

1

P. caledoniciis

SMIB 2

DW12

S He des Pins

22°51’S, 167°13’E; 445-460

IM-2010-11205

1

P. caledoniciis

SMIB 2

DW09

S He des Pins

22°55’S, 167°16’E; 475-500

IM-2010-11206

1

P. caledoniciis

SMIB 3

DW26

SW lie des Pins

22°55’S, 167°16’E; 450

IM-2010-11207

1

P. caledoniciis

SMIB 3

DW20

Jumeau Ouest Bank

23°40’S, 168°00’E; 280

IM-2010-11208

1

P. caledoniciis

SMIB 3

DW27

SW lie des Pins

22°55’S, 167°18’E; 457

IM-2010-11209

1

P. caledoniciis

SMIB 3

DW28

SW lie des Pins

22°46’S, 167°11’E; 394

IM-2010-11210

1

P. caledoniciis

SMIB 3

DW06

Eponge Bank

24°55’S, 168 0 21’E; 505

IM-2010-11211

1

P. caledoniciis

SMIB 3

DW25

SW lie des Pins

22°56’S, 167°15’E; 437

IM-2010-11212

2

P. caledoniciis

CHALCAL 2

DW82

Norfolk Ridge

23°14’S, 168°04’E; 304

IM-2010-11213

1

P. caledoniciis

MUSORSTOM 4

CP213

S Grande-Terre

22°51’S, 167°12’E; 405-430

IM-2010-11214

1

P. caledoniciis

MUSORSTOM 4

DW212

S Grande-Terre

22°47’S, 167°10’E; 375-380

IM-2010-11215

1

P. caledoniciis

MUSORSTOM 4

DW229

S Grande-Terre

22°52’S, 167°13’E; 445-460

IM-2010-11216

5

P. caledoniciis

MUSORSTOM 4

CP214

S Grande-Terre

22°54’S, 167°14’E;425-440

IM-2010-11217

6

P. caledoniciis

SMIB 4

DW65

He des Pins

22°56’S, 167°14’E; 400-420

IM-2010-11218

1

P. caledoniciis

SMIB 4

DW57

Antigonia Bank

23°21’S, 168°04’E; 210-260

IM-2010-11219

1

P. caledoniciis

VAUBAN

DR24

22°48’S, 167°09’E; 355-360

IM-2010-11220

6

P. caledoniciis

VAUBAN

DR15

22°49’S, 167°12’E; 390-395

IM-2010-11221

2

P. caledoniciis

444

18°15’S, 162°59’E; 300-350

IM-2010-11222

1

P. deforgesi

MUSORSTOM 5

DC361

Chesterfield

19°52’S, 158°38’E; 400

IM-2010-11223

3

P. deforgesi

CHALCAL1

DC32

Chesterfield

19°43’S, 158°33’E; 350

IM-2010-11224

1

P. deforgesi

CHALCAL1

CP08

Chesterfield

19°44’S, 158°35’E; 348

IM-2010-11225

1

P. deforgesi

MUSORSTOM 5

DW338

Chesterfield

19°52’S, 158°40’E; 540-580

IM-2010-11226

3

P. deforgesi

MUSORSTOM 5

DC379

Chesterfield

19°53’S, 158°39’E; 370-400

IM-2010-11227

5

P. deforgesi

MUSORSTOM 5

DC361

Chesterfield

19°52’S, 158°38’E; 400

IM-2010-11228

1

P. deforgesi

MUSORSTOM 5

DC361

Chesterfield

19°52’S, 158°38’E; 400

IM-2010-11229

1

P. deforgesi

MUSORSTOM 5

DW337

Chesterfield

19°54’S, 158°38’E; 412-430

IM-2010-11230

1

P. deforgesi

MUSORSTOM 5

DC361

Chesterfield

19°52’S, 158°38’E; 400

IM-2010-11231

1

P. deforgesi

MUSORSTOM 5

DC378

Chesterfield

19°54’S, 158°38’E; 355

IM-2010-11232

1

P. deforgesi

EBISCO

CP2579

N Bellona

20°21’S, 158°40’E; 440-455

IM-2010-11233

2

P. deforgesi

EBISCO

CP2596

Chesterfield

19°43’S, 158°37’E; 382-386

IM-2010-11234

1

P. deforgesi

EBISCO

DW2606

Chesterfield

19°37’S, 158°42’E; 442-443

IM-2010-11235

3

P. deforgesi

EBISCO

DW2564

NW Bellona

20°25’S, 158 0 41’E; 333-386

20

N

9

3

1

17

21

2

3

1

4

3

1

3

4

3

1

2

5

2

1

2

4

3

5

1

8

1

4

1

2

1

ANSEEUW P. et al.. New species of Pleurotomariidae

SPECIES MNHN EXPEDITION STATION GENERAL AREA COORDINATES; DEPTH (in) BOLD ID GENBANK#

P. deforgesi

EBISCO

CP2595

Chesterfield

19°44’S, 158°35’E; 345-377

P. deforgesi

EB1SC0

DW2577

N Bellona

20°20’S, 158°39’E; 399-602

P. deforgesi

EBISCO

DW2576

N Bellona

20°20’S, 158°43’E; 390-394

P. pseudogramdosus

EBISCO

DW2530

S Nova Bank

22°48’S, 159°23’E; 338-343

P. pseudogramdosus

EBISCO

CP2531

S Nova Bank

22°47’S, 159°23’E; 330-340

P. pseudogramdosus

EBISCO

CP2503

Capel Bank

24°48’S, 159°46’E; 366-380

P. pseudogranulosus

EBISCO

DW2533

N Nova Bank

22°18’S, 159°28’E; 360-370

P. pseudogramdosus

MUSORSTOM 5

DW304

Nova Bank

22°10’S, 159°25’E; 385-420

P. pseudogramdosus

MUSORSTOM 5

DW300

Nova Bank

22°48’S, 159°24’E; 450

P. pseudogramdosus

MUSORSTOM 5

DW301

Nova Bank

22°07’S, 159°25’E; 487-610

P. pseudogramdosus

MUSORSTOM 5

DW299

Nova Bank

22°48’S, 159°24’E; 360-390

P. wareni

MUSORSTOM 6

DW407

Loyalty Ridge

20°41’S, 167°07’E; 360

P. wareni

HALICAL 1

DW01

Grand Passage

18°56’S, 163°24’E; 380-400

P. wareni

HALICAL 1

DW04

Grand Passage

18°55’S, 163°24’E; 350-365

P. wareni

HALICAL 1

DW03

Grand Passage

18°53’S, 163°24’E; 350-380

P. wareni

MUSORSTOM 4

CC173

Grand Passage

19°02’S, 163°19’E; 250-290

P. wareni

MUSORSTOM 6

DC402

Loyalty Ridge

20°30’S, 166°49’E; 520

P. wareni

BERYX 11

CP51

Norfolk Ridge

23°45’S, 168°17’E; 390-400

P. wareni

TERRASSES

DW3101

Norfolk Ridge

22°59’S, 168 0 21’E; 320-390

P. wareni

TERRASSES

DW3110

Norfolk Ridge

23°02’S, 168°16’E; 270-310

P. wareni

NORFOLK 2

CP2095

Kaimon Maru Bank

24°46’S, 168°10’E; 283-310

P. wareni

SMIB 5

DW85

NW Walpole

22°20’S, 168°42’E; 240-260

P. wareni

LITHIST

CP14

Jumeau Ouest Bank

23°42’S, 168°16’E; 378-402

P. wareni

SMIB 8

DW160

Kaimon Maru Bank

24°47’S, 168°08’E; 280-282

P. wareni

CONCALIS

DW2968

Grand Passage

18°14’S, 163°02’E; 247-256

P. wareni

CONCALIS

CP3006

Grand Passage

18°32’S, 163°08’E; 400

P. wareni

CONCALIS

CP2975

Grand Passage

18°15’S, 163°06’E; 297-316

P. wareni

CONCALIS

DW2946

Grand Passage

19°02’S, 163°27’E; 276-277

P. wareni

BATHUS 4

DW924

Grand Passage

18°55’S, 163°24’E; 344-360

P. wareni

BATHUS 4

CP938

Grand Passage

19°00’S, 163°26’E; 280-288

P. wareni

BATHUS 4

DW925

Grand Passage

18°55’S, 163°24’E; 370-405

P. wareni

BATHUS 4

DW926

Grand Passage

18°57’S, 163°25’E; 325-330

P. wareni

BATHUS 4

DW931

Grand Passage

18°55’S, 163°24’E; 360-377

P. wareni

NORFOLK 1

DW1709

Norfolk Ridge

23°43’S, 168°16’E; 380-389

P. wareni

NORFOLK 1

DW1658

Norfolk Ridge

23°27’S, 167°50’E; 320-336

P. wareni

NORFOLK 1

DW1729

Norfolk Ridge

23°2rS, 168°16’E; 340-619

P. wareni

SMIB 8

DW189

Antigonia Bank

23°18’S, 168°06’E; 400-402

P. wareni

BATHUS 4

DW924

Grand Passage

18°55’S, 163°24’E; 344-360

P. wareni

MUSORSTOM 4

CP 193

Grand Passage

18°56’S, 163°23’E; 430

P. wareni

MUSORSTOM 4

DW196

Grand Passage

18°55’S, 163°24’E; 460

P. wareni

MUSORSTOM 4

DW222

S Grande-Terre

22°58’S, 167°33’E; 410-440

P. wareni

MUSORSTOM 6

DW407

Loyalty Ridge

20°4rS, 167°07’E; 360

P. wareni

MUSORSTOM 6

DW473

Loyalty Ridge

21°09’S, 167°55’E; 236

P. wareni

MUSORSTOM 4

250

P. wareni

SMIB 6

DW121

Grand Passage

18°58’S, 163°26’E; 315

P. wareni

BATHUS 3

CP812

Norfolk Ridge

23°43’S, 168°16’E; 391-440

21

European Journal of Taxonomy 134: 1-23 (2015)

MNHNID

N

SPECIES

MNHN EXPEDITION

STATION

GENERALAREA

COORDINATES; DEPTH (ir

IM-2010-11337

1

P. wareni

VOLSMAR

DW40

Loyalty Ridge

22°19’S, 168°41’E; 275-295

IM-2010-11340

6

P. wareni

CONCALIS

DW2977

Grand Passage

18°15’S, 163°07’E; 326-368

IM-2010-11341

6

P. wareni

CONCALIS

DW3025

Grand Passage

18°57’S, 163°23’E; 396-400

IM-2010-11342

6

P. wareni

CONCALIS

DW2978

Grand Passage

18°16’S, 163°04’E; 360-400

IM-2010-11343

4

P. wareni

CONCALIS

DW3024

Grand Passage

18°57’S, 163°22’E; 349-370

IM-2010-11344

2

P. wareni

CONCALIS

DW2943

Grand Passage

18°57’S, 163°23’E; 380-430

IM-2010-11345

1

P. wareni

CONCALIS

DW2983

Grand Passage

18°0rS, 163°02’E; 367-430

IM-2010-11346

1

P. wareni

CONCALIS

DW2951

Grand Passage

18°58’S, 163°25’E; 300

1M-2010-11347

1

P. wareni

CONCALIS

DW2945

Grand Passage

19°00’S, 163°26’E; 297-310

IM-2010-11348

1

P. wareni

CONCALIS

DW2944

Grand Passage

18°59’S, 163°24’E; 320-335

1M-2010-11349

11

P. wareni

CONCALIS

DW2947

Grand Passage

19°02’S, 163°26’E; 272-284

IM-2010-11350

14

P. wareni

CONCALIS

DW2974

Grand Passage

18°15’S, 163°06’E; 283-326

IM-2010-11351

25

P. wareni

CONCALIS

DW2979

Grand Passage

18°16’S, 162°54’E; 350

BOLD ID

GENBANK #

OUTGROUPS

Bayerotrochus africanus

AM049330.1

Mikadotrochns beyrichii

AM049331.1

Bayerotrochus midas

AY296820.1

P. amabUis

AY923929.1

Bayerotrochus midas

AY923930.1

Mikadotrochns beyrichii

EU530109.1

Entemnotrochus

L78910.1

adansoniamts

Entemnotrochus rumphii

L78911.1

Bayerotrochus teramachii

L78912.1

Bayerotrochus midas

L78913.1

P. maureri

L78914.1

P. quoyanus

L78915.1

P. lucaya

L78916.1

Haliotis tuberculata

JQ894846.1

22

ANSEEUW P. et al.. New species of Pleurotomariidae

Table 2. Shell characters in the Perotrochus caledonicus species complex. Measurements correspond to

mean values (with ranges) calculated on N specimens. Values for P. deforge si are taken from Metivier

(1990).

Character

Perotrochus caledonicus

Perotrochus wareni sp.

Perotrochus

nov.

pseudogranulosus sp. nov.

deforgesi

General appearance

trochoid with regularly

increasing weakly convex

whorls

conical, with weakly

convex to rather

straight-sided whorls

rather conical, with weakly

convex to straight-sided

whorls

depressed turbiniform

H/D ratio

1.00 (0.92-1.00, N = 7)

1.05 (1.01-1.09, N = 6)

0.87 (0.84-0.92, N = 9)

0.76 (N = 20)

Shell weight /

thickness

light, thin shell

very heavy, thick shell

light, thin shell

thick, heavy shell

Apical whorls, profile

acute sharp / straight conical

(with no whitish cord

protruding on surface)

more depressed, with

whitish cord protruding

near suture of apical

whorls

straight conical

depressed

Dominant teleoconch

sculpture

macroscopically generally

unbeaded to slightly beaded,

straight spiral cords

macroscopically finely

but distinctly beaded

spiral cords, variable

strength with crenulated

periphery

distinct finely light beaded

spiral cords, with angled

periphery

strongly beaded spiral

ribs above and under

selenizone, with

angled periphery

Teleoconch

fine diverging radiating

no diverging radiating

fine microsculpture

microsculpture

threads

microsculpture

radiating threads

Nacreous coverage

of inner slit lips at

aperture edge

nearly completely covered

by nacre (90%), small area

with porcellaneous layer

parallel to slit margins

partially covered by

nacre (75%), leaving

V-shaped area with

porcellaneous layer

uncovered (25%)

nearly completely covered

by nacre (85%), small area

with porcellaneous layer

parallel to slit margins

partially covered

by nacre (80%),

porcellaneous

layer parallel to slit

uncovered

Slit width

wide

very narrow

rather wide

long, 1 /6 th basal diameter

(1/5.7-1/6.6, N = 7)

shorter, 1/7* basal

long, about l/5 th basal

long, 1/5* basal

diameter (N = 7)

Slit length

diameter (1/6.5-1/7.6,

N = 6)

diameter (1/5.4-1/5.8,

N = 8)

Whorl surface

dull

metallic lustre

slightly lustruous

Checker-board colour

pattern in areas below

selenizone

not always clearly marked;

rather weak colour intensity,

not limited to the area

below selenizone

more irregularly

sized and spaced

colour blocks of less

contrasting intensity

very regularly spaced

checkerboard color

markings of deeper colour

intensity; overall deeper

colour of teleoconch

orange blotches of

variable intensity

in areas above

selenizone, not below

Surface of umbilical

callus pad on basal

disc

medium large

(34%-29.1-35.2,

N = 6), generally smooth

or weakly ridged radially in

large adults

very large (45% -

38.0-48.9, N = 6), ridged

radially, sharply edged

at margin

small (29% - 19.6-40.5,

N = 9), finely ridged

radially, with clearly raised

edge

large (36% - N = 7),

clearly ridged radially

and with raised edge

Basal disc profile

generally rather convex

flattened

rather flattened

23