European Journal of Taxonomy 129: 1-15

http://dx.doi.org/10.5852/ejt.2015.129

This work is licensed under a Creative Commons Attribution 3.0 License.

ISSN 2118-9773

www. europeanj ournaloftaxonomy. eu

2015 • Lorenz F. & Puillandre N.

Research article

urn:lsid:zoobank.org:pub:A76F290B-C259-4B2D-8068-AFB3B294BF12

Conus hughmorrisoni, a new species of cone snail from New Ireland,

Papua New Guinea (Gastropoda: Conidae)

Felix LORENZ 1 & Nicolas PUILLANDRE 2

1 Fr.-Ebert-Str 12, 35418 Buseck, Germany.

Email: [email protected]

2 Institut de Systematique, Evolution, Biodiversite ISYEB - UMR 7205 -

CNRS, MNHN, UPMC, EPHE, Museum national d’Histoire naturelle, Sorbonne Universites,

57 rue Cuvier, CP26, F-75005, Paris, France.

Email: [email protected] (corresponding author); tel: +33 140793173.

1 urn:lsid:zoobank.org:author:44070096-9354-4DF0-88ED-0D5D6CD2AB81

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

Abstract. Based on newly collected material from the Kavieng Lagoon Biodiversity Survey, we

describe a new species of cone snail. Conus hughmorrisoni sp. nov., from the vicinity of Kavieng, New

Ireland, Papua New Guinea. It closely resembles the New Caledonian C. exiguus and the Philippine

C. hanshassi , but differs from these species by having more numerous shoulder tubercles, by the shell’s

sculpturing and details of the color pattern. We also sequenced a fragment of the mitochondrial COI

gene of five specimens collected alive. All possessed very similar sequences (genetic distances < 0.3%),

different from all the COI sequences of cone snails available in GenBank (genetic distances > 10%).

Keywords. COI mitochondrial gene, Conidae, Conus hughmorrisoni sp. nov., new species. New Ireland.

Lorenz F. & Puillandre N. 2015. Conus hughmorrisoni, a new species of cone snail from New Ireland, Papua New

Guinea (Gastropoda: Conidae). European Journal of Taxonomy 129: 1-15. http://dx.doi.org/10.5852/eit.2015.129

Introduction

We are now standing on over 250 years of species descriptions, and apart from vertebrates and butterflies,

few animal groups have attracted as much attention from taxonomists as cone snails (Dance 1986). They

have always been prized by shell collectors and studied by amateur conchologists as well as malacologists

for their remarkable forms, colors, and biogeographical and ecological patterns. They are present in all

the tropical regions, sometimes abundant (and thus easy to collect) in shallow waters, and can be found

up to 700 m deep. Cone snails, as the other members of the superfamily Conoidea, are characterized by

a venom apparatus. The venom, comprising up to 200 different toxins (“conotoxins”), specific to each

species of cone snails (Dutertre et al. 2013; Violette et al. 2012), is injected in the prey (worms, molluscs

or fish) by a highly modified harpoon-shaped radular tooth. The relatively recent discovery of conotoxins

and their therapeutical applications is also fueling the enthusiasm of the scientific community for cone

snails, and the discovery of a new species is always a warranty to discover new toxins.

European Journal of Taxonomy 129: 1-15 (2015)

Given its relatively recent origin - 55 MY (Duda & Kohn 2005) - cone snails (i.e., Conidae) is one of

the most diverse group of marine invertebrates, with currently 820 species considered as valid (WoRMS,

1 Oct. 2014). New species are described regularly, and among the taxa Bacher analysed (Bacher 2012),

the number of species described per taxonomist has declined since about 1900 in all but one taxon: the

cone snails. In 2014 alone, 45 new species were described (WoRMS, 1 Oct. 2014). In the last 30 years,

an average of 11.5 new species have been described each year, totaling nearly 42% of all valid species

since 1758.

However, if new species descriptions are published often and regularly, they generally correspond to a

form that was known for decades by malacologists but never described as a new species; discovering

an unknown form of cone snail in shallow water is thus relatively unco mm on and remarkable. In this

article we describe a new species of cone snail, Conus ( Splinoconus ) hughmorrisoni sp. nov. (following

the classification of Puillandre et al. 2015), discovered during the Kavieng Lagoon Biodiversity Survey

(“KAVIENG 2014”) in Papua New Guinea (June 2014), during which numerous new species have been

discovered, some of them described already ( Ahyong 2014). On the third day of diving, a little Conus

Linnaeus, 1758 crawling across a piece of coral at 11 m was caught, and during the following weeks,

several further specimens were collected. Additional shells were found in the vicinity of where the species

was first discovered, confirming the consistency of the shell characteristics, different from all the known

species of cone snails. The sequencing of a fragment of the mitochondrial COI gene also confirmed that

the new species is different from the species for which a COI gene is available in public databases.

Material and methods

Part of the material was collected during the Kavieng Lagoon Biodiversity Survey in Papua New Guinea

(June 2014; Principal Investigators: Philippe Bouchet, Jeff Kinch), as part of the Our Planet Reviewed

expeditions. Some additional specimens come from private collections. Specimens collected alive were

microwaved (Galindo et al. 2014) to remove the body from the shell and a piece of foot tissue was

preserved in ethanol. A buccal complex was dissected to isolate the radular sac, which was then treated

with a solution of commercially available bleach until soft tissues were completely dissolved. The radula

was then rinsed in several shifts of distilled water, air dried and mounted for further SEM examination.

DNA was extracted using the Epmotion 5075 robot (Eppendorf), following the manufacturers’

recommendations. A fragment of the cytochrome oxidase subunit I (COI) was amplified using universal

primers LCO1490/HCO2198 (Folmer et al. 1994). PCR reactions were performed in 25 pi, containing

3 ng of DNA, IX reaction buffer, 2.5 m M MgC12, 0.26 m M dNTP, 0.3 m M of each primer, 5% DMSO,

and 1.5 units of Qbiogene Q-Bio Taq. Amplification consisted of an initial denaturation step at 94°C for

4 min, followed by 35 cycles of denaturation at 94°C for 30 s, annealing at 50°C for COI, followed by

extension at 72°C for 1 min. The final extension was at 72°C for 5 min. PCR products were purified and

sequenced by the Eurofins sequencing facility. Specimens are registered in the MNHN collections and

sequences were deposited in BOLD (Barcode of Life Datasystem) and GenBank (Table 1).

COI sequences of cone snails were downloaded from GenBank and aligned with the newly produced

COI sequences using using Muscle 3.8.31 (Edgar 2004). Given the size of the dataset (more than 1700

sequences), a first quick analysis was performed to reduce the dataset. A neighbor-joining (NJ) analysis

with Kimura-2-parameters (K2P) genetic distances was performed using MEGA 5 (Tamura et al. 2011)

to roughly identify the group in which the new species clustered. Closely related sequences in the NJ tree

(i.e., corresponding to the species belonging to the Floraconus, Leporiconus and Splinoconus clades, as

defined in Puillandre et al. 2014), together with sequences of more distant species (used as outgroups),

were then retained in the final dataset. Bathytoma neocaledonica Puillandre, Sysoev, Olivera, Couloux

& Bouchet, 2010 (Conoidea, Borsoniidae), was used as distant outgroup to root the tree. The final

dataset was analysed using a Bayesian approach as implemented in MrBayes 3.2 (Huelsenbeck et al.

2001), with two runs each consisting of three Markov chains of 10000000 generations, each with a

LORENZ F. & PUILLANDRE N., Conus hughmorrisoni , a new species of cone snail

Table 1 . List of specimens used in the final dataset for phylogenetic analyses.

Species

MNHN voucher number

Status

BOLD ID

GenBank

Accession Number

Conus hughmorrisoni sp. nov.

MNHN IM-2013-53462

Holotype

C0N01794-15

KR070759

Conus hughmorrisoni sp. nov.

MNHN IM-2013-53463

Paratype 2

C0N01795-15

KR070758

Conus hughmorrisoni sp. nov.

MNHN IM-2013-53464

Paratype 3

C0N01796-15

KR070757

Conus hughmorrisoni sp. nov.

MNHN IM-2013-53771

Paratype 1

C0N01793-15

KR070760

Conus hughmorrisoni sp. nov.

MNHN IM-2013-53772

Paratype 4

C0N01792-15

KR070761

Conus balteatus G. B. Sowerby I, 1833

KJ550134

Conus biliosus (Roding, 1798)

KJ549870

Conus boeticus Reeve, 1844

KJ550139

Conus capitanellus Fulton, 1938

KJ549891

Conus coffeae Gmelin, 1791

KJ550178

Conus corallinus Kiener, 1847

KJ550163

Conus distans Hwass in Bruguiere, 1792

KJ550205

Conus fumigatus Hwass in Bruguiere, 1792

KJ550268

Conus glans Hwass in Bruguiere, 1792

KJ549918

Conus granum Rockel & Fischoder, 1985

KJ549922

Conus hamamotoi Yoshiba & Koyama, 1984

KJ550283

Conus Ienavati da Motta & Rockel, 1982

KJ550324

Conus luteus G. B. Sowerby I, 1833

KJ549942

Conus mozambicus Hwass in Bruguiere, 1792

KJ550075

Conus queenslandis da Motta, 1984

KJ550432

Conus roseorapum G. Raybaudi & da Motta, 1990

KJ550544

Conus sazanka Shikama, 1970

KJ550444

Conus shikamai Coomans, Moolenbeek & Wils, 1985

KJ549989

Conus sp.

KJ550327

Conus tenuistriatus G. B. Sowerby 11, 1858

KJ550476

Conus tinianus Hwass in Bruguiere, 1792

KJ550002

Conus tribblei Walls, 1977

KJ550510

Conus viola Cq mohorsky, 1977

KJ550010

Conus voluminalis Reeve, 1843

KJ550525

Conasprella pagoda (Kiener, 1847)

EU015729

Californiconus californicus (Reeve, 1844)

KJ549878

Profundiconus vaubani (Rockel & Moolenbeek, 1995)

KJ550518

Bathytoma neocaledonica Puillandre, Sysoev, Olivera, Couloux & Bouchet, 2010

EU015653

sampling frequency of one tree each 500 generations. Each codon position of the COI gene was treated

as an unlinked partition, each following a 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 1.4.1 (Rambaut & Drummond 2007) and analyses were terminated

when ESS values were all greater than 200. A consensus tree was then calculated after omitting the first

b25% trees as burn-in.

European Journal of Taxonomy 129: 1-15 (2015)

Abbreviations

FL = Collection Felix Lorenz, Germany

MNHN = Museum national d’Histoire naturelle, Paris

PNG = Papua New Guinea

Results

Five specimens of Conus hughmorrisoni sp. nov. were successfully sequenced for a 658 bp fragment

of the COI gene. The K2P genetic distances among the five specimens are very low (0.2-0.3%) and

correspond to genetic distances generally considered as intraspecific distances in cone snails; conversely

all the genetic distances with other known cone snail species are large (> 10%) and correspond to

genetic distances generally considered as interspecific distances in cone snails (e.g. Duda et al. 2008;

Puillandre et al. 2011). The preliminary analysis using the NJ method suggests that the new species

belongs to the Splinoconus clade, as defined in Puillandre et al. (2014). Consequently, the dataset was

then limited to species belonging to this clade, together with additional species from closely related

clades ( Leporiconus , Floraconus ) and more distantly related cone snails. In the resulting tree (Fig. 1),

the five specimens of Conus hughmorrisoni sp. nov. are grouped in a highly supported clade (Posterior

Probability = 1). The species clusters in the Splinoconus clade (Posterior Probability = 1). The new

species is described below.

Bathytoma neocaledonica

-- Profundiconus vaubani

- Californicomis californicus

- Conasprella pagoda

0.98

0.96

0.95

— Conus distans

— Conus glans

— Conus granum

i- Conus pomareae

Conus coffeae

Conus tenuistriatus

96r

- Conus mozambicus

Conus balteatus

- Conus tinianus

— Conus hamamotoi

0.98

0.99

0.95

- Conus capitanellus

1 1 — Conus corallinus

I— Conus sazanka

- Conus fumigatus

0.2 subst./site

— Conus roseorapum

- Conus lenavati

1 r Conus queenslandis

I— Conus tribblei

- Conus viola

- Conus biliosus

1 |

Conus shikamai

- Conus boeticus

Conus voluminalis

- Conus sp.

IM-2013-5 3462

IM-2013-53464

IM-2013-5 3463

IM-2013-53771

IM-2013-53772

Leporiconus

Floraconus

Splinoconus

Conus

hughmorrisoni

n. sp.

Fig. 1. Bayesian phylogenetic tree obtained with the COI gene. Posterior Probabilities (> 0.9) are shown

above nodes).

LORENZ F. & PUILLANDRE N., Conus hughmorrisoni , a new species of cone snail

Class Gastropoda Cuvier, 1795

Subclass Caenogastropoda Cox, 1960

Order Neogastropoda Wenz, 1938

Superfamily Conoidea Fleming, 1822

Family Conidae Fleming, 1822

Genus Conus Linnaeus, 1758

Conus (Splinoconus) hughmorrisoni sp. nov.

um:lsid:zoobank.org:act:4105F23A-B87D-4D62-842A-476B9E889C24

Table 1; Figs 2-A

Etymology

This new species is named in honour of Hugh Morrison from Perth, Western Australia. He is a well

known malacologist, shell dealer and pioneer scuba diver. He led the team of divers on board the MV

PNG Explorer during the Kavieng Lagoon Biodiversity Survey. He is among the leading experts in

Australian shells, and a dear friend of the first author.

Type material examined

Holotype and paratypes 1-5 are deposited in the MNHN.

Holotype

PAPUA NEW GUINEA: MNHN-IM-2013-53462, 20.4 mm. New Ireland, Kavieng Lagoon, E side of

Wadei Island, 02°40.3’S, 150°39.UE, 9-11 m deep (stationKR70), Fig. 2A, BOLD ID C0N01794-15,

GenBank accession number (COI sequence) KR070759.

Paratypes

PAPUA NEW GUINEA: paratype 1, MNHN-IM-2013-53771, 13.25 mm, New Ireland, Kavieng

Lagoon, NW point of Ungan Island, 02°38.8’S, 150°39.7’E, 3-12 m deep (station KR80), Fig. 2B-

C, BOLD ID C0N01793-15, GenBank accession number (COI sequence): KR070760; paratype 2,

MNHN-IM-2013-53463, 12.55 mm, New Ireland, Kavieng Lagoon, E side of Wadei Island, 02°40.3’S,

150°39.EE, 9-11 m deep (station KR70), Figs 2D, 3A-B, BOLD ID C0N01795-15, GenBank

accession number (COI sequence) KR070758; paratype 3, MNHN-IM-2013-53464, 16.5 mm. New

Ireland, Kavieng Lagoon, E side of Wadei Island, 02°40.3’S, 150 o 39.UE, 9-11 m deep (station KR70),

Figs 2E, 3C-F, BOLD ID C0N01796-15, GenBank accession number (COI sequence) KR070757;

paratype 4, MNHN-IM-2013-53772, 10.8 mm. New Ireland, Kavieng Lagoon, NW point of Ungan

Island, 02°38.8’S, \59°29TE, 3-12 m deep (station KR80), Fig. 2F-G, BOLD ID C0N01792-

15, GenBank accession number (COI sequence) KR070761; paratype 5, MNHN-IM-2000-27955,

14.15 mm, New Ireland, Kavieng Lagoon, NW point of Ungan Island, 02°38.8’S, 150°39.7 , E, 3-12 m

deep (station KR70), Fig. 2H; paratype 6, 21.6 mm, Nusaum Is., New Ireland, FL, Fig. 4A; paratype 7,

19.0 mm, Ungan Is., New Ireland, FL, Fig. 4B; paratype 8, 19.1 mm, Ungan Is., New Ireland, FL, Fig.

4C; paratype 9, 19.6 mm, Wadei Is., New Ireland, FL, Fig. 4D; paratype 10, 18.3 mm, Nusaum Is., New

Ireland, FL, Fig. 4E; paratype 11, 18.1 mm, Nusaum Is., New Ireland, Coll. Hugh Morrison, Fig. 4F.

Type locality

Papua New Guinea, New Ireland, Kavieng Lagoon, E side of Wadei Island, 02°40.3’S, 150°39.UE,

9-11 m deep (station KR70).

Description

The shell of the holotype is rather small and lightweight. The last whorl is moderately broad and conical.

The aperture is equally narrow throughout. The spire is pointed, acutely stepped, the outline very slightly

European Journal of Taxonomy 129: 1-15 (2015)

Fig. 2. Conushughmorrisoni sp. nov. A. Holotype, 20.4mm. B-C. Paratype 1,13.25 mm. B. Teleoconch.

C. Protoconch (scale bar = 0.5 mm). D. Paratype 2, 12.55 mm. E. Paratype 3, 16.5 mm. F-G. Paratype 4,

10.8 mm F. Teleoconch. G. Top view. H. Paratype 5, 14.15 mm. All pictures by Manuel Caballer

Gutierrez (credits project E-Recolnat, MNHN).

LORENZ F. & PUILLANDRE N., Conus hughmorrisoni, a new species of cone snail

Fig. 3. Conus hughmorrisoni sp. nov. A-B. Paratype 2, 12.55 mm. C. Operculum of paratype 5.

D-F. Radula of paratype 3.

European Journal of Taxonomy 129: 1-15 (2015)

Fig. 4. Conus hughmorrisoni sp. nov. A. Paratype 6, 20.8 mm B. Paratype 7, 21.6 mm C. Paratype 8,

19.0 inm D. Paratype 9, 19.1 mm E. Paratype 10, 19.6 mm F. Paratype 11, 18.3 mm

LORENZ F. & PUILLANDRE N., Conus hughmorrisoni , a new species of cone snail

concave. The suture is narrow and shallow. The protoconch is smooth, bulbous, of about 2 whorls, measuring

about 0.55 mm in width. It is missing or strongly corroded in all specimens studied (Fig. 2C). The first

two postnuclear whorls have no spiral grooves, and their shoulders are shallow and without discernible

tubercles. There are three shallow incised striae on the postnuclear sutural ramps, and two to three broader,

less distinct striae along the angle of the shoulder and below on the adapical end of the last whorl. The last

adult whorl and the preceeding whorl show 15-17 prominent tubercles each (Fig. 2G); their interstices are

deeply indented, also below the shoulder, dorsally on the last adult whorl, forming an undulating outline.

The number of tubercles increases by one or two in the preceeding earlier whorls, gradually weakening,

finally disappearing towards the protoconch. The shell seems glossy and smooth in the half below the spire,

but there are regularly spaced, shallow, axially striate spiral grooves between spiral ribbons on its entire

surface. These are enhanced by intermittent tubercles which gradually become more prominent and denser

towards the abapical end, and directly above the lip. The anal notch is rounded and rather shallow.

The protoconch is pale yellow. The ground colour of the teleoconch is white, with purple on parts of the

last whorl. The abapical end is stained with darker purple. The spire is white with occasionaly discrete

brown axial lines. The last adult whorl shows irregular brown patches above a pale purple mid-dorsal

area without darker pattern; these patches can eventually be connected to form an irregular spiral band.

There are numerous, evenly spaced narrow spiral rows of white and brown dots. They appear as rows

of white spots within the darker blotches and as discrete red-brown spots in the paler, unblotched areas.

The tubercles on the spiral ribbons are white, which is especially obvious in the darker stained abapical

end. The interior of the shell is purplish brown.

The paratypes agree with the holotype in all morphological aspects and show basically no variation

in shape or the development of the spiral tubercles. In some specimens, the spiral grooves are less

developed adapically. The coloration, however, varies considerably: in some specimens, the last adult

whorl is rather rich purple and the darker stain of the abapical end is less obvious. The spire is generally

white; darker dashes and lines are always discrete. The color of the darker dorsal blotches varies from

orange to black. In most shells it is purplish brown, green in one specimen. The spiral rows of white and

brown dots are reduced in the paler specimens. The darker blotches above the paler mid-dorsal zone

can be fused to a compact dark band with irregular outline; in other specimens there are irregular axial

flames in which the borders seem fringed by the white component of the axial lines.

The periostracum is reddish brown and thin. The animal has a brown foot with discrete yellow dashes,

the crawling surface is pale brown (Fig. 3A-B). The edge of the transparent orange siphon is framed

with black. The operculum is illustrated on Fig. 3C. The radular tooth is of the vermivorous kind, rather

slender, and of small relative size (L/TL = 72) (Fig. 3C-F). The anterior portion is much shorter than

the posterior section (TL/APL = 2.7-2.8). Waist evident. Apical barb present, opposing a rounded blade

which covers most of the anterior portion of the tooth (100 BL/APL = 84 %). There are no denticles

present in serration. Instead, there are 6-7 raised irregular marginal undulations arranged in one row,

ending in a terminating cusp. These structures seem to correspond to precursors of denticles in a primitive

serration (protoserration). Base axially elongated, with a small basal spur present, pointing upwards.

Measurements of the shells are provided in Table 2 (abbreviations: L = shell length; TL = radular tooth

length; APL = anterior portion length; BL = blade length).

Distribution and habitat

Conus hughmorrisoni sp. nov. is so far known only from a small area between Kavieng and New

Hannover Island to the west of New Ireland, Papua New Guinea: east side of Wadei Island, 02°40.3’S,

150°39.EE (Station KR70), at 9-11 m, on sand and rubble; NW point of Ungan Island, 02°38.8’S,

150°39.7’E (Station KR80), at 3-12 m, on flat sand and rubble slope with larger corals; S side of

Nusaum Island, 02°38.38ES, 150°38.436’E, at 5-24 m, active coral reef 8-15 m, coarse rubble slope

European Journal of Taxonomy 129: 1-15 (2015)

Table 2. Measurements (in mm) of shells of Conus hughmorrisoni sp. nov., Conus exiguus Lamarck,

1810 (taken from Rocket et al. 1995) and Conus hanshassi (Lorenz & Barbier, 2012). L = shell length;

MD = maximum diameter; AH = aperture height; HMD = height of maximum diameter; RD = relative

diameter of last whorl (MD/AH); PMD = position of maximum diameter of last whorl (HMD/AH);

RSH = relative spire height ((L-AH)/L).

AH HMD

PMD

RSH

C. hughmorrisoni sp. nov. MNHN-IM-2013-53462 (Holotype)

20.4

10.2

16.4

14.8

0.62

0.90

0.20

C. hughmorrisoni sp. nov. MNHN-IM-2013-53771 (Paratype 1)

13.3

7.2

10.6

9.9

0.68

0.93

0.21

C. hughmorrisoni sp. nov. MNHN-IM-2013-53463 (Paratype 2)

12.6

6.4

8.7

0.64

0.87

0.21

C. hughmorrisoni sp. nov. MNHN-IM-2013-53464 (Paratype 3)

16.5

8.3

12.9

11.4

0.64

0.88

0.22

C. hughmorrisoni sp. nov. MNHN-IM-2013-53772 (Paratype 4)

10.8

5.7

8.7

7.6

0.65

0.87

0.19

C. hughmorrisoni sp. nov. MNHN-IM-2000-27955 (Paratype 5)

14.2

7.4

10.6

0.62

0.88

0.16

Mean values for C. hughmorrisoni

0.65

0.89

0.20

C. exiguus Lamarck, 1810

16-54

0.57-0.67

0.83-0.95

0.09-0.20

C. hanshassi (Lorenz & Barbier, 2012) MNHN-IM-2000-24814

(Holotype)

23.4

10.9

17.4

15.4

0.63

0.89

0.26

Fig. 5. Map of the Kavieng Region showing the different sampling sites. Black circles: stations with

sequenced material; grey circles: stations with other material.

LORENZ F. & PUILLANDRE N., Conus hughmorrisoni , a new species of cone snail

to gently sloping bottom at 21 m; NW side of Ral Island, 02°36.373’S, 150 o 38.518’E, at 4-25 m, slope

with coral conglomerate (Fig. 5). Further specimens have been collected in a similar habitat and depth

in the vicinity of the islands of Wadei, Ungan and Nusaum.

Remarks

There are two species of Conidae which should be compared to Conus hughmorrisoni sp. nov.:

Conus (. Phasmoconus) exiguus Lamarck, 1810, a highly variable species known only from New

Caledonia (Fig. 6A-D). Some of its formae can be similar to C. hughmorrisoni sp. nov. in shape, the

tuberculate spire, and by having rows of spiral tubercles. Also, the general color pattern can be quite

similar. However, the number of tubercles on the shoulder differs. In C. exiguus and its variations (e.g.

the smaller form Conus cabritii Bernardi, 1858), there are 12 to 14 tubercles associated with the last

adult whorl, in C. hughmorrisoni sp. nov. there are 15 to 17. The suture of C. exiguus is deeper and

the incised sutural striae are more distinct, especially on the early postnuclear whorls. In C. exiguus ,

the pattern of the last adult whorl is continued on the spire, which hence is often dark blotched. The

spire of C. hughmorrisoni sp. nov. is usually untinted or shows only discrete spots or lines. The darker

patches of C. hughmorrisoni sp. nov. are crossed by the narrow spiral bands of white and dark brown

intermittent spots that can be replaced, in some cases, by rows of irregular white dashes. The resulting

effect caused by this banding, minute white specks within the darker blotches, and fine brown spots in

the paler areas of the dorsum, give the pattern of C. hughmorrisoni sp. nov. a more delicate look.

Conus (Stratego conus) hanshassi (Lorenz & Barbier, 2012), from Siargao Island in the Philipines, is

so far known from only three specimens (Fig. 6E-F). It is somewhat narrower, with a taller spire. Like

in C. exiguus , the number of tubercles (12-13) along the shoulder of the last adult whorl is lower than

in C. hughmorrisoni sp. nov. The color pattern is compact, sparser and extends onto the spire in C.

hanshassi , in which the narrow spiral bands of white and darker spots of C. hughmorrisoni sp. nov.

are replaced by numerous fine brown spots arranged in spiral lines across the last adult whorl. As a

consequence, the darker blotches of C. hanshassi do not show white specks as in C. hughmorrisoni

sp. nov.

The ratios, based on measurements of the shells (Table 2), do not show any differences between

C. hughmorrisoni sp. nov., C. exiguus and C. hanshassi , except for the RSH ratio of C. hanshassi , which

is slighlty higher than for both the other species, albeit based on only one specimen of C. hanshassi.

Discussion

The species Conus hughmorrisoni sp. nov. is molecularly different from all the species of Conus

available in GenBank, and in this regard molecular data does not contradict the fact that it corresponds

to a new species. However, the species morphologically resembles two other cone snails: Conus

exiguus and Conus hanshassi. The radula is known for one of them, Conus exiguus (Tucker & Tenorio

2009); it suggests that C. exiguus and C. hughmorrisoni sp. nov. are related, but different species (M.

Tenorio pers. com.). The tooth of C. exiguus is broader than that of C. hughmorrisoni sp. nov., and does

not show with clarity (under the optical microscope) any protoserration. The base of the tooth of C.

exiguus is more rounded than in C. hughmorrisoni sp. nov. in which it is axially elongated. Concerning

Conus hanshassi , the radula is unknown but M. Tenorio (pers. com.) considers that it could be a

Rolaniconus Tucker & Tenorio, 2009, a genus-level taxon synonymized with Str ate go conus da Motta,

1991 by Puillandre et al. (2015). Although it remains to be confirmed by a molecular analysis, both

the characters of the shell (for C. exiguus and C. hanshassi) and of the radular tooth (for C. exiguus)

suggest that C. hughmorrisoni sp. nov., C. exiguus and C. hanshassi are three different species.

European Journal of Taxonomy 129 : 1-15 ( 2015 )

Fig. 6. A Conus exiguus var. optimus Sowerby, 1913, 35.3 mm. Point Parme, New Caledonia. FL.

B. Conus exiguus var. bougei Sowerby, 1907,20.3 mm. Poum, N. New Caledonia. FL. C. Conus exiguus

var. cabritii Bernardi, 1858, 22.0 mm. Northern New Caledonia. FL. D. Conus sp. cf. exiguus, 18.0 mm.

Apia, Western Samoa. From Rockel et al. (1995), pi. 72, figs 14-15. E-F. Conus hanshassi (Lorenz &

Barbier, 2012). E. 23.4 mm Siargao Is., Philippines. Holotype, MNHN-IM-2000-24814. F. 22.9 mm.

Siargao Is., Philippines. Paratype 1, FL.

LORENZ F. & PUILLANDRE N., Conus hughmorrisoni, a new species of cone snail

Geographically, C. hughmorrisoni sp. nov. connects the distribution areas of C. hanshassi and C. exiguus.

These three species seem to represent close allies along a line that marks the distribution of several other

species-complexes of the famliy (e.g. australis, coccineus, corallinus, monachus, mucronatus,proximus,

and their closer relatives). The continuation of this line eastwards is suggested by an undescribed species

illustrated in Rockel et al. (1995, pi. 72, figs 14-15) from Western Samoa. This shell has a rather flat

spire with few, prominent tubercles and a heavily tuberculate last adult whorl. Its color pattern resembles

that of C. exiguus. We have little doubt that additional related species will be discovered along the line

between the Philippines, Papua New Guinea, New Caledonia and Samoa. The paucispiral protoconch of

C. hughmorrisoni sp. nov. and C. hanshassi suggests intracapsular development or at least an abbreviated

planctonic stage that limits the dispersal of larvae over greater distances (Jablonski & Lutz 1980). The

same seems to be true for C. exiguus from New Caledonia, whose radiation into numerous local and

bathymetric forms (and eventually sibling species, see Rockel et al. 1995) may be the result of such

isolation based on limited dispersal.

Acknowledgements

Part of the material was collected during the Kavieng Lagoon Biodiversity Survey in Papua New

Guinea (June 2014; Principal Investigators: Philippe Bouchet, Jeff Kinch), as part of the Our Planet

Reviewed expeditions organized jointly by Museum national d'Histoire naturelle (MNHN), Pro-Natura

International (PNI) and Institut de Recherche pour le Developpement (IRD), with support from Papua

New Guinea’s National Fisheries Authority. The organizers acknowledge supporting funding from the

Total Foundation, the Laboratoire d’Excellence Diversites Biologiques et Culturelles (LabEx BCDiv,

ANR-10-LABX-0003-BCDiv), the Programme Investissement d’Avenir (ANR-ll-IDEX-0004-02), the

Fonds Pacifique, and CNRS’ Institut Ecologie et Environnement (INEE). The expedition was endorsed

by the New Ireland Provincial Administration and operated under a Memorandum of Understanding

with the University of Papua New Guinea (UPNG). This project was partly supported by the Service de

Systematique Moleculaire (UMS 2700 CNRS-MNHN) and by the project CONOTAX, funded by the

French ANR (grant number ANR-13-JSV7-0013-01). The authors also thank Virginie Heros, Barbara

Buge, Julien Brisset, Manuel Caballer Gutierrez, Laetitia Aznar-Cormano and Gilberto Marani for

their help in curating the specimens, Alexander Fedosov and Manolo Tenorio for radula preparation

and analysis, Dietmar and Ange Amon (Lissenung Island, Kavieng), Dr. Marco Chiapponi (Chiapponi

Lorenz Seashell Foundation, Lecco, Italy), Dr. Michael A. Mont (Molluscan Science Foudation, Owings

Mills, Maryland, USA), Jean Pierre Barbier, Jana Kratzsch, and Alan J. Kohn and two anonymous

referees for their co mm ents on a previous version of the manuscript.

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