Profundiconus puillandrei, Tenorio & Castelin, 2016

Tenorio, Manuel J. & Castelin, Magalie, 2016, Genus Profundiconus Kuroda, 1956 (Gastropoda, Conoidea): Morphological and molecular studies, with the description of five new species from the Solomon Islands and New Caledonia, European Journal of Taxonomy 173, pp. 1-45 : 28-33

Profundiconus puillandrei sp. nov.

urn:lsid:zoobank.org:act:55807BF8-D984-42C1-B039-F3EBD8EB2C13

Figs 12A–J , 13A–D , 14

Conus ikedai – Poppe 2008: pl. 615, fig. 1a–b (non C. ikedai Ninomiya, 1987 ).

Conus smirna – Marshall 1981: 499, fig. 3j (non C. smirna Bartsch & Rehder, 1943 ).

Conus sp. C – Röckel et al. 1995b: 585, fig. 49.

Profundiconus n. sp. g . – Puillandre et al. 2014: Supplementary Material 1 (unfigured).

Etymology

This new species is dedicated to Dr. Nicolas Puillandre, mollusc curator at the MNHN. Dr. Puillandre has a long and outstanding trajectory in the study of the phylogeny of the Conoidea . The naming of the new species after him recognises his important contributions to the taxonomy of Conoidea at the molecular level.

Type material examined

Holotype

NEW CALEDONIA: 43.2 × 18.0 mm, R/V Alis, NORFOLK 1 Expedition, st. DW 1707, Banc Jumeau Est, Norfolk Ridge , 23°43' S, 168°16' E, 381–493 m ( MNHN IM-2000-30771 ; Fig. 12 A ).

Paratypes

NEW CALEDONIA: 39.6 × 17.3 mm, R/V Alis, NORFOLK 2 Expedition, st. DW 2072, Banc Aramis, Norfolk Ridge , 25°21' S, 168°57' E, 1000–1006 m ( MNHN IM-2000-30772 ; paratype 1; Fig. 12B ); 40.7 × 16.8 mm, R/V Alis, NORFOLK 2 Expedtion , st. DW 2077, Banc Zorro , Norfolk Ridge , 25°21' S, 168°19' E, 666–1000 m ( MNHN IM-2000-30773 ; paratype 2; Fig. 12 C ) ; 35.0 × 15.5 mm, R/V Alis, NORFOLK 2 Expedition, st. DW 2068, Banc Porthos , Norfolk Ridge , 25°20' S, 168°57' E, 680–980 m ( MNHN IM-2000-30774 ; paratype 3; Fig. 12D ) ; 31.7 × 13.2 mm, R/V Alis, NORFOLK 2 Expedition, st. DW 2066, Banc Athos , Norfolk Ridge , 25°17' S, 168°55' E, 834–870 m ( MNHN IM-2000- 30775 ; paratype 4; Fig. 12E, H ) ; 43.6 × 18.1 mm, R/V Alis, NORFOLK 2 Expedition, st. DW 2074, Banc Zorro , Norfolk Ridge , 25°24' S, 168°20' E, 623–691 m ( MNHN IM-2000-30776 ; paratype 5; Fig. 12F ) ; 57.3 × 24.6 mm, R/V Alis, EXBODI Expedition, st. DW 3907, Récifs de l’Astrolabe-Nord Ouest , 19°50' S, 165°33' E, 608–671 m ( MNHN IM-2009-31320 ; paratype 6; Fig. 12G ; GenBank accession number (cox1 sequence): KT874752 ) ; 45.2 × 18.5 mm, R/V Alis, NORFOLK 2 Expedition, st. DW 2054, Banc Jumeau Est , Norfolk Ridge , 23°40' S, 168°15' E, 736–800 m ( MNHN IM-2000- 30778 ; paratype 7; Fig. 12I ) ; 44.8 × 18.2 mm, R/V Alis, BATHUS 3 Expedition, st. DW 776, Loyalty Ridge , 24°44' S, 170°08' E, 770–830 m ( MNHN IM-2000-30779 ; paratype 10) ; 49.9 × 21.6 mm, R/V Alis, TERRASSES Expedition, st. DW 3045, Mont J, Ride des Loyautés , 23°48' S, 169°46' E, 660– 710 m ( MNHN IM-2009-18221 ; paratype 11; GenBank accession number (cox1 sequence): KJ550484 ) ; 37.0 × 17.2 mm, R/V Alis, BATHUS 3 Expedition, st. DW 776, Loyalty Ridge , 24°44' S, 170°08' E, 770–830 m ( MNCN 15.05/60171 ; paratype 12) .

NEW ZEALAND: 38.9 × 17.5 mm, R/V Tangaroa , st. K861, Kermadec Ridge, 30°36.5' S, 178°22.5' W, 1030 m ( NIWA 99587; paratype 8; Fig. 12J ); 44.9 × 19.5 mm, R/V Tangaroa , st. K831, Kermadec Ridge, 29°51.5' S, 178°10.5' W, 965 m ( NIWA 99588; paratype 9).

Type locality

NEW CALEDONIA: Banc Jumeau Est , Norfolk Ridge, 381–493 m, 23°43' S, 168°16' E (NORFOLK 1 st. DW1707).

Other material examined

NEW CALEDONIA: 33.2 × 14.8 mm, R/V “Alis”, NORFOLK 2 Expedition, st. DW 2057, Banc Introuvable, Norfolk Ridge, 24°40' S, 168°39' E, 555–565 m ( MNHN IM- 2007-34865; GenBank accession number (cox1 sequence): KJ550262 ). Note: This specimen, matching the holotype, was photographed (http://coldb.mnhn.fr/catalognumber/mnhn/im/2007-34865) and sequenced, but the shell was destroyed in the process and is no longer available.

Additionally, we examined 31 more specimens from 15 uncataloged MNHN lots collected at several stations in Norfolk Ridge and Loyalty Ridge, New Caledonia, by the R/V “Alis” in the course of several campaigns. Several specimens of shells in private collections collected in Balut Is., Philippines at 100– 150 m, showing the conchological features of Profundiconus puillandrei sp. nov., were also examined.

Description

Morphometric parameters: S L = 29–57 mm; RD = 0.53–0.62; RSH = 0.22–0.29; PMD = 0.81–0.90.

Shell moderately small to medium sized (maximum length 57.0 mm). Shell profile ventricosely conical, with high spire. Spire profile sigmoid to slightly concave. Protoconch multispiral, with 3–3.5 whorls, white to yellow-brown. Last whorl of larval shell shows minute axial ridges. Early teleoconch whorls with nodules which are often indistinct after whorls 5 to 6, but may persist, forming nodulose ridge reaching shoulder on last whorl. Sutural ramp flat to slightly concave, with very fine striae and arcuate threads becoming obsolete in late whorls. Shoulder with distinct ridge, usually smooth, although nodulose or even strongly nodulose in some specimens. Last whorl with convex sides adapically, then almost straight and slightly concave abapically. Last whorl smooth or with very fine striae becoming more evident towards base. Spire and last whorl patternless, white to pale straw-yellow in colour. Columella white. Aperture pale yellow or white. Periostracum yellow, thin and translucent. Operculum with serrations.

Radular tooth examined in holotype ( Fig. 13A ), in paratypes 3 ( Fig. 13B–D ), 6 and 9, and in an additional non-type specimen. 48 to 62 teeth in radular sac. Radular tooth small for size of shell: its total length relative to shell length S L /T L = 75–105. Anterior portion shorter than posterior section of tooth (T L/AP L = 3.1–3.6). With one barb and pointed blade which covers 50–62% of anterior portion of tooth. External cusp present, extending between 64 and 90% of length of anterior portion of tooth. External cusp laterally widened and serrated, with 4–5 small denticles. Large adapical opening occupying most of anterior portion of tooth (100AO L/AP L = 64–75). Fringe of closely spaced projections pointing towards apex immediately below waist. Shaft fold present. Large and prominent basal spur on top of slanted base of tooth.

Distribution and habitat

New Caledonia (Norfolk Ridge and Loyalty Ridge) and New Zealand (Kermadec Ridge), at depths from 380 to 1100 m ( Fig. 14 ). Several empty shells matching P. puillandrei sp. nov. from Balut Is., Mindanao, Philippines, have been examined. The identity of these specimens from the Philippines (allocated in several private collections) could not be confirmed by radular or molecular studies, but the conchological features seem consistent with the identification of these specimens (often labelled as Conus cf. ikedai , or Conus darkini “albinistic”) as P. puillandrei sp. nov. This is a feasible possibility given the multispiral protoconch of this species (suggesting a planktonic larval development), and might represent a significant range extension to the Philippines.

Remarks

Profundiconus puillandrei sp. nov. has been dredged alive from 1030–1180 m off Curtis Island, Kermadec Ridge, New Zealand (identified as Conus smirna ; see Marshall 1981). This observation makes this species one of the deepest-living ones among the known cone snails. P. puillandrei sp. nov. was initially identified as “giant” P. vaubani ( Fig. 1I ). Apart from being smaller in size, P. vaubani has a paucispiral protoconch of 1.75 whorls and a ridge at the shoulder with axial costae, which are absent in the case of P. puillandrei sp. nov. The shell pattern of P. vaubani consists of light brown axial streaks from base to spire, whereas the shell of P. puillandrei sp. nov. is patternless. The radular teeth of P. vaubani ( Fig. 2A–C ) and P. puillandrei sp. nov. ( Fig. 13 ) are superficially similar, but the tooth of P. vaubani has a much larger relative size, with S L/T L = 27–31 compared to 75–105 for P. puillandrei sp. nov. The new species can also be compared with P. profundorum ( Fig. 1A, L ) and P. smirnoides ( Fig. 1D ). Like P. puillandrei sp. nov, these two species have multispiral protoconchs.There are no significant differences in shell shape among these species: ANCOVA for MD, HMD and SH, using species hypothesis as a factor and S L as covariate, did not yield statistically significant results. However, they differ significantly in average shell length: puillandrei S L 39.82 mm, profundorum S L 96.93 mm (t = 10.28, p = 2.13 × 10-12; U = 0, p = 5.15 × 10-7), smirnoides S L 76.09 mm (t = -10.20, p = 5.25 × 10-10; U = 0, p = 2.5 × 10-5).

The shell of P. puillandrei sp. nov. has a distinct shoulder ridge, usually smooth but some times nodulose, which is absent in P. profundorum and less developed and always smooth in P. smirnoides . The shell of P.puillandrei sp.nov. is patternless, whereas both P.profundorum and P. smirnoides exhibit a characteristic pattern of broad, pale brown spiral bands on each side of centre, often interrupted by creamy white axial streaks in the case of P. smirnoides . The morphology of the radular tooth of P. puillandrei sp. nov. ( Fig. 13 ) and of P. smirnoides ( Fig. 9H ) is very different. The radular tooth of P. profundorum is unknown, preventing its comparison with the tooth of P. puillandrei sp. nov.

Most of the shells of P. puillandrei sp. nov. examined were not nodulose at the shoulder ridge ( Fig. 12 ). About 10% of the specimens studied had a nodulose spire and shoulder ridge (i.e., paratype 6, Fig. 12G ), coming mainly from Loyalty Ridge. These include two of the sequenced individuals, which, however, were genetically similar to the ones with a smooth ridge. Moreover, nodulose and non-nodulose specimens exhibit analogous radular and protoconch morphology. The presence of nodules at the spire and shoulder causes an apparent difference in shape, which is possibly the main source of intraspecific phenotypic variability within this species. Nodulose specimens may resemble a small P. teramachii (e.g., paratype 6, Fig. 12G ), a distinct but related species as inferred from the tree in Fig. 5 . The shell of P. teramachii is also patternless and has a ground colour and protoconch morphology similar to that of P. puillandrei sp. nov. The radular tooth of P. teramachii ( Fig. 2D–F ) is also similar to that of P. puillandrei sp. nov. ( Fig. 13 ). However, P. teramachii attains a larger size (S L = 55–111 mm), has a lower spire (RSH 0.11–0.22 versus 0.22–0.29 in P. puillandrei sp. nov.) and usually exhibits a broadly carinate shoulder. P. puillandrei sp. nov. is phylogenetically related to P. neotorquatus stat. nov., P. neocaledonicus sp. nov., P.teramachii and P. smirnoides .

Figures

Fig. 1. [opposite page] A. Profundiconus profundorum (Kuroda, 1956) (MJT coll., Okezoko, Kochi Prefecture, Japan, 350 m), 75.0 mm. B. Profundiconus teramachii (Kuroda, 1956) (MJT coll., South China Sea, trawled in 400 m), 111.3 mm. C. Profundiconus neotorquatus (da Motta, 1984) (MJT coll., NW coast of Madagascar, dredged in 600–800 m), 78.4 mm. D. Profundiconus smirnoides Tenorio, 2015 (holotype, MNHN IM-2009-18220, off Ile des Pins, New Caledonia, 480–500 m), 71.8 mm. E. Conilithes antidiluvianus (Bruguière, 1792), Upper Pliocene, Piacenzian Stage (MJT coll., Pedrera Anna, Molins de Rey, Barcelona, Spain), 35.3 mm. F. Profundiconus emersoni (Hanna, 1963) (LACM 146906, off Isla Santa Maria (Charles), Galápagos Is., Ecuador, 310 m), 33.7 mm. G. Profundiconus tuberculosus (Tomlin, 1937) (MJT coll., Miura, Sagami Bay, Kanagawa Prefecture, Japan, 100 m), 13.7 mm. H. Profundiconus loyaltiensis (Röckel & Moolenbeek, 1995) (holotype, MNHN IM-2000- 2545, Ride des Loyauté, New Caledonia, 480 m), 21.8 mm. I. Profundiconus vaubani (Röckel & Moolenbeek,1995)(holotype,MNHNIM-2000-3455,NorfolkRidge,NewCaledonia,435m),25.8mm. J. Profundiconus kanakinus (Richard, 1983) (MNHN, South New Caledonia, 410–440 m), 19.5 mm. K. Profundiconus cakobaui Moolenbeek et al., 2008 (holotype, MNHN IM-2000-21030, Somo-somo Strait,South ofVanua Levu, Fiji, 426-487m),18.9 mm.L.Profundiconusprofundorum (Kuroda, 1956) (MJT coll., South China Sea, trawled in 500–600 m), 109.9 mm. Scalebars = 10 mm.

Fig. 2. — A–C. Radular teeth extracted from Profundiconus vaubani (Röckel & Moolenbeek, 1995), with major parts and structures labelled. A. Optical photograph (MNHN, uncataloged, S L 20.6 mm). B. SEM photograph(MNHN, uncataloged, S L 26.0 mm).C. Enlargement of the middle section of B. — D–F. Radular teeth extracted from Profundiconus teramachii (Kuroda, 1956). D. Optical photograph (MNHN IM-2013-44602, S L 82.7 mm). E. SEM photograph (MNHN IM-2013-50257, S L 76.7 mm). F. Enlargement of the middle section of E.

Fig. 5. Likelihood phylogenetic tree of Profundiconus Kuroda, 1956 based on a subsample of the mitochondrial cox1 dataset produced by Puillandre et al. (2014). Posterior probabilities and bootstrap valuesare indicatedfor eachnode (whenPP± 0.95%and BP± 75%,respectively).Forclarity purposes, external branches of phylogenetically highly supported species are highlighted in red in the consensus tree.Cox1sequencesarelabelledusingtheMNHNvoucheridentificationnumber,theGenbankaccession number(when available), thespeciesnameand thenumberof specimens (oridentical haplotypes)when greater than 1.

Fig. 9. — A–D. Profundiconus virginiae sp. nov. A. Holotype, 42.5 × 19.4 mm. B. Paratype 1, 33.3 × 16.5 mm. C. Paratype 1, enlargement of the spire. D. Paratype 2 (fragment), 16.3 × 13.0 mm. — E. Profundiconus cf. cakobaui Moolenbeek et al., 2008 (MJT coll., Balut Is., Mindanao, Philippines, 450–500 m), 35.8 × 16.3 mm. — F. Profundiconus frausseni (Tenorio & Poppe, 2004) (holotype, NMP,Aliguay Island, Philippines, 100–150 m), 47.0 ×21.0 mm. — G. Radular tooth of Profundiconus virginiae sp. nov. from the holotype. — H. Radular tooth, Profundiconus smirnoides Tenorio, 2015 (MNHN IM-2000-30058, Norfolk Ridge, New Caledonia), S L 81.4 mm. Scale bars = 10 mm unless otherwise indicated.

Fig. 13. Radular teeth extracted from Profundiconus puillandrei sp. nov. A. Optical photomicrograph (holotype, MNHN IM-2000-30771, S L 43.2 mm). B. SEM photomicrograph (paratype 3, MNHN IM- 2000-30774, S L 35.0 mm). C. Enlargement of the external cusp. D. Enlargement of the middle section of B.

Fig. 14. Distribution map for Profundiconus puillandrei sp. nov. Red circles indicate the points where the species has been collected.

Fig.12.Profundiconuspuillandreisp.nov.A.Holotype,43.2×18.0mm.B.Paratype1,39.6×17.3mm. C. Paratype 2, 40.7 × 16.8 mm. D. Paratype 3, 35.0 × 15.5 mm. E. Paratype 4, 43.6 × 18.1 mm. F. Paratype 5,43.6 ×18.1 mm. G. Paratype6, 57.3 × 24.6 mm. H. Paratype4, enlargement of the spire. I. Paratype 7, 45.2 × 18.5 mm. J. Paratype 8, 38.9 × 17.5 mm. Scale bars = 10 mm unless otherwise indicated.