Abstract

A new genus-level classification of the Conoidea is presented, based on the molecular phylogeny of Puillandre et al. in the accompanying paper. Fifteen lineages are recognized and ranked as families to facilitate continuity in the treatment of the names Conidae (for ‘cones’) and Terebridae in their traditional usage. The hitherto polyphyletic ‘Turridae’ is now resolved as 13 monophyletic families, in which the 358 currently recognized genera and subgenera are placed, or tentatively allocated: Conorbidae (2 (sub)genera), Borsoniidae (34), Clathurellidae (21), Mitromorphidae (8), Mangeliidae (60), Raphitomidae (71), Cochlespiridae (9), Drilliidae (34), Pseudomelatomidae (=Crassispiridae) (59), Clavatulidae (14), Horaiclavidae new family (28), Turridae s. s. (16) and Strictispiridae (2). A diagnosis with description of the shell and radulae is provided for each of these families.

INTRODUCTION

The history of classification of the Conoidea can be divided in two mains periods. During the first period, from Adams & Adams (1853), Bellardi (1875), Fischer (1887) and Cossmann (1896) to the 1980s (e.g. Powell, 1942, 1966; Morrison, 1965; McLean, 1971), classifications were based solely on the morphology of the ‘hard parts’ (shell and radula). Contradictions were commonplace and none of these classifications satisfied the malacological community, even if some of them were adopted by other authors (e.g. Kilburn, 1983, 1985, 1986, 1988, 1991, 1992, 1993, 1994, 1995; Chang, 1995, 2001). Actually, only the Conidae (cone snails) and Terebridae (auger snails) were confidently recognized as well-defined groups, all the other constituents being lumped in the artificial group ‘turrids’, more or less separated into different subfamilies. Indeed, numerous cases of homoplasy for the morpho-anatomical characters used in these pre-1990s classifications are known (Taylor, 1994; Kantor, Medinskaya & Taylor, 1997; Fedosov & Kantor, 2008; Holford et al., 2009), thus rendering earlier classifications questionable. Protoconch characters, in particular, have been much used and much misunderstood (Bouchet, 1990). Inevitably, different authors emphasized different types of characters (radula, foregut anatomy, shape of the shell, protoconch): when one taxonomist recognized a group based on protoconch similarities and ignored the radula morphology, another one stated that the similarity of the protoconch was the result of convergence, and split the group into as many groups as there were radula types. This contributed to the conoideans earning their reputation of ‘taxonomic nightmare’.

In 1993, based on anatomical characters (and not just the radula), Taylor, Kantor and Sysoev proposed a completely revised classification that recognized six families. One of their major results was that Conus, hitherto a genus in its own monotypic family Conidae, became one of many genera in an expanded family Conidae, which also now contained genera formerly placed in ‘Turridae’, thus confirming Thiele's (1929) intuitive classification. The Taylor, Kantor & Sysoev (1993) classification formed the basis of the system published by Bouchet & Rocroi (2005; Fig. 1A). Most recently, a new shell-and-radula classification of the Toxoglossa, with emphasis on Conus and other ‘cone-like’ genera, has been proposed by Tucker & Tenorio (2009; Fig. 1C), who recognized two superfamilies: the Turroidea, grouping together the Turridae of Taylor et al. (1993), plus the Terebridae, Drilliidae, Pseudomelatomidae and Strictispiridae, and the Conoidea, corresponding to the Conidae of Taylor et al. (1993) except for several of their constituents having been raised to family level.

Figure 1.

Comparison of the last two conoidean classifications with the new classification proposed in this article. A.Taylor et al. (1993) and Bouchet & Rocroi (2005). 1Elevated to familial rank by Bouchet & Rocroi (2005). 2Absent in Taylor et al. (1993). B. Proposed classification. C.Tucker & Tenorio (2009). Fossil taxa.

Figure 1.

Comparison of the last two conoidean classifications with the new classification proposed in this article. A.Taylor et al. (1993) and Bouchet & Rocroi (2005). 1Elevated to familial rank by Bouchet & Rocroi (2005). 2Absent in Taylor et al. (1993). B. Proposed classification. C.Tucker & Tenorio (2009). Fossil taxa.

Based on a dataset of 57 genera and molecular sequences of fragments of one mitochondrial (COI) and three nuclear (28S, 18S and H3) genes, Puillandre et al. (2008) published the first molecular phylogeny of Conoidea. Even though ranks differed, most of the lineages defined by Taylor et al. (1993) were also retrieved in this molecular approach. The new molecular phylogeny of Puillandre et al. (2011) is based on a dataset of 102 conoidean genera (87 ‘turrids’, 5 cones and 10 terebrids) and sequences of three gene fragments (COI, 12S rRNA and 16S rRNA). Because of the high congruency between the classification based on anatomical characters and the molecular tree obtained, we are inclined to think that our understanding of the phylogeny of the Conoidea has now reached a stable position, and that a new classification is warranted. The classification presented below (Fig. 1B) aims to transform the molecular phylogeny into an operational classification by: (1) presenting revised morphological diagnoses for the now redefined families of Conoidea; (2) allocating (sometimes tentatively) all Recent genera recognized or used in current literature to the corresponding family.

MATERIAL AND METHODS

Genus-group names have been allocated to the newly defined families based on the following critera: Our aim was to provide an exhaustive list of conoidean genus-group names based on Recent type species or reasonably recognized as Recent in the literature. Genera present exclusively in the fossil record are not included. In very few cases, obviously erroneous attribution of Recent species to entirely fossil genera also excluded these genera from our listing. Junior homonyms and nomina nuda are also not included because they are unavailable for nomenclatural purposes. When the status of a nominal genus is uncertain, we adopted a ‘valid until synonymized’ approach; that is, a name is regarded valid (irrespective of our opinion) if it has not been synonymized in literature. The names of synonyms are placed after corresponding valid names (the synonyms applied to the valid subgenus name are positioned after the subgenus name). All the genus and subgenus names are also listed in alphabetic order in Appendix 1.

  • (1) 224 genera were assigned to a family based on shell characters, and phenetic resemblance to those genera with radula and/or molecular characters available; those are marked 1.

  • (2)103 genera were classified on the basis of radula morphology (both our own and published data), and congruence between radula and molecular characters for those genera that were sequenced; these are marked 2.

  • (3)98 genera were classified in a family on the basis of the molecular data; these are marked 3.

  • (4)63 genera and subgenera do not fall easily into the morphological groups resulting from the preceding steps, but they have tentatively been assigned to a family as a working hypothesis; these genera are preceded by a question mark.

  • (5)173 subgenera and/or synonyms are also listed; subgenera are in parentheses, synonyms in square brackets.

Radulae

To illustrate the range of radular types in the molecularly defined clades, the radulae were extracted as far as possible from the specimens used for the molecular analyses or from conspecific specimens (after rehydration when soft parts had been dried), cleaned with diluted bleach, rinsed in distilled water, mounted on stubs, air dried, coated with gold-palladium, and investigated with a JEOL JSM 840A scanning electron microscope. Some of the previously photographed radulae (examined by YuIK and/or J.D. Taylor) were additionally illustrated.

For radular descriptions we mostly followed the terminology accepted and discussed by Kantor & Taylor (2000). In radular formulae, the parentheses indicate partial or complete fusion of lateral and rachidian teeth (for more details, see Kantor, 2006).

Ranking and nomenclature

Although the names Conoidea and Toxoglossa are used interchangeably in the taxonomic literature, we have avoided the name Toxoglossa because (i) it is not typified and cannot be used for a family–group name, and (ii) many of the included taxa do not have a toxoglossate radula. Within the Conoidea, ranking of the clades was determined by a conservative approach, thus retaining the names Terebridae and Conidae s. s. (the latter including the cone snails Profundiconus, Californiconus, Conasprella, Conus and Taranteconus) in their accustomed usage at family rank. An alternative would have been to recognize only two families, Conidae sensuTaylor et al. (1993) and Turridae (including the Turridae, Terebridae, Drilliidae and Pseudomelatomidae), with the resulting inconvenience that Conidae s. s. and Terebridae, although monophyletic, would lose their traditional usage and the vast associated literature dealing with these names that largely ignores the Taylor et al. (1993) and Bouchet & Rocroi (2005) classifications.

Fourteen clades of rank equivalent to Conidae s. s. and Terebridae are recognizable from the molecular phylogenetic tree (Puillandre et al., 2011). Forty-three family–group names within Conoidea are nomenclaturally available (Appendix 2), of which five are based on a genus with a fossiltype species (Andoniinae Vera-Pelaez, 2002; Cryptoconinae Cossmann, 1896; Hemiconidae Tucker & Tenorio, 2009; Johnwyattidae Serna, 1979; Siphopsinae Le Renard, 1995). These five cannot be applied to a molecular clade and will not be used in our classification. Names were applied to clades based on the position of their type genus in the tree. If more than one family–group name was applicable, the valid name was determined by priority. If the type genus of a nominal family name was not sequenced, application of the name was determined by reference to the morphologically most similar genus used in the analysis.

Because the molecular taxon sampling is still too patchy for such levels, we have abstained from extending the classification below family level (i.e. subfamilies, tribes), even when some molecular clades obviously match previously recognized ‘subfamilies’ (e.g. Californiconinae, Oenopotinae, Zemaciinae, Zonulispirinae). They are all included in a family (Appendix 2), without precluding their usefulness and potential taxonomic validity.

Illustration of voucher material

Each family (except the Conidae and Terebridae, already extensively covered and illustrated in other recent works, e.g. Röckel, Korn & Kohn, 1995; Terryn, 2007; Tucker & Tenorio, 2009) is illustrated by one or several shells, radulae and protoconchs, covering the morphological variability of the group. As far as possible, specimens used for the molecular analyses were used for illustration. However, since a substantial part of the morphological variability was not covered by our dataset, shells and radulae of other specimens are also illustrated.

SYSTEMATICS

Superfamily CONOIDEA Fleming, 1822

Family Conidae Fleming, 1822

[=Conilithidae Tucker & Tenorio, 2009, new synonym; =Taranteconidae Tucker & Tenorio, 2009, new synonym]

Diagnosis: Shell medium-sized to large or very large, normally 20–50 mm, up to 170 mm high, conical or biconical, with narrow aperture and short siphonal canal. Shell with considerable internal remodelling due to inner wall resorption. Spiral sculpture usually developed, axial sculpture absent or in form of shoulder tuberculation. Anal sinus on subsutural ramp, shallow to moderately deep. Operculum present, small, with terminal nucleus. Radula of marginal hypodermic teeth, generally harpoon-shaped, barbed at tip, often with complex inner structure of folds and serration, base small and swollen, tooth canal opening (sub)terminally, rarely laterally. Subradular membrane vestigial. Teeth can be attached to the membrane by long or very long flexible ligament. Tooth wall forms several overlapping layers.

Remarks: in Conidae and the other families that are included in this major clade (family Conidae sensuTaylor et al., 1993), the radula consists only of marginal teeth that are usually enrolled with completely overlapping edges (hypodermic). Teeth at their formation in the radular sac are already enrolled and they are attached to the radular membrane only by their bases, sometimes through a long flexible ligament. In the ‘turrids’ with enrolled teeth, these are attached to the membrane along most of its length (Kantor & Taylor, 2000). A molecular phylogeny of the Conidae is currently in preparation (C. Meyer, personal communication).

Included genera: Conorbidae de Gregorio, 1880

  • Africonus Petuch, 19751

  • Asprella Schaufuss, 18691

    •  [= Embrikena Iredale, 1937]

    •  [= Fumiconus da Motta, 1991]

    •  [= Graphiconus da Motta, 1991]

  • AustroconusTucker & Tenorio, 20091

  • BathyconusTucker & Tenorio, 20091

  • CalamiconusTucker & Tenorio, 20091

  • Calibanus da Motta, 19911

  • CaliforniconusTucker & Tenorio, 20093

  • Chelyconus Mörch, 18521

  • ConasprellaThiele, 19293

  • ConasprelloidesTucker & Tenorio, 20091

  • Conus Linnaeus, 17583

    •  [= Coronaxis Swainson, 1840]

    •  [= Cucullites Herrmannsen, 1847]

    •  [= Cucullus Röding, 1798]

  • Cylindrus Batsch, 17891 (For authorship of Cylindrus, see Dubois & Bour (2010))

  •   [= Cylinder Montfort, 1810]

  •   [= Cylindrella Swainson, 1840]

  • Dalliconus Tucker & Tenorio, 20091

  • Darioconus Iredale, 19301

  •   [= Erythroconus da Motta, 1991]

  •   [= Regiconus Iredale, 1930]

  • Dauciconus Cotton, 19451

    •  [= Cariboconus Petuch, 1991]

  • Dendroconus Swainson, 18401

    •  [= Clebula Iredale, 1930]

  • Ductoconus da Motta, 19911

  • Dyraspis Iredale, 19491

  • Endemoconus Iredale, 19311

    •  [= Kermasprella Powell, 1958]

    •  [= Mamiconus Cotton & Godfrey, 1932]

  • EremiconusTucker & Tenorio, 20091

  • Eugeniconus da Motta, 19911

  • Floraconus Iredale, 19301

  • Fulgiconus da Motta, 19911

    •  [= Thoraconus da Motta, 1991]

  • Fusiconus da Motta, 19911

  • Gastridium Modeer, 17931

    •  [= Rollus Montfort, 1810]

    •  [= Tuliparia Swainson, 1840]

  • GenuanoconusTucker & Tenorio, 20091

  • GladioconusTucker & Tenorio, 20091

  • GlobiconusTucker & Tenorio, 20091

  • Gradiconus da Motta, 19911

  • Harmoniconus da Motta, 19911

  • Hermes Montfort, 18101

    •  [= Theliconus Swainson, 1840]

  • Jaspidiconus Petuch, 20031

  • Kalloconus da Motta, 19911

  • Kenyonia Brazier, 18961

  • Ketyconus da Motta, 19911

  • Kioconus da Motta, 19911

    •  [= Ongoconus da Motta, 1991]

  • KohniconusTucker & Tenorio, 20091

  • Kuradoconus Shikama & Habe, 19681

  • Lamniconus da Motta, 19911

  • Lautoconus Monterosato, 19231

  • Leporiconus Iredale, 19301

  • Leptoconus Swainson, 18401

  • Lilliconus Raybaudi Massilia, 19941

  • Lindaconus Petuch, 20021

    •  [= Calusaconus Petuch, 2003]

    •  [= Spuriconus Petuch, 2003]

    •  [= Utriculus Schumacher, 1817]

  • Lithoconus Mörch, 18521

    •  [= Tesselliconus da Motta, 1991]

  • Lividoconus Wils, 19701

    •  [= Splinoconus da Motta, 1991]

  • MiliariconusTucker & Tenorio, 20091

  • Monteiroconus da Motta, 19911

  • NataliconusTucker & Tenorio, 20091

  • Parviconus Cotton & Godfrey, 19321

  • Perplexiconus Tucker & Tenorio, 20091

  • Phasmoconus Mörch, 18521

  • Pionoconus Mörch, 18521

    •  [= Heroconus da Motta, 1991]

    •  [= Socioconus da Motta, 1991]

    •  [= StrioconusThiele, 1929]

  • Plicaustraconus Moolenbeek 20081

  • Profundiconus Kuroda, 19563

    •  [= Lizaconus da Motta, 1991]

  • Protoconus da Motta, 19911 [invalid name]

    •  [= Seminoleconus Petuch, 2003]

  • ProtostrioconusTucker & Tenorio, 20091

  • PseudoconorbisTucker & Tenorio, 20091

  • PseudolilliconusTucker & Tenorio, 20091

  • Pseudonoduloconus Tucker & Tenorio, 20091

  • Puncticulis Swainson, 18401

  • Purpuriconus da Motta, 19911

    •  [= Magelliconus da Motta, 1991]

  • Pyruconus Olsson, 19671

  • Quasiconus Tucker & Tenorio, 20091

  • Rhizoconus Mörch, 18521

  • Rhombiconus Tucker & Tenorio, 20091

    •  [= Rhombus Montfort, 1810]

  • Rolaniconus Tucker & Tenorio, 20091

  • Sciteconus da Motta, 19911

  • Stellaconus Tucker & Tenorio, 20091

  • Stephanoconus Mörch, 18521

  • Strategoconus da Motta, 19911

  • Taranteconus Azuma, 19723

    •  [= Cornutoconus Suzuki, 1972]

  • Textilia Swainson, 18401

  • Trovaoconus Tucker & Tenorio, 20091

  • Turriconus Shikama & Habe, 19681

  • Varioconus da Motta, 19911

  • Viminiconus Tucker & Tenorio, 20091

  • Virgiconus Cotton, 19451

  • Virroconus Iredale, 19301

  • Vituliconus da Motta, 19911

  • Ximeniconus Emerson & Old, 19621

  • Yeddoconus Tucker & Tenorio, 20091

(Figs 2O, 22C)

Figure 2.

Shells. AN. Borsoniidae. A.Typhlomangelia sp.**, MNHN IM200717931, Solomon Islands, SALOMON 2, st. CP2269, 07°45.1′S, 156°56.3′E, 768–890 m, SL 14.2 mm. B. Borsoniidae gen. 1**, MNHN IM200717911, Philippines, PANGLAO 2005, st. CP2333, 9°38.2′N, 123°43.5′E, 584–596 m, SL 17.1 mm (radula: Fig. 3D). C.Darbya lira Bartsch, 1934, USNM 810565, off Puerto Rico, SL 19.4 mm. D.Borsonia sp.**, MNHN IM200717932, Solomon Islands, SALOMON 2, st. CP2197, 08°24.4′S, 159°22.5′E, 897–1057 m, male, SL 24.6 mm (radula: Fig. 3B). E.Bathytoma neocaledonica Puillandre et al., 2010**, paratype, MNHN IM200717857, Coral Sea, EBISCO, st. CP2551, SL 44 mm (radula: Fig. 3C). F.Zemacies excelsa Sysoev & Bouchet, 2001**, MNHN IM200911056, New Caledonia, Musorstom 4, st. DW226, 22°47′S, 167°22′E, 395 m, SL 38 mm. G.Tomopleura reevii (C. B. Adams, 1850)**, MNHN IM200717875, Philippines, PANGLAO 2004, st. T26, 09°43.3′N, 123°48.8′E, 123–135 m, SL 7.7 mm. H.Microdrillia cf. optima (Thiele, 1925)**, MNHN IM200717887, Philippines, PANGLAO 2004, st. T36, 09°29.3′N, 123°51.5′E, 95–128 m, SL unknown (broken). I.Borsonellopsis chrysothemis (Dall, 1919), SIO (Scripps Institution of Oceanography) M2530, San Diego Trough, off San Diego Co., California, 1244 m, SL 19.3 mm. J.Belaturricula turrita (Strebel, 1908), USNM 897559, South Georgia Island, 54.3°S, 35.6°W, 238–247 m, SL 49.8 mm. K.Glyptaesopus oldroydi (Arnold, 1903). 10 m, off Punta Abreojos, Baja California, Mexico, SL 8.3 mm. L.Tropidoturris fossata fossata (Sowerby, 1903), holotype, SAMC A346, off Cape Vidal, Zululand, 146–182 m, SL 20.4 mm. M.Genota mitriformis (Wood, 1828)**, MNHN IM200742293, Angola, 30–50 m, SL 30 mm (radula: Fig. 3A). N.Ophiodermella inermis (Hinds, 1843), USNM 56092, W coast of North America, SL 40.7 mm. O: Conorbidae. O.Benthofascis lozoueti Sysoev & Bouchet, 2001**, MNHN IM200742331, New Caledonia, NORFOLK 2, st. DW2147, 22°50′S, 167°16′E, 496 m, SL 16.5 mm. Abbreviation and symbols: SL, shell length; * , sequenced species; **, sequenced specimen. Photo credits: B. Buge (A, B, D, E, G, H), M.G. Harasewych and D. Tippett (C, J, N), R.N. Kilburn (L), J. McLean (I, K).

Figure 2.

Shells. AN. Borsoniidae. A.Typhlomangelia sp.**, MNHN IM200717931, Solomon Islands, SALOMON 2, st. CP2269, 07°45.1′S, 156°56.3′E, 768–890 m, SL 14.2 mm. B. Borsoniidae gen. 1**, MNHN IM200717911, Philippines, PANGLAO 2005, st. CP2333, 9°38.2′N, 123°43.5′E, 584–596 m, SL 17.1 mm (radula: Fig. 3D). C.Darbya lira Bartsch, 1934, USNM 810565, off Puerto Rico, SL 19.4 mm. D.Borsonia sp.**, MNHN IM200717932, Solomon Islands, SALOMON 2, st. CP2197, 08°24.4′S, 159°22.5′E, 897–1057 m, male, SL 24.6 mm (radula: Fig. 3B). E.Bathytoma neocaledonica Puillandre et al., 2010**, paratype, MNHN IM200717857, Coral Sea, EBISCO, st. CP2551, SL 44 mm (radula: Fig. 3C). F.Zemacies excelsa Sysoev & Bouchet, 2001**, MNHN IM200911056, New Caledonia, Musorstom 4, st. DW226, 22°47′S, 167°22′E, 395 m, SL 38 mm. G.Tomopleura reevii (C. B. Adams, 1850)**, MNHN IM200717875, Philippines, PANGLAO 2004, st. T26, 09°43.3′N, 123°48.8′E, 123–135 m, SL 7.7 mm. H.Microdrillia cf. optima (Thiele, 1925)**, MNHN IM200717887, Philippines, PANGLAO 2004, st. T36, 09°29.3′N, 123°51.5′E, 95–128 m, SL unknown (broken). I.Borsonellopsis chrysothemis (Dall, 1919), SIO (Scripps Institution of Oceanography) M2530, San Diego Trough, off San Diego Co., California, 1244 m, SL 19.3 mm. J.Belaturricula turrita (Strebel, 1908), USNM 897559, South Georgia Island, 54.3°S, 35.6°W, 238–247 m, SL 49.8 mm. K.Glyptaesopus oldroydi (Arnold, 1903). 10 m, off Punta Abreojos, Baja California, Mexico, SL 8.3 mm. L.Tropidoturris fossata fossata (Sowerby, 1903), holotype, SAMC A346, off Cape Vidal, Zululand, 146–182 m, SL 20.4 mm. M.Genota mitriformis (Wood, 1828)**, MNHN IM200742293, Angola, 30–50 m, SL 30 mm (radula: Fig. 3A). N.Ophiodermella inermis (Hinds, 1843), USNM 56092, W coast of North America, SL 40.7 mm. O: Conorbidae. O.Benthofascis lozoueti Sysoev & Bouchet, 2001**, MNHN IM200742331, New Caledonia, NORFOLK 2, st. DW2147, 22°50′S, 167°16′E, 496 m, SL 16.5 mm. Abbreviation and symbols: SL, shell length; * , sequenced species; **, sequenced specimen. Photo credits: B. Buge (A, B, D, E, G, H), M.G. Harasewych and D. Tippett (C, J, N), R.N. Kilburn (L), J. McLean (I, K).

Diagnosis (from Tucker & Tenorio, 2009): “Radular tooth: anterior fold is usually present; basal spur is directed toward the apex of the tooth or parallel with the tooth base; the waist, base and C-fold are absent; terminating cusps, serratins and accessory process are also absent. Shell characters: the interior of the shell is extensively remodelled including the columellar region; nodules are absent but cords may be present. Shells can be squatly conical to elongated or biconical.”

Remarks: The type genus of the family is Conorbis Swainson, 1840, based on an Eocene type species. The only Recent species that was believed to belong to Conorbis, Conus coromandelicus E.A. Smith, 1894, was made the type of the new genus Pseudoconorbis Tucker & Tenorio, 2009, and included by them in their family Conilithidae (herein in Conidae). In the molecular phylogeny of Puillandre et al. (2011), what we have identified as ‘Benthofascis’ constitutes an independent lineage (i.e. not included in any other family-ranked clade), characterized by a long branch. This certainly explains why its position is not the same in the Maximum Likelihood analyses (sister group of the Borsoniidae) and in the Bayesian analyses (included in the Borsoniidae). Benthofascis lozoueti (the sequenced species) differs from other species of the genus in the absence of inner shell wall resorption (N. Puillandre, personal observations), but the phylogenetic and taxonomic value of this character remains unverified. The problem could be resolved by sequencing other species of Benthofascis, which have inner wall resorption. Until then we prefer to keep Benthofascis lozoueti within this genus and to apply the name Conorbidae to this lineage.

Included genera:

Artemidiconus da Motta, 19911

Benthofascis Iredale, 1936 3

Borsoniidae A. Bellardi, 1875

[=Zemaciinae Sysoev, 2003, new synonym]

(Figs 2A–N, 3)

Figure 3.

Radulae. Borsoniidae. A.Genota mitriformis (Wood, 1828)**, MNHN IM200742293 (shell: Fig. 2M). B.Borsonia sp.**, MNHN IM200717932 (shell: Fig. 2D). C.Bathytoma neocaledonica Puillandre et al., 2010**, MNHN IM200717857 (shell: Fig. 2E). D. Borsoniidae gen. 1**, MNHN IM200717911 (shell: Fig. 2B). Symbol: **, sequenced specimen.

Figure 3.

Radulae. Borsoniidae. A.Genota mitriformis (Wood, 1828)**, MNHN IM200742293 (shell: Fig. 2M). B.Borsonia sp.**, MNHN IM200717932 (shell: Fig. 2D). C.Bathytoma neocaledonica Puillandre et al., 2010**, MNHN IM200717857 (shell: Fig. 2E). D. Borsoniidae gen. 1**, MNHN IM200717911 (shell: Fig. 2B). Symbol: **, sequenced specimen.

Diagnosis: Shell small to large (5–80 mm), fusiform to biconic, sometimes with strong to obsolete columellar pleats. Sculpture usually well developed, axial ribs sometimes obsolete to absent. Siphonal canal short to moderately long. Anal sinus on subsutural ramp, deep. Protoconch when multispiral with up to five whorls, initially smooth and then with arcuate axial riblets, when paucispiral up to two smooth whorls. Operculum with terminal nucleus, fully developed to missing. Radula of hypodermic marginal teeth that usually have a weakly developed solid basal part, often attached to the ligament (marked by an arrow on Fig. 3C). Tooth canal opening (sub)terminally or, sometimes, laterally. At their tip teeth can have weak to rather strong barb(s) (Genota, Fig. 3A). Overlapping of the tooth edges is weak (Fig. 3C). In Zemacies, the radula is completely absent.

Remarks: This is a rather heterogeneous group. Obviously, it is not fully resolved as it is based on molecular data and comprises rather conchologically different clades. This could be explained by the fact that many taxa of this group are among most ancient of conoideans, known since the Palaeocene (Zemacies, Borsonia, Tomopleura) or Eocene (Bathytoma, Genota, Microdrillia). The loss of apomorphies by mutation could be more important for old taxa (Puillandre et al., 2011).

The names Borsoniinae Bellardi, 1875, and Pseudotominae Bellardi, 1875, were established simultaneously. As First Revisers, under Art. 24 of the ICZN Code, we here give precedence to the former over the latter.

Included genera: Clathurellidae H. Adams & A. Adams, 1858

  • ?Apaturris Iredale, 19171

  • AphanitomaBellardi, 18751

  • Asthenotoma Harris & Burrows, 18911

  • Austroturris Laseron, 19541

  • Bathytoma Harris & Burrows, 18913

    •  (Micantapex Iredale, 1936)

    •  (Parabathytoma Shuto, 1961)

    •  (Riuguhdrillia Oyama, 1951)

  • ?Belaturricula Powell, 19512

  • Borsonella Dall, 19082

  • BorsonellopsisMcLean, 19711

  • Borsonia Bellardi, 18393

    •  (Boettgeriola Wenz, 1943)

  • Cordieria Rouault, 18481

  • Darbya Bartsch, 19441

  • Diptychophlia Berry, 19641

  • Drilliola Cossmann, 19032

  • Filodrillia Hedley, 19221

  • GenotaH. Adams & A. Adams, 18533

    •  [= Genotia Fischer, 1883]

  • ?Glyptaesopus Pilsbry & Olsson, 19412

  • Heteroturris Powell, 19671

  • MaoritomellaPowell, 19422

    •  [= Narraweena Laseron, 1954]

  • Microdrillia Casey, 19033

    •  [= Acropota Nordsieck, 1977]

  • Ophiodermella Bartsch, 19442

  • Paraborsonia Pilsbry, 19221

  • Phenatoma Finlay, 19242

  • Pulsarella Laseron, 19542

  • Retidrillia McLean, 20001

  • Suavodrillia Dall, 19082

  • Tomopleura Casey, 19043

  • TropidoturrisKilburn, 19862

  • ?Typhlodaphne Powell, 19511

  • Typhlomangelia G.O. Sars, 18783

    •  [= Viridoturris Powell, 1964]

    •  [= Vexithara Finlay, 1926]

  • Zemacies Finlay, 19263

(Figs 4A–F, 5)

Figure 4.

Shells. A–F. Clathurellidae. A.Nannodiella ravella (Hedley, 1922)*, MNHN IM200742350, Philippines, PANGLAO 2004, st. T9, 09°33.5′N, 123°49.5′E, 97–120 m, SL 5.8 mm (radula: Fig. 5A). B.Comarmondia gracilis (Montagu, 1803), MNHN, Le Brusc, Cap Sicié, Provence, France, 40–100 m, SL 23 mm. C.Etrema cf. tenera (Hedley, 1899)**, MNHN IM200717869, Philippines, PANGLAO 2004, st. S21, 09°41.7′N, 123°50.9′E, 4–12 m, SL unknown (broken). D.Glyphostoma rostrata Sysoev & Bouchet, 2001, MNHN, New Caledonia, BATHUS 4, st. DW896, 20°16′S, 163°52′E, 315–350 m, SL 21.5 mm. E.Lienardia nigrotincta (Montrouzier, 1872)*, MNHN, Touho, New Caledonia, 20°45.2′S, 165°16.3′E, intertidal, SL 6.9 mm (radula: Fig. 5B). F.Strombinoturris crockeri Hertlein & Strong, 1951, LACM 747–37, off Isabel Island, Mexico, 18–33 m, SL 47.5 mm. G–I: Mitromorphidae. G.Lovellona atramentosa (Reeve, 1849)*, MNHN, Philippines, PANGLAO 2004, st. M2, 09°32.8′N, 123°45.9′E, 0–2 m, SL 9.0 mm (radula: Fig. 6A). H.Mitromorpha metula (Hinds, 1843)*, MNHN IM200742339, Philippines, PANGLAO 2004, st. B8, 09°37.1′N, 123°46.1′E, 3 m, SL 3.1 mm (radula: Fig. 6B). I.Anarithma sp.*, MNHN, Philippines, PANGLAO 2004, st. S5, 09°37.1′N, 123°46.1′E, 2–4 m, SL 6.9 mm (radula: Fig. 6C). Abbreviation and symbols: SL, shell length; * , sequenced species; **, sequenced specimen. Photo credits: B. Buge (C), M.G. Harasewych and D. Tippett (F), P. Maestrati (D, E).

Figure 4.

Shells. A–F. Clathurellidae. A.Nannodiella ravella (Hedley, 1922)*, MNHN IM200742350, Philippines, PANGLAO 2004, st. T9, 09°33.5′N, 123°49.5′E, 97–120 m, SL 5.8 mm (radula: Fig. 5A). B.Comarmondia gracilis (Montagu, 1803), MNHN, Le Brusc, Cap Sicié, Provence, France, 40–100 m, SL 23 mm. C.Etrema cf. tenera (Hedley, 1899)**, MNHN IM200717869, Philippines, PANGLAO 2004, st. S21, 09°41.7′N, 123°50.9′E, 4–12 m, SL unknown (broken). D.Glyphostoma rostrata Sysoev & Bouchet, 2001, MNHN, New Caledonia, BATHUS 4, st. DW896, 20°16′S, 163°52′E, 315–350 m, SL 21.5 mm. E.Lienardia nigrotincta (Montrouzier, 1872)*, MNHN, Touho, New Caledonia, 20°45.2′S, 165°16.3′E, intertidal, SL 6.9 mm (radula: Fig. 5B). F.Strombinoturris crockeri Hertlein & Strong, 1951, LACM 747–37, off Isabel Island, Mexico, 18–33 m, SL 47.5 mm. G–I: Mitromorphidae. G.Lovellona atramentosa (Reeve, 1849)*, MNHN, Philippines, PANGLAO 2004, st. M2, 09°32.8′N, 123°45.9′E, 0–2 m, SL 9.0 mm (radula: Fig. 6A). H.Mitromorpha metula (Hinds, 1843)*, MNHN IM200742339, Philippines, PANGLAO 2004, st. B8, 09°37.1′N, 123°46.1′E, 3 m, SL 3.1 mm (radula: Fig. 6B). I.Anarithma sp.*, MNHN, Philippines, PANGLAO 2004, st. S5, 09°37.1′N, 123°46.1′E, 2–4 m, SL 6.9 mm (radula: Fig. 6C). Abbreviation and symbols: SL, shell length; * , sequenced species; **, sequenced specimen. Photo credits: B. Buge (C), M.G. Harasewych and D. Tippett (F), P. Maestrati (D, E).

Figure 5.

Radulae. Clathurellidae. A.Nannodiella ravella (Hedley, 1922)*, MNHN IM200742350 (shell: Fig. 4A). B.Lienardia nigrotincta (Montrouzier, 1872)*, MNHN (shell: Fig. 4E). C.Lienardia jousseaumei (Hervier, 1896), MNHN, Philippines, PANGLAO 2004, st. B7, 9°35.9′N, 123°51.8′E, 4–30 m. Symbol: *, sequenced species.

Figure 5.

Radulae. Clathurellidae. A.Nannodiella ravella (Hedley, 1922)*, MNHN IM200742350 (shell: Fig. 4A). B.Lienardia nigrotincta (Montrouzier, 1872)*, MNHN (shell: Fig. 4E). C.Lienardia jousseaumei (Hervier, 1896), MNHN, Philippines, PANGLAO 2004, st. B7, 9°35.9′N, 123°51.8′E, 4–30 m. Symbol: *, sequenced species.

Diagnosis: Shell small- to medium-sized (5–40, usually 10–20 mm), fusiform to broadly fusiform, subsutural ramp usually unclearly separated. Sculpture mostly strong, usually cancellate, subsutural fold lacking. Shell surface often microgranular. Apertural armature often well developed, in the form of pleats and denticles on both inner and outer lips; no true columellar pleats. Siphonal canal well expressed, moderately long. Anal sinus on subsutural ramp, deep, often (sub)tubular. Protoconch typically multispiral, up to six whorls, generally smooth with keeled last whorls; when paucispiral, protoconch usually smooth or spirally striated, sometimes with remaining carination in last portion. Operculum always absent. Radula of hypodermic marginal teeth that usually have relatively small solid basal part (Fig. 5). Distinct ligaments not found. At their tip, teeth can have a weak barb.

Included genera: Mitromorphidae Casey, 1904, new rank

  • Acrista Hedley, 19221

  • ?Adanaclava Bartsch, 19501

  • ?Clathromangelia Monterosato, 18841

  • Clathurella Carpenter, 18573

  • Comarmondia Monterosato, 18441

    •  [= Bellatula Strand, 1929]

  • Corinnaeturris Bouchet & Warén, 19802

  • Crockerella Hertlein & Strong, 19512

  • Etrema Hedley, 19183

    •  [= Iraquetrema Dance & Eames, 1966]

    •  (Etremopa Oyama, 1953)

    •  (EtremopsisPowell, 1942)

  • Euclathurella Woodring, 19281

  • Glyphostoma Gabb, 18722

    •  (Euglyphostoma Woodring, 1970)

    •  (Glyphostomops Bartsch, 1934)

  • Lienardia Jousseaume, 18841

    •  (Thetidos Hedley, 1899)

  • Nannodiella Dall, 19193

  • Paraclathurella Boettger, 18951

  • Pseudoetrema Oyama, 19531

  • Strombinoturris Hertlein & Strong, 19512

  • Turrella Laseron, 19541

(Figs 4F–H, 6)

Figure 6.

Radulae. Mitromorphidae. A.Lovellona atramentosa (Reeve, 1849)*, MNHN (shell: Fig. 4G). B.Mitromorpha metula (Hinds, 1843), MNHN IM200742339 (shell: Fig. 4H). C.Anarithma sp.*, MNHN (shell: Fig. 4 I). Symbol: *, sequenced species.

Figure 6.

Radulae. Mitromorphidae. A.Lovellona atramentosa (Reeve, 1849)*, MNHN (shell: Fig. 4G). B.Mitromorpha metula (Hinds, 1843), MNHN IM200742339 (shell: Fig. 4H). C.Anarithma sp.*, MNHN (shell: Fig. 4 I). Symbol: *, sequenced species.

Diagnosis: Shell small- to medium-sized, 3–30 mm, usually 5–10 mm high, biconic, of mitriform shape. Sculpture rather smooth, with dominant spiral elements. Aperture narrow, with or without 1–3 columellar pleats, sometimes with denticles within. Siphonal canal short or indistinct. Anal sinus from indistinct to rather shallow indentation on weakly pronounced subsutural ramp. Protoconch multispiral or paucispiral, up to 4.5 smooth whorls. No operculum. Radula of hypodermic, marginal, relatively short, awl-shaped teeth with large swollen solid basal part (Fig. 6). Distinct ligaments present, short. Tooth canal opening subterminally or laterally. At their tip, teeth can have a weak barb (Fig. 6C).

Remarks: Mitrolumninae was established as a substitute name for Diptychomitrinae. Mitromorphidae and Mitrolumninae were published the same year but Mitromorphidae (19 May 1904) has priority over Mitrolumninae (31 August 1904).

Included genera: Mangeliidae P. Fischer, 1883, new rank

  • Anarithma Iredale, 19181

  • Arielia Shasky, 19611

    •  (Vexiariella Shuto, 1983)

  • Lovellona Iredale, 19173

  • Maorimorpha Powell, 19391

  • MitrellatomaPowell, 19421

  • Mitromorpha Carpenter, 18653

    •  [= Cymakra Gardner, 1937]

    •  [= Helenella Casey, 1904]

    •  [= ?Itia Marwick, 1931]

    •  [= Mitrithara Hedley, 1922]

    •  [= Mitrolumna Bucquoy, Dautzenberg & Dollfus, 1883]

  • Scrinium Hedley, 19221

[= Oenopotinae Bogdanov, 1987, new synonym]

(Figs 7, 8)

Figure 7.

Shells. Mangeliidae. A.Bela clarae Peňas & Rolán, 2008, paratype, MNHN Moll 21396, Sitges, Vallcarca, Barcelona, Spain, 45–60 m, SL 7 mm. B.Macteola segesta (Duclos in Chenu, 1850), MNHN, LIFOU 2000, st. 1425, 20°46.8′S, 167°07.2′E, 4–5 m, SL 4.6 mm. C.Benthomangelia cf. trophonoidea (Schepman, 1913)**, MNHN IM200717835, Vanuatu, BOA1, st. CP2462, 16°38′S, 167°57′E, 618–641 m, SL 21 mm (radula: Fig. 8A). D.Citharomangelia sp., MNHN, Futuna Island, MUSORSTOM 7, st. DW504, 14°20′S, 178°04′E, 300–390 m, SL 15.4 mm. E.Anticlinura sp.**, MNHN IM200742513, Solomon Islands, SALOMON 2, st. CP2182, 08°47.0′S, 159°37.9′E, 762–1060 m, SL 17.3 mm (radula: Fig. 8E). F.Eucithara cf. coronata (Hinds, 1843)**, MNHN IM200717900, Philippines, PANGLAO 2004, st. B8, 09°37.1′N, 123°46.1′E, 3 m, SL 8.7 mm (radula: Fig. 8D). G.Leiocithara sp., MNHN, New Caledonia, BATHUS 2, st. DW719, 22°48′S, 167°16′E, 444–445 m, SL 5.4 mm. H.Oenopota uschakovi Bogdanov, 1985, holotype, ZIN 37974/1, Iturup Island, Kurile Islands, 44°24.2′N, 147°09′01″E. 160 m, SL 10.3 mm. I.Genotina adamii (Bozzetti, 1994), MNHN, LIFOU 2000, st. DW1650, 20°54.15′S, 167°01.7′E, 120–150 m, SL 20.0 mm. J.Vitjazinella multicostata Sysoev, 1988, Paratype, ZMMU 22356, Izu-Bonin Trench, 33°18′N, 149°46′E., 6096 m, SL 7.2 mm. K.Oenopotella ultraabyssalis ultraabyssalis Sysoev, 1988, holotype, ZMMU Lc-22681, Kurile-Kamchatka trench, 49°29′N, 158°41′E, 7210–7230 m, SL 12.8 mm. L.Pseudorhaphitoma sp., Coral Sea, Chestefield Plateau, 19 53′S, 158 40′E, 410 m, SL 9.7 mm. M.Paraspirotropis simplicissima (Dall, 1907), ZIN, off Shikotan I., S Kurile Islands, 1450–1530 m, SL 28.0 mm. N.Heterocithara sp.**, MNHN IM200717884, Philippines, PANGLAO 2004, st. L46, 09°30.9′N, 123°41.2′E, 90–110 m, SL 3.4 mm. O.Toxicochlespira pagoda Sysoev & Kantor, 1990**, MNHN IM200717925, Solomon Islands, SALOMON 2, st. CP2227, 6°37.2′N, 156°12.7′E, 508–522 m, SL 18.0 mm (radula: Fig. 8B). Abbreviation and symbol: SL, shell length; **, sequenced specimen. Photo credits: B. Buge (F, O), P. Maestrati (B, D, G, I, L).

Figure 7.

Shells. Mangeliidae. A.Bela clarae Peňas & Rolán, 2008, paratype, MNHN Moll 21396, Sitges, Vallcarca, Barcelona, Spain, 45–60 m, SL 7 mm. B.Macteola segesta (Duclos in Chenu, 1850), MNHN, LIFOU 2000, st. 1425, 20°46.8′S, 167°07.2′E, 4–5 m, SL 4.6 mm. C.Benthomangelia cf. trophonoidea (Schepman, 1913)**, MNHN IM200717835, Vanuatu, BOA1, st. CP2462, 16°38′S, 167°57′E, 618–641 m, SL 21 mm (radula: Fig. 8A). D.Citharomangelia sp., MNHN, Futuna Island, MUSORSTOM 7, st. DW504, 14°20′S, 178°04′E, 300–390 m, SL 15.4 mm. E.Anticlinura sp.**, MNHN IM200742513, Solomon Islands, SALOMON 2, st. CP2182, 08°47.0′S, 159°37.9′E, 762–1060 m, SL 17.3 mm (radula: Fig. 8E). F.Eucithara cf. coronata (Hinds, 1843)**, MNHN IM200717900, Philippines, PANGLAO 2004, st. B8, 09°37.1′N, 123°46.1′E, 3 m, SL 8.7 mm (radula: Fig. 8D). G.Leiocithara sp., MNHN, New Caledonia, BATHUS 2, st. DW719, 22°48′S, 167°16′E, 444–445 m, SL 5.4 mm. H.Oenopota uschakovi Bogdanov, 1985, holotype, ZIN 37974/1, Iturup Island, Kurile Islands, 44°24.2′N, 147°09′01″E. 160 m, SL 10.3 mm. I.Genotina adamii (Bozzetti, 1994), MNHN, LIFOU 2000, st. DW1650, 20°54.15′S, 167°01.7′E, 120–150 m, SL 20.0 mm. J.Vitjazinella multicostata Sysoev, 1988, Paratype, ZMMU 22356, Izu-Bonin Trench, 33°18′N, 149°46′E., 6096 m, SL 7.2 mm. K.Oenopotella ultraabyssalis ultraabyssalis Sysoev, 1988, holotype, ZMMU Lc-22681, Kurile-Kamchatka trench, 49°29′N, 158°41′E, 7210–7230 m, SL 12.8 mm. L.Pseudorhaphitoma sp., Coral Sea, Chestefield Plateau, 19 53′S, 158 40′E, 410 m, SL 9.7 mm. M.Paraspirotropis simplicissima (Dall, 1907), ZIN, off Shikotan I., S Kurile Islands, 1450–1530 m, SL 28.0 mm. N.Heterocithara sp.**, MNHN IM200717884, Philippines, PANGLAO 2004, st. L46, 09°30.9′N, 123°41.2′E, 90–110 m, SL 3.4 mm. O.Toxicochlespira pagoda Sysoev & Kantor, 1990**, MNHN IM200717925, Solomon Islands, SALOMON 2, st. CP2227, 6°37.2′N, 156°12.7′E, 508–522 m, SL 18.0 mm (radula: Fig. 8B). Abbreviation and symbol: SL, shell length; **, sequenced specimen. Photo credits: B. Buge (F, O), P. Maestrati (B, D, G, I, L).

Figure 8.

Radulae. Mangeliidae. A.Benthomangelia trophonoidea (Schepman, 1913), MNHN IM200717835 (shell: Fig. 7C). B.Toxicochlespira pagoda Sysoev & Kantor, 1990**, MNHN IM200717925 (shell: Fig. 7O). C. Mangeliinae gen. 2. MNHN IM200910331, Philippines, PANGLAO 2004, st. S26, 9°41.50′N, 123°51.00′E, 21 m, SL unknown (broken). D.Eucithara cf. coronata (Hinds, 1843)**, MNHN IM200717900 (shell: Fig. 7F). E.Anticlinura sp.**, MNHN IM200742513 (shell: Fig. 7E). F. Mangeliidae gen. sp., MNHN, Philippines, PANGLAO 2004, st. T26, 9°43.3′N, 123°48.8′E, 123–135 m. G.Mangelia powisiana (Dautzenberg, 1887). Plymouth, England, after Taylor et al. (1993). Symbol: **, sequenced specimen.

Figure 8.

Radulae. Mangeliidae. A.Benthomangelia trophonoidea (Schepman, 1913), MNHN IM200717835 (shell: Fig. 7C). B.Toxicochlespira pagoda Sysoev & Kantor, 1990**, MNHN IM200717925 (shell: Fig. 7O). C. Mangeliinae gen. 2. MNHN IM200910331, Philippines, PANGLAO 2004, st. S26, 9°41.50′N, 123°51.00′E, 21 m, SL unknown (broken). D.Eucithara cf. coronata (Hinds, 1843)**, MNHN IM200717900 (shell: Fig. 7F). E.Anticlinura sp.**, MNHN IM200742513 (shell: Fig. 7E). F. Mangeliidae gen. sp., MNHN, Philippines, PANGLAO 2004, st. T26, 9°43.3′N, 123°48.8′E, 123–135 m. G.Mangelia powisiana (Dautzenberg, 1887). Plymouth, England, after Taylor et al. (1993). Symbol: **, sequenced specimen.

Diagnosis: Shell small to medium size, 3–30 mm, usually 6–12 mm, oval to low- or high-fusiform, usually with comparatively low spire, often with a shoulder angulation. Spiral and axial sculpture both well developed. Shell surface often bearing microsculpture of spirally aligned granules. Subsutural ramp usually not separated sculpturally. Anal sinus on subsutural ramp, shallow to rather deep, rarely tubular. Aperture normally not constrained with strong outgrowths or callus pads, rarely denticulate. Siphon rather short to moderately long. Protoconch typically multispiral, of up to five whorls, with axially ribbed protoconch II; spiral cords on protoconch II present or absent. When paucispiral, protoconch usually spirally lirate. Operculum present, with terminal nucleus (Oenopotinae), or normally absent. Radula of marginal teeth of very variable morphology. Teeth can be from semi-enrolled (Fig. 8F, G) to true hypodermic. Frequently side projections around the base are present and there is large irregularly shaped ‘root’ projecting from the base. Barbs may be present, from small to very large (Fig. 8C) or absent (Fig. 8B). Tooth canal opening laterally.

Included genera: Raphitomidae A. Bellardi, 1875

  • Acmaturris Woodring, 19281

  • Agathotoma Cossmann, 18992

  • Anticlinura Thile, 19343

    •  [= Clinuromella Beets, 1943]

  • Apispiralia Laseron, 19541

  • Apitua Laseron, 19541

  • Bactrocythara Woodring, 19281

  • Bela Gray, 18472

    •  [= Ginnania Monterosato, 1884]

    •  [= Smithiella Monterosato, 1890]

  • BellacytharaMcLean, 19712

  • Benthomangelia Thiele, 19253

  • Brachycythara Woodring, 19281

  • ?Cacodaphnella Pilsbry & Lowe, 19321

  • CitharomangeliaKilburn, 19922

  • Cryoturris Woodring, 18281

  • Curtitoma Bartsch, 19412

    •  [= Nematoma Bartsch, 1941]

    •  [= Widalli Bogdanov, 1986]

  • Cytharopsis A. Adams, 18651

  • Eucithara Fischer, 18833

    •  [= ?Cythara Schumacher, 1817]

  • ?Euryentmema Woodring, 19281

  • Fehria van Aartsen, 19881

  • Genotina Vera-Peláez, 20041

  • GingicitharaKilburn, 19922

  • Glyphoturris Woodring, 19281

  • Granotoma Bartsch, 19412

  • Granoturris Fargo, 19531

  • Guraleus Hedley, 19182

  •   (Euguraleus Cotton, 1947)

  • Hemicythara Kuroda & Oyama in Kuroda, Habe & Oyama, 19711

  • Heterocithara Hedley, 19223

  • Ithycythara Woodring, 19282

  • Kurtzia Bartsch, 19442

  • Kurtziella Dall, 19182

  • Kurtzina Bartsch, 19442

  • Leiocithara Hedley, 19221

  • Liracraea Odhner, 19242

  • ?Macteola Hedley, 19181

  • Mangelia Risso, 18262

    •  [= Cyrtocythara Nordsieck, 1977]

    •  [= Cytharella Monterosato, 1875]

    •  [= Lyromangelia Nordsieck, 1977]

    •  [= Mangiliella Bucqoy, Dautzenberg & Dollfus, 1883]

    •  [= Rissomangelia Monterosato, 1917]

    •  [= Rugocythara Nordsieck, 1977]

    •  [= Villiersiella Monterosato, 1884]

  • Marita Hedley, 19221

  • Mitraguraleus Laseron, 19541

  • ?Neoguraleus Powell, 19392

  • ?Notocytharella Hertlein & Strong, 19551

  • Obesotoma Bartsch, 19412

  • Oenopota Mörch, 18523

    •  (Nodotoma Bartsch, 1941)

  • Oenopotella Sysoev, 19881

  • PapillocitharaKilburn, 19921

  • ?Paraspirotropis Sysoev & Kantor, 19842

  • Perimangelia McLean, 20001

  • Platycythara Woodring, 19281

  • Propebela Iredale, 19182

    •  [= AntiguraleusPowell, 1942]

    •  [= ?Belalora Powell, 1951]

    •  [= Cestoma Bartsch, 1941]

    •  [= Funitoma Bartsch, 1941]

    •  [= ?Lorabela Powell, 1951]

    •  [= Turritomella Bartsch, 1941]

  •   (Canetoma Bartsch, 1941)

  • Pseudorhaphitoma Boettger, 18952

  • Pyrgocythara Woodring, 19282

  • Rubellatoma Bartsch & Rehder, 19391

  • Saccharoturris Woodring, 19281

  • Stellatoma Bartsch & Rehder, 19391

  • Suturocythara García, 20081

  • Tenaturris Woodring, 19282

  • Toxicochlespira Sysoev & Kantor, 19903

  • ?Venustoma Bartsch, 19411

  • Vitjazinella Sysoev, 19881

  • ?Vitricythara Fargo, 19531

(Figs 9, 10)

Figure 9.

Shells. Raphitomidae. A.Hemilienardia calcicincta (Melvill & Standen, 1895)*, Philippines, PANGLAO 2004, st. B14, 09°38.5′N, 123°49.2′E, 2–4 m, SL 4.6 mm. B.Hemilienardia malleti (Recluz, 1852), MNHN, New Caledonia, MONTROUZIER, st. 1245, 20°45.2′S, 165°16.3′E, intertidal, SL 4.4 mm. C.Rimosodaphnella sp.**, MNHN IM200717836, Vanuatu, BOA1, st. CP2462, 16°37.5′S, 167°57.4′E, 618–641 m, SL 15.1 mm. D.Veprecula sp.**, MNHN IM200717883, Philippines, PANGLAO 2004, st. L46, 09°30.9′N, 123°41.2′E, 90–110 m, SL 3.4 mm. E.Glyphostomoides sp.**, MNHN IM200717892, Philippines, PANGLAO 2004, st. T39, 09°30′N, 123°50.4′E, 100–138 m, SL 7.0 mm. F.Aliceia sp., MNHN, Wallis Islands, MUSORSTOM 7, st. DW604, 13°21′S, 176°08′W, 415–420 m, SL 4.0 mm. G.Thatcheriasyrinx orientis (Melvill, 1904), MNHN, New Caledonia, 20°49′S, 165°19′E, 105–110 m, SL 3.6 mm. H.Tritonoturris cumingii (Powys in Powys & Sowerby, 1835), MNHN, New Caledonia, SL 20 mm. I.Rocroithys niveus Sysoev & Bouchet, 2001, holotype, MNHN, Vanuatu, 15°08′S 167°12′E, 312–314 m, SL 25.5 mm. J.Pleurotomella sp.**, MNHN IM200717848, Coral Sea, EBISCO, st. DW2625, 20°05′S, 160°19′E, 627–741 m, SL 20.6 mm. K.Taranis sp.**, MNHN IM200742296, Philippines, AURORA 07, st. CP2749, 15°56.6′N, 121°50.2′E, 473 m, SL 6 mm. A07 1717. L.Nepotilla sp., MNHN, New Caledonia, SMIB8, st. DW146–147, 24°55′S, 168°22′, 508–532 m, SL 1.5 mm. M.Eucyclotoma cymatodes (Hervier, 1899)**, MNHN IM200717903, Philippines, PANGLAO 2004, st. S12, 09°29.4′N, 123°56.0′E, 6–8 m, SL 5.0 mm. N.Buccinaria pendula Bouchet & Sysoev, 1997, MNHN, New Caledonia, BATHUS 1, st. DE697, 20°34′S, 164°58′E, 570–650 m, SL 7.2 mm. O.Microdaphne morrisoni Rehder, 1980, MNHN, New Caledonia, SL 4.5 mm. P.Cryptodaphne sp., MNHN, New Caledonia, MUSORSTOM 4, st. DW150, 19°07′S, 163°22′E, 110 m, SL 11.2 mm. Q.Kermia aureotincta (Hervier, 1897)**, MNHN IM200717878, Philippines, PANGLAO 2004, st. B25, 09°29′N, 123°56.1′E, 16 m, SL 4.5 mm. R.Teretiopsis cf. hyalina Sysoev & Bouchet, 2001**, MNHN IM200717845, Coral Sea, EBISCO, st. CP2651, 21°29′S, 162°36′E, 883–957 m, SL 22.8 mm. S.Daphnella itonis Sysoev & Bouchet, 2001, holotype, MNHN, New Caledonia, 22°48′S 167°16′E, 444–445 m, SL 66.0 mm. T.Phymorhynchus major Warén & Bouchet, 2001, holotype, MNHN, Site Barbecue, East Pacific Rise, 09°50′N, 104°17′W, 2505 m, SL 72.0 mm. U.Gymnobela sp.**, MNHN IM200717841, Coral Sea, EBISCO, st. CP2648, 21°32′S, 162°30′E, 750–458 m, SL 25.7 mm. V.Thatcheria mirabilis Angas, 1877**, MNHN IM200717924, Solomon Islands, SALOMON 2, st. CP2184, 08°16.9′S, 159°59.7′E, 464–523 m, SL 32.5 mm. Abbreviation and symbols: SL, shell length; *, sequenced species; **, sequenced specimen. Photo credits: B. Buge (C–E, J, Q, R, U, V), P. Maestrati (F, G, L, P).

Figure 9.

Shells. Raphitomidae. A.Hemilienardia calcicincta (Melvill & Standen, 1895)*, Philippines, PANGLAO 2004, st. B14, 09°38.5′N, 123°49.2′E, 2–4 m, SL 4.6 mm. B.Hemilienardia malleti (Recluz, 1852), MNHN, New Caledonia, MONTROUZIER, st. 1245, 20°45.2′S, 165°16.3′E, intertidal, SL 4.4 mm. C.Rimosodaphnella sp.**, MNHN IM200717836, Vanuatu, BOA1, st. CP2462, 16°37.5′S, 167°57.4′E, 618–641 m, SL 15.1 mm. D.Veprecula sp.**, MNHN IM200717883, Philippines, PANGLAO 2004, st. L46, 09°30.9′N, 123°41.2′E, 90–110 m, SL 3.4 mm. E.Glyphostomoides sp.**, MNHN IM200717892, Philippines, PANGLAO 2004, st. T39, 09°30′N, 123°50.4′E, 100–138 m, SL 7.0 mm. F.Aliceia sp., MNHN, Wallis Islands, MUSORSTOM 7, st. DW604, 13°21′S, 176°08′W, 415–420 m, SL 4.0 mm. G.Thatcheriasyrinx orientis (Melvill, 1904), MNHN, New Caledonia, 20°49′S, 165°19′E, 105–110 m, SL 3.6 mm. H.Tritonoturris cumingii (Powys in Powys & Sowerby, 1835), MNHN, New Caledonia, SL 20 mm. I.Rocroithys niveus Sysoev & Bouchet, 2001, holotype, MNHN, Vanuatu, 15°08′S 167°12′E, 312–314 m, SL 25.5 mm. J.Pleurotomella sp.**, MNHN IM200717848, Coral Sea, EBISCO, st. DW2625, 20°05′S, 160°19′E, 627–741 m, SL 20.6 mm. K.Taranis sp.**, MNHN IM200742296, Philippines, AURORA 07, st. CP2749, 15°56.6′N, 121°50.2′E, 473 m, SL 6 mm. A07 1717. L.Nepotilla sp., MNHN, New Caledonia, SMIB8, st. DW146–147, 24°55′S, 168°22′, 508–532 m, SL 1.5 mm. M.Eucyclotoma cymatodes (Hervier, 1899)**, MNHN IM200717903, Philippines, PANGLAO 2004, st. S12, 09°29.4′N, 123°56.0′E, 6–8 m, SL 5.0 mm. N.Buccinaria pendula Bouchet & Sysoev, 1997, MNHN, New Caledonia, BATHUS 1, st. DE697, 20°34′S, 164°58′E, 570–650 m, SL 7.2 mm. O.Microdaphne morrisoni Rehder, 1980, MNHN, New Caledonia, SL 4.5 mm. P.Cryptodaphne sp., MNHN, New Caledonia, MUSORSTOM 4, st. DW150, 19°07′S, 163°22′E, 110 m, SL 11.2 mm. Q.Kermia aureotincta (Hervier, 1897)**, MNHN IM200717878, Philippines, PANGLAO 2004, st. B25, 09°29′N, 123°56.1′E, 16 m, SL 4.5 mm. R.Teretiopsis cf. hyalina Sysoev & Bouchet, 2001**, MNHN IM200717845, Coral Sea, EBISCO, st. CP2651, 21°29′S, 162°36′E, 883–957 m, SL 22.8 mm. S.Daphnella itonis Sysoev & Bouchet, 2001, holotype, MNHN, New Caledonia, 22°48′S 167°16′E, 444–445 m, SL 66.0 mm. T.Phymorhynchus major Warén & Bouchet, 2001, holotype, MNHN, Site Barbecue, East Pacific Rise, 09°50′N, 104°17′W, 2505 m, SL 72.0 mm. U.Gymnobela sp.**, MNHN IM200717841, Coral Sea, EBISCO, st. CP2648, 21°32′S, 162°30′E, 750–458 m, SL 25.7 mm. V.Thatcheria mirabilis Angas, 1877**, MNHN IM200717924, Solomon Islands, SALOMON 2, st. CP2184, 08°16.9′S, 159°59.7′E, 464–523 m, SL 32.5 mm. Abbreviation and symbols: SL, shell length; *, sequenced species; **, sequenced specimen. Photo credits: B. Buge (C–E, J, Q, R, U, V), P. Maestrati (F, G, L, P).

Figure 10.

Radulae. Raphitomidae. A.Buccinaria pendula Bouchet & Sysoev, 1997, MNHN, New Caledonia, BATHUS 4, st. CP948, 20°33′S, 164°57′E, 533–610 m. B.Daphnella pulvisculus Chino, 2006, MNHN, New Caledonia, BATHUS 4, st. DW927, 18°56′S, 163°22′E, 444–452 m. C.Gymnobela yoshidai Kuroda & Habe in Habe, 1962, MNHN, Norfolk ridge, NORFOLK 2, st. DW2058, 24°40′S, 168°40′E, 591–1032 m. D.Kermia irretita (Hedley, 1899), MNHN, New Caledonia, LIFOU 2000, st. 1419, 20°55.6′S, 167°04.5′E, 0–5 m. E.Daphnella cladara Sysoev & Bouchet, 2001, MNHN, Norfolk Ridge, LITHIST, st. CP09, 24°53′S, 168°22′E, 518–540 m. F.Daphnella mitrellaformis (Nomura, 1940), MNHN, New Caledonia, LIFOU 2000, st. DW1649, 20°54′S, 167°01′E, 150–200 m. G.Miowateria sp., MNHN, Fidji, MUSORSTOM 10, st. CP1354, 17°43′S, 178°55′E, 959–963 m. H.Spergo fusiformis (Kuroda & Habe in Habe, 1962), MNHN, Tongatapu, Tonga, BORDAU 2, st. CP1566, 21°02′S, 175°18′W, 530–531 m.

Figure 10.

Radulae. Raphitomidae. A.Buccinaria pendula Bouchet & Sysoev, 1997, MNHN, New Caledonia, BATHUS 4, st. CP948, 20°33′S, 164°57′E, 533–610 m. B.Daphnella pulvisculus Chino, 2006, MNHN, New Caledonia, BATHUS 4, st. DW927, 18°56′S, 163°22′E, 444–452 m. C.Gymnobela yoshidai Kuroda & Habe in Habe, 1962, MNHN, Norfolk ridge, NORFOLK 2, st. DW2058, 24°40′S, 168°40′E, 591–1032 m. D.Kermia irretita (Hedley, 1899), MNHN, New Caledonia, LIFOU 2000, st. 1419, 20°55.6′S, 167°04.5′E, 0–5 m. E.Daphnella cladara Sysoev & Bouchet, 2001, MNHN, Norfolk Ridge, LITHIST, st. CP09, 24°53′S, 168°22′E, 518–540 m. F.Daphnella mitrellaformis (Nomura, 1940), MNHN, New Caledonia, LIFOU 2000, st. DW1649, 20°54′S, 167°01′E, 150–200 m. G.Miowateria sp., MNHN, Fidji, MUSORSTOM 10, st. CP1354, 17°43′S, 178°55′E, 959–963 m. H.Spergo fusiformis (Kuroda & Habe in Habe, 1962), MNHN, Tongatapu, Tonga, BORDAU 2, st. CP1566, 21°02′S, 175°18′W, 530–531 m.

Diagnosis: Shell very variable in shape and size, from buccinoid to ovate or to elongate-fusiform or high-cylindrical and from about 2 to over 140 mm high. Sculpture variously developed: from nearly smooth shell to well-developed spiral and axial elements, subsutural ramp, when separated from rest of the whorl, usually smooth or with traces of anal sinus growth. Apertural armature rarely well expressed, inner lip usually smooth. Anal sinus either subsutural, shaped as a ‘reversed-L’, or on subsutural ramp, asymmetric, very shallow to rather deep, typically not constrained but sometimes even almost tubular. Siphon short to rather long. Protoconch typically planktotrophic, multispiral, of 2.5–6.5 whorls, protoconch I often spirally striated, protoconch II with diagonally cancellated sculpture. There are many variations of the basic pattern of the protoconch II sculpture, though a combination of spiral and axial elements is retained, with the latter usually more pronounced, but protoconchs with a spiral keel and axial pillars (Fig. 23) depart from this pattern. Paucispiral protoconch with predominantly spiral striation. No operculum. Radula of hypodermic marginal teeth of very variable morphology. Teeth can be relatively very long (to 12.8% of shell length in Daphnella cladara, Fig. 10E) to very short and reduced (0.13% of shell length in Spergo fusiformis, Fig. 10H). Solid base of the teeth from very small (Fig. 10E, G) to rather large (Fig. 10B, F), sometimes side projections around the base are present (Fig. 10F). Barb(s) present or absent. At least in one species cockscomb-shaped structure is present near the tooth tip (Fig. 10G). Tooth canal opening laterally or subapically. Reduction in radular apparatus and other structures of the anterior foregut is frequent, grading to complete absence.

Remarks: This is the largest and most variable taxon in the Conoidea, as concerns the number of species, with the largest vertical range (intertidal to hadal depths).

Included genera: CochlespiridaePowell, 1942

  • Abyssobela Kantor & Sysoev, 19861

  • ?Abyssothauma Sysoev, 19961

  • Acamptodaphne Shuto, 19711

  • Acanthodaphne Bonfitto & Morassi, 20061

  • Aliceia Dautzenberg & Fischer, 18971

  • Asperdaphne Hedley, 19221

    •  (Aspertilla Powell, 1944)

  • Austrodaphnella Laseron, 19541

  • ?Austropusilla Laseron, 19541

    •  (Metaclathurella Shuto, 1983)

  • Bathybela Kobelt, 19052

    •  [= Bathypota Nordsieck, 1968]

  • Buccinaria Kittl, 18872

    •  [= Ootomella Bartsch, 1933]

    •  [= Pionotoma Kuroda, 1952]

  • Cenodagreutes E.H. Smith, 19671

  • Clathurina Melvill, 19171

  • ClinuraBellardi, 18751

  • CryptodaphnePowell, 19421

  • Daphnella Hinds, 18443

    •  [= Eudaphnella Bartsch, 1933]

    •  [= Paradaphne Laseron, 1954]

    •  (Diaugasma Melvill, 1917)

    •  (Hemidaphne Hedley, 1918)

  • Eubela Dall, 18891

  • Eucyclotoma Boettger, 18953

    •  [= Turrhyssa Dall, 1924]

  • ?Exomilus Hedley, 19181

  • Famelica Bouchet & Warén, 19801

  • Favriella Hornung, 19201

  • Glyphostomoides Shuto, 19833

  • Gymnobela Verrill, 18843

    •  [= Majox Nordsieck, 1968]

    •  [= Watsonaria Nordsieck, 1968]

    •  (Theta Clarke, 1959)

  • Hemilienardia Boettger, 18953

  • Isodaphne Laseron, 19541

  • Kermia Olivier, 19153

  • Kuroshiodaphne Shuto, 19651

  • Leiosyrinx Bouchet & Sysoev, 20012

  • Lusitanops Nordsieck, 19681

  • Magnella Dittmer, 19601

  • MicrodaphneMcLean, 19711

  • Microgenia Laseron, 19541

  • Mioawateria Vella, 19541

  • Neopleurotomoides Shuto, 19711

  • Nepotilla Hedley, 19182

  • Pagodidaphne Shuto, 19831

  • Paramontana Laseron, 19542

  • Philbertia Monterosato, 18842

    •  [= Leufroyia Monterosato, 1884]

    •  [= Lineotoma Nordsieck, 1977]

    •  [= Peratotoma Harris & Burrows, 1891]

  • Phymorhynchus Dall, 19082

  • Pleurotomella Verrill, 18733

    •  [= Azorilla Nordsieck, 1968]

    •  [= Azorita Nordsieck, 1968]

    •  [= AnomalotomellaPowell, 1966]

    •  [= Fusidaphne Laseron, 1954]

    •  [= Tasmadaphne Laseron, 1954]

  •   (Cyrtoides Nordsieck, 1968)

  • Pontiothauma E.A. Smith, 18952

  • Pseudodaphnella Boettger, 18952

    •  (Qii Zhang, 1995)

  • Raphitoma Bellardi, 18483

  • Rimosodaphnella Cossmann, 19153

  • Rocroithys Sysoev & Bouchet, 20012

  • Spergo Dall, 18952

    •  (Speoides Kuroda & Habe in Habe, 1962)

  • Stilla Finlay, 19261

  • Taranidaphne Morassi & Bonfitto, 20011

  • Taranis Jeffreys, 18673

    •  [= ?Allo Lamy, 1934]

    •  [= ?Feliciella Lamy, 1934]

    •  [= Fenestrosyrinx Finlay, 1926]

  • ?Teleochilus Harris, 18971

    •  [= Litachilus Powell, 1944]

  • Teretia Norman, 18882

  • Teretiopsis Kantor & Sysoev, 19893

  • Thatcheria Angas, 18773

    •  [= Cochlioconus Yokoyama, 1928]

  • ?Thatcheriasyrinx Powell, 19691

  • Thatcherina Vera-Peláez, 19981

  • ?Thesbia Jeffreys, 18672

  • Tritonoturris Dall, 19243

  • TruncadaphneMcLean, 19711

  • Tuskaroria Sysoev, 19881

  • Typhlosyrinx Thiele, 19252

  • Veprecula Melvill, 19173

    •  [= Mordica Dall, 1924]

  • Vepridaphne Shuto, 19831

  • XanthodaphnePowell, 19422

  • Zenepos Finlay, 19281

(Figs 11A–D, 12)

Figure 11.

Shells. A–D. Cochlespiridae. A.Nihonia maxima Sysoev, 1997, holotype, MNHN, Indonesia, 09°23′S 131°09′E, 275–246 m, SL 128 mm. B.Sibogasyrinx sp.**, MNHN IM200717701, Vanuatu, BOA1, st. CP2432, 14°59.7′S, 166°55.0′E, SL 46.0 mm (radula: Fig. 12В, С). C.Aforia multispiralis Dell, 1990, USNM 870023, N of Livingston Island, South Shetland Islands, 62°S, 61.1°W, 1437 m, SL 45.6 mm. D.Cochlespira pulchella (Schepman, 1913)**, MNHN IM200717920, PANGLAO 2005, st. CP2340, 09°29.4′N, 123°44.4′E, SL 22.0 mm. E. Strictispiridae. E.Strictispira solida (C.B. Adams, 1850), USNM 900425, E side of Marquesas Keys, Florida Keys, SL 17.5 mm. F–L. Drilliidae. F.Plagiostropha caledoniensis Wells, 1995, S New Caledonia, SL 12.2 mm. G.Imaclava unimaculata (Sowerby I, 1834)* subadult, det. J.A. Todd, NHMUK MOEA 20100529, Gulf of Panama, JTD-00–46, 08°31.37′N, 79°05.79′W, 24–25 m, SL 25.5 mm. H.Clavus canalicularis (Röding, 1798)*, MNHN IM200742345, Philippines, PANGLAO 2004, st. B11, 09°29.4′N, 123°56.0′E, 2–4 m, SL 19 mm. I.Cruziturricula arcuata (Reeve, 1843)* subadult, det. J. A. Todd, YK; NHMUK 20100542, Gulf of Panama, JTD-00–34, 08°26.24′N, 79°09.14′W, 66–68 m, SL = 17.6 mm (radula: Fig. 13E). J.Conopleura striata Hinds, 1844*, New Caledonia, MONTROUZIER, st. 1311, 20°40.4′S, 164°14.9′E, 10–60 m, SL 15.6 mm. K.Iredalea pupoidea (H. Adams, 1872)*, MNHN, New Caledonia, MONTROUZIER, st. 1240, 20°46.5′S, 165°14′E-165°15′E, 2 m, SL 6.2 mm. L.Splendrillia sp.**, MNHN IM200717847, Coral Sea, EBISCO, st. DW2617, 20°06′S, 160°22′E, 427–505 m, SL 16.4 mm (radula: Fig. 13A, B). Abbreviation and symbols: SL, shell length; *, sequenced species; **, sequenced specimen. Photo credits: B. Buge (B, D, F), A. Fedosov (I, G), M.G. Harasewych and D. Tippett (C, E),

Figure 11.

Shells. A–D. Cochlespiridae. A.Nihonia maxima Sysoev, 1997, holotype, MNHN, Indonesia, 09°23′S 131°09′E, 275–246 m, SL 128 mm. B.Sibogasyrinx sp.**, MNHN IM200717701, Vanuatu, BOA1, st. CP2432, 14°59.7′S, 166°55.0′E, SL 46.0 mm (radula: Fig. 12В, С). C.Aforia multispiralis Dell, 1990, USNM 870023, N of Livingston Island, South Shetland Islands, 62°S, 61.1°W, 1437 m, SL 45.6 mm. D.Cochlespira pulchella (Schepman, 1913)**, MNHN IM200717920, PANGLAO 2005, st. CP2340, 09°29.4′N, 123°44.4′E, SL 22.0 mm. E. Strictispiridae. E.Strictispira solida (C.B. Adams, 1850), USNM 900425, E side of Marquesas Keys, Florida Keys, SL 17.5 mm. F–L. Drilliidae. F.Plagiostropha caledoniensis Wells, 1995, S New Caledonia, SL 12.2 mm. G.Imaclava unimaculata (Sowerby I, 1834)* subadult, det. J.A. Todd, NHMUK MOEA 20100529, Gulf of Panama, JTD-00–46, 08°31.37′N, 79°05.79′W, 24–25 m, SL 25.5 mm. H.Clavus canalicularis (Röding, 1798)*, MNHN IM200742345, Philippines, PANGLAO 2004, st. B11, 09°29.4′N, 123°56.0′E, 2–4 m, SL 19 mm. I.Cruziturricula arcuata (Reeve, 1843)* subadult, det. J. A. Todd, YK; NHMUK 20100542, Gulf of Panama, JTD-00–34, 08°26.24′N, 79°09.14′W, 66–68 m, SL = 17.6 mm (radula: Fig. 13E). J.Conopleura striata Hinds, 1844*, New Caledonia, MONTROUZIER, st. 1311, 20°40.4′S, 164°14.9′E, 10–60 m, SL 15.6 mm. K.Iredalea pupoidea (H. Adams, 1872)*, MNHN, New Caledonia, MONTROUZIER, st. 1240, 20°46.5′S, 165°14′E-165°15′E, 2 m, SL 6.2 mm. L.Splendrillia sp.**, MNHN IM200717847, Coral Sea, EBISCO, st. DW2617, 20°06′S, 160°22′E, 427–505 m, SL 16.4 mm (radula: Fig. 13A, B). Abbreviation and symbols: SL, shell length; *, sequenced species; **, sequenced specimen. Photo credits: B. Buge (B, D, F), A. Fedosov (I, G), M.G. Harasewych and D. Tippett (C, E),

Figure 12.

Radulae. Cochlespiridae. A.Cochlespira radiata (Dall, 1889), MNHN, SE Brazil, after Kantor & Taylor, 2000. B–C.Sibogasyrinx sp.**, MNHN IM200717701 (shell: Fig. 11B). Symbol: **, sequenced specimen.

Figure 12.

Radulae. Cochlespiridae. A.Cochlespira radiata (Dall, 1889), MNHN, SE Brazil, after Kantor & Taylor, 2000. B–C.Sibogasyrinx sp.**, MNHN IM200717701 (shell: Fig. 11B). Symbol: **, sequenced specimen.

Diagnosis: Shell of moderate size, about 20–30 mm, up to 100 mm high, high-pagodiform to fusiform, with a tall spire and usually a long siphonal canal. Axial sculpture poorly developed or absent, subsutural ramp usually smooth. Anal sinus deep, on subsutural ramp. Protoconch paucispiral, smooth. Operculum with terminal nucleus. Radula 1–0-R-0–1. Rachidian broad, subrectangular or arched, with a single rather large cusp (Fig. 12), rarely absent (some Aforia). Marginal teeth duplex, with well developed accessory limb.

Remarks: This is a clade with poor congruence between molecular and shell characters. The contents of most genera need revision and the family limits remain uncertain.

Sibogasyrinx was proposed as a subgenus of Leucosyrinx for two species characterized by a low position of the peripheral angle. During our examination of radulae of species provisionally attributed to Leucosyrinx, two distinct radulae types were found, differing in the presence of the rachidian, without apparent congruence with shell morphology. In Sibogasyrinx pyramidalis, the type species of Sibogasyrinx, as well as in the sequenced specimen (Fig. 11B), the radula is characterized by the presence of a well-developed rachidian. Although the sequenced specimen is conchologically closer to typical Leucosyrinx, the molecular sequence suggests affinities to Cochlespira.

Included genera: Drilliidae Olsson, 1964

  • ?Aforia Dall, 18892

  •   [= Irenosyrinx Dall, 1908]

  •   [= Danilacarina Bozzetti, 1997]

    •  (?Abyssaforia Sysoev & Kantor, 1987)

    •  (?Dallaforia Sysoev & Kantor, 1987)

    •  (?Steiraxis Dall, 1896)

  • ?Apiotoma Cossmann, 18891

  • ?Clavosurcula Schepman, 19131

  • Cochlespira Conrad, 18653

  •   [= Ancistrosyrinx Dall, 1881]

  •   [= Coronasyrinx Powell, 1944]

  •   [= Pagodasyrinx Shuto, 1969]

  • ?Nihonia MacNeil, 19611

  •  Sibogasyrinx Powell, 19693

(Figs 11F–L, 13)

Figure 13.

Radulae. Drilliidae. A–B.Splendrillia sp.**, MNHN IM200717847 (shell: Fig. 11L). C.Clavus exasperatus (Reeve, 1843), MNHN, New Caledonia, LIFOU 2000, st. 1420, 20°47.7′S, 167°09.35′E, 4–5 m. D.Imaclava pilsbryi (Bartsch, 1950), after Kantor & Taylor, 2000. E.Cruziturricula arcuata (Reeve, 1843)**, NHMUK MOEA 20100541, Gulf of Panama, JTD-00–34, 08°26.24′N, 79°09.14′W, 66–68 m (shell: Fig. 11I). Symbol: **, sequenced specimen.

Figure 13.

Radulae. Drilliidae. A–B.Splendrillia sp.**, MNHN IM200717847 (shell: Fig. 11L). C.Clavus exasperatus (Reeve, 1843), MNHN, New Caledonia, LIFOU 2000, st. 1420, 20°47.7′S, 167°09.35′E, 4–5 m. D.Imaclava pilsbryi (Bartsch, 1950), after Kantor & Taylor, 2000. E.Cruziturricula arcuata (Reeve, 1843)**, NHMUK MOEA 20100541, Gulf of Panama, JTD-00–34, 08°26.24′N, 79°09.14′W, 66–68 m (shell: Fig. 11I). Symbol: **, sequenced specimen.

Diagnosis: Shell small- to medium-sized, usually 15–25 mm, up to 50 mm high, with a rather high spire and usually truncated base. Spiral sculpture often obsolete. Anal sinus on subsutural ramp, deep, (sub)symmetrical, sometimes tubular. Protoconch usually paucispiral (up to two whorls), smooth or abapically carinate. Operculum with terminal nucleus. Radular formula 1-1-R-1-1, rarely 1-1-0-1-1, central tooth small, from narrow unicuspid (Fig. 13B) to subrectangular with additional cusps (Fig. 13C), rarely reduced to completely absent. Lateral teeth broad, pectinate, and arched, marginal teeth from simple flat and sharply pointed (Fig. 13A) to duplex with slightly thickened edges and to loosely enrolled with the small barb near the tip (Imaclava, Fig. 13D).

Remarks: The genera Cruziturricula and Fusiturricula form an unsupported group that is sister to Drilliidae. In the studied Cruziturricula arcuata (Reeve, 1843), the very characteristic radula differs from the Drilliidae: 1-0-0-0-1. Marginal teeth are loosely enrolled with little overlap of the edges, with two barbs on the tip and a tongue-shape extension at the base (Fig. 13E). The type species of Fusiturricula, Turris fusinella Dall, 1908, is different from what is currently conceived as belonging to that genus (e.g. Williams, 2006), but those species are similar to Cruziturricula sensu auctt. The only data on the radula was provided by Powell (1966: 31), who stated (without mentioning the species): “Radula of marginals only, wishbone-type, but long and narrow”. Although Cruziturricula and Fusiturricula definitely do not belong in Drilliidae and may represent a new family, they are here provisionally placed in Drilliidae for lack of a better alternative.

Included genera: PseudomelatomidaeMorrison, 1965

  • ?AcinodrilliaKilburn, 19881

  • Agladrillia Woodring, 19283

  • Bellaspira Conrad, 18681

  • CalliclavaMcLean, 19713

  • Cerodrillia Bartsch & Rehder, 19393

    •  (Lissodrillia Barsch and Rehder, 1943)

  • Clavus Monfort, 18103

    •  [= Clavicantha Swainson, 1840]

    •  [= Eldridgea Bartsch, 1934]

    •  [= Tylotia Melvill, 1917]

  • Clathrodrillia Dall, 19183

  • Conopleura Hinds, 18443

  • Crassopleura Monterosato, 18841

  • ?Cruziturricula Marks, 19513

  • Cymatosyrinx Dall, 18892

  • Douglassia Bartsch, 19341

  • Drillia Gray, 18383

  • Elaeocyma Dall, 19182

  • Eumetadrillia Woodring, 19281

  • Fenimorea Bartsch, 19341

  • ?Fusiturricula Woodring, 19283

    •  (?Fusisyrinx Bartsch, 1934)

  • Globidrillia Woodring, 19282

  • Imaclava Bartsch, 19443

  • Iredalea Oliver, 19153

    •  [= Brephodrillia Pilsbry & Lowe, 1932]

  • Kylix Dall, 19192

  • Leptadrillia Woodring, 19282

  • Neodrillia Bartsch, 19432

  • OrrmaesiaKilburn, 19882

  • ?Paracuneus Laseron, 19541

  • Plagiostropha Melvill, 19272

  • Splendrillia Hedley, 19223

  • Spirotropis G.O. Sars, 18782

  • ?Stenodrillia Korobkov, 19551

  • Syntomodrillia Woodring, 19282

  • Tylotiella Habe, 19642

(Figs 14–16)

Figure 14.

Shells. Pseudomelatomidae. A.Comitas sp.**, MNHN IM200717918, Philippines, PANGLAO 2005, st. CP2388, 09°26.9′N, 123°34.5′E, 762–786 m, SL 113.8 mm (radula: Fig. 15C, D). B.Knefastia tuberculifera (Broderip & Sowerby, 1829)*, NHMUK MOEA 20100534, Gulf of Panama, JTD-00-18, 08°19.50′N, 78°47.71′W, 25–32 m, SL 36.6 mm. C.Otitoma sp.**, MNHN IM200717905, Philippines, PANGLAO 2005, st. CP2348, 09°26.6′N, 123°52.5′E, 196–216 m, SL 17.2 mm. D.Pyrgospira aenone (Dall, 1919)**, det. J. A. Todd, YK, NHMUK MOEA 20100539, Gulf of Panama, JTD-00–18, 08°19.50′N, 78°47.71′W, 25–32 m. SL 36.2 mm. E.Inquisitor sp.**, MNHN IM200717851, Coral Sea, EBISCO, st. DW2625, 20°05′N, 160°19′E, 627–741 m, SL 32.2 mm (radula: Fig. 16F). F.Funa incerta (Smith, 1877)**, NHMUK MOEA 20100553, st. 70, off southern Hong Kong, SL 42.8 mm. G.Leucosyrinx sp.**, MNHN IM200717846, Coral Sea, EBISCO, st. CP2600, 19°38′S, 158°46′E, 603–630 m, SL 27.1 mm (radula: Fig. 16A). H.Hindsiclava alesidota (Dall, 1889)**, det. J. A. Todd, NHMUK MOEA 20100524, Lower Florida Keys, JTD-01-15, 24°33.47′N, 81°07.72′W, 117–148 m SL 37.9 mm. I.Conorbela antarctica (Strebel, 1908) USNM 894465, Zavodovski Island, South Sandwich Islands, 56.27°S, 27.5°W, 208–375 m, SL 23.6 mm. J.Carinodrillia dichroa Pilsbry & Lowe, 1932*, det. J. A. Todd, NHMUK MOEA 20100531, Gulf of Panama, JTD-00-18, 08°19.50′N, 78°47.71′W, 25–32 m, SL 23.0 mm (radula: Fig. 16C). K.Cheungbeia robusta (Hinds, 1839)*, NHMUK MOEA 20100558, st. 70, off southern Hong Kong, South China Sea, SL 21.2 mm (radula: Fig. 16E). L.Plicisyrinx vitjazi Sysoev t Kantor, 1986, holotype, ZMMU Lc-22382, E of northern Honshu, Japan, 40°55.2′N, 144°53.3′E, 3880–4000 m, SL 13.9 mm. M.Maesiella maesae McLean & Poorman, 1971, LACM 1513, holotype, Guaymas, Sonora, Mexico, SL 9.2 mm. N.Kurilohadalia elongata Sysoev & Kantor, 1986, holotype, ZMMU Lc-22401, Kurile-Kamchatka Trench, 43°55′ N, 149°47′ E, 8080–8160 m, SL 12.5 mm. O.Crassispira quadrilirata (E.A. Smith, 1882)**, MNHN IM200717755, Philippines, PANGLAO 2004, st. L46, 09°30.9′N, 123°41.2′E, 90–110 m, SL 14.0 mm. P.Epideira sibogae (Schepman, 1913), MNHN, Vanuatu, MUSORSTOM 8, st. DW1097, 15°05′S, 167°11′E, 281–288 m, SL 24.1 mm. Q.Tiariturris spectabilis Berry, 1958**, det. J. A. Todd, NHMUK MOEA 20100540, Gulf of Panama, JTD-00-34, 08°26.24′N, 79 09.14′W, 66–68 m SL 48.7 mm (radula: Fig. 15A). R.Pseudomelatoma penicillata Carpenter, 1864, USNM 130590, California, SL 28.1 mm. S.Megasurcula carpenteriana (Gabb, 1865), USNM 523713, California, SL 84.1 mm. T.Antiplanes sanctiioannis (E.A. Smith, 1875), ZMMU Lc-17096, off northern Kurile Islands, 49°40.9′N, 154°41′E, 142 m, SL 37.2 mm. U.Naudedrillia sp., New Caledonia, VAUBAN, st. 3, 22°17′S, 167 °12′E, 390 m, SL 19.0 mm. V.Zonulispira sp.*, NHMUK MOEA 20100537, det. J. A. Todd, Gulf of Panama, JTD-00-18, 08°19.50′N, 78°47.71′W, 25–32 m, SL 21.2 mm (radula: Fig. 16B). Abbreviation and symbols: SL, shell length; *, sequenced species; **, sequenced specimen. Photo credits: B. Buge (A, G), A. Fedosov (B–F, H, J, K, Q), M.G. Harasewych and D. Tippett (R, S), P. Maestrati (U), J. McLean (M).

Figure 14.

Shells. Pseudomelatomidae. A.Comitas sp.**, MNHN IM200717918, Philippines, PANGLAO 2005, st. CP2388, 09°26.9′N, 123°34.5′E, 762–786 m, SL 113.8 mm (radula: Fig. 15C, D). B.Knefastia tuberculifera (Broderip & Sowerby, 1829)*, NHMUK MOEA 20100534, Gulf of Panama, JTD-00-18, 08°19.50′N, 78°47.71′W, 25–32 m, SL 36.6 mm. C.Otitoma sp.**, MNHN IM200717905, Philippines, PANGLAO 2005, st. CP2348, 09°26.6′N, 123°52.5′E, 196–216 m, SL 17.2 mm. D.Pyrgospira aenone (Dall, 1919)**, det. J. A. Todd, YK, NHMUK MOEA 20100539, Gulf of Panama, JTD-00–18, 08°19.50′N, 78°47.71′W, 25–32 m. SL 36.2 mm. E.Inquisitor sp.**, MNHN IM200717851, Coral Sea, EBISCO, st. DW2625, 20°05′N, 160°19′E, 627–741 m, SL 32.2 mm (radula: Fig. 16F). F.Funa incerta (Smith, 1877)**, NHMUK MOEA 20100553, st. 70, off southern Hong Kong, SL 42.8 mm. G.Leucosyrinx sp.**, MNHN IM200717846, Coral Sea, EBISCO, st. CP2600, 19°38′S, 158°46′E, 603–630 m, SL 27.1 mm (radula: Fig. 16A). H.Hindsiclava alesidota (Dall, 1889)**, det. J. A. Todd, NHMUK MOEA 20100524, Lower Florida Keys, JTD-01-15, 24°33.47′N, 81°07.72′W, 117–148 m SL 37.9 mm. I.Conorbela antarctica (Strebel, 1908) USNM 894465, Zavodovski Island, South Sandwich Islands, 56.27°S, 27.5°W, 208–375 m, SL 23.6 mm. J.Carinodrillia dichroa Pilsbry & Lowe, 1932*, det. J. A. Todd, NHMUK MOEA 20100531, Gulf of Panama, JTD-00-18, 08°19.50′N, 78°47.71′W, 25–32 m, SL 23.0 mm (radula: Fig. 16C). K.Cheungbeia robusta (Hinds, 1839)*, NHMUK MOEA 20100558, st. 70, off southern Hong Kong, South China Sea, SL 21.2 mm (radula: Fig. 16E). L.Plicisyrinx vitjazi Sysoev t Kantor, 1986, holotype, ZMMU Lc-22382, E of northern Honshu, Japan, 40°55.2′N, 144°53.3′E, 3880–4000 m, SL 13.9 mm. M.Maesiella maesae McLean & Poorman, 1971, LACM 1513, holotype, Guaymas, Sonora, Mexico, SL 9.2 mm. N.Kurilohadalia elongata Sysoev & Kantor, 1986, holotype, ZMMU Lc-22401, Kurile-Kamchatka Trench, 43°55′ N, 149°47′ E, 8080–8160 m, SL 12.5 mm. O.Crassispira quadrilirata (E.A. Smith, 1882)**, MNHN IM200717755, Philippines, PANGLAO 2004, st. L46, 09°30.9′N, 123°41.2′E, 90–110 m, SL 14.0 mm. P.Epideira sibogae (Schepman, 1913), MNHN, Vanuatu, MUSORSTOM 8, st. DW1097, 15°05′S, 167°11′E, 281–288 m, SL 24.1 mm. Q.Tiariturris spectabilis Berry, 1958**, det. J. A. Todd, NHMUK MOEA 20100540, Gulf of Panama, JTD-00-34, 08°26.24′N, 79 09.14′W, 66–68 m SL 48.7 mm (radula: Fig. 15A). R.Pseudomelatoma penicillata Carpenter, 1864, USNM 130590, California, SL 28.1 mm. S.Megasurcula carpenteriana (Gabb, 1865), USNM 523713, California, SL 84.1 mm. T.Antiplanes sanctiioannis (E.A. Smith, 1875), ZMMU Lc-17096, off northern Kurile Islands, 49°40.9′N, 154°41′E, 142 m, SL 37.2 mm. U.Naudedrillia sp., New Caledonia, VAUBAN, st. 3, 22°17′S, 167 °12′E, 390 m, SL 19.0 mm. V.Zonulispira sp.*, NHMUK MOEA 20100537, det. J. A. Todd, Gulf of Panama, JTD-00-18, 08°19.50′N, 78°47.71′W, 25–32 m, SL 21.2 mm (radula: Fig. 16B). Abbreviation and symbols: SL, shell length; *, sequenced species; **, sequenced specimen. Photo credits: B. Buge (A, G), A. Fedosov (B–F, H, J, K, Q), M.G. Harasewych and D. Tippett (R, S), P. Maestrati (U), J. McLean (M).

Figure 15.

Radulae. Pseudomelatomidae. A.Tiariturris spectabilis Berry, 1958**, NHMUK MOEA 20100540, Gulf of Panama, JTD-00-34, 08°26.24′N, 79 09.14′W, 66–68 m (shell: Fig. 14Q). B.Comitas onokeana vivens Dell, 1956, MNHN, New Caledonia, MONTROUZIER, st. 1269, after Kantor & Taylor (2000). C, D. Comitas sp.**, MNHN IM200717918 (shell: Fig. 14A). E, F.Crassiclava turricula (Sowerby, 1834), Off Nacascola, West side of Bahia Culebra, Costa Rica, after Kantor et al. (1997). Symbol: **, sequenced specimen.

Figure 15.

Radulae. Pseudomelatomidae. A.Tiariturris spectabilis Berry, 1958**, NHMUK MOEA 20100540, Gulf of Panama, JTD-00-34, 08°26.24′N, 79 09.14′W, 66–68 m (shell: Fig. 14Q). B.Comitas onokeana vivens Dell, 1956, MNHN, New Caledonia, MONTROUZIER, st. 1269, after Kantor & Taylor (2000). C, D. Comitas sp.**, MNHN IM200717918 (shell: Fig. 14A). E, F.Crassiclava turricula (Sowerby, 1834), Off Nacascola, West side of Bahia Culebra, Costa Rica, after Kantor et al. (1997). Symbol: **, sequenced specimen.

Figure 16.

Radulae. Pseudomelatomidae. A.Leucosyrinx sp.**, MNHN IM200717846 (shell: Fig. 14G). B.Zonulispira sp.**, NHMUK MOEA 20100536, Gulf of Panama, JTD-00-18, 08°19.50′N, 78°47.71′W, 25–32 m (shell: Fig. 14V). C.Carinodrillia dichroa Pilsbry & Lowe, 1932**, NHMUK MOEA 20100530, Gulf of Panama, JTD-00-18, 08°19.50′N, 78°47.71′W, 25–32 m (shell: Fig. 1 J). D.Ptychobela suturalis (Gray, 1838)**, det. J.A. Todd, YK, NHMUK MOEA 20100560, off southern Hong Kong, Sta. 71. E.Cheungbeia robusta (Hinds, 1839)*, NHMUK MOEA 20100557, coll. B. Morton, off southern Hong Kong, Sta. 70 (shell: Fig. 14K). F.Inquisitor sp.**, MNHN IM200717851 (shell: Fig. 14E). Symbols: *, sequenced species; **, sequenced specimen.

Figure 16.

Radulae. Pseudomelatomidae. A.Leucosyrinx sp.**, MNHN IM200717846 (shell: Fig. 14G). B.Zonulispira sp.**, NHMUK MOEA 20100536, Gulf of Panama, JTD-00-18, 08°19.50′N, 78°47.71′W, 25–32 m (shell: Fig. 14V). C.Carinodrillia dichroa Pilsbry & Lowe, 1932**, NHMUK MOEA 20100530, Gulf of Panama, JTD-00-18, 08°19.50′N, 78°47.71′W, 25–32 m (shell: Fig. 1 J). D.Ptychobela suturalis (Gray, 1838)**, det. J.A. Todd, YK, NHMUK MOEA 20100560, off southern Hong Kong, Sta. 71. E.Cheungbeia robusta (Hinds, 1839)*, NHMUK MOEA 20100557, coll. B. Morton, off southern Hong Kong, Sta. 70 (shell: Fig. 14K). F.Inquisitor sp.**, MNHN IM200717851 (shell: Fig. 14E). Symbols: *, sequenced species; **, sequenced specimen.

Diagnosis: Shell small to rather large, 15–100 mm high, claviform to fusiform. Spiral and axial sculpture generally well developed, often strong. Subsutural fold often present. Anal sinus on subsutural ramp, usually moderately deep to very deep, often constrained by callus rendering anal sinus subtubular. Protoconch usually paucispiral, sometimes multispiral, with up to three whorls, smooth or sometimes axially or spirally sculptured on later whorls. Operculum with terminal nucleus.

Radula very variable; four types are recorded:

1-(1-R-1)-1 – Comitas type (includes also Knefastia and Antiplanes). The central formation is rather variable in degree of development of the rachidian tooth and fusion of three teeth. In some Comitas (Fig. 15C, D), the central formation looks like single well-defined tooth [as in Comitas murrawolga (Garrard, 1961)], while in Comitas onokeana vivens Dell, 1956, Knefastia and Antiplanes, the rachidian is totally reduced and the formation appears as two poorly developed paired plates (reduced laterals) (Fig. 15B). Marginal teeth in Comitas and Knefastia flat, broadly oval, with thickened edges and teeth tips and without pronounced accessory limb. In Antiplanes marginal teeth narrowly elongate, with well-developed accessory limb.

1-1-0-1-1 – Crassiclava type. Differs from the Comitas type by the better defined laterals (Fig. 15E, F) that are low, arcuate and sharply curved towards the midline of the ribbon.

1-0-R-0-1 – Pseudomelatoma type (also includes Hormospira and Tiariturris, Fig. 15A). Rachidian with strong cusp and subrectangular base (contrary to the other types with a central formation formed by fused lateral and rachidian). Despite neither ontogeny nor folding of the radula have been examined, we tentatively treat this structure as a true rachidian. Marginal teeth simple, solid and strongly curved, attached to the membrane by rather a narrow base and free along most of their length.

1-0-0-0-1 – Most genera of the family. Marginal teeth elongated, narrow, flat, with thickened edge (e.g. Funa, Carinodrillia, Fig. 16C), or trough-shaped in transverse section, sometimes with small barb near the tip (Cheungbeia, Fig. 16E), which may become semi-enrolled (e.g. Pyrgospira, Pilsbryspira, Zonulispira, Fig. 16B), to nearly hollow, where limbs overlap at significant length of the teeth (Ptychobela, Fig. 16D).

Remarks: Anatomically, Pseudomelatomidae is the most variable family of Conoidea. Most genera were formerly included in the subfamily Crassispirinae, but the nomenclaturally valid name for this clade is Pseudomelatomidae. Its constituents includes several taxa that were previously recognized as separate (sub)families: Zonulispirinae, characterized by semi-enrolled marginal radular teeth (a character found in several branches of the clade), and Pseudomelatominae, defined on the basis of the very characteristic solid marginal teeth and strongly developed rachidian.

The genus Leucosyrinx has long been a convenient genus for placement of turreted-fusiform species, mostly from deep water of the Atlantic and Indo-Pacific. Currently, it is a mixture of species of probably different taxonomic position, and the membership of the genus needs revision. The type species from the northern Atlantic, Leucosyrinx verrilli (Dall, 1881), has a radula consisting of only duplex, rather robust marginal teeth (Powell, 1966: text fig. B12), a radula type also found by us in Indo-Pacific species (Fig. 16A). A second type of radula is found in Sibogasyrinx, originally described as a subgenus of Leucosyrinx, and which clusters in the molecular tree with the Cochlespiridae (see under that family). In our study, true Leucosyrinx appears to be sister to the Pseudomelatomidae, but this relationship has poor support. As a working hypothesis, we tentatively include Leucosyrinx in the Pseudomelatomidae. Clavatulidae Gray, 1853

  • Included genera:

  • ?Abyssocomitas Sysoev & Kantor, 19862

  • Aguilaria Taylor & Wells, 19942

  • ?AnticomitasPowell, 19421

  • AntimelatomaPowell, 19421

  • Antiplanes Dall, 19022

    •  [= Rectiplanes Bartsch, 1944]

    •  [= Rectisulcus Habe, 1958]

  • Austrotoma Finlay, 19241

  • ?Benthodaphne Oyama, 19621

  • Burchia Bartsch, 19442

  • Buridrillia Olsson, 19422

  • CalcatodrilliaKilburn, 19882

  • Carinodrillia Dall, 19193

  • ?Carinoturris Bartsch, 19442

  • Cheungbeia Taylor & Wells, 19943

  • Comitas Finlay, 19263

  • Compsodrillia Woodring, 19282

    •  (Mammillaedrillia Kuroda and Oyama in Kuroda, Habe & Oyama, 1971)

  • ?Conorbela Powell, 19512

  • Conticosta Laseron, 19541

  • CrassiclavaMcLean, 19712

  • Crassispira Swainson, 18403

    •  (Crassispirella Bartsch & Rehder, 1939)

  • Cretaspira Kuroda & Oyama in Kuroda, Habe & Oyama, 19711

  • Dallspira Bartsch, 19502

  • DoxospiraMcLean, 19712

  • Epideira Hedley, 19182

    •  [= Epidirona Hedley, 1931]

  • FunaKilburn, 19883

  • GibbaspiraMcLean, 19712

  • GlossispiraMcLean, 19712

  • Hindsiclava Hertlein & Strong, 19553

    •  [= Turrigemma Berry, 1958]

  • Hormospira Berry, 19582

  • Inquisitor Hedley, 19183

  • Knefastia Dall, 19193

  • ?Kurilohadalia Sysoev & Kantor, 19862

  • Kurodadrillia Azuma, 19751

  • ?Leucosyrinx Dall 18893

  • Lioglyphostoma Woodring, 19282

  • Lioglyphostomella Shuto, 19701

  • MaesiellaMcLean, 19712

  • Megasurcula Casey, 19042

  • ?Meggittia Ray, 19771

  • Miraclathurella Woodring, 19282

  • Monilispira Bartsch & Rehder, 19392

  • NaudedrilliaKilburn, 19882

  • Otitoma Jousseaume, 18983

    •  [= Thelecytharella Shuto, 1969]

  • ?ParacomitasPowell, 19421

  • PilsbryspiraMcLean, 19713

  •   (NymphispiraMcLean, 1971)

  • Plicisyrinx Sysoev & Kantor, 19862

  • Pseudomelatoma Dall, 19183

  • ?Pseudotaranis McLean, 19952

  • Ptychobela Thiele, 19253

    •  [= ?Brachytoma Swainson, 1840]

  • PyrgospiraMcLean, 19713

  • ?Rhodopetoma Bartsch, 19442

  • Sediliopsis Petuch, 19881

  • ?Shutonia Van der Bijl, 19931

  • Striospira Bartsch, 19502

  • Tiariturris Berry, 19583

  • Viridrillia Bartsch, 19431

    •  [= Viridrillina Bartsch, 1943]

  • Zonulispira Bartsch, 19503

(Figs 17A–F, 18)

Figure 17.

Shells. A–F. Clavatulidae. A.Gemmuloborsonia colorata (Sysoev & Bouchet, 2001)**, MNHN IM200717849, Coral Sea, EBISCO, st. DW2619, 20°06′S, 160 °23′E, 490–550 m, SL 45.5 mm (radula: Fig. 18F). B.Pusionella compacta Strebel, 1914**, MNHN IM200717830, Cacuaco, Bengo Prov., Angola, 5–10 m, SL 24.1 mm (radula: Fig. 18C). C.Clavatula xanteni Nolf & Verstraeten, 2006**, MNHN IM200717829, Ilha de Luanda, Luanda Prov., Angola, 40–60 m, SL 20.5 mm (radula: Fig. 18E). D.Perrona subspirata (von Martens, 1902)**, MNHN IM200717833, Moçâmedes, Namibe Prov., Angola, 5–10 m, SL 24.9 mm. E.Turricula javana (Linnaeus, 1767), S India, SL 63 mm. F.Fusiturris undatiruga (Bivona, 1838), Sierra Leone, 60–80 m, SL 43 mm. G–R: Horaiclavidae. G.Marshallena philippinarum (Watson, 1882), MNHN Philippines, PANGLAO 2005, st. CP2334, 09°37.5′N, 123°40.2′E, 606–631 m, SL 26 mm. H.Horaiclavus splendidus (A. Adams, 1867)**, MNHN IM200717840, Coral Sea, EBISCO, st. DW2631,21°03′S, 160°44′E, 372–404 m, SL 18.0 mm (radula: Fig. 19D). I.Ceritoturris pupiformis (Smith, 1884)**, MNHN IM200717888, Philippines, PANGLAO 2004, st. T36, 09°29.3′N, 123°51.5′E, 95–128 m, SL 3.4 mm (radula: Fig. 19C). J.Inkinga sp., New Caledonia, NORFOLK 1, st. DW1737, 22°52′S, 167°12′E, 343–400 m, SL 11.1 mm. (radula: Fig. 19B). K. Horaiclavidae gen. 1**, MNHN IM200742501, Solomon Islands, SALOMON 2, st. CP2219, 07°58.3′S, 157°34.4′E, 650–836 m, SL 10.1 mm. L.Buchema interpleura (Dall & Simpson, 1901), USNM 900896, Lemon Cay, San Blas Islands, Panama, Caribbean Sea, 2 m, SL 10.0 mm. M.Carinapex minutissima (Garrett, 1873), MNHN, New Caledonia, LIFOU 2000, st. 1457, 20°46.8′S, 167°02.75′E, 5–10 m, SL 3.1 mm. N.Paradrillia sp.**, MNHN IM200742475, Philippines, PANGLAO 2005, st. CP2396 09°36.3′N, 123°42.0′E, 609–673 m, SL 11.5 mm (radula: Fig. 19A). O.Haedropleura septangularis MNHN, Finistère, France, 20–30 m, SL 11 mm. P.Anguloclavus sp.**, MNHN IM200717908, PANGLAO 2005, st. CP2332, 09°38.8′N, 123°45.9′E, SL 19.0 mm. Q.Iwaoa sp., New-Caledonia, MUSORSTOM6, st. DW489, 20°48′S, 167°06′E, 700 m, SL 13.5 mm. R.Anacithara themeropsis (Melvill & Standen, 1896), MNHN, LIFOU 2000, st. 1427, 20°47.6′S, 167°10.2′E, 10 m, SL 4.5 mm. Abbreviation and symbols: SL, shell length; * , sequenced species; **, sequenced specimen. Photo credits: B. Buge (B–D, H, I, P, O), M.G. Harasewych and D. Tippett (L), P. Maestrati (J, Q, R).

Figure 17.

Shells. A–F. Clavatulidae. A.Gemmuloborsonia colorata (Sysoev & Bouchet, 2001)**, MNHN IM200717849, Coral Sea, EBISCO, st. DW2619, 20°06′S, 160 °23′E, 490–550 m, SL 45.5 mm (radula: Fig. 18F). B.Pusionella compacta Strebel, 1914**, MNHN IM200717830, Cacuaco, Bengo Prov., Angola, 5–10 m, SL 24.1 mm (radula: Fig. 18C). C.Clavatula xanteni Nolf & Verstraeten, 2006**, MNHN IM200717829, Ilha de Luanda, Luanda Prov., Angola, 40–60 m, SL 20.5 mm (radula: Fig. 18E). D.Perrona subspirata (von Martens, 1902)**, MNHN IM200717833, Moçâmedes, Namibe Prov., Angola, 5–10 m, SL 24.9 mm. E.Turricula javana (Linnaeus, 1767), S India, SL 63 mm. F.Fusiturris undatiruga (Bivona, 1838), Sierra Leone, 60–80 m, SL 43 mm. G–R: Horaiclavidae. G.Marshallena philippinarum (Watson, 1882), MNHN Philippines, PANGLAO 2005, st. CP2334, 09°37.5′N, 123°40.2′E, 606–631 m, SL 26 mm. H.Horaiclavus splendidus (A. Adams, 1867)**, MNHN IM200717840, Coral Sea, EBISCO, st. DW2631,21°03′S, 160°44′E, 372–404 m, SL 18.0 mm (radula: Fig. 19D). I.Ceritoturris pupiformis (Smith, 1884)**, MNHN IM200717888, Philippines, PANGLAO 2004, st. T36, 09°29.3′N, 123°51.5′E, 95–128 m, SL 3.4 mm (radula: Fig. 19C). J.Inkinga sp., New Caledonia, NORFOLK 1, st. DW1737, 22°52′S, 167°12′E, 343–400 m, SL 11.1 mm. (radula: Fig. 19B). K. Horaiclavidae gen. 1**, MNHN IM200742501, Solomon Islands, SALOMON 2, st. CP2219, 07°58.3′S, 157°34.4′E, 650–836 m, SL 10.1 mm. L.Buchema interpleura (Dall & Simpson, 1901), USNM 900896, Lemon Cay, San Blas Islands, Panama, Caribbean Sea, 2 m, SL 10.0 mm. M.Carinapex minutissima (Garrett, 1873), MNHN, New Caledonia, LIFOU 2000, st. 1457, 20°46.8′S, 167°02.75′E, 5–10 m, SL 3.1 mm. N.Paradrillia sp.**, MNHN IM200742475, Philippines, PANGLAO 2005, st. CP2396 09°36.3′N, 123°42.0′E, 609–673 m, SL 11.5 mm (radula: Fig. 19A). O.Haedropleura septangularis MNHN, Finistère, France, 20–30 m, SL 11 mm. P.Anguloclavus sp.**, MNHN IM200717908, PANGLAO 2005, st. CP2332, 09°38.8′N, 123°45.9′E, SL 19.0 mm. Q.Iwaoa sp., New-Caledonia, MUSORSTOM6, st. DW489, 20°48′S, 167°06′E, 700 m, SL 13.5 mm. R.Anacithara themeropsis (Melvill & Standen, 1896), MNHN, LIFOU 2000, st. 1427, 20°47.6′S, 167°10.2′E, 10 m, SL 4.5 mm. Abbreviation and symbols: SL, shell length; * , sequenced species; **, sequenced specimen. Photo credits: B. Buge (B–D, H, I, P, O), M.G. Harasewych and D. Tippett (L), P. Maestrati (J, Q, R).

Figure 18.

Radulae. Clavatulidae. A, B.Turricula nelliae (E.A. Smith, 1877)**, NHMUK MOEA 20100551, Danang, Vietnam. C.Pusionella compacta Strebel, 1914**, MNHN IM200717830 (shell: Fig. 17B). D.Toxiclionella tumida (Sowerby, 1870), South Africa, after Kantor & Taylor (2000). E.Clavatula xanteni Nolf & Verstraeten, 2006**, MNHN IM200717829 (shell: Fig. 17C). F.Gemmuloborsonia colorata (Sysoev & Bouchet, 2001)**, MNHN IM200717849 (shell: Fig. 17A). Symbol: **, sequenced specimen. Arrow, see text.

Figure 18.

Radulae. Clavatulidae. A, B.Turricula nelliae (E.A. Smith, 1877)**, NHMUK MOEA 20100551, Danang, Vietnam. C.Pusionella compacta Strebel, 1914**, MNHN IM200717830 (shell: Fig. 17B). D.Toxiclionella tumida (Sowerby, 1870), South Africa, after Kantor & Taylor (2000). E.Clavatula xanteni Nolf & Verstraeten, 2006**, MNHN IM200717829 (shell: Fig. 17C). F.Gemmuloborsonia colorata (Sysoev & Bouchet, 2001)**, MNHN IM200717849 (shell: Fig. 17A). Symbol: **, sequenced specimen. Arrow, see text.

Diagnosis: Shell medium-sized to rather large (usually 15–30 mm, up to 85 mm high), broad-fusiform to turreted-fusiform, with high spire and usually moderately long siphonal canal. Subsutural ramp usually well developed, with very shallow to rather deep anal sinus situated on its lower part or shifted abapically, with apex at almost a peripheral position. Sculpture variously developed, from almost smooth shell surface to well-developed axial ribs and spiral cords. Protoconch only known as paucispiral, up to c. 2.5 smooth whorls. Operculum with medio-lateral nucleus. Radular formula 1-(1-R-1)-1. The central formation is composed of very thin, broad, plate-like lateral teeth and a small, but sometimes strong, narrow rachidian. The central formation is variously developed, sometimes appearing as a pronounced tooth (Fig. 18B), sometimes clearly consisting of three elements (Clionella sinuata, see Taylor et al., 1993; Fig. 18A, B), to its nearly complete reduction (Fig. 18C). Marginal teeth usually duplex, with sharp-edged major limb and a deep socket where an accessory limb is inserted, often with angulation distal to the socket (arrow in Fig. 18E). In Toxiclionella the marginal teeth are hypodermic, loosely enrolled, attached along their length to radular membrane, having two barbs at the tip and a tooth canal opening subterminally (Fig. 18D).

Remarks: The genus Gemmuloborsonia represents a sister group to the (Clavatulidae + Horaiclavidae) clade, but this node is not well supported. Because it resembles much more the Clavatulidae than the Horaiclavidae in terms of shell and radular characters (Fig. 18F), Gemmuloborsonia is provisionally included in the former family. If further studies support that hypothesis, then the diagnosis of the Clavatulidae should be amended to account for the presence of weak columellar pleats and multispiral Turridae-type protoconch present in Gemmuloborsonia. Horaiclavidae new family

  • Included genera:

  • Benthoclionella Kilburn, 19742

  • CaliendrulaKilburn, 19851

  • Clavatula Lamarck, 18013

  • Clionella Gray, 18472

  • ?Fusiturris Thiele, 19291

    •  [= Tyrrhenoturris Coen, 1929]

  • ?Gemmuloborsonia Shuto, 19893

  • Makiyamaia Kuroda in MacNeil, 19611

  • Perrona Schumacher, 18173

    •  (Tomellana Wenz, 1943)

  • Pusionella Gray, 18473

    •  [= Netrum Philippi, 1850]

  • Scaevatula Gofas, 19891

  • ToxiclionellaPowell, 19662

  • Trachydrillida Nolf & Swinnen, 20101

  • Turricula Schumacher, 18173

  •   [= SurculaH. & A. Adams, 1853]

(Figs 17G–R, 19)

Figure 19.

Radulae. Horaiclavidae. A.Paradrillia sp.**, MNHN IM200742475 (shell: Fig. 17N). B.Inkinga sp., MNHN (shell: Fig. 17 J). C.Ceritoturris pupiformis (E.A. Smith, 1884)**, MNHN IM200717888 (shell: Fig. 17I). D.Horaiclavus splendidus (A. Adams, 1867)**, MNHN IM200717840 (shell: Fig. 17H). Symbol: **, sequenced specimen. Arrows, see text.

Figure 19.

Radulae. Horaiclavidae. A.Paradrillia sp.**, MNHN IM200742475 (shell: Fig. 17N). B.Inkinga sp., MNHN (shell: Fig. 17 J). C.Ceritoturris pupiformis (E.A. Smith, 1884)**, MNHN IM200717888 (shell: Fig. 17I). D.Horaiclavus splendidus (A. Adams, 1867)**, MNHN IM200717840 (shell: Fig. 17H). Symbol: **, sequenced specimen. Arrows, see text.

Type genus: Horaiclavus Oyama, 1954.

Diagnosis: Shell generally small, 5–25 mm, usually 7–15 mm high, shortly claviform, with relatively low spire and a short, truncated, poorly differentiated siphonal canal. Subsutural ramp usually poorly differentiated. Axial sculpture almost always present, usually as strong sinuate ribs. Spiral sculpture normally weak or obsolete, often with glossy shell surface. Anal sinus on subsutural slope, weak to moderately deep, often constrained by callus. Protoconch of up to 3.5 medially carinate but otherwise smooth whorls when multispiral, but usually paucispiral and smooth. Operculum with terminal nucleus. Radular formula: 1-0-0-0-1. Marginal teeth duplex, with lanceolate major limb and usually narrow accessory limb, which is inserted in a shallow socket. Major limb often with angulation lateral to the place of accessory limb insertion. Rarely (some Paradrillia, cf. Kilburn, 1988, Fig. 17; Inkinga, Fig. 19B) the additional limb is of similar size to the major limb and the teeth become trough-shaped in transverse section, with a collar near the base (arrows in Fig. 19B). In several species of Horaiclavus the radular apparatus is absent.

Remarks: This family shares many characters with Pseudomelatomidae, conchologically differing by a small stout shell with short siphonal canal and usually poorly developed spiral sculpture.

Radulae also are rather similar to many representatives of Pseudomelatomidae and no clear cut distinctions were found. Genera currently included in Horaiclavidae have been usually included in the Crassispiridae (= Pseudomelatomidae), and the clear molecular-based division between the two clades seems to be not so clearly reflected in shell-based distinction. Therefore, the generic composition of the family is somewhat provisional and needs confirmation by further molecular data and/or a detailed analysis of conchological and radular characters.

Included genera: TurridaeH. Adams & A. Adams, 1853

  • Anacithara Hedley, 19223

  • Anguloclavus Shuto, 19833

  • ?AoteadrilliaPowell, 19422

  • ?Austrocarina Laseron, 19541

  • Austrodrillia Hedley, 19181

  • Buchema Corea, 19342

  • Carinapex Dall, 19243

  • Ceritoturris Dall, 19243

  • Coronacomitas Shuto, 19831

  • Cytharoclavus Kuroda & Oyama in Kuroda, Habe & Oyama, 19711

  • ?Darrylia García, 20081

  • Graciliclava Shuto, 19831

  • Haedropleura Bucquoy, Dautzenberg & Dollfus, 18832

  • Horaiclavus Oyama, 19543

  • Inodrillia Bartsch, 19431

    •  [= Inodrillara Bartsch, 1943]

    •  [= Inodrillina Bartsch, 1943]

  • InkingaKilburn, 19881

  • Iwaoa Kuroda, 19401

  • ?Marshallena Allan, 19272

    •  [= Sugitanitoma Kuroda, 1959]

  • MauidrilliaPowell, 19422

  • ?Micropleurotoma Thiele, 19292

  • Naskia Sysoev & Ivanov, 19851

  • NqumaKilburn, 19882

  • Paradrillia Makiyama, 19403

    •  [= VexitominaPowell, 1942]

    •  [= Alticlavatula MacNeil, 1960]

  • ?Pseudexomilus Powell, 19441

  • PsittacodrilliaKilburn, 19881

  • StriatoguraleusKilburn, 19941

  • ?Thelecythara Woodring, 19281

(Figs 20 and 21)

Figure 20.

Shells. Turridae. A.Xenuroturris legitima Iredale, 1929**, MNHN IM200717684, Vanuatu, SANTO 2006, st. DR087, 15°38.5′S, 167°15.1′E, 13 m, SL 57.0 mm (radula: Fig. 21C). B.Iotyrris cingulifera (Lamarck, 1822)**, MNHN IM200717685, Vanuatu, SANTO 2006, st. FS84, 15°33.6′S, 167°16.6′E, 8–9 m, SL 15.5 mm (radula: Fig. 21D). C.Decollidrillia nigra Habe & Ito, 1965, ZMMU uncatalogued, southern Kurile Islands, SL 12.8 mm. D.Lophiotoma acuta (Perry, 1811)**, MNHN IM200717860, Philippines, PANGLAO 2004, st. R44, 09°33.3′N, 123°43.9′E, 2 m, SL 44.0 mm. E.Turridrupa acutigemmata (E.A. Smith, 1877), New Caledonia, 46 m, SL 26.5 mm. F.Turris babylonia (Linnaeus, 1758)**, MNHN IM200717754, Philippines, PANGLAO 2004, st. R42, 09°37.1′N, 123°52.6′E, 8–22 m, SL 79.4 mm. G.Gemmula rarimaculata (Kuroda & Oyama, 1971)**, MNHN IM200717838, Coral Sea, EBISCO, st. DW2533, 22°18′S, 159°28′E, 360–370 m, SL 13.7 mm. H.Ptychosyrinx chilensis Berry, 1968, USNM 870005, S of Coquimbo, Chile, 31.1° S, 71.8° W, 179–187 m, SL 21.1 mm. I.Lucerapex cf. casearia (Hedley & Petterd, 1906)**, MNHN IM200742448, Philippines, PANGLAO 2005, st. CP2363, 09°06.0′N, 123°25.0′E, 437–439 m, SL 21.0 mm (radula: Fig. 21 E). J.Cryptogemma corneus (Okutani, 1966), ZIN 58809/1, off Shikotan I., Kurile Islands, 1450–1530 m, 12.3 mm. Abbreviation and symbol: SL, shell length; **, sequenced specimen.

Figure 20.

Shells. Turridae. A.Xenuroturris legitima Iredale, 1929**, MNHN IM200717684, Vanuatu, SANTO 2006, st. DR087, 15°38.5′S, 167°15.1′E, 13 m, SL 57.0 mm (radula: Fig. 21C). B.Iotyrris cingulifera (Lamarck, 1822)**, MNHN IM200717685, Vanuatu, SANTO 2006, st. FS84, 15°33.6′S, 167°16.6′E, 8–9 m, SL 15.5 mm (radula: Fig. 21D). C.Decollidrillia nigra Habe & Ito, 1965, ZMMU uncatalogued, southern Kurile Islands, SL 12.8 mm. D.Lophiotoma acuta (Perry, 1811)**, MNHN IM200717860, Philippines, PANGLAO 2004, st. R44, 09°33.3′N, 123°43.9′E, 2 m, SL 44.0 mm. E.Turridrupa acutigemmata (E.A. Smith, 1877), New Caledonia, 46 m, SL 26.5 mm. F.Turris babylonia (Linnaeus, 1758)**, MNHN IM200717754, Philippines, PANGLAO 2004, st. R42, 09°37.1′N, 123°52.6′E, 8–22 m, SL 79.4 mm. G.Gemmula rarimaculata (Kuroda & Oyama, 1971)**, MNHN IM200717838, Coral Sea, EBISCO, st. DW2533, 22°18′S, 159°28′E, 360–370 m, SL 13.7 mm. H.Ptychosyrinx chilensis Berry, 1968, USNM 870005, S of Coquimbo, Chile, 31.1° S, 71.8° W, 179–187 m, SL 21.1 mm. I.Lucerapex cf. casearia (Hedley & Petterd, 1906)**, MNHN IM200742448, Philippines, PANGLAO 2005, st. CP2363, 09°06.0′N, 123°25.0′E, 437–439 m, SL 21.0 mm (radula: Fig. 21 E). J.Cryptogemma corneus (Okutani, 1966), ZIN 58809/1, off Shikotan I., Kurile Islands, 1450–1530 m, 12.3 mm. Abbreviation and symbol: SL, shell length; **, sequenced specimen.

Figure 21.

Radulae. Turridae. A.Turridrupa cf. armillata (Reeve, 1845), MNHN IM200740773, Coral Sea, EBISCO, st. DW2607, 19°33′S, 158°40′E, 400–413 m. B.Gemmula kieneri (Doumet, 1840), MNHN, Vanuatu, MUSORSTOM 8, st. CP1123, 15°07′S, 166°55′E, 262–352 m. C.Xenuroturris legitima Iredale, 1929, MNHN IM200717684 (shell: Fig. 20A). D.Iotyrris cingulifera (Lamarck, 1822)**, MNHN IM200717685 (shell: Fig. 20B). E.Lucerapex cf. casearia (Hedley & Petterd, 1906)**, MNHN IM200742448 (shell: Fig. 20I). F.Turris crispa (Lamarck, 1816), MNHN, Ile Ouen-Baie du Prony, st. 80, 22°31S, 166°28E, 33 m, SS 39 mm. Symbols: **, sequenced specimen.

Figure 21.

Radulae. Turridae. A.Turridrupa cf. armillata (Reeve, 1845), MNHN IM200740773, Coral Sea, EBISCO, st. DW2607, 19°33′S, 158°40′E, 400–413 m. B.Gemmula kieneri (Doumet, 1840), MNHN, Vanuatu, MUSORSTOM 8, st. CP1123, 15°07′S, 166°55′E, 262–352 m. C.Xenuroturris legitima Iredale, 1929, MNHN IM200717684 (shell: Fig. 20A). D.Iotyrris cingulifera (Lamarck, 1822)**, MNHN IM200717685 (shell: Fig. 20B). E.Lucerapex cf. casearia (Hedley & Petterd, 1906)**, MNHN IM200742448 (shell: Fig. 20I). F.Turris crispa (Lamarck, 1816), MNHN, Ile Ouen-Baie du Prony, st. 80, 22°31S, 166°28E, 33 m, SS 39 mm. Symbols: **, sequenced specimen.

Diagnosis: shell of medium to large size (usually 20–30 mm, up to 110 mm high), short- to high-fusiform, usually with a high spire and a long (rarely short and truncated) siphonal canal. Axial sculpture weak or absent. Anal sinus on whorl periphery. Protoconch typically multispiral, up to six whorls, protoconch I smooth, protoconch II with arcuate axial riblets; reduced paucispiral protoconch usually smooth, may have arcuate axial riblets. Operculum fully developed, with terminal nucleus. Radular formula typically 1-(1:R:1)-1. Small and narrow rachidian and plate-like laterals are fused together, together constituting a central formation of different development (Kantor, 2006), varying from a well-defined broad central tooth (Fig. 21B) to a tooth clearly formed of three elements (Fig. 21A) through a gradual reduction of rachidian and/or laterals to complete absence (Fig. 21D). Marginals duplex, of variable morphology, from broadly oval and flattened (Fig. 21E, F) with nearly equally developed limbs to awl-shaped and divided only in basal part (Fig. 21A). In most cases, the major limb is large and knife-shaped, while the accessory limb is dorsal and more weakly developed (Fig. 21). In Iotyrris, marginal teeth have equally developed limbs that form a shallow broad trough (Fig. 21D).

Remarks: This group is well defined by its usually narrowly fusiform shell with obsolete axial sculpture and peripheral anal sinus. However, the genus Lucerapex, although fully conforming conchologically, occupies a position on the tree (sister group to Turridae + Terebridae) that excludes it from the Turridae. It is nevertheless tentatively included here in Turridae.

Included genera: StrictispiridaeMcLean, 1971

  • Cryptogemma Dall, 19182

  • Decollidrillia Habe & Ito, 19651

  • Epidirella Iredale, 19131

    •  [= Austrogemmula Laseron, 1954]

  • Gemmula Weinkauff, 18753

    •  [= Eugemmula Iredale, 1931]

    •  (Pinguigemmula MacNeil, 1960)

  • Iotyrris Medinskaya & Sysoev, 20013

  • Kuroshioturris Shuto, 19611

  • Lophiotoma Casey, 19043

    •  [= Lophioturris Powell, 1964]

  • ?Lucerapex Iredale, 19363

  • Polystira Woodring, 19283

    •  [= Oxytropa Glibert, 1955]

  • Ptychosyrinx Thiele, 19253

    •  [= Bathybermudia Haas, 1949]

  • Turridrupa Hedley, 19223

  • Turris Batsch, 17893 (For authorship of Turris, see Dubois & Bour (2010))

    •  [= Pleurotoma Lamarck, 1799]

    •  [= Pleurotomus Blainville, 1810]

    •  (AnnulaturrisPowell, 1966)

  • Unedogemmula MacNeil, 19612

  • Xenuroturris Iredale, 19293

  •   [= Clamturris Iredale, 1931]

(Figs 11E, 22A, B)

Figure 22.

Radulae. A, B. Strictispiridae.Strictispira paxillus (Reeve, 1845), British Virgina Islands, ANSP. A. teeth detached from the membrane and showing the large median flange. B. Two teeth, showing the large median flange attached to the membrane. Photo submit: J.D. Taylor (A. B). C. Conorbidae.Benthofascis lozoueti Sysoev & Bouchet, 2001, MNHN IM200742331 (shell: Fig. 2O).

Figure 22.

Radulae. A, B. Strictispiridae.Strictispira paxillus (Reeve, 1845), British Virgina Islands, ANSP. A. teeth detached from the membrane and showing the large median flange. B. Two teeth, showing the large median flange attached to the membrane. Photo submit: J.D. Taylor (A. B). C. Conorbidae.Benthofascis lozoueti Sysoev & Bouchet, 2001, MNHN IM200742331 (shell: Fig. 2O).

Figure 23.

Protoconchs. Only multispiral protonchs are illustrated. A. Drilliidae. Iredalea sp. (= aureolinea), MNHN, New Caledonia, LIFOU 2000, st. 1441, 20°46.4′S, 167°02.0′E, 20 m. B. Pseudomelatomiidae. Inquisitor cf. nodicostatusKilburn, 1988, MNHN, New Caledonia, MUSORSTOM 6, st. DW391, 20°47′S, 167°06′E, 390 m. C. Clavatulidae. Clavatula strebeli Knudsen, 1952, MNHN, Ilha de Luanda, Angola, 40–60 m. D. Horaiclavidae. Ceritoturris pupiformis (E.A. Smith, 1884)*, MNHN, New Caledonia, MONTROUZIER, st.1249, 20°49′S, 165°19′E, 80–140 m. E. Turridae. Gemmula sp., MNHN IM200740766, Coral Sea, EBISCO, DW2625, 20°05′S, 160°20′E, 627–741 m. F. Borsoniidae. Microdrillia fastosa (Hedley, 1907), MNHN, New Caledonia, SMIB 8, st. DW146, 24°55′S, 168°22′E, 514–522 m. G, H. Clathurellidae. G.Clathurella curtisiana (Hedley, 1922), MNHN, New Caledonia, MONTROUZIER, st.1250, 20°46.7′S, 165°13.7′E, 3–6 m. H.Nannodiella acricula (Hedley, 1922), MNHN, New Caledonia, MONTROUZIER, st.1261, 20°46′S, 165°15′E, 45–56 m. I. Mitromorphidae. Mitromorpha metula** (Hinds, 1843), MNHN IM200717898, Philippines, PANGLAO 2004, st. B8, 9°37.1′N, 123°46.1′E, 3 m. J–L. Mangeliidae. J.Benthomangelia sp., MNHN, Fiji, BORDAU 1, st. CP1407, 16°40′S, 179°39′E, 499–529 m. K.Neoguraleus sp., MNHN, New Caledonia, CONCALIS, st. DW2982, 18°02′S, 163°04′E, 320–337 m. L.Pyrgocythara notabilis (E.A. Smith, 1888), MNHN, New Caledonia, LIFOU 2000, st. 1429, 20°47.5′S, 167°07.1′E, 8–18 m. M–Q. Raphitomidae. M.Neopleurotomoides sp., MNHN, New Caledonia, BIOCAL, st. CP75, 22°19′S, 167°23′E. N.Famelica sp., MNHN, New Caledonia, BATHUS 4, st. DW914, 18°49′S, 163°15′E, 600–616 m. O.Tritonoturris cumingi (Powys, 1835), MNHN, Touho, New Caledonia, st. 1357, 22°30.2′S, 166°26,4′E, 22–35 m. P.Veprecula sp., MNHN, New Caledonia, MONTROUZIER, st. 1306, 20°39.1′S, 164°12.4′E, 11–13 m. Q.Pleurotomella packardi Verrill, 1872, MNHN, INCAL, Porcupine abyssal plain, st. WS10, 47°27′N, 09°40′W, 4354 m. Symbol: *, sequenced species; **, sequenced specimen.

Figure 23.

Protoconchs. Only multispiral protonchs are illustrated. A. Drilliidae. Iredalea sp. (= aureolinea), MNHN, New Caledonia, LIFOU 2000, st. 1441, 20°46.4′S, 167°02.0′E, 20 m. B. Pseudomelatomiidae. Inquisitor cf. nodicostatusKilburn, 1988, MNHN, New Caledonia, MUSORSTOM 6, st. DW391, 20°47′S, 167°06′E, 390 m. C. Clavatulidae. Clavatula strebeli Knudsen, 1952, MNHN, Ilha de Luanda, Angola, 40–60 m. D. Horaiclavidae. Ceritoturris pupiformis (E.A. Smith, 1884)*, MNHN, New Caledonia, MONTROUZIER, st.1249, 20°49′S, 165°19′E, 80–140 m. E. Turridae. Gemmula sp., MNHN IM200740766, Coral Sea, EBISCO, DW2625, 20°05′S, 160°20′E, 627–741 m. F. Borsoniidae. Microdrillia fastosa (Hedley, 1907), MNHN, New Caledonia, SMIB 8, st. DW146, 24°55′S, 168°22′E, 514–522 m. G, H. Clathurellidae. G.Clathurella curtisiana (Hedley, 1922), MNHN, New Caledonia, MONTROUZIER, st.1250, 20°46.7′S, 165°13.7′E, 3–6 m. H.Nannodiella acricula (Hedley, 1922), MNHN, New Caledonia, MONTROUZIER, st.1261, 20°46′S, 165°15′E, 45–56 m. I. Mitromorphidae. Mitromorpha metula** (Hinds, 1843), MNHN IM200717898, Philippines, PANGLAO 2004, st. B8, 9°37.1′N, 123°46.1′E, 3 m. J–L. Mangeliidae. J.Benthomangelia sp., MNHN, Fiji, BORDAU 1, st. CP1407, 16°40′S, 179°39′E, 499–529 m. K.Neoguraleus sp., MNHN, New Caledonia, CONCALIS, st. DW2982, 18°02′S, 163°04′E, 320–337 m. L.Pyrgocythara notabilis (E.A. Smith, 1888), MNHN, New Caledonia, LIFOU 2000, st. 1429, 20°47.5′S, 167°07.1′E, 8–18 m. M–Q. Raphitomidae. M.Neopleurotomoides sp., MNHN, New Caledonia, BIOCAL, st. CP75, 22°19′S, 167°23′E. N.Famelica sp., MNHN, New Caledonia, BATHUS 4, st. DW914, 18°49′S, 163°15′E, 600–616 m. O.Tritonoturris cumingi (Powys, 1835), MNHN, Touho, New Caledonia, st. 1357, 22°30.2′S, 166°26,4′E, 22–35 m. P.Veprecula sp., MNHN, New Caledonia, MONTROUZIER, st. 1306, 20°39.1′S, 164°12.4′E, 11–13 m. Q.Pleurotomella packardi Verrill, 1872, MNHN, INCAL, Porcupine abyssal plain, st. WS10, 47°27′N, 09°40′W, 4354 m. Symbol: *, sequenced species; **, sequenced specimen.

Diagnosis: Shell medium-sized, to 20 mm, claviform. Spiral and axial sculpture well developed, shoulder with a marked subsutural fold. Anal sinus deep, laterally directed. Parietal callus well developed. Protoconch paucispiral in all species examined, smooth. Operculum leaf-shaped, with terminal nucleus. Radula consisting of a pair of solid awl-shaped marginal teeth that have a prominent flange, located above the base of the tooth and firmly attached to the radular membrane.

Remarks: We did not obtain any material of Strictispiridae for molecular analysis and the position of the family group remains unclear. The shell resembles that of Pseudomelatomidae and the shape of the radular marginal teeth is also somewhat similar to Pseudomelatoma, Hormospira and Tiariturris, but it differs in the absence of the rachidian. Among the examined species of Strictispira, S. paxillus is characterised by rather unusual characters, such as the absence of the venom gland together with very large and powerful odontophore (Kantor & Taylor, 1994). Until molecular data are available, we conservatively treat this family as valid, following Taylor et al. (1993).

Included genera: Terebridae Mörch, 1852

Diagnosis: Shell medium-sized to large, 8–270 mm, usually 30–100 mm high, auger-shaped, with high to very high multiwhorled spire and flattened shell profile, aperture relatively small. Siphonal canal short, anal sinus not pronounced. Protoconch with up to 5 smooth whorls when multispiral. Radular formula 1–0-0-0-1 but radular apparatus absent in many species. Marginal teeth range from solid and curved to hypodermic, with or without small barb at the tip. Hypodermic teeth without solid bases. In some species (e.g. Impages hectica) the walls of the teeth are penetrated by numerous holes.

Included genera (After Bratcher & Cernohorsky, 1987; Terryn, 2007; Terryn & Holford, 2008):

  • Cinguloterebra Oyama, 19613

  • Clathroterebra Oyama, 19613

  • Duplicaria Dall, 19081

    •  [= Diplomeriza Dall, 1919]

    •  [= Myurellisca Bartsch, 1923]

    •  [= Pervicacia Iredale, 1924]

  • Euterebra Cotton & Godfrey, 19323

    •  [= Gradaterebra Cotton & Godfrey, 1932]

    •  [= Partecosta Dance & Eames, 1966]

  • Granuliterebra Oyama, 19611

  • HastulaH. & A. Adams, 18533

    •  [= Acuminia Dall, 1908]

    •  [= Egentelaria Rehder, 1980]

    •  [= Hastulina Oyama, 1961]

  • Hastulopsis Oyama, 19613

  • Impages E.A. Smith, 18731

  • Microtrypetes Pilsbry & Lowe, 19321

  • Myurella Hinds, 18453

    •  [= Decorihastula Oyama, 1961]

  • Oxymeris Dall, 19033

    •  [= Acus Gray, 1847 (non Müller, 1774)]

    •  [= Nototerebra Cotton, 1947]

  • Pellifronia Terryn & Holford, 20081

  • Perirhoe Dall, 19081

    •  [= Abretiella Bartsch, 1923]

    •  [= Dimidacus Iredale, 1929]

  • Pristiterebra Taki & Oyama, 19541

    •  [= Laeviacus Oyama, 1961]

  • Strioterebrum Sacco, 18913

    •  [= Brevimyurella Oyama, 1961]

    •  [= Punctoterebra Bartsch, 1923]

  • Terebra Bruguière, 17893

    •  [= Myurellina Bartsch, 1923]

    •  [= Panaterebra Olsson, 1967]

    •  [= Paraterebra Woodring, 1928]

    •  [= Subula Schumacher, 1817]

    •  [= Terebrum Montfort, 1810]

  • Terenolla Iredale, 19293

  • Triplostephanus Dall, 19081

ACKNOWLEDGEMENTS

The authors thank B.A. Marshall, M.G. Harasewych, R.N. Kilburn, M.J. Tenorio, J.K. Tucker and D. Tippett for constructive comments and help in completing the lists of genera and subgenera. A.E. Fedosov, J.D. Taylor, J. McLean, M.G. Harasewych and D.L. Tippett supplied shell and radula pictures. J.A. Todd provided specimens for dissection and photography. Barbara Buge curated and photographed the voucher specimens in MNHN.

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APPENDIX 1: INDEX OF GENERA AND SUBGENERA

Genus-group taxa removed from Conoidea are preceded by an asterisk.

(Sub) genus Family 
Abretiella Terebridae 
Abyssaforia ?Cochlespiridae 
Abyssobela Raphitomidae 
Abyssocomitas ?Pseudomelatomidae 
Abyssothauma ?Raphitomidae 
Acamptodaphne Raphitomidae 
Acanthodaphne Raphitomidae 
Acinodrillia ?Drilliidae 
Acmaturris Mangeliidae 
Acrista Clathurellidae 
Acropota Borsoniidae 
Acuminia Terebridae 
Acus Terebridae 
Adanaclava ?Clathurellidae 
Aforia ?Cochlespiridae 
Africonus Conidae 
Agathotoma Mangeliidae 
Agladrillia Drilliidae 
Aguilaria Pseudomelatomidae 
Aliceia Raphitomidae 
Allo ?Raphitomidae 
Alticlavatula Horaiclavidae 
Anacithara Horaiclavidae 
Anarithma Mitromorphidae 
Ancistrosyrinx Cochlespiridae 
Anguloclavus Horaiclavidae 
Annulaturris Turridae 
Anomalotomella Raphitomidae 
Anticlinura Mangeliidae 
Anticomitas ?Pseudomelatomidae 
Antiguraleus Mangeliidae 
Antimelatoma Pseudomelatomidae 
*Antimitra Buccinidae 
Antiplanes Pseudomelatomidae 
Aoteadrillia ?Horaiclavidae 
Apaturris ?Borsoniidae 
Aphanitoma Borsoniidae 
Apiotoma ?Cochlespiridae 
Apispiralia Mangeliidae 
Apitua Mangeliidae 
Arielia Mitromorphidae 
Artemidiconus Conorbidae 
Asperdaphne Raphitomidae 
Aspertilla Raphitomidae 
Asprella Conidae 
Asthenotoma Borsoniidae 
Austrocarina ?Horaiclavidae 
Austroconus Conidae 
Austrodaphnella Raphitomidae 
Austrodrillia Horaiclavidae 
Austrogemmula Turridae 
Austropusilla ?Raphitomidae 
Austrotoma Raphitomidae 
Austroturris Borsoniidae 
Azorilla Raphitomidae 
Azorita Raphitomidae 
Bactrocythara Mangeliidae 
Bathybela Raphitomidae 
Bathybermudia Turridae 
*Bathyclionella Buccinidae 
Bathyconus Conidae 
Bathypota Raphitomidae 
Bathytoma Borsoniidae 
Bela Mangeliidae 
Belalora ?Mangeliidae 
Belaturricula ?Borsoniidae 
Bellacythara Mangeliidae 
Bellaspira Drilliidae 
Bellatula Clathurellidae 
*Belomitra Buccinidae 
Benthoclionella Clavatulidae 
Benthodaphne ?Pseudomelatomidae 
Benthofascis Conorbidae 
Benthomangelia Mangeliidae 
Boettgeriola Borsoniidae 
Borsonella Borsoniidae 
Borsonellopsis Borsoniidae 
Borsonia Borsoniidae 
Brachycythara Mangeliidae 
Brachytoma ?Pseudomelatomidae 
Brephodrillia Drilliidae 
Brevimyurella Terebridae 
Buccinaria Raphitomidae 
Buchema Horaiclavidae 
Burchia Pseudomelatomidae 
Buridrillia Pseudomelatomidae 
Cacodaphnella ?Mangeliidae 
Calamiconus Conidae 
Calcatodrillia Pseudomelatomidae 
Calibanus Conidae 
Caliendrula Clavatulidae 
Californiconus Conidae 
Calliclava Drilliidae 
Calusaconus Conidae 
Canetoma Mangeliidae 
Cariboconus Conidae 
Carinapex Horaiclavidae 
Carinodrillia Pseudomelatomidae 
Carinoturris ?Pseudomelatomidae 
Cenodagreutes Raphitomidae 
Ceritoturris Horaiclavidae 
Cerodrillia Drilliidae 
Cestoma Mangeliidae 
Chelyconus Conidae 
Cheungbeia Pseudomelatomidae 
Cinguloterebra Terebridae 
Citharomangelia Mangeliidae 
Clamturris Turridae 
Clathrodrillia Drilliidae 
Clathromangelia ?Clathurellidae 
Clathroterebra Terebridae 
Clathurella Clathurellidae 
Clathurina Raphitomidae 
Clavatula Clavatulidae 
Clavicantha Drilliidae 
Clavosurcula ?Cochlespiridae 
Clavus Drilliidae 
Clebula Conidae 
Cleospira Strictispriridae 
Clinura Raphitomidae 
Clinuromella Mangeliidae 
Clionella Clavatulidae 
Cochlespira Cochlespiridae 
Cochlioconus Raphitomidae 
Comarmondia Clathurellidae 
Comitas Pseudomelatomidae 
Compsodrillia Pseudomelatomidae 
Conasprella Conidae 
Conasprelloides Conidae 
Conopleura Drilliidae 
Conorbela ?Pseudomelatomidae 
Conticosta Pseudomelatomidae 
Conus Conidae 
Cordieria Borsoniidae 
Corinnaeturris Clathurellidae 
Cornutoconus Conidae 
Coronacomitas Horaiclavidae 
Coronasyrinx Cochlespiridae 
Coronaxis Conidae 
Crassiclava Pseudomelatomidae 
Crassispira Pseudomelatomidae 
Crassispirella Pseudomelatomidae 
Crassopleura Drilliidae 
Cretaspira Pseudomelatomidae 
Crockerella Clathurellidae 
Cruziturricula ?Drilliidae 
Cryoturris Mangeliidae 
Cryptodaphne Raphitomidae 
Cryptogemma Turridae 
*Cryptomitra Buccinidae 
Cucullites Conidae 
Cucullus Conidae 
Curtitoma Mangeliidae 
Cylinder Conidae 
Cylindrella Conidae 
Cylindrus Conidae 
Cymakra Mitromorphidae 
Cymatosyrinx Drilliidae 
Cyrtocythara Mangeliidae 
Cyrtoides Raphitomidae 
Cythara ?Mangeliidae 
Cytharella Mangeliidae 
Cytharoclavus Horaiclavidae 
Cytharopsis Mangeliidae 
Dallaforia ?Cochlespiridae 
Dalliconus Conidae 
Dallspira Pseudomelatomidae 
Danilacarina Cochlespiridae 
Daphnella Raphitomidae 
*Daphnellopsis Muricidae 
Darbya Borsoniidae 
Darioconus Conidae 
Darrylia ?Horaiclavidae 
Dauciconus Conidae 
Decollidrillia Turridae 
Decorihastula Terebridae 
Dendroconus Conidae 
Diaugasma Raphitomidae 
Dimidacus Terebridae 
Diplomeriza Terebridae 
Diptychophlia Borsoniidae 
Douglassia Drilliidae 
Doxospira Pseudomelatomidae 
Drillia Drilliidae 
Drilliola Borsoniidae 
Ductoconus Conidae 
Duplicaria Terebridae 
Dyraspis Conidae 
Egentelaria Terebridae 
Elaeocyma Drilliidae 
Eldridgea Drilliidae 
Embrikena Conidae 
Endemoconus Conidae 
Epideira Pseudomelatomidae 
Epidirella Turridae 
Epidirona Pseudomelatomidae 
Eremiconus Conidae 
Erythroconus Conidae 
Etrema Clathurellidae 
Etremopa Clathurellidae 
Etremopsis Clathurellidae 
Eubela Raphitomidae 
Eucithara Mangeliidae 
Euclathurella Clathurellidae 
Eucyclotoma Raphitomidae 
Eudaphnella Raphitomidae 
Eugemmula Turridae 
Eugeniconus Conidae 
Euglyphostoma Clathurellidae 
Euguraleus Mangeliidae 
Eumetadrillia Drilliidae 
Euryentmema ?Mangeliidae 
Euterebra Terebridae 
Exomilus ?Raphitomidae 
Famelica Raphitomidae 
Favriella Raphitomidae 
Fehria Mangeliidae 
Feliciella ?Raphitomidae 
Fenestrosyrinx Raphitomidae 
Fenimorea Drilliidae 
Filodrillia Borsoniidae 
Floraconus Conidae 
Fulgiconus Conidae 
Fumiconus Conidae 
Funa Pseudomelatomidae 
Funitoma Mangeliidae 
Fusiconus Conidae 
Fusidaphne ?Raphitomidae 
Fusisyrinx ?Drilliidae 
Fusiturricula ?Drilliidae 
Fusiturris ?Clavatulidae 
Gastridium Conidae 
Gemmula Turridae 
Gemmuloborsonia ?Clavatulidae 
Genota Borsoniidae 
Genotia Borsoniidae 
Genotina Mangeliidae 
Genuanoconus Conidae 
Gibbaspira Pseudomelatomidae 
Gingicithara Mangeliidae 
Ginnania Mangeliidae 
Gladioconus Conidae 
Globiconus Conidae 
Globidrillia Drilliidae 
Glossispira Pseudomelatomidae 
Glyphostoma Clathurellidae 
Glyphostomoides Raphitomidae 
Glyphostomops Clathurellidae 
Glyphoturris Mangeliidae 
Glyptaesopus ?Borsoniidae 
Graciliclava Horaiclavidae 
Gradaterebra Terebridae 
Gradiconus Conidae 
Granotoma Mangeliidae 
Granoturris Mangeliidae 
Granuliterebra Terebridae 
Graphiconus Conidae 
Guraleus Mangeliidae 
Gymnobela Raphitomidae 
Haedropleura Horaiclavidae 
Harmoniconus Conidae 
Hastula Terebridae 
Hastulina Terebridae 
Hastulopsis Terebridae 
Hauturua Drilliidae 
Helenella Mitromorphidae 
Hemicythara Mangeliidae 
Hemidaphne Raphitomidae 
Hemilienardia Raphitomidae 
Hermes Conidae 
Heroconus Conidae 
Heterocithara Mangeliidae 
Heteroturris Borsoniidae 
Hindsiclava Pseudomelatomidae 
Horaiclavus Horaiclavidae 
Hormospira Pseudomelatomidae 
Imaclava Drilliidae 
Impages Terebridae 
Inkinga Horaiclavidae 
Inodrillara Horaiclavidae 
Inodrillia Horaiclavidae 
Inodrillina Horaiclavidae 
Inquisitor Pseudomelatomidae 
Iotyrris Turridae 
Iraquetrema Clathurellidae 
Iredalea Drilliidae 
Irenosyrinx Cochlespiridae 
Isodaphne Raphitomidae 
Ithycythara Mangeliidae 
Itia ?Mitromorphidae 
Iwaoa Horaiclavidae 
Jaspidiconus Conidae 
*Jumala Buccinidae 
Kalloconus Conidae 
Kenyonia Conidae 
Kermasprella Conidae 
Kermia Raphitomidae 
Ketyconus Conidae 
Kioconus Conidae 
Knefastia Pseudomelatomidae 
Kohniconus Conidae 
Kuradoconus Conidae 
Kurilohadalia ?Pseudomelatomidae 
Kurodadrillia Pseudomelatomidae 
Kuroshiodaphne Raphitomidae 
Kuroshioturris Turridae 
Kurtzia Mangeliidae 
Kurtziella Mangeliidae 
Kurtzina Mangeliidae 
Kylix Drilliidae 
Laeviacus Terebridae 
Lamniconus Conidae 
Lautoconus Conidae 
Leiocithara Mangeliidae 
Leiosyrinx Raphitomidae 
Leporiconus Conidae 
Leptadrillia Drilliidae 
Leptoconus Conidae 
Leucosyrinx ?Pseudomelatomidae 
Leufroyia Raphitomidae 
Lienardia Clathurellidae 
Lilliconus Conidae 
Lindaconus Conidae 
Lineotoma Raphitomidae 
Lioglyphostoma Pseudomelatomidae 
Lioglyphostomella Pseudomelatomidae 
Liracraea Mangeliidae 
Lissodrillia Drilliidae 
Litachilus Raphitomidae 
Lithoconus Conidae 
Lividoconus Conidae 
Lizaconus Conidae 
Lophiotoma Turridae 
Lophioturris Turridae 
Lorabela ?Mangeliidae 
Lovellona Mitromorphidae 
Lucerapex ?Turridae 
Lusitanops Raphitomidae 
Lyromangelia Mangeliidae 
Macteola ?Mangeliidae 
Maesiella Pseudomelatomidae 
Magelliconus Conidae 
Magnella Raphitomidae 
Majox Raphitomidae 
Makiyamaia Clavatulidae 
Mamiconus Conidae 
MammillaedrilliPseudomelatomidae 
Mangelia Mangeliidae 
Mangiliella Mangeliidae 
Maorimorpha Mitromorphidae 
Maoritomella Borsoniidae 
Marita Mangeliidae 
Marshallena ?Horaiclavidae 
Mauidrillia Horaiclavidae 
Megasurcula Pseudomelatomidae 
Meggittia ?Pseudomelatomidae 
Metaclathurella Raphitomidae 
Micantapex Borsoniidae 
Microdaphne Raphitomidae 
Microdrillia Borsoniidae 
Microgenia Raphitomidae 
Micropleurotom?Horaiclavidae 
Microtrypetes Terebridae 
Miliariconus Conidae 
Mioawateria Raphitomidae 
Miraclathurella Pseudomelatomidae 
Mitraguraleus Mangeliidae 
Mitrellatoma Mitromorphidae 
Mitrithara Mitromorphidae 
Mitrolumna Mitromorphidae 
Mitromorpha Mitromorphidae 
Monilispira Pseudomelatomidae 
Monteiroconus Conidae 
Mordica Raphitomidae 
Myurella Terebridae 
Myurellina Terebridae 
Myurellisca Terebridae 
Nannodiella Clathurellidae 
Narraweena Borsoniidae 
Naskia Horaiclavidae 
Nataliconus Conidae 
Naudedrillia Pseudomelatomidae 
Nematoma Mangeliidae 
Neodrillia Drilliidae 
Neoguraleus ?Mangeliidae 
Neopleurotomoides Raphitomidae 
Nepotilla Raphitomidae 
Netrum Clavatulidae 
Nihonia ?Cochlespiridae 
Nodotoma Mangeliidae 
Notocytharella ?Mangeliidae 
Nototerebra Terebridae 
Nquma Horaiclavidae 
Nymphispira Pseudomelatomidae 
Obesotoma Mangeliidae 
Oenopota Mangeliidae 
Oenopotella Mangeliidae 
Ongoconus Conidae 
Ootomella Raphitomidae 
Ophiodermella Borsoniidae 
Orrmaesia Drilliidae 
Otitoma Pseudomelatomidae 
Oxymeris Terebridae 
Oxytropa Turridae 
Pagodasyrinx Cochlespiridae 
Pagodidaphne Raphitomidae 
Panaterebra Terebridae 
Papillocithara Mangeliidae 
Parabathytoma Borsoniidae 
Paraborsonia Borsoniidae 
Paraclathurella Clathurellidae 
Paracomitas ?Pseudomelatomidae 
Paracuneus ?Drilliidae 
Paradaphne Raphitomidae 
Paradrillia Horaiclavidae 
Paramontana Raphitomidae 
Paraspirotropis ?Mangeliidae 
Paraterebra Terebridae 
Partecosta Terebridae 
Parviconus Conidae 
Pellifronia Terebridae 
Peratotoma Raphitomidae 
Perimangelia Mangeliidae 
Perirhoe Terebridae 
Perplexiconus Conidae 
Perrona Clavatulidae 
Pervicacia Terebridae 
Phasmoconus Conidae 
Phenatoma Borsoniidae 
Philbertia Raphitomidae 
Phymorhynchus Raphitomidae 
Pilsbryspira Pseudomelatomidae 
Pinguigemmula Turridae 
Pionoconus Conidae 
Pionotoma Raphitomidae 
Plagiostropha Drilliidae 
Platycythara Mangeliidae 
Pleurotoma Turridae 
Pleurotomella Raphitomidae 
Pleurotomus Turridae 
Plicaustraconus Conidae 
Plicisyrinx Pseudomelatomidae 
Polystira Turridae 
Pontiothauma Raphitomidae 
Pristiterebra Terebridae 
Profundiconus Conidae 
Propebela Mangeliidae 
Protoconus Conidae 
Protostrioconus Conidae 
Pseudexomilus ?Horaiclavidae 
Pseudoconorbis Conidae 
PseudodaphnellRaphitomidae 
Pseudoetrema Clathurellidae 
PseudolilliconuConidae 
PseudomelatomPseudomelatomidae 
Pseudonoduloconus Conidae 
Pseudorhaphitoma Mangeliidae 
Pseudotaranis ?Pseudomelatomidae 
Psittacodrillia Horaiclavidae 
Ptychobela Pseudomelatomidae 
Ptychosyrinx Turridae 
Pulsarella Borsoniidae 
Puncticulis Conidae 
Punctoterebra Terebridae 
Purpuriconus Conidae 
Pusionella Clavatulidae 
Pyrgocythara Mangeliidae 
Pyrgospira Pseudomelatomidae 
Pyruconus Conidae 
Qii Raphitomidae 
Quasiconus Conidae 
Raphitoma Raphitomidae 
Rectiplanes Pseudomelatomidae 
Rectisulcus Pseudomelatomidae 
Regiconus Conidae 
Regidrillia Horaiclavidae 
Retidrillia Borsoniidae 
Rhizoconus Conidae 
Rhodopetoma ?Pseudomelatomidae 
Rhombiconus Conidae 
Rhombus Conidae 
Rimosodaphnella Raphitomidae 
Rissomangelia Mangeliidae 
Riuguhdrillia Borsoniidae 
Rocroithys Raphitomidae 
Rolaniconus Conidae 
Rollus Conidae 
Rubellatoma Mangeliidae 
Rugocythara Mangeliidae 
Saccharoturris Mangeliidae 
Scaevatula Clavatulidae 
Sciteconus Conidae 
Scrinium Mitromorphidae 
Sediliopsis Pseudomelatomidae 
Seminoleconus Conidae 
Shutonia ?Pseudomelatomidae 
Sibogasyrinx Cochlespiridae 
Smithiella Mangeliidae 
Socioconus Conidae 
Speoides Raphitomidae 
Spergo Raphitomidae 
Spirotropis Drilliidae 
Splendrillia Drilliidae 
Splinoconus Conidae 
Spuriconus Conidae 
Steiraxis ?Cochlespiridae 
*Steironepion Columbellidae 
Stellaconus Conidae 
Stellatoma Mangeliidae 
Stenodrillia ?Drilliidae 
Stephanoconus Conidae 
Stilla Raphitomidae 
Strategoconus Conidae 
Striatoguraleus Horaiclavidae 
Strictispira Strictispriridae 
Strioconus Conidae 
Striospira Pseudomelatomidae 
Strioterebrum Terebridae 
Strombinoturris Clathurellidae 
Suavodrillia Borsoniidae 
Subula Terebridae 
Sugitanitoma Horaiclavidae 
Surcula Clavatulidae 
*Surculina Turbinellidae 
Suturocythara Mangeliidae 
Syntomodrillia Drilliidae 
Taranidaphne Raphitomidae 
Taranis Raphitomidae 
Taranteconus Conidae 
Tasmadaphne Raphitomidae 
Teleochilus ?Raphitomidae 
Tenaturris Mangeliidae 
Terebra Terebridae 
Terebrum Terebridae 
Terenolla Terebridae 
Teretia Raphitomidae 
Teretiopsis Raphitomidae 
Tesselliconus Conidae 
Textilia Conidae 
Thatcheria Raphitomidae 
Thatcheriasyrinx ?Raphitomidae 
Thatcherina Raphitomidae 
Thelecythara Horaiclavidae 
Thelecytharella Pseudomelatomidae 
Theliconus Conidae 
Thesbia ?Raphitomidae 
Theta Raphitomidae 
Thetidos Clathurellidae 
Thoraconus Conidae 
Tiariturris Pseudomelatomidae 
Tomellana Clavatulidae 
Tomopleura Borsoniidae 
Toxiclionella Clavatulidae 
Toxicochlespira Mangeliidae 
Trachydrillida Clavatulidae 
Triplostephanus Terebridae 
Tritonoturris Raphitomidae 
Tropidoturris Borsoniidae 
Trovaoconus Conidae 
Truncadaphne Raphitomidae 
Tuliparia Conidae 
Turrella Clathurellidae 
Turrhyssa Raphitomidae 
Turriconus Conidae 
Turricula Clavatulidae 
Turridrupa Turridae 
Turrigemma Pseudomelatomidae 
*Turrijaumelia Columbellidae 
Turris Turridae 
Turritomella Mangeliidae 
Tuskaroria Raphitomidae 
Tylotia Drilliidae 
Tylotiella Drilliidae 
Typhlodaphne ?Borsoniidae 
Typhlomangelia Borsoniidae 
Typhlosyrinx Raphitomidae 
Tyrrhenoturris Clavatulidae 
Unedogemmula Turridae 
Utriculus Conidae 
Varioconus Conidae 
Venustoma ?Mangeliidae 
Veprecula Raphitomidae 
Vepridaphne Raphitomidae 
Vexiariella Mitromorphidae 
Vexithara Borsoniidae 
Vexitomina Horaiclavidae 
Villiersiella Mangeliidae 
Viminiconus Conidae 
Virgiconus Conidae 
Viridoturris Borsoniidae 
Viridrillia Pseudomelatomidae 
Viridrillina Pseudomelatomidae 
Virroconus Conidae 
Vitjazinella Mangeliidae 
Vitricythara Mangeliidae 
Vituliconus Conidae 
Watsonaria Raphitomidae 
Widalli Mangeliidae 
Xanthodaphne Raphitomidae 
Xenuroturris Turridae 
Ximeniconus Conidae 
Yeddoconus Conidae 
Zemacies Borsoniidae 
Zenepos Raphitomidae 
*Zetekia Columbellidae 
Zonulispira Pseudomelatomidae 
(Sub) genus Family 
Abretiella Terebridae 
Abyssaforia ?Cochlespiridae 
Abyssobela Raphitomidae 
Abyssocomitas ?Pseudomelatomidae 
Abyssothauma ?Raphitomidae 
Acamptodaphne Raphitomidae 
Acanthodaphne Raphitomidae 
Acinodrillia ?Drilliidae 
Acmaturris Mangeliidae 
Acrista Clathurellidae 
Acropota Borsoniidae 
Acuminia Terebridae 
Acus Terebridae 
Adanaclava ?Clathurellidae 
Aforia ?Cochlespiridae 
Africonus Conidae 
Agathotoma Mangeliidae 
Agladrillia Drilliidae 
Aguilaria Pseudomelatomidae 
Aliceia Raphitomidae 
Allo ?Raphitomidae 
Alticlavatula Horaiclavidae 
Anacithara Horaiclavidae 
Anarithma Mitromorphidae 
Ancistrosyrinx Cochlespiridae 
Anguloclavus Horaiclavidae 
Annulaturris Turridae 
Anomalotomella Raphitomidae 
Anticlinura Mangeliidae 
Anticomitas ?Pseudomelatomidae 
Antiguraleus Mangeliidae 
Antimelatoma Pseudomelatomidae 
*Antimitra Buccinidae 
Antiplanes Pseudomelatomidae 
Aoteadrillia ?Horaiclavidae 
Apaturris ?Borsoniidae 
Aphanitoma Borsoniidae 
Apiotoma ?Cochlespiridae 
Apispiralia Mangeliidae 
Apitua Mangeliidae 
Arielia Mitromorphidae 
Artemidiconus Conorbidae 
Asperdaphne Raphitomidae 
Aspertilla Raphitomidae 
Asprella Conidae 
Asthenotoma Borsoniidae 
Austrocarina ?Horaiclavidae 
Austroconus Conidae 
Austrodaphnella Raphitomidae 
Austrodrillia Horaiclavidae 
Austrogemmula Turridae 
Austropusilla ?Raphitomidae 
Austrotoma Raphitomidae 
Austroturris Borsoniidae 
Azorilla Raphitomidae 
Azorita Raphitomidae 
Bactrocythara Mangeliidae 
Bathybela Raphitomidae 
Bathybermudia Turridae 
*Bathyclionella Buccinidae 
Bathyconus Conidae 
Bathypota Raphitomidae 
Bathytoma Borsoniidae 
Bela Mangeliidae 
Belalora ?Mangeliidae 
Belaturricula ?Borsoniidae 
Bellacythara Mangeliidae 
Bellaspira Drilliidae 
Bellatula Clathurellidae 
*Belomitra Buccinidae 
Benthoclionella Clavatulidae 
Benthodaphne ?Pseudomelatomidae 
Benthofascis Conorbidae 
Benthomangelia Mangeliidae 
Boettgeriola Borsoniidae 
Borsonella Borsoniidae 
Borsonellopsis Borsoniidae 
Borsonia Borsoniidae 
Brachycythara Mangeliidae 
Brachytoma ?Pseudomelatomidae 
Brephodrillia Drilliidae 
Brevimyurella Terebridae 
Buccinaria Raphitomidae 
Buchema Horaiclavidae 
Burchia Pseudomelatomidae 
Buridrillia Pseudomelatomidae 
Cacodaphnella ?Mangeliidae 
Calamiconus Conidae 
Calcatodrillia Pseudomelatomidae 
Calibanus Conidae 
Caliendrula Clavatulidae 
Californiconus Conidae 
Calliclava Drilliidae 
Calusaconus Conidae 
Canetoma Mangeliidae 
Cariboconus Conidae 
Carinapex Horaiclavidae 
Carinodrillia Pseudomelatomidae 
Carinoturris ?Pseudomelatomidae 
Cenodagreutes Raphitomidae 
Ceritoturris Horaiclavidae 
Cerodrillia Drilliidae 
Cestoma Mangeliidae 
Chelyconus Conidae 
Cheungbeia Pseudomelatomidae 
Cinguloterebra Terebridae 
Citharomangelia Mangeliidae 
Clamturris Turridae 
Clathrodrillia Drilliidae 
Clathromangelia ?Clathurellidae 
Clathroterebra Terebridae 
Clathurella Clathurellidae 
Clathurina Raphitomidae 
Clavatula Clavatulidae 
Clavicantha Drilliidae 
Clavosurcula ?Cochlespiridae 
Clavus Drilliidae 
Clebula Conidae 
Cleospira Strictispriridae 
Clinura Raphitomidae 
Clinuromella Mangeliidae 
Clionella Clavatulidae 
Cochlespira Cochlespiridae 
Cochlioconus Raphitomidae 
Comarmondia Clathurellidae 
Comitas Pseudomelatomidae 
Compsodrillia Pseudomelatomidae 
Conasprella Conidae 
Conasprelloides Conidae 
Conopleura Drilliidae 
Conorbela ?Pseudomelatomidae 
Conticosta Pseudomelatomidae 
Conus Conidae 
Cordieria Borsoniidae 
Corinnaeturris Clathurellidae 
Cornutoconus Conidae 
Coronacomitas Horaiclavidae 
Coronasyrinx Cochlespiridae 
Coronaxis Conidae 
Crassiclava Pseudomelatomidae 
Crassispira Pseudomelatomidae 
Crassispirella Pseudomelatomidae 
Crassopleura Drilliidae 
Cretaspira Pseudomelatomidae 
Crockerella Clathurellidae 
Cruziturricula ?Drilliidae 
Cryoturris Mangeliidae 
Cryptodaphne Raphitomidae 
Cryptogemma Turridae 
*Cryptomitra Buccinidae 
Cucullites Conidae 
Cucullus Conidae 
Curtitoma Mangeliidae 
Cylinder Conidae 
Cylindrella Conidae 
Cylindrus Conidae 
Cymakra Mitromorphidae 
Cymatosyrinx Drilliidae 
Cyrtocythara Mangeliidae 
Cyrtoides Raphitomidae 
Cythara ?Mangeliidae 
Cytharella Mangeliidae 
Cytharoclavus Horaiclavidae 
Cytharopsis Mangeliidae 
Dallaforia ?Cochlespiridae 
Dalliconus Conidae 
Dallspira Pseudomelatomidae 
Danilacarina Cochlespiridae 
Daphnella Raphitomidae 
*Daphnellopsis Muricidae 
Darbya Borsoniidae 
Darioconus Conidae 
Darrylia ?Horaiclavidae 
Dauciconus Conidae 
Decollidrillia Turridae 
Decorihastula Terebridae 
Dendroconus Conidae 
Diaugasma Raphitomidae 
Dimidacus Terebridae 
Diplomeriza Terebridae 
Diptychophlia Borsoniidae 
Douglassia Drilliidae 
Doxospira Pseudomelatomidae 
Drillia Drilliidae 
Drilliola Borsoniidae 
Ductoconus Conidae 
Duplicaria Terebridae 
Dyraspis Conidae 
Egentelaria Terebridae 
Elaeocyma Drilliidae 
Eldridgea Drilliidae 
Embrikena Conidae 
Endemoconus Conidae 
Epideira Pseudomelatomidae 
Epidirella Turridae 
Epidirona Pseudomelatomidae 
Eremiconus Conidae 
Erythroconus Conidae 
Etrema Clathurellidae 
Etremopa Clathurellidae 
Etremopsis Clathurellidae 
Eubela Raphitomidae 
Eucithara Mangeliidae 
Euclathurella Clathurellidae 
Eucyclotoma Raphitomidae 
Eudaphnella Raphitomidae 
Eugemmula Turridae 
Eugeniconus Conidae 
Euglyphostoma Clathurellidae 
Euguraleus Mangeliidae 
Eumetadrillia Drilliidae 
Euryentmema ?Mangeliidae 
Euterebra Terebridae 
Exomilus ?Raphitomidae 
Famelica Raphitomidae 
Favriella Raphitomidae 
Fehria Mangeliidae 
Feliciella ?Raphitomidae 
Fenestrosyrinx Raphitomidae 
Fenimorea Drilliidae 
Filodrillia Borsoniidae 
Floraconus Conidae 
Fulgiconus Conidae 
Fumiconus Conidae 
Funa Pseudomelatomidae 
Funitoma Mangeliidae 
Fusiconus Conidae 
Fusidaphne ?Raphitomidae 
Fusisyrinx ?Drilliidae 
Fusiturricula ?Drilliidae 
Fusiturris ?Clavatulidae 
Gastridium Conidae 
Gemmula Turridae 
Gemmuloborsonia ?Clavatulidae 
Genota Borsoniidae 
Genotia Borsoniidae 
Genotina Mangeliidae 
Genuanoconus Conidae 
Gibbaspira Pseudomelatomidae 
Gingicithara Mangeliidae 
Ginnania Mangeliidae 
Gladioconus Conidae 
Globiconus Conidae 
Globidrillia Drilliidae 
Glossispira Pseudomelatomidae 
Glyphostoma Clathurellidae 
Glyphostomoides Raphitomidae 
Glyphostomops Clathurellidae 
Glyphoturris Mangeliidae 
Glyptaesopus ?Borsoniidae 
Graciliclava Horaiclavidae 
Gradaterebra Terebridae 
Gradiconus Conidae 
Granotoma Mangeliidae 
Granoturris Mangeliidae 
Granuliterebra Terebridae 
Graphiconus Conidae 
Guraleus Mangeliidae 
Gymnobela Raphitomidae 
Haedropleura Horaiclavidae 
Harmoniconus Conidae 
Hastula Terebridae 
Hastulina Terebridae 
Hastulopsis Terebridae 
Hauturua Drilliidae 
Helenella Mitromorphidae 
Hemicythara Mangeliidae 
Hemidaphne Raphitomidae 
Hemilienardia Raphitomidae 
Hermes Conidae 
Heroconus Conidae 
Heterocithara Mangeliidae 
Heteroturris Borsoniidae 
Hindsiclava Pseudomelatomidae 
Horaiclavus Horaiclavidae 
Hormospira Pseudomelatomidae 
Imaclava Drilliidae 
Impages Terebridae 
Inkinga Horaiclavidae 
Inodrillara Horaiclavidae 
Inodrillia Horaiclavidae 
Inodrillina Horaiclavidae 
Inquisitor Pseudomelatomidae 
Iotyrris Turridae 
Iraquetrema Clathurellidae 
Iredalea Drilliidae 
Irenosyrinx Cochlespiridae 
Isodaphne Raphitomidae 
Ithycythara Mangeliidae 
Itia ?Mitromorphidae 
Iwaoa Horaiclavidae 
Jaspidiconus Conidae 
*Jumala Buccinidae 
Kalloconus Conidae 
Kenyonia Conidae 
Kermasprella Conidae 
Kermia Raphitomidae 
Ketyconus Conidae 
Kioconus Conidae 
Knefastia Pseudomelatomidae 
Kohniconus Conidae 
Kuradoconus Conidae 
Kurilohadalia ?Pseudomelatomidae 
Kurodadrillia Pseudomelatomidae 
Kuroshiodaphne Raphitomidae 
Kuroshioturris Turridae 
Kurtzia Mangeliidae 
Kurtziella Mangeliidae 
Kurtzina Mangeliidae 
Kylix Drilliidae 
Laeviacus Terebridae 
Lamniconus Conidae 
Lautoconus Conidae 
Leiocithara Mangeliidae 
Leiosyrinx Raphitomidae 
Leporiconus Conidae 
Leptadrillia Drilliidae 
Leptoconus Conidae 
Leucosyrinx ?Pseudomelatomidae 
Leufroyia Raphitomidae 
Lienardia Clathurellidae 
Lilliconus Conidae 
Lindaconus Conidae 
Lineotoma Raphitomidae 
Lioglyphostoma Pseudomelatomidae 
Lioglyphostomella Pseudomelatomidae 
Liracraea Mangeliidae 
Lissodrillia Drilliidae 
Litachilus Raphitomidae 
Lithoconus Conidae 
Lividoconus Conidae 
Lizaconus Conidae 
Lophiotoma Turridae 
Lophioturris Turridae 
Lorabela ?Mangeliidae 
Lovellona Mitromorphidae 
Lucerapex ?Turridae 
Lusitanops Raphitomidae 
Lyromangelia Mangeliidae 
Macteola ?Mangeliidae 
Maesiella Pseudomelatomidae 
Magelliconus Conidae 
Magnella Raphitomidae 
Majox Raphitomidae 
Makiyamaia Clavatulidae 
Mamiconus Conidae 
MammillaedrilliPseudomelatomidae 
Mangelia Mangeliidae 
Mangiliella Mangeliidae 
Maorimorpha Mitromorphidae 
Maoritomella Borsoniidae 
Marita Mangeliidae 
Marshallena ?Horaiclavidae 
Mauidrillia Horaiclavidae 
Megasurcula Pseudomelatomidae 
Meggittia ?Pseudomelatomidae 
Metaclathurella Raphitomidae 
Micantapex Borsoniidae 
Microdaphne Raphitomidae 
Microdrillia Borsoniidae 
Microgenia Raphitomidae 
Micropleurotom?Horaiclavidae 
Microtrypetes Terebridae 
Miliariconus Conidae 
Mioawateria Raphitomidae 
Miraclathurella Pseudomelatomidae 
Mitraguraleus Mangeliidae 
Mitrellatoma Mitromorphidae 
Mitrithara Mitromorphidae 
Mitrolumna Mitromorphidae 
Mitromorpha Mitromorphidae 
Monilispira Pseudomelatomidae 
Monteiroconus Conidae 
Mordica Raphitomidae 
Myurella Terebridae 
Myurellina Terebridae 
Myurellisca Terebridae 
Nannodiella Clathurellidae 
Narraweena Borsoniidae 
Naskia Horaiclavidae 
Nataliconus Conidae 
Naudedrillia Pseudomelatomidae 
Nematoma Mangeliidae 
Neodrillia Drilliidae 
Neoguraleus ?Mangeliidae 
Neopleurotomoides Raphitomidae 
Nepotilla Raphitomidae 
Netrum Clavatulidae 
Nihonia ?Cochlespiridae 
Nodotoma Mangeliidae 
Notocytharella ?Mangeliidae 
Nototerebra Terebridae 
Nquma Horaiclavidae 
Nymphispira Pseudomelatomidae 
Obesotoma Mangeliidae 
Oenopota Mangeliidae 
Oenopotella Mangeliidae 
Ongoconus Conidae 
Ootomella Raphitomidae 
Ophiodermella Borsoniidae 
Orrmaesia Drilliidae 
Otitoma Pseudomelatomidae 
Oxymeris Terebridae 
Oxytropa Turridae 
Pagodasyrinx Cochlespiridae 
Pagodidaphne Raphitomidae 
Panaterebra Terebridae 
Papillocithara Mangeliidae 
Parabathytoma Borsoniidae 
Paraborsonia Borsoniidae 
Paraclathurella Clathurellidae 
Paracomitas ?Pseudomelatomidae 
Paracuneus ?Drilliidae 
Paradaphne Raphitomidae 
Paradrillia Horaiclavidae 
Paramontana Raphitomidae 
Paraspirotropis ?Mangeliidae 
Paraterebra Terebridae 
Partecosta Terebridae 
Parviconus Conidae 
Pellifronia Terebridae 
Peratotoma Raphitomidae 
Perimangelia Mangeliidae 
Perirhoe Terebridae 
Perplexiconus Conidae 
Perrona Clavatulidae 
Pervicacia Terebridae 
Phasmoconus Conidae 
Phenatoma Borsoniidae 
Philbertia Raphitomidae 
Phymorhynchus Raphitomidae 
Pilsbryspira Pseudomelatomidae 
Pinguigemmula Turridae 
Pionoconus Conidae 
Pionotoma Raphitomidae 
Plagiostropha Drilliidae 
Platycythara Mangeliidae 
Pleurotoma Turridae 
Pleurotomella Raphitomidae 
Pleurotomus Turridae 
Plicaustraconus Conidae 
Plicisyrinx Pseudomelatomidae 
Polystira Turridae 
Pontiothauma Raphitomidae 
Pristiterebra Terebridae 
Profundiconus Conidae 
Propebela Mangeliidae 
Protoconus Conidae 
Protostrioconus Conidae 
Pseudexomilus ?Horaiclavidae 
Pseudoconorbis Conidae 
PseudodaphnellRaphitomidae 
Pseudoetrema Clathurellidae 
PseudolilliconuConidae 
PseudomelatomPseudomelatomidae 
Pseudonoduloconus Conidae 
Pseudorhaphitoma Mangeliidae 
Pseudotaranis ?Pseudomelatomidae 
Psittacodrillia Horaiclavidae 
Ptychobela Pseudomelatomidae 
Ptychosyrinx Turridae 
Pulsarella Borsoniidae 
Puncticulis Conidae 
Punctoterebra Terebridae 
Purpuriconus Conidae 
Pusionella Clavatulidae 
Pyrgocythara Mangeliidae 
Pyrgospira Pseudomelatomidae 
Pyruconus Conidae 
Qii Raphitomidae 
Quasiconus Conidae 
Raphitoma Raphitomidae 
Rectiplanes Pseudomelatomidae 
Rectisulcus Pseudomelatomidae 
Regiconus Conidae 
Regidrillia Horaiclavidae 
Retidrillia Borsoniidae 
Rhizoconus Conidae 
Rhodopetoma ?Pseudomelatomidae 
Rhombiconus Conidae 
Rhombus Conidae 
Rimosodaphnella Raphitomidae 
Rissomangelia Mangeliidae 
Riuguhdrillia Borsoniidae 
Rocroithys Raphitomidae 
Rolaniconus Conidae 
Rollus Conidae 
Rubellatoma Mangeliidae 
Rugocythara Mangeliidae 
Saccharoturris Mangeliidae 
Scaevatula Clavatulidae 
Sciteconus Conidae 
Scrinium Mitromorphidae 
Sediliopsis Pseudomelatomidae 
Seminoleconus Conidae 
Shutonia ?Pseudomelatomidae 
Sibogasyrinx Cochlespiridae 
Smithiella Mangeliidae 
Socioconus Conidae 
Speoides Raphitomidae 
Spergo Raphitomidae 
Spirotropis Drilliidae 
Splendrillia Drilliidae 
Splinoconus Conidae 
Spuriconus Conidae 
Steiraxis ?Cochlespiridae 
*Steironepion Columbellidae 
Stellaconus Conidae 
Stellatoma Mangeliidae 
Stenodrillia ?Drilliidae 
Stephanoconus Conidae 
Stilla Raphitomidae 
Strategoconus Conidae 
Striatoguraleus Horaiclavidae 
Strictispira Strictispriridae 
Strioconus Conidae 
Striospira Pseudomelatomidae 
Strioterebrum Terebridae 
Strombinoturris Clathurellidae 
Suavodrillia Borsoniidae 
Subula Terebridae 
Sugitanitoma Horaiclavidae 
Surcula Clavatulidae 
*Surculina Turbinellidae 
Suturocythara Mangeliidae 
Syntomodrillia Drilliidae 
Taranidaphne Raphitomidae 
Taranis Raphitomidae 
Taranteconus Conidae 
Tasmadaphne Raphitomidae 
Teleochilus ?Raphitomidae 
Tenaturris Mangeliidae 
Terebra Terebridae 
Terebrum Terebridae 
Terenolla Terebridae 
Teretia Raphitomidae 
Teretiopsis Raphitomidae 
Tesselliconus Conidae 
Textilia Conidae 
Thatcheria Raphitomidae 
Thatcheriasyrinx ?Raphitomidae 
Thatcherina Raphitomidae 
Thelecythara Horaiclavidae 
Thelecytharella Pseudomelatomidae 
Theliconus Conidae 
Thesbia ?Raphitomidae 
Theta Raphitomidae 
Thetidos Clathurellidae 
Thoraconus Conidae 
Tiariturris Pseudomelatomidae 
Tomellana Clavatulidae 
Tomopleura Borsoniidae 
Toxiclionella Clavatulidae 
Toxicochlespira Mangeliidae 
Trachydrillida Clavatulidae 
Triplostephanus Terebridae 
Tritonoturris Raphitomidae 
Tropidoturris Borsoniidae 
Trovaoconus Conidae 
Truncadaphne Raphitomidae 
Tuliparia Conidae 
Turrella Clathurellidae 
Turrhyssa Raphitomidae 
Turriconus Conidae 
Turricula Clavatulidae 
Turridrupa Turridae 
Turrigemma Pseudomelatomidae 
*Turrijaumelia Columbellidae 
Turris Turridae 
Turritomella Mangeliidae 
Tuskaroria Raphitomidae 
Tylotia Drilliidae 
Tylotiella Drilliidae 
Typhlodaphne ?Borsoniidae 
Typhlomangelia Borsoniidae 
Typhlosyrinx Raphitomidae 
Tyrrhenoturris Clavatulidae 
Unedogemmula Turridae 
Utriculus Conidae 
Varioconus Conidae 
Venustoma ?Mangeliidae 
Veprecula Raphitomidae 
Vepridaphne Raphitomidae 
Vexiariella Mitromorphidae 
Vexithara Borsoniidae 
Vexitomina Horaiclavidae 
Villiersiella Mangeliidae 
Viminiconus Conidae 
Virgiconus Conidae 
Viridoturris Borsoniidae 
Viridrillia Pseudomelatomidae 
Viridrillina Pseudomelatomidae 
Virroconus Conidae 
Vitjazinella Mangeliidae 
Vitricythara Mangeliidae 
Vituliconus Conidae 
Watsonaria Raphitomidae 
Widalli Mangeliidae 
Xanthodaphne Raphitomidae 
Xenuroturris Turridae 
Ximeniconus Conidae 
Yeddoconus Conidae 
Zemacies Borsoniidae 
Zenepos Raphitomidae 
*Zetekia Columbellidae 
Zonulispira Pseudomelatomidae 

Appendix 2: List of all the family and subfamily names available for the Conoidea.

(see Bouchet and Rocroi, 2005 for details). The status of names including only fossil genera is not discussed

Acidae Gray, 1853 (nomenclaturally invalid) 
Andoniinae Vera-Pelaez, 2002 (fossil) 
Belinae Bellardi, 1875 (synonym of Mangeliidae) 
Borsoniidae Bellardi, 1875 
Brachytominae Thiele, 1929 (nomen dubium) 
Californiconinae Tucker & Tenorio, 2009 (synonym of Conidae) 
Clathurellidae H. Adams & A. Adams, 1858 
Clavatulidae Gray, 1853 
Clavinae Casey 1904 (nomenclaturally invalid) 
Clionellidae Stimpson, 1865 (synonym of Clavatulidae) 
Cochlespiridae Powell, 1942 
Conidae Fleming, 1822 
Conilithidae Tucker & Tenorio, 2009 (synonym of Conidae) 
Conorbidae de Gregorio 1880 
Conulinae Rafinesque, 1815 (nomenclaturally invalid) 
Crassispirinae McLean, 1971 (synonym of Pseudomelatomidae) 
Cryptoconinae Cossmann, 1896 (fossil) 
Cytharinae Thiele, 1929 (synonym of Mangeliidae) 
Daphnellinae Casey, 1904 (synonym of Raphitomidae) 
Defranciinae Gray, 1853 (nomenclaturally invalid) 
Diptychomitrinae Bellardi, 1888 (synonym of Mitromorphidae) 
Drilliidae Olsson, 1964 
Hemiconidae Tucker & Tenorio, 2009 (fossil) 
Horaiclavidae fam. nov. 
Johnwyattidae Serna, 1979 (fossil) 
Lorinae Thiele, 1925 sensu Opinion 666 (synonym of Clathurellidae) 
Lorinae Thiele, 1925 sensu Thiele (synonym of Mangeliidae) 
Mangeliidae Fischer, 1883 
Melatomidae Gill, 1871 (synonym of Clavatulidae) 
Mitrolumnidae Sacco, 1904 (synonym of Mitromorphidae) 
Mitromorphidae Casey, 1904 
Oenopotinae Bogdanov, 1987 (synonym of Mangeliidae) 
Pervicaciinae Rudman, 1969 (synonym of Terebridae) 
Pleurotominae Gray, 1838 (synonym of Turridae) 
Pleurotomellinae Nordsieck, 1968 (synonym of Raphitomidae) 
Pseudomelatomidae Morrison, 1965 
Pseudotominae Bellardi, 1875 (synonym of Borsoniidae) 
Puncticuliinae Tucker & Tenorio, 2009 (synonym of Conidae) 
Pusionellinae Gray, 1853 (synonym of Clavatulidae) 
Raphitomidae Bellardi, 1875 
Siphopsinae Le Renard, 1995 (fossil) 
Speightiidae Powell, 1942 (fossil; doubtfully Conoidea) 
Strictispiridae McLean, 1971 
Taraninae Casey, 1904 (synonym of Raphitomidae) 
Taranteconidae Tucker & Tenorio, 2009 (synonym of Conidae) 
Terebridae Mörch, 1852 
Thatcheriidae Powell, 1942 (synonym of Raphitomidae) 
Turriculinae Powell, 1942 (synonym of Clavatulidae) 
Turridae H. Adams & A. Adams, 1853 
Zemaciinae Sysoev, 2003 (synonym of Borsoniidae) 
Zonulispirinae McLean, 1971 (synonym of Pseudomelatomidae) 
Acidae Gray, 1853 (nomenclaturally invalid) 
Andoniinae Vera-Pelaez, 2002 (fossil) 
Belinae Bellardi, 1875 (synonym of Mangeliidae) 
Borsoniidae Bellardi, 1875 
Brachytominae Thiele, 1929 (nomen dubium) 
Californiconinae Tucker & Tenorio, 2009 (synonym of Conidae) 
Clathurellidae H. Adams & A. Adams, 1858 
Clavatulidae Gray, 1853 
Clavinae Casey 1904 (nomenclaturally invalid) 
Clionellidae Stimpson, 1865 (synonym of Clavatulidae) 
Cochlespiridae Powell, 1942 
Conidae Fleming, 1822 
Conilithidae Tucker & Tenorio, 2009 (synonym of Conidae) 
Conorbidae de Gregorio 1880 
Conulinae Rafinesque, 1815 (nomenclaturally invalid) 
Crassispirinae McLean, 1971 (synonym of Pseudomelatomidae) 
Cryptoconinae Cossmann, 1896 (fossil) 
Cytharinae Thiele, 1929 (synonym of Mangeliidae) 
Daphnellinae Casey, 1904 (synonym of Raphitomidae) 
Defranciinae Gray, 1853 (nomenclaturally invalid) 
Diptychomitrinae Bellardi, 1888 (synonym of Mitromorphidae) 
Drilliidae Olsson, 1964 
Hemiconidae Tucker & Tenorio, 2009 (fossil) 
Horaiclavidae fam. nov. 
Johnwyattidae Serna, 1979 (fossil) 
Lorinae Thiele, 1925 sensu Opinion 666 (synonym of Clathurellidae) 
Lorinae Thiele, 1925 sensu Thiele (synonym of Mangeliidae) 
Mangeliidae Fischer, 1883 
Melatomidae Gill, 1871 (synonym of Clavatulidae) 
Mitrolumnidae Sacco, 1904 (synonym of Mitromorphidae) 
Mitromorphidae Casey, 1904 
Oenopotinae Bogdanov, 1987 (synonym of Mangeliidae) 
Pervicaciinae Rudman, 1969 (synonym of Terebridae) 
Pleurotominae Gray, 1838 (synonym of Turridae) 
Pleurotomellinae Nordsieck, 1968 (synonym of Raphitomidae) 
Pseudomelatomidae Morrison, 1965 
Pseudotominae Bellardi, 1875 (synonym of Borsoniidae) 
Puncticuliinae Tucker & Tenorio, 2009 (synonym of Conidae) 
Pusionellinae Gray, 1853 (synonym of Clavatulidae) 
Raphitomidae Bellardi, 1875 
Siphopsinae Le Renard, 1995 (fossil) 
Speightiidae Powell, 1942 (fossil; doubtfully Conoidea) 
Strictispiridae McLean, 1971 
Taraninae Casey, 1904 (synonym of Raphitomidae) 
Taranteconidae Tucker & Tenorio, 2009 (synonym of Conidae) 
Terebridae Mörch, 1852 
Thatcheriidae Powell, 1942 (synonym of Raphitomidae) 
Turriculinae Powell, 1942 (synonym of Clavatulidae) 
Turridae H. Adams & A. Adams, 1853 
Zemaciinae Sysoev, 2003 (synonym of Borsoniidae) 
Zonulispirinae McLean, 1971 (synonym of Pseudomelatomidae)