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Genus Profunditerebra new genus (Fig. 9 A–E) Type species: Profunditerebra papuaprofundi n. sp. Zoobank registration: urn:lsid:zoobank.org:act: A5A78A3D-AB11- 41C3-86F0-7C1451512FF4 Definition: Includes all species included in clade E3 of Modica et al. (2019) and those that show a combination of conchological, anatomical and distribution characteristics closely comparable with Profunditerebra papuaprofundi n. sp. or any genetically proven member of the clade. Diagnosis: Diagnostic nucleotide combinations provided in Table 8. Shell: Small to medium-sized (< 45 mm), with multispiral protoconch and slender siphonal canal. Whorls subcylindrical or flattened. Sculpture varying greatly; axial sculpture of rounded ribs, weak to very strong and elevated; spiral sculpture typically limited to striae in interstices and a subsutural groove, but may be absent or represented by strong cords generating overall cancellate pattern. Anatomy: Proboscis, radula, venom gland and salivary glands present. Radula of duplex marginal teeth (Fig. 9E). Accessory proboscis structure absent. Distribution: Almost exclusively deep water (typically 150–600 m) of tropical Indo-West Pacific from Taiwan to New Caledonia; one species known from shallow water of South Australia. Remarks: Profunditerebra is the only terebrine genus with duplex radular teeth similar to those in Pellifroniinae; however, members of the two groups can easily be differentiated by shell characters. Species of Profunditerebra can be separated into four clusters based on their shell morphology; however, only the first cluster corresponds to a phylogenetic lineage. Profunditerebra papuaprofundi n. sp. group. Shell uniformly coloured white to dark brown; heavily sculptured, with distinct subsutural groove and subcylindrical whorls. Sculpture of strong rounded ribs forming row of prominent nodules on subsutural band. Spiral sculpture of fine striae limited to interstices between axials, or overriding them to form fine continuous cords (including on subsutural band), or coarse cancellate pattern. Species of Neoterebra (e.g. N. armillata) exhibit a similar sculpture, but can be differentiated from Profunditerebra by their flattened whorls. The Punctoterebra textilis group also includes some similar forms (see Remarks on Punctoterebra). Profunditerebra orientalis group. Shell with flattened whorl profile; variegated colour pattern. Sculpture of fine orthocline ribs intersected by cords of about equal strength or slightly weaker than axials, to form delicate cancellate pattern. Subsutural band demarcated by shallow punctuate groove. Species of this group can easily be differentiated from congeners that either lack spiral elements, or display an overall much coarser sculpture. Both P. orientalis and P. hiscocki closely resemble species of Maculauger, from which they can confidently be differentiated only by anatomy (all studied species of Maculauger lack a radula) or molecular characters. Profunditerebra anseeuwi group. Shell with flattened whorl outline; subsutural band represented by short ribs or indentation. Sculpture of axial ribs, pronounced throughout whorl height, but weakening and becoming obsolete on later whorls; spiral sculpture absent. Species of this group are conchologically close to Duplicaria and to the Punctoterebra teramachii group; however, they can be differentiated from both by the overall weaker sculpture. In addition, these lineages can confidently be differentiated by the radular teeth morphology, Duplicaria having solid and recurved teeth, Profunditerebra duplex teeth and Punctoterebra flat teeth. Profunditerebra brazieri stands apart from other species in the genus, being endemic to temperate shallow water off Tasmania. Morphologically, it is close to the P. anseeuwi group, but has a very weakly defined subsutural band. Etymology: Name refers to the considerable depths at which almost all known species of the genus occur. Gender feminine. Included species: Profunditerebra anseeuwi (Terryn, 2005) 1 n. comb.; P. brazieri (Angas, 1871) 1 n. comb.; P. evelynae (Clench & Aguayo, 1939) 3 n. comb.; P. macclesfieldensis n. sp. Malcolm, Terryn & Fedosov 1 n. sp.; P. orientalis (Aubry, 1999) 1 n. comb.; P. papuaprofundi n. sp. Malcolm, Terryn & Fedosov 1 n. sp.; P. poppei (Terryn, 2003) 1 n. comb.; P. hiscocki (Sprague, 2004) 1 n. comb.

Genus Perirhoe Dall, 1908 (Fig. 10E) Type species: Perirhoe circumcincta (Deshayes, 1857); OD. Diagnosis: Shell small to medium-sized (< 50 mm), solid; axial sculpture weak to obsolete; spiral sculpture of irregular punctate groves; subsutural band weakly defined. Distribution: S Pacific, particularly New Caledonia. Remarks: Perirhoe is close to the genus Oxymeris in terms of shell proportions; however, it can be readily differentiated from species of the latter by the distinct spiral grooves throughout the whorls’ height. The sculpture pattern of widely set spiral grooves in Perirhoe resembles that in some species of Terebra, but the latter group has notably more slender shells, with higher spire and less robust siphonal canal. Included species: Perirhoe circumcincta (Deshayes, 1857) 4; Perirhoe valentinae (Aubry, 1999) 4 n. comb.

Genus Perirhoe Dall, 1908 (Fig. 10E) Type species: Perirhoe circumcincta (Deshayes, 1857); OD. Diagnosis: Shell small to medium-sized (< 50 mm), solid; axial sculpture weak to obsolete; spiral sculpture of irregular punctate groves; subsutural band weakly defined. Distribution: S Pacific, particularly New Caledonia. Remarks: Perirhoe is close to the genus Oxymeris in terms of shell proportions; however, it can be readily differentiated from species of the latter by the distinct spiral grooves throughout the whorls’ height. The sculpture pattern of widely set spiral grooves in Perirhoe resembles that in some species of Terebra, but the latter group has notably more slender shells, with higher spire and less robust siphonal canal. Included species: Perirhoe circumcincta (Deshayes, 1857) 4; Perirhoe valentinae (Aubry, 1999) 4 n. comb., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 22, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["DALL, W. H. 1908. Subdivisions of the Terebridae. Nautilus, 21: 124 - 125."]}

Genus Hastula H. Adams & A. Adams, 1858 (Fig. 6 A–E) Type species: Buccinum strigilatum Linnaeus, 1758; SD, Cossmann (1896). Synonyms: Impages E.A. Smith, 1873 (type species Terebra caerulescens Lamarck, 1822 = Buccinum hecticum Linnaeus, 1758; SD, Cossmann, 1896); Acuminia Dall, 1908 (type species Buccinum lanceatum Linnaeus, 1767; OD); Hastulina Oyama, 1961 (type species Terebra casta Hinds, 1844; OD); Egentelaria Rehder, 1980 (type species Terebra stylata Hinds, 1844; OD). Definition: Includes all species in clade D of Modica et al. (2019), and those that show a combination of conchological, anatomical and distribution characteristics closely comparable with Hastula strigilata or any genetically proven member of the clade. Diagnosis: Diagnostic nucleotide combinations provided for entire clade with exception of four specimens of H. albula (see Material and Methods) and, separately, for subclade H. albula – H. natalensis – H. aff. casta of clade D (Table 4). Shell: Small to medium-sized, shiny, 12–90 mm. Protoconch paucispiral or multispiral, about 1–5 whorls. Axial sculpture of crenulations on subsutural portion of whorl, or fine ribs on part or throughout whorl height; spiral sculpture of single subsutural groove, occasionally with additional row of punctations. Siphonal canal stout, very short, with well-developed fasciole. Aperture elongate to rounded. Genus Hastula (except four specimens of H. albula) Species/sequences analysed 18/91 Diagnostic nucleotides: 28: A, 67: T, 95: C, 181: T, 221: C, 222: C, 253: A, 581: T, 598: A Subclade H. albula – H. natalensis – H. aff. casta Anatomy: Rhynchodeal introvert, proboscis, venom gland, salivary glands and radula well developed. Radula of hypodermic marginal teeth attached to thin, reduced membrane; walls of marginal teeth often with multiple perforations (Fig. 6E, F) Species/sequences analysed 3/17 Diagnostic nucleotides 49: G, 347: C, 349: T, 517: T Distribution: Indo-Pacific, West Africa, Caribbean. Remarks: Species of Hastula are rather easy to recognize among Terebridae by their typically shiny shells, with fine close-set axial ribs and usually by the lack of a clearly defined subsutural band. The shell morphology in the Partecosta macleani group closely resembles that of Hastula; however, the latter is usually larger at maturity, while shells of comparable size can be differentiated by texture—a polished surface in Hastula vs dull in Partecosta. Furthermore, species of Hastula possess a venom gland and hypodermic radular teeth, whereas Partecosta species lack a venom gland and their radulae teeth are solid. Included species: Hastula aciculina (Lamarck, 1822) 2; H. acumen (Deshayes, 1859) 1; H. alboflava Bratcher, 19882; H. albula (Menke, 1843) 1; H. androyensis Bozzetti, 20082; H. anomala (Gray, 1834) 2; H. anosyana (Bozzetti, 2016) 2; H. apicitincta (G. B. Sowerby III, 1900) 2; H. bacillus (Deshayes, 1859) 2; H. casta (Hinds, 1844) 1; H. celidonota (Melvill & Sykes, 1898) 2; H. cernohorskyi Burch, 19652; H. cinerea (Born, 1778) 1; H. continua Deshayes, 18592; H. crossii (Deshayes, 1859) 1 ∗; H. cuspidata (Hinds, 1844) 2; H. denizi Rolán & Gubbioli, 20002; H. engi Malcolm & Terryn, 20172; H. escondida (Terryn, 2006) 2; H. exacuminata Sacco, 18912; H. filmerae (G. B. Sowerby III, 1906) 2; H. hamamotoi Tsuchida & Tanaka, 19992; H. hastata (Gmelin, 1791) 1; H. hectica (Linnaeus, 1758) 1; H. imitatrix (Auffenberg & Lee, 1988) 2; H. inconstans (Hinds, 1844) 2; H. knockeri (E. A. Smith, 1872) 2; H. lanceata (Linnaeus, 1767) 1; H. leloeuffi Bouchet, 19832; H. lepida (Hinds, 1844) 2; H. marqueti (Aubry, 1994) 2; H. maryleeae R. D. Burch, 19652; H. matheroniana (Deshayes, 1859) 1; H. nana (Deshayes, 1859) 2; H. natalensis (E. A. Smith, 1903) 1 ∗; H. parva (Baird, 1873) 1; H. penicillata (Hinds, 1844) 1; H. philippiana (Deshayes, 1859) 2; H. puella (Thiele, 1925) 1; H. raphanula (Lamarck, 1822) 1; H. rufopunctata (E. A. Smith, 1877) 2; H. salleana (Deshayes, 1859) 1; H. sandrogorii Ryall, Terryn & Rosado, 20172; H. solida (Deshayes, 1857) 1; H. strigilata (Linnaeus, 1758) 1; H. stylata (Hinds, 1844) 1; H. tenuicolorata Bozzetti, 20081; H. tiedemani Burch, 19652; H. venus Aubry, 20082; H. verreauxi (Deshayes, 1857) 1; H. westralica (Aubry, 1999) 2; H. daniae (Aubry, 2008) 3.

Neoterebra guadeloupensis Malcolm, Terryn & Fedosov newspecies (Fig. 13A, B) Type material: Holotype: MNHN IM-2013-61448, 1 lv, 17.2 mm; S Marie-Galante, Guadeloupe, 15 ° 50 'N, 61 ° 19 'W, 305–312 m (N/O Antea, KARUBENTHOS 2015 Stn DW4638). Paratypes 1–3: off Fajou, Guadeloupe, 16 ° 22 'N, 61 ° 34 ', 80 m (KARUBENTHOS 2012 Stn GD02). Paratype 1: MNHN IM-2013-20542, 1 lv, 6.5 mm. Paratype 2: MNHN IM-2013-20541, 1 lv, 5.7 mm. Paratype 3: MNHN IM-2013-20539, 1 lv, 10.4 mm. Paratype 4: MNHN IM-2013-9110, 1 lv, 7 × 2.1 mm; Port Louis, Guadeloupe, 16 ° 22 ' 46 'N, 61 ° 34 'W, 66 m (KARUBENTHOS 2012 Stn GD35). Zoobank registration: urn:lsid:zoobank.org:act: 57E995EE-24F0- 47A0-9DDC-E9D54948B056 Diagnostic nucleotide positions: Table 11. Description: Shell small. Protoconch multispiral, transparent with 3 whorls tinged yellow; protoconch/teleoconch transition clear-cut with visible sinusigera. Teleoconch of holotype with 15 whorls; overall whorl outline straight to convex. Weak axial sculpture of 15 narrow axial ribs with wider interspaces, increasing with maturity to 25 on penultimate whorl. Deep subsutural groove crossed obliquely by ribs. Subsutural band with strong elongate nodes; subsequently, 2 strongly projecting spiral cords at top and middle of whorl dominating sculpture, forming reticulate pattern of deep indentations with thin ribs; the upper cord projects in outline beyond subsutural band. On mature whorls interspaces sometimes with 1 or 2 weak spiral striae. Reticulate sculpture continuing on shell base. Aperture long with weak callus; columella with 2 weak folds. Shell lacks any pattern; coloured in shades of white to yellow. Habitat: Deep water, 60– 310 m. Distribution: Only known from off Guadeloupe. Etymology: Named after the type locality, the French Caribbean island of Guadeloupe. Remarks: Anumber of species of Terebridae have been described from the Western Atlantic and Caribbean Sea that have a similar strong reticulate sculpture. These include N. limatula that has more than 2 strong spiral cords in addition to the subsutural band (Fig. 13C) and a paucispiral protoconch. Neoterebra colombiensis is very similar to N. guadeloupensis n. sp., but the spiral sculpture in the former is less dominant, giving a more uniform reticulate sculpture with nodes at the intersections. In addition, N. colombiensis has a paucispiral protoconch in comparison with the multispiral protoconch of N. guadeloupensis n. sp. Neoterebra simonei is similar in colour and sculpture, but has 3 spiral cords in addition to the subsutural band and has a distinctive paucispiral protoconch of 1.5 whorls. Other similar species from the region, e.g. N. intumescyra and N. alagoensis (Fig. 9F), all have paucispiral protoconchs and different sculpture.

Maculauger sudchinensis Malcolm, Terryn & Fedosov newspecies (Fig. 13D, E) Type material: Holotype: MNHN IM-2013-61887, 1 lv, 19.9 mm; N Macclesfield Bank, S China Sea, 16 ° 07 'N, 114 ° 23 'E, 161 m (ZhongSha 2015 Stn DW4144). Paratypes: N Macclesfield Bank, S China Sea, 16 ° 08 'N, 114 ° 19 'E, 180–226 m (ZhongSha 2015 Stn CP4145). Paratype 1: MNHN IM-2013-61902, 1 lv, 20.2 mm. Paratype 2: MNHN IM-2013-61895, 1 lv, 20.7 mm. Other examined material: N Macclesfield Bank, S China Sea, 16 ° 07 'N, 114 ° 19 'E, 218–281 m (ZhongSha 2015 Stn CP4148), 2 dd. Zoobank registration: urn:lsid:zoobank.org:act: BB5F5C93-C9FE- 44F0-B1EC-C00AA31EFF23 Diagnostic nucleotide positions: Table 11. Description: Shell to 21 mm; narrow apical angle. Protoconch paucispiral, 1.5 translucent whorls. Teleoconch of holotype with 14 whorls; outline of whorls slightly convex. Subsutural band formed of deep pits. Axial ribs narrow with wide interspaces; ribs c. 18–20 on penultimate whorl, straight on subsutural band, becoming oblique at subsutural groove, then curved on abapical part of whorls. Axial ribs intersected by 2–3 evenly spaced spiral grooves. Aperture long, with straight columella. Ground colour chalkywhite, sometimes with vague brown marks on subsutural band. Distribution: Only known from N Macclesfield Bank. Habitat: Depths of 160– 220 m. Etymology: Named after the South China Sea. Remarks: Maculauger campbelli has a similar protoconch and slender overall shape with slightly convex whorls, but in comparison with M. sudchinensis n. sp. has a more continuous subsutural groove, less pronounced sculpture, with 5–7 spiral grooves and a distinctive pattern of straw colour with random square spots. Maculauger sudchinensis n. sp. is similar to Terebra helichrysum (Fig. 13F), which however has a much shorter curved columella, a multispiral protoconch of 3.5 whorls, shorter whorls and a mottled orange colour pattern. Terebra levantina (Fig. 13G) has similar sculpture, but its whorls are much shorter, with 4–6 spiral grooves and it has a mottled colour pattern.

Profunditerebra papuaprofundi Malcolm, Terryn & Fedosov newspecies (Fig. 12A, B) Type material: Holotype: MNHN IM-2013-58123, 1 lv, 19.0 mm; off New Ireland, Papua New Guinea, 2 ° 21 'S, 150 ° 38 'E, 496–609 m (KAVIENG 2014 Stn CP4422). Paratype 1: MNHN IM-2013-45571, 1 lv, 29.5 mm; W Buka I., N Bougainville, Papua New Guinea, 5 ° 43 'S, 154 ° 03 'E, 490–530 m (MADEEP Stn DW4278), Paratype 2: MNHN IM-2013-59946, 1 lv, 25.6 mm; same locality. Paratype 3: MNHN IM-2013-59944, 1 lv, 13.7 mm; same locality. Paratype 4: MNHN IM-2013-59945, 1 lv, 16.3 mm; W Buka I., N Bougainville, 5 ° 35 'S, 153 ° 58 'E, 623–640 m (MADEEP Stn DW4271),. Paratype 5: MNHN IM-2013-45509, 1 lv, 27.1 mm; W Buka I., N Bougainville, 5 ° 37 'S, 153 ° 59 'E, 421 m (MADEEP Stn DW4270). Paratype 6: MNHN IM-2013-30574, 1 lv, 33 mm (broken); SW Santa Isabel I, Solomon Islands, 8 ° 24 'S, 159 ° 27 'E, 362–432 m (SALOMON 2 Stn CP2193). Other material examined: Taiwan, 15 ° 03 'N; 116 ° 31 'E (NANHAI 2014 Stn DW4102), IM-2013-52276, 1 lv. Zoobank registration: urn:lsid:zoobank.org:act: ABAFB4A7-FCD6- 4632-A07C-E9945D21EA7A Diagnostic nucleotide positions: see Table 11. Description (holotype): Shell of moderate size, to 42 mm. Protoconch of 4 narrow, conical whorls with small nucleus. Teleoconch of 12 whorls; teleoconch whorl outline convex constricted abapically at the suture. Subsutural groove represented by weak indentation on early whorls becoming sharper groove cutting through the ribs on mature whorls. Sculpture of strong slightly convex narrow axial ribs on subsutural band aligned with those on succeeding portion of whorls, varying significantly between specimens in spacing, and in number from 18 to 25 (20 ribs on penultimate whorl). No spiral sculpture in interspaces except for striae on base below periphery. Last whorl cup-shaped, becoming more elongate on mature specimens. Aperture wide with slight callus; columella straight with 2 folds. Colour off-white with tinges of yellow or pink with shiny white columella. Habitat: Depths of 350– 600 m. Distribution: Papua New Guinea, Solomon Islands and Taiwan. Etymology: Named after its deep-water habitat in Papua New Guinea. Remarks: The only sequenced specimen of P. papuaprofundi n. sp. with intact protoconch was designated as the holotype. It is notably smaller than the sequenced paratypes, which reach shell lengths of 42 mm. The larger specimens demonstrate the increased curvature of ribs and increased number and density of ribs to 25 on the penultimate whorl. Profunditerebra papuaprofundi n. sp. is phylogenetically closest to P. orientalis and P. anseeuwi. Profunditerebra anseeuwi has a weaker subsutural groove, straight and oblique ribs and a mottled colour pattern. Profunditerebra orientalis has more axial ribs, a mottled colour pattern, and a sculpture of ribs and spiral cords creating nodes at their intersections. Specimens of P. papuaprofundi n. sp. with more numerous curving ribs are similar to Maculauger cinctella and Terebra textilis (Salvador & Pickering, 2017). However, M. cinctella (Fig. 12C) has a shorter protoconch of 2.5 whorls, a much narrower noded subsutural band and many spiral striae within the interspaces between the ribs. In turn, T. textilis (Fig. 12D) differs in having a spiral groove defined by deep punctations or partially cutting the ribs, and has several spiral striae in the interspaces between the ribs., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 25, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["SALVADOR, A. & PICKERING, J. 2017. Type catalogue of Terebridae (Mollusca, Gastropoda, Conoidea) in the Natural History Museum, London, U. K. Zootaxa, 4250: 101 - 142."]}

Genus Hastulopsis Oyama, 1961 (Fig. 10B) Type species: Terebra melanacme E. A. Smith, 1873; OD. Diagnosis: Shell small to medium-sized (to 60 mm), shiny, faintly axially ribbed, with numerous fine spiral incisions. Subsutural band generally not spirally decorated, bordered by relatively deep incision (Terryn, 2007). Distribution: Indian and western Pacific Oceans. Remarks: The genus comprises two groups of shallow-water species, one found from the eastern coast of Africa to the Arabian Gulf and another in waters of mainland Japan to Australia. Similar, but deeper-water species are found in the genus Maculauger n. gen., most of which can be recognized by having a row of large spots on the periphery. The species of Myurella have much stronger and more distinct sculpture features, and generally a brown basal area. Included species: Hastulopsis baliensis Terryn & Dekker, 20174; H. blanda (Deshayes, 1859) 4; H. cebuensis Gargiulo, 20144; H. elialae (Aubry, 1994) 4; H. gotoensis (E. A. Smith, 1879) 4; H. loisae (E. A. Smith, 1903) 4; H. maestratii Terryn & Rosado, 20114; H. marmorata (Deshayes, 1859) 4; H. masirahensis Terryn & Rosado, 20164; H. melanacme (E. A. Smith, 1873) 4; H. mirbatensis Terryn & Rosado, 20164; H. suspensa (E. A. Smith, 1904) 4; H. whiteheadae (Aubry & Marquet, 1995) 4., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 21, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["OYAMA, K. 1961. On some new facts of the taxonomy of Terebridae. Venus, 21: 176 - 189.","TERRYN, Y. 2007. A collectors' guide to Recent Terebridae (Mollusca: Neogastropoda). Conchbooks, Hackenheim."]}

Genus Gradaterebra Cotton & Godfrey, 1932 (Fig. 10D) Type species: Gradaterebra scalariformis Cotton & Godfrey, 1932; OD. Diagnosis: Shell small (< 25 mm), often dull-coloured, with flattened tobulbouswhorls; axialsculptureof prominentribs; spiralsculpture absent, microscopic or represented by fine cords; last whorl inflated. Distribution: Indian and SW Pacific Oceans. Remarks. The genus mainly comprises small species from Australia and South Africa, which can be differentiated based on adult size < 25 mm. Included species: Gradaterebra scalariformis (Cotton & Godfrey, 1932) 4 n. comb.; G. assecla (Iredale, 1924) 4 n. comb.; G. capensis (E. A. Smith, 1873) 4 n. comb.; G. easmithi (Aubry, 1999) 4 n. comb.; G. kowiensis (Turton, 1932) 4 n. comb.; G. lightfooti (E. A. Smith, 1899) 4 n. comb.; G. ninfae (Campbell, 1961) 4 n. comb.; G. pilsbryi (Aubry, 1999) 4 n. comb.; G. planecosta (Barnard, 1958) 4 n. comb.; G. severa (Melvill, 1897) 4 n. comb.; G. sorrentense (Aubry, 1999) 4 n. comb.; G. taylori (Reeve, 1860) 4 n. comb.

Genus Pellifronia Terryn & Holford, 2008 (Fig. 2 A–D) Type species: Terebra jungi Lai, 2001; OD. Definition: Includes species with combination of conchological, anatomical and distributional characteristics closely comparable with Pellifronia jungi, including all species in clade A1 of Modica etal. (2019) and any species proven to be a member of this clade. Diagnosis: Diagnostic nucleotide combinations in Table 1. Shell: Medium-sized (25–50 mm), elongate, light brown. Protoconch brown, paucispiral, cyrtoconoid, 1.5–2.5 whorls. Coarse sculpture of pronounced ribs, orthocline or prosocline, sometimes arcuate on late teleoconch whorls. Axial ribs often thickened in their adapical and/or abapical part(s), giving slightly concave outline to spire whorls. Last adult whorl with flattened adapical portion and short, somewhat angulate transition to shell base. Siphonal canal moderately long, recurved at tip, shallowly notched; siphonal fasciole weak or indistinct. Aperture elongate, curved. Anatomy: Rhynchodeal introvert small; proboscis, radular sac, odontophore, salivary and venom glands well developed. Acessory proboscis structure absent. Radula with well-developed membrane, bearing 2 rows of wide marginal teeth of duplex type. Distribution: Indo-Pacific, bathyal depths. Remarks: This genus differs from the closely related Bathyterebra in having stronger axial sculpture, a more flattened whorl outline and the last adult whorl constricted to the siphonal canal with visible angulation. Species of Pellifronia can be distinguished by their coarse sculpture and swollen rib ends with slightly raised subsutural area. Similar species in other genera, e.g. Neoterebra puncturosa, N. crenifera and Granuliterebra bathyrhaphe, normally have a sharper nodular structure on the subsutural band or rib endings, or can be separated by the lack of radula (Neoterebra). Included species: Pellifronia brianhayesi (Terryn & Sprague, 2008) 2; P. jungi (Lai, 2001) 1.

Genus Maculauger new genus (Fig. 7E, F) Type species: Terebra pseudopertusa Bratcher & Cernohorsky, 1985. Zoobank registration: urn:lsid:zoobank.org:act: 3565DB17-E4EF- 478B-8757-3A682BB8F6CF Definition: Includes all species included in clade E5A of Modica et al. (2019) and those that show a combination of conchological, anatomical and distribution characteristics closely comparable with Maculauger pseudopertusus or any genetically proven member of the clade. Diagnosis: Diagnostic nucleotide combinations in Table 10. Shell: Small to medium-sized (< 60 mm); typically, orange-brown with irregular maculations on subsutural band. Whorl outline flattened or gently convex. Subsutural band lightly ribbed or flattened, separated from lower portion of whorl by narrow groove or row of punctations. Axial and spiral sculpture of about equal strength, producing cancellate sculpture below subsutural band. Anatomy: Proboscis, radula, venom gland and accessory proboscis structure absent; salivary glands present or absent. Distribution: Indo-Pacific; subtidal and bathyal depths to about 600 m. Remarks: The new genus Maculauger comprises several species, formerly placed in Hastulopsis, that can be differentiated from similar species in Myurella by the equal prominence of spiral and axial spiral elements, producing a fine cancellate sculpture. Hastulopsis species have very weak sculpture and are found in the Indian Ocean or Japan. Some species of Maculauger, such as M. campbelli, are conchologically very close to Profunditerebra orientalis —a case where only anatomical and molecular characters can ensure correct identification (see Remarks on Profunditerebra). Etymology: The name combines ‘macula’ (Latin: spot, blotch), referring to the commonly maculated subsutural band in the species of the genus, with ‘auger’, the English epithet for Terebridae (‘auger shells’). Gender masculine. Included species: Maculauger alveolatus (Hinds, 1844) 1 n. comb.; M. campbelli (R. D. Burch, 1965) 1 n. comb.; M. castigatus (A. H. Cooke, 1885) 1 n. comb.; M. cinctella (Deshayes, 1859) 3 n. comb.; M. minipulcher (Bozzetti, 2008) 1 n. comb.; M. pseudopertusus (Bratcher & Cernohorsky, 1985) 1 n. comb.; M. sudchiniensis n. sp. Malcolm, Terryn & Fedosov 1 n. sp.

Genus Neoterebra new genus (Fig. 9 F–H) Type species: Terebra specillata Hinds, 1844. Zoobank registration: urn:lsid:zoobank.org:act: 68993F36-2AD1- 4DDE-A853-8832B4A9EE30 Definition: Includes all species included in clade E4 of Modica et al. (2019) and those that show a combination of conchological, anatomical and distribution characteristics closely comparable with Neoterebra specillata or any genetically proven member of the clade. Diagnosis: Diagnostic nucleotide combinations in Table 9. Shell: Small to medium-sized (< 60 mm); uniformly creamy-white to light brown or with pattern of brown blotches; pointed multispiral or paucispiral protoconch, 1.5 to about 3 whorls. Axial sculpture of orthocline or arcuate ribs, forming axially elongated subsutural nodules on well-defined band. Spiral sculpture from weak striae in interspaces between ribs to distinct cords, overriding ribs, often forming nodes at intersection or on lower portion of ribs. Last whorl typically abruptly constricted to siphonal canal. Canal clearly separated from shell base, bearing pronounced fasciole. Anatomy: Radula, proboscis, venom gland typically absent. Radula, when present, of hypodermic type. Salivary glands typically present. Accessory proboscis structure absent. Distribution: Tropical East Pacific, Caribbean, off Brazil; intertidal to upper bathyal depths, but typically shallow water. Remarks: Shallow-water species of Neoterebra differ from their congeners from deep water of Brazil and the Caribbean, being larger at maturity and sharing characteristic sculpture of strong axial ribs that form elevated, sometimes axially elongated nodules on the subsutural band. Caribbean species have a weaker subsutural band; those from deep water commonly lack a recognizable subsutural band and display a fine cancellate pattern of narrow axial riblets, intersected by spiral cords of equal strength. Although the affinity of Panamic and Caribbean species is suggested by the analysis of COI, further studies are needed to document relationships and delineate major groupings within the genus. Etymology: The name is a combination of neo (Latin: new) and Terebra, referring to the fact that species of this new genus only occur in the New World, constituting the majority of the terebrid fauna in the region. Included species: Neoterebra acrior (Dall, 1889) 2 n. comb.; N. alagoensis (Lima, Tenorio & Barros, 2007) 2 n. comb.; N. alba (Gray, 1834) 2 n. comb.; N. allyni (Bratcher & Burch, 1970) 2 n. comb.; N. angelli (J. Gibson-Smith & W. Gibson-Smith, 1984) 2 n. comb.; N. arcas (Abbott, 1954) 2 n. comb.; N. armillata (Hinds, 1844) 1 n. comb.; N. assu (Simone, 2012) 2 n. comb.; N. berryi (Campbell, 1961) 2 n. comb.; N. biminiensis (Petuch, 1987) 2 n. comb.; N. brandi (Bratcher & Burch, 1970) 2 n. comb.; N. brasiliensis (E. A. Smith, 1873) 2 n. comb.; N. bridgesi (Dall, 1908) 3 n. comb.; N. carolae (Bratcher, 1979) 2 n. comb.; N. churea (Campbell, 1964) 2 n. comb.; N. colombiensis (Simone & Gracia, 2006) 2 n. comb.; N. concava (Say, 1826) 2 n. comb.; N. corintoensis (Pilsbry & Lowe, 1932) 2 n. comb.; N. crassireticula (Simone, 1999) 2 n. comb.; N. crenifera (Deshayes, 1859) 1 n. comb.; N. curacaoensis (De Jong & Coomans, 1988) 2 n. comb.; N. dislocata (Say, 1822) 1 n. comb.; N. doellojuradoi (Carcelles, 1953) 2 n. comb.; N. elata (Hinds, 1844) 1 n. comb.; N. frigata (Hinds, 1844) 2 n. comb.; N. glauca (Hinds, 1844) 2 n. comb.; N. glossema (Schwengel, 1942) 1 n. comb.; N. guadeloupensis n. sp. Malcolm, Terryn & Fedosov 1 n. sp.; N. guayaquilensis (E. A. Smith, 1880) 2 n. comb.; N. hancocki (Bratcher & Burch, 1970) 2 n. comb.; N. hemphilli (Vanatta, 1924) 2 n. comb.; N. hondurasiensis (Gargiulo, 2016) 2 n. comb.; N. intertincta (Hinds, 1844) 3 n. comb.; N. intumescyra (Lima, Tenorio & Barros, 2007) 2 n. comb.; N. jacquelinae (Bratcher & Burch, 1970) 2 n. comb.; N. juanica (Dall & Simpson, 1901) 2 n. comb.; N. lamyi (Terryn, 2011) 2 n. comb.; N. larvaeformis (Hinds, 1844) 1 n. comb.; N. leptapsis (Simone, 1999) 2 n. comb.; N. limatula (Dall, 1889) 1 n. comb.; N. lucana (Dall, 1908) 2 n. comb.; N. mugridgeae (García, 1999) 2 n. comb.; N. nassula (Dall, 1889) 2 n. comb.; N. pacei (Petuch, 1987) 2 n. comb.; N. panamensis (Dall, 1908) 2 n. comb.; N. pedroana (Dall, 1908) 2 n. comb.; N. petiveriana (Deshayes, 1857) 2 n. comb.; N. plicata (Gray, 1834) 2 n. comb.; N. protexta (Conrad, 1846) 2 n. comb.; N. puncturosa (Berry, 1959) 1 n. comb.; N. rancheria (Bratcher, 1988) 2 n. comb.; N. riosi (Bratcher & Cernohorsky, 1985) 2 n. comb.; N. roperi (Pilsbry & Lowe, 1932) 3 n. comb.; N. rushii (Dall, 1889) 2 n. comb.; N. sanjuanense (Pilsbry & Lowe, 1932) 2 n. comb.; N. shyana (Bratcher & Burch, 1970) 2 n. comb.; N. simonei (Lima, Tenorio & Barros, 2007) 2 n. comb.; N. specillata (Hinds, 1844) 1 n. comb.; N. spirosulcata (Simone, 1999) 2 n. comb.; N. sterigma (Simone, 1999) 2 n. comb.; N. sterigmoides (Simone & Gracia, 2006) 1 n. comb.; N. stohleri (Bratcher & Burch, 1970) 2 n. comb.; N. tiarella (Deshayes, 1857) 3 n. comb.; N. variegata (Gray, 1834) 1 n. comb.; N. vinosa (Dall, 1889) 2 n. comb., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 18-20, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059.","DALL, W. H. 1908. Subdivisions of the Terebridae. Nautilus, 21: 124 - 125."]}

Genus Microtrypetes Pilsbry & Lowe, 1932 (Fig. 10F) Type species: Microtrypetes iola Pilsbry & Lowe, 1932; OD. Diagnosis: Shell small (< 20 mm), dull, with turreted outline due to subcylindrical to clearly cylindrical whorl outline. Sculpture of strong, straight to finely curving axial ribs and fine spiral striae. Subsutural band indistinct or weakly defined. Siphonal canal long, curved. Distribution: E Pacific. Remarks: The genus can be differentiated from other New World genera of Terebridae by the stepped appearance of the spire, due to the cylindrical or subcylindrical whorl outline, by the generally indistinct subsutural band and by the siphonal canal that is rather long for the family. Included species: Microtrypetes iola Pilsbry & Lowe, 19324; M. polypenus (Pilsbry & Lowe, 1932) 4 n. comb. Descriptionsof newspecies

Genus Microtrypetes Pilsbry & Lowe, 1932 (Fig. 10F) Type species: Microtrypetes iola Pilsbry & Lowe, 1932; OD. Diagnosis: Shell small (< 20 mm), dull, with turreted outline due to subcylindrical to clearly cylindrical whorl outline. Sculpture of strong, straight to finely curving axial ribs and fine spiral striae. Subsutural band indistinct or weakly defined. Siphonal canal long, curved. Distribution: E Pacific. Remarks: The genus can be differentiated from other New World genera of Terebridae by the stepped appearance of the spire, due to the cylindrical or subcylindrical whorl outline, by the generally indistinct subsutural band and by the siphonal canal that is rather long for the family. Included species: Microtrypetes iola Pilsbry & Lowe, 19324; M. polypenus (Pilsbry & Lowe, 1932) 4 n. comb. Descriptionsof newspecies, Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 22-23, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844

Partecosta bozzettii Malcolm & Terryn new species (Figs 3K, 11E) Type material: Holotype: MNHN IM- 2009-10163, lv, 12.5 mm; Port d’Ehoala, SMadagascar, 25 ° 03 ' 43.9 ''S, 46 ° 57 ' 42.9 ''E, 3–4 m (ATIMO VATAE Stn TP29). Paratype 1: MNHN IM-2009- 10162, lv, 11.8 mm; type locality. Zoobank registration: urn:lsid:zoobank.org:act: 6E3E67B5-1B55- 4A99-8D39-6517DA4D065C Diagnostic nucleotide positions: Table 11. Description: Shell small. Protoconch paucispiral, 1 whorl, with broad nucleus. Holotype 9 teleoconch whorls; outline of whorls weakly convex. No subsutural band, or demarcation thereof, except for minor indentation of ribs; no visiblespiral sculpture. Axial sculpture of almost straight, thin ribs, 14 on penultimate whorl, stretching across whorl height; interspaces wider than ribs. Last whorl elongate, tapering towards base. Columella straight, aperture elongate. Shell colour off-white with darker-tinged spiral band adapically on whorls and below periphery; brownish blotches between the axial ribs abapically on whorls; inner columella dark brown or purplish. Habitat: In sand at 3– 4 m. Distribution: Known only from type locality. Etymology: The species honours Luigi Bozzetti (Italy), who has contributed tremendously to the knowledge of the molluscan fauna of SMadagascar. Remarks: The species resembles P. trilineata (Fig. 11F), but differs by its more globose and convex whorls and by having a single row of brown blotches instead of two. In addition, the protoconch of P. trilineata is differently shaped, having two whorls and a medium nucleus. The phylogenetic analysis of Modica et al. (2019) shows significant genetic distance between the two species. Partecosta bozzettii n. sp. shows some resemblance to P. albofuscata, but again differs by its more convex whorls, distinct axial ribbing and shell colour pattern. Partecosta bozzettii n. sp. is similar in general morphology and shell colour pattern to Hastula daniae (Fig. 2G) from West Africa, but the latter has a multispiral protoconch and more globose mature whorls.

Genus Terebra Bruguière, 1789 (Figs 4, 5) Type species: Buccinum subulatum Linnaeus, 1767; SD (Lamarck, 1799: 71). Synonyms: Triplostephanus Dall, 1908 (type species Terebra triseriata Gray, 1834; OD); Cinguloterebra Oyama, 1961 (type species Terebra hedleyana Pilsbry, 1905 = Terebra adamsii E.A. Smith, 1873; OD); Dimidacus Iredale, 1929 (new name for Terebrina Bartsch, 1923, non Rafinesque, 1815; type species Terebra cingulifera Lamarck, 1822, by typification of replacement name); Myurellina Bartsch, 1923 (type species Terebra ornata Gray, 1834; OD); Subula Schumacher, 1817 (unnecessary replacement name for Terebra; type species Buccinum subulatum, by typification of replacement name); Terebrina Bartsch, 1923 (invalid: junior homonym of Terebrina Rafinesque, 1815; type species Terebra cingulifera Lamarck, 1822, OD); Terebrum Montfort, 1810 (unnecessary emendation of Terebra Bruguière, 1789; type species Buccinum subulatum Linnaeus, 1767, by typification of replaced name); Paraterebra Woodring, 1928 (type species Terebra texana Dall, 1898; OD); Panaterebra Olsson, 1967 (type species Terebra robusta Hinds, 1844; OD). Definition: Primarily defined based on phylogenetic analysis of Modica etal. (2019); includes all species in clade Cof Modica etal. (2019), and species showing resemblance in shell morphology, anatomy and distribution to Terebra subulata or any genetically proven member of this clade. Diagnosis: Diagnostic nucleotide combinations (provided separately for the seven clades of the genus inferred by Modica et al. (2019) in Table 3. Shell: Elongate, medium-sized to large, reaching 192 mm; truncated base abruptly constricted to siphonal canal, sometimes with distinct angulation. Sculpture from very weak, limited to often obsolete subsutural groove, to very strong. Subsutural band simple, or consisting of two closely set cords, often gemmate. Lower whorl portion smooth or with varying sculpture, with spiral elements being as strong or stronger than axials. Shell base separated from siphonal canal by concave waist. Aperture elongate to quadrangular. Anatomy: Rhynchodeal introvert, proboscis, venom gland, salivary glands and radula well developed. Radula of hypodermic marginal teeth, with only bases attached to strongly reduced membrane. Accessory proboscis structure present or absent. Distribution: Indo-Pacific, West Africa, tropical East Pacific, Caribbean and Western Atlantic. Remarks: The genus Terebra is here established with boundaries consistent with clade Cof Modica etal. (2019). This clade is subdivided into eight lineages, referred to as C1–C7 (Fig. 4), their relationships being unresolved. Clades C1, C2, C3-2, C4 and C7 are well supported; clade C5 comprises one species only— T. argus. Because of the extreme heterogeneity of clade C, our attempts to provide a molecular diagnosis failed and, therefore, we compiled separate diagnoses for the inferred lineages within clade C (Table 3). Although lineages C3-1 and C3-2 do not constitute a supported clade, this arrangement is not rejected by the phylogenetic analysis and we have grouped them together based on biogeographic considerations. The genus Terebra is also very heterogeneous in regard to shell morphology (Fig. 5); some common morphotypes can be distinguished, but their correspondence with inferred phylogenetic groupings is far from perfect. Below we provide a brief overview of these morphotypes. Typical Terebra are characterized by a weakly sculptured or smooth, medium-sized shell, with gently convex or subcylindrical whorls, often bearing a pattern of large spots. Species referable to this morphotype are included in clades C1 (T. subulata and T. guttata), C4 (T. ornata) and C6 (T. argus). This morphotype is closest in appearance to Oxymeris and differentiation can be difficult. In most cases Terebra species can be recognized by the narrower anterior notch; however, for definite allocation, examination of the foregut is advised to determine whether proboscis and radula are present. The fenestrata morphotype comprises the most heavily sculptured species, with a strong, often elevated, gemmate subsutural band that is closely followed by a second spiral band of similar morphology, but usually weaker and/or narrower. The lower portion of the whorl is sculptured with spiral cords, or has a cancellate pattern. Species exhibiting this morphotype include clade C2, and some strikingly similar species in clades C7 (T. aff. adamsii, T. fenestrata MNHN IM-2013-14908, T. anilis) and C8 (T. cumingi species complex). Strong, often elevated, gemmate or beaded spiral elements readily differentiate this morphotype from most terebrid genera. However, a similar pattern is present in Granuliterebra, the species of which are differentiated by their single subsutural band and usually by predominantly axial sculpture on the lower portion of the whorl. The Triplostephanus morphotype unifies species with a very high spire and very small, quadrate aperture. Whorls are flattened or slightly concave due to a raised subsutural band and the lower part of the whorl is sculptured by regular cords, while axial sculpture is not developed. Most species in clade C5 exhibit this morphotype; however, species with subcylindrical whorls and stepped spire outline (e.g. T. laevigata and T. funiculata) are found in C5 as well. The combination of a very high spire and flattened whorl outline makes this morphotype very distinctive among Terebridae. While the three described morphotypes represent extremes of the conchological diversity of Terebra, there is a plethora of intermediate forms. For example, T. taurina and T. fujitai of clade C3 have a divided ribbed subsutural band, similar to the fenestrata morphotype, although notably lower. Similarly, a divided, nodulose subsutural band is present in T. corrugata, a species close to T. subulata in clade C1. The apparent lack of phylogenetic congruence in the distribution of these morphotypes across clade C precludes assignment of any taxonomic status to them. Moreover, the general lack of homogeneity in shell characteristics within the subclades of Terebra makes their establishment as supraspecific taxonomic units pointless as well. Taking into consideration the great conchological variation within Terebra, it is difficult to suggest general diagnostic criteria for this genus. However, it should be noted that the predominantly spiral sculpture below the subsutural band allows differentiation of Terebra from most other terebrid genera. Similarly, a divided subsutural band has not been recorded in any species outside Terebra. These characters, in combination with the presence of a complete complex of foregut organs and radular teeth of hypodermic type, allow for reliable differentiation of Terebra species. Included species: Terebra achates Weaver, 19602; T. adamsii (E. A. Smith, 1873) 2; T. albocancellata Bratcher, 19883; T. albomarginata Deshayes, 18591 ∗; T. amanda Hinds, 18441; T. anilis (Röding, 1798) 1; T. archimedis Deshayes, 18591 ∗; T. argosyia Olsson, 19712; T. argus Hinds, 18441; T. babylonia Lamarck, 18221; T. balabacensis Aubry & Picardal, 20113; T. bellanodosa Grabau & King, 19283; T. binii (Aubry, 2014) 2; T. boucheti (Bratcher, 1981) 2; T. bratcherae Cernohorsky, 19873; T. caddeyi (Bratcher & Cernohorsky, 1982) 2; T. caelata Adams & Reeve, 18501 ∗; T. castaneostriata Kosuge, 19793; T. cingulifera Lamarck, 18221; T. circinata Deshayes, 18572 ∗; T. cognata E. A. Smith, 18773; T. commaculata (Gmelin, 1791) 2; T. connelli (Bratcher & Cernohorsky, 1985) 2; T. consobrina Deshayes, 18571; T. corrugata Lamarck, 18221; T. cossignanii Aubry, 20082; T. cumingii (Deshayes, 1857) 1; T. deshayesii Reeve, 18602; T. donpisori Terryn, 20173; T. elliscrossi (Bratcher, 1979) 1; T. erythraeensis Terryn & Dekker, 20173; T. eximia (Deshayes, 1859) 2; T. fenestrata (Hinds, 1844) 1; T. fernandae Aubry, 19953; T. fernandesi Bouchet, 19833; T. fijiensis (E. A. Smith, 1873) 3; T. floridana (Dall, 1889) 3; T. formosa Deshayes, 18571; T. fujitai (Kuroda & Habe, 1952) 1; T. funiculata Hinds, 18441; T. gabriellae Aubry, 20083; T. gaiae Aubry, 20083; T. giorgioi Aubry, 19993; T. grayi E. A. Smith, 18773; T. guineensis Bouchet, 19833; T. guttata (Röding, 1798) 1; T. helichrysum Melvill & Standen, 19033; T. histrio Deshayes, 18573; T. hoaraui (Drivas & Jay, 1988) 1; T. insalli (Bratcher & Burch, 1976) 2; T. irregularis Thiele, 19253; T. jacksoniana (Garrard, 1976) 3; T. jenningsi (Burch, 1965) 1; T. knudseni Bratcher, 19832; T. laevigata Gray, 18341; T. lauretanae Tenison-Woods, 18783; T. levantina Aubry, 19993; T. ligata Hinds, 18442; T. lillianae Withney, 19762 ∗; T. lima (Deshayes, 1857) 2; T. lindae Petuch, 19872; T. mamillata (Watson, 1886) 2; T. mariesi (E. A. Smith, 1880) 2; T. marrowae (Bratcher & Cernohorsky, 1982) 2; T. montgomeryi Burch, 19652; T. neglecta (Poppe, Tagaro & Terryn, 2009) 2; T. nodularis Deshayes, 18593; T. noumeaensis Aubry, 19993; T. ornata Gray, 18341; T. pellyi E. A. Smith, 18773; T. picta Hinds, 18443; T. polygonia Reeve, 18603; T. praelonga Deshayes, 18592; T. pretiosa Reeve, 18422; T. pseudopicta Aubry, 20083; T. pseudoturbonilla Talavera, 19753; T. punctatostriata Gray, 18342; T. punctum (Poppe, Tagaro & Terryn, 2009) 2; T. quoygaimardi Cernohorsky & Bratcher, 19761; T. raybaudii Aubry, 19932; T. reticularis (Pecchioli in Sacco, 1891) 3; T. robusta Hinds, 18441; T. rosae Aubry, 20153; T. russetae (Garrard, 1976) 3; T. salisburyi Drivas & Jay, 19982; T. stearnsii Pilsbry, 18911; T. straminea Gray, 18341 ∗; T. subangulata Deshayes, 18593; T. subulata (Linnaeus, 1767) 1; T. succinea Hinds, 18443; T. swobodai Bratcher, 19813; T. tagaroae Terryn, 20172; T. taiwanensis Aubry, 19993; T. taurina (Lightfoot, 1786) 1; T. tessellata Gray, 18342; T. tricolor G. B. Sowerby I, 18251; T. triseriata (Gray, 1834) 1; T. twilae Bouchet, 19833; T. unicolor Preston, 19081 ∗; T. vanuatuensis Aubry, 19993; T. vappereaui Tröndlé, Boutet & Terryn, 20132; T. venilia Tenison-Woods, 18793; T. vicdani (Kosuge, 1981) 2; T. virgo Schepman, 19133; T. waikikiensis (Pilsbry, 1921) 2., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 9-12, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["LAMARCK, J. B. de 1799. Prodrome d'une nouvelle classification des coquilles. Memoires de la Societe d'Histoire Naturelle de Paris, 1: 63 - 91.","DALL, W. H. 1908. Subdivisions of the Terebridae. Nautilus, 21: 124 - 125.","OYAMA, K. 1961. On some new facts of the taxonomy of Terebridae. Venus, 21: 176 - 189.","MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059."]}

Bathyterebra zhongshaensis Malcolm, Terryn & Fedosov newspecies (Fig. 2H) Type material: Holotype: MNHN IM-2013-61800, 17.4 mm; South China Sea, Helen Shoal, 19°13'N, 113°55'E, 470–494 m (ZhongSha 2015 Stn DW4138). Zoobank registration: urn:lsid:zoobank.org:act: C8461C7A-BF53- 48D4-BE1B-EF23F17413F8 Diagnostic nucleotide positions: Table 11. Description: Shell small, with orthocline spire, wide apical angle. Protoconch wide, paucispiral, 2.5 whorls, with small nucleus. Teleoconch 9 whorls. Early whorls convex in outline with strong, sharply convex ribs; subsequent whorls with narrow constriction formed by indentation, with straight ribs below suture; suture distinct, ragged; mature whorls with less convex outline, weaker and more numerous ribs, 30 on penultimate whorl. Spiral sculpture of very weak threads. Last whorl bulbous at base, but elongate with wide aperture. Columella straight, with weak fold. Overall colour white, with extensive patches of light-tan axial flammules on mature whorls. Habitat: Depths of 470– 494 m. Distribution: Only known from type locality. Etymology: Named for the Zhongsha sand bars, which form a subsurface atoll on top of the Macclesfield Bank. Remarks: Bathyterebra zhongshaensis n. sp. is similar to the deep-water species, B. coriolisi and B. benthalis. Bathyterebra coriolisi (Fig. 2E) is similar in colour, in the sculpture of the early whorls and in the shape of its aperture. However, B. coriolisi differs in having a much narrower shape, with a strong subsutural groove and fewer ribs on the early whorls, creating a band with nodes. Its subsutural groove becomes weaker, with just a slight indentation on mature whorls. Bathyterebra benthalis (Fig. 2I) is similar in colour and in overall sculpture of the shell, but differs from B. zhongshaensis n. sp. in the outline of its mature whorls being flat and the ribs on the early whorls extremely convex, creating nodes that dominate the profile of the early whorls., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 23, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844

Genus Granuliterebra Oyama, 1961 (Fig. 10A) Type species: Terebra bathyrhaphe E.A. Smith, 1875; OD. Diagnosis: Shell of small size (< 36 mm), slender, sculptured by strong ribs, bearing nodules on subsutural band and on bottom of ribs, giving concave outline to whorls. Protoconch of 2.5–4 whorls. Distribution: Indian and western Pacific Oceans. Remarks: Species in the genus Pellifronia are similar in having a concave whorl outline, but have notably weaker nodules and a less clearly defined subsutural band. Species with similar nodulose sculpture exist in the genus Neoterebra, but generally these have a wider apical angle or a very shiny texture. Duplicaria tricincta has similar sculpture, but including two raised, thin, spiral cords. Included species: Granuliterebra bathyrhaphe (E. A. Smith, 1875) 4; G. constricta (Thiele, 1925) 4; G. eddunhami Terryn & Holford, 20084; G. oliverai Terryn & Holford, 20084; G. palawanensis (Aubry & Picardal, 2011) 4; G. persica (E. A. Smith, 1877) 4., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 21, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["OYAMA, K. 1961. On some new facts of the taxonomy of Terebridae. Venus, 21: 176 - 189.","TERRYN, Y. & HOLFORD, M. 2008. The Terebridae of Vanuatu with a revision of the genus Granuliterebra Oyama, 1961. Visaya, Supplement, 3: 1 - 118."]}

Genus Neoterebra new genus (Fig. 9 F–H) Type species: Terebra specillata Hinds, 1844. Zoobank registration: urn:lsid:zoobank.org:act: 68993F36-2AD1- 4DDE-A853-8832B4A9EE30 Definition: Includes all species included in clade E4 of Modica et al. (2019) and those that show a combination of conchological, anatomical and distribution characteristics closely comparable with Neoterebra specillata or any genetically proven member of the clade. Diagnosis: Diagnostic nucleotide combinations in Table 9. Shell: Small to medium-sized (< 60 mm); uniformly creamy-white to light brown or with pattern of brown blotches; pointed multispiral or paucispiral protoconch, 1.5 to about 3 whorls. Axial sculpture of orthocline or arcuate ribs, forming axially elongated subsutural nodules on well-defined band. Spiral sculpture from weak striae in interspaces between ribs to distinct cords, overriding ribs, often forming nodes at intersection or on lower portion of ribs. Last whorl typically abruptly constricted to siphonal canal. Canal clearly separated from shell base, bearing pronounced fasciole. Anatomy: Radula, proboscis, venom gland typically absent. Radula, when present, of hypodermic type. Salivary glands typically present. Accessory proboscis structure absent. Distribution: Tropical East Pacific, Caribbean, off Brazil; intertidal to upper bathyal depths, but typically shallow water. Remarks: Shallow-water species of Neoterebra differ from their congeners from deep water of Brazil and the Caribbean, being larger at maturity and sharing characteristic sculpture of strong axial ribs that form elevated, sometimes axially elongated nodules on the subsutural band. Caribbean species have a weaker subsutural band; those from deep water commonly lack a recognizable subsutural band and display a fine cancellate pattern of narrow axial riblets, intersected by spiral cords of equal strength. Although the affinity of Panamic and Caribbean species is suggested by the analysis of COI, further studies are needed to document relationships and delineate major groupings within the genus. Etymology: The name is a combination of neo (Latin: new) and Terebra, referring to the fact that species of this new genus only occur in the New World, constituting the majority of the terebrid fauna in the region. Included species: Neoterebra acrior (Dall, 1889) 2 n. comb.; N. alagoensis (Lima, Tenorio & Barros, 2007) 2 n. comb.; N. alba (Gray, 1834) 2 n. comb.; N. allyni (Bratcher & Burch, 1970) 2 n. comb.; N. angelli (J. Gibson-Smith & W. Gibson-Smith, 1984) 2 n. comb.; N. arcas (Abbott, 1954) 2 n. comb.; N. armillata (Hinds, 1844) 1 n. comb.; N. assu (Simone, 2012) 2 n. comb.; N. berryi (Campbell, 1961) 2 n. comb.; N. biminiensis (Petuch, 1987) 2 n. comb.; N. brandi (Bratcher & Burch, 1970) 2 n. comb.; N. brasiliensis (E. A. Smith, 1873) 2 n. comb.; N. bridgesi (Dall, 1908) 3 n. comb.; N. carolae (Bratcher, 1979) 2 n. comb.; N. churea (Campbell, 1964) 2 n. comb.; N. colombiensis (Simone & Gracia, 2006) 2 n. comb.; N. concava (Say, 1826) 2 n. comb.; N. corintoensis (Pilsbry & Lowe, 1932) 2 n. comb.; N. crassireticula (Simone, 1999) 2 n. comb.; N. crenifera (Deshayes, 1859) 1 n. comb.; N. curacaoensis (De Jong & Coomans, 1988) 2 n. comb.; N. dislocata (Say, 1822) 1 n. comb.; N. doellojuradoi (Carcelles, 1953) 2 n. comb.; N. elata (Hinds, 1844) 1 n. comb.; N. frigata (Hinds, 1844) 2 n. comb.; N. glauca (Hinds, 1844) 2 n. comb.; N. glossema (Schwengel, 1942) 1 n. comb.; N. guadeloupensis n. sp. Malcolm, Terryn & Fedosov 1 n. sp.; N. guayaquilensis (E. A. Smith, 1880) 2 n. comb.; N. hancocki (Bratcher & Burch, 1970) 2 n. comb.; N. hemphilli (Vanatta, 1924) 2 n. comb.; N. hondurasiensis (Gargiulo, 2016) 2 n. comb.; N. intertincta (Hinds, 1844) 3 n. comb.; N. intumescyra (Lima, Tenorio & Barros, 2007) 2 n. comb.; N. jacquelinae (Bratcher & Burch, 1970) 2 n. comb.; N. juanica (Dall & Simpson, 1901) 2 n. comb.; N. lamyi (Terryn, 2011) 2 n. comb.; N. larvaeformis (Hinds, 1844) 1 n. comb.; N. leptapsis (Simone, 1999) 2 n. comb.; N. limatula (Dall, 1889) 1 n. comb.; N. lucana (Dall, 1908) 2 n. comb.; N. mugridgeae (García, 1999) 2 n. comb.; N. nassula (Dall, 1889) 2 n. comb.; N. pacei (Petuch, 1987) 2 n. comb.; N. panamensis (Dall, 1908) 2 n. comb.; N. pedroana (Dall, 1908) 2 n. comb.; N. petiveriana (Deshayes, 1857) 2 n. comb.; N. plicata (Gray, 1834) 2 n. comb.; N. protexta (Conrad, 1846) 2 n. comb.; N. puncturosa (Berry, 1959) 1 n. comb.; N. rancheria (Bratcher, 1988) 2 n. comb.; N. riosi (Bratcher & Cernohorsky, 1985) 2 n. comb.; N. roperi (Pilsbry & Lowe, 1932) 3 n. comb.; N. rushii (Dall, 1889) 2 n. comb.; N. sanjuanense (Pilsbry & Lowe, 1932) 2 n. comb.; N. shyana (Bratcher & Burch, 1970) 2 n. comb.; N. simonei (Lima, Tenorio & Barros, 2007) 2 n. comb.; N. specillata (Hinds, 1844) 1 n. comb.; N. spirosulcata (Simone, 1999) 2 n. comb.; N. sterigma (Simone, 1999) 2 n. comb.; N. sterigmoides (Simone & Gracia, 2006) 1 n. comb.; N. stohleri (Bratcher & Burch, 1970) 2 n. comb.; N. tiarella (Deshayes, 1857) 3 n. comb.; N. variegata (Gray, 1834) 1 n. comb.; N. vinosa (Dall, 1889) 2 n. comb.

Genus Pellifronia Terryn & Holford, 2008 (Fig. 2 A–D) Type species: Terebra jungi Lai, 2001; OD. Definition: Includes species with combination of conchological, anatomical and distributional characteristics closely comparable with Pellifronia jungi, including all species in clade A1 of Modica etal. (2019) and any species proven to be a member of this clade. Diagnosis: Diagnostic nucleotide combinations in Table 1. Shell: Medium-sized (25–50 mm), elongate, light brown. Protoconch brown, paucispiral, cyrtoconoid, 1.5–2.5 whorls. Coarse sculpture of pronounced ribs, orthocline or prosocline, sometimes arcuate on late teleoconch whorls. Axial ribs often thickened in their adapical and/or abapical part(s), giving slightly concave outline to spire whorls. Last adult whorl with flattened adapical portion and short, somewhat angulate transition to shell base. Siphonal canal moderately long, recurved at tip, shallowly notched; siphonal fasciole weak or indistinct. Aperture elongate, curved. Anatomy: Rhynchodeal introvert small; proboscis, radular sac, odontophore, salivary and venom glands well developed. Acessory proboscis structure absent. Radula with well-developed membrane, bearing 2 rows of wide marginal teeth of duplex type. Distribution: Indo-Pacific, bathyal depths. Remarks: This genus differs from the closely related Bathyterebra in having stronger axial sculpture, a more flattened whorl outline and the last adult whorl constricted to the siphonal canal with visible angulation. Species of Pellifronia can be distinguished by their coarse sculpture and swollen rib ends with slightly raised subsutural area. Similar species in other genera, e.g. Neoterebra puncturosa, N. crenifera and Granuliterebra bathyrhaphe, normally have a sharper nodular structure on the subsutural band or rib endings, or can be separated by the lack of radula (Neoterebra). Included species: Pellifronia brianhayesi (Terryn & Sprague, 2008) 2; P. jungi (Lai, 2001) 1., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 5-6, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["TERRYN, Y. & HOLFORD, M. 2008. The Terebridae of Vanuatu with a revision of the genus Granuliterebra Oyama, 1961. Visaya, Supplement, 3: 1 - 118.","MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059."]}

Genus Myurellopsis new genus (Fig. 7 G–I) Type species: Terebra undulata Gray, 1834. Zoobank registration: urn:lsid:zoobank.org:act: EC3F14B2-3595- 482B-9890-399BA78A7631 Definition: Includes all species included in the clade E5B of Modica etal. (2019) and those that show a combination of conchological, anatomical and distribution characteristics closely comparable with Myurellopsis undulata or any genetically proven member of the genus. Diagnosis: Diagnostic nucleotide combinations in Table 10. Shell: Small to medium-sized (< 50 mm), pinkish or orange, often with white subsutural band. Protoconch multispiral. Sculpture of strong, rounded, slightly undulating ribs, polished in appearance. Row of punctations with groove partially cutting ribs forming subsutural band. Interstices between ribs deep, narrow, bearing regular striae, often darkly coloured. Adapical parts of ribs forming one row of rounded nodules, separated from their lower portions by shallow depression or partial groove. Siphonal canal moderately long, tapering, with weakly developed fasciole. Anatomy: Two distinct states—either all typical foregut structures present, or all lacking; in latter case, accessory proboscis structure present. Distribution: Throughout Indo-Pacific; intertidal to upper bathyal depths (to 358 m), typically shallow. Remarks: Species in Myurellopsis n. gen. appear conchologically closest to Myurella species, but can be differentiated by the sharper, more elevated ribs, often with nodes on the subsutural band or top of the whorls. Etymology: The name refers to the close resemblance between its members and species in the genus Myurella. Gender feminine. Included species: Myurellopsis alisi (Aubry, 1999) 3 n. comb.; M. columellaris (Hinds, 1844) 1 n. comb.; M. guphilae (Poppe, Tagaro & Terryn, 2009) 1 n. comb.; M. joserosadoi (Bozzetti, 2001) 1 n. comb.; M. kilburni (R. D. Burch, 1965) 1 n. comb.; M. monicae (Terryn, 2005) 2 n. comb.; M. nathaliae (Drivas & Jay, 1988) 1 n. comb.; M. parkinsoni (Bratcher & Cernohorsky, 1976) 1 n. comb.; M. paucistriata (E. A. Smith, 1873) 1 n. comb.; M. undulata (Gray, 1834) 1 n. comb.; M. vaubani (Aubry, 1999) 3 n. comb. Genera defined based on conchological characters

Partecosta bozzettii Malcolm & Terryn new species (Figs 3K, 11E) Type material: Holotype: MNHN IM- 2009-10163, lv, 12.5 mm; Port d’Ehoala, SMadagascar, 25 ° 03 ' 43.9 ''S, 46 ° 57 ' 42.9 ''E, 3–4 m (ATIMO VATAE Stn TP29). Paratype 1: MNHN IM-2009- 10162, lv, 11.8 mm; type locality. Zoobank registration: urn:lsid:zoobank.org:act: 6E3E67B5-1B55- 4A99-8D39-6517DA4D065C Diagnostic nucleotide positions: Table 11. Description: Shell small. Protoconch paucispiral, 1 whorl, with broad nucleus. Holotype 9 teleoconch whorls; outline of whorls weakly convex. No subsutural band, or demarcation thereof, except for minor indentation of ribs; no visiblespiral sculpture. Axial sculpture of almost straight, thin ribs, 14 on penultimate whorl, stretching across whorl height; interspaces wider than ribs. Last whorl elongate, tapering towards base. Columella straight, aperture elongate. Shell colour off-white with darker-tinged spiral band adapically on whorls and below periphery; brownish blotches between the axial ribs abapically on whorls; inner columella dark brown or purplish. Habitat: In sand at 3– 4 m. Distribution: Known only from type locality. Etymology: The species honours Luigi Bozzetti (Italy), who has contributed tremendously to the knowledge of the molluscan fauna of SMadagascar. Remarks: The species resembles P. trilineata (Fig. 11F), but differs by its more globose and convex whorls and by having a single row of brown blotches instead of two. In addition, the protoconch of P. trilineata is differently shaped, having two whorls and a medium nucleus. The phylogenetic analysis of Modica et al. (2019) shows significant genetic distance between the two species. Partecosta bozzettii n. sp. shows some resemblance to P. albofuscata, but again differs by its more convex whorls, distinct axial ribbing and shell colour pattern. Partecosta bozzettii n. sp. is similar in general morphology and shell colour pattern to Hastula daniae (Fig. 2G) from West Africa, but the latter has a multispiral protoconch and more globose mature whorls., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 24, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059."]}

Genus Bathyterebra new genus (Fig. 2 E–I) Type species: Terebra benthalis Dall, 1889. Zoobank registration: urn:lsid:zoobank.org:act: EF1F9047-25AB- 4400-A1D4-C705B757E114 Definition: Includes species with combination of conchological, anatomical and distributional characteristics closely comparable with Bathyterebra benthalis, including all species in clade A2 of Modica etal. (2019) and any species proven to be a member of this clade. Diagnosis: Diagnostic nucleotide combinations in Table 1. Shell: Small, 12–30 mm, elongate, white to tan. Protoconch paucispiral, cyrthoconoid, about 1.5 glossy whorls. Sculpture of fine dense orthocline ribs, interrupted by depression in adapical portion of whorl. Axial ribs weakening towards anterior throughout their height on mature whorls, often forming weak nodules bordering adapical depression, giving distinctly angulated outline to early spire whorls; late teleoconch whorls gently convex. Last adult whorl evenly convex, transitioning to shell base without visible angulation. Siphonal canal stout, slightly recurved at tip, shallowly notched; siphonal fasciole weak or strong. Aperture elongate, wide. Anatomy: Rhynchodeal introvert small; proboscis, radular sac, salivary and venom glands well developed. Accessory proboscis structure absent. Radula with well-developed membrane, bearing 2 rows of wide marinal teeth of duplex type. Distribution: South China Sea, New Caledonia, Caribbean; bathyal depths. Etymology: Name refers to bathyal depths, at which three known species of the genus were sampled. Gender feminine. Remarks: Bathyterebra spp. can be differentiated from members of most other genera of Terebridae by the combination of relatively faint sculpture lacking spiral elements, weakly defined subsutural band and relatively high last adult whorl with convex outline. In shell proportions the genus resembles Oxymeris; however, the latter has more flattened whorls and typically can be easily differentiated by a larger size at maturity. Included species: Bathyterebra benthalis (Dall, 1889) 1 n. comb.; B. coriolisi (Aubry, 1999) 1 n. comb.; B. zhongshaensis Malcolm, Terryn & Fedosov 1 n. sp.

Genus Gradaterebra Cotton & Godfrey, 1932 (Fig. 10D) Type species: Gradaterebra scalariformis Cotton & Godfrey, 1932; OD. Diagnosis: Shell small (< 25 mm), often dull-coloured, with flattened tobulbouswhorls; axialsculptureof prominentribs; spiralsculpture absent, microscopic or represented by fine cords; last whorl inflated. Distribution: Indian and SW Pacific Oceans. Remarks. The genus mainly comprises small species from Australia and South Africa, which can be differentiated based on adult size < 25 mm. Included species: Gradaterebra scalariformis (Cotton & Godfrey, 1932) 4 n. comb.; G. assecla (Iredale, 1924) 4 n. comb.; G. capensis (E. A. Smith, 1873) 4 n. comb.; G. easmithi (Aubry, 1999) 4 n. comb.; G. kowiensis (Turton, 1932) 4 n. comb.; G. lightfooti (E. A. Smith, 1899) 4 n. comb.; G. ninfae (Campbell, 1961) 4 n. comb.; G. pilsbryi (Aubry, 1999) 4 n. comb.; G. planecosta (Barnard, 1958) 4 n. comb.; G. severa (Melvill, 1897) 4 n. comb.; G. sorrentense (Aubry, 1999) 4 n. comb.; G. taylori (Reeve, 1860) 4 n. comb., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 21-22, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844

Genus Duplicaria Dall, 1908 (Fig. 3 A–E) Synonyms: Diplomeriza Dall, 1919 (unnecessary new name for Duplicaria Dall, 1908, treated by Dall as homonym of nonexistent ‘ Duplicaria Rafinesque, 1833 ’; type species Buccinum duplicatum Linnaeus, 1758, by typification of replaced name); Myurellisca Bartsch 1923, type species Myurella duplicatoides Bartsch, 1923, OD); Pervicacia Iredale, 1924 (type species Terebra ustulata Deshayes, 1857, OD). Type species: Buccinum duplicatum Linnaeus, 1758; OD. Definition: Primarily defined based on phylogenetic analysis, including all species in clade F1 of Modica etal. (2019) and, in addition, species that show shell morphology, anatomical or genetic characteristics closely comparable with Duplicaria duplicata or any genetically proven member of genus. Diagnosis. Diagnostic nucleotide combinations in Table 2. Shell: Elongate, small to large, 20–100 mm, with wide aperture, predominant axial sculpture, either continuous from suture to suture or interrupted by shallow to deep subsutural indentation. Spiral structure obsolete, sometimes microscopic striae or weak band joining nodes. Anatomy: Rhynchodeal introvert small; salivary glands and radular sac present; radula of solid curved marginal teeth attached to strong membrane. Venom gland, proboscis and accessory proboscis structure absent. Distribution: Tropical and subtropical Indo-Pacific, West Africa. Remarks: According to shell morphology, three distinct clusters can be delineated: Duplicaria duplicata group. Small to large shells, 25–100 mm; paucispiral protoconch; sculpture of strong axial ribs. Deep narrow spiral groove delineating distinct flattened subsutural band; no spiral sculpture. Whorl outline flattened, or shouldered due to elevated ribs below subsutural band. Duplicaria tristis group. Small shells, 10–30 mm; paucispiral protoconch; sculpture of strong axial ribs, subsutural area weakly impressed. Duplicaria tricincta group. Small shells, 10–25 mm; paucispiral protoconch; sculpture of strong spiral ribs. Amore convex shape with outline flattened due to elevation of ribs below subsutural area. Ribs weakening on mature whorls. Included species: Duplicaria albozonata (E. A. Smith, 1875) 2; D. australis (E. A. Smith, 1873) 2; D. badia (Deshayes, 1859) 2; D. bernardii (Deshayes, 1857) 1; D. brevicula (Deshayes, 1859) 1; D. concolor (E. A. Smith, 1873) 2; D. copula (Hinds, 1844) 2; D. costellifera (Pease, 1869) 2; D. crakei (Burch, 1965) 2; D. duplicata (Linnaeus, 1758) 1; D. dussumierii (Kiener, 1839) 2; D. evoluta (Deshayes, 1859) 2; D. fictilis (Hinds, 1844) 2; D. gemmulata (Kiener, 1839) 3; D. helenae (Hinds, 1844) 3; D. herberti n. sp. Malcolm, Terryn & Fedosov 1; D. hiradoensis (Pilsbry, 1921) 2; D. jukesi (Deshayes, 1857) 2; D. juliae (Aubry, 1999) 2; D. kieneri (Deshayes, 1859) 2; D. kirai (Oyama, 1962) 2; D. koreana (Yoo, 1976) 2; D. morbida (Reeve, 1860) 2; D. mozambiquensis Bratcher & Cernohorsky, 19821; D. silvanae (Aubry, 1999) 3; D. similis (E. A. Smith, 1873) 2; D. sowerbyana (Deshayes, 1857) 2; D. tricincta (E. A. Smith, 1877) 1; D. tristis (Deshayes, 1859) 1; D. ustulata (Deshayes, 1857) 2; D. veronicae (Nicolay & Angioy, 1993) 2., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 7-9, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["DALL, W. H. 1908. Subdivisions of the Terebridae. Nautilus, 21: 124 - 125.","MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059."]}

Genus Pristiterebra Oyama, 1961 (Fig. 10C) Type species: Terebra tsuboiana Yokoyama, 1922 †; OD. Synonym: Laeviacus Oyama, 1961 (type species Terebra chibana Yokoyama, 1922; OD). Diagnosis: Shell small to medium-sized (20–65 mm). Whorls flattened to convex, with predominant pustulose or seldom cancellate sculpture. Whorl periphery shifted abapically, leading to truncated appearance of shell, with shell base abruptly constricted to short siphonal canal. Aperture wide, especially in anterior portion. Distribution: Western Pacific. Remarks: Acharacteristic truncated outline with a short siphonal canal and broad aperture allow for easy recognition of Pristiterebra among other terebrid genera. Species in Pristiterebra are nearest to the nodulose species of Neoterebra (N. tuberculosa, N. glauca, N. petiveriana), which can be recognized by the presence of a distinct subsutural groove. Some species of Gradaterebra and Partecosta have similar shell outline, but both can be differentiated from Pristiterebra by dominant axial sculpture or, rarely, complete lack of sculpture, and (in Partecosta) by generally smaller size. Included species: Pristiterebra bifrons (Hinds, 1844) 4; P. frausseni Poppe, Tagaro & Terryn, 20094 n. comb.; P. miranda (E. A. Smith, 1873) 4 n. comb.; P. pustulosa (E. A. Smith, 1879) 4 n. comb.; P. tuberculosa (Hinds, 1844) 4 n. comb., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 21, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["OYAMA, K. 1961. On some new facts of the taxonomy of Terebridae. Venus, 21: 176 - 189."]}

Genus Hastula H. Adams & A. Adams, 1858 (Fig. 6 A–E) Type species: Buccinum strigilatum Linnaeus, 1758; SD, Cossmann (1896). Synonyms: Impages E.A. Smith, 1873 (type species Terebra caerulescens Lamarck, 1822 = Buccinum hecticum Linnaeus, 1758; SD, Cossmann, 1896); Acuminia Dall, 1908 (type species Buccinum lanceatum Linnaeus, 1767; OD); Hastulina Oyama, 1961 (type species Terebra casta Hinds, 1844; OD); Egentelaria Rehder, 1980 (type species Terebra stylata Hinds, 1844; OD). Definition: Includes all species in clade D of Modica et al. (2019), and those that show a combination of conchological, anatomical and distribution characteristics closely comparable with Hastula strigilata or any genetically proven member of the clade. Diagnosis: Diagnostic nucleotide combinations provided for entire clade with exception of four specimens of H. albula (see Material and Methods) and, separately, for subclade H. albula – H. natalensis – H. aff. casta of clade D (Table 4). Shell: Small to medium-sized, shiny, 12–90 mm. Protoconch paucispiral or multispiral, about 1–5 whorls. Axial sculpture of crenulations on subsutural portion of whorl, or fine ribs on part or throughout whorl height; spiral sculpture of single subsutural groove, occasionally with additional row of punctations. Siphonal canal stout, very short, with well-developed fasciole. Aperture elongate to rounded. Genus Hastula (except four specimens of H. albula) Species/sequences analysed 18/91 Diagnostic nucleotides: 28: A, 67: T, 95: C, 181: T, 221: C, 222: C, 253: A, 581: T, 598: A Subclade H. albula – H. natalensis – H. aff. casta Anatomy: Rhynchodeal introvert, proboscis, venom gland, salivary glands and radula well developed. Radula of hypodermic marginal teeth attached to thin, reduced membrane; walls of marginal teeth often with multiple perforations (Fig. 6E, F) Species/sequences analysed 3/17 Diagnostic nucleotides 49: G, 347: C, 349: T, 517: T Distribution: Indo-Pacific, West Africa, Caribbean. Remarks: Species of Hastula are rather easy to recognize among Terebridae by their typically shiny shells, with fine close-set axial ribs and usually by the lack of a clearly defined subsutural band. The shell morphology in the Partecosta macleani group closely resembles that of Hastula; however, the latter is usually larger at maturity, while shells of comparable size can be differentiated by texture—a polished surface in Hastula vs dull in Partecosta. Furthermore, species of Hastula possess a venom gland and hypodermic radular teeth, whereas Partecosta species lack a venom gland and their radulae teeth are solid. Included species: Hastula aciculina (Lamarck, 1822) 2; H. acumen (Deshayes, 1859) 1; H. alboflava Bratcher, 19882; H. albula (Menke, 1843) 1; H. androyensis Bozzetti, 20082; H. anomala (Gray, 1834) 2; H. anosyana (Bozzetti, 2016) 2; H. apicitincta (G. B. Sowerby III, 1900) 2; H. bacillus (Deshayes, 1859) 2; H. casta (Hinds, 1844) 1; H. celidonota (Melvill & Sykes, 1898) 2; H. cernohorskyi Burch, 19652; H. cinerea (Born, 1778) 1; H. continua Deshayes, 18592; H. crossii (Deshayes, 1859) 1 ∗; H. cuspidata (Hinds, 1844) 2; H. denizi Rolán & Gubbioli, 20002; H. engi Malcolm & Terryn, 20172; H. escondida (Terryn, 2006) 2; H. exacuminata Sacco, 18912; H. filmerae (G. B. Sowerby III, 1906) 2; H. hamamotoi Tsuchida & Tanaka, 19992; H. hastata (Gmelin, 1791) 1; H. hectica (Linnaeus, 1758) 1; H. imitatrix (Auffenberg & Lee, 1988) 2; H. inconstans (Hinds, 1844) 2; H. knockeri (E. A. Smith, 1872) 2; H. lanceata (Linnaeus, 1767) 1; H. leloeuffi Bouchet, 19832; H. lepida (Hinds, 1844) 2; H. marqueti (Aubry, 1994) 2; H. maryleeae R. D. Burch, 19652; H. matheroniana (Deshayes, 1859) 1; H. nana (Deshayes, 1859) 2; H. natalensis (E. A. Smith, 1903) 1 ∗; H. parva (Baird, 1873) 1; H. penicillata (Hinds, 1844) 1; H. philippiana (Deshayes, 1859) 2; H. puella (Thiele, 1925) 1; H. raphanula (Lamarck, 1822) 1; H. rufopunctata (E. A. Smith, 1877) 2; H. salleana (Deshayes, 1859) 1; H. sandrogorii Ryall, Terryn & Rosado, 20172; H. solida (Deshayes, 1857) 1; H. strigilata (Linnaeus, 1758) 1; H. stylata (Hinds, 1844) 1; H. tenuicolorata Bozzetti, 20081; H. tiedemani Burch, 19652; H. venus Aubry, 20082; H. verreauxi (Deshayes, 1857) 1; H. westralica (Aubry, 1999) 2; H. daniae (Aubry, 2008) 3., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 12-14, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["COSSMANN, M. 1896. Essais de paleoconchologie comparee, Vol. 2. Author and Comptoir Geologique, Paris.","DALL, W. H. 1908. Subdivisions of the Terebridae. Nautilus, 21: 124 - 125.","OYAMA, K. 1961. On some new facts of the taxonomy of Terebridae. Venus, 21: 176 - 189.","MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059."]}

Subfamily PERVICACIINAE Rudman, 1969 (Fig. 3) Type genus: Pervicacia Iredale, 1924 (= Duplicaria Dall, 1908) Shell: Elongate, 10–100 mm, with high spire, predominant axial sculpture, wide aperture. Anatomy: Rhynchodeal introvert rather small; salivary glands and radular sac present; radula of solid curved marginal teeth attached to strong membrane. Venom gland, proboscis and accessory proboscis structure absent., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 7, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["RUDMAN, W. B. 1969. Observations on Pervicacia tristis (Deshayes, 1859) and comparison with other toxoglossan gastropods. Veliger, 12: 53 - 64.","DALL, W. H. 1908. Subdivisions of the Terebridae. Nautilus, 21: 124 - 125."]}

Genus Oxymeris Dall, 1903 (Fig. 6 F–H) Type species: Buccinum maculatum Linnaeus, 1758; OD. Synonyms: Abretia H. & A. Adams, 1853 (invalid, junior homonym of Abretia Rafinesque, 1814; type species Terebra cerithina Lamarck, 1822; SD, Dall, 1908); Abretiella Bartsch, 1923 (replacement name for Abretia H. & A. Adams, 1853; type species Terebra cerithina Lamarck, 1822; by typification of replaced name); Nototerebra Cotton, 1947 (type species Terebra albida Gray, 1834; OD). Definition: Includes all species in clade B of Modica et al. (2019), and those that show a combination of conchological, anatomical and distribution features closely comparable with Oxymeris maculata or any genetically proven member of the clade. Diagnosis: Diagnostic nucleotide combinations in Table 5. Shell: Medium-sized to large, reaching 273 mm; weakly sculptured, typically with orthoconoid spire and rather wide aperture. Protoconch multispiral, 3–3.75 whorls. Sculpture of subsutural band, smooth or with regular crenulations; juvenile sculpture often consisting of axial ribs. Whorls flattened in outline; last adult whorl inflated; siphonal canal short, stout. Aperture wide, elliptic to rounded. Anatomy: Rhynchodeal introvert very large, proboscis absent or vestigial, salivary glands absent or very small, radula venom gland and accessory proboscis structure absent. Distribution: Indo-Pacific, tropical East Pacific, West Africa. Remarks: Species of Oxymeris are among the largest terebrids and can usually be easily identified by their large, shiny shells, with an orthoconoid or even slightly acuminate spire, often with a bulbous last whorl, a very wide aperture and a very short siphonal canal separated from the shell base by a deep groove. Nevertheless, some smaller species (e.g. O. cerithina) have a spire with convex sides and superficially resemble medium-sized Punctoterebra and Myurella species. However, in most cases (except O. crenulata), the Oxymeris species can be recognized by the complete lack of axial sculpture other than growth lines, while axial elements of varying strength and shape are present in Punctoterebra, Myurella and most other terebrid genera. Finally, widely spaced deep spiral grooves differentiate Perirhoe from Oxymeris. Included species: Oxymeris albida (Gray, 1834) 2; O. areolata (Link, 1807) 1; O. barbieri (Aubry, 2008) 2; O. caledonica (G. B. Sowerby III, 1909) 1; O. cerithina (Lamarck, 1822) 1; O. chlorata (Lamarck, 1822) 1; O. consors (Hinds, 1844) 1; O. crenulata (Linnaeus, 1758) 1; O. dillwynii (Deshayes, 1859) 2; O. dimidiata (Linnaeus, 1758) 1; O. fatua (Hinds, 1844) 2; O. felina (Dillwyn, 1817) 1; O. gouldi (Deshayes, 1857) 2; O. lineopunctata (Bozzetti, 2008) 2; O. maculata (Linnaeus, 1758) 1; O. ngai Thach, 20162; O. senegalensis (Lamarck, 1822) 2; O. strigata (G. B. Sowerby I, 1825) 1; O. suffusa (Pease, 1869) 2; O. swinneni Terryn & Ryall, 20142; O. trochlea (Deshayes, 1857) 2; O. troendlei (Bratcher, 1981) 1., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 14, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["DALL, W. H. 1908. Subdivisions of the Terebridae. Nautilus, 21: 124 - 125.","MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059."]}

Genus Profunditerebra new genus (Fig. 9 A–E) Type species: Profunditerebra papuaprofundi n. sp. Zoobank registration: urn:lsid:zoobank.org:act: A5A78A3D-AB11- 41C3-86F0-7C1451512FF4 Definition: Includes all species included in clade E3 of Modica et al. (2019) and those that show a combination of conchological, anatomical and distribution characteristics closely comparable with Profunditerebra papuaprofundi n. sp. or any genetically proven member of the clade. Diagnosis: Diagnostic nucleotide combinations provided in Table 8. Shell: Small to medium-sized (< 45 mm), with multispiral protoconch and slender siphonal canal. Whorls subcylindrical or flattened. Sculpture varying greatly; axial sculpture of rounded ribs, weak to very strong and elevated; spiral sculpture typically limited to striae in interstices and a subsutural groove, but may be absent or represented by strong cords generating overall cancellate pattern. Anatomy: Proboscis, radula, venom gland and salivary glands present. Radula of duplex marginal teeth (Fig. 9E). Accessory proboscis structure absent. Distribution: Almost exclusively deep water (typically 150–600 m) of tropical Indo-West Pacific from Taiwan to New Caledonia; one species known from shallow water of South Australia. Remarks: Profunditerebra is the only terebrine genus with duplex radular teeth similar to those in Pellifroniinae; however, members of the two groups can easily be differentiated by shell characters. Species of Profunditerebra can be separated into four clusters based on their shell morphology; however, only the first cluster corresponds to a phylogenetic lineage. Profunditerebra papuaprofundi n. sp. group. Shell uniformly coloured white to dark brown; heavily sculptured, with distinct subsutural groove and subcylindrical whorls. Sculpture of strong rounded ribs forming row of prominent nodules on subsutural band. Spiral sculpture of fine striae limited to interstices between axials, or overriding them to form fine continuous cords (including on subsutural band), or coarse cancellate pattern. Species of Neoterebra (e.g. N. armillata) exhibit a similar sculpture, but can be differentiated from Profunditerebra by their flattened whorls. The Punctoterebra textilis group also includes some similar forms (see Remarks on Punctoterebra). Profunditerebra orientalis group. Shell with flattened whorl profile; variegated colour pattern. Sculpture of fine orthocline ribs intersected by cords of about equal strength or slightly weaker than axials, to form delicate cancellate pattern. Subsutural band demarcated by shallow punctuate groove. Species of this group can easily be differentiated from congeners that either lack spiral elements, or display an overall much coarser sculpture. Both P. orientalis and P. hiscocki closely resemble species of Maculauger, from which they can confidently be differentiated only by anatomy (all studied species of Maculauger lack a radula) or molecular characters. Profunditerebra anseeuwi group. Shell with flattened whorl outline; subsutural band represented by short ribs or indentation. Sculpture of axial ribs, pronounced throughout whorl height, but weakening and becoming obsolete on later whorls; spiral sculpture absent. Species of this group are conchologically close to Duplicaria and to the Punctoterebra teramachii group; however, they can be differentiated from both by the overall weaker sculpture. In addition, these lineages can confidently be differentiated by the radular teeth morphology, Duplicaria having solid and recurved teeth, Profunditerebra duplex teeth and Punctoterebra flat teeth. Profunditerebra brazieri stands apart from other species in the genus, being endemic to temperate shallow water off Tasmania. Morphologically, it is close to the P. anseeuwi group, but has a very weakly defined subsutural band. Etymology: Name refers to the considerable depths at which almost all known species of the genus occur. Gender feminine. Included species: Profunditerebra anseeuwi (Terryn, 2005) 1 n. comb.; P. brazieri (Angas, 1871) 1 n. comb.; P. evelynae (Clench & Aguayo, 1939) 3 n. comb.; P. macclesfieldensis n. sp. Malcolm, Terryn & Fedosov 1 n. sp.; P. orientalis (Aubry, 1999) 1 n. comb.; P. papuaprofundi n. sp. Malcolm, Terryn & Fedosov 1 n. sp.; P. poppei (Terryn, 2003) 1 n. comb.; P. hiscocki (Sprague, 2004) 1 n. comb., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 17-18, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059."]}

Genus Duplicaria Dall, 1908 (Fig. 3 A–E) Synonyms: Diplomeriza Dall, 1919 (unnecessary new name for Duplicaria Dall, 1908, treated by Dall as homonym of nonexistent ‘ Duplicaria Rafinesque, 1833 ’; type species Buccinum duplicatum Linnaeus, 1758, by typification of replaced name); Myurellisca Bartsch 1923, type species Myurella duplicatoides Bartsch, 1923, OD); Pervicacia Iredale, 1924 (type species Terebra ustulata Deshayes, 1857, OD). Type species: Buccinum duplicatum Linnaeus, 1758; OD. Definition: Primarily defined based on phylogenetic analysis, including all species in clade F1 of Modica etal. (2019) and, in addition, species that show shell morphology, anatomical or genetic characteristics closely comparable with Duplicaria duplicata or any genetically proven member of genus. Diagnosis. Diagnostic nucleotide combinations in Table 2. Shell: Elongate, small to large, 20–100 mm, with wide aperture, predominant axial sculpture, either continuous from suture to suture or interrupted by shallow to deep subsutural indentation. Spiral structure obsolete, sometimes microscopic striae or weak band joining nodes. Anatomy: Rhynchodeal introvert small; salivary glands and radular sac present; radula of solid curved marginal teeth attached to strong membrane. Venom gland, proboscis and accessory proboscis structure absent. Distribution: Tropical and subtropical Indo-Pacific, West Africa. Remarks: According to shell morphology, three distinct clusters can be delineated: Duplicaria duplicata group. Small to large shells, 25–100 mm; paucispiral protoconch; sculpture of strong axial ribs. Deep narrow spiral groove delineating distinct flattened subsutural band; no spiral sculpture. Whorl outline flattened, or shouldered due to elevated ribs below subsutural band. Duplicaria tristis group. Small shells, 10–30 mm; paucispiral protoconch; sculpture of strong axial ribs, subsutural area weakly impressed. Duplicaria tricincta group. Small shells, 10–25 mm; paucispiral protoconch; sculpture of strong spiral ribs. Amore convex shape with outline flattened due to elevation of ribs below subsutural area. Ribs weakening on mature whorls. Included species: Duplicaria albozonata (E. A. Smith, 1875) 2; D. australis (E. A. Smith, 1873) 2; D. badia (Deshayes, 1859) 2; D. bernardii (Deshayes, 1857) 1; D. brevicula (Deshayes, 1859) 1; D. concolor (E. A. Smith, 1873) 2; D. copula (Hinds, 1844) 2; D. costellifera (Pease, 1869) 2; D. crakei (Burch, 1965) 2; D. duplicata (Linnaeus, 1758) 1; D. dussumierii (Kiener, 1839) 2; D. evoluta (Deshayes, 1859) 2; D. fictilis (Hinds, 1844) 2; D. gemmulata (Kiener, 1839) 3; D. helenae (Hinds, 1844) 3; D. herberti n. sp. Malcolm, Terryn & Fedosov 1; D. hiradoensis (Pilsbry, 1921) 2; D. jukesi (Deshayes, 1857) 2; D. juliae (Aubry, 1999) 2; D. kieneri (Deshayes, 1859) 2; D. kirai (Oyama, 1962) 2; D. koreana (Yoo, 1976) 2; D. morbida (Reeve, 1860) 2; D. mozambiquensis Bratcher & Cernohorsky, 19821; D. silvanae (Aubry, 1999) 3; D. similis (E. A. Smith, 1873) 2; D. sowerbyana (Deshayes, 1857) 2; D. tricincta (E. A. Smith, 1877) 1; D. tristis (Deshayes, 1859) 1; D. ustulata (Deshayes, 1857) 2; D. veronicae (Nicolay & Angioy, 1993) 2.

Neoterebra guadeloupensis Malcolm, Terryn & Fedosov newspecies (Fig. 13A, B) Type material: Holotype: MNHN IM-2013-61448, 1 lv, 17.2 mm; S Marie-Galante, Guadeloupe, 15 ° 50 'N, 61 ° 19 'W, 305–312 m (N/O Antea, KARUBENTHOS 2015 Stn DW4638). Paratypes 1–3: off Fajou, Guadeloupe, 16 ° 22 'N, 61 ° 34 ', 80 m (KARUBENTHOS 2012 Stn GD02). Paratype 1: MNHN IM-2013-20542, 1 lv, 6.5 mm. Paratype 2: MNHN IM-2013-20541, 1 lv, 5.7 mm. Paratype 3: MNHN IM-2013-20539, 1 lv, 10.4 mm. Paratype 4: MNHN IM-2013-9110, 1 lv, 7 × 2.1 mm; Port Louis, Guadeloupe, 16 ° 22 ' 46 'N, 61 ° 34 'W, 66 m (KARUBENTHOS 2012 Stn GD35). Zoobank registration: urn:lsid:zoobank.org:act: 57E995EE-24F0- 47A0-9DDC-E9D54948B056 Diagnostic nucleotide positions: Table 11. Description: Shell small. Protoconch multispiral, transparent with 3 whorls tinged yellow; protoconch/teleoconch transition clear-cut with visible sinusigera. Teleoconch of holotype with 15 whorls; overall whorl outline straight to convex. Weak axial sculpture of 15 narrow axial ribs with wider interspaces, increasing with maturity to 25 on penultimate whorl. Deep subsutural groove crossed obliquely by ribs. Subsutural band with strong elongate nodes; subsequently, 2 strongly projecting spiral cords at top and middle of whorl dominating sculpture, forming reticulate pattern of deep indentations with thin ribs; the upper cord projects in outline beyond subsutural band. On mature whorls interspaces sometimes with 1 or 2 weak spiral striae. Reticulate sculpture continuing on shell base. Aperture long with weak callus; columella with 2 weak folds. Shell lacks any pattern; coloured in shades of white to yellow. Habitat: Deep water, 60– 310 m. Distribution: Only known from off Guadeloupe. Etymology: Named after the type locality, the French Caribbean island of Guadeloupe. Remarks: Anumber of species of Terebridae have been described from the Western Atlantic and Caribbean Sea that have a similar strong reticulate sculpture. These include N. limatula that has more than 2 strong spiral cords in addition to the subsutural band (Fig. 13C) and a paucispiral protoconch. Neoterebra colombiensis is very similar to N. guadeloupensis n. sp., but the spiral sculpture in the former is less dominant, giving a more uniform reticulate sculpture with nodes at the intersections. In addition, N. colombiensis has a paucispiral protoconch in comparison with the multispiral protoconch of N. guadeloupensis n. sp. Neoterebra simonei is similar in colour and sculpture, but has 3 spiral cords in addition to the subsutural band and has a distinctive paucispiral protoconch of 1.5 whorls. Other similar species from the region, e.g. N. intumescyra and N. alagoensis (Fig. 9F), all have paucispiral protoconchs and different sculpture., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 25-26, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844

Subfamily PELLIFRONIINAE new subfamily (Fig. 2) Type genus: Pellifronia Terryn & Holford, 2008 Zoobank registration: urn:lsid:zoobank.org:act: E827742C-E1B3- 42DD-B5EB-5D7EAA6E2C95 Diagnosis: Diagnostic nucleotide combinations in Table 1. Shell: Small to medium-sized (12–50 mm), with predominant axial sculpture. Early teleoconch whorls often with angled profile, later ones flattened or even slightly concave to convex. Sculpture of numerous close-set, evenly spaced, ribs, or forming raised nodules bordering upper or lower suture. Spiral sculpture absent or of very shallow, wide subsutural depression. Anatomy: Rhynchodeal introvert small; proboscis, radular sac, odontophore, salivary and venom glands well developed. Radula with well-developed membrane, bearing 2 rows of wide marginal teeth of duplex type. Accessory proboscis structure absent. Remarks: This subfamily comprises five described species in two genera, Bathyterebra and Pellifronia, that occur in the Indo-Pacific and Caribbean at bathyal depths. The anatomy of the studied species shows characters that are believed to be ancestral for Terebridae in general, i.e. weakly developed rhynchodeal introvert and radular teeth of duplex type (Castelin et al., 2012).

Genus Partecosta Dance & Eames, 1966 (Fig. 3 F–O) Type species: Strioterebrum wilkinsi Dance & Eames, 1966 (= Terebra fuscobasis E.A. Smith, 1877); OD. Definition: Includes species with combination of conchological, anatomical and distributional characteristics closely comparable with Strioterebrum wilkinsi, including all species in clade F2 of Modica et al. (2019) and any species proven to be a member of this clade. Diagnosis: Diagnostic nucleotide combinations in Table 2. Shell: Small, 10–30 mm, pale or tan, often with spiral bands and dark basal area. Protoconch 1.5–3 whorls. Predominant sculpture of axial ribs spanning whorl height, resulting in even, slightly convex whorl outline, typically interrupted by depression or narrow groove forming subsutural row of nodules. Spiral sculpture indistinct or of fine regular striae. Siphonal canal very short, stout, indistinctly or shallowly notched, with strong fasciole. Aperture wide, pyriform. Anatomy: Rhynchodeal introvert small; salivary glands and radular sac present; radula of solid curved marginal teeth attached to strong membrane. Venom gland, proboscis and accessory proboscis structure absent. Distribution: Indian Ocean; intertidal and shallow subtidal. Remarks: Three clusters with varying sculpture patterns can be recognized: Partecosta fuscobasis group: sculpture of strong ribs and clearly delineated subsutural band forming distinct nodules at intersections with ribs (Fig. 3 F–I). Closely comparable with New World genus Neoterebra, but can be differentiated by smaller size, wider aperture, usually with wide anterior notch, and solid curved radular teeth. Partecosta trilineata group: glossy shells sculptured by strong continuous ribs; no distinct spiral sculpture or subsutural band (Fig. 3K, L). Very similar sympatric forms are known in genus Punctoterebra (i.e. Punctoterebra solangeae Bozzetti, 2015 group) that can only be reliably differentiated by anatomical characters (radula lacking in P. solangeae; of solid curved teeth in Partecosta spp.). Partecosta macleani group: flattened whorl outline, very short robust siphonal canal; very weak sculpture of rounded, indistinct subsutural nodules extended into even less distinct axial folds on whorl periphery (Fig. 3M). Some species of Hastula are close to P. macleani, but are commonly larger, have polished surface and retain distinct ribbing at least on adapical portions of whorls. Included species: Partecosta albofuscata (Bozzetti, 2008) 1 n. comb.; P. bozzettii n. sp. Malcolm, Terryn & Fedosov 1; P. fuscolutea Bozzetti, 20082 n. comb.; P. macleani (Bratcher, 1988) 1 n. comb.; P. nassoides (Hinds, 1844) 1 n. comb.; P. sandrinae (Aubry, 2008) 1 n. comb.; P. trilineata (Bozzetti, 2008) 1 n. comb.; P. varia (Bozzetti, 2008) 1 n. comb.; P. fuscobasis (E.A. Smith, 1877) 2 n. comb.; P. fuscocincta (E.A. Smith, 1877) 2 n. comb.; P. herosae (Terryn & Rosado, 2011) 2 n. comb.; P. macandrewii (E.A. Smith, 1877) 2 n. comb.; P. padangensis (Thiele, 1925) 2 n. comb.; P. tantilla (E.A. Smith, 1873) 2 n. comb.; P. tenera (Hinds, 1844) 3 n. comb; P. veliae (Aubry, 1991) 3 n. comb., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 9, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059."]}

Genus Oxymeris Dall, 1903 (Fig. 6 F–H) Type species: Buccinum maculatum Linnaeus, 1758; OD. Synonyms: Abretia H. & A. Adams, 1853 (invalid, junior homonym of Abretia Rafinesque, 1814; type species Terebra cerithina Lamarck, 1822; SD, Dall, 1908); Abretiella Bartsch, 1923 (replacement name for Abretia H. & A. Adams, 1853; type species Terebra cerithina Lamarck, 1822; by typification of replaced name); Nototerebra Cotton, 1947 (type species Terebra albida Gray, 1834; OD). Definition: Includes all species in clade B of Modica et al. (2019), and those that show a combination of conchological, anatomical and distribution features closely comparable with Oxymeris maculata or any genetically proven member of the clade. Diagnosis: Diagnostic nucleotide combinations in Table 5. Shell: Medium-sized to large, reaching 273 mm; weakly sculptured, typically with orthoconoid spire and rather wide aperture. Protoconch multispiral, 3–3.75 whorls. Sculpture of subsutural band, smooth or with regular crenulations; juvenile sculpture often consisting of axial ribs. Whorls flattened in outline; last adult whorl inflated; siphonal canal short, stout. Aperture wide, elliptic to rounded. Anatomy: Rhynchodeal introvert very large, proboscis absent or vestigial, salivary glands absent or very small, radula venom gland and accessory proboscis structure absent. Distribution: Indo-Pacific, tropical East Pacific, West Africa. Remarks: Species of Oxymeris are among the largest terebrids and can usually be easily identified by their large, shiny shells, with an orthoconoid or even slightly acuminate spire, often with a bulbous last whorl, a very wide aperture and a very short siphonal canal separated from the shell base by a deep groove. Nevertheless, some smaller species (e.g. O. cerithina) have a spire with convex sides and superficially resemble medium-sized Punctoterebra and Myurella species. However, in most cases (except O. crenulata), the Oxymeris species can be recognized by the complete lack of axial sculpture other than growth lines, while axial elements of varying strength and shape are present in Punctoterebra, Myurella and most other terebrid genera. Finally, widely spaced deep spiral grooves differentiate Perirhoe from Oxymeris. Included species: Oxymeris albida (Gray, 1834) 2; O. areolata (Link, 1807) 1; O. barbieri (Aubry, 2008) 2; O. caledonica (G. B. Sowerby III, 1909) 1; O. cerithina (Lamarck, 1822) 1; O. chlorata (Lamarck, 1822) 1; O. consors (Hinds, 1844) 1; O. crenulata (Linnaeus, 1758) 1; O. dillwynii (Deshayes, 1859) 2; O. dimidiata (Linnaeus, 1758) 1; O. fatua (Hinds, 1844) 2; O. felina (Dillwyn, 1817) 1; O. gouldi (Deshayes, 1857) 2; O. lineopunctata (Bozzetti, 2008) 2; O. maculata (Linnaeus, 1758) 1; O. ngai Thach, 20162; O. senegalensis (Lamarck, 1822) 2; O. strigata (G. B. Sowerby I, 1825) 1; O. suffusa (Pease, 1869) 2; O. swinneni Terryn & Ryall, 20142; O. trochlea (Deshayes, 1857) 2; O. troendlei (Bratcher, 1981) 1.

Genus Maculauger new genus (Fig. 7E, F) Type species: Terebra pseudopertusa Bratcher & Cernohorsky, 1985. Zoobank registration: urn:lsid:zoobank.org:act: 3565DB17-E4EF- 478B-8757-3A682BB8F6CF Definition: Includes all species included in clade E5A of Modica et al. (2019) and those that show a combination of conchological, anatomical and distribution characteristics closely comparable with Maculauger pseudopertusus or any genetically proven member of the clade. Diagnosis: Diagnostic nucleotide combinations in Table 10. Shell: Small to medium-sized (< 60 mm); typically, orange-brown with irregular maculations on subsutural band. Whorl outline flattened or gently convex. Subsutural band lightly ribbed or flattened, separated from lower portion of whorl by narrow groove or row of punctations. Axial and spiral sculpture of about equal strength, producing cancellate sculpture below subsutural band. Anatomy: Proboscis, radula, venom gland and accessory proboscis structure absent; salivary glands present or absent. Distribution: Indo-Pacific; subtidal and bathyal depths to about 600 m. Remarks: The new genus Maculauger comprises several species, formerly placed in Hastulopsis, that can be differentiated from similar species in Myurella by the equal prominence of spiral and axial spiral elements, producing a fine cancellate sculpture. Hastulopsis species have very weak sculpture and are found in the Indian Ocean or Japan. Some species of Maculauger, such as M. campbelli, are conchologically very close to Profunditerebra orientalis —a case where only anatomical and molecular characters can ensure correct identification (see Remarks on Profunditerebra). Etymology: The name combines ‘macula’ (Latin: spot, blotch), referring to the commonly maculated subsutural band in the species of the genus, with ‘auger’, the English epithet for Terebridae (‘auger shells’). Gender masculine. Included species: Maculauger alveolatus (Hinds, 1844) 1 n. comb.; M. campbelli (R. D. Burch, 1965) 1 n. comb.; M. castigatus (A. H. Cooke, 1885) 1 n. comb.; M. cinctella (Deshayes, 1859) 3 n. comb.; M. minipulcher (Bozzetti, 2008) 1 n. comb.; M. pseudopertusus (Bratcher & Cernohorsky, 1985) 1 n. comb.; M. sudchiniensis n. sp. Malcolm, Terryn & Fedosov 1 n. sp., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 20, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059."]}

Profunditerebra macclesfieldensis Malcolm, Terryn & Fedosov newspecies (Fig. 12E, F) Type material: Holotype: MNHN IM-2013-61875, 1 lv, 22.4 mm; N Macclesfield Bank; South China Sea, 16 ° 07 'N, 114 ° 23 'E, 161 m (ZhongSha 2015 Stn DW4144). Paratype 1: MNHN IM- 2013-61877, 1 lv, 22.2 mm; type locality. Paratype 2: MNHN IM- 2013-61995, 1 lv, 27.2 mm; type locality. Paratype 3: MNHN IM- 2013-58887, 1 lv, 19.2 mm; New Ireland, 2 ° 26 'S, 149 ° 55 'E, 240–242 m (KAVIENG 2014 Stn DW4485). Paratype 4: 1 dd, 23.1 mm; South China Sea. YT, 200 m. Paratypes 5–7: YT, 3 lv, 20.6–29.5 mm; off Aliguay I., Philippines, dredged at 150–200 m. Zoobank registration: urn:lsid:zoobank.org:act: F1081F38-DB65- 4EDF-B73C-8743D92009A9 Diagnostic nucleotide positions: Table 11. Description: Shell to 30 mm. Protoconch multispiral, 4 translucent amber whorls, small nucleus. Holotype of 12 teleoconch whorls. Teleoconch whorls with straight outline. Subsutural band defined by continuous punctate groove. Axial ribs thin compared with interspaces, gradually becoming more numerous, resulting in about 25 ribs on penultimate whorl of holotype. Ribs only intersected by punctate groove, extended onto flat subsutural band. On last whorl, ribs fade at periphery. No spiral sculpture visible between ribs. Aperture long, creating elongate rounded shape to last whorl. Columella straight, with callus in mature specimens. Colour brown to white with some brown markings. Habitat: Depths of 160– 240 m. Distribution: South China Sea, Philippines and Papua New Guinea. Etymology: Named after the type locality, the Macclesfield Bank in the South China Sea. Remarks: In terms of shell morphology, P. macclesfieldensis n. sp. is most similar to P. anseeuwi (Fig. 12G), Duplicaria duplicata (Fig. 3A) and Punctoterebra teramachii (Fig. 8B). Profunditerebra anseeuwi is the most similar in shell morphology; it has a much shorter columella, creating a cup-shape to the last whorl; its ribs are set at an oblique angle; its colour is white, with broad brown spiral bands between the ribs and dark spots on the subsutural band. Duplicaria duplicata differs in having a paucispiral protoconch of 1.5 whorls, a shorter aperture, and its ribs are broad and closely packed. Punctoterebra teramachii has a much narrower apical angle, the profile of its whorls is more rounded and the subsutural band narrower., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 25, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844

Genus Punctoterebra Bartsch, 1923 (Fig. 8) Type species: Terebra nitida Hinds, 1844; OD. Definition: Primarily defined based on phylogenetic analysis of Modica etal. (2019), comprising species that belong to clade E2 of Modica etal. (2019) and additionally species that show shell morphology, anatomical or genetic characteristics closely comparable with Punctoterebra nitida or any genetically proven member of the clade. Diagnosis. Diagnostic nucleotide combinations are provided separatelyforthefourphylogeneticclustersof the Punctoterebra clade inferred by Modica et al. (2019) in Table 7. Shell: Elongate, small to medium-sized (< 50 mm), with high spire and predominantly axial sculpture. Protoconch paucispiral or multispiral, of varying morphology. Axial ribs strong on spire whorls, sometimes becoming obsolete on last whorl. Spiral sculpture limited to a weak punctuate groove delineating subsutural band, or well-developed and represented by regular striae in interspaces between axial ribs. Siphonal canal demarcated from shell base by distinct concavity; often slightly recurved. Anatomy: Proboscis, venom gland, radula and salivary glands typically present; radula, when present, of flat marginal teeth. Accessory proboscis structure absent. Distribution: Indo-Pacific; intertidal to upper bathyal depths (to 405 m), but typically shallow water. Remarks: With regard to shell morphology, species of this genus form three distinct groups that are consistent with three of the phylogenetic lineages inferred by Modica et al. (2019). Punctoterebra nitida group. Shell small to medium-sized (< 50 mm), polished, with subsutural band demarcated by deep punctures or short grooves; sculpture predominantly of axial ribs. The small species of the group (P. plumbea and P. solangeae) are conchologically very close to some species of Partecosta (see Remarks on the latter genus). Punctoterebra teramachii group. Shells medium-sized (< 45 mm), with dominant sculpture of orthocline axial ribs; spiral sculpture absent; subsutural band clearly defined. Species of this group are conchologically close to Duplicaria, from which they can be differentiated by the punctate subsutural groove and narrower aperture, and by radular teeth that are solid and recurved in Duplicaria, but flat (or absent) in Punctoterebra. Punctoterebra textilis group. Shell medium-sized (< 50 mm), heavily sculptured, with gently convex whorls. Sculpture of arcuate ribs; interspaces bearing distinct regular striae; subsutural band demarcated by punctuate groove of varying strength. Syphonal canal typically recurved, separated from shell base by distinct waist or deep groove. This group is rather heterogeneous in shell morphology, with its core formed by heavily sculptured species of the P. textilis – P. succincta complex. Similar heavily sculptured shells characterize typical Profunditerebra species; however, these can be distinguished by more rounded ribs, a row of nodules on the subsutural band and by straight siphonal canal. The heavily sculptured species of Terebra in the T. amanda group can be readily differentiated by their divided subsutural bands. The inferred membership of P. lineaperlata in the P. textilis group is, however, unexpected and needs further analysis. Included species: Punctoterebra arabella (Thiele, 1925) 2 n. comb.; P. baileyi (Bratcher & Cernohorsky, 1982) 1 n. comb.; P. ballina (Hedley, 1915) 2 n. comb.; P. caliginosa (Deshayes, 1859) 1 n. comb.; P. castaneofusca (Thiele, 1925) 1 n. comb.; P. contracta (E. A. Smith, 1873) 1 n. comb.; P. exiguoides (Schepman, 1913) 3 n. comb.; P. fuscotaeniata (Thiele, 1925) 2 n. comb.; P. illustris (Malcolm & Terryn, 2012) 2 n. comb.; P. isabella (Thiele, 1925) 2 n. comb.; P. japonica (E. A. Smith, 1873) 3 n. comb.; P. lineaperlata (Terryn & Holford, 2008) 1 n. comb.; P. livida (Reeve, 1860) 3 n. comb.; P. longiscata (Deshayes, 1859) 2 n. comb.; P. nitida (Hinds, 1844) 1 n. comb.; P. paucincisa (Bratcher, 1988) 3 n. comb.; P. plumbea (Quoy & Gaimard, 1833) 1 n. comb.; P. polygyrata (Deshayes, 1859) 1 n. comb.; P. rosacea (Pease, 1869) 3 n. comb.; P. roseata (Adams & Reeve, 1850) 1 ∗ n. comb.; P. solangeae (Bozzetti, 2015) 1 n. comb.; P. souleyeti (Deshayes, 1859) 1 ∗ n. comb.; P. succincta (Gmelin, 1791) 1 n. comb.; P. swainsoni (Deshayes, 1859) 2 n. comb.; P. teramachii (Burch, 1965) 1 n. comb.; P. textilis (Hinds, 1844) 1 n. comb.; P. trismacaria (Melvill, 1917) 1 n. comb.; P. turrita (E. A. Smith, 1873) 1 n. comb.; P. turschi (Bratcher, 1981) 2 n. comb., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 15-17, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059.","TERRYN, Y. & HOLFORD, M. 2008. The Terebridae of Vanuatu with a revision of the genus Granuliterebra Oyama, 1961. Visaya, Supplement, 3: 1 - 118."]}

Genus Myurellopsis new genus (Fig. 7 G–I) Type species: Terebra undulata Gray, 1834. Zoobank registration: urn:lsid:zoobank.org:act: EC3F14B2-3595- 482B-9890-399BA78A7631 Definition: Includes all species included in the clade E5B of Modica etal. (2019) and those that show a combination of conchological, anatomical and distribution characteristics closely comparable with Myurellopsis undulata or any genetically proven member of the genus. Diagnosis: Diagnostic nucleotide combinations in Table 10. Shell: Small to medium-sized (< 50 mm), pinkish or orange, often with white subsutural band. Protoconch multispiral. Sculpture of strong, rounded, slightly undulating ribs, polished in appearance. Row of punctations with groove partially cutting ribs forming subsutural band. Interstices between ribs deep, narrow, bearing regular striae, often darkly coloured. Adapical parts of ribs forming one row of rounded nodules, separated from their lower portions by shallow depression or partial groove. Siphonal canal moderately long, tapering, with weakly developed fasciole. Anatomy: Two distinct states—either all typical foregut structures present, or all lacking; in latter case, accessory proboscis structure present. Distribution: Throughout Indo-Pacific; intertidal to upper bathyal depths (to 358 m), typically shallow. Remarks: Species in Myurellopsis n. gen. appear conchologically closest to Myurella species, but can be differentiated by the sharper, more elevated ribs, often with nodes on the subsutural band or top of the whorls. Etymology: The name refers to the close resemblance between its members and species in the genus Myurella. Gender feminine. Included species: Myurellopsis alisi (Aubry, 1999) 3 n. comb.; M. columellaris (Hinds, 1844) 1 n. comb.; M. guphilae (Poppe, Tagaro & Terryn, 2009) 1 n. comb.; M. joserosadoi (Bozzetti, 2001) 1 n. comb.; M. kilburni (R. D. Burch, 1965) 1 n. comb.; M. monicae (Terryn, 2005) 2 n. comb.; M. nathaliae (Drivas & Jay, 1988) 1 n. comb.; M. parkinsoni (Bratcher & Cernohorsky, 1976) 1 n. comb.; M. paucistriata (E. A. Smith, 1873) 1 n. comb.; M. undulata (Gray, 1834) 1 n. comb.; M. vaubani (Aubry, 1999) 3 n. comb. Genera defined based on conchological characters, Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 20-21, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059."]}

Subfamily PELLIFRONIINAE new subfamily (Fig. 2) Type genus: Pellifronia Terryn & Holford, 2008 Zoobank registration: urn:lsid:zoobank.org:act: E827742C-E1B3- 42DD-B5EB-5D7EAA6E2C95 Diagnosis: Diagnostic nucleotide combinations in Table 1. Shell: Small to medium-sized (12–50 mm), with predominant axial sculpture. Early teleoconch whorls often with angled profile, later ones flattened or even slightly concave to convex. Sculpture of numerous close-set, evenly spaced, ribs, or forming raised nodules bordering upper or lower suture. Spiral sculpture absent or of very shallow, wide subsutural depression. Anatomy: Rhynchodeal introvert small; proboscis, radular sac, odontophore, salivary and venom glands well developed. Radula with well-developed membrane, bearing 2 rows of wide marginal teeth of duplex type. Accessory proboscis structure absent. Remarks: This subfamily comprises five described species in two genera, Bathyterebra and Pellifronia, that occur in the Indo-Pacific and Caribbean at bathyal depths. The anatomy of the studied species shows characters that are believed to be ancestral for Terebridae in general, i.e. weakly developed rhynchodeal introvert and radular teeth of duplex type (Castelin et al., 2012)., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 5, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["TERRYN, Y. & HOLFORD, M. 2008. The Terebridae of Vanuatu with a revision of the genus Granuliterebra Oyama, 1961. Visaya, Supplement, 3: 1 - 118.","CASTELIN, M., PUILLANDRE, N., KANTOR, Y. I., MODICA, M. V., TERRYN, Y., CRUAUD, C., BOUCHET, P. & HOLFORD, M. 2012. Macroevolution of venom apparatus innovations in auger snails (Gastropoda; Conoidea; Terebridae). Molecular Phylogenetics and Evolution, 64: 21 - 44."]}

Genus Bathyterebra new genus (Fig. 2 E–I) Type species: Terebra benthalis Dall, 1889. Zoobank registration: urn:lsid:zoobank.org:act: EF1F9047-25AB- 4400-A1D4-C705B757E114 Definition: Includes species with combination of conchological, anatomical and distributional characteristics closely comparable with Bathyterebra benthalis, including all species in clade A2 of Modica etal. (2019) and any species proven to be a member of this clade. Diagnosis: Diagnostic nucleotide combinations in Table 1. Shell: Small, 12–30 mm, elongate, white to tan. Protoconch paucispiral, cyrthoconoid, about 1.5 glossy whorls. Sculpture of fine dense orthocline ribs, interrupted by depression in adapical portion of whorl. Axial ribs weakening towards anterior throughout their height on mature whorls, often forming weak nodules bordering adapical depression, giving distinctly angulated outline to early spire whorls; late teleoconch whorls gently convex. Last adult whorl evenly convex, transitioning to shell base without visible angulation. Siphonal canal stout, slightly recurved at tip, shallowly notched; siphonal fasciole weak or strong. Aperture elongate, wide. Anatomy: Rhynchodeal introvert small; proboscis, radular sac, salivary and venom glands well developed. Accessory proboscis structure absent. Radula with well-developed membrane, bearing 2 rows of wide marinal teeth of duplex type. Distribution: South China Sea, New Caledonia, Caribbean; bathyal depths. Etymology: Name refers to bathyal depths, at which three known species of the genus were sampled. Gender feminine. Remarks: Bathyterebra spp. can be differentiated from members of most other genera of Terebridae by the combination of relatively faint sculpture lacking spiral elements, weakly defined subsutural band and relatively high last adult whorl with convex outline. In shell proportions the genus resembles Oxymeris; however, the latter has more flattened whorls and typically can be easily differentiated by a larger size at maturity. Included species: Bathyterebra benthalis (Dall, 1889) 1 n. comb.; B. coriolisi (Aubry, 1999) 1 n. comb.; B. zhongshaensis Malcolm, Terryn & Fedosov 1 n. sp., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 6-7, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059."]}

Family TEREBRIDAE Mörch, 1852 Synonym: Acusidae Gray, 1853 Shell: Elongate or conical, multiwhorled, with very high spire and relatively small last whorl. Aperture small, with anterior siphonal notch, or very short straight siphonal canal. Anal sinus indistinct. Anatomy: Operculum always present, horny, rounded or leaf-shaped, with terminal nucleus. Head with two very short close-set eye tentacles with eye situated near tip. Rhynchodaeal introvert always present, small to very large. Proboscis, radula, salivary glands, accessory salivary glands, venom gland and accessory proboscis structure present or absent. Radula, when present, composed of marginal teeth only, varying in shape from flat, solid and recurved, semi-enrolled or duplex, to hollow hypodermic., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on page 5, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844

Profunditerebra macclesfieldensis Malcolm, Terryn & Fedosov newspecies (Fig. 12E, F) Type material: Holotype: MNHN IM-2013-61875, 1 lv, 22.4 mm; N Macclesfield Bank; South China Sea, 16 ° 07 'N, 114 ° 23 'E, 161 m (ZhongSha 2015 Stn DW4144). Paratype 1: MNHN IM- 2013-61877, 1 lv, 22.2 mm; type locality. Paratype 2: MNHN IM- 2013-61995, 1 lv, 27.2 mm; type locality. Paratype 3: MNHN IM- 2013-58887, 1 lv, 19.2 mm; New Ireland, 2 ° 26 'S, 149 ° 55 'E, 240–242 m (KAVIENG 2014 Stn DW4485). Paratype 4: 1 dd, 23.1 mm; South China Sea. YT, 200 m. Paratypes 5–7: YT, 3 lv, 20.6–29.5 mm; off Aliguay I., Philippines, dredged at 150–200 m. Zoobank registration: urn:lsid:zoobank.org:act: F1081F38-DB65- 4EDF-B73C-8743D92009A9 Diagnostic nucleotide positions: Table 11. Description: Shell to 30 mm. Protoconch multispiral, 4 translucent amber whorls, small nucleus. Holotype of 12 teleoconch whorls. Teleoconch whorls with straight outline. Subsutural band defined by continuous punctate groove. Axial ribs thin compared with interspaces, gradually becoming more numerous, resulting in about 25 ribs on penultimate whorl of holotype. Ribs only intersected by punctate groove, extended onto flat subsutural band. On last whorl, ribs fade at periphery. No spiral sculpture visible between ribs. Aperture long, creating elongate rounded shape to last whorl. Columella straight, with callus in mature specimens. Colour brown to white with some brown markings. Habitat: Depths of 160– 240 m. Distribution: South China Sea, Philippines and Papua New Guinea. Etymology: Named after the type locality, the Macclesfield Bank in the South China Sea. Remarks: In terms of shell morphology, P. macclesfieldensis n. sp. is most similar to P. anseeuwi (Fig. 12G), Duplicaria duplicata (Fig. 3A) and Punctoterebra teramachii (Fig. 8B). Profunditerebra anseeuwi is the most similar in shell morphology; it has a much shorter columella, creating a cup-shape to the last whorl; its ribs are set at an oblique angle; its colour is white, with broad brown spiral bands between the ribs and dark spots on the subsutural band. Duplicaria duplicata differs in having a paucispiral protoconch of 1.5 whorls, a shorter aperture, and its ribs are broad and closely packed. Punctoterebra teramachii has a much narrower apical angle, the profile of its whorls is more rounded and the subsutural band narrower.

Genus Myurella Hinds, 1845 (Fig. 7 A–D) Type species: Terebra affinis Gray, 1834; SD, Cossmann (1896). Synonyms: Clathroterebra Oyama, 1961 (type species Terebra fortunei Deshayes, 1857; OD); Decorihastula Oyama, 1961 (type species Terebra affinis Gray, 1834; OD); Terenolla Iredale, 1929 (type species Terebra pygmaea Hinds, 1844; OD). Definition: Primarily defined based on phylogenetic analysis of Modica etal. (2019), comprising species in clade E1 of Modica etal. (2019) and additionally those that show shell morphology, anatomical or genetic characteristics closely comparable with Myurella affinis or any genetically proven member of the clade. Diagnosis: Diagnostic nucleotide combinations in Table 6. Shell: very small to medium-sized, with elongated, with high orthoconoid spire; sculpture of arcuate or undulating ribs, fine and narrow to flat and wide. Anatomy: Rhynchodeal introvert very large, proboscis, radula and venom gland absent, salivary glands and accessory proboscis structure present in few species. Distribution: Indo-Pacific; intertidal to upper bathyal depths. Remarks: In regard to shell morphology, four distinct groups can be recognized: Myurella affinis group. Shell medium-sized; axial sculpture of low, wide, flattened ribs; subsutural band with elongate nodules separated by punctations, which are also pronounced in interspaces between axial ribs. Similar species that were earlier classified in Myurella, but do not belong in the T. affinis clade (i.e. M. undulata, M. columellaris and M. kilburni) are here reclassified in the genus Myurellopsis. Typical Myurella species can be recognized by the lower axials and shallower interstices between them. Because of the overall flatter axials, the subsutural band is visually continuous in Myurella; on the contrary, in Myurellopsis, due to the more elevated ribs, the subsutural band is represented by a row of nodules. Myurella pseudofortunei group. Shell elongate, of variable size (17–60 mm), with very high, slender spire, evenly convex whorls, indistinct subsutural band and narrow, clearly arcuate axial ribs; interspaces with regular striae. Species of this group exhibit a very distinctive morphotype, which has little in common with other species here attributed to Myurella. Terebrids with this morphology were previously placed in the genus Clathroterebra; however, in the analysis of Modica etal. (2019), they formed several lineages unrelated to Clathroterebra; morphological differences between these lineages are still to be studied. Myurella pygmaea group. Shell small, c. 10 mm; convex spire profile, straight sides to whorls; very short, tapering siphonal canal. Axial sculpture of numerous fine oblique ribs; subsutural band and other spiral elements absent. The group includes one species, M. pygmaea, formerly classified in the monotypic genus Terenolla. The very small but rather solid, finely ribbed shell is easily recognizable among terebrids; its recognition is further aided by the pattern of flesh-colour background with contrasting dark brown spiral line just below the suture. Myurella amoena group. Shell medium-sized; sculptured with fine to strong ribs, subsutural band defined by line of punctuations, spiral sculpture of narrow grooves or cords. The species of this group form a well-supported clade and exhibit a shallow-water morphotype that was formerly classified in Hastulopsis. Similar deeper-water species are found in Maculauger n. gen., most of which can be recognized by having a row of large spots on the periphery. The other similar species remain within Hastulopsis, comprising a number of Indian Ocean and Japanese endemic species. Species of the M. amoena group have strong but narrow ribs that are notably stronger than spiral elements. In Myurellopsis the ribs are about as strong as in the M. amoena group, but are wider and rounded, with interstices narrower than the ribs. Both Hastulopsis and Maculauger comprise species with axial and spiral elements of comparable strength, resulting in a rather cancellate pattern, and a continuous subsutural band. Included species: Myurella affinis (Gray, 1834) 1; M. amoena (Deshayes, 1859) 1; M. andamanica (Melvill & Sykes, 1898) 1 ∗; M. bilineata (Sprague, 2004) 1; M. brunneobandata (Malcolm & Terryn, 2012) 1; M. burchi (Bratcher & Cernohorsky, 1982) 1; M. conspersa (Hinds, 1844) 1; M. dedonderi (Terryn, 2003) 1; M. eburnea (Hinds, 1844) 1; M. flavofasciata (Pilsbry, 1921) 1; M. fortunei (Deshayes, 1857) 1; M. mactanensis (Bratcher & Cernohorsky, 1982) 2; M. mindanaoensis (Aubry, 2008) 1; M. multistriata (Schepman, 1913) 1; M. nebulosa (G. B. Sowerby I, 1825) 1; M. pertusa (Born, 1778) 1; M. picardali (Aubry, 2011) 2; M. pseudofortunei (Aubry, 2008) 1; M. pygmaea (Hinds, 1844) 1 n. comb.; M. russoi (Aubry, 1991) 2; M. suduirauti (Terryn & Conde, 2004) 2; M. wellsilviae (Aubry, 1994) 1., Published as part of Fedosov, Alexander E, Malcolm, Gavin, Terryn, Yves, Gorson, Juliette, Modica, Maria Vittoria, Holford, Mandë & Puillandre, Nicolas, 2019, Phylogenetic classification of the family Terebridae (Neogastropoda: Conoidea), pp. 359-388 in Journal of Molluscan Studies The Malacological Society of London 85 (4) on pages 14-15, DOI: 10.1093/mollus/eyz004, http://zenodo.org/record/4469844, {"references":["COSSMANN, M. 1896. Essais de paleoconchologie comparee, Vol. 2. Author and Comptoir Geologique, Paris.","OYAMA, K. 1961. On some new facts of the taxonomy of Terebridae. Venus, 21: 176 - 189.","MODICA, M. V., GORSON, J., FEDOSOV, A. E., MALCOLM, G., TERRYN, Y., PUILLANDRE, N. & HOLFORD, M. 2019. Macroevolutionary analyses suggest environmental factors, not venom apparatus, play key role in Terebridae marine snail diversification. Systematic Biology. DOI: https: // doi. org / 10.1093 / sysbio / syz 059."]}