taxonID	type	description	language	source
03908790FF93FFF3B024779DD149B431.taxon	description	Although the specimen had been positioned for obtaining sagittal sections, the proboscis was diverted and sectioned nearly in frontal plane (Fig. 6 A, D). Proboscis moderately long, broad, conical, strongly contracted, so that its wall forms several telescopic folds (Fig. 6 A, C, D). Mouth opening into medium long buccal tube, spanning around 1 / 3 of proboscis length and leading into broad buccal cavity. Buccal cavity subdivided into two chambers by the large circular fold of its wall (Fig. 6 D, arrow); both chambers lined with tall, densely ciliated epithelium, epithelial cells with large basal or central nuclei. Wall of the posterior chamber bearing very tall longitudinal folds (Fig. 6 D – F). Wide anterior oesophagus leaving buccal cavity dorsally, forming rather long loop within proboscis, leaving proboscis and narrowing before passing through the nerve ring and then widening before transition to the posterior oesophagus (Fig. 6 D). Distinct valve-like structure, identifiable as a valve of Leiblein although lacking a ciliary cone, delimiting an anterior and a posterior oesophagus (Fig. 6 G, vl). Anterior oesophagus empty in sectioned specimen, posterior oesophagus filled with food material (Fig. 6 D, G) in which, however, no detectable remnants were found. After the oesophageal valve, tall and lightly stained epithelium of anterior oesophagus changing to low epithelium formed by small cubic cells. Anterior oesophagus lined with thin layer of muscular fibres, which become notably stronger in posterior oesophagus (Fig. 6 G). Thin and strongly convoluted salivary ducts adjoining anterior oesophagus in front of passage through nerve ring and passing forward being embedded into its muscular wall, leaving oesophageal wall closer to the proboscis tip and entering buccal mass. Paired odontophore cartilages well developed, not fusing with strong lateral odontophore protractor muscles attached to odontophore ventrally (Fig. 6 D). Strong and massive odontophore retractor adjoining the columellar muscle shortly behind the nerve ring. Radular diverticulum opening into buccal cavity ventrally in posterior position, behind thick circular fold of the buccal cavity BI, Bayesian inference; ML, maximum likelihood; n / a, not applicable. * Taxa represented by single specimen. wall, forming massive ‘ lip’, longer dorsally (Fig. 6 D, E, bl). After entering buccal cavity wall, distal salivary ducts passing inside that lip laterally on both sides of radular diverticulum opening (Fig. 6 D, E), and opening in the buccal cavity ventrally, underneath the semi-circular fold (Fig. 6 F). Epiproboscis absent. Salivary gland large, unpaired, situated to the left of posterior oesophagus and dorsally to the nerve ring, histologically represented by a compactly packed, strongly convoluted tube with one layer of tall highly vacuolar, obviously glandular, cells and thin outer connective tissue layer (Fig. 6 A, B). Condylomitra tuberosa (PANGLAO 2004, Stn B 5) Proboscis moderately long, leaf-shaped, anteriorly flattened dorso-ventrally (Fig. 7 A – H) and cylindrical at its base (Fig. 7 I). On transversal sections, proboscis with two lateral lobes at mid-length and ventral part with rounded axial keel. Proboscis wall with thick cuticle and nearly cubic epithelium, with large basal nuclei; epithelium underlined by a layer of circular muscle fibres (Fig. 7 C, E, H). Lateral lobes occupied by complex interlacement of dorso-ventral, oblique and longitudinal muscular fibres; two blood vessels (Fig. 7 F, bv) running along margins of proboscis (Fig. 7 B, D, F, G). Central part of proboscis occupied with oesophagus, anterior buccal mass and epiproboscis complex forming a single stem. Mouth opening anteriorly on the dorsal side of the proboscis in short dorsal groove; mouth surrounded by small peristome rim (Fig. 7 A). Compact buccal mass situated shortly after mouth, furnished with thick muscular walls and triangular lumen; radular diverticulum opening ventrally into buccal cavity (Fig. 7 B, C). Anterior oesophagus quadrangular on transversal sections, lying dorsally in the anterior part of the proboscis, attached to proboscis wall by thin muscular fibres (Fig. 7 D – H). Dorsal and ventral walls of oesophagus very thin, lateral walls with two large longitudinal folds jutting out into the lumen of oesophagus. Lateral folds of oesophagus disappearing at mid-proboscis height and thick layer of muscular fibres evenly covering it (Fig. 7 I). Radular diverticulum rather short; odontophore formed by two small cartilages (Fig. 7 D, E), not fused anteriorly. Odontophore retractor very strong, lying in a muscular sheath in its anterior portion (Fig. 7 F – H), and dividing into two symmetrical branches at mid-proboscis length (Fig. 7 I). Epiproboscis short, formed by densely packed longitudinal muscle fibres, with no epithelium. Epiproboscis lying ventrally from radular diverticulum, enclosed inside its own muscular sheath, which is attached to the radular diverticulum along most of its length (Fig. 7 D – H). Both epiproboscis and its sheath originating from the ventral part of the radular retractor sheath (Fig. 7 G, H). Epiproboscis sheath opening ventrally on the anterior border of the buccal mass. Salivary ducts passing forward along the anterior oesophagus, embedded into lateral folds of oesophagus wall (Fig. 7 D – I); salivary gland itself not sectioned. Salivary ducts shifting ventrally in the walls of the buccal cavity adjoining the epiproboscis sheath. More distally, salivary ducts running backwards along the epiproboscis sheath to its proximal part, where they enter the epiproboscis and run through its entire length to open at its tip (Fig. 7 E).	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FF99FFF7B0257324D19EB09E.taxon	description	Decisions on the generic assignment of individual species are based on: 1 molecular evidence and radula morphology or molecular data only, 2 shell characters and radula morphology and 3 solely shell characters. In ‘ Species included’ sections, type species of recognized genera are shown in bold.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FF98FFF7B09874CAD63DB446.taxon	materials_examined	Type species: Charitodoron euphrosyne Tomlin, 1932 (junior subjective synonym of Columbella barbara Thiele, 1925); OD. Species included: Charitodoron agulhasensis (Thiele, 1925) 3, C. alcyone Lussi, 20093, C. barbara (Thiele, 1925) 3, C. bathybius (Barnard, 1959) 1; C. rosadoi Kilburn, 19953, C. thalia Tomlin, 19323, C. veneris (Barnard, 1964) 2.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FF98FFF7B3AA76DAD0D2B298.taxon	description	(FIG. 8) Diagnosis: Shell small (13 – 35 mm), fusiform, white or pale, sometimes with dark bands, or covered with brown periostracum. Protoconch white, bulbous, of about 1.5 smooth glossy whorls and with large nucleus. Suture distinct, impressed. Spire tall, orthoconoid; spire whorls evenly convex, smooth or sculptured with weak spiral grooves or cords. Sometimes distinct cancellate sculpture developed. Siphonal canal moderately long, robust or tapering, sometimes slightly recurved at its tip; siphonal notch shallow. Aperture wide, elongate; outer aperture lip smooth, evenly convex. Inner lip with no columellar folds. Head – foot pale, with sometimes nebulose dark bands. Tentacles moderately long with eyes situated laterally at about mid-length. Siphon long. Osphradium as long as gill, attaining about 2 / 3 of the length of the latter. Proboscis well developed, without distinct proboscis rim; epiproboscis absent. Radula with narrow rachidian, bearing five subequal cusps, and wide, multicuspidate laterals, with cusps being more dense and stout proximally and becoming more slender towards radula margins; lateral 1 / 5 portion of teeth smooth. Mid-oesophagus muscular; mid-gut gland absent. Salivary gland single with paired ducts opening in buccal tube ventrally and anterior to radular diverticulum. Remarks: The newly proposed family Charitodoronidae is distinctive in many ways. The shell bears no columellar folds and rather resembles a buccinoid shell; for this reason, many Charitodoron species were originally erroneously placed in Columbella. The bulbous protoconch is unusual for Mitridae and suggests a non-planktotrophic development, consistent with the narrow distribution range. Despite Charitodoron being close to mitrids in radula morphology, it does not possess the key morphological innovation of the Mitridae, the epiproboscis. The obvious morphological distinctiveness of Charitodoron and its position in the molecular tree as a sister group to all other Mitridae justify its recognition in a separate family.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FF9BFFF4B21B7779D771B2CB.taxon	description	(FIGS 9, 10 A) Type species: Voluta mitra Linnaeus, 1758, by tautonymy. Synonym: Tiarella Swainson, 1840. Type species: Voluta papalis Linnaeus, 1758; SD (Gray, 1847) Papalaria Dall, 1915. Type species: Voluta episcopalis Linnaeus, 1758 [= Mitra mitra (Linnaeus, 1758)]; SD, Coan (1966: 131). Diagnosis: Shell medium sized to large (50 – 170 mm), thick-walled, fusiform or elongate-fusiform, with truncated base and colour pattern of multiple rounded or squarish red spots or bands on white background. Protoconch multispiral, of about three smooth slightly convex whorls. Suture deeply impressed. Teleoconch whorls gently convex or angulated adapically, bearing a row of short spiny bumps at shoulder. Rest of whorl surface sculptured with weak, evenly spaced, spiral grooves, well pronounced on early spire whorls and often vanishing later. Siphonal canal very short with robust siphonal fasciole delineated from shell base by deep depression. Aperture rather wide, with deep siphonal notch; outer apertural lip nearly straight adapically and strongly convex in its lower portion, bearing numerous fine denticles throughout its length or limited to its abapical margin. Inner lip reflected, with three to four subequal oblique columellar folds. Radula with rather narrow rachidian bearing six subequal cusps, and very wide laterals bearing cusps throughout their width (Fig. 10 A). Cusps rather strong near rachidian and becoming progressively weaker laterally. Distribution: Indo-Pacific, shallow intertidal and subtidal to upper bathyal depths, sand and rocky bottom. Species included: Mitra mitra (Linnaeus, 1758) 1, Mitra deprofundis Turner, 20013, Mitra papalis (Linnaeus, 1758) 1, Mitra stictica (Link, 1807) 3. Remarks: A critical reassessment of the contents of the genus Mitra is one of the main outcomes of the present study, and our phylogenetic analysis demonstrates that the name Mitra may be confidently applied to a handful of species only. The redefined genus Mitra is morphologically distinctive because of the conservative shell shape with rather high spire and low aperture, presence of denticles on the outer aperture lip and a recognizable colour pattern. At the same time, its radular morphology is overall the same as in many other mitrid clades and adds little to the diagnosis of the genus. Quasimitra cardinalis and Q. puncticulata resemble species of Mitra. However, Q. cardinalis has a notably wider shell with lower spire, and Q. puncticulata lacks the characteristic colour pattern of Mitra and bears no denticle on the outer aperture lip. Nebularia incompta also resembles Mitra in shell proportions and the presence of denticles on the margin of the outer lip, but has a stronger sculpture and its siphonal canal is not clearly delineated from the shell base; besides, it has a typical Nebularia - type colour pattern instead of a Mitra - type. Some Strigatella species, especially S. imperialis, also closely resemble Mitra in the morphology of the abapical part of the shell (aperture and siphonal canal), but they have a notably lower spire and lack the Mitra - type colour pattern as well.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FF9BFFC8B0967482D1A0B137.taxon	description	(FIGS 10 B, C, 11) Type species: Calcimitra kingtsio Huang, 2011; OD. Diagnosis: Shell medium sized to large (35 – 125 mm), elongate-fusiform, with high orthoconoid or slightly acuminate spire and slender, tapering siphonal canal. Protoconch narrowly conical, multispiral, of about three smooth convex whorls. Suture deeply impressed or canaliculate. Spire whorls flattened to evenly convex, smooth or sculptured with weak and irregular, or distinct, evenly set cords. Interspaces between spiral cords sometimes ornamented with fine collabral growth lines or dense riblets sometimes forming fine cancellate sculpture pattern. Siphonal canal long, tapering, with shallow or indistinct siphonal notch. Aperture elongate, its outer lip smooth, evenly convex. Inner aperture lip often calloused, sometimes reflected, bearing three to four fine columellar folds. Shell white, pale to orange, sometimes bearing pattern of reddish or brownish axial bands; fresh specimens often covered with brown periostracum. Radula with rather narrow rachidian, bearing six to eight subequal cusps; laterals wide, bearing 12 – 20 pointed cusps (Fig. 10 B, C), two proximal cusps rather short, weaker than the next five ones that are strongest; marginal five to eight cusps gradually diminishing. Distribution: Indo-Pacific, bathyal depths, mud, sometimes on sunken wood. Species included: Calcimitra arnoldeyasi (Poppe, Tagaro & Salisbury, 2010) 3 comb. nov., C. brinkae (Salisbury & Kilburn, 1996) 3 comb. nov., C. brunetta (Chino & Herrmann, 2015) 3 comb. nov., C. chiangfucius (Huang & Salisbury, 2017) 2 comb. nov., C. christinae (Poppe, 2008) 1 comb. nov., C. chuoi (Huang & Salisbury, 2017) 3 comb. nov., C. glaphyria (Turner, 2001) 1 comb. nov., C. hilli (Cernohorsky, 1976) 3 comb. nov., C. invicta (Huang & Salisbury, 2017) 1 comb. nov., C. kingtsio Huang, 20113, C. labecula (Herrmann & Dekkers, 2009) 1 comb. nov., C. lussii (Turner & Salisbury, 2007) 3 comb. nov., C. marrowi (Turner, 2001) 3 comb. nov., C. meyeriana (Salisbury, 1992) 3 comb. nov., C. morchii (A. Adams, 1855) 2 comb. nov., C. philosopha (Huang & Salisbury, 2017) 2 comb. nov., C. poppei (Guillot de Suduiraut, 2000) 3 comb. nov., C. salva (Turner, 2001) 3 comb. nov., C. subflava (Kuroda & Habe, 1971) 3 comb. nov., C. taiwanbale (Huang & Salisbury, 2017) 3 comb. nov., C. triplicata (Martens, 1904) 1 comb. nov., C. verweyi (Knudsen, 1970) 3 comb. nov. Remarks: The five species included in clade 13 of the COI - based tree show considerable variation in shell morphology, both intra- and interspecifically. Six sequenced specimens of C. glaphyria display a range of sculpture types, from smooth shells resembling C. kingtsio to distinct, cancellated ones, and differently sculptured specimens can be found in the same general locality. While typical Calcimitra are medium-sized, thin-walled shells, our phylogenetic analysis placed ‘ Mitra ’ invicta in the same phylogenetic clade. The latter is a recently described species representative of a group of large deep-water mitrids (here referred to as the C. morchii complex) that possess heavy, strongly sculptured shells (Huang & Salisbury, 2017). The close affinity of these conchologically diverse species is consistent with a generally conservative radular morphology in C. kingtsio (Huang, 2011), C. morchii, C. chiangfucius, C. philosopha (Huang & Salisbury, 2017), C. glaphyria and C. invicta (herein). The intraspecific variation in sculpture pattern demonstrated by C. glaphyria challenges the status of the species in the C. morchii complex accepted by Huang & Salisbury (2017). The distinction of C. chiangfucius, C. taiwanbale and C. invicta is based on subtle differences in shell morphology; they may prove to represent variants of a single species, and more comprehensive sampling is necessary to clarify species boundaries. Magnamitra sandrogorii strongly resembles some of the larger species of Calcimitra, and the distinction of these two genera based on shell characters is problematic. The spiral sculpture in M. sandrogorii is even stronger than in species of the C. morchii complex, and the main distinction between the two genera is in the radula. In Magnamitra, the rachidian has a very strong unpaired central cusp that notably exceeds lateral cusps, whereas all studied species of Calcimitra possess a rachidian with equal or subequal cusps. Furthermore, rachidian and laterals have comparable width in M. sandrogorii, whereas in species of Calcimitra the width of the rachidian does not exceed half of the laterals width. Species of Profundimitra gen. nov. superficially resemble Calcimitra, but the latter genus can be recognized by the generally larger shells with proportionally longer siphonal canal and often calloused inner aperture lip. The radulae of species of Calcimitra and Profundimitra gen. nov. differ greatly, the latter having rachidian and marginals of about equal width. Species of Eumitra are also similar to Calcimitra in shell proportions and sculpture; however, Eumitra can be distinguished by its columellar folds, which are reduced to not more than two very weak folds, while in Calcimitra there are four (very rarely three) distinct ones. Some species of Calcimitra, in particular C. triplicata, resemblance the Miocene fossil genus Clifdenia, both in whorl outline with evenly rounded inflated shoulders, and in shape and orientation of columellar plaits (A. Beu, personal communication). Based on this resemblance, we tentatively allocate Clifdenia to the subfamily Mitrinae.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFA7FFC9B22C7783D051B4E8.taxon	description	(FIG. 12) Type species: † Mitra alokiza Tenison-Woods 1879; SD (Ludbrook, 1958: 71). Diagnosis: Shell small to medium sized (15 – 58 mm), fusiform, thin walled, with high orthoconoid spire and slender siphonal canal. Protoconch very low, cyrtoconoid, paucispiral, with glossy whorls and rather small nucleus in all known species. Suture distinct, impressed. Early teleoconch whorls subcylindrical, later evenly convex, sometimes resulting in a slightly acuminate spire profile. Shell smooth, sometimes with very weak axial folds on first teleoconch whorl or fine grooves on shell base and siphonal canal. Siphonal canal, slender, tapering. Siphonal notch absent. Aperture elongate, outer lip evenly convex, smooth. Inner aperture lip either smooth, bearing one very weak or sometimes two distinct, but weak, columellar folds. Shell uniformly pale to light brown. Radula with very small, narrow rachidian, bearing five short robust cusps, only slightly extending beyond preceding tooth margin, but forming ridges, notably elevated above surface of rachidian. Laterals wide with eight to ten strong, pointed subequal cusps. Distribution: New Caledonia, Coral Sea and the Philippines at bathyal depths, and a further undescribed species referable to Eumitra from Indonesia. Fossil species widely distributed in the Miocene of Europe and in the Australia – New Zealand region. Recent species included: Eumitra apheles Lozouet, 19912, E. caledonica Lozouet, 19913, E. imbricata Lozouet, 19913, E. richeri Lozouet, 19913. ‘ Eumitra ’ suduirauti Bozzetti, 19973. Remarks: Eumitra was not included in our molecular analysis, and our understanding of this genus is largely based on published data (Lozouet, 1991), although additional material from the West Pacific has been examined. The placement of Eumitra in the newly circumscribed Mitrinae is thus tentative and based on the plesiomorphic radula of Eumitra and the presence of a well-developed epiproboscis (YK, unpublished data). Based on shell characters, Eumitra resembles Calcimitra and Profundimitra gen. nov.; however, in the latter two genera species commonly have three or more strong columellar folds. Besides, all studied species of Profundimitra gen. nov. are characterized by a radula with a rachidian as broad as the laterals. Species of Eumitra lacking shell sculpture somewhat resemble the costellariid genera Latiromitra and Costapex (Fedosov et al., 2017), which however, can be distinguished by the presence of an operculum, although reduced, and a longer siphonal canal (Latiromitra) or three or more distinct columellar folds (Costapex). Moreover, in both Latiromitra and Costapex, the early teleoconch whorls are sculptured with strong axial ribs that are absent in Eumitra. ‘ Eumitra ’ suduirauti Bozzetti, 1997 (Fig. 12 E) shows some features (such as a wide aperture, especially wide in its anterior portion, a distinct subsutural ramp, pronounced on late teleoconch whorls, as well, as on early spire whorls) that are uncommon not only in the genus Eumitra, but in the family Mitridae in general. However, as no data on radula morphology or DNA of this odd species are available, we maintain it in the genus Eumitra pending a better placement to be determined in future. All known species of Eumitra share a paucispiral protoconch that distinguishes them from most other mitrid genera. Nevertheless, given the wide distribution of Eumitra in the fossil record, we cannot rule out the possibility of multiple transitions to non-planktotrophic development in the evolutionary history of the genus, and thus some members of Eumitra might have retained planktotrophic development, and thus a multispiral protoconch. GENUS NEOTIARA FEDOSOV, HERRMANN, KANTOR &	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFA6FFC9B26D7281D66DB5DA.taxon	description	(FIGS 10 D, E, 13) Type species: Mitra lens W. Wood, 1828; OD, herein. Diagnosis: Shell small to large (20 – 75 mm), solid, coarsely sculptured, broadly fusiform to about turriform. Protoconch multispiral, rather wide, cyrtoconoid, of about 3.5 smooth, convex whorls. Spire high; suture deeply impressed. Teleoconch whorls flattened to strongly convex or distinctly shouldered. Sculpture of slightly prosocline, wide and broadly spaced axial folds, or rather dense ribs; axials overridden by spiral cords which are typically wide and flattened, but may be narrow and elevated. Rounded or spirally elongated beads sometimes present at the intersection of spiral and axial sculptural elements. Siphonal canal rather short, stout, with weak or no fasciole. Siphonal notch shallow or indistinct. Outer aperture lip evenly convex, or its adapical half about straight, and abapical half strongly convex. Inner aperture lip with three to four strong columellar folds, adapicalmost strongest. Shell dull, light to dark brown, sometimes with well-developed periostracum. Radula with broad laterals, bearing robust, short and blunt cusps on their inner portion, turning gradually to fine serration on outer half (Fig. 10 D, E). Rachidian with robust, very short and deeply rooted cusps, their tips merely reaching anterior margin of adjacent tooth base. Species included: Neotiara crenata (Broderip, 1836) 3 comb. nov., N. fultoni (E. A. Smith, 1892) 3 comb. nov., N. gausapata (Reeve, 1845) 3 comb. nov., N. inca (d’Orbigny, 1841) 3 comb. nov., N. lens (Wood, 1828) 1 comb. nov., N. muricata (Broderip, 1836) 3 comb. nov., N. nodulosa (Gmelin, 1791) 1 comb. nov., N. pallida (Nowell-Usticke, 1959) 3 comb. nov., N. rupicola (Reeve, 1844) 3 comb. nov., N. sphoni (Shasky & Campbell, 1964) 3 comb. nov. Etymology: The name refers to the now disused name Tiara Swainson, 1831, one of the genera first established in what later became the family Costellariidae. Remarks: According to Cernohorsky (1976: 454), the type of M. lens, which used to be in Mawe’s collection, can no longer be traced. In order to fix the identity of the species and of the genus Neotiara gen. nov., we here designate our sequenced specimen BAU 00800 from Panama City, Panama, measuring 40.0 × 15.5 mm (Fig. 13 A), as neotype of M. lens, MNHN IM- 2000 - 33145. As circumscribed here, the genus Neotiara can be recognized among New World Mitridae primarily by the well developed, typically coarse sculpture with distinct axial elements. Other Caribbean and Panamic Mitridae of the genera Isara, Subcancilla, Probata and Atrimitra lack axial sculpture, and the spiral elements are either lacking as well, or represented by fine cords. As indicated by our phylogenetic analysis, the Panamic Neotiara lens and N. aff. inca, and the Caribbean N. nodulosa are closely related and Neotiara is at present the only mitrid genus with both Pacific and Atlantic species. Although only three species are included in our phylogenetic analysis, we classify all heavily sculptured American mitrids in Neotiara gen. nov.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFA6FFCFB08D738ED268B5E8.taxon	description	(FIGS 10 F, 14 A – D) Type species: Mitra carinata Swainson, 1824; SD, Wenz (1943: 1292). Diagnosis: Shell small (25 – 35 mm), elongate to fusiform, usually light brown to olive. Protoconch narrowly conical of about 2.5 – 3 smooth convex whorls. Suture distinct, impressed. Early teleoconch whorls evenly convex, with weak cancellate sculpture, later whorls typically distinctly carinated. Sculpture of weak grooves on late spire whorls, sometimes missing on last adult whorl. Last adult whorl high, with shell base gradually extended into tapering siphonal canal, typically sculptured with wide and low rounded cords. Siphonal notch weakly or moderately shallow. Aperture elongate, narrow, its outer lip smooth, gently convex or flattened. Inner lip with three fine, oblique columellar folds. Rachidian with five, equally developed, pointed, median cusps, flanked by two notably smaller marginal cusps (Fig. 10 F). Lateral teeth multicuspidate, third proximal cusp strongest, and subsequent cusps gradually diminishing; distal third of tooth smooth, with no cusps. Distribution: West Africa, continental shelf, on mud. Species included: Ziba carinata (Swainson, 1824) 1, Z. gambiana (Dohrn, 1861) 3, Z. ogoouensis Biraghi, 19843. Remarks: The present study has led to a drastic reconsideration of the position and contents of the genus Ziba. Based on the results of our phylogenetic analysis, Ziba is now classified in the subfamily Mitrinae and its contents exclude all the Indo-Pacific and West American species hitherto referred to Ziba by Cernohorsky (1991). An array of Indo-Pacific Ziba species are here transferred to Imbricaria, whereas the Panamic Ziba attenuata is reclassified in Subcancilla, neither of these genera showing an affinity to the true Ziba, which ends up being confined to the tropical eastern Atlantic; to our knowledge, there is no Indo-Pacific species referable to this genus. The radula of Z. carinata shows a plesiomorphic morphology which is commonly found in Mitridae and, although supporting placement of Ziba in Mitrinae, adds little to understanding its relationships within the subfamily. The moderately supported lineage Z. carinata – Episcomitra inferred in the 3 - mit tree (though absent in other analyses) may reflect an affinity that makes sense from the biogeographical point of view; however, further studies are needed to determine the relationships of Ziba.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFA0FFCFB3EF73D1D735B6B9.taxon	description	(FIGS 10 G, H, 14 E – H) Type species: Mitra zonata Marryat, 1818; M. Diagnosis: Shell small to large (15 – 110 mm), fusiform or elongate-fusiform, smooth, with well-developed periostracum. Protoconch large, paucispiral, cyrtoconoid, of about two smooth glossy convex whorls. Spire tall or rather low; suture impressed. Spire whorls evenly convex in outline, sculpture represented by fine, dense, collabral lines. Last adult whorl typically evenly convex with rather short siphonal canal bearing a distinct fasciole, separated from shell base by deep groove. Siphonal notch deep or rather shallow. Outer aperture lip evenly convex, or convex in its adapical portion and straight throughout most of its length. Inner aperture lip with four, fine, subequal columellar folds. Shell covered with periostracum, light brown to dark brown or olive-green, uniform or with multiple dark freckles, or (typically) adapical 2 / 3 of spire whorls slightly lighter than their abapical portions. Inside of aperture white or orange. Rachidian with six to eight pointed cusps, increasing in length towards centre; laterals wide, bearing rather long and pointed cusps throughout their length (Fig. 10 G, H).	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFA3FFCCB24C7531D7B2B727.taxon	description	(FIGS 10 I – M, 15) Type species: Mitra sanguinolenta Lamarck, 1811; OD, herein. Diagnosis: Shell small to large (20 – 90 mm), solid, broadly fusiform, vividly coloured. Protoconch pointed, narrowly conical, of about 3.5 slightly convex, smooth whorls. Suture deeply impressed. Shell proportions variable, spire moderately high to rather low, with ortho- or cyrtoconoid profile. Spire whorls evenly convex to subcylindrical, sometimes distinctly shouldered. Sculpture of very weak and low, flat or rounded, cords separated by shallow depressions or fine grooves. Subsutural area smooth or with one row of short spiny warts or tubercles, giving whorl a somewhat shouldered appearance. Siphonal canal short, with well-developed fasciole usually separated from shell base by narrow concave waist. Siphonal notch deep. Aperture elongate, wide or moderately wide. Outer lip smooth, inner lip with four distinct columellar folds. Rachidian narrow (1 / 3 of laterals width) to rather wide (exceeding half of laterals width), with five to eight subequal cusps (Fig. 10 I – M). Laterals rather wide, with ten to 14 widely set and triangular, or dense and rather long, pointed cusps distributed throughout tooth width, proximal one to two cusps notably smaller than succeeding ones. Distribution: Indo-Pacific, upper subtidal depths, mostly on sand. Species included: Quasimitra albocarnea (Bozzetti, 2016) 3 comb. nov., Q. barbieri (Poppe & Tagaro, 2006) 3 comb. nov., Q. bovei (Kiener, 1838) 3 comb. nov., Q. brettinghami (E. A. Smith, 1906) 3 comb. nov., Q. cardinalis (Gmelin, 1791) 1 comb. nov., Q. floccata (Reeve, 1844) 3 comb. nov., Q. fulgurita (Reeve, 1844) 3 comb. nov., Q. lacunosa (Reeve, 1844) 3 comb. nov., Q. lamarckii (Deshayes, 1832) 3 comb. nov., Q. latruncularia (Reeve, 1844) 3 comb. nov., Q. leforti (Turner, 2007) 3 comb. nov., Q. manuellae (Cossignani & Cossignani, 2006) 3 comb. nov., Q. marmorea (Turner, 2007) 3 comb. nov., Q. nubila (Gmelin, 1791) 3 comb. nov., Q. nympha (Reeve, 1845) 3 comb. nov., Q. propinqua (A. Adams, 1853) 3 comb. nov., Q. pseudobovei (T. Cossignani & V. Cossignani, 2005) 3 comb. nov., Q. puncticulata (Lamarck, 1811) 1 comb. nov., Q. punctostriata (A. Adams, 1855) 3 comb. nov., Q. raphaeli (Drivas & Jay, 1990) 3 comb. nov., Q. rinaldii (Turner, 1993) 3 comb. nov., Q. roselineae (Martin & Salisbury, 2013) 3 comb. nov., Q. rossiae (Reeve, 1844) 3 comb. nov., Q. sanguinolenta (Lamarck, 1811) 1 comb. nov., Q. sarmientoi (Poppe, 2008) 3 comb. nov., Q. solida (Reeve, 1844) 3 comb. nov., Q. sophiae (Crosse, 1862) 1 comb. nov., Q. stossieri (Herrmann, 2016) 3 comb. nov., Q. structilis (Herrmann & Salisbury, 2016) 3 comb. nov., Q. variabilis (Reeve, 1844) 3 comb. nov. Etymology: Latin quasi, meaning appearing like, which refers to the overall shell resemblance with species of the genus Mitra. Gender feminine. Remarks: The genus Quasimitra gen. nov. is defined primarily based on molecular characters, and there are no apomorphies or clear diagnostic morphological features that support its recognition. Shell proportions and sculpture vary extensively, with two extremes represented by Q. cardinalis and Q. sophiae, the close affinity of which is evidenced by the molecular data. The radulae of studied species generally retain a plesiomorphic morphology and also do not present any notable feature. We tentatively assign Mitra solida Reeve, 1844 to Quasimitra, because of its characteristic sculpture and colour pattern, although varying notably in shell proportions. Cernohorsky treated Vicimitra prosphora Iredale, 1929 as a junior synonym of M. solida, despite V. prosphora has a smooth, uniformly pale shell, and only superficially resembles M. solida and the other sequenced species here assigned to Quasimitra. At present, we treat V. prosphora as a separate species in a monotypic genus Vicimitra, but the name Vicimitra may prove to be a senior synonym of Quasimitra. Some species of Strigatella, like S. luctuosa or S. aurantia, bear a remarkable resemblance to Quasimitra in shell outline and can only be distinguished by the generally more flattened spire whorls; at the same time, there is a consistent difference in radular morphology between the two groups: the laterals bear cusps throughout their width in Quasimitra, whereas the distal third of laterals is devoid of cusps in the species of Strigatella mentioned.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFA2FFCDB3E175E3D6F7B729.taxon	description	(FIGS 16 A – E, 17 A – D) Type species: Voluta filaris Linnaeus, 1771; OD. Diagnosis: Shell small to medium sized (12 – 65 mm), fusiform to elongate-fusiform, with high spire and rather short aperture. Protoconch pointed, narrowly conical, of 2.5 – 3.5 smooth, slightly convex whorls to large, cyrtoconoid, of nearly five smooth whorls. Suture distinct, impressed. Teleoconch whorls evenly convex in outline, with dominating spiral sculpture represented by regular, elevated, widely interspaced cords that are fine or (usually) strong, sometimes slightly gemmate. Interspaces between cords further sculptured by fine spiral grooves intersected by axial grooves of equal strength, resulting in fine, cancellated background pattern. Siphonal canal moderately long, tapering or robust, with well-developed fasciole, delimited from shell base by distinct narrow waist, or extended concave stretch of shell base. Aperture rather short, elongate, with distinct siphonal notch. Outer lip gently convex, inner lip with three to four fine columellar folds, subequal or adapical strongest. Shell usually white or light with contrasting spiral cords, either uniformly dark brown or dashed-brown. Rachidian from narrow (about 40 % of laterals width) with 11 short, blunt cusps to almost as wide as laterals, arched, with 20 + pointed cusps. Laterals broad, bearing 20 – 35 equal, pointed cusps set on narrow, straight or arched, plate (Fig. 17 A – D). Distribution: Indo-Pacific, perhaps West Africa (D. hebes), shallow subtidal to bathyal depths, on sand. Species included: Domiporta aglais 3 (Li & Zhang, 2005), D. carnicolor (Reeve, 1844) 1, D. circula (Kiener, 1838) 1 comb. nov., D. daidaleosa (Li & Li, 2005) 3 comb. nov., D. filaris (Linnaeus, 1771) 1, D. gloriola (Cernohorsky, 1970) 3, D. granatina (Lamarck, 1811) 1, D. hebes (Reeve, 1845) 2 comb. nov., D. latistriata (Herrmann & Salisbury, 2012) 3 comb. nov., D. lichtlei (Herrmann & Salisbury, 2012) 3 comb. nov., C. manoui (Huang, 2011) 3 comb. nov., D. praestantissima (Röding, 1798) 1, D. shikamai Habe, 19803, D. sigillata (Azuma, 1965) 3, D. valdacantamessae Maxwell, Dekkers, Berschauer & Congdon, 20173. Remarks: The genus Domiporta is a rather compact and conchologically moderately variable group; however, the colour and sculpture pattern in some species are close to those found in imbricariine genera. For example, D. circula and D. praestantissima are superficially very much like Imbricaria as circumscribed here, while D. granatina and D. carnicolor resemble species of Neocancilla. It was probably this general similarity that led Cernohorsky to assign Domiporta to the subfamily Imbricariinae. Nevertheless, the species of Domiporta mentioned can be recognized by the higher spire, more convex whorls, and the presence of a concavity which delimits the shell base from the siphonal canal. Among Mitrinae, Roseomitra gen. nov.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFADFFC0B0D47007D18CB411.taxon	description	(FIGS 16 F – I, 17 E, F) Type species: Mitra millepunctata G. B. Sowerby III, 1889; OD, herein. Diagnosis: Shell small to medium sized (10 – 45 mm), thin walled, fusiform with rather low spire and high aperture. Protoconch pointed, narrowly conical, of three or more smooth, glossy, slightly convex whorls. Suture distinct, slightly impressed. Spire rather low, orthoconoid; spire whorls with convex outline, sculptured with narrow, rounded, widely set cords. Interspaces between spiral cords bearing finer and lower secondary cords, or fine, widely interspaced, axial grooves. Last adult whorl very high, evenly convex, with shell base extended without clear distinction into straight, stout siphonal canal. Siphonal fasciole not pronounced, siphonal notch shallow or indistinct. Aperture high and moderately wide, its outer lip evenly convex or strongly convex adapically and nearly straight in its mid-width. Inner lip with five fine folds, adapicalmost strongest and succeeding ones gradually diminishing. Shell white, cream or pink, with brown blotches or with regular brown dots or dashes on spiral cords. Radula with narrow rachidian bearing five to eight subequal cusps and wide laterals with narrow base and 15 – 21 moderately long pointed cusps (Fig. 17 E, F). Distribution: Indo-Pacific, shallow subtidal to bathyal depths, on sand or mud. Species included: Roseomitra citharoidea (Dohrn, 1862) 3 comb. nov., R. earlei (Cernohorsky, 1977) 3 comb. nov., R. fluctuosa (Herrmann & Salisbury, 2013) 3 comb. nov., R. honkeri (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., R. incarnata (Reeve, 1845) 3 comb. nov., R. millepunctata (G. B. Sowerby III, 1889) 1 comb. nov., R. reticulata (A. Adams, 1853) 3 comb. nov., R. rosacea (Reeve, 1845) 1 comb. nov., R. roseovitta (Huang, 2011) 3 comb. nov., R. strangei (Angas, 1867) 3 comb. nov., R. tagaroae (Poppe, 2008) 3 comb. nov. Etymology: The genus name refers to the pink (ish) colour that characterizes some species, such as R. incarnata or R. rosacea. Gender feminine. Remarks: As the type specimen of M. millepunctata is lost [Cernohorsky (1991) noted that the type specimen cannot be traced, and the first author failed to find it in NHMUK as well], we here designate our sequenced specimen IM- 2013 - 40686, measuring 30.0 × 11.4 mm (Fig. 16 F) as neotype of M. millepunctata. This specimen was collected off Noc-nocan Is., Central Philippines, at the type locality of Mitra terryni, a junior synonym (Herrmann, 2017) of M. millepunctata. Species of the genus Roseomitra gen. nov. can be recognized by the combination of rather thin-walled shell with proportionally low spire and high aperture, distinctly rounded whorls and Domiporta - like sculpture. The shell proportions with low spire differentiate Roseomitra gen. nov. from closely related Domiporta and most other mitrid genera. Species of Pterygia and Imbricariopsis have a proportionally even higher aperture, but in these genera the shell is conical or ovate, whereas it is fusiform in Roseomitra gen. nov. In turn, Condylomitra tuberosa, C. bernhardina and Quasimitra sophiae have a notably stronger sculpture compared to Roseomitra spp. GENUS PROFUNDIMITRA FEDOSOV, HERRMANN,	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFAFFFC6B2037267D182B09E.taxon	description	(FIGS 17 G, H, 18 A – H) Type species: Profundimitra taylori sp. nov.; OD, herein. Diagnosis: Small shell (20 – 35 mm), thin walled, fusiform or elongate-fusiform, with high orthoconoid or slightly acuminate spire. Protoconch pale, semi-translucent, narrowly conical with three or more smooth whorls (usually strongly eroded). Suture adpressed, sometimes shallowly canaliculated. Teleoconch whorls gently convex, early teleoconch whorls sometimes subcylindrical. Sculpture variable, from very weak, represented by indistinct spirally aligned punctures, to deep spiral grooves separating flat, rounded or distinctly gemmate spiral cords. Siphonal canal rather long, straight, tapering, not notched at its tip. Aperture elongate. Outer lip very thin, convex, smooth or slightly undulating. Inner lip with three columellar folds, usually fine but well pronounced, sometimes very weak. Shell white, pale to yellowish, sometimes covered by light brown periostracum. Radula with laterals and rachidian of equal width, with evenly convex anterior margin bearing 12 – 14 equal pointed cusps (Fig. 17 G, H). Distribution: Indo-Pacific, bathyal depths, mostly on mud. Species included: Profundimitra abyssicola (Schepman, 1911) 3 comb. nov., Profundimitra planofilum (Huang, 2011) 1 comb. nov., P. taylori sp. nov. 1. Etymology: The name Profundimitra refers to the fact that all known species of this new genus inhabit deep water. Gender feminine. Remarks: Whereas all the sequenced species of Calcimitra were identifiable to known species, the new deep-water genus Profundimitra (as well as the related Fusidomiporta) appears to represent a kind of Mitridae terra incognita. Of four species recognized in our phylogenetic analysis, only P. planofilum has a name. The latter was initially described in Cancilla, and indeed the sculpture pattern and light brown marks on spiral cords of P. planofilum strikingly resemble the pattern seen in species of Cancilla. Profundimitra planofilum, however, differs from Cancilla by the shell base outline, which is distinctly concave, and by a slightly higher spire. The immature lectotype of C. abyssicola figured by Cernohorsky (1991) is very close to species of the Profundimitra clade in shell characteristics, and we tentatively assign this species to Profundimitra. It should be noted, though, that C. abyssicola as circumscribed by Cernohorsky combined a variety of forms, some of them [like the specimen figured by Cernohorsky (1970: pl. 6, fig. 7), which is undoubtedly conspecific with our specimens IM- 2007 - 35711 and IM- 2007 - 35618] referable to Profundimitra gen. nov., but most likely not conspecific with the lectotype, while others obviously represent true Cancilla or Imbricaria as circumscribed here. The genus Profundimitra gen. nov. can be recognized by the elongate, typically weakly sculptured shell, either white or pale, sometimes covered with light brown periostracum. The genus Calcimitra is closest to Profundimitra gen. nov. in shell morphology; however, species in the latter genus have proportionally higher spire and, correspondingly, lower last adult whorl and siphonal canal. Another genus which seems superficially close to Profundimitra gen. nov. is Eumitra, which however is characterized by a smooth columella, typically with no folds or with only one or two distinct folds, while species of Profundimitra gen. nov. have three or more columellar folds, albeit sometimes slightly sunken in the aperture. Another notable difference between the two genera is the morphology of protoconch: it is narrow and narrowly conical in Profundimitra, very distinctly demarcated from the much wider first teleoconch whorl, whereas the protoconch of Eumitra is very low, cyrtoconoid, and not clearly delimited from teleoconch. Moreover, Profundimitra gen. nov. has a characteristic radula morphology with rachidian and laterals of equal width and similar morphology, as opposed to the very small, narrow rachidian and wide laterals of Eumitra. GENUS FUSIDOMIPORTA FEDOSOV, HERRMANN,	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFA9FFC6B20376D2D637B275.taxon	description	(FIGS 17 I, 18 I – L) Type species: Fusidomiporta ponderi sp. nov.; OD, herein. Diagnosis: As the genus is established primarily based on the molecular data, and its recognition based on the morphology is problematic, we do not provide a formal diagnosis. Description (based on type species): Shell small (about 25 mm), fusiform, heavily sculptured, with rather high spire and stout siphonal canal. Protoconch multispiral, of three or more whorls usually strongly corroded or missing. Suture canaliculated; subsutural shelf well developed, rather narrow, bordered by projecting margin of adapical spiral cord. Sculpture of strong, elevated, gemmate spiral cords separated by deep interspaces; microsculpture of fine growth lines in interspaces between spiral cords. Spiral cords become distinctly beaded on penultimate and last adult whorls, forming rows of closely set, axially expanded, glossy granules. Siphonal rather long and stout, slightly recurved. Aperture elongate, its outer lip gently convex throughout its height, wavy, reflecting arrangement of spiral cords. Inner lip strongly calloused, reflected, with three strong oblique columellar folds at its mid-height. Rachidian and laterals multicuspidate, of about same width. Distribution: Indo-Pacific, bathyal depths in about 400 – 500 m. Species included: Fusidomiporta ponderi sp. nov. 1,? F. suturata (Reeve, 1845) 3 comb. nov. Etymology: The name Fusidomiporta reflects the close affinity of the new genus to Domiporta and refers to the fusiform shell of the type species. Gender feminine. Remarks: Morphologically, F. ponderi sp. nov. falls within the range of variation seen in Profundimitra gen. nov., both in the shell and radula, and we even initially thought that specimen IM- 2013 - 40665 (now the holotype of F. ponderi) was conspecific with IM- 2007 - 35711 (an unknown species of Profundimitra). The molecular distinctiveness of F. ponderi sp. nov. was thus totally unexpected. However, none of the single-gene analyses placed F. ponderi sp. nov. in the same molecular clade as Profundimitra, and a Fusidomiporta – Profundimitra grouping is only present in the three gene-based tree. Thus, the genus is largely defined by molecular characters; the morphological basis for the separation between Fusidomiporta and Profundimitra needs to be more carefully elaborated, for which more specimens need to be examined. A sculpture pattern close to that present in Fusidomiporta gen. nov. is also observed in species of Gemmulimitra gen. nov. (G. neocaledonica sp. nov. and G. boucheti); however, Gemmulimitra differs notably from Fusidomiporta in radula morphology, showing a classical mitrid radula with a narrow rachidian (Fig. 20 F, G). GENUS CANCILLOPSIS FEDOSOV, HERRMANN,	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFA9FFC7B0AB740BD1AEB211.taxon	description	(FIG. 18 M) Type species: Cancilla meimiaoae Huang & Salisbury, 2017; OD, herein. Diagnosis: As the genus is established primarily based on the molecular data, and its recognition based on the morphology is problematic, we do not provide a formal diagnosis. Description (based on the type species, adapted from Huang & Salisbury (2017): Shell medium sized (50 – 82 mm), fusiform, with high spire and long, stout siphonal canal. Protoconch multispiral, cyrtoconoid, of about three glossy whorls. Teleoconch of up to ten gently convex whorls; suture distinct. Sculpture of rounded spiral cords, intersected by closely set, slightly prosocline axial riblets, forming fine beaded sculpture on early spire whorls. On last adult whorl cords more widely set, and beaded sculpture indistinct, as axial riblets are well pronounced in interspaces between spiral cords only. Siphonal canal stout, deeply notched, slightly recurved at its tip. Aperture elongate, its outer lip evenly convex throughout its length. Inner lip calloused, with four closely set fine columellar folds at its mid-height. Shell white with irregular yellow or brownish blotches. Distribution: Indo-Pacific, bathyal depths. Species included: Cancillopsis meimiaoae (Huang & Salisbury, 2017) 1 comb. nov., † C. acuminata (Shuto, 1969) 3 comb. nov. Etymology: The name Cancillopsis refers to the close resemblance between C. meimiaoae, the type species of the new genus, and species of Cancilla. Gender feminine. Remarks: Cancillopsis meimiaoae shows a notable resemblance to the species of the imbricariine genus Cancilla as well as to the species of Profundimitra gen. nov. and Fusidomiporta gen. nov., while showing no phylogenetic affinity to either of them. A visible concavity at the transition of shell base to the siphonal canal distinguishes the new genus from similarly sculptured Cancilla species. We hypothesize that Cancillopsis gen. nov. is more closely related to the Domiporta group of Mitrinae, and so we provisionally place it in Mitrinae. However, a more comprehensive taxon sampling of the deep-water Mitridae is obviously needed to delineate these convergent deep-water lineages and to clarify affinities of Cancillopsis gen. nov. GENUS PSEUDONEBULARIA FEDOSOV, HERRMANN,	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFA8FFC7B22D7467D628B5F3.taxon	description	(FIGS 19, 20 A – E) Type species: Mitra tornata Reeve, 1845; OD, herein. Diagnosis: Shell small (9 – 30 mm), fusiform or broadly fusiform to biconical, vividly coloured. Protoconch pointed, narrowly conical, of about three slightly convex, glossy whorls. Suture slightly impressed, often indistinct. Teleoconch whorls evenly convex or flattened, sometimes giving the spire a distinctly stepped profile. Sculpture of strong, rounded spiral cords, regularly interspaced and covering the entire shell or, rarely, limited to shell base. Interspaces between cords concave, smooth or bearing fine, dense riblets. Siphonal canal moderately long, stout, siphonal notch shallow or absent. Aperture rather narrow, elongate, with undulating outer lip, sometimes bearing rounded denticles on its inside. Inner lip calloused, often reflected, with three to four strong columellar folds, adapicalmost strongest. Radula with broad laterals, bearing equal, rather long, widely set cusps throughout their width; rachidian not exceeding half the width of the laterals, multi- or unicuspidate (Fig. 20 A – E). Distribution: Indo-Pacific, intertidal to upper subtidal depths, rocky flats, reef crevices and coral rubble. Species included: Pseudonebularia atjehensis (Oostingh, 1939) 1 comb. nov., P. chrysalis (Reeve, 1844) 3 comb. nov., P. cingulata (A. Adams, 1853) 3 comb. nov., P. connectens (Dautzenberg & Bouge, 1923) 1 comb. nov., P. cucumerina (Lamarck, 1811) 3 comb. nov., P. cuyosae (Poppe, 2008) 3 comb. nov., P. damasomonteiroi (Cossignani & Cossignani, 2007) 3 comb. nov., P. doliolum (Küster, 1839) 3 comb. nov., P. dovpeledi (Turner, 1997) 3 comb. nov., P. fraga (Quoy & Gaimard, 1833) 2 comb. nov., P. gracilefragum (Turner, 2007) 2 comb. nov., P. indentata (G. B. Sowerby II, 1874) 3 comb. nov., P. kantori (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., P. kilburni (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., P. lienardi (G. B. Sowerby II, 1874) 3 comb. nov., P. maesta (Reeve, 1845) 1 comb. nov., P. oliverai (Poppe, 2008) 3 comb. nov., P. pediculus (Lamarck, 1811) 1 comb. nov., P. perdulca (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., P. proscissa (Reeve, 1844) 3 comb. nov., P. rotundilirata (Reeve, 1844) 3 comb. nov., P. rubiginea (A. Adams, 1855) 3 comb. nov., P. rubritincta (Reeve, 1844) 3 comb. nov., P. rueppellii (Reeve, 1844) 3 comb. nov., P. rutila (A. Adams, 1853) 3 comb. nov., P. sarinoae (Poppe, 2008) 3 comb. nov., P. semiferruginea (Reeve, 1845) 3 comb. nov., P. silviae (Turner, 2007) 3 comb. nov., P. tabanula (Lamarck, 1811) 1 comb. nov., P. tornata (Reeve, 1845) 1 comb. nov., P. wareni (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., P. willani (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., P. yayanae (Huang, 2011) 3 comb. nov. Etymology: Pseudonebularia means ‘ false Nebularia ’ and refers to the long accepted, and erroneous, placement of the included species in Nebularia, which it resembles superficially. Gender feminine. Remarks: Pseudonebularia gen. nov. comprises a group of popular, colourful shallow water or even intertidal mitres that are traditionally referred to Nebularia but, as demonstrated by our analysis, are not related to it, as Nebularia is recovered in a sister position to Pterygia. Species of Pseudonebularia gen. nov. typically have a small, but broad and robust, brightly coloured shell. While red and variegated biconical shells in species of the P. fraga group are quite distinctive, more elongated species, for example P. atjehensis, resemble species of Domiporta or Imbricaria as newly circumscribed here. The elongated Pseudonebularia species, however, can be distinguished by generally broader, rounded spiral cords that bear no darker lines on their crests. The three species of Pseudonebularia studied anatomically show a significant disparity in radular morphology, with rachidian varying considerably from bow-shaped with multiple densely set cusps to very narrow, but robust, bearing a single short, pointed cusps on a bulbous shaft. GENUS GEMMULIMITRA FEDOSOV, HERRMANN,	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFA8FFC5B0B57386D7ECB5C5.taxon	description	(FIGS 20 F, G, 21) Type species: Gemmulimitra neocaledonica sp. nov.; OD, herein. Diagnosis: As the genus is established primarily based on the molecular data, and its recognition based on the morphology is problematic, we do not provide a formal diagnosis. Description (based on the type species): Shell small (about 20 – 25 mm), elongate-fusiform to almost biconical. Protoconch multispiral, of three or more slightly convex, glossy whorls. Suture canaliculated. Subsutural ramp forming distinct shelf, giving spire a stepped appearance. Spire whorls sculptured with strong, closely set, spiral cords overriding low indistinct axial ribs to form series of prominent beads. On later teleoconch whorls, interspaces between gemmate spiral cords broadened, with axial ribs becoming more distinct and sculpture rather reticulate. Interspaces between axial ribs and spiral cords forming deep quadrangular depressions with microsculpture of fine axial growth lines. Shell base extended to rather long, tapering, slightly notched siphonal canal. Aperture narrow, elongated. Outer lip wavy, convex in adapical portion and straight below mid-height; inside of outer lip smooth. Inner lip strongly calloused, sometimes reflected, with four closely set fine columellar folds. Rachidian narrow, bearing eight strong, moderately long, pointed cusps; laterals wide, bearing 16 + strong, pointed, rather widely set cusps. Distribution: Indo-Pacific, subtidal to bathyal depths. Species included: Gemmulimitra aliciae (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., G. apprimapex (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., G. avenacea (Reeve, 1845) 1 comb. nov., G. boucheti (Cernohorsky, 1988) 2 comb. nov., G. duplilirata (Reeve, 1845) 3 comb. nov., G. edgari (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., G. gonatophora (Sturany, 1903) 3 comb. nov., G. hansturneri (E. Guillot de Suduiraut & E. G. Guillot de Suduiraut, 2009) 3 comb. nov., G. margaritata (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., G. neocaledonica sp. nov. 1, G. rubiginosa (Reeve, 1844) 3 comb. nov., G. solanderi (Reeve, 1844) 3 comb. nov., G. strongae (Poppe, Tagaro & Salisbury, 2009) 1 comb. nov. Etymology: The genus name refers to the sculpture of gemmate cords that characterizes many species in this newly recognized lineage, including the type species G. neocaledonica. Remarks: Gemmulimitra neocaledonica sp. nov. and G. avenacea that together constitute the Gemmulimitra clade on multi-gene trees have little in common in shell morphology, and thus this clade is at present defined solely based on the molecular characteristics. The third species confidently included in the genus is G. strongae that is close to G. neocaledonica sp. nov. in the COI - based tree. The other species here attributed to Gemmulimitra are tentatively classified in this genus based on their general resemblance to the type species or to Mitra avenacea, rather than on any specific diagnostic feature. Therefore, the species included in Gemmulimitra constitute two conchologically distinctive groups: species in the first group have a typical sculpture of gemmate cords, such as G. neocaledonica sp. nov., G. boucheti (Fig. 21 C), G. strongae (Fig. 21 D) and G. gonatophora (holotype figured by Albano et al., 2017), while species in the second group have continuous, smooth spiral cords, such as G. avenacea (Fig. 21 E), G. edgari and G. rubiginosa (Fig. 21 F). Species of the first group closely resemble Fusidomiporta ponderi sp. nov., whereas species of the second group resemble species of Pseudonebularia, from which, however, they can be distinguished by a more elongated shell with more flattened spire whorls. The radula of Gemmulimitra is plesiomorphic and phylogenetically uninformative. More studies are needed to better define the boundaries of Gemmulimitra gen. nov. and elaborate its diagnosis. SUBFAMILY STRIGATELLINAE TROSCHEL, 1869 (FIGS 22, 23)	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFAAFFD8B098739AD7F2B4ED.taxon	materials_examined	Type species: Mitra zebra Lamarck, 1811; SD, Gray (1847: 141). Synonyms: Chrysame H. Adams & A. Adams, 1853; Type species: Mitra coronata Lamarck, 1811; SD, Cox (1927: 91). Phaeomitra Martens, 1880; Type species: Mitra fulva Swainson, 1829 (= Mitra coffea Schubert & Wagner, 1829); SD, Coan (1966: 131). Diagnosis: Shell small (12 – 40 mm), broadly fusiform, solid and stout in appearance, protoconch pointed, narrowly conical, of about 2.5 to three smooth, very slightly convex whorls. Suture deeply impressed. Spire whorls gently convex or flattened to subcylindrical, sometimes with narrow subsutural area, giving spire a somewhat telescopic outline. Sculpture typically not developed; sometimes shell sculptured with very faint to strong rounded spiral cords over all the shell surface or limited to shell base. Last adult whorl usually strongly convex, siphonal canal typically short and stout to moderately long, with distinct siphonal fasciole. Siphonal notch shallow or absent. Aperture elongate, narrow; outer lip straight adapically and strongly convex in its abapical portion, often thickened adapically, forming a callous inner varix, constricting aperture. Inner lip with three to five fine oblique columellar folds. Shell orange to dark brown, typically with a pattern of white bands and blotches. Radula tri- or monoseriate. In triseriate radulae, rachidian narrow, with five to seven very short and robust cusps, unpaired central cusp always present (Fig. 23); in uniseriate radulae, rachidian bearing 20 – 25 pointed cusps with unpaired central cusp distinctly exceeding others. Laterals broad, wavy, with their distal halves smooth and seven to ten rather short and robust cusps situated proximally. Distribution: Indo-Pacific and Panamic provinces, intertidal and subtidal depths, mostly on hard substrates (rocky shores, reef crevices or coral rubble). Species included: Strigatella abacophora (Melvill, 1888) 3 comb. nov., S. amaura (Hervier, 1897) 3, S. ambigua (Swainson, 1829) 3 comb. nov., S. assimilis (Pease, 1868) 3, S. aurantia (Gmelin, 1791) 1 comb. nov., S. auriculoides (Reeve, 1845) 3, S. aurora (Dohrn, 1861) 2 comb. nov., S. colombelliformis (Kiener, 1838) 2, S. coronata (Lamarck, 1811) 1 comb. nov., S. crassicostata (G. B. Sowerby II, 1874) 3 comb. nov., S. decurtata (Reeve, 1844) 1, S. fasciolaris (Deshayes, 1834) 3 comb. nov., S. flavocingulata (Lamy, 1938) 3, S. fulvescens (Broderip, 1836) 1 comb. nov., S. holkosa (B. Q. Li, 2005) 3 comb. nov., S. imperialis (Röding, 1798) 1 comb. nov., S. litterata (Lamarck, 1811) 1, S. luctuosa (A. Adams, 1853) 1, S. lugubris (Swainson, 1821) 1 comb. nov., S. nana (Reeve, 1844) 3 comb. nov., S. paupercula (Linnaeus, 1758) 1, S. pica (Dillwyn, 1817) 3, S. retusa (Lamarck, 1811) 3, S. scutulata (Gmelin, 1791) 1 comb. nov., S. subruppeli (Finlay, 1927) 3 comb. nov., S. telescopium (Reeve, 1844) 3, S. testacea (Broderip, 1836) 3 comb. nov., S. ticaonica (Reeve, 1844) 3 comb. nov., S. tristis (Broderip, 1836) 1, S. vexillum (Reeve, 1844) 1, S. vultuosa (Reeve, 1845) 3 comb. nov.,? S. coffea (Schubert & J. A. Wagner, 1829) 2 comb. nov.,? S. pudica (Pease, 1860) 2,? S. tabida (Herrmann & Salisbury, 2013) 3 comb. nov., S. zebra (Lamarck, 1811) 1. Remarks: The sculpture of strong rounded cords of some species of the Strigatella clade may lead to a wrong generic allocation; for instance, Cernohorsky (1976) placed S. vexillum, S. aurantia, S. lugubris and even S. fulvescens in Nebularia, whereas our phylogenetic analysis unequivocally places them in Strigatella. There are three conchologically more or less distinct groups of Strigatella: (1) a S. paupercula group with typical broadly fusiform Strigatella shell, (2) a S. fulvescens group with more elongate shell, distinctly recurved siphonal canal and lack of inner callous varix on the outer aperture lip and (3) a S. coronata group with a fusiform shell also lacking a callous varix on the outer aperture lip, and also distinguished by a uniserial radula. Strigatella coronata and S. lugubris form a very distinct clade in our phylogenetic tree, and the name Chrysame is available for it. Given its distinctive morphology and its long branch on the molecular tree, the S. coronata group could be treated as a subgenus of Strigatella. However, we prefer to abstain from recognizing Chrysame, as it would necessitate to also recognize as subgenera the other two Strigatella groups – a level of taxonomic splitting that we are not currently prepared to defend. SUBFAMILY IMBRICARIINAE TROSCHEL, 1867 (FIGS 24 – 30)	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFB7FFD9B07472E9D65FB1F2.taxon	description	(FIGS 24, 25 A – H) Type species: Imbricaria conica Schumacher, 1817 (= Mitra conularis Lamarck, 1811); M. Synonym: Conoelix Swainson, 1821. Type species: Conoelix lineatus Swainson, 1821 (= M. conularis Lamarck, 1811); OD. Diagnosis: Shell small to medium sized (10 – 55 mm), fusiform, narrowly biconical to nearly conical, uniformly coloured or with variegated pattern. Protoconch narrowly conical, of three or more slightly convex or flattened, smooth and glossy whorls. Early teleoconch whorls forming narrowly conical apex; later whorls widening at equal rate, resulting in fusiform or biconical shell, or (typically) at increasing rate, resulting in acuminated spire and slightly inflated or conical last adult whorl. Early spire whorls sculptured with rounded spiral cords; late whorls smooth or with spiral cords, the latter wide and rounded or fine and sharp, sometimes bearing regular tubercles. Interspaces between cords smooth or with fine, dense riblets. Siphonal canal tapering, with distinct siphonal notch. Aperture elongate, narrow to slit-like. Outer apertural lip convex in its adapical portion; inner lip with four to five fine columellar folds gradually diminishing towards siphonal canal. Rachidian with four to seven cusps, with one to two marginal pairs of cusps notably weaker than the two or three medial cusps; unpaired central cusp when present equal or weaker than two flanking cusps (Fig. 25 A – H). Lateral teeth broad, with seven to 12 triangular or spindle-like, long and pointed subequal cusps, gradually increasing in size towards rachidian, with one cusp strongest, sometimes very long, greatly exceeding others, resembling a tin-opener (Fig. 25 D – F).	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFB6FFDFB3AE7144D6F0B15B.taxon	distribution	Distribution: Indo-Pacific, intertidal to bathyal depths, mostly on soft bottoms, mud, silty or well-washed sand. Species included: Imbricaria amoena (A. Adams, 1853) 3 comb. nov., I. annulata (Reeve, 1844) 3 comb. nov., I. armonica (T. Cossignani & V. Cossignani, 2005) 3 comb. nov., I. astyagis (Dohrn, 1860) 3 comb. nov., I. bacillum (Lamarck, 1811) 3 comb. nov., I. baisei (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., I. bantamensis (Oostingh, 1939) 1 comb. nov., I. bellulavaria (Dekkers, Herrmann, Poppe & Tagaro, 2014) 3 comb. nov., I. cernohorskyi (Rehder & Wilson, 1975) 3 comb. nov., I. cloveri (Cernohorsky, 1971) 3 comb. nov., I. conularis (Lamarck, 1811) 1, I. flammea (Quoy & Gaimard, 1833) 3 comb. nov., I. flammigera (Reeve, 1844) 1 comb. nov., I. fulgetrum (Reeve, 1844) 1 comb. nov., I. hidalgoi (G. B. Sowerby III, 1903) 3 comb. nov., I. hrdlickai (Salisbury, 1994) 1 comb. nov., I. insculpta (A. Adams, 1851) 1 comb. nov., I. interlirata (Reeve, 1844) 2 comb. nov., I. intersculpta (G. B. Sowerby II, 1870) 3 comb. nov., I. kermadecensis (Cernohorsky, 1978) 3 comb. nov., I. maui (Kay, 1979) 3 comb. nov., I. nadayaoi (Bozzetti, 1997) 3 comb. nov., I. philpoppei (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., I. polycincta (Turner, 2007) 3 comb. nov., I. pretiosa (Reeve, 1844) 2 comb. nov., I. pugnaxa (Poppe, Tagaro & Salisbury, 2009) 1 comb. nov., I. rufilirata (A. Adams & Reeve, 1850) 3 comb. nov., I. ruberorbis (Dekkers, Herrmann, Poppe & Tagaro, 2014) 3 comb. nov., I. rufogyrata (Poppe, Tagaro & Salisbury, 2009) 1 comb. nov., I. salisburyi (Drivas & Jay, 1990) 2 comb. nov., I. tahitiensis (Herrmann & Salisbury, 2012) 3 comb. nov., I. verrucosa (Reeve, 1845) 1 comb. nov., I. yagurai, (Kira, 1959) 1 comb. nov., I. zetema (Dekkers, Herrmann, Poppe & Tagaro, 2014) 3 comb. nov. Remarks: The genus Imbricaria is, in the present classification, undergoing a dramatic rearrangement: whereas species traditionally classified in Imbricaria, like I. punctata and I. olivaeformis, that are conchologically close to I. conularis, are excluded from the genus, numerous Indo-Pacific species earlier assigned to Ziba and Subcancilla are now transferred to Imbricaria. As circumscribed here, Imbricaria appears rather heterogeneous morphologically, with shell varying from conical to narrowly fusiform, most species having a notably higher spire than I. conularis. Shell sculpture may be faint, like in the type species, or represented by strong elevated cords, and depressions between cords are often sculptured with dense riblets. Moreover, species of Imbricaria display a remarkable disparity in radular morphology, as seen in the degree of cusp differentiation in both the rachidian and lateral teeth. In this respect, the eight studied species form a continuous grade from radulae with subequal cusps (presumably a plesiomorphic state, found in I. conularis and I. pugnaxa – see ‘ Discussion’) to more derived ones with one or two cusps on each tooth greatly exceeding the others.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFB0FFDDB0A47775D1BCB341.taxon	description	(FIGS 25 I – M, 26) Type species: Mitra isabella Swainson, 1831; SD, Herrmannsen (1846: 166). Diagnosis: Shell small to rather large (15 – 110 mm), fusiform, with high aperture and long, tapering siphonal canal. Protoconch conical, of about three very slightly convex smooth whorls. Spire whorls evenly convex to subcylindrical; suture distinct, impressed. Shell sculptured with spiral cords that are slightly gemmate or with finely dentate margins, very wide and flat, separated by narrow grooves, or with strong, narrow and elevated cords separated by broad depressions bearing regular, very fine riblets. Shell base gradually extended into rather long, tapering siphonal canal. Siphonal fasciole not pronounced, notch deep or shallow. Aperture elongate, narrow; outer aperture lip smooth, gently convex adapically, and flattened in its lower portion. Inner lip with four oblique columellar folds, adapicalmost strongest. Shell pale or cream, typically with multiple darker spiral strokes on crests of spiral cords. Radula with narrow rachidian, usually bearing four cusps, of which central pair notably stronger (Fig. 25 I – M). Lateral teeth with seven to ten pointed triangular cusps, second proximal cusp greatly exceeding others, and subsequently gradually weakening towards margins of radula. Distribution: Indo-Pacific, bathyal depths, mostly on mud. Species included: Cancilla baeri (Turner & Cernohorsky, 2003) 1, C. fibula Poppe, Tagaro & Salisbury, 20091, C. heinickei (Salisbury & Guillot de Suduiraut, 2003) 3 comb. nov., C. herrmanni Dekkers, 20143, C. isabella (Swainson, 1831) 2, C. liliformis Huang & Salisbury 20173, C. rehderi (Webb, 1958) 1, C. rikae Guillot de Suduiraut, 20043, C. schepmani (Salisbury & Guillot de Suduiraut, 2003) 1 comb. nov., C. turneri Poppe, Tagaro & Salisbury, 20093. Remarks: The typification of Cancilla renders the identity of the genus a little weak: the specimen of M. isabella originally illustrated by Swainson (1831) was selected as lectotype by Cernohorsky (1991), but, like many types of Swainson relevant to this study, this specimen is lost (Cernohorsky, 1991). Taking in consideration frequent parallelisms in shell morphology demonstrated in Mitridae, the utility of Swainson’s original illustration for taxonomic decisions is limited and it will be desirable in the future to designate a neotype with associated molecular data. The radula of C. isabella was studied by Salisbury & Huang (2015). Although the two specimens dissected by these authors differ notably in shell proportions, their radulae are morphologically close to each other, and rather peculiar. The rachidian bears two pairs of long pointed cusps (the lateral cusps being notably weaker than the central ones), flanked by two pairs of very weak cusps; the laterals bear long pointed cusps that progressively increase in size towards tooth mid-width, whereas the distal halves bear no cusps and show only weak serration (Salisbury & Huang 2015: figs 1.1, 3.2), a pattern comparable to the laterals in Scabricola, which are however still more robust. Of all Cancilla species studied by us, C. baeri is probably closest to C. isabella: the specimen of C. baeri MNHN IM- 2013 - 58853 (Fig. 26 A) resembles one of the two specimens of C. isabella studied by Salisbury & Huang (2015: fig. 3.2 b) in shell proportions, whorl outline and shape of the siphonal canal. The other studied Cancilla species – the C. rehderi – C. fibula complex and C. schepmani – are generally smaller, with less convex whorls and stouter siphonal canal, which is not clearly delineated from the shell base. Therefore, the genus Cancilla, as we circumscribe it, is rather heterogeneous and can be subdivided by shell morphology into two conventional groups: ‘ robust’ Cancilla with species such as C. isabella, C. baeri and C. chuoi, and ‘ slender’ Cancilla with species like C. fibula, C. rehderi, C. schepmani and C. liliformis. The relationships of these two conchological groups is supported by (1) inclusion of C. baeri in the Cancilla clade (clade 24 – 7) in the COI - based tree and (2) occurrences of very similar radular morphologies in ‘ slender’ species of the C. fibula – C. rehderi complex (Fig. 25 I, K) and in ‘ robust’ C. baeri IM- 2013 - 58853 (Fig. 25 L). It should be noted, however, that the radula of C. schepmani (Fig. 25 M), with a morphology closest to the radulae of Strigatella or of the mitrine Neotiara nodulosa (Fig. 10 E), is thoroughly different from the radulae of other studied Cancilla species. Therefore, Cancilla appears to be very heterogeneous in radular morphology as well. Taking into account this observed radular disparity among sequenced (and thus confirmed related) Cancilla species, we believe that the divergent radular morphology of C. isabella does not contradict its affinity to the other species studied herein. Species of Domiporta can be distinguished from the ‘ robust’ Cancilla species by the sculpture pattern with narrow, elevated, widely interspaced spiral cords, whereas a distinct concavity at the transition of shell base to siphonal canal distinguishes species of Domiporta from ‘ slender’ Cancilla species. Moreover, the two genera differ greatly in radular morphology. Besides, there is a clear resemblance between some species of Imbricaria and ‘ slender’ Cancilla: I. fulgetrum and C. rehderi / fibula (especially our immature specimen IM- 2013 - 19073) or the I. yagurai group and C. cf. fibula (represented by IM- 2013 - 40646). In these pairs, the species of Cancilla can be recognized by the more convex outline of the shell base, which gives the shell a rather fusiform shape compared to the rather biconical shell of Imbricaria. The new genus Cancillopsis can be differentiated from the ‘ robust’ Cancilla species by the higher spire with rather subcylindrical whorls, and from the ‘ slender’ Cancilla species by the presence of a concavity at the base of the siphonal canal. Finally, the radula with rachidian and lateral teeth of similar morphology differentiates Profundimitra gen. nov. and Fusidomiporta gen. nov. from Cancilla. GENUS IMBRICARIOPSIS FEDOSOV, HERRMANN,	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFB2FFD2B22D7537D67CB61D.taxon	description	(FIGS 27, 28 A) Type species: Conoelix punctatus Swainson, 1821; OD, herein. Diagnosis: Shell small (10 – 20 mm), conical, uniformly pale or with dark tip of siphonal canal. Protoconch pointed multispiral, narrowly conical, dark. Suture distinct, impressed. Spire very short, less than 20 % of total shell height, conical or distinctly acuminate, starting with several subcylindrical whorls. Last adult whorl very tall, sculptured with regular, distinctly punctate, spiral grooves. Siphonal notch shallow or moderately deep. Aperture very tall and narrow, slit-like. Outer aperture lip thick, flattened, smooth inside; inner lip with five fine oblique folds situated on abapical portion of columella. Radula triserial; rachidian with six cusps, paired central cusps greatly exceeding those situated laterally (Fig. 28 A). Lateral teeth with five cusps, second cusp from rachidian very robust, notably exceeding the others. Distribution: Indo-Pacific, intertidal and upper subtidal depths, on sand patches and in reef crevices. Species included: Imbricariopsis conovula (Quoy & Gaimard, 1833) 2 comb. nov., I. punctata (Swainson, 1821) 1 comb. nov., I. vanikorensis (Quoy & Gaimard, 1833) 2 comb. nov.,? I. carbonacea (Hinds, 1844) 3 comb. nov. Etymology: The name Imbricariopsis reflects the resemblance of the species in this newly recognized lineage with the type species of Imbricaria, I. conularis, that has long distracted mitrid taxonomists. Gender feminine. Remarks: Few mitrids are characterized by an almost involute conical shell, and these were formerly allocated to either Imbricaria or Pterygia. However, we found Imbricaria as traditionally construed to be paraphyletic, with I. punctata forming a lineage of its own, not closely related to I. conularis. Although a single species has been sequenced, we hypothesize a close relationship between I. punctata, I. conovula and I. vanikorensis, largely based on shell characters. Imbricariopsis gen. nov., as we circumscribe it, constitutes a rather distinctive group of small shallow-water mitrids: species of Imbricariopsis can be recognized by conical, weakly sculptured, or not sculptured at all, pale shells with a minute, pointed apex (that is much lower than in I. conularis). The radulae of the three species that we assign to Imbricariopsis gen. nov. were illustrated by Cernohorsky (1970) and are again figured here; all three share the same general morphology of lateral radular teeth, with one robust major and several minor cusps situated on both sides of the major one.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFBCFFD3B20F75A9D084B59A.taxon	description	(FIGS 28 D – F, 29 A – F) Type species: Mitra serpentina Lamarck, 1811 (= Voluta variegata Gmelin, 1791); SD, Gray (1847: 141). Diagnosis: see under ‘ Remarks’. Distribution: Indo-Pacific, upper subtidal depths, on sand. Species included: Scabricola bicolor (Swainson, 1824) 3, S. desetangsii (Kiener, 1838) 1, S. olivaeformis (Swainson, 1821) 1, S. padangensis (Thiele, 1925) 2, S. variegata (Gmelin, 1791) 1,? Scabricola barrywilsoni (J. Cate, 1968) 3,? S. albina A. Adams, 18533,? S. caerulea (Reeve, 1844) 3,? S. condei Guillot de Suduiraut, 2 0 0 1 3, S. cor i acea (Reeve, 1 8 4 5) 2,? S. b a ck ae (Cernohorsky, 1973) 3,? S. dampierensis Salisbury & Heinicke, 19983,? S. dianneae (Salisbury & Guillot de Suduiraut, 2003) 3 comb. nov.,? S. eximia (A. Adams, 1853) 3,? S. geigeri Poppe, Tagaro & Salisbury, 20093,? S. gilbertsoni (Cate, 1968) 3 comb. nov.,? S. guttata (Swainson, 1824) 3 comb. nov.,? S. hayashii (Kira, 1959) 3,? S. incisa (A. Adams & Reeve, 1850) 3,? S. ivanmarrowi Marrow, 20163,? S. lavoisieri Guillot de Suduiraut, 20023,? S. lorenzi (Poppe & Tagaro, 2006) 3 comb. nov.,? S. mariae (A. Adams, 1853) 3,? S. martini Poppe & Tagaro, 2006 3,? S. petiti Poppe & Tagaro, 20063,? S. parkinsoni Salisbury & Wolff, 2005 3,? S. potensis (Montrouzier, 1858) 3,? S. sowerbyi d’Orbigny, 18523,? S. splendidula Salisbury & Guillot de Suduiraut, 20033,? S. vicdani Cernohorsky, 19813,? S. yaekoae (Habe & Kosuge, 1966) 3. Remarks: The genus Scabricola as commonly understood is extremely heterogeneous. It combines such dissimilar forms as S. yaekoae, which in shell outline is close to Mitra mitra, and S. potensis (Montrouzier, 1858), with its ovate shell resembling species of Pterygia. The range of sculpture patterns is equally considerable: whereas most species of Scabricola are characterized by wide and low cords or punctate grooves, some (e. g. S. padangensis) have a ‘ Subcancilla - type’ sculpture pattern (Cernohorsky, 1991) of narrow, high cords, and still others, like S. coriacea, show a distinctly crenulated sculpture. In the treatment of Cernohorsky, the more slender species of Scabricola, with weaker sculpture, were separated in the subgenus Scabricola (Swainsonia). It should be noted, though, that the boundaries of the two subgenera appear blurred, primarily because S. variegata (the type species of Scabricola) is conchologically closer to Swainsonia than to many species of Scabricola (Scabricola). One of the important diagnostic characters of Scabricola is the morphology of the radula lateral teeth, unique for the Mitridae, ‘ with 3 – 5 strong cusps on the inward facing angular plate’ (Cernohorsky, 1991: 115) (Fig. 28 D – S. variegata). This radular morphology is shared by seven species of Scabricola (Scabricola) and Scabricola (Swainsonia) studied by Cernohorsky (1970), whereas radulae with divergent morphologies are found in S. desetangsii (Fig. 28 E) and S. coriacea (Cernohorsky, 1970: fig. 120): the former has unicuspidate laterals, the latter lacks the major cusp on laterals which, therefore, have a somewhat intermediate morphology between typical Scabricola and Neocancilla. Because the boundaries of the genus remain poorly understood, we are only confident about the placement of those species for which molecular and radular morphology data are available, and the remaining species are placed in Scabricola with a question mark. Given this major uncertainty on its contents, we refrain from giving a formal diagnosis of the genus.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFBCFFD1B0597026D2A2B584.taxon	description	(FIGS 28 B, C, 29 G – I) Type species: Mitra fissurata Lamarck, 1811; by typification of replaced name (Mitrella Swainson, 1831 non- Risso, 1826). Synonym: Mitrella Swainson, 1831 (not Risso, 1826). Type species: Mitra fissurata Lamarck, 1811; SD, Cossmann (1899: 152). Diagnosis: Shell small to medium sized (15 – 65 mm), fusiform or elongate-biconical, weakly sculptured and often polished in appearance. Protoconch narrowly conical, of about three smooth, weakly convex, whorls. Spire ortho- or cyrtoconoid, spire whorls evenly convex to flattened; suture distinct, impressed. Sculpture of regular, shallow and widely spaced punctate spiral grooves, either covering the entire shell surface, or limited to the adapical whorl portion, or lacking entirely. Axial sculpture usually not developed or, when present, consisting of numerous, fine and low, rounded folds, developed on the spire but indistinct on the periphery of the last adult whorl. Shell base typically with convex outline, continuing into short and stout, deeply notched, siphonal canal. Siphonal fasciole indistinct or moderately developed. Aperture elongate, narrow to moderately wide. Outer aperture lip straight or gently evenly convex adapically; anterior edge of outer lip strongly convex, sometimes slightly drawn abapically. Inner lip with four to five fine or strong, subequal columellar folds. Shell light brown, orange or white, typically with intricate scalelike pattern of fine light or white lines. Scabricola - type radula with comb-like rachidian bearing six long, slen- der cusps, the four central ones notably exceeding the two marginal ones (Fig. 28 B, C). Lateral teeth with very strong, robust cusp, giving tooth the appearance of a tin-opener, the serrate inner surface of tooth bearing three to four weaker cusps. Base of laterals with curved upper margin, forming a socket interlocking laterals of adjacent rows. Distribution: Indo-Pacific, upper subtidal depths, on sand. Species included: Swainsonia biconica Bozzetti, 20113, S. casta (Gmelin, 1791) 1, S. ekerae Cernohorsky, 19733, S. fissurata (Lamarck, 1811) 2, S. fusca (Swainson, 1824) 1, S. limata (Reeve, 1845) 3, S. newcombii (Pease, 1869) 3, S. ocellata (Swainson, 1831) 3. Remarks: Many contemporary authors have treated Swainsonia as a subgenus of Scabricola (Cernohorsky, 1970, 1991; Pechar, Prior & Parkinson, 1980; but see Poppe & Tagaro, 2008), but our results indicate that it should be elevated to full genus. This treatment of Scabricola and Swainsonia, however, contradicts the phylogenetic hypothesis implied by radular morphology. The typical Scabricola radula (as described by Cernohorsky, 1970) is present in Scabricola variegata, Swainsonia fusca and Swainsonia casta, while the radulae of Scabricola desetangsii and S. olivaeformis show a divergent morphology.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFBEFFD6B3ED7021D15AB59F.taxon	description	(FIGS 28 G – I, 30) Type species: Voluta papilio Link, 1807; OD. Diagnosis: Shell small to medium sized (10 – 65 mm), fusiform, with short and stout siphonal canal. Protoconch pointed, narrowly conical, of about three slightly convex glossy whorls. Suture deeply impressed to canaliculate. Spire orthoconoid, teleoconch whorls evenly convex. Sculpture of wide spiral cords, low or slightly elevated adapically, or bearing rows of fine beads. Spiral cords separated by narrow deep groves and intersected by axial riblets of similar strength, forming regular cancellate sculpture. Siphonal canal short and robust, gradually merging to shell base, although separated from it by narrow depression at about the level of the second columellar fold. Siphonal notch deep. Aperture elongate; outer aperture lip evenly convex along most of its length. Inner aperture lip with four strong columellar folds, adapicalmost strongest. Shell cream, pale or orange, usually with contrasting darker bands. Radula with narrow rachidian bearing six blunt, very robust cusps; paired central cusps greatly exceeding the others (Fig. 28 G – I). Lateral teeth with four or five very short and blunt cusps situated proximally, distal portions of teeth without cusp. Distribution: Indo-Pacific, intertidal and upper subtidal depths, mostly on sand. Species included: Neocancilla antoniae (H. Adams, 1870) 3, N. arenacea (Dunker, 1852) 3, N. clathrus (Gmelin, 1791) 2, N. hartorum Poppe, Salisbury & Tagaro, 20153, N. hemmenae (Salisbury & Heinicke, 1993) 3, N. kayae Cernohorsky, 19783, N. maculosa (Gmelin, 1791) 1, N. madagascariensis Herrmann, 20173, N. papilio (Link, 1807) 1, N. rufescens (A. Adams, 1853) 1, N. takiisaoi (Kuroda, 1959) 2, N. waikikiensis (Pilsbry, 1921) 2. Remarks: The genus Neocancilla is one of the few lineages the composition of which is left rather unchanged after our phylogenetic analysis. The group is well delineated, thanks to the characteristic shell and conservative radular morphology. The radula of the West African ‘ Neocancilla ’ hebes depicted by Cernohorsky (1991) rather resembles the radula of Strigatellinae and led Cernohorsky to question the relation of that species to the Indo-Pacific Neocancilla. Here, we remove this species from Neocancilla and provisionally place it in Domiporta. Species of Domiporta are closest to Neocancilla primarily because of similar shell proportions and sculpture pattern. Species of Neocancilla, however, can be distinguished by the shell base and siphonal canal, which form a continuous, usually slightly convex, outline with no concave waist. Besides, there are consistent differences between the two genera in radular morphology: in Neocancilla, the radula has few short, blunt and robust cusps on the rachidian and laterals, while in Domiporta the rachidian and laterals bear multiple pointed cusps that are not differentiated in size or shape. The Scabricola – Swainsonia grade is closely related to Neocancilla, and the species of these three genera are conchologically similar. However, shells of Scabricola and Swainsonia can be differentiated by a notably weaker sculpture, with no or very weak axial elements; besides, most species of Scabricola are wider, with a higher last adult whorl compared to Neocancilla.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFB9FFD7B20373CDD21EB590.taxon	description	(FIGS 31, 32 A – C) Type species: Voluta dactylus Linnaeus, 1767; SD, Dall (1915: 52). Synonym: Cylindromitra P. Fischer, 1884. Type species: Cylindra coronata Schumacher, 1817 (objective synonym of Voluta crenulata Gmelin, 1791); by typification of replaced name (Cylindra Schumacher, 1817, non-Illiger, 1802; itself by M) Acuticylindra Iredale, 1929. Type species: Voluta nucea Gmelin, 1791; OD. Diagnosis: Shell small to medium sized (20 – 70 mm), ovate or ovate-elongate, with very low spire and tall aperture. Protoconch cyrtoconoid, with about three smooth evenly convex whorls. Spire height attaining only about 10 – 15 % of total shell height, with only three to four whorls exposed, and a proportionally very high last adult whorl. Suture adpressed. Last adult whorl convex or strongly convex, smooth or with dominating spiral or cancellate or crenulate sculpture. Spiral elements represented by fine grooves or cords that can be fine or wide, undulating, rounded or resembling scales tiling in posterior direction. Siphonal notch distinct. Aperture elongate, usually narrow. Outer aperture lip convex, smooth or denticulate, inner lip calloused, bearing six to eight subequal columellar folds. Shell off-white, whitish or brownish or with a pattern of red or orange bands. Radula very narrow, consisting of rachidian only, bow-shaped, bearing multiple cusps, or with a single blunt medial cusp and lateral serration (Fig. 32 A – C). Distribution: Indo-Pacific, subtidal, mostly on coral rubble, one species occurs on mud. Species included: Pterygia arctata (G. B. Sowerby II, 1874) 3, P. conus (Gmelin, 1791) 3, P. crenulata (Gmelin, 1791) 2, P. dactylus (Linnaeus, 1767) 1, P. fenestrata (Lamarck, 1811) 1, P. gorii (Turner, 2007) 3, P. japonica Okutani & Matsukuma, 19823, P. jeanjacquesi Bozzetti, 20103, P. morrisoni Marrow, 20163, P. nucea (Gmelin, 1791) 2, P. purtymuni Salisbury, 19983, P. scabricula (Linnaeus, 1767) 3, P. sinensis (Reeve, 1844) 1, P. undulosa (Reeve, 1844) 3. Remarks: The genus Pterygia is morphologically distinctive thanks to its ovate shell with multiple columellar folds, and monoserial radula. In the present study, we leave the contents of the genus unchanged; however, some species currently classified in other genera (e. g. Scabricola potensis) may actually belong to Pterygia. Conical shells resembling those of Pterygia are found in Imbricaria and Imbricariopsis gen. nov., which are, however, phylogenetically very distant from Pterygia and are characterized by a triserial radulae.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFB8FFD5B20A702FD025B614.taxon	description	(FIGS 32 D – F, 33) Type species: Mitra contracta Swainson, 1820; SD, Herrmannsen (1847: 110). Synonyms: Dibaphus Philippi, 1847. Type species: Mitra edentula Swainson, 1823; M. Mutyca H. Adams & A. Adams, 1853. Type species: Mitra ancillides Broderip, 1836; SD, Wenz (1943: 1296). Mitroidea Pease, 1865. Type species: Mitroidea multiplicata Pease, 1865; M.? Mauritia H. Adams, 1869. Type species: Mauritia barclayi H. Adams, 1869; M. Diagnosis: Shell small to large (15 – 150 mm), fusiform to elongate-fusiform. Protoconch pointed, narrowly conical, of three or more slightly convex, smooth and glossy whorls. Teleoconch whorls usually flattened in outline, with orthoconoid or acuminate spire. Sculpture variable, from indistinct to strong and deep groves separating flattened or rounded, sometimes undulating, cords. Siphonal canal robust, short to moderately long, sometimes with distinct fasciole, giving siphonal canal a recurved appearance. Aperture elongate, its outer lip often thickened in its adapical portion. Margin of outer lip smooth or bearing fine denticles throughout its length. Inner lip with four to five delicate folds of subequal strength. Shell pale, usually with a pattern of brown bands and blotches. Radula triserial, with narrow rachidian and laterals of varying width. Rachidian with five cusps of which a central unpaired cusp greatly exceeds the others in strength (Fig. 32 D – F). Laterals bearing six to 12 cusps, second to fourth from rachidian cusp being strongest, and succeeding ones gradually diminishing. Distribution: Indo-Pacific, intertidal and upper subtidal depths, rocky shores and reef crevices. Species included: Nebularia acuminata (Swainson, 1824) 1 comb. nov., N. aegra (Reeve, 1845) 3 comb. nov., N. ancillides (Broderip, 1836) 3 comb. nov., N. baerorum (Poppe & Tagaro, 2010) 3, N. bellula (A. Adams, 1853) 3 comb. nov., N. chrysostoma (Broderip, 1836) 3, N. coarctata (Reeve, 1844) 3, N. contracta (Swainson, 1820) 1, N. deynzeri (Cernohorsky, 1980) 3 comb. nov., N. dondani (Cernohorsky, 1985) 3 comb. nov., N. edentula (Swainson, 1823) 3 comb. nov., N. eremitarum (Röding, 1798) 1 comb. nov., N. fastigium (Reeve, 1845) 3 comb. nov., N. ferruginea (Lamarck, 1811) 2, N. gourgueti (Poppe, Salisbury & Tagaro, 2015) comb. nov., N. guidopoppei (Thach, 2016) 3 comb. nov., N. incompta (Lightfoot, 1786) 1 comb. nov., N. inquinata (Reeve, 1844) 3, N. kamehameha (Pilsbry, 1921) 3 comb. nov., N. multiplicata (Pease, 1865) 3 comb. nov., N. nebulosa (Broderip, 1836) 3 comb. nov., N. nivea (Broderip, 1836) 3 comb. nov., N. pellisserpentis (Reeve, 1844) 3 comb. nov., N. petrosa (G. B. Sowerby II, 1874) 3 comb. nov., N. pyramis (Wood, 1828) 3, N. semperi (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., N. thachi (Turner, 2007) 3, N. ustulata (Reeve, 1844) 3 comb. nov. Remarks: Our phylogenetic analysis has led to a considerable rearrangement of the genus Nebularia. The ‘ lesser’ Nebularia species (N. fraga and N. tornata groups) appeared unrelated to the true Nebularia typified by N. contracta. On the other hand, some large Mitra species (M. incompta, M. eremitarum) showed a close affinity to N. contracta, and so did ‘ Strigatella ’ acuminata. The genus Nebularia, as circumscribed herein, is supported by several distinctive morphological shell (colour pattern, a thickened outer aperture lip and an often recurved tip of the siphonal canal) and radula characters (a very strong unpaired central cusp on the rachidian). The same features characterize a number of species classically included in Mitra (M. ancillides, M. coarctata, M. edentula, M. ferruginea, M. nebulosa, M. nivea, M. ustulata), which we thus classify in Nebularia. As a consequence, we also consider the nominal genera Dibaphus, Mitroidea and Mutyca to be synonyms of Nebularia. Both shell proportions and development of sculpture vary considerably among the species of Nebularia, the two extremes being N. incompta (Fig. 33 D) and N. ancillides (Fig. 33 G). An even more cylindrical shell characterizes Mauritia barclayi, which we tentatively place in Nebularia as well.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFBAFFAAB03170BAD12CB340.taxon	description	(FIGS 34 – 36) Diagnosis: Shell small to large (15 – 125 mm) mm, biconical, fusiform to turriform, pale to dark brown due to well-developed periostracum. Shell smooth, or sculptured with fine to very strong spiral cords. Suture distinct, impressed. Spire moderately to very high; teleoconch whorls gently convex, distinctly shouldered due to pronounced spiral sculpture, or flattened. Sculpture absent, or represented by regular spiral cords, ranging from wide and flattened to rounded, or narrow, elevated, separated by wide, concave interspaces. Microsculpture of fine collabral growth lines. Siphonal canal short and stout with well-developed fasciole, to moderately long, strait and tapering. Siphonal notch rarely deep, commonly shallow or absent. Aperture elongate, varying from narrow to moderately wide; outer aperture lip evenly convex or straight in its adapical portion and strongly convex anteriorly. Inner lip with four columellar folds, the anteriormost notably weaker than succeeding. Rachidian narrow, less than half as wide as the laterals, bearing five or six cusps of varying shape (Figs 33 A – C). Laterals wide, with eight to 40 cusps, of varying strength.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFC5FFA8B3DD7575D2A1B72F.taxon	description	(FIGS 34, 35 A – C) Type species: Mitra bulimoides Reeve, 1845 [= M. glabra Swainson, 1821, after Cernohorsky (1976)]; SD, Cossmann (1899: 153). Synonym: Fuscomitra Pallary, 1900. Type species: Mitra fusca Pallary, 1900 (= Mitra cornea Lamarck, 1811); SD, Cox (1936: 66). Pallary referred to the type species as ‘ Mitra (Fuscomitra) fusca (Swainson) Reeve’, that is, the species identified by Reeve (1844) as M. fusca Swainson, 1824. Reeve did not establish a new species, and there is no such thing as ‘ M. fusca Reeve, 1844 ’. Instead, this is a case of a deliberate misidentification of a nominal species used for the fixation of the type species and, under Art. 67.13, Pallary is deemed to have established a new nominal species, M. fusca Pallary, 1900. Diagnosis: Shell medium sized to large (30 – 100 mm), fusiform to turriform, lightly sculptured, light or dark brown due to well-developed periostracum. Suture distinct, impressed. Spire high to very high; teleoconch whorls gently convex to flattened, sculptured by regular or occasional spiral grooves, or smooth, often with a microsculpture of fine collabral growth lines. Siphonal canal short with well-developed fasciole to moderately long and tapering. Siphonal notch shallow or absent. Aperture elongate, rather wide; outer aperture lip evenly convex or straight in its adapical portion and strongly convex anteriorly. Inner lip with four columellar folds, the anteriormost notably weaker. Rachidian narrow, less than half as wide as the laterals, bearing five or six cusps of varying shape (Fig. 33 A – C). Laterals wide, with eight to 40 cusps, subequal and densely set throughout, or proximal strongest and subsequent ones weakening towards tooth edge. Species included: Isara aerumnosa (Melvill, 1888) 3 comb. nov., I. aikeni (Lussi, 2009) 3 comb. nov., I. antillensis (Dall, 1889) 3 comb. nov., I. badia (Reeve, 1844) 3 comb. nov., I. beui (Thach, 2016) 3 comb. nov., I. carbonaria (Swainson, 1822) 1 comb. nov., I. chalybeia (Reeve, 1844) 3 comb. nov., I. chinensis (Gray, 1834) 3 comb. nov., I. cookii (G. B. Sowerby II, 1874) 3 comb. nov., I. cornea (Lamarck, 1811) 1 comb. nov., I. declivis (Reeve, 1844) 3 comb. nov., I. gabonensis (Biraghi, 1984) 3 comb. nov., I. glabra (Swainson, 1821) 1 comb. nov., I. goreensis (Melvill, 1925) 3 comb. nov., I. joostei (Lussi, 2009) 3 comb. nov., I. lenhilli (Petuch, 1988) 3 comb. nov., I. midwayensis (Kosuge, 1979) 3 comb. nov., I. nigra (Gmelin, 1791) 1 comb. nov., I. pele (Cernohorsky, 1970) 3 comb. nov., I. peterclarksoni (Marrow, 2013) 3 comb. nov., I. picta (Reeve, 1844) 3 comb. nov., I. slacksmithae (Marrow, 2013) 3 comb. nov., I. straminea (A. Adams, 1853) 1 comb. nov., I. swainsonii (Broderip, 1836) 3 comb. nov., I. turtoni (E. A. Smith, 1890) 3 comb. nov., I. ulala (Garcia, 2011) 3 comb. nov. Distribution: Indo-Pacific, West Africa and Mediterranean, Caribbean, from upper subtidal to upper bathyal depths. Remarks: The type species of Isara, M. bulimoides (Fig. 34 B), was considered by Cernohorsky (1976) a synonym of M. glabra, and indeed shows conchological resemblance to it. Cernohorsky (1976) provided no whereabouts of the type specimen of M. glabra Swainson, 1821, and designated two unnumbered figures on plate 2 of the ‘ Exotic Conchology ’ as the lectotype of M. glabra. Although this designation is nomenclaturally valid, ICZN Art. 74.4 makes it clear that this designation is to be treated as designation of the specimen illustrated. As this specimen cannot currently be traced, we herein designate the holotype of M. bulimoides NHMUK 1966656 as neotype of M. glabra. Thus, M. bulimoides Reeve, 1845 becomes an objective synonym of the M. glabra Swainson 1821. The genus Isara, as circumscribed herein, is a group largely defined based on molecular characters. Although the included species share a similarity in shell morphology (weak sculpture and, in many cases, well-developed periostracum), the shell proportions are drastically different in I. straminea and I. glabra, and their relatedness would hardly be guessed based on overall shell appearance. The sequenced Isara species altogether show an unusually broad distribution: Azores and Mediterranean (I. cornea), West Africa (I. nigra), Caribbean (I. straminea) and Western Australia (I. glabra and I. carbonaria). Based on the overall shell morphology, we tentatively assign M. chinensis to Isara, as well as an array of Australian species (M. badia, M. chalybeia, M. peterclarcksoni and M. slacksmithae). The species of Isara are commonly large, and their protoconchs are often corroded in (sub) adults; we managed to find an intact protoconch only in juveniles of I. chalybeia from southwestern Australia. It is characterized by a shape unusual for Mitridae – paucispiral, cyrtoconoid, broad, with a very small nucleus, and strongly convex whorls. Isara chalybeia lives in subtropical / warm temperate waters and we doubt that this protoconch morphology also occurs in the tropical species of Isara, which are expected to have the plesiomorphic multispiral protoconch present throughout the family. Although we assign to Isara the Eastern Pacific Mitra swainsonii, the distribution of Isara as inferred from the phylogenetic analysis does not include the East Pacific. Whereas no ‘ smooth’ Atrimitra species from the Panamic province was included in our phylogenetic analysis, the obvious resemblance between the Panamic ‘ Mitra ’ orientalis, and ‘ M. ’ caliginosa, and the Atlantic I. nigra and I. cornea, should also be noted. Another species which also remains unstudied, and may prove to be related to Isara, is Atrimitra idae. At present, we treat these New World species as Atrimitra, pending availability of molecular data.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFC7FFAEB04F7257D192B295.taxon	description	(FIGS 35 D – F, 36) Type species: Mitra sulcata Swainson, 1825; OD. Diagnosis: Shell small to large (15 – 125 mm), fusiform, with white or light background colour and contrasting brown lines on crests of spiral cords. Protoconch smooth, with slightly convex or flattened whorls, either pointed, narrowly conical, of about three whorls, or bulbous, of about two whorls. Suture indistinct. Spire moderately high, spire whorls evenly convex, sometimes shouldered in appearance due to strong elevated spiral cords that are rather sharp and widely set. Interspaces between spiral cords sculptured with fine, dense riblets, or smooth with microsculpture of very fine growth lines. Shell base triangular with straight, tapering siphonal canal not delineated from shell base. Siphonal notch deep. Aperture high, about half of shell height, narrow, parallel-sided. Outer aperture lip very gently convex, smooth. Inner lip with three fine, oblique columellar folds. Thin light brown periostracum sometimes present. Radula with narrow rachidian and laterals two to four times wider than rachidian (Fig. 35 D – F), both multicuspidate, either with dense pointed cusps of subequal strength or with one cusp on laterals notably stronger than others and rachidian bearing symmetrical paired cusps also differentiated in length and strength. Distribution: Tropical East Pacific, Caribbean, subtidal and bathyal depths. Species included: Subcancilla attenuata (Broderip, 1836) 1, S. belcheri (Hinds, 1843) 3 comb. nov., S. calodinota (S. S. Berry, 1960) 3, S. candida (Reeve, 1845) 3, S. directa (Berry, 1960) 3, S. edithrexae (Sphon, 1976) 3, S. erythrogramma (Tomlin, 1931) 1, S. funiculata (Reeve, 1844) 3, S. gigantea (Reeve, 1844) 3, S. haneti (Petit de la Saussaye, 1852) 3, S. hindsii (Reeve, 1844) 3, S. joapyra Simone & Cunha, 20123, S. larranagai (Carcelles, 1947) 3, S. leonardi (Petuch, 1990) 1 comb. nov., S. leonardhilli Petuch, 19872, Subcancilla lindae Petuch, 19873, S. lopesi Matthews & Coelho, 19693, S. malleti (Petit de la Saussaye, 1852) 3, S. phorminx (S. S. Berry, 1969) 3, † S. scrobiculata (Brocchi, 1814) 3, S. sulcata (Swainson, 1825) 2. Remarks: Cernohorsky (1991) remarked that the placement of the New World S. sulcata and of Indo-Pacific species in the same genus Subcancilla was inconsistent with their disparate radular morphology; however, no alternative was suggested. Thorsson & Salisbury (2008: 3) stressed that the classification of the Panamic and Caribbean species of Subcancilla, including the type species S. sulcata, has not been addressed sufficiently, and that the ‘ exact placement of Subcancilla into the family Mitridae remain [ed] to be absolutely determined’. Based on our phylogenetic analysis, we demonstrate a close affinity of three East Pacific and one Caribbean species of Subcancilla, and reconsider the contents of the genus, so that only New World species are here assigned to it. The species of Subcancilla can be readily distinguished from other American mitrids by their characteristic sculpture of narrow, elevated, usually widely interspaced, spiral cords, often bearing a brown line on their crests; the same sculptural pattern characterizes Indo-Pacific species in the genera Domiporta and Imbricaria. Members of these three genera are also characterized by similar shell proportions, so that their shells share the same characteristic habitus that has long confused conchologists. Our results demonstrate that the three genera represent three major phylogenetic lineages of Mitridae in the subfamilies Isarinae, Mitrinae and Imbricariinae, and their resemblance is obviously a result of convergence. The radula differs remarkably in the three studied species, with those of S. attenuata and S. cf. leonardhilli being overall unremarkable underived mitrid radulae, whereas that of S. erythrogramma demonstrates signs of cusp differentiation, resembling imbricariine radulae. The small and unconspicious radula of S. sulcata pictured by Cernohorsky (1991: fig. 79) undoubtedly belongs to the first type.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFC1FFAEB3FE74D3D79CB128.taxon	materials_examined	Type species: † Voluta carolinensis Conrad 1840: 387; M. Diagnosis: As no material of Pleioptygma was available for the present study, we do not provide our own diagnosis to this enigmatic and poorly studied genus, and refer to Quinn (1989). Species included: Pleioptygma helenae (Radwin & Bibbey, 1972) 2. Distribution: Caribbean, subtidal to upper bathyal depths. Remarks: For remarks on the placement of Pleioptygma, see section ‘ Implications of morphological studies for the taxonomy of the Mitridae’ below.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFC1FFAFB0AB77C7D629B5A4.taxon	description	(FIGS 37, 38 A – C) Type species: Mitra peculiaris Reeve, 1845; OD, herein. Diagnosis: Shell small (7 – 20 mm), elongate-fusiform, with characteristic pattern formed by light adapical portions of teleoconch whorls, gradually transiting to yellow or orange peripheries. Protoconch narrowly conical or cyrtoconoid, of three to almost five, evenly convex, smooth and glossy whorls. Suture distinct, slightly impressed. Early spire whorls with flattened or slightly convex outlines, later gently convex or distinctly carinated at the suture. Except for subsutural carina, sculpture limited to very weak spiral grooves on shell base and siphonal canal, thus the shell appears smooth. Siphonal canal tapering, siphonal notch shallow or indistinct. Aperture elongate, rather narrow; outer lip smooth, convex. Inner lip with four to five fine oblique columellar folds. Rachidian narrow, bearing either six cusps of comparable size, or four cusps with two medial notably stronger than two laterals. Laterals wide, bearing multiple cusps, of which second and third from rachidian are strongest. Distribution: Indo-Pacific, upper subtidal depths, reef crevices and coral rubble. Species included: Carinomitra peculiaris (Reeve, 1845) 1 comb. nov., C. saltata (Pease, 1865) 2 comb. nov., C. typha (Reeve, 1845) 1 comb. nov. Etymology: The name Carinomitra refers to the presence of the distinct keel (Latin carina) in at least two species of this newly recognized lineage, including the type species C. peculiaris. Remarks: The genus Carinomitra is conchologically distinctive due to the carinated or adapically inflated adult whorls and the characteristic colour pattern. The radulae of the three studied species exhibit notable disparity. In C. peculiaris, the rachidian bears multiple cusps with a central unpaired cusp; conversely, in C. saltata and C. typha, the unpaired central cusp of the rachidian is absent, and the paired central cusps greatly exceed the paired lateral cusps. This variation in radula morphology parallels that observed in Imbricaria and Cancilla (see discussion below). GENUS CONDYLOMITRA FEDOSOV, HERRMANN,	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFC0FFACB0B573DDD091B425.taxon	description	(FIGS 38 D, E, 39) Type species: Mitra tuberosa Reeve, 1845; OD, herein. Diagnosis: Shell small (10 – 25 mm), ovate to almost conical, heavily sculptured, with pointed conical apex. Protoconch pointed, narrowly conical, of about three very slightly convex, smooth whorls. Spire whorls convex; last adult whorl proportionally very high, sculptured with densely set, gemmate spiral cords, one to three adapical cords very strong, forming prominent, spirally elongated rounded tubercles, directed slightly upwards. Siphonal canal very short, with well-developed fasciole, siphonal notch distinct. Aperture elongate, rather narrow, widening anteriorly. Outer apertural lip gently convex or straight, inner lip with four closely set columellar folds. Radula very narrow, monoserial, with rachidian bearing strong, unpaired medial cusp, flanked with two to eight smaller pointed cusps on each side. Proboscis dorso-ventrally flattened. Distribution: Indo-Pacific, intertidal and upper subtidal depths, mostly on hard substrates. S p e c i e s i n c l u d e d: C o n d y l o m i t r a b e r n h a r d i n a (Röding, 1798) 1 comb. nov., C. tuberosa (Reeve, 1845) 1 comb. nov. Etymology: The name Condylomitra combines the Latin word condylus, meaning knob, with the suffix Mitra, in reference to the characteristic shell sculpture of tubercles that is shared by the species in this newly recognized lineage. Gender feminine. Remarks: Although each on a quite long branch, C. bernhardina and C. tuberosa form a clade in our molecular trees and share features of the shell and radula, and the characteristic sculpture pattern allows a rather easy recognition of this group. Because of their superficial similarity, we initially considered Strigatella lugubris and ‘ Mitra ’ pudica to be possibly related to Condylomitra; however, the former is closely related to Strigatella coronata in the Strigatella clade, and the latter, although it was not sequenced, is excluded from Condylomitra by its triserial radula.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFC3FFACB0497280D7A9B611.taxon	materials_examined	Type species: Magnamitra sandrogorii Huang & Salisbury, 2017; OD. Diagnosis: As no material on M. sandrogorii was available for the present study, we do not provide our own diagnosis for this recently described genus, and refer to Huang & Salisbury (2017). Species included: Magnamitra sandrogorii Huang & Salisbury, 20172. Distribution: Indo-Pacific, bathyal depths. Remarks: See remarks under Calcimitra.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFC3FFADB0A070A1D167B1F0.taxon	description	(FIG. 40 A – C) Type species: Mitra idae Melvill, 1893; OD. Diagnosis: As no material on Atrimitra was available for the present study, we do not provide a diagnosis for this poorly studied genus. Species included: Atrimitra caliginosa (Reeve, 1844) 3 comb. nov., A. catalinae (Dall, 1919) 2 comb. nov., A. effusa (Broderip, 1836) 3, A. idae (Melvill, 1893) 3, A. orientalis (Griffith & Pidgeon, 1834) 3 comb. nov., A. semigranosa (Martens, 1897) 3 comb. nov. Distribution: Tropical East Pacific, subtidal depths. Remarks: see remarks under Isara.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFC2FFADB21577C5D643B180.taxon	description	(FIG. 40 D) Type species: Vicimitra prosphora Iredale, 1929; M. Diagnosis: As no material on V. prosphora was available for the present study, we do not provide a diagnosis for this genus. Species included: Vicimitra prosphora Iredale, 19293 (Fig. 40 D). Distribution: New South Wales, Australia. Remarks: See remarks under Quasimitra.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFC2FFA2B0B877C5D2E8B4F4.taxon	description	(FIGS 38 E, 40 E, F) Type species: Mitra espinosai Sarasúa, 1978; OD. Diagnosis: Shell small to medium sized (15 – 45 mm), broadly fusiform, light brown, sometimes with occasional white flakes. Protoconch white, pointed, of about 2.5 smooth glossy whorls. Suture distinct, impressed. Teleoconch whorls slightly convex, sculptured with fine, low, evenly interspaced spiral cords. Siphonal canal stout, tapering, siphonal notch shallow. Aperture elongate, outer lip convex, thickened adapically. Inner apertural lip with five fine columellar folds, weakening gradually towards siphonal canal. Radula triserial, rachidian narrow, with eight pointed cusps, two medial longest, those situated laterally becoming gradually weaker. Laterals with 15 + cusps, strong and long compared to length of tooth base. Species included: Probata barbadensis (Gmelin, 1791) 1 comb. nov., P. espinosai (Sarasúa, 1978) 3. Remarks: Mitra espinosai, the type species of Probata, has been considered a junior synonym of Mitra barbadensis by Cernohorsky (1991). Indeed, the immature holotype of the former very much resembles M. barbadensis and, even if not conspecific, they are definitely closely related. Thus, we tentatively allocate the name Probata to the M. barbadensis clade. The affinities of Probata remain unclear and obviously need further studies. In terms of sculpture pattern, typical Atrimitra, as typified by Mitra idae, resembles Probata; however, the latter differs notably in shell proportions and number of columellar folds. The overall plesiomorphic radula of P. barbadensis (Fig. 38 F) resembles the radula of many other lineages of Mitridae and gives no clue as to where the genus should be assigned.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFCDFFA2B3C672C9D62DB1EB.taxon	description	(FIG. 40 G) Type species: Mitra florida Gould, 1856; OD. Diagnosis: As no material on Dibaphimitra florida was available for the present study, we do not provide our own diagnosis to this poorly studied genus, and refer to Cernohorsky (1970). Species included: Dibaphimitra florida (Gould, 1856) 2. Distribution: Caribbean, subtidal to upper bathyal depths. Remarks: The radula of D. florida pictured by Cernohorsky (1970) is overall a typical mitrid radula, with a narrow rachidian and very wide nulticuspidate laterals. Although this morphology unequivocally places Dibaphimitra in Mitridae, it is inconclusive regarding the subfamilial placement of D. florida. More studies are needed to characterize the affinities of this enigmatic Caribbean lineage. DESCRIPTION OF NEW SPECIES PROFUNDIMITRA TAYLORI FEDOSOV, HERRMANN,	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFCDFFA3B0BD779ED1A1B5A8.taxon	description	(FIGS 17 G, 18 A – C) Type data: Holotype: MNHN IM- 2013 - 40651, Papua New Guinea, North of Rabaul, BIOPAPUA Stn CP 3671, 04 ° 04 ′ S, 151 ° 56 ′ E, 585 – 601 m, lv, 24.15 mm (Figs 17 G, 18 A, B). Paratype: MNHN IM- 2013 - 58429, KAVIENG 2014 Stn CP 4449, 02 ° 10 ′ S, 150 ° 11 ′ E, 623 – 908 m, lv, 35.5 mm. Other material examined: Papua New Guinea, BIOPAPUA Stn 3736, off Lancasay, 08 ° 14 ′ S 150 ° 32 ′ E, 760 – 769 m, 1 lv (21.5 mm); PAPUA NUIGINI Stn CP 3979, North of Bagabag Is., 04 ° 44 ′ S 146 ° 11 ′ E, 540 – 580 m, 1 lv (MNHN IM- 2013 - 19698, 25.75 mm); Stn CP 3994, North from Cape King William, 06 ° 00 ′ S 147 ° 35 ′ E, 648 – 652 m, 1 lv (MNHN IM- 2013 - 19833, 33.6 mm); MADEEP Stn DW 4321 08 ° 39 ′ S 151 ° 47 ′ E, 612 m, 1 lv (MNHN IM- 2013 - 45873).? NEW CALEDONIA, BATHUS 3 Stn DW 790, 23 ° 49 ′ S 169 ° 48 ′ E, 685 – 715 m, 17.9 mm; Norfolk Ridge, BATHUS 4 Stn DW 920 18 ° 45 ′ S 163 ° 17 ′ E, 610 – 620 m, 15.1 mm (Fig. 18 C). Description: Shell medium sized (holotype 24.15 mm, paratype 35.5 mm), fusiform, with high, cyrtoconoid, slightly tilted spire and rather short siphonal canal. Protoconch pointed, subcylindrical, of two or more whorls (corroded in all specimens studied). Teleoconch of 7.5 gently convex whorls, suture linear, distinctly impressed. First teleoconch whorls with reticulate sculpture formed by four low, very weak, rounded spiral cords, intersected by even weaker rounded spiral folds. Interstices rather deep, forming characteristic pattern of four spirally aligned punctures, well pronounced on two succeeding whorls and rather gentle, widely spaced on later whorls. Penultimate whorl with three distinct spiral rows of punctures on adapical portion, followed by three very weak rows below. Last adult whorl evenly convex, gradually extending to rather short, straight tapering siphonal canal. Adapical portion and periphery of last adult whorl sculptured with ten spiral rows of punctures. On shell base spiral rows of punctures becoming denser, as well as punctures themselves merging to form narrow grooves on abapical part of shell base and strong, oblique grooves on siphonal canal. Siphonal canal slender, moderately long, tapering, not notched at its tip. Aperture rather low, lanceolate, its outer lip thin, evenly convex. Inner apertural lip evenly concave in its adapical half and straight below, with three oblique columellar folds at mid-height, slightly deepened in aperture and weak in appearance. Shell colouration uniformly pale-yellowish; microsculpture of very fine collabral growth lines. Radula (of holotype) 0.7 – 0.75 mm long, 0.14 mm wide, consisting of about 85 rows. Rachidian about 40 µm and laterals 45 µm wide, all three similar, with evenly convex anterior margin, bearing 12 – 13 moderately long, pointed subequal cusps, those at teeth margins smaller than others. Distribution: Bismarck Sea off mainland New Guinea and New Ireland; East of New Caledonia, Norfolk Ridge, 580 – 760 m. Etymology: The species named after our friend and colleague John D. Taylor of the NHMUK in recognition of his immense contribution to malacology, and in particular to our knowledge of mitrid feeding biology. Remarks: Profundimitra taylori sp. nov. can be readily distinguished from its congeners by its almost smooth shell with faint columellar folds; both features are rather characteristic of Eumitra. Members of the latter genus, however, are characterized by more convex whorls, often with subcylindrical early teleoconch whorls. Smooth species of Calcimitra, such as C. labecula, resemble P. taylori sp. nov. in shell proportions and sculpture; however, Calcimitra species can be recognized by their generally larger shells with strong columellar folds and, typically, by the presence of a siphonal notch. Finally, the smooth shallow-water Quasimitra sarmientoi, although close to P. taylori sp. nov. in shell proportions and lack of sculpture, can be differentiated by the intensely coloured shell and distinct siphonal notch in adult specimens. Apart from the mentioned shell characters, the characteristic radula with similar laterals and rachidian differentiates P. taylori sp. nov. and its congeners from most other mitrids. FUSIDOMIPORTA PONDERI FEDOSOV, HERRMANN,	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFCCFFA0B22773C1D191B434.taxon	description	(FIGS 17 I, 18 I, K) Type data: Holotype: MNHN IM- 2013 - 40665, Papua New Guinea, BIOPAPUA Stn CP 3692, SE of Manus I., 02 ° 10 ′ S, 147 ° 19 ′ E, 408 – 448 m, lv, 25.2 mm (Fig. 18 I, K). Paratype: BIOPAPUA Stn CP 3629, Gulf of Huon, South of Lae, 06 ° 57 ′ S, 147 ° 08 ′ E, 240 – 269 m, dd, 26.15 mm. Other material examined: BIOPAPUA Stn DW 3735, 08 ° 15 ′ S, 150 ° 34 ′ E, 450 – 500 m, 21.4 mm; Stn data lost, 28.5 mm; Stn DW 3771, 05 ° 34 ′ S, 151 ° 33 ′ E, 295 – 422 m, 9.2 mm. Description: Shell medium sized (holotype 25.2 mm), fusiform, heavily sculptured, with rather high spire and elongated aperture. Protoconch pointed, cyrtoconoid, of three or more glossy, convex whorls (usually strongly corroded or missing). Teleoconch of six or more slightly convex whorls, suture distinctly canaliculated. Subsutural shelf well developed, rather narrow, slightly inclined towards axis, bordered by projecting margin of adapical spiral cord. Early teleoconch whorls corroded, third and subsequent teleoconch whorls with five very strong, elevated, gemmate spiral cords separated by deep interspaces. Spiral cords subequal, adapical and abapical cords slightly weaker than intermediate ones, microsculpture of fine growth lines in interspaces between spiral cords; abapical cord of previous whorl partly superposed by adapical cord of succeeding whorl. Gemmules on spiral cords axially aligned, connected by low and weak axial riblets in interspaces between spiral cords. Spiral cords become distinctly beaded on penultimate and last adult whorls, forming rows of closely set, rounded, glossy granules. Five spiral cords on adapical portion of last adult whorl succeeded by six cords on shell base and five cords on siphonal canal, of which last three oblique, weaker than preceding, irregularly gemmate. Siphonal canal delineated from shell base by slightly widened interspace with low wavy spiral thread. Siphonal canal rather long and stout, slightly recurved. Aperture elongate, its outer lip gently convex throughout its height, wavy, reflecting arrangement of spiral cords. Inside of outer lip with eight strong lirae. Inner lip strongly calloused, reflected, retaining weak reticulate sculpture adapically, with three strong oblique columellar folds at mid-height. Shell uniformly white or pale. Radula of holotype 0.76 – 0.8 mm long, 0.19 – 0.20 mm wide, composed of about 65 transverse rows of uniform, delicate, multicuspidate teeth. Rachidian about 60 µm wide, with 17 cusps, of which two marginals very weak, and the others subequal, long and pointed. Laterals about 50 µm wide, with slightly convex anterior margin, bearing 14 – 15 pointed, rather long cusps. Distribution: Papua New Guinea, North of main island, 269 – 450 m. Etymology: The species is named after another giant in malacology, Winston Ponder of the AMS, whose early work, including his studies on mitriform families, set a milestone in our understanding of neogastropod evolution. Remarks: The identification of Fusidomiporta ponderi sp. nov. is uneasy, mainly because of its striking resemblance to (still undescribed) species of Profundimitra gen. nov., represented in our analysis by specimens IM- 2007 - 35618 and IM- 2007 - 35711 (both mislaid). The single live-taken specimen of F. ponderi sp. nov. seems to have a proportionally slightly lower spire and a somewhat stepped appearance of spire whorls, with more elevated spiral cords. At the same time, the distinctive and heavy sculpture pattern makes it quite recognizable from among most Mitridae. In shell proportions and whorl outline, F. ponderi sp. nov. is very close to F. suturata; however, the latter has lower and wider spiral cords with narrower grooves between them. Some deep-water species of Gemmulimitra gen. nov. have a somewhat similar, beaded sculpture, which however is nowhere as strong as in F. ponderi sp. nov. Besides, Gemmulimitra boucheti has a proportionally broader shell with more convex whorls compared to F. ponderi sp. nov. On the contrary, G. neocaledonica sp. nov. has a rather biconical shell with subcylindrical whorls and a rather reticulate sculpture pattern. However, at this moment, many of the deep-water mitrids are still represented in our data set by too few specimens to assess their intraspecific variation and elaborate robust species delimitation criteria. GEMMULIMITRA NEOCALEDONICA FEDOSOV,	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
03908790FFCFFFB8B3C2724DD27DB777.taxon	materials_examined	Type data: Holotype: MNHN IM- 2013 - 40655, New Caledonia, Kouakoué Canyon, EXBODI Stn 3821, 21 ° 53 ′ S, 166 ° 50 ′ E, 211 – 440 m, lv, 24.25 mm (Fig. 21 A, B). Paratypes: New Caledonia, EXBODI, Stn 3800, East coast off Thio, 21 ° 32 ′ S, 166 ° 22 ′ E, 279 – 282 m, 1 lv (Paratype 3, 21.5 mm); Stn 3807, East coast off Toupeti, 21 ° 43 ′ S, 166 ° 36 ′ E, 352 – 372 m, 1 lv (Paratype 2, 26.9 mm); Stn 3810, East coast off Toupeti, 21 ° 44 ′ S, 166 ° 38 ′ E, 384 – 385 m, 2 lv (Paratype 1, 27.35 mm; Paratype 4, 24.0 mm); Stn 3814, Passe du Solitaire, 21 ° 49 ′ S, 166 ° 44 ′ E, 331 – 344 m, 1 lv (Paratype 5, 20.4 mm). Other material examined: New Caledonia, EXBODI Stn 3795, 21 ° 32 ′ S, 166 ° 21 ′ E, 240 - 245 m, 1 lv (16.4 mm); Stn 3800, 21 ° 32 ′ S, 166 ° 22 ′ E, 279 - 282 m, 12 lv (16.7, 17.1, 17.3, 15.0, 16.2, 16.2, 16.55, 16.8, 15.05, 23.1, 17.65, 17.8 mm); Stn 3805, 21 ° 42 ′ S, 166 ° 34 ′ E, 302 m, 1 lv (20.9 mm); Stn 3806, 21 ° 42 ′ S, 166 ° 34 ′ E, 307 – 309 m, 7 lv (22.0, 18.4, 15.9, 16.3, 14.0, 9.15, 9.8 mm); Stn 3807, 21 ° 43 ′ S, 166 ° 36 ′ E, 352 – 372 m, 11 lv (22.6, 20.1, 21.05, 20.7, 20.0, 19.2, 17.4, 20.9, 18.1, 18.2, 19.65 mm); Stn 3810, 21 ° 44 ′ S, 166 ° 38 ′ E, 384 – 385 m, 8 lv (25.5, 25.85, 29.3, 25.5, 27.4, 21.65, 25.5, 23.3 mm); Stn 3814, 21 ° 49 ′ S, 166 ° 44 ′ E, 331 – 344 m, 6 lv (22.8, 21.5, 20.6, 21.6, 18.9, 14.1 mm); Stn 3821, 21 ° 53 ′ S, 166 ° 50 ′ E, 211 – 440 m, 2 lv (25.85 and 25.15 mm); Stn 3823, 21 ° 55 ′ S, 166 ° 55 ′ E, 246 – 255 m, 2 lv (15.5 and 14.5 mm); Stn 3825, 21 ° 58 ′ S, 166 ° 59 ′ E, 349 – 405 m, 1 lv (16.2 mm); Stn 3828, 22 ° 00 ′ S, 167 ° 01 ′ E, 300 – 302 m, 1 lv (16.4 mm); Stn 3829, 22 ° 02 ′ S, 167 ° 05 ′ E, 350 – 360 m, 1 lv (23.0 mm); TERRASSES Stn 3095, 22 ° 02 ′ S, 167 ° 06 ′ E, 320 – 380 m, 1 lv (21.5 mm). Papua New Guinea, BIOPAPUA Stn CP 3634, 07 ° 29 ′ S, 147 ° 31 ′ E, 279 – 290 m, 1 dd (21.75 mm). Description: Shell small to medium sized (holotype 24.25 mm), elongate-fusiform to almost biconical, heavily sculptured, with narrow aperture and stepped appearance of late teleoconch whorls. Protoconch pointed, cyrtoconoid, translucent, of three or more slightly convex, glossy whorls. Protoconch / teleoconch transition distinct. Teleoconch of about nine whorls; suture canaliculated. Subsutural ramp forming distinct shelf, flat or even slightly inclined inside, giving teleoconch whorls a subcylindrical profile. Whorl periphery slightly convex, early spire whorls sculptured with three strong, closely set, spiral cords overriding low indistinct axial ribs to form series of prominent, axially aligned, beads. On third teleoconch whorl, adapical spiral cord splits into two, and succeeding spire whorls sculptured with four gemmate cords, adapical cord bordering subsutural ramp wavy, slightly narrower than others, delineated from second cord by narrower interspace. On later teleoconch whorls, interspaces between gemmate spiral cords gradually broadening, axial ribs becoming more distinct and sculpture rather reticulate. Last adult and penultimate whorls with 26 axial ribs each. Interspaces between axial ribs and spiral cords forming deep quadrangular depressions with microsculpture of fine co-axial growth lines, sometimes retaining fragments of periostracum forming squamiform projections. Shell base extended to moderately long, tapering, slightly notched siphonal canal. Four gemmate spiral cords on adapical portion of last adult whorl, and 12 cords on shell base and siphonal canal. Aperture narrow, elongated; outer lip wavy, convex in adapical portion and straight below mid-height. Inside of outer lip smooth. Inner lip strongly calloused, sometimes reflected, retaining weak reticulate sculpture adapically, with four closely set fine columellar folds at mid-height. Coloration uniformly cream, protoconch white. Radula (of Paratype 5) 0.62 mm long, 0.13 mm wide, of about 75 rows. Rachidian narrow, about 20 µm wide, bearing eight strong, moderately long, pointed cusps. Laterals attaining 47 µm in width, with straight anterior margin bearing 16 + strong, pointed, rather widely set cusps. Distribution: New Caledonia, Papua New Guinea, 255 – 384 m. Etymology: The species epithet refers to the type locality. Remarks: While the specimens from New Caledonia show a fine reticulate sculpture pattern on the late teleoconch whorls, the specimen from Papua New Guinea is characterized by denser axial ribs and in overall shell morphology is somewhat intermediate between such typical G. neocaledonica sp. nov. and Fusidomiporta ponderi sp. nov. The latter species is the one that resembles most G. neocaledonica sp. nov., but it can be differentiated by its notably stronger spiral cords and rather fusiform shell. The combination of elongate-biconical shell with subcylindrical whorl profile and characteristic reticulate pattern on late whorls allows for easy recognition of G. neocaledonica sp. nov. Other species of Gemmulimitra gen. nov. differ either in shell shape or in sculpture pattern, and G. avenacea, despite being the closest to G. neocaledonica sp. nov. in our multi-gene analyses, shows no morphological resemblance to it whatsoever. IMPLICATIONS OF MORPHOLOGICAL STUDIES FOR THE TAXONOMY OF THE MITRIDAE According to our earlier results (Fedosov et al., 2015), the families Mitridae, Charitodoronidae and Pyramimitridae form a well-supported phylogenetic group, which we designate as a superfamily Mitroidea, although they are rather heterogeneous morphologically, as summarized in Table 4. The relationships between the three mitroidean families are clearly established, but the affinities of Mitroidea remain uncertain. The phylogenetic analysis of Fedosov et al. (2015) suggested that Mitroidea may be a sister group to Conoidea; this affinity, however, is not supported by any shared morphological features and needs to be further investigated. The morphological distinctiveness of the Mitridae has been appreciated by many authors and, when anatomical descriptions scattered in many papers are brought together, the family ranks among the better studied among neogastropods. Beside multiple illustrations of radulae (Risbec, 1928; Cernohorsky, 1970, 1976, 1991), data on the general body morphology and / or foregut anatomy are available, among Mitrinae, for Mitra mitra (Ponder, 1972), Ziba carinata (Simone & Turner, 2010), Pseudonebularia cucumerina (Risbec, 1928) and Episcomitra zonata (Vayssière, 1901); among Strigatellinae, for Strigatella paupercula (Ponder, 1972), S. retusa and S. scutulata (Risbec, 1928); among Imbricariinae, for Imbricariopsis conovula (Ponder, 1972); among Cylindromitrinae, for Pterygia crenulata (Risbec, 1928); among Isarinae, for Isara cornea (Harasewych, 2009); and for Atrimitra idae (West, 1990), A. catalinae (West, 1991) and Condylomitra tuberosa (herein), that we treat as incertae sedis. All the species that were studied with sufficient attention revealed an epiproboscis (referred to as ‘ tube à venin ’ by Risbec, 1928), either in combination with a radula of underived morphology (in most of the species mentioned) or with a uniserial radula (Pterygia crenulata, Condylomitra tuberosa). Meanwhile, an epiproboscis is not present in the Charitodoronidae, the Neogastropoda clade closest to the Mitridae. There is no doubt that the epiproboscis represents an autapomorphy of the Mitridae and can be used for the anatomical circumscription of the family. In this connection, the morphology of the Caribbean Pleioptygma helenae, as addressed by Quinn (1989), deserves special attention. Quinn’s description of the foregut anatomy raises many questions; in particular, the position of the buccal mass is not described explicitly, as well as the nature of the ‘ proboscis bulb ’, and we believe that the homologies of these organs may have been misinterpreted by him. This assumption is further reinforced by the fact that the two specimens dissected by Quinn displayed some variation in foregut morphology, which may be an artefact of poor fixation (with the use of rhum suggested by the author!). Some parts of the description are of special interest. In particular, the ‘ proboscis bulb attached to the cephalic cavity floor by a broad, rather thick band of muscles originating in the foot’ (Quinn, 1989: 14) more likely refers to the large buccal mass and a radula / odontophore retractor muscle. Then the peculiar introvert, which ‘ invaginates and runs back through the outer tube / sheath and enters the proboscis bulb [? = buccal mass] ’ may not be anything but an epiproboscis. Undoubtedly, the anatomy of Pleioptygma helenae is peculiar and its homologies need to be clarified based on investigations of additional material. However, given its rarity and the small chance of obtaining live-taken specimens in the near future, we dare reassess the systematics of Pleioptygma based on currently available data. Several lines of evidence support a placement of Pleioptygma as a separate subfamily within the Mitridae, including (1) the mitriform shell, (2) the radula with rachidian and laterals of about equal morphology – similar to the one in Domiporta and (3) the presence of a structure that, based on topology and morphology, is closely comparable to the mitrid epiproboscis, and most likely is an epiproboscis. EVOLUTION OF RADULAR MORPHOLOGY IN THE MITRIDAE Our studies revealed only two major radula types in the family Costellariidae, with a very simple scenario of two independent transitions from plesiomorphic to derived (Fedosov et al., 2017). The situation is incomparably more complex in the Mitridae. Here, we recognize six major types of radula; each is referred to a genus that typically represents it (Fig. 41). 1 – Mitra type: radula with laterals notably wider than rachidian tooth and bearing multiple equal or subequal cusps. This underived radula morphology is widespread across the Mitridae tree. It characterizes the family Charitodoronidae, most Mitrinae (except the Domiporta group and some Pseudonebularia), most Isarinae, as well as the incertae sedis genera Probata, Carinomitra, Atrimitra (Cernohorsky, 1970; West 1990, 1991) and Dibaphimitra (Bayer, 1942). 1 a – Strigatella type: radula with slightly curved laterals bearing short robust cusps on their medial convex portion and lacking cusps laterally; rachidian always with central unpaired cusp. This type of radula is found in all Strigatellinae (except in the S. lugubris – S. coronata clade). The Strigatella type of radula represents a modified Mitra type, and intermediate morphologies with Strigatella - like laterals but different rachidian are found outside Strigatellinae in Neotiara nodulosa (Mitrinae) and Cancilla schepmani (Imbricariinae). 1 b – Nebularia type: a slightly modified version of the Mitra type, with a characteristic rachidian bearing only five cusps, the central unpaired cusp being notably enlarged. This type of morphology characterizes the genus Nebularia as circumscribed herein. 2 – Profundimitra type: very wide rachidian, roughly attaining the width of laterals and bearing equal number of cusps. This type of radula characterizes the mitrine genera Profundimitra and Fusidomiporta and is also found in some Domiporta (but not in its type species, D. filaris), Pseudonebularia maesta and, outside Mitrinae, in the genus Pleioptygma (Quinn, 1989). 3 – Imbricaria type: radula characterized by moderately wide multicuspidate laterals, with one of the medial cusps notably enlarged compared to the others. Cusps of laterals are also differentiated in size, with a central unpaired cusp retained, although often reduced in size. This type of radula is found in all Imbricaria species except I. fulgetrum. 3 a – Cancilla type: it resembles the Imbricaria type, but the laterals are more than twice as wide as the rachidian, and the central unpaired cusp of the rachidian is absent. This type of radula is found in Cancilla isabella (Salisbury & Huang, 2015) and other species of this genus studied herein, as well as in Imbricariopsis, Subcancilla erythrogramma and Imbricaria fulgetrum. 4 – Scabricola type: radula characterized by strongly modified laterals in which the lateralmost cusp is greatly enlarged to form a robust spine. Typically, the rachidian is comb-like (Scabricola variegata, Swainsonia spp.) but it may also bear a single strong cusp. This type of radula is found in species of Scabricola and Swainsonia. 5 – Neocancilla type: radula characterized by a rachidian with a pair of very robust and blunt central cusps, and laterals bearing few short and robust cusps in their medial portions. This type of radula is known in all species of Neocancilla. The radulae of Scabricola olivaeformis and S. coriacea have morphologies somewhat intermediate between the Scabricola type and the Neocancilla type. 6 – Pterygia type: very narrow uniserial radulae remarkable by the complete loss of laterals. Rachidians of varying morphology, but usually flattened, with serrated margins. Radulae of this type are found in all species of the genera Pterygia and Condylomitra, and in the Strigatella lugubris – Strigatella coronata clade of Strigatella. Although the diversity of radular morphologies in the family Mitridae is impressive, the understanding of its evolution is greatly hampered by homoplasies. Radula morphology does not clearly align with the inferred phylogenetic groupings of the Mitridae (Fig. 41) because of an ubiquitous retention, in advanced lineages, of the plesiomorphic state and because of multiple convergences. In 15 genera of Mitridae, all studied species presented an underived Mitra - type radula. Seven genera appeared heterogeneous in radula morphology, combining several general types. Thus, specific apomorphies can be identified in only few cases, and in even fewer cases the apomorphic state is shared by all members of the lineage. Only eight genera (Profundimitra, Fusidomiporta, Neocancilla, Pterygia, Nebularia, Condylomitra and, with some reservations, Imbricariopsis and Swainsonia) are supported by distinct apomorphies in radular morphology. Among them, only Nebularia and Neocancilla show autapomorphic radula types. Undoubtedly, the subfamily Imbricariinae shows the greatest diversity of radular morphologies, and the pattern of radular evolution is most obvious and consistent in that subfamily. The progressive differentiation of cusps on both the laterals and the rachidian is noteworthy, along with a general tendency to the reduction of the number of cusps. The radulae of Cancilla spp. are closest to the underived Mitra type, and alternate courses of radula transformation are observed in the Imbricaria and Scabricola – Neocancilla clades. In addition to a blurred phylogenetic signal of radula characters, our understanding of radula evolution in the Mitridae is impeded by a lack of evidence on the functionality of different types of radula, as there are no data on the possible adaptive value of different morphologies and selection pressures that have led to the emergence of the observed diversity. Further studies on the functional morphology and biochemistry of mitrid secretions would in this respect open new perspectives. TAXONOMY OF MITRIDAE: HISTORICAL CONSIDERATIONS AND NEW ARRANGEMENT The current Mitridae species list (as indexed in World Register of Marine Species. consulted on 19 August 2017) comprises 402 accepted Recent species, plus the three new species described in the present work. The baseline of Mitridae systematics is the fundamental revisions by Walter Cernohorsky, who first dealt with the subfamily Mitrinae (Cernohorsky, 1976), and later with the Imbricariinae and Cylindromitrinae (Cernohorsky, 1991). Cernohorsky’s monographs had a profound impact on mitrid taxonomy and systematics, despite the general lumping attitude prevailing in his time, which resulted in an extensive taxonomic graveyard for many of the species-level taxa he accepted as valid. Several cases of abusive synonymization are demonstrated in the present study. For instance, Mitra morchii A. Adams, 1855, considered by Cernohorsky a synonym of Cancilla isabella, is, based on Huang & Salisbury (2017) and on our data, not related to Cancilla or even to Imbricariinae. Another example is Mitra millepunctata G. B. Sowerby III, 1889 synonymized (Cernohorsky, 1976) with Domiporta carnicolor despite notable disparity in shell proportions, and later described as Mitra terryni Poppe, 2008 (Herrmann, 2017). Altogether the number of valid species of Mitridae was greatly diminished by Cernohorsky, but the rate of species description increased notably in the following decades and over 100 species (i. e. almost 30 % of the currently accepted number of species) were described since 1991. This burst of activity in mitrid taxonomy is also notable for being almost entirely accounted for by the amateur community: only ten out of 112 species described in the last 25 years were described by academics. Amateurs perhaps more reluctantly establish new supraspecific taxa, and only two genera Calcimitra Huang, 2011 and Magnamitra Huang & Salisbury, 2017 were established in the last 25 years, whereas 65 species described over this period were originally placed in Mitra. A concise review of changes in mitrid genus-level taxonomy was presented by Cernohorsky (1970), in a study that itself contained the description of four new genera and subgenera (Domiporta, Dibaphimitra, Neocancilla and Sohlia). In the present study, we have re-assessed the genus-level systematics of Recent Mitridae based on a combination of molecular and morphological data. The genus Charitodoron is segregated in the newly established family Charitodoronidae. The 26 genera of the revised Mitridae comprise six subfamilies: Mitrinae (with 14 genera), Strigatellinae (with the single genus Strigatella), Imbricariinae (with six genera), Cylindromitrinae (with the genera Pterygia and Nebularia), Isarinae new subfamily (with the genera Isara and Subcancilla) and Pleioptygmatinae (with the single genus Pleioptygma); seven genera, Atrimitra Dall, 1918, Carinomitra gen. nov., Condylomitra gen. nov., Dibaphimitra Cernohorsky, 1970, Magnamitra Huang & Salisbury, 2017, Vicimitra Iredale, 1929 and Probata Sarasúa, 1989 are treated as incertae sedis. Undoubtedly the most revolutionary change in the taxonomy of the Mitridae is the falling apart of Mitra with the genus in its former taxonomic extension now reassigned to 14 genera, of which six are new. Other noteworthy changes are the transfer of the formerly mitrine Nebularia to Cylindromitrinae; of Strigatella to its own, newly recognized, subfamily Strigatellinae; of the formerly imbricariine genera Ziba and Domiporta to Mitrinae; and of Subcancilla to the newly established subfamily Isarinae. The genera Mitra and Ziba now include a much reduced species diversity, as all the Indo-Pacific species earlier placed in Ziba are now transferred to Imbricaria. The contents of Imbricaria is also expanded as a result of the transfer of the Indo-Pacific species of Subcancilla, which appear unrelated to the New World species of that genus. The 32 genera now recognized in the family Mitridae increase considerably the previously accepted genus-level diversity (19 genera, including Charitodoron, indexed in WoRMS as of August 2017). Still, the placement of several sequenced species remains uncertain. Lineage 3 represented by MNHN IM- 2007 - 35623 and lineage 16 represented by MNHN IM- 2007 - 30270 did not cluster with any other lineage and were not successfully sequenced for genetic markers other than COI. Thus, they at present remain ‘ hanging’ in the list of unallocated Mitridae and may potentially represent two more new genera. PHYLOGEOGRAPHIC PATTERNS IN THE MITRIDAE It has been widely known that the Indo-West Pacific harbours the greatest diversity of mitrid species and lineages, and therefore a good representation of IP localities at various depths was a primary requirement to our sampling. Eighty eight of the 103 inferred species of Mitridae and Charitodoronidae in our data set originate from Indo-Pacific localities. Besides, three species were sampled from the Mediterranean and NE Atlantic, two from West Africa, four from the Caribbean and six from the Panamic province (Fig. 42). Twenty-three mitrid genera were sampled in the Indo-Pacific, of which 19 do not occur outside the Indo-Pacific according to our data. Moreover, the subfamilies Imbricariinae and Cylindromitrinae, as circumscribed herein, are represented solely by Indo-Pacific forms. At least three genera (Episcomitra, Isara and Ziba) are found in West Africa, with the first two also inhabiting the Mediterranean. The New World fauna of Mitridae includes at least eight genera in four subfamilies: Mitrinae (Neotiara), Strigatellinae (Strigatella), Isarinae (Isara, Subcancilla) and Pleioptygmatinae (Pleioptygma), plus the genera Atrimitra, Dibaphimitra and Probata that we treat as incertae sedis. The six genera Atrimitra, Dibaphimitra, Neotiara, Pleioptygma, Probata and Subcancilla are currently thought to be endemic to Panamic and / or the Caribbean, although a close affinity of Panamic and Caribbean species was herein confirmed only in the genera Neotiara and Subcancilla. Of all studied genera, Charitodoron probably demonstrates the narrowest range, being confined to deep waters in the Mozambique Chanel and off South Africa. According to some authors (e. g. Obura, 2012), starting in the Eocene, this area served as a main refuge for relict lineages of Tethyan origin which, for some reason, did not give rise to new radiations. The long branch that separates Charitodoron on the molecular tree, some characters such as the underived morphology (with only the radula indicating its relatedness to Mitridae), and its low diversity in the Recent fauna, all point to the relict nature of Charitodoron. The paucispiral, bulbous protoconch of Charitodoron indicates non-planktotrophic development, which correlates well with its restricted distribution. Conversely, the distribution of Isara is the widest among mitrid genera: this is the only genus recorded in more than two major zoogeographical regions of the shelf (as defined by Briggs & Bowen, 2012) – Indo-West Pacific, East Atlantic and West Atlantic (Fig. 42) – and it may be also present in the East Pacific (see remarks under Isara). Isara species contribute significantly to mitrid diversity in the peripheral Indo-Pacific (South Australia, presumably South Africa) (see Fig. 8), but they are rare and not really diverse in the Central Indo-West Pacific. In our understanding, Isara is undoubtedly an old and underived lineage of Mitridae, and this pattern may be interpreted in two different ways. According to one possible scenario, it was once widely distributed in tropical seas and subsequently was replaced by younger and derived lineages in the Central Indo-West Pacific while maintaining its diversity in peripheral areas. The other possible scenario suggests that the primarily Indo-Pacific Isara, once ‘ forced’ into subtropical waters by growing competition with younger evolutionary lineages of Mitridae, has adapted to new temperatures and, through this, was capable of spreading beyond the biogeographical limits of the Indo-Pacific. Although most known species of Strigatella occur in the Indo-Pacific, an amphi- Pacific distribution characterizes this genus, with at least one species inhabiting shallow water in the Panamic province ending up close to the Indo-Pacific species in both molecular and morphological characters. Finally, if ‘ Mitra ’ hebes is related to the species of Domiporta, as suggested by shell and radular morphology, the range of that genus would also include West Africa. The generally low overlap in regional lists of mitrid genera may be interpreted as a consequence of the relatively late major diversification of mitrid lineages, dating back to the time when modern biogeographical barriers were already established, and largely separate evolutionary radiations having taken place in (1) Late Tethys – Paratethys – Indo-Pacific, (2) West Africa and (3) the New World. We have attempted to circumscribe the bathymetric distribution of the newly delineated genera, based primarily on the locality data of sequenced specimens. Given the patchiness of our data, we arbitrarily divided the sampled depth range from 0 to about 1800 m into four depth intervals: (1) from 0 to 40 m, (2) from 41 to 80 m, (3) from 81 to 300 m and (4)> 300 m. Of the 103 species of Mitridae studied herein based on molecular characters, 72 were sampled from the 0 - to 40 - m interval, seven from the 41 - to 80 - m interval, nine from the 81 - to 300 - m interval, and 20 from depths in excess of 300 m. This accounting does not, however, strictly reflects changes in species richness with depths, as it is strongly biased by the distribution of collecting efforts (Fig. 43). Intertidal and upper subtidal zones (at diveable depths) as well as outer slopes from about 200 down to about 1800 m were sampled incomparably better than the mesophotic and abyssal zones. Therefore, our data do not necessarily demonstrate a drop in species richness in the 41 - to 300 - m interval. Nevertheless, despite the sampling bias, there is an obvious pattern of greater mitrid diversity in shallow water, decreasing with depth. This pattern can be explained by a greater diversity of habitats, often fragmented and intermixed at a small scale, in shallow-water ecosystems. Another related factor is the multitude of biological interactions in these shallow habitats, driven by the complex ecological structure of reefs and associated habitats, providing a wealth of niches to be explored by mitrids and their preys. The bathymetric distribution of the newly delineated mitrid genera is shown in Figure 43 based on our results (black vertical bars) and literature (grey vertical bars). Because of the very fragmentary data, some of the displayed results are inconsistent, like the disjunct bathymetric range of Gemmulimitra and Subcancilla. We in fact fully expect that many genera, which in our results are restricted to the 0 - to 40 - m-depth interval, in fact reach deeper, probably down to 80 – 120 m. Nevertheless, some general conclusions can be drawn from the observed bathymetric distribution of the genera. The representation of mitrid genera in shallow water is notably higher than at depths in excess of 100 – 150 m, which mirrors the distribution of species diversity discussed above. Whereas Gemmulimitra, Imbricaria, Isara, Roseomitra and Subcancilla have extensive bathymetric ranges, with congeneric species sampled from the intertidal down to 300 – 400 m, the mitrid fauna from the greatest depths is essentially represented by specialized lineages (treated here as genera), that do not occur in shallower water. The genera Calcimitra, Cancilla, Cancillopsis, Fusidomiporta and Profundimitra represented in our data set by 15 species (plus two species of Charitodoron) and the genus Eumitra (not included in our phylogenetic studies) constitute several separate radiations that thrive in the deep sea. Interpretation of the fossil mitrid taxa is difficult, primarily due to the fact that similar shell features, such as shell shape or sculpture pattern, were demonstrated to have evolved convergently in unrelated Recent lineages of the family. The inferred topology of the Mitridae tree, with unresolved early polytomy, further complicates tracing the evolution of shell features and the placement of fossil forms. Cernohorsky (1970, 1976) suggested that the most Recent genera can be traced back to Miocene; however, given the drastic rearrangement of the genus-level taxonomy of the family herein, the validity of Cernohorsky’s statement needs to be revisited. Whereas the Oligocene Clifdenia is here considered a possible relative of the Recent Calcimitra, relationships of the genera Dentimitra Koenen, 1890, Fusimitra Conrad, 1855 and Pseudocancilla Staadt, 1913 are highly speculative. The early Miocene Austroimbricaria gracilior (Ihering, 1897), with a low spire and strong columellar folds, is clearly a mitrid; however, it is unclear to which of the conchologically similar Pterygia or Imbricariopsis it may be close. WORLD DIVERSITY OF MITRIDAE Of 103 species included in our molecular data set, 89 were identified with confidence – these constitute 22 % of the described diversity of the family Mitridae. Of the remaining 15 species, four were either not identified (‘ sp. ’) or were attributed to a described species with an indication of some disparity in morphological or molecular characters (referred to as ‘ cf. ’); ten proved to be unnamed – although three of them showed some similarity to described species and were allocated to tentative species complexes (referred to as ‘ aff. ’). Remarkably, of the 60 mitrid species from the Indo-Pacific sampled in the 0 - to 40 - m-depth interval, no new species were identified with confidence. The proportion of unidentified / undescribed species grows with increasing depth and, of 20 species from depths greater than 300 m in the analysis, only 11 were identified with confidence, and seven (i. e. 35 %) were undoubtedly new, all representing previously undescribed lineages of Mitrinae. Three of these species were here described as Profundimitra taylori gen. et sp. nov., Fusidomiporta ponderi gen. et sp. nov. and Gemmulimitra neocaledonica gen. et sp. nov. Thus, an estimate of 30 – 100 species of Mitridae remaining to be described from the deep waters of the Indo-Pacific seems reasonable to us. Recognition of new deep-water mitrid species may be hampered by the high intraspecific variability of shell sculpture and, simultaneously, rampant convergences leading to hardly distinguishable shells in not closely related species. Likewise, the radula appears generally useless for species delimitation, as in most cases it retains plesiomorphic morphology, and the significance of slight variations in the number of cusps is unknown. Thus, molecular data will remain essential for a reliable recognition of new deep water taxa of Mitridae. Whereas a significant proportion of the deep-water mitrids remains undescribed, the overall diversity of the family in deep-water pales in comparison with the well-documented diversity of the family in shallow water. These contrasting proportions of described Mitridae in shallow vs. deep water can be viewed as a result of recent efforts by collectors and amateur taxonomists. Since the first half of the 19 th century and the iconic monograph by Reeve (1844 – 1845), the comparatively easily accessible species from shallow water are collected steadily throughout many Indo-Pacific locales and then carefully scrutinized by a dedicated community of amateurs, leading to the present day state of mitrid taxonomy, with a pretty well-inventoried shallow-water fauna. Undoubtedly, there are areas with high local endemism that still hold undescribed species, especially in peripheral locales (e. g. southern Australia, South and East Africa and the Arabian Sea), but this is not the general situation. The diversity of mitrids in the Mediterranean is low, and all three species known from that sea were included in the present study. Conversely, we have included only a limited number of species from West Africa and the New World, which makes any judgment on diversity in these regions untimely. Two Panamic species in our data set were not confidently identified but were represented by subadults, the identification of which is always troublesome, and the uncertainty of our identifications cannot be viewed as an indication of undescribed species. A more thorough sampling in both the Panamic and Caribbean region is definitely required to assess the proportion of undescribed species in these areas and uncover the relationships of their endemic phylogenetic lineages.	en	Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria, Bouchet, Philippe (2018): The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda). Zoological Journal of the Linnean Society 183: 253-337
