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MARSENIA OKEN, 1823 (FIGS 2, 3, 5H, I, 8D, 9K, L, 10M–O, 11L) Marsenia Oken, 1823: columns 458, 460; type species Bulla haliotoidea Montagu, 1803 = Helix perspicua Linnaeus, 1758 by monotypy. Included species: Marsenia herberti (Drivas & Jay, 1990) comb. nov., Marsenia perspicua (Linnaeus, 1758), Marsenia affinis Bergh, 1886 [taxon inquirendum], Marsenia cabulana Bergh, 1886 [taxon inquirendum], Marsenia dubia Bergh,1886 [taxoninquirendum], Marseniagemma Bergh, 1875 [taxon inquirendum], Marsenia isabellina Bergh, 1875 [taxon inquirendum], Marsenia perspicua var. lara Bergh, 1899 [taxon inquirendum]. Description: Body of small to medium size for the subfamily, 0.3–10.0 cm total length. Shell thin, weakly calcified; ear shaped, low to high spire, with expanded aperture; smooth or weakly sculptured by axial growth lines; completely enclosed by the mantle or presenting a small dorsal fissure. Periostracum not visible. Protoconch of 1.2–1.6 whorls; protoconch I 0.6–0.75 whorls, nucleus diameter 100–250 μm, smooth, with subsutural axial folds; protoconch II with or without axial growth lines; protoconch–teleoconch boundary not always distinct. Mantle flat or dome shaped, outline rounded; thick or thin, with or without tubercles on dorsum; with anterior siphon folds; texture smooth/wrinkled/jellylike; colour highly variable, almost transparent to white, grey, beige, yellow, orange, red, violet, brown, often patterned with dots, streaks or patches of colour. Penis to the right of the right cephalic tentacle; with or without a lateral subterminal papilla. Vas deferens with or without a free loop in haemocoel. Radula reduced taenioglossate, with formula 0:1:1:1:0; rachidian tooth base bifurcated; rachidian cusp with several external denticles; lateral teeth elongated, with a pointed triangular external cusp, with several denticles on both sides. Jaws short to elongated. Distribution: Indo-West Pacific (Madagascar, Reunion, Red Sea, Taiwan, Philippines, Papua New Guinea, Tasmania, New Caledonia, Hawaii), Mediterranean Sea (Corsica and Alboran Sea), Caribbean Sea (Martinique and French Guiana); 0–214 m deep. Remarks: The type species, Marsenia perspicua, is part of a complex of at least two cryptic species. This widely distributed lineage is present in the tropical areas of the Indo-Pacific and the Atlantic, and in the Mediterranean Sea. It would be difficult to recognize this genus from other lamellarines with a low spire and moderately calcified shell, if it was not for its radula, characterized by a rachidian tooth bifurcated at the base, with small denticles on both sides and elongated lateral teeth with pointed, triangular and external cusps, with small denticles on both sides. In one species (Marsenia sp. L45), a specimen with a shell not completely enclosed by the mantle but presenting a small dorsal fissure (similar to that of some Variolipallium specimens) was observed. We think that, based on the original descriptions, several taxa inquirenda described by Bergh (1886b) from Philippines and Cape Verde material (Marsenia affinis, Marsenia cabulana, Marsenia dubia, Marsenia gemma, Marsenia isabellina and Marsenia perspicua var. lara) might belong to this genus., Published as part of Fassio, Giulia, Stefani, Matteo, Russini, Valeria, Buge, Barbara, Bouchet, Philippe, Treneman, Nancy, Malaquias, Manuel António E., Schiaparelli, Stefano, Modica, Maria Vittoria & Oliverio, Marco, 2023, Neither slugs nor snails: a molecular reappraisal of the gastropod family Velutinidae, pp. 924-964 in Zoological Journal of the Linnean Society 197 (4) on pages 956-957, DOI: 10.1093/zoolinnean/zlac091, http://zenodo.org/record/7814306, {"references":["Oken L. 1823. Litterarischer Anzeiger. Etwas uber den Pariser Konigs-Garten, IV. ISIS 13: 441 - 469.","Bergh LSR. 1886 b. Nudibranchien. Nachtrage und Erganzungen. In: Semper C, ed. Reisen im Archipel der Philippinen. Malacologische Untersuchungen. Suppl. 3, 129 - 225, pls M - R. Wiesbaden: C. W. Kreidel's Verlag."]}

HAINOTINAE FASSIO, BOUCHET, SCHIAPARELLI & OLIVERIO SUBFAM. NOV. (FIGS 2, 3, 4G, 6B) Zoobank registration: urn: lsid: zoobank. org:act: A356DC93-B469-4DC0-B075-ED51CAEBDAA4 Type genus: Hainotis F. Riedel, 2000. Included genera: Hainotis F. Riedel, 2000, Mysticoncha J. K. Allan, 1936. Description: Body of small to medium size, 0.5– 5.0 cm total length. Shell thin to very thin, weakly calcified, ear shaped, high spire, with expanded aperture, smooth, with or without growth lines, completely enclosed by the mantle. Periostracum thin to moderately developed and hairy. Protoconch of 1.7–2.1 whorls; protoconch I of 0.6 whorls, smooth or with weak growth striations, nucleus diameter 86 µm; protoconch–teleoconch boundary not always distinct. Mantle dome shaped, outline rounded or polygonal (low ridges dividing the mantle into six areas, from a raised hexagonal area at the centre of the dorsum); thin to thick; with anterior elongated siphonal fold; colour highly variable, almost transparent to white, grey, beige, yellow, orange, red, violet, brown or black, frequently patterned with dots and/or polygonal shape lines. Penis and vas deferens unknown. Radula taenioglossate, formula 2:1:1:1:2; rachidian tooth elongated, with rectangular base; rachidian cusp with two external denticles on each side; lateral teeth elongated, with a pointed external cusp, with one denticle on the internal side and two on the external side; marginals narrow, with one small denticle on the internal side. Jaws elongated, with masticatory denticles. Distribution: North-eastern Pacific (from Oregon to northern Mexico), South Australia, New Zealand; 0–45 m deep. Remarks: This subfamily is represented in the phylogenetic tree only by specimens of Hainotis sharonae described from the coast of California (type locality: Anaheim Bay, Orange County, CA, USA; Willett, 1939). This lineage is diagnosed by its shell with a higher spire, a radular formula of 2:1:1:1:2 with elongated central and lateral teeth, and an elongated anterior siphonal fold. Pending further molecular analysis, we suggest also including the genus Mysticoncha in this subfamily because its radular formula and shape, shell shape, siphon shape and temperate distribution (South Australia, Victoria and New Zealand) are consistent with those of the type genus, Hainotis., Published as part of Fassio, Giulia, Stefani, Matteo, Russini, Valeria, Buge, Barbara, Bouchet, Philippe, Treneman, Nancy, Malaquias, Manuel António E., Schiaparelli, Stefano, Modica, Maria Vittoria & Oliverio, Marco, 2023, Neither slugs nor snails: a molecular reappraisal of the gastropod family Velutinidae, pp. 924-964 in Zoological Journal of the Linnean Society 197 (4) on pages 949-950, DOI: 10.1093/zoolinnean/zlac091, http://zenodo.org/record/7814306, {"references":["Willett G. 1939. Description of a new mollusk from California. The Nautilus 52: 123 - 124, pl. 9 figs 1, 1 a-b."]}

LAMELLARIINAE D’ORBIGNY, 1841 (FIGS 2, 3, 4H–J, 5A–I, 7A–D, 8A–D, 9C–L, 10D–L, 11D–L) Lamellariinae d’Orbigny, 1841: 200; type genus Lamellaria Montagu, 1816. Coriocellidae Troschel, 1848: 545; type genus Coriocella Blainville, 1824. Lamellariidae d’Orbigny, 1841: 200; type genus Lamellaria Montagu, 1816. Marseniidae Leach in Gray, 1842: 268 [as Marseniadae]; type genus Marsenia Oken, 1823. Included genera: Calyptoconcha Bouchet & Warén, 1993, Coriocella Blainville, 1824, Djiboutia Vayssière, 1912, Lamellaria Montagu, 1816, Marseniella Bergh, 1886, Marsenia Oken, 1823, Pacifica Fassio, Bouchet & Oliverio gen. nov., Variolipallium Fassio, Bouchet & Oliverio gen. nov. Description: Body of small to medium size for the family, 0.5–10.0 cm total length. Shell thin to very thin, from strongly calcified to membranaceus; ear shaped, low to high spire, with expanded aperture; smooth or weakly sculptured by axial growth lines; completely enclosed by the mantle or with a small dorsal fissure. Periostracum thin to not visible. Protoconch of one to two whorls; protoconch I of 0.46–1.20 whorls, smooth, with subsutural axial folds, nucleus diameter 54–250 μm; protoconch II with or without axial growth lines; protoconch–teleoconch boundary not always distinct. Mantle flat or dome shaped, outline rounded or polygonal; thick or thin, with or without dorsal warts or tubercles; with anterior siphon fold; texture smooth, wrinkled, jelly- or velvet-like; colour highly variable, almost transparent to white, grey, beige, yellow, orange, red, violet, blue, dark green, brown or black, often patterned with dots, lines or irregular colour patches. Penis to the right of the right cephalic tentacle, with or without a lateral subterminal papilla.Vas deferens with or without a free loop or, less frequently, several folds in the haemocoel. Radula reduced taenioglossate, formula 0:1:1:1:0; rachidian tooth base bifurcated or rectangular; rachidian cusp with or without several external denticles; lateral teeth elongated, with a single external pointed cusp, or composed of an external, pointed and bold cusp plus a internal truncated projection; in both cases with or without several denticles. Jaws homogeneous, short to elongated. Distribution: Temperate and tropical regions worldwide; 0–4500 m deep. Remarks: Lamellariines are diagnosed by their reduced taenioglossate radula lacking marginals (formula 0:1:1:1:0), a mantle completely enclosing the shell in most species (the only exception so far being Variolipallium regium) and an anterior siphon. The name Echinospira was introduced by Krohn, 1853 for Echinospira diaphana Krohn, 1853, based on a (velutinid?) larva collected in the plankton of the Messina Strait (Italy). Based on the original description, we have no clues confidently to identify this animal, which might be one of the cryptic species in either of the two genera (Lamellaria and Marsenia) present in the Central Mediterranean. Within the examined material, we have recognized the following six lineages that merit recognition at genus level., Published as part of Fassio, Giulia, Stefani, Matteo, Russini, Valeria, Buge, Barbara, Bouchet, Philippe, Treneman, Nancy, Malaquias, Manuel António E., Schiaparelli, Stefano, Modica, Maria Vittoria & Oliverio, Marco, 2023, Neither slugs nor snails: a molecular reappraisal of the gastropod family Velutinidae, pp. 924-964 in Zoological Journal of the Linnean Society 197 (4) on page 951, DOI: 10.1093/zoolinnean/zlac091, http://zenodo.org/record/7814306, {"references":["d'Orbigny A. 1841. Voyage dans l'Amerique meridionale (le Bresil, la republique orientale de l'Uruguay, la republique Argentine, la Patagonie, la republique du Chili, la republique de BoliVia, la republique du Perou), execute pendant les annees 1826, 1827, 1828, 1829, 1830, 1831, 1832 et 1833, Vol. 5. Paris: Bertrand & Strasbourg, Levrault, 1 - 48, 73 - 128, (xliii + 758 pp., 85 plates) (publication dates after Sherborn & Griffin, 1934).","Montagu G. 1816. An account of some new and rare marine British shells and animals. Transactions of the Linnean Society of London 11: 179 - 204, pls 12 - 14 [' 1815 '; 24 January 1816].","Troschel FH. 1848. Mollusca, Gastropoda. In: Wiegmann AFA, Ruthe JF, eds. Handbuch der Zoologie, 3 rd edn. Berlin: Luderitz, 536 - 568.","de Blainville HMD. 1824. Mollusques, Mollusca (Malacozoaires) In: Cuvier F, ed. Dictionnaire des Sciences Naturelles, Vol. 32. Strasbourg & Paris: Levrault & Le Normant, 1 - 392.","Gray JE. 1842. Molluscs. In: Synopsis of the contents of the British Museum, 44 th edn. London: British Museum, 48 - 92, iv + 308 p.","Oken L. 1823. Litterarischer Anzeiger. Etwas uber den Pariser Konigs-Garten, IV. ISIS 13: 441 - 469.","Bouchet P, Waren A. 1993. Revision of the Northeast Atlantic bathyal and abyssal Mesogastropoda. Bollettino Malacologico Suppl. 3: 579 - 840.","Vayssiere A. 1912. Recherches zoologiques et anatomiques sur les Opisthobranches de la Mer Rouge et du Golfe d'Aden. Deuxieme Partie. Annales de la Faculte des Sciences de l'UniVersite de Marseille 20 Supplement: 5 - 157."]}

DJIBOUTIA VAYSSIÈRE, 1912 (FIGS 2, 3, 5C, 8A, 9E, 10I, 11I) Djiboutia Vayssière, 1912: 121; type species Djiboutia Ʋerrucosa Vayssière, 1912 by monotypy. Included species: Djiboutia australis (Basedow, 1905) comb. nov., Djiboutia sibogae (Bergh, 1908) comb. nov., Djiboutia Ʋerrucosa Vayssière, 1912. Description: Body of small to medium size for the subfamily, 0.5–3.3 cm total length. Shell thin to very thin, weakly calcified; ear shaped, low spire, with expanded aperture; smooth or weakly sculptured by axial growth lines; completely enclosed by the mantle. Periostracum not visible. Protoconch of 1.30–1.75 whorls; nucleus diameter 120–145 μm; protoconch I 0.75–1.00 whorls, smooth, with subsutural axial folds; protoconch II with axial growth lines; protoconch–teleoconch boundary not always distinct. Mantle flat or dome shaped, outline rounded; thin; with anterior siphon folds; texture smooth; colour highly variable, almost transparent to white, grey, beige, yellow, orange, red, violet, brown, black, often patterned with dots or patches of colour. Penis to the right of the right cephalic tentacle; with or without a lateral subterminal papilla. Vas deferens without a free loop in haemocoel. Radula reduced taenioglossate, formula 0:1:1:1:0; rachidian tooth base bifurcated; rachidian cusp smooth and slim; lateral teeth elongated, composed of a smooth, external, pointed and bold cusp, plus a smooth internal truncated projection. Jaws short. Distribution: Indo-West Pacific (Madagascar, Gulf of Tadjoura, Papua New Guinea, Sulawesi, southern Australia and Tasmania, New Caledonia); 0–101 m deep. Remarks: The key character to recognize this genus is the rachidian tooth (bifurcated, smooth and slim, without denticles) that represents a unique combination in this family. Based on this character and on an overall similar morphology, we suggest that Lamellaria australis and Marsenia sibogae belong to this genus., Published as part of Fassio, Giulia, Stefani, Matteo, Russini, Valeria, Buge, Barbara, Bouchet, Philippe, Treneman, Nancy, Malaquias, Manuel António E., Schiaparelli, Stefano, Modica, Maria Vittoria & Oliverio, Marco, 2023, Neither slugs nor snails: a molecular reappraisal of the gastropod family Velutinidae, pp. 924-964 in Zoological Journal of the Linnean Society 197 (4) on pages 954-955, DOI: 10.1093/zoolinnean/zlac091, http://zenodo.org/record/7814306, {"references":["Vayssiere A. 1912. Recherches zoologiques et anatomiques sur les Opisthobranches de la Mer Rouge et du Golfe d'Aden. Deuxieme Partie. Annales de la Faculte des Sciences de l'UniVersite de Marseille 20 Supplement: 5 - 157.","Bergh LSR. 1908. The Opisthobranchia of South Africa. Transactions of the South African Philosophical Society 17: 1 - 144."]}

MARSENIELLA BERGH, 1886 Marseniella Bergh, 1886a: 14; type species Marseniella borealis Bergh, 1886 by monotypy. Included species: Marseniella borealis Bergh, 1886. Description: Body of medium size for the subfamily, 2.2 cm total length. Shell thin, membranaceus, weakly calcified; ear shaped, low spire, last whorl wide and detached from the spire at the back, with expanded aperture; completely enclosed by the mantle. Periostracum unknown. Protoconch unknown. Mantle flat, outline rounded; with coarse and fine knots on the dorsum; with anterior siphon folds; texture unknown; colour unknown. Penis to the right of the right cephalic tentacle; with a lateral subterminal papilla. Vas deferens with a free loop and several folds in the haemocoel. Radula reduced taenioglossate, formula 0:1:1:1:0; rachidian tooth base bifurcated; rachidian cusp with several denticles on both sides; lateral tooth elongated, with a pointed triangular external cusp, with several denticles on both sides. Jaws short. Distribution: Known only from Florø, north of Bergen (Norway); depth unknown. Remarks: Bergh (1886a, b, 1887) described this monotypic genus from a single specimen collected in Norwegian waters. Bergh described it as very similar to velutinids that we call here Lamellaria and Marsenia, but different enough to deserve a separate name. The radular formula and the ‘V’-arched rachidian tooth confirm that it belongs to the Lamellariinae. The rachidian tooth with denticle on both sides and a lateral tooth elongated, with a pointed triangular external cusp, with several denticles on both sides might be compatible with the genus Marsenia (but we have not found any Marsenia in Norwegian waters, from where we have examined only Lamellaria specimens). The vas deferens producing several folds in the haemocoel has been observed in the genus Lamellaria, but we do not consider this character as diagnostic at the genus level. Above all, Bergh (1886a, b, 1887) described for Marsenia borealis a very unusual corneous shell, only partly calcified, somehow similar in texture to that of Onchidiopsis. The described shape is also unusual, flatter than Lamellaria and Marsenia, with a very wide last whorl detached from the spire at the back (a shell shape not observed in any other velutinid examined so far) (Bergh 1887: pl. X figs 1, 2). For these reasons, pending further analysis, we provisionally keep Marseniella as a valid genus in the subfamily Lamellariinae, not present in our molecular dataset., Published as part of Fassio, Giulia, Stefani, Matteo, Russini, Valeria, Buge, Barbara, Bouchet, Philippe, Treneman, Nancy, Malaquias, Manuel António E., Schiaparelli, Stefano, Modica, Maria Vittoria & Oliverio, Marco, 2023, Neither slugs nor snails: a molecular reappraisal of the gastropod family Velutinidae, pp. 924-964 in Zoological Journal of the Linnean Society 197 (4) on page 957, DOI: 10.1093/zoolinnean/zlac091, http://zenodo.org/record/7814306, {"references":["Bergh LSR. 1886 a. Report on the Marseniadae collected by H. M. S. Challenger during the years 1873 - 1876. Report on the Scientific Results of the Voyage of H. M. S. Challenger during the years 1873 - 76. Zoology 15 (part 41): 1 - 24, pl. 1.","Bergh LSR. 1887. Nudibranchien. Nachtrage und Erganzungen. In: Semper C, ed. Reisen im Archipel der Philippinen. Malacologische Untersuchungen. Suppl. 4, 226 - 289, pls S - V, X - Z, AE. Wiesbaden: C. W. Kreidel's Verlag."]}

VARIOLIPALLIUM FASSIO, BOUCHET & OLIVERIO GEN. NOV. (FIGS 2, 3, 4H–J, 7B, 9D, 10E, F, 11E, F) Zoobank registration: urn: lsid: zoobank. org:act: 170D2A4B-E62D-4612-9737-FFC39B639EFA Type species: Variolipallium regium Fassio, Bouchet & Oliverio sp. nov. Included species: Variolipallium cerebroides (Hutton, 1882) comb. nov., Variolipallium elatum (Strebel, 1906) comb. nov., Variolipallium leptoconcha (Bergh, 1907) comb. nov., Variolipalliumnodosum (Ev.Marcus, 1987) comb. nov., Variolipallium patagonicum (E. A. Smith, 1881) comb. nov. and Variolipallium regium Fassio, Bouchet, Oliverio sp. nov. Description: Body of small to medium size for the subfamily, 0.5–3.0 cm total length. Shell very thin, weakly calcified to membranaceus; ear shaped, high spired, with expanded aperture; smooth or weakly sculptured by axial growth lines; completely enclosed by the mantle or presenting a small dorsal fissure. Periostracum not visible. Protoconch of 1.24–1.70 whorls; protoconch I of 0.48– 1.20 whorls, nucleus diameter 100–150 μm, smooth, with subsutural axial folds; protoconch II with axial growth lines; protoconch–teleoconch boundary not always distinct. Mantle flat or dome shaped, outline rounded or polygonal; thick to thin, often with a few to several dorsal tubercles; with anterior siphon fold; texture smooth to wrinkled (resembling the convolutions of a brain); colour variable, almost transparent to white, grey, yellow, orange to red, pink to violet, light blue, brown, dark green, occasionally patterned with spots. Cephalic tentacle tips can be white, lime yellow or almost transparent. Penis to the right of the right cephalic tentacle, with or without a lateral subterminal papilla. Vas deferens without a free loop in haemocoel. Radula reduced taenioglossate, formula 0:1:1:1:0; rachidian tooth base bifurcated; rachidian cusp with few to several, very small to pronounced external denticles; lateral teeth elongated, with a pointed triangular external cusp, with few to several, very small to pronounced denticles on both sides or only on the internal side. Jaws homogeneous, short to elongated. Distribution: Tropical West Pacific (South China Sea, Papua New Guinea, New Caledonia), New Zealand; Caribbean; tip of South America (Chile, Argentina, Falkland Islands), South Africa; 42–1573 m deep. Etymology: From Variola, the Latin name for smallpox, and pallium, meaning ‘mantle’, referring to the coloured small tubercles on the mantle of the type species and other members of the group. Gender neuter. Remarks: Variolipallium can be diagnosed by a membranaceous to weakly calcified shell with high spire (similar to that of Calyptoconcha and Marseniopsis) and a bifurcated rachidian tooth (like Lamellaria, Marsenia, etc.), although in Variolipallium the ‘V’-shaped base of the rachidian tooth can be comparatively less marked. All the species included in this genus have been found at depths> 95 m. The dorsal appearance of the mantle varies from wrinkled, with several tubercles (resembling the convolutions of a brain), to rather smooth and studded with fewer small tubercles of different colour, to completely smooth. Several species (Variolipallium cerebroides, Variolipallium cf. patagonicum, Variolipallium cf. elatum, Variolipallium sp. L10, Variolipallium sp. L11, Variolipallium sp. L12, Variolipallium sp. L14 and Variolipallium regium) include specimens with a small to very small dorsal mantle fissure, occasionally marked by a small black spot (visible also in preserved specimens). Lamellaria patagonica E. A. Smith, 1881 (type locality: Trinidad Channel, Chile, in 54 m) and Lamellaria elata Strebel, 1906 (type locality: Puerto Condor, Chile) belong to this genus based on their fragile and high-spired shells, smooth mantle and radular formula and shape. Variolipallium patagonicum and Variolipallium elatum can be distinguished by a more rapidly expanding first whorl in the former and, consequently, a higher shell in the latter, also resulting in a flatter body in Variolipallium patagonicum and a more globose one in Variolipallium elatum. Lamellariaampla Strebel, 1906 (Ushuaia, Argentina) might also belong here (based on shell shape and shell consistency), but further analysis of the type material is necessary, because the radula, in particular, is not described in the original description. However, the general description, in particular the jelly-like mantle, wrinkled, grey with darker spots, would also be compatible with a position in Marseniopsinae. Lamellaria leptoconcha (South Africa, Cape Point, in 1097–1280 m) is included in Variolipallium based on the membranaceus texture of the shell and the shape of the rachidian tooth. Bergh (1907: pl. IX, fig. 18) sketched a rachidian tooth with a not marked but visible ‘V’-shaped base (similar to our SEM photographs; Fig. 10E, F), and in the description underlined that the difference between this species and Marsenia leptolemma (= Calyptoconcha pellucida) was the shape of the rachidian tooth (that in Calyptoconcha indeed has a squared base). Lamellaria cerebroides (Auckland, New Zealand) is included in Variolipallium based on the radular formula and shape, and the wrinkled dorsum appearance. Lamellaria Ʋerrucosa (= Lamellaria nodosa) (Auckland Islands) is also included in this genus based on the membranaceus texture of the shell, the rugose appearance of the dorsum, and the radular formula and shape. Odhner (1924) described it as being similar to Marseniopsis mollis, but the radular formula of Lamellaria Ʋerrucosa clearly indicates that it belongs to the subfamily Lamellariinae. The original description of Lamellaria ophione Gray, 1850 is general (fitting most lamellariine genera) and does not report any information regarding diagnostic characters. However, considering the type locality (Auckland, New Zealand), there is a chance that this species also belongs to this genus, because this is the only lamellarine genus recorded in this area so far.

CALYPTOCONCHA BOUCHET & WARÉN, 1993 (FIGS 2, 3, 7A, 9C, 10D, 11D) Calyptoconcha Bouchet & Warén, 1993: 742. Type species Lamellaria pellucida A. E. Verrill, 1880, by original designation. Included species: Calyptoconcha branca (Simone, 2004) comb. nov., Calyptoconcha capensis (Bergh, 1907) comb. nov., Calyptoconcha pellucida (A. E. Verrill, 1880). Description: Body of small to medium size for the subfamily, 1–4 cm total length. Shell very thin, weakly calcified to membranaceus; ear shaped, high spired with expanded aperture; smooth or weakly sculptured by axial growth lines; completely enclosed by the mantle. Periostracum not visible. Protoconch of 1.77–1.80 whorls; protoconch I of 0.58–1.00 whorls, smooth, nucleus diameter 54–75 μm; protoconch II with axial growth lines; protoconch– teleoconch boundary not always distinct. Mantle flat or dome shaped, outline rounded; thick to very thick, with anterior and right lateral siphon folds; texture smooth or wrinkled; colour variability unknown. Penis to the right of the right cephalic tentacle, with a subterminal lateral papilla. Vas deferens with a free loop in haemocoel. Radula reduced taenioglossate, formula 0:1:1:1:0; rachidian tooth base squared; rachidian cusp with several small external denticles; lateral teeth elongated, triangular with a single external pointed cusp, with several small denticles on both sides. Jaws homogeneous, short, with a small vertical protrusion in the centre. Distribution: Northern temperate Atlantic, Alboran Sea, South Africa, Brazil, Uruguay; 58–4500 m deep. R e m a r k s: T h e g e n u s C a l y p t o c o n c h a c a n b e distinguished by the combination of a reduced taenioglossate radula (formula 0:1:1:1:0), a rachidian tooth with a squared base, and a non-calcified internal vestigial shell (often broken in dredged specimens and difficult to remove from the body) with a comparatively higher spire. The protoconch is also different from the rest of the subfamily in having a small diameter of the nucleus, a small diameter of the first whorl and lacking subsutural axial folds on protoconch I. The jaws show a diagnostic small vertical protrusion in the centre. Based on the radular formula, the shape of the rachidian tooth and composition of the shell, we include in this genus the South African Marsenia capensis (Cape Point, 239–1463 m). We also suggest including Lamellaria branca (type locality: Brazil, off Sao Paulo state, in 78 m) based on its radula and shell shape, although the shell seems to be calcified compared with the other two congeners, which might be related to its shallower water habitat., Published as part of Fassio, Giulia, Stefani, Matteo, Russini, Valeria, Buge, Barbara, Bouchet, Philippe, Treneman, Nancy, Malaquias, Manuel António E., Schiaparelli, Stefano, Modica, Maria Vittoria & Oliverio, Marco, 2023, Neither slugs nor snails: a molecular reappraisal of the gastropod family Velutinidae, pp. 924-964 in Zoological Journal of the Linnean Society 197 (4) on pages 951-952, DOI: 10.1093/zoolinnean/zlac091, http://zenodo.org/record/7814306, {"references":["Bouchet P, Waren A. 1993. Revision of the Northeast Atlantic bathyal and abyssal Mesogastropoda. Bollettino Malacologico Suppl. 3: 579 - 840.","Simone LRL. 2004. Morphology and phylogeny of the Cypraeoidea (Mollusca, Caenogastropoda). Rio de Janeiro: Papel Virtual Editoria.","Bergh LSR. 1907. The Opisthobranchiata of South Africa. In: Marine InVestigations of South Africa 5 (1) [Dated 1908]. Page (s): 106, pl. 9, figs 17 - 20; pl. 10, figs 1, 2. Transactions of the South African Philosophical Society 17: 1 - 144, pls 1 - 14."]}

PACIFICA LENTIGINOSA FASSIO, BOUCHET & OLIVERIO SP. NOV. (FIGS 2, 3, 5A, 7C, 9E, 10G, 11G) Zoobank registration: urn: lsid: zoobank. org:act: 210F01F5-C95D-4F27-A63D-9715015E67EE Type material: Holotype: MNHN-IM-2009-16140 (Sud Madagascar, Lavanono West sector, ATIMO VATAE/ BP21, 25°23.1–23.2ʹS, 44°51.4–51.6ʹE, 20 m deep, female). Material examined: MNHN-IM-2009-16140 (Sud Madagascar, Lavanono West sector, ATIMO VATAE/ BP21, 25°23.1–23.2ʹS, 44°51.4–51.6ʹE, 20 m deep, female). Description: Body of medium size for the genus, 1 cm total length. Shell very thin, weakly calcified; ear shaped, low spire, with expanded aperture; weakly sculptured by axial growth lines; completely enclosed by the mantle. Periostracum not visible. Protoconch of 1.6 whorls; nucleus diameter 110 μm; protoconch I 1.2 whorls, smooth, with subsutural axial folds; protoconch II with axial growth lines; protoconch–teleoconch boundary distinct. Mantle dome shaped, outline rounded; thin; with anterior siphon folds; texture slightly wrinkled; dorsal mantle colour light violet, patterned with several small white and beige spots, fewer bigger orange spots and some brown patches. Penis unknown; vas deferens unknown. Radula reduced taenioglossate, formula 0:1:1:1:0; rachidian tooth base bifurcated; rachidian cusp with three external long denticles on each side; lateral teeth elongated, with a pointed triangular external cusp, with four or five long denticles on each side. Jaws short; with an external protrusion on the upper side. Distribution: So far known only from the type locality (southern Madagascar), 20 m deep. Etymology: From the Latin adjective lentiginosa, meaning ‘freckled’, referring to the dorsal coloration of the holotype. Remarks: This species is known only from shallow waters of southern Madagascar. Owing to the high variability in shell shape and mantle coloration in this family, the most reliable way to identify this species remains by molecular analysis.

PACIFICA FASSIO, BOUCHET & OLIVERIO GEN. NOV. (FIGS 2, 3, 5A, B, 7C, D, 9E, F, 10G, H, 11G, H) Zoobank registration: urn: lsid: zoobank. org:act: BC75A004-F0A8-40C5-9636-2B3CFDC96EEF Type species: Pacifica lentiginosa Fassio, Bouchet & Oliverio sp. nov. Included species: Pacifica indecora (Bergh, 1886) comb. nov., Pacifica lentiginosa Fassio, Bouchet & Oliverio sp. nov. Description: Body of small size for the subfamily, 0.5– 1.0 cm total length. Shell very thin, weakly calcified; ear shaped, low spire, with expanded aperture; smooth or weakly sculptured by axial growth lines; completely enclosed by the mantle. Periostracum not visible. Protoconch of 1.6–1.8 whorls; nucleus diameter 110–135 μm; protoconch I 1.1–1.2 whorls, smooth, with subsutural axial folds; protoconch II with axial growth lines;protoconch–teleoconchboundarynotalwaysdistinct. Mantle flat or dome shaped, outline rounded; thin; with anterior siphon folds; texture smooth; colour almost transparent to white, grey, yellow, orange, pink, violet, brown, patterned with many small and big spots or colour patches. Penis to the right of the right cephalic tentacle; with a lateral subterminal papilla. Vas deferens with or without a free loop in haemocoel. Radula reduced taenioglossate, formula 0:1:1:1:0; rachidian tooth base bifurcated; rachidian cusp with three or four external long denticles on each side; lateral teeth elongated, with a pointed triangular external cusp, with four to seven long denticles on each side side. Jaws short, with an external protrusion on the upper side. Distribution: Indo-West Pacific (Madagascar, Philippines, Papua New Guinea, New Caledonia, Vanuatu); 20–244 m deep. Etymology: From the Latin adjective pacifica, meaning ‘peaceful’, as a symbolic quest for peace from the scientific community. Gender feminine. Remarks: This lineage includes a few Indo-West Pacific species with a peculiar radula (both rachidian and lateral teeth with long denticles) and a unique jaw shape, small and with an external protrusion on the upper side. The two specimens photographed alive showed a dorsal mantle with several coloured spots. Marsenia indecora Bergh, 1886 (Philippines Sea) belongs to this genus because of its peculiar jaw shape and radular shape. Bergh (1886b) himself regarded this species as remarkably different from all other species he described from the Philippines (that are now included in the genus Marsenia)., Published as part of Fassio, Giulia, Stefani, Matteo, Russini, Valeria, Buge, Barbara, Bouchet, Philippe, Treneman, Nancy, Malaquias, Manuel António E., Schiaparelli, Stefano, Modica, Maria Vittoria & Oliverio, Marco, 2023, Neither slugs nor snails: a molecular reappraisal of the gastropod family Velutinidae, pp. 924-964 in Zoological Journal of the Linnean Society 197 (4) on pages 953-954, DOI: 10.1093/zoolinnean/zlac091, http://zenodo.org/record/7814306, {"references":["Bergh LSR. 1886 b. Nudibranchien. Nachtrage und Erganzungen. In: Semper C, ed. Reisen im Archipel der Philippinen. Malacologische Untersuchungen. Suppl. 3, 129 - 225, pls M - R. Wiesbaden: C. W. Kreidel's Verlag."]}

MARSENIOPSINAE FASSIO, BOUCHET, SCHIAPARELLI & OLIVERIO SUBFAM. NOV. (FIGS 2, 3, 4A, B, 6A, 9A, 10A, 11A, M) Zoobank registration: urn: lsid: zoobank. org:act: 0FF5505F-49D2-4814-8E81-CA559340917E Type genus: Marseniopsis Bergh, 1866. Included genera: Marseniopsis Bergh, 1866, Lamellariopsis Vayssière, 1906 (probably a synonym of Marseniopsis Bergh, 1866). Description: Body of small to medium size for the family, 0.5–11.5 cm total length. Shell very thin, weakly calcified to membranaceus; ear shaped, high spired, with expanded aperture; smooth or weakly sculptured by axial growth lines; enclosed by the mantle. Periostracum thin to not visible. Protoconch of 0.76–1.9.0 whorls; protoconch I of 0.25–0.73 whorls, with or without granular sculpture, nucleus diameter 150–875 μm; protoconch II with or without marked longitudinal ribs; protoconch– teleoconch boundary not always distinct. Mantle flat or dome shaped, outline rounded or polygonal; often thick, but sometimes thin, with or without warts; with anterior siphonal fold; texture smooth to wrinkled or jelly-like; colour highly variable, almost transparent to white, grey, beige, yellow, orange, red or brown, frequently patterned with dots and/or polygonal lines. Penis to the right of the right cephalic tentacle; with or without a lateral subterminal papilla. Vas deferens with a free loop in haemocoel. Radula taenioglossate, formula 2: 1:1: 1: 2; rachidian tooth elongated, with rectangular base, elongated; rachidian cusp with three or four small to pronounced external denticles on each side; lateral teeth elongated, with pointed triangular external cusp, with two or three small to pronounced denticles on the internal side and one or two small to large denticles on the external side; marginals narrow, without denticles. Jaws elongated, composed of scale-like elements. Distribution: Southern Ocean and south Argentinian waters; 75–668 m deep. Remarks: Fassio et al. (2019) suggested that the radiation of velutinids in Antarctica might represent a lineage worthy of taxonomic recognition at subfamily level. This is here confirmed, and the geographical range is now extended outside the Southern Ocean to Argentina, with the record of M5 (= new MOTU P) from north of Isla de los Estados (USNM 1137366, 54°27ʹ19.1″S, 63°52ʹ39.4″W, 108 m depth). Marseniopsinae can be diagnosed by the radular formula combined with the elongated shape of the rachidian tooth with a rectangular base, and by an ear-shaped shell, not well calcified, with a high spire. Lamellariopsis is probably a synonym of Marseniopsis, but the genetic analysis of type or topotypical material is needed to check it. Two types of protoconchs, potentially indicating a difference in the length of the larval phase, have been described for this subfamily (see Fassio et al., 2019)., Published as part of Fassio, Giulia, Stefani, Matteo, Russini, Valeria, Buge, Barbara, Bouchet, Philippe, Treneman, Nancy, Malaquias, Manuel António E., Schiaparelli, Stefano, Modica, Maria Vittoria & Oliverio, Marco, 2023, Neither slugs nor snails: a molecular reappraisal of the gastropod family Velutinidae, pp. 924-964 in Zoological Journal of the Linnean Society 197 (4) on page 949, DOI: 10.1093/zoolinnean/zlac091, http://zenodo.org/record/7814306, {"references":["Fassio G, Modica MV, Alvaro MC, Buge B, Salvi D, Oliverio M, Schiaparelli S. 2019. An Antarctic flock under the Thorson's rule: diversity and larval development of Antarctic Velutinidae (Mollusca: Gastropoda). Molecular Phylogenetics and EVolution 132: 1 - 13."]}

FAMILY VELUTINIDAE GRAY, 1840 Velutinidae Gray, 1840: 90; type genus Velutina J. Fleming, 1820. Included subfamilies: Hainotinae Fassio, Bouchet, Schiaparelli & Oliverio subfam. nov., Lamellariinae d’Orbigny, 1841, Marseniopsinae Fassio, Bouchet, Schiaparelli & Oliverio subfam. nov., Velutininae Gray, 1840. Description: Small to medium size for the superfamily, 0.5–11.5 cm total length. Shell exposed to completely Marseniella Uniform Norway? Marsenia Uniform Indo-West Pacific, Mediter- ranean Sea, Caribbean Sea – 214 0 Lamellaria Uniform and Tropical Temperate eastern Pacific Tropical, Atlantic, north-eastern Atlantic, Mediterra- Sea nean, North Sea 61 0 – Coriocella Uniform Indo-West Pacific – 0 18 Djiboutia Uniform Indo-West Pacific –101 0 Pacifica Uniform Indo-West Pacific 20 – 244 Variolipallium Uniform Tropical West New, Pacific Zealand,, Caribbean South tip of, America Africa South 1573 – 42 Calyptoconcha Uniform Northern temperate, Atlantic, Sea Alboran South, Africa Uru- Brazil, guay 58 – 4500 Lamellariinae Uniform Worldwide temperate tropical, regions 4500 0 – Velutininae Scale-like elements, Arctic worldwide temperate regions 0– 1200 Hainotinae? North-eastern, South Pacific, Australia New Zealand 0 45 –, Continued Marseniopsinae Scale-like elements Southern Ocean Argen- south tinian waters 668 75 –. Table 2 Character Composition Geograph- distri- ical bution m (Depth) Table 2. Continued enclosed by the mantle, thin to very thin, from strongly calcified to membranaceus; ear, shield or cap shaped, low to high spired, with expanded aperture; smooth or weakly sculptured by axial growth lines. Periostracum from thick and hairy to not visible. Protoconch of 0.76–2.10 whorls; protoconch I of 0.25–1.20 whorls, smooth or with microgranules, with or without subsutural axial folds, nucleus diameter 54–875 μm; protoconch II with or without marked axial ribs or growth lines; protoconch–teleoconch boundary not always distinct. Echinospira planktotrophic larva with double larval shell: the outer periostracal planispiral, smooth and rounded or strongly carinate, the inner helicoid. Mantle flat (Fig. 4C) or dome shaped (Fig. 4A), outline from above rounded (Fig. 4J) or polygonal (Fig. 4B, H); thick or thin, with or without dorsal warts or tubercles; with or without anterior (inhalant) and right lateral (exhalant) siphon folds; texture smooth, wrinkled, jelly-like or velvet-like; colour highly variable, almost transparent to white, grey, beige, yellow, orange, red, violet, blue, dark green, brown, black, often patterned. Penis to the right of the right cephalic tentacle; with or without a lateral subterminal papilla; with or without tip of the seminal duct protruding from the penis tip. Vas deferens with or without a free loop in haemocoel. Radula taenioglossate (with formula 2:1:1:1:2) or reduced taenioglossate (if lacking marginal teeth, formula 0:1:1:1:0); rachidian tooth base rectangular (broad or elongated) or bifurcated (inverted V-shape), rachidian cusp with or without several external denticles; lateral teeth broad or elongated, with a pointed, triangular, internal or external cusp or one external cusp plus one truncated projection, with or without denticles; marginals, when present, narrow, with or without denticles. Jaw s of v a r i a b l e s h a p e, s h o r t t o e l o n g at e d; homogeneous or composed of scale-like elements; with or without uniform masticatory denticles. Distribution: Worldwide, from shallow to abyssal waters (0–4500 m). Remarks: Velutinids differ from the other two velutinoidean families (Triviidae and Eratoidae) chiefly in their thin to very thin, helicoid shell, with expanded aperture (vs. solid, cowry-like, with narrow aperture in triviids and eratoids) and the planispiral outer layer of the echinospira larval shell (vs. helicoid in triviids and eratoids). Also, the siphon is proportionally shorter in velutinids than in triviids and eratoids. Many velutinid species are reported to live and feed on ascidians (Wilson, 1998), and the colour and texture of the dorsal mantle can mimic the ascidian host. Both hermaphroditic and gonochoristic species are reported (Wilson, 1998). Egg capsules are flask shaped (Diehl, 1956; Fretter & Graham, 1962) and are laid in holes in the tunic of the ascidians (Peck et al., 2006; Fassio et al., 2019)., Published as part of Fassio, Giulia, Stefani, Matteo, Russini, Valeria, Buge, Barbara, Bouchet, Philippe, Treneman, Nancy, Malaquias, Manuel António E., Schiaparelli, Stefano, Modica, Maria Vittoria & Oliverio, Marco, 2023, Neither slugs nor snails: a molecular reappraisal of the gastropod family Velutinidae, pp. 924-964 in Zoological Journal of the Linnean Society 197 (4) on pages 945-948, DOI: 10.1093/zoolinnean/zlac091, http://zenodo.org/record/7814306, {"references":["Gray JE. 1840. Shells of molluscous animals, pp. 105 - 152. In: Synopsis of the contents of the British Museum. ed. 42. London: G. Woodfall. 370 p.","d'Orbigny A. 1841. Voyage dans l'Amerique meridionale (le Bresil, la republique orientale de l'Uruguay, la republique Argentine, la Patagonie, la republique du Chili, la republique de BoliVia, la republique du Perou), execute pendant les annees 1826, 1827, 1828, 1829, 1830, 1831, 1832 et 1833, Vol. 5. Paris: Bertrand & Strasbourg, Levrault, 1 - 48, 73 - 128, (xliii + 758 pp., 85 plates) (publication dates after Sherborn & Griffin, 1934).","Wilson B. 1998. Superfamily Velutinoidea. In: Beesley PL, Ross GJB, Wells A, eds. Mollusca: the southern synthesis. Fauna of Australia, Vol. 5. Melbourne: CSIRO Publishing, part B, 786 - 790.","Diehl VM. 1956. Die Raubschnecke Velutina Velutina als Feind und Bruteinmieter der Ascidie Styela coriacea. Kieler Meeresforschungen 12: 180 - 185.","Fretter V, Graham A. 1962. British prosobranch molluscs. Their functional anatomy and ecology. London: Ray Society.","Peck L, Clarke A, Chapman A. 2006. Metabolism and development of pelagic larvae of Antarctic gastropods with mixed reproductive strategies. Marine Ecology Progress Series 318: 213 - 220.","Fassio G, Modica MV, Alvaro MC, Buge B, Salvi D, Oliverio M, Schiaparelli S. 2019. An Antarctic flock under the Thorson's rule: diversity and larval development of Antarctic Velutinidae (Mollusca: Gastropoda). Molecular Phylogenetics and EVolution 132: 1 - 13."]}

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The family Teredinidae (shipworms) contains 70-plus species of boring bivalves specialized to live in and digest wood. Traditional means of species identification and taxonomy of this group encounter numerous challenges, often compounded by the diverse and dynamic nature of shipworm ecology and distribution. Modern integrative taxonomic methods are shedding new light on this complex group, from delineating cryptic species to resolving phylogenetic relationships within the family. This study reported new sequence data from shipworm species rafted from the western to eastern Pacific Ocean in woody marine debris resulting from the Japanese tsunami of 2011. Eight species of shipworms were found in this debris and tissue from five species was collected. Partial nuclear ribosomal 18S rRNA gene sequences were obtained from Bankia bipennata (Turton, 1819), Bankia carinata (Gray, 1827), Psiloteredo sp., Teredora princesae (Sivickis, 1928), and Teredothyra smithi (Bartsch, 1927). A 658 base pair fragment of COI was successfully sequenced from Psiloteredo sp. and T. princesae specimens from tsunami debris, as well as Psiloteredo megotara (Hanley, 1848) from Europe and Nototeredo norvagica (Spangler, 1792) from Scandinavia. Psiloteredo sp. is very similar morphologically to the North Atlantic Ocean P. megotara; however, these two species are genetically distinct with a 12.8% K2P distance in their COI sequences. The transport of shipworms across the North Pacific Ocean in woody debris generated by a tsunami shows that major geologic events can connect previously isolated geographic areas and provide the opportunity for the establishment of invasive species and subsequent speciation.

The Tohoku tsunami of March 2011 ejected a vast amount of debris into the Pacific Ocean. Wood boring shipworms (Bivalvia: Teredinidae) were either already present in, or settled on, the wooden fraction of this debris, offering a unique opportunity to study shipworm diversity in rafted wood of a known origin and time of ocean entry. Lumber and other wood began appearing on Central Pacific (Hawaiian Islands) and Eastern Pacific beaches in 2013. Eighty pieces of wood Japanese Tsunami Marine Debris (JTMD) consisting of construction beams, trees, milled logs, and wood from vessels or maritime structures were analyzed. Six shipworm species resident in the coastal waters of Japan were found: Bankia bipennata (Turton, 1819), Bankia carinata (Gray, 1827), Teredothyra smithi (Bartsch, 1927), Psiloteredo sp., Lyrodus takanoshimensis (Roch, 1929), and Teredo navalis Linnaeus, 1758. Two pelagic species, Teredora princesae (Sivickis, 1928) and Uperotus clava (Gmelin, 1791), were acquired by JTMD wood in the transoceanic voyage. Several of these wood items were discovered soon after stranding and contained live shipworms. Up to five shipworm species were found in any one wooden object. The present work represents the first study of the diversity and abundance of shipworms transported across an ocean basin in a large woody debris field.