LAMELLARIINAE D’ORBIGNY, 1841

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, Zoological Journal of the Linnean Society 197 (4), pp. 924-964 : 951

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.

Figures

Figure 2. Phylogenetic relationships of the family Velutinidae (Bayesian tree based on the combined dataset), with clades (A–L) collapsed by genera. Numbers at nodes indicate branch support values [posterior probability (PP) and ultrafast bootstrap (UFb) values, respectively]; support values are shown only when at least one of them is ≥ 80%; white circles at nodes indicate maximum support (PP = 1, Ufb = 100). On the right side of the tree, for each genus is reported a representative photograph of the shell, a schematic representation of the radula, the depth range (in metres) and the geographical distribution.

Figure 3. Phylogenetic relationships of the family Velutinidae (Bayesian tree based on the combined dataset) showing clades A–L expanded. Numbers at nodes indicate branch support values [posterior probability (PP) and ultrafast bootstrap (UFb) values, respectively]; support values are shown only when at least one of them is ≥ 80%; white circles at nodes indicate maximum support (PP = 1, Ufb = 100). Boxes indicate hypothetical species; dashed lines indicate possible alternative species partitions. Coloured boxes on the right indicate genera, and white boxes indicate subfamilies. Letters at the end of sample codes broadly indicate the area of origin:AK, Alaska; ANT, Antarctica; AR, Argentina; BC, British Columbia; BRE, Bretagne; CA, California; CH, China; COR, Corse; CR, Costa Rica; FL, Florida; GF, French Guiana; GP, Guadeloupe; HI, Hawaii; IT, Italy; MG, Madagascar; MQ, Martinique; MX, Mexico; MZ, Mozambique; NC, New Caledonia; NL, Newfoundland; NO, Norway; OR, Oregon; PA, Panama; PF, French Polynesia; PG, Papua New Guinea; PH, Philippines; QLS, Queensland; RE, Reunion; SA, Saudi Arabia; SP, Spain; TAS, Tasmania; TW, Taiwan; VU, Vanuatu; WA, Washington State; ZNZ, Zanzibar.

Figure 4. A, Marseniopsis sp. (M9), MNA 5376. B, Marseniopsis conica, MNA 5361. C, Marsenina zadei, LACM 182861. D, Marsenina rhombica, LACM 182862. E, Limneria prolongata, LACM 182858. F, Velutina sp., LACM 182869. G, Hainotis sharonae. H, Variolipallium regium sp. nov., MNHN-IM-2013-69285, holotype. I, Variolipallium sp. (L3), MNHN-IM-2013-69605. J, Variolipallium sp. (L10), MNHN-IM-2013-69280. Photograph credits: A, B, MNA, Stefano Schiaparelli; C–F, Nancy Treneman; G, iNaturalist, Robin Gwen Agarwal; H–J, MNHN, Philippe Maestrati. For the sequenced specimens, the corresponding species code (see Fig. 3) is reported in parentheses.

Figure 5. A, Pacifica lentiginosa sp. nov., MNHN-IM-2009-16140, holotype. B, Pacifica sp. (L18), MNHN-IM-2007-33407. C, Djiboutia sp. (L22), MNHN-IM-2019-9053. D, Coriocella sp., UF-IZ-523481. E, Coriocella sp., MNHN-IM-2019-26097. F, Lamellaria mopsicolor, MNHN-IM-2013-73086. G, Lamellaria latens complex (L41), MNHN-IM-2019-6391. H, Marsenia herberti, MNHN-IM-2019-8219. I, Marsenia perspicua complex (L55), MNHN-IM-2019-13522. Photograph credits: A–C, F, MNHN, Philippe Maestrati; D, Florida Museum of Natural History—Invertebrate Zoology, Gustav Paulay; E, MNHN, Giulia Fassio; G, I, MNHN, Gilles Devauchelle; H, MNHN, Laurent Charles. For the sequenced specimens, the corresponding clade code (see Fig. 3) is reported in parentheses.

Figure 7. Velutinidae shells. A, Calyptoconcha pellucida, BAU 3758. B, Variolipallium cerebroides, NIWA 74086.A. C, Pacifica lentiginosa sp. nov., MNHN-IM-2009-16140, holotype. D, Pacifica sp. (L16), MNHN-IM-2013-58543. Scale bars: A, B, 5 mm; C, D, 1 mm. For the sequenced specimens, the corresponding species code (see Fig. 3) is reported in parentheses.

Figure 8. Velutinidae shells. A, Djiboutia sp. (L26), MNHN-IM-2019-7668. B, Coriocella sp. (L27), MNHN-IM-2013-84230. C, Lamellaria latens complex (L41), MNHN-IM-2019-14179. D, Marsenia perspicua complex (L55), MNHN-IM-2019-14574. Scale bars: A, C, D, 1 mm; B, 5 mm. For the sequenced specimens, the corresponding species code (see Fig. 3) is reported in parentheses.

Figure 9. Velutinidae protoconchs. A, Marseniopsis conica, BAS 03-764. B, Marsenina rhombica, BAU 4229. C, Calyptoconcha pellucida, BAU 3758. D, Variolipallium cerebroides, NIWA 74086.A. E, Pacifica lentiginosa sp. nov., MNHN-IM-2009-16140, holotype.F, Pacifica sp. (L16), MNHN-IM-2013-58543. G, Djiboutia sp. (L22), MNHN-IM-2019-9053. H, Coriocella sp. (L27), MNHN-IM-2013-84230. I, Lamellaria latens complex (L41), MNHN-IM-2019-14179. J, Lamellaria mopsicolor, MNHN-IM-2013-73086. K, Marsenia perspicua complex (L55), MNHN-IM-2019-14574. L, Marsenia sp. (L42), MNHN-IM-2019-7641. Scale bars: A–L, 500 µm. For the sequenced specimens, the corresponding species code (see Fig. 3) is reported in parentheses. Arrows indicate protoconch I – protoconch II boundary.

Figure 10. Velutinidae radulae. A, Marseniopsis mollis, CE 6137. B, Marsenina rhombica, BAU 4229. C, Velutina cf. coriacea, LACM 182870. D, Calyptoconcha pellucida, BAU 3751. E, Variolipallium regium sp. nov., MNHN-IM-2013-69285, holotype. F, Variolipallium sp. (L10), MNHN-IM-2013-69280. G, Pacifica lentiginosa sp. nov., MNHN-IM-2009-16140, holotype. H, Pacifica sp. (L15), MNHN-IM-2013-59969. I, Djiboutia sp. (L22), MNHN-IM-2019-9053. J, Coriocella sp. (L27), MNHN-IM-2013-84230. K, Lamellaria mopsicolor, MNHN-IM-2013-73086. L, Lamellaria latens complex (L41), MNHN-IM-2019-14113. M, Marsenia herberti, MNHN-IM-2019-8503. N, Marsenia sp. (L45), MNHN-IM-2013-59960. O, Marsenia perspicua complex (L55), MNHN-IM-2019-5980. Scale bars: A–O, 100 µm. For the sequenced specimens, the corresponding species code (see Fig. 3) is reported in parentheses.

Figure 11. Velutinidae jaws. A, M, Marseniopsis mollis, CE 6152. B, N, O, Marsenina rhombica, BAU 2339. C, Limneria prolongata, BAU 4228. D, Calyptoconcha pellucida, BAU 3758. E, Variolipallium regium sp. nov., MNHN-IM-2013-69285, holotype. F, Variolipallium sp. (L10), MNHN-IM-2013-69280. G, Pacifica lentiginosa sp. nov., MNHN-IM-2009-16140, holotype.H, Pacifica sp. (L19), MNHN-IM-2009-16136.I,Djiboutia sp.(L22), MNHN-IM-2019-9053.J,Coriocella cf.jayi (L28), MNHN-IM-2009-16119. K, Lamellaria latens complex (L40), BAU 2536.2. L, Marsenia sp. (L45), MNHN-IM-2013-59960. Scale bars: H, M–O, 100 µm; A–G, I–L, 500 µm. For the sequenced specimens, the corresponding species code (see Fig. 3) is reported in parentheses.