Mitromorpha (Mitrolumna) columbellaria (Scacchi, 1836)

(Figs 1–5; 6A–C; 7; 27A–E; 28A)

Mitra columbellaria Scacchi, 1836: 10, figs 12, 13 (31 December: ICZN 1999: Art. 21.3, see Cretella et al. 2005: 131)

Mitra obsoleta Philippi, 1836: 230 (31 December: ICZN 1999: Art. 21.3, see Cretella et al. 2005: 115) non Brocchi, 1814 nec Bronn, 1831

Mitra columbellaria Scacchi: Philippi 1844:195, pl. XXVII, fig. 17

Mitra leontocroma Brusina, 1866: 34 –35

Columbella greci Philippi: Monterosato 1878:103 –104

Columbella greci var. lactea Monterosato, 1878:103 –104

? Columbella greci var. unifasciata Monterosato, 1878:103 –104

Columbella greci var. minor Monterosato, 1878:103 –104

Columbella greci var. granulosa Monterosato, 1878:103 –104

Mitrolumna olivoidea Cantraine: Bucquoy, Dautzenberg & Dollfus 1883: 121, pl. XV, figs 33–35

Mitrolumna olivoidea var. major Bucquoy, Dautzenberg & Dollfus, 1883: 122, pl. XV, figs 36–37

Mitrolumna olivoidea var. minor Bucquoy, Dautzenberg & Dollfus, 1883: 122 ex Monterosato ms.

Mitrolumna olivoidea var. granulosa Bucquoy, Dautzenberg & Dollfus, 1883: 122, pl. XV, figs 38–39 ex Monterosato ms.

Mitrolumna olivoidea var. lactea Bucquoy, Dautzenberg & Dollfus, 1883: 122 ex Monterosato ms.

Mitrolumna major Locard, 1886:109, 542 ex Bucquoy, Dautzenberg & Dollfus ms

Mitrolumna granulosa Locard, 1886:109, 542 ex Monterosato ms. non (Delle Chiaie, 1827)

Mitrolumna algeriana Pallary, 1900: 266, pl. VI, fig. 6 ex Monterosato ms

Mitra olivoidea Cantraine, 1835: 391 ‘pars’ sensu Cernohorsky 1970: pl. 12, fig. 8 et 1975, figg. 55 e 56)

Mitrolumna algeriana Monterosato: Pallary, 1900

Mitrolumna algeriana Monterosato: Nordsieck 1977

Mitrolumna olivoidea Cantraine: Nordsieck 1977

Mitrolumna olivoidea Cantraine: Chirli 1997:27, pl. VII, fig 8 e 10

Mitromorpha (Mitrolumna) mediterranea Mifsud, 2001:10 –11, pl. 1, figs 13–16, plate 2, figs 18a,18b, 19, 20, pl. 3, fig 40 (left).

Mitromorpha mediterranea Mifsud: Gofas, Moreno & Salas 2011: 329

Type material. M. columbellaria (Scacchi, 1836), neotype here designated (H 6.95 mm, W 3.21 mm), Capri Is., (MCZR, Monterosato collection, M- 21-17319) (Fig. 1 G–I). M. leontocroma, lectotype (H 6.9 mm, W 3.15 mm, CNHM-2088c), here designated, and 3 paralectotypes (CNHM-2088a,b,d). Brusina reported three specimens (H 7 mm, W 3 mm), but the type series consists of four shells, of which one is a broken encrusted subadult (CNHM- 2088b), hence probably not cited by the author (Figs 1 L–N; 5A–I). Mitromorpha major (Locard, 1866 ex BDD ms) type material not found at MCZR, MNHN, MCL. Mitromorpha granulosa (Locard, 1886 ex Monterosato ms) not found at MCZR, MNHN, MCL. Mitromorpha mediterranea Mifsud, 2001, holotype (paralectotype of M. olivoidea Cantraine: Cernohorsky 1970 pl. 12, fig. 9 IRSN-IG3354), H 5.4 mm, W 2.7 mm (Fig. 1 J, K) and 17 paratypes (IRSN-IG3354), Mediterranean; 1 paratype, Algeçiras, Spain, coll. Mifsud; 1 paratype (white specimen), St. Thomas Bay, Malta, coll. Mifsud. Mitromorpha algeriana (Pallary, 1900 ex Monterosato ms), neotype, here designated (H 5.75 mm, W 2.85 mm), Oran, Algeria, 5–10 m depth, viii.1977 (Bruno Amati legit) (MNHN IM- 2000-27714) (Fig. 1 D).

Type locality. M. columbellaria (Scacchi) Capri Is., Italy. M. leontocroma (Brusina) Croatia, “southern Dalmatia ( Lesina, Lissa, Lagosta e Ragusa)” (Brusina 1866: 3, 34–35). M. major (Locard), Les Martigues, France. M. granulosa (Locard) Cannes, France. M. algeriana (Pallary) Oran, Algeria. M. mediterranea Mifsud, Mediterranean Sea, originally not designated, type material from ‘Mediterraneo’.

Other material esamined. Italy: Bosa Marina, 35 m, 3 sh (CS); “Ennio Falco” cave, Porto Conte, Sardinia, 5–15 m, viii.2003, 3 sh (MO); Castelsardo, 50 m, 4 sh (AR); Spargi Is., Sardinia, 18–20 m, 13. xii.1993, 1 sh (MO); Maddalena Is., Sardinia, beached, 1979, 1 sh (BA); Maddalena Is., Sardinia, 80 m, vii.1984, 1 sh (IN), unspecified depth, 1 sh (BA), 7 sh (CS); Caprera Is., Sardinia, 20 m, vi.1995, 8 sh (3 white) (CS); Santa Teresa di Gallura, Sassari, Sardegna, 5 sh (PC); Tuscan Arcipelago, unspecified locality and depth, 4 sh (SB-MS), 4 sh (PC); Gorgona Is., 400 m. ix.1979. 2 sh (IN); Elba Is., 40 m, 3 sh (AR); Giglio Is., 20 m, 4 sh (CS), 25–35 m, 4 sh (MO); Giannutri Is., 25 m, 32 sh (CS), 40 m, 8 sh (CS), 35 m, 7 sh (CS), 40 m, 2011, 29 sh (CS), 47 m, v.1982, 9 sh (MO), 48 m, 7 sh (MO); Tor Paterno Shoal, unspecified depth, 1998, 1 sh (INC), 30 m, v.1986, 4 sh (IN), 45 m, 1 sh (BA); Torre Astura, unspecified depth, 1977, 3 sh (BA); Cape Circeo, 150 m, 1 sh (CS); Ponza Is., 60 m, 4 sh (CS); Ventotene Is., 22 m, 1 sh (CS); Procida Is., Baia del Carboncio, Napoli, 1 sh (PC); Procida, Secca delle Formiche, 4 sh (PC); Capri Is., Grotta Azzurra, 14 m, 5 sh (CS); Capri Is., unspecified depth, 6 sh (MCZR, Monterosato collection, M- 21-17319); Cape Palinuro, unspecified depth, 7 sh (SB-MS), 45–55 m, 12 sh (SB-MS); Marina di Camerota, 30 m, 5 sh (CS); Cannizzaro, Sicily, 43 m, 5 sh (BA), 30 m, 1 sh (BA), 45 m, 70 sh (CS), 35 sh (PC); Scilla, 42 m, 46 sh (CS), 30 m, 4 sh (CS), 60 m, 27 sh (AR), 40 m, 3 sh (AR), 49 sh (PC), 217 sh (PC); Villa San Giovanni, loc. Santa, Reggio Calabria, 3 sh (PC); Tricase, 20 m, 1987, 1 sh (CS); Scilla, San Gregorio, 1 sh (PC); Scilla, Dente di Skilla, 8 sh (PC); Messina, Sicily, 20–50 m, fishing by-catch, 1 lv subadult (DS); Pozzallo, Porri Is., 25 m, viii.1985, 1 sh (IN); Siracusa, Sicily, 3 sh (MCZR, coll. Monterosato, cabinet M drawer 16, labelled olivoidea); Brucoli beach, Augusta, Siracusa, Sicily, 3 sh (PC); Lipari Is., Sicily, (ex Tiberi), 2 lv (MCZR, coll. Monterosato, M- 21-17325); Sicily, several localities (ex Brugnone), 14 sh (MCZR, coll. Monterosato, M- 21-17325); Vendicari, Sicily, 24 m, vii.1985, 4 sh (IN), 29 m vii.1985, 4 sh (IN); Acitrezza, Sicily, 32 sh (PC); Portopalo, Sicily, 19 m, 2 lv (IN); Secca Catanesi, Portopalo, Sicily, 18 m, ix.1998, 12 sh (IN); Salina Is., Sicily, 35 m, vii.2002, 8 sh (MO), 3 sh (BA), 46 m, 3 sh (CS); “ Isola delle Correnti”, Sicily, 12 m, vii.1986, 7 sh (white) (IN); Capo Passero, Sicily, 21 m, vii.1983, 3 sh (IN); Capo Asparano, Sicily, unspecified depth, ix.1985, 10 sh (BA); Talbot Bank, Sicily Strait, 25 m, 3 sh (CS); Pantelleria Is., 40–60 m, 2 sh (1 juv. white) (AR); Pantelleria Is., 10–24 m, vii.1983, 4 sh (IN); Lampedusa Is., Punta Cappellone, Sicily, 4 sh (PC); Cala Lampedusa Is., 60 m 15 sh (AR); Cinque Denti, Pantelleria Is., 10–30 m, viii.1983, 4 sh (IN); Trapani, beached, iv.1983, 1 sh (IN); Punta Altarella, Levanzo Is., Sicily, 20–30 m, 3. v.1991, 4 sh (MO), 31 m 3. vi.1991, 4 sh (BA); Marettimo Is, Sicily, 25–45 m, 4–20. vi.1998, 1 sh (MO), 1 sh (CS); S. Vito lo Capo, Sicily, 3 sh (MCZR, coll. Monterosato, M- 21-17319 labelled crenulata). Portocesareo, beached, viii.1973, 5 sh (IN); Pliocene (Zanclean) Pietrafitta, Siena (Mauro Brunetti, Rioveggio, at http:// www.naturamediterraneo.com/forum/topic.asp? TOPIC _ID=136044 last accessed on 11/JAN/2015).

France: Coast of Provence (ex Sollier), 3 sh (MCZR, coll. Monterosato, M- 21-17319 labelled M. olivoidea); Roussillon, 11 sh (MCZR, coll. Monterosato, M- 21-17319 labelled M. olivoidea); Porquerolles, 1 sh (MNHN, labelled Mitrolumna granulosa coll. Locard ‘1892 p.50’); Porquerolles, 3 sh (MNHN labelled Mitrolumna major coll. Locard ‘1892 p.50’); Toulon, 3 sh (MNHN labelled Mitrolumna major coll. Locard ‘1892 p.50’); “Secca del Toro”, Cervicales Is., Corsica, 25 m, 31 sh (CS); Centuri Port, Corsica, 30–40 m, 3 sh (CS); Capo Corso, Corsica, 440 m, 1 sh (CS).

Spain: Tarifa, 30 m, 2012, 14 sh (SB-MS); Algeciras, port, viii.1980, 6 sh (IN), 1sh (PC); Getares, beached, viii.1985, 8 sh (IN), 2 sh (PC); Roquetas del Mar, Almeria, 2 sh (PC); Benidorm, Alicante, Punta de Albir, 3 sh (PC); Punta Mona, Malaga, 40 m, 19 sh (SB-MS); Cabo de Palos, 5 m, viii.1980, 19 sh (IN); La Herradura, Granada, ix. 2006, 35 m, 35 sh (AR), 1sh (PC).

Greece: Paros Is., 3 sh (MCZR, coll. Monterosato, M- 21-17319); Lagonissi, beached, vi.1982, 9 sh (IN); Aghia Pelaghia, Crete, 1 m, vi. 1982, 1 lv (IN); Creta Is., Georgopulis, 1 sh (PC).

Turkey: Kas, 50 m, 1992, 5 sh (MO); Aydincik, 9 m, vii.1989, 2 sh (IN).

Cyprus: Cape Greco Protaras, ix.2011, 1 sh (BA).

Lebanon: Ramkine Island, 13–14 m, v/ vi.2000, 8 sh (MO); Ras El Chakaa, cave sediment, 3–4.5 m, 04. vi.2000, 2 sh (MO); Saida, 31 m, 5. vi.2000, 1 sh; El Kassmieh, 44 m, 25. x.1999, 2 sh (MO).

Slovenija: Portoroz, unspecified depth, 1977, 4 sh (BA).

Croatia: Ragusa, 2 sh (MCZR, M- 21-17319 coll. Monterosato, ex Coen, ex Brusina).

Malta: Malta, unspecified locality and depth, 1 sh (BA); St. Thomas Bay, beached, 1 sh (CS).

Algeria: Algeria, 2 sh (MCZR, type cabinet, ex coll. Monterosato, labelled Mitrolumna algeriana v. cingulosa ms. Monterosato); Oran, Algeria, 9 sh (MCZR, coll. Monterosato M- 21-17319), two labels handwritten by Pallary reading: “Golfe d’Oran 40/ 60 m Mitrolumna olivoidea Cantr. ” and “fonds corall. Oran 2 Mitrolumna olivoidea v. major BDD”; one label handwritten by Monterosato reading: “ Mitrolumna maxima Mont. Mers el Kébir 50 m Pallary 900” (Fig. 1 E, F)

Tunisia: Tunis, 2 sh (MNHN coll. B.D.D. labelled M. olivoidea v. granulosa).

No locality, 3 sh (MCZR coll. Monterosato M- 21-17319); No locality, 2 sh (MCZR, coll. Monterosato M- 21- 17319); No locality, 1 sh (MCZR, coll. Monterosato M- 21-17319); No locality, 2 lv (MCZR coll. Monterosato [ex Lecheis], M- 21-17319 labelled olivoidea); ‘varie località del Mediterraneo’, 3 sh (MNHN coll. B.D.D. labelled M. olivoidea).

Distribution and habitat. Pliocene, Zanclean of Pietrafitta, Siena (Mauro Brunetti at http:// www.naturamediterraneo.com/forum/topic.asp?TOPIC_ID=136044, as M. mediterranea). Pliocene of Romagna, Campania and Sicily and Pleistocene (Selinuntian) of Sicily (Chirli 1997: 27, pl. 7, figs 8, 10 as Mitrolumna olivoidea). Recent: entire Mediterranean Sea.

Empty shells in bioclastic sediments from 1–60 m; shells from deeper bottoms (80–440 m) probably drifted from shallower habitat (Bonfitto et al. 1994a: 134; Bonfitto et al. 1994b: 141; Smriglio & Mariottini 1996: 191). Living on rocky bottoms with photophylous algae (Gofas, Moreno & Salas 2011: 329), and amidst the rhizomes of Posidonia oceanica (as M. olivoidea: Templado & Llanso 1981). Found sympatric, often at different depths, with M. olivoidea, M. karpathoensis, M. wilhelminae, M. alyssae n. sp., M. bogii n. sp., M. mariottinii n. sp., M. mifsudi n. sp., M. nofronii n. sp. and M. cossyrae n. sp.

Description. (in parentheses the data of the neotype) Shell of large size for the genus (Figs 1–5; 27A–E), height 3.85–9.4 mm (6.95 mm), width 2.15–4.75 mm (3.21 mm). Solid, fusiform, biconic, slender, H/ W=1.980–2.190 (2.165). Protoconch paucispiral (Figs 6A–C; 28A) shiny, of 1.15–1.5 (1.3) convex whorl, d 0.15–0.25 mm (0.175), Do 0.32–0.40 mm (0.375), DM 0.50–0.60 mm (0.55), h 0.35–0.62 mm (0.45), nucleus with fine spiral threads, interrupted and randomly arranged, with rare granules in the interspaces. Remaining whorls sculptured by numerous microgranules. Protoconch-teleoconch boundary well marked, slightly flexuose, opisthocline. Teleoconch of 3.8–5.6 (4.8) moderately convex whorls. Sculpture starting with 3 (3) spiral cordlets crossed by 12–17 (15) axial ribs on the first whorl. Additional spiral cordlets between the two adapical ones, and between the suprasutural one and the suture. Last whorl sculptured over the entire surface by 16–28 (24) spiral cordlets, flat, not equidistant, of variable size, of which 4–6 (6), exceptionally up to 8, above the aperture. Axial ribs of variable strength producing nodules at the intersection with the spirals. A wider interspace between the two adapical cordlets, of which the subsutural usually stronger and lighter. Two columellar folds, the posterior larger and frequently bifid on the outer tip (Fig. 6D). Growth lines visible over the entire surface. Outer lip sharp, thickened internally with 6–9 (9) denticles, the second or third posterior larger. Anal sinus shallow. Siphonal canal short, and wide. Coloration monochrome, dark to light brown, rarely white (Figs 2 D; 3O–Q) ( var. lactea Monterosato, 1878), commonly with a lighter (almost white) band on the adapical cordlet, sometimes with darker or lighter spirals. Protoconch of the same color as the teleoconch, often darker than the teleoconch. Soft parts very variable in coloration: light background with yellow stripes (Scaperrotta et al. 2012); beige to light orange with many white speckles over the surface, and black eyes (Gofas et al. 2011:326, fig. s.n.); white translucid background with many light yellowish speckles over the entire surface (as M. olivoidea: Templado & Llanso 1981); yellow monochrome (Mifsud 2001; Italo Nofroni, Roma pers. comm.); yellow with warty surface over the foot (Fig. 1 O) (Danilo Scuderi, Catania pers. comm.).

Remarks. Mitra columbellaria Scacchi, 1836 and Mitra obsoleta Philippi, 1836 have been both dated to 31 December by Cretella et al. 2005: 115, 131 (ICZN 1999: Art. 21.3). As first revisors, we consider Mitra columbellaria Scacchi, 1836 as having priority over Mitra obsoleta Philippi, 1836 (ICZN 1999: Art. 24.2).

The frequent observation of small, yet fully adult specimens, even within the same population, prompted the description of size varieties ( minor, maxima [Monterosato in schedis] and major, raised to species by Locard 1886). The presence of a sexual dimorphism is a hypothesis to test by examination of live collected specimens.

The study of a large number of samples of Mitromorpha columbellaria from all over the Mediterranean Sea, showed a great variability in size, sculture and color. We have spotted at least 5 distinct morphs. Morph A, corresponding to the description of M. columbellaria (Scacchi), with axial sculpture only on the first whorls, and spirals over the entire surface (Figs 1 G–I; 3J–R; 4C–L). Morph B, corresponding to M. leontocroma (Brusina), M. granulosa (Locard) (partim) and M. algeriana (Pallary), with axial and spiral sculture over the entire surface, and tubercles at the intersections (Figs 1 A–C, D–F, L–N; 2A–C, F–I; 4A,B). Morph C, corresponding to M. major (Locard) (partim) and M. mediterranea (Mifsud), with axials and spirals sculpture of equal strength over the entire surface, without tubercles (Figs 1 J, K; 2D, E, J–K). Morph D, with axial and spiral sculpture over the entire surface with evident nodules at the intersections, and with the exception of the extreme base and the last half whorl where only the spirals are present (Figs 3 A–G; 4A). Morph E, almost lacking any axial sculpture (Fig. 3 H, I). All adult specimens of M. columbellaria we have examined (> 900 adult shells from the entire range) had 4–6 spirals over the aperture, with the exception of only two specimens from Capo Palinuro with 8 spirals (collected with other 17 shells with 4–6 spirals).

Mitra columbellaria Scacchi, 1836 has been regarded (e.g.: Locard 1886a: 101; Monterosato 1872: 50; 1874: 276) as a junior synonym of Mitra greci Philippi, 1844, despite being an older name, and of M. olivoidea sensu auctores (Seguenza 1880, BDD 1883, Dautzenberg 1883, Locard 1886a: 101, Cernohorsky 1970, Sabelli et al. 1990, Chirli 1997). The type material, formerly at MZUN (Cretella et al. 2005) is now lost. To stabilize the use of the name, we have selected a neotype from the Gulf of Napoli, fitting the original description and the concept of M. columbellaria of the historical authors (Fig. 1 G–I). M. wilhelminae can be distiguished from M. columbellaria by its lighter coloration with darker spiral striae and alternate dark and light blotches, its more inflated outline, and the weaker sculpture; M. wilhelminae morph A can be separated also by the stouter outline and the usually higher number of spirals over the aperture (8–9 v. 4–6 in most columbellaria). Juveniles of M. wilhelminae, have also a more inflated outline (nearly keeled at the periphery) than those of M. columbellaria . See under M. mariottinii n. sp., M. mifsudi n. sp., M. tricolorata n. sp. and M. alyssae n. sp. for distinction with M. columbellaria .

M. leontocroma was originally compared also to M. typostigma Brusina, 1866 (: 67, a replacement name for Mitra striata Brusina, 1865, due to the presumed existence of a homonym by Lamarck which we could not identify; however there are also at least a Mitra striata Lea, 1833, and a M. striata Gray, 1839). According to the presence of three columellar folds and the teleoconch surface with a pitted sculpture (Brusina 1865: 14), M. typostigma was quite probably based on a juvenile of Mitra cornicula (Linnaeus, 1758) .

Specimens of morph B can at first sight be confused with M. mariottinii, which can be diagnosed by the spiral sculpture over the entire surface and the axials usually only on the first whorls (rarely on the entire shell), with large nodules at the intersections (and not clear cut square interspaces with small tubercles); finally, M. mariottinii has always 2–3 colored spirals, the first and often the second subsutural darker, and the suprasutural white, often interrupted.

The type material of Mitrolumna algeriana Pallary, 1900 has not been found at MNHN, MCZR and NHM. In the Monterosato collection (MCZR) are two specimens labelled “ M. algeriana var cingulosa ” (a varietal name never published Fig. 1 E, F), quite probably not types, from Algeria. They fit the original description, corresponding to the morph A along with the nominal taxa M. granulosa (Delle Chiaje), M. leontocroma (Brusina) and M. granulosa (Locard), albeit are not well preserved. We have designated one specimen from Oran (ex BA) perfectly congruent with the original description, as neotype of Mitrolumna algeriana Pallary, 1900 to stabilize the use of the name (Fig. 1 D). The samples of M. major Locard and M. granulosa Locard found in the MNHN cannot be considered as types (they correspond to the material in Locard 1892) but are of help in confirming the concept of Locard on this taxon (Fig. 2 A–C, F–I, L–O).