Leufroyia concinna (Scacchi, 1836)

(Figs 9-13)

Pleurotoma concinna Scacchi, 1836: 13, fig. 18.

Fusus boothi J. Smith, 1839: 98 .

Defrancia leufroyi var. carnosula Jeffreys, 1867: 367 .

Pleurotoma linearis var. major-violacea Monterosato, 1875: 270.

Homotoma michaudi Bellardi, 1877: 276 . Raphitoma (Leufroyia) scacchii De Casa & Hallgass, 1979: 6, pl. 1; non Bellardi, 1847.

Lineotoma concinna form maderensis Nordsieck & Talavera, 1979: 166, no. 34.

Clathurella (Cirillia) linearis var. concinna – Coen 1933: 72, no. 480.

Clathurella (Cyrillia) linearis concinna – Kobelt 1905: 368, pl. 96, figs 6, 7.

Clathurella concinna – Bucquoy et al., 1883: 98, pl. 14, fig. 5 – Tryon 1890: 277, pl. 34, fig. 85 – Locard 1886: 116; 1892: 69 – Locard & Caziot 1899: 63.

Defrancia leufroyi – sensu Jeffreys, 1867: 366-368 non Michaud, 1828 – sensu Norman, 1879: 30, no. 237 non Michaud, 1828.

Leufroyia concinna – Monterosato 1884: 134 – Bellini 1929: 32 – Trono 2006: 65 – Mazziotti et al. 2008: 78.

Peratotoma (Leufroyia) concinna – Cerulli-Irelli 1910: 60 (252), pl. 5 (36), fig. 47 – Cipolla 1914: 61 (175), pl. 3 (14), figs 25, 26.

Peratotoma michaudi – Tropeano et al. 1984: 55.

Philbertia concinna – Van Straaten 1960: 112.

Philbertia leufroyi boothi – E.H. Smith 1967: 176-187.

Philbertia leufroyi – sensu Hubendick & Warén 1974: 28, fig. 222 non Michaud, 1828 – sensu Lebour 1934: 553 (veliger figured) non Michaud, 1828.

Pleurotoma (Defrancia) concinna – Monterosato 1877a: 43, pl. 5, fig. 1; 1877b: 336; 1877c: 425; 1878: 106; 1880: 230.

Pleurotoma (Leufroyia) michaudi – Sacco 1904: 53, pl. 13, figs 55, 56.

Pleurotoma concinna – Calcara 1839a: 5; 1839b: 17 – Philippi 1844: 166 – Petit de la Saussaye 1852: 133 – Weinkauff 1868: 133 [in the synonymy of Murex linearis Montagu, 1803] – Aradas & Benoit 1876: 251 [in the synonymy of Murex linearis Montagu, 1803] – Carus 1893: 251 [in the synonymy of Murex linearis Montagu, 1803] – Cretella et al. 2005: 115, 121.

Pleurotoma concinnum [sic] – Sandri & Danilo 1856: 137, no. 141 [in the synonymy of Murex linearis, Montagu, 1803].

Pleurotoma leufroyi – sensu Forbes & Hanley 1853: 468-470 non Michaud, 1828.

Raphitoma (Cirillia) concinna – Nordsieck 1968: 178, pl. 30, fig. 94.62.

Raphitoma (Leufroyia) concinna – Nordsieck 1977: 60, pl. 20, fig. 159 – Piani 1980: 156 – Luque & Templado 1981: 22 – Templado & Llanso 1981: 36 – Sabelli et al. 1990: 44, 216.

Raphitoma concinna – Parenzan 1970: 210, pl. 44, fig. 851 – Ghisotti 1972: 85 – Bogi, Coppini & Margelli 1980: 14, fig. 1 – Borja 1987: 217 – Poppe & Goto 1991: 174, pl. 1, fig. 20 – Koutsoubas et al. 1992: 73 – Cachia, Mifsud & Sammut 1993: 34 – Giribet & Peñas 1994: 52; 1997: 52 – Panetta & Imperatrice 1994: 39 – Tabanelli & Segurini 1994: 12 – Arduino et al. 1995: 94 (figured) – Consolado Macedo 1996: 121, fig. 152 – Borja & Muxika 2001: 76 – Cachia, Mifsud & Sammut 2001: 64, pl. 9, fig. 10 – Basso & Brusoni 2004: 40 – Öztürk, Buzzurro & Benli 2004: 59 – Repetto et al. 2005: 39, 216, fig. 888 – Peñas et al. 2006: 43; 2009: 25 – Oliver Baldoví 2007: 39 – Vardala-Theodorou & Nicolaidou 2007: 65 – Cecalupo et al. 2008: 32 – Cossignani & Ardovini 2011: 31, 324, 325 (figured) – Gofas, Moreno & Salas 2011: 339 (figured) – Manousis 2012: 178 (only central figure) – Oliver et al. 2012: 55; 2015: 109 – Trono & Macrì 2013: 35 – Ceulemans et al. 2018: 113 – Manousis et al. 2018: 11, fig. 6a-c.

Raphitoma leufroyi – Fretter & Graham 1985: 538-540, fig. 371 non Michaud, 1828 – Ardovini & Cossignani 2004: 225 non Michaud, 1828 [only the first shell of 2nd row in the figure] – Manousis 2012: 179 (lower figure) non Michaud, 1828 – Wigham & Graham 2018: 137, fig. 161 non Michaud, 1828.

Raphitoma michaudi – Chirli 1997: 84, pl. 24, figs 1-4.

TYPE MATERIAL. — Pleurotoma concinna Scacchi: presumably lost (Cretella et al. 2005). Fusus boothi J. Smith: presumably lost (Kathie Way pers. comm.). J. Smith’s material was at the Anderson Museum (Glasgow) closed on 1887, thereafter moved at the Hunterian Museum (Glasgow) where this sample has not been found. Defrancia leufroyi var. carnosula Jeffreys:Types not found (Warén 1980: 33). Pleurotoma linearis var. major-violacea Monterosato: not found at the Museo Civico di Zoologia, Rome. Homotoma michaudi Bellardi: it should be at the Museo di Storia Naturale diTorino, currently not accessible. Raphitoma (Leufroyia) scacchii De Casa & Hallgass: private collection Alessandro Hallgass (Rome). Lineotoma concinna form maderensis Nordsieck & Talavera:SMF, Senckenberg Museum, Frankfurt/M (Germany) not seen.

TYPE LOCALITY. — Pleurotoma concinna Scacchi: “ In sinu Neapolitano et Tarentino parum frequens ” [scarcely frequent in the gulfs of Napoli and Taranto] (Scacchi 1836). Fusus boothi J. Smith: Oban and Skye (Scotland). Defrancia leufroyi var. carnosula Jeffreys: shetland. Pleurotoma linearis var. major-violacea Monterosato: Atlantic Ocean. Homotoma michaudi Bellardi: Pliocene of Zinola, Albenga and Vallone Torsero (Italy). Raphitoma (Leufroyia) scacchii De Casa & Hallgass: Sardinia and La Maddalena Island. Lineotoma concinna form maderensis Nordsieck & Talavera: Porto Santo (Madeira).

MATERIAL EXAMINED. — Atlantic. Norway • 1 lv; 60°33’N, 4°52’12”E; ZMBN-H-3-69 • 1 lv; 60°18’N, 5°10’48”E; ZMBN-E-23-67 • 1 lv; 60°13’48”N, 5°12’E; ZMBN-020209-F. Iceland • 1 sh; Reykjavík; coll. KEL. Great Britain • 1 sh; Clyde (Scotland); USNM 305163 • 2 sh; Aber- deen Bank; 57°02’N, 01°12’W; 58 m, 27.VII.2011; coll. SWI • 1 sh; Cumbrae I.; Jeffreys coll. under “ leufroyi ”; USNM 190583 • 3 sh; North Sea, England; 53°1’N, 2°10’E; leg. S. Hanley; SMNH 70481 • 1 sh; Cornwall; leg. A. Gardiner in 1919; coll. FEN • 11 sh; Fal- mouth; 50°9’N, 5°4’W; leg. A. Gardiner in 1919; SMNH 70479 • 1 sh; Falmouth; 50°9’N, 5°4’W; SMNH 70480 • 1 sh; Herm I.; Jeffreys coll. under “ leufroyi ”; USNM 190585 • 2 sh; Guernsey I.; Jeffreys coll. under “ leufroyi ”; USNM 190584. France • 1 sh; Roscoff; coll. DSC • 1 sh; Ouessant I; coll. DSC. Spain • 1 sh; Gijòn; coll. GON • 1 sh; 10 km NNE Cap Finisterre; 42°55’N, 9°17’W; SMHN 73165A • 1 sh; La Coruña; coll. GUB • 1 sh; El Ferrol; SMHN 73164A. Portugal • 1 sh; Sesimbra; SMNH 24101 • 1 sh; Faro; SMNH 25009 • 1 sh; Sagres; SMNH 73211. Canary Islands • 1 sh; Lanzarote I., Puerto del Carmen; coll. DSC • 1 sh; Lanzarote I., Arrecife; coll. DSC. Mediterranean. Spain • 1 sh; Malaga; coll. PUS • 2 sh; Formentera I.; SMHN 70486A • 1 sh; Cadaqués; coll. PUS • 1 sh; Sitges; coll. PUS. France • 1 sh; Bouche du Rhone; coll. PUS • 2 sh; S. Raphael, Le Dramont; coll. HOA • 1 sh; Antibes, Le Graillon; coll. HOA • 1 sh; Antibes, Le Graillon; BAU-1880.1 • 2 sh; Provence; “C[oste] di Provenza”; MCZR-M • 1 lv; Figuerolles, La Ciotat; 43°09’53”N, 5°35’45”E; 15 m; BAU-2263.1. Corsica • 1 sh; Baie de Calvi; 7 m; SMNH 73171J • 1 sh; Ajaccio; coll. CRO • 1 lv; Corsica2019; Cap Corse; MNHN-IM-2019-4050. Sardinia • 1 sh; Porto Conte; coll. OLI • 1 sh; Isola Rossa; coll. DSC • 2 sh; S. Teresa di Gallura; 6 m; coll. CRO • 1 sh; Cagliari; coll. PIS • 8 sh; Sant’Antioco; coll. PIS. Sicily • 1 sh; Palermo Gulf; coll. GIR • 6 sh; Palermo; MCZR-M 17111 • 1 sh; Isola delle Femmine; coll. SER • 1 sh; Marettimo I., Punta S. Simone; 40 m; coll. PAO • 4 sh; Catania; MCZR-M 17342 • 1 sh; Ognina; coll. GER • 2 sh; Cannizzaro; coll. GER • 1 sh; Acitrezza; coll. CRO • 1 sh; Acicastello; 38 m; coll. PAG • 1 sh; Messina; 6 m; coll. PAG. Tunisia • 1 sh; Kerkennah Islands; NMR 2633. Italy • 5 sh; Bergeggi; 38 m; coll. REP • 1 sh; Capraia I.; coll. GOR • 1 sh; Elba I.; 40 m; coll. BAR • 1 sh; Elba I., Punta dell’Acqua Bona; coll. GOR • 9 sh; Golfi di Baratti; 7 m; coll. PAO • 3 sh; Formiche di Grosseto; coll. PAO • 2 sh; Porto Santo Stefano; coll. PAG • 6 sh; Secca delle Vedove; 80-100 m; coll. PAO • 1 sh; Castiglioncello; coll. MAR • 1 sh; Tor Paterno; coll. GER • 2 sh; Ostia; coll. PIE • 1 sh; Circeo; coll. NAP • 1 sh; Sorrento; coll. DUR • 1 sh; SMR; Giannutri I.; coll. • 1 sh; MCZR-M 16700; Napoli • 1 sh; Capri I.; sub nomine “ Philbertia (Leufroyia) leufroyi albida BDD”; HUJ coll. Coen no. 8090; • 4 sh; Capri I.; coll. CRO • 5 sh; Maratea; coll. CAR • 1 sh; Calabria, Costa Viola; 36 m; coll. PAO • 3 sh; Scilla; coll. CRO • 1 sh; Scilla; 50 m; coll. VAZ • 15 sh; Lazzaro; coll. VAZ • 1 sh; Campomarino; coll. CRO • 1 sh; Marina di Ugento; coll. MAC • 2 sh; S. Isidoro; coll. TRO • 1 sh; Porto Badisco; coll. MAC • 2 sh; Otranto. Croatia; coll. MAC • 1 sh; Dugi Otok I., Veli rat; as concinna var. grata ms; MCZR-M17342 • 5 sh; Pag I., Košljun bay; 2-5 m; coll. PRK • 1 sh; Molat I.; amidst red coral colonies; 35-50 m; coll. PRK • 3 sh; Rivanj I.; 18-30 m; coll. PRK • 1 sh; Ugljan I., Karantun; 1-3 m; coll. PRK • 8 sh; Sukošan; 2-7 m; coll. PRK • 3 sh; Dugi Otok I., Mežanj; 10 m; coll. UGA • 18 sh; Dugi Otok I., Veli Garmenjak; 32-40 m; coll. PRK • 5 sh; Dugi Otok I., Nozdre bay; 1-10 m; coll.PRK • 29 sh; Murter I., Kosirina bay; 1-7 m; coll. PRK • 1 sh; Žirje I.; amidst red coral colonies; 70-100 m; coll. PRK • 6 sh; Primošten, Šparadići; 2-6 m; coll. PRK • 20 sh; Sevid; 2-6 m; coll. PRK • 2 sh; Voluja; 30-50 m; coll. PRK • 6 sh; Jabuka I.; 15-35 m; coll. STA • 13 sh; Jabuka I.; 15-35 m; coll. UGA • 4 sh; Čiovo I., Fumija; 10 m; coll. UGA • 7 sh; Šolta I.; 3-15 m; coll. STA • 2 sh; Šolta I.; 3-15 m; coll. UGA • 10 sh; Split; 1-6 m; coll. PRK • 1 sh; Split; 10 m; coll. UGA • 15 sh; Brač I., Milna; 7-10 m; coll. PRK • 19 sh; Brač I., Milna; 7-10 m; coll. STA • 1 sh; Brač I., Maslinova bay; 3-15 m; coll. PRK • 74 sh; Brač I., Maslinova bay; 3-15 m; coll. STA • 14 sh; Omiš, Balića Rat; 4-8 m; coll. PRK • 1 sh; Brela; 5 m; coll. PRK • 18 sh; Lastovo I.; amidst red coral colonies, 70-100 m; coll. PRK • 6 sh; Lastovo I.; amidst red coral colonies, 70-100 m; coll. STA • 1 sh; Lastovo I.; 50 m; coll. BAR • 5 sh; Pelješac, Prapratno; 2-6 m; coll. PRK • 2 sh; Mljet I., Pod Škoji; 1-3 m; coll. PRK • 48 sh; Mljet I., amidst red coral colonies; 70-100 m; coll. PRK • 1 lv; Bio- grad; 43°55’51”N, 15°26’42”E; BAU-2254.1. Greece • 3 sh; Kato Achaia; coll. CRO • 1 sh; Sane; coll. CRO • 1 sh; Crete Island, Elounda; coll. DSC • 2 sh; Korinthiakos Gulf; 60-120 m; coll. KON • 4 sh; Saronic Gulf; 30-70 m; coll. KON • 2 sh; Pylos; 6 m; coll. PAG. Turkey • 1 sh; unprecised locality; coll. CGS • 1 sh; Göçek, Sarsala Bay; coll. CGS • 1 sh; Didim; 50-100 m; coll. CGS • 1 sh; Izmir Bay; 28 m; coll. OZT • 1 sh; Datça; coll. TRI • 1 sh; Mersin, Soguksu; coll. TRI .

A B D C

DISTRIBUTION. — Early-late Pliocene: Northern Italy (Sacco 1904); Altavilla, Palermo, Italy (Cipolla 1914); Pliocene, Piedmont, Italy (Tropeano et. al. 1984); Rio Albonello, Emilia-Romagna (Tabanelli & Segurini 1994); Tuscany, Italy (Chirli 1997).

Recent: Northeastern Atlantic from Norway to Canary Islands. The entire Mediterranean. Common on rocky bottoms at 1-100 m depth. Collected alive under stones or amidst algae, and in sciaphilous habitats, including amidst red coral colonies where it is quite frequent. It often lives in sympatry/syntopy with other raphitomids, also with other Leufroyia spp.; in shallow water it is frequently found with L. leufroyi even under the same stone or amidst the same algae.

ORIGINAL DESCRIPTION. — “ Pleurotoma concinna Nobis (18) Testa albo-sordida, lineis transversis rubro-fuscis inconcinne ornata; transversim striata; per longum costata; apertura dilute violacea. Alta lin. 7-8. P. C. similis at minor, gracilior, striis transversis rariobus et diversa colorum pictura. In sinu neapolitano et tarentino parum frequens. Consule” Fig. 18 (Scacchi 1836:13).

DESCRIPTION

Shell

Solid and fusiform, of medium size for the genus. Height: 9.0- 15.17 mm (mean 11.8 mm, SD: 1.69); width: 3.3-7.91 mm (mean 5.32 mm, SD: 0.81); H/W: 1.84-2.77 (mean 2.24, SD: 0.18).

Protoconch

Multispiral with 2.4 to 2.9 (mean 2.66, SD: 0.14) convex whorls and small nucleus (d: 140 µm). Protoconch I of 1.0 whorl, diameter 265 µm, covered by dense cancellate sculpture; protoconch II with axial threads under the suture and less dense and diagonally cancellate sculpture on rest of whorl. A keel at the end of last whorl of varying length or absent. Protoconch-teleoconch boundary of flexuose, opisthocline growth lines. Colour usually brown with white nucleus, sometimes purplish-brown, rarely very light with pinkish hue, never dark brown or blackish. Transition from protoconch to teleoconch well visible by contrasting colours, brown vs pink or purple.

Teleoconch

Of 5.2-7.0 convex whorls (mean 6.16, SD: 0.48), with deep and distinct wavy sutures and prominent axial sculpture; suture area slightly thickened abapically and white coloured with brown dots or lines; whole external surface covered with relatively thick periostracum; dense and rough axial riblets (growth lines) present on ribs, interspaces, and on subsutural ramps; microgranules over the whole surface, scattered on 2-3 adapical whorls and dense on others.

Axial sculpture of 10-16 (mean 12.93, SD: 1.53)] strong, equidistant, orthocline or slightly opisthocline ribs slightly broader, equal or narrower than interspaces.

Spiral sculpture of 9-16 narrow and low cords (mean 11.53, SD: 1.74) above the aperture, of which 3-4 weak on subsutural ramp, remaining of almost equal strength and equally spaced, or with few additional thin cordlets between the strongest cords, occasionally one thin cordlet between all pairs of strong cords; interspaces 2-4 times broader than cords; on first adapical whorl 4 spiral cords almost equal in strength and one thin subsutural above them.

Siphonal fasciole with 6-9 (mean 7.86, SD: 0.75) strong and slightly nodulose cords, stronger than cords of body-whorl.

Cancellation rectangular to squared, with very elongate and slightly elevated tubercles at intersections.

Subsutural ramp narrow and inclined, covered with dense growth marks of the anal sinus, crossed by 3-4 weak spiral cordlets.

Columella simple, straight medially and arcuate or angled posteriorly. Siphonal canal wide and short, posterior canal deep and moderately wide.

Outer lip thickened and smooth internally, with edge crenated by spiral cords.

Height of aperture: 3.46-6.42 mm (mean 4.86 mm, SD: 0.74); A/H: 35.92-46.79% (mean 41.13%, SD: 2.50).

Coloration

Background colour light, from whitish to pale brownish, with pinkish, purple or purplish-brown areas present always on 1-3 adapical whorls, often inside aperture, on subsutural ramps and in interspaces between axial ribs, frequently darker above a median spiral brown cord. Some spiral cords completely or partially dark brown or reddish-brown, occasionally light brown, orange or yellowish, rarely only slightly darker than background; a broad light spiral band almost devoid of blotches at aproximately 1/3 to 1/2 of whorl height; a brown band often below periphery. Cords on siphonal fasciole light with some tubercles of intense brown or brownish-orange colour. Aperture white or pinkish.

Soft parts

Foot very long, broad and deeply bilobed anteriorly and with recurved anterolateral corners, narrowly tapering posteriorly. Siphon cylindrical with ventral slit, widely opening anteriorly, very extensile. Head small with a pair of long cylindrical tentacles, black eyes on bulges about halfway their length, distal part slightly longer and much narrower than basal. Operculum absent, toxoglossate radula present (G.O. Sars 1878: pl. VIII, fig. 2).

Colour translucent white with minute white speckles densely covering upper side of foot, scattered or absent on sole, head and along neck. Tentacles translucent white, with or without white speckles along entire lenght. Siphon translucent white or yellowish-white, with less dense white speckles.

REMARKS

L. concinna has been largely misidentified as L. leufroyi by many authors, old and modern.

The maximum height of examined shells is 16 mm, but shells larger than 14 mm are rare. The height and maximum diameter of the protoconch depend on the number of whorls; in the specimen of Fig. 13, with 2.75 protoconch whorls, the protoconch is 580 µm high and 504 µm wide. The diameter of nucleus and first whorl are small and similar to L. leufroyi, smaller than in L. erronea and L. villaria . The colour of protoconch varies geographically, mostly brown in some areas (e.g. Adriatic Sea), purple or purplish-brown in others (e.g.Tyrrhenian Sea). The sexes are separated (male and female reproductive systems are described by E.H. Smith (1967) under the name Philbertia leufroyi boothi), but we did not find any evidence of bimodality in morphometrics, thus guessing no size dimorphism is present.

Diagnostic features of L. concinna from all other Leufroyia spp. are the constant presence of pinkish or purple areas on parts of the shell, the numerous brown spiral cords on all whorls recalling the colour pattern of Cyrillia linearis (Montagu, 1803), and the more intense brown colour of tubercles on the siphonal fasciole (although shells tend to fade out after some years from collection). A few shells may look more homogeneously coloured with very light spiral cords, but also in such cases the pinkish colour of at least the first teleoconch whorl is always diagnostic. We have observed that specimens stored in 96-100% ethanol does not change significantly intensity of colours after some years, whilst those stored in 70% ethanol or formalin do fade out rapidly and significantly. There is some geographic variation in colour and sculpture: almost uniformly dark brown shells are not known from the Mediterranean; some Atlantic specimens have stronger and broader spiral cords than Mediterranean ones, also the axial ribs can be very weak or absent on the body whorl (HØisaeter 2016). Some specimens from Canary Islands are smaller (max. 10 mm), more slender and with a colour pattern recalling L. leufroyi (Fig. 10 I-K). The northermost record (HØisaeter 2016) is from southern Lofoten Islands at 67°10.7’N, 14°20.3’E. The southernmost record is from Canary Islands. We have no confirmed records from the Açores and Madeira.