Leufroyia leufroyi ( Michaud, 1828 )

Giannuzzi-Savelli, Riccardo, Pusateri, Francesco, Prkić, Jakov, Bartolini, Stefano, Russini, Valeria, Fassio, Giulia & Oliverio, Marco, 2020, Revision of Mediterranean and NE Atlantic Raphitomidae (Gastropoda, Conoidea) 8: The genus Leufroyia Monterosato, 1884, Zoosystema 42 (22), pp. 433-473 : 438-446

Leufroyia leufroyi ( Michaud, 1828)

( Figs 4-8 )

Pleurotoma leufroyi Michaud, 1828: 121 , pl. 1, figs 5, 6.

Pleurotoma zonalis Delle Chiaje, 1831 : pl. 84, fig. 1.

Pleurotoma cyrilli O.G. Costa, 1844: 54 .

Murex caudicula Nardo, 1847: 59 ex Chiereghini ms – Brusina 1870: 158.

Clathurella leufroyi var. albida Bucquoy, Dautzenberg & Dollfus, 1883: 96 .

Clathurella mirabilis Locard, 1891: 68 .

Clathrurella [sic] concina [sic] – sensu Ostroumoff 1896: 59 non Scacchi, 1836.

Clathurella leufroyi – Bucquoy et al. 1883: 95, pl. 14, figs 3, 4 – Carus 1893: 427, no. 16 – Claudon 1902: 189 – Kobelt 1905: 365, pl. 96, figs 4, 5 – Coen 1933: 74, no. 482 – Dautzenberg 1889: 28 – G.O. Sars 1878: 361 – Gignoux 1913: 93 – J.T. Marshall 1912: 297 – Locard & Caziot 1899: 62 – Locard 1886: 115; 1892: 68, fig. 54; 1899: 49 – Nobre 1936: 52, pl. 32, fig. 4; 1938: 133, pl. 19, fig. 8 – Norman 1899: 137 – Paulus & Mars 1942: 76 – Sykes 1906: 186 – Tryon 1890: 276, pl. 18, fig. 33 – Watson 1886: 350.

Defrancia leufroyi – Appelius 1869: 138 – Caruana-Gatto & Despott 1919: 25 – Cocconi 1873: 59 – Coppi 1869: 165 – Despott 1919: 181 – Foresti 1874: 71 – H. Adams & A. Adams 1853: 96 – Jeffreys 1870: 82 – Monterosato 1872: 51 – Seguenza 1873: 298; 1875: 208 – Weinkauff 1868: 132 sp. 4; 1873: 11.

Homotoma leufroyi – Bellardi 1877: 274, no. 13.

Leufroyia leufroyi – Monterosato 1884: 134 – Di Stefano 1889: 101 – Pallary 1900: 257 – Bellini 1929: 31 – Montefameglio et al. 1979: 191 – Nordsieck & Garcia Talavera 1979: 165, pl. 41, fig. 33 – Orlando & Palazzi 1985: 44, no. 344 – Öztürk, Buzzurro & Benli 2004: 59 – Brunetti & Vecchi 2005: 3 – Brunetti & Della Bella 2006: 118, 121 – Trono 2006: 65 – Mazziotti et al. 2008: 78 – Cecalupo et al. 2008: pl. 70, no. 19 – Vazzana 2010: 71; 2011: 60.

Mangelia leufroyi – Forbes & Hanley 1853: 468 pl. CXIII, figs 6, 7; pl. RR, fig. 1 – Malm 1855: 41 – Jeffreys 1856: 187; 1859: 199 – G. B. Sowerby II 1859: pl. 19, fig. 11 – Capellini 1860: 48 – Doderlein 1864: 102 – P. Fischer 1878: 189, no. 500 – Scalia 1900: 25; 1907: 37 – Zelinskaya et al. 1968: 224, pl. 50, figs 40, 41.

Mangilia (Clathurella) leufroyi – Cossmann 1896: 124.

Mangilia (Leufroyia) leufroyi – Dollfus & Dautzenberg 1886: 102.

Mangilia leufroyi – Friedberg 1912: pl. 14, fig. 8 – Meznerics 1933: 320 – Friedberg 1951: 222, pl. 14, fig. 8.

Peratotoma (Leufroyia) leufroyi – Sacco 1904: 53, pl. 13, figs 52, 53 – Cerulli-Irelli 1910: 59 (251), pl. 5 (36), fig. 46 – Cipolla 1914: 70 (174), pl. 3 (14), Fig. 24A, B.

Peratotoma leufroyi – Bałuk 2003: 73, pl. 29, fig. 12.

Philbertia (Leufroya) leufroyi – Thiele 1929: 370 – Steuer 1939: 44, fig. 18 – Devidts 1959: 104 – Priolo 1967: 702.

Philbertia (Leufroyia) leufroyi f. praecedens – Glibert, 1954: 60, pl. 7, fig. 8.

Philbertia leufroyi – Odhner 1931: 21 – Mars 1956: 44 – Pasteur- Humbert 1962: 104, pl. 39, fig. 169 – Powell 1966: 16, fig. 155F – Cadée 1968: 115 – Thiriot-Quiévreux 1972: 558, pl. 4, figs A-F – McMillan 1973: 56 – Spada et al., 1973: 55.

Pleurotoma (Clathurella) leufroyi – Watson 1897: 304, no. 249.

Pleurotoma inflata – sensu Philippi 1836: 197, pl. XI, fig. 24 non De Cristofori & Jan, 1832.

Pleurotoma lefroyi [sic] – Deshayes 1835: 275 – Forbes 1844: 157 – Aradas & Benoit 1876: 251, no. 665.

Pleurotoma leufroyi – Potiez & Michaud 1838: 445 – Kiener 1839: 70, pl. 24, fig. 3 – Reeve 1843: pl. 16, sp. 131 – Philippi 1844:165, pl. 11, fig. 24 – Requien 1848: 73, no. 522 – D’Orbigny 1852: 172 – Petit de la Saussaye 1852: 187; 1869: 153 – Hörnes 1854: 373, pl. 40, Fig. 16 A-C – Danilo & Sandri 1856: 137, no. 140s – Tallack 1861: 314 – Brusina 1866: 64 – Caruana 1867: 49 – Klećak 1873: 36 – Monterosato 1874: 277; 1875: 44, no. 744; 1878: 46; 1880: 230 – Nyst 1878:50 (1); 1881: pl. 3, Fig. 14A, B – Dewalque 1880: 478 – Locard 1891 b: 11 – Tesch 1912: 90 – Nobre 1931: 79, pl. 32, fig. 4.

Raphitoma (Leufroyia) leufroyi – Wenz 1943: 1453, fig. 4114 – Shikama & Horikoshi 1963: 123, pl. 101, fig. 5 – Brébion 1964: 611, pl. 15, fig. 5 – Nordsieck 1968: 179, pl. 30, fig. 94.70; 1977: 60, pl. 20, fig. 157 – De Casa & Hallgass 1979: 6 pl. 1 – Montefameglio et al. 1979: 191 – Piani 1980: 156 – Templado & Llanso 1981: 36 – Van Aartsen et al. 1984: 45, fig. 219 – Sabelli et al. 1990: 44, 216 – Cavallo & Repetto 1992: 26, 146, fig. 404 (WI) – Cossignani et al. 1992: 26, fig. 179 – Mifsud 1993: 6 – Vera-Peláez et al. 1999: 13.

Raphitoma concinna – Manousis 2012: 178 [top and low figures], non Scacchi, 1836.

Raphitoma erronea – Arduino et al. 1995: 94 (figured) non Monterosato, 1884.

Raphitoma leufroyi – Bellardi 1847: 89 – Sismonda 1847: 36 – Ruggieri 1967: 314 – Albergoni & Spada 1969: 158, 161 – Parenzan 1970: 210, pl. 45, fig. 853 – Spada 1971: 131 – Ghisotti 1972: 85 – Lozet J.-B. & Dejean-Arrecgros 1977:112, fig. 163 – D’Angelo & Gargiullo 1978: 153 (figured) – Biagi & Corselli 1978: 11 – Bogi, Coppini & Margelli 1980: 14, fig. 2 – Terreni 1981: 41, no. 336 – Corselli, 1981: 16 – Berardelli et al. 1982:330 – Idato, Fresi & Russo 1983: 112 – Poppe & Goto 1991: 44,167, pl. 35, fig. 22, pl. 1, fig. 25 – Barash & Danin 1992: 161, fig. 184 – Cachia, Mifsud & Sammut 1993: 34 – Delamotte & Vardala- Theodorou 1994: 137, fig. 7 – Arduino et al. 1995: 94 (figured) – Hayward, Wigham & Yonow 1995: 536, fig. 10.17 – Giribet & Peñas 1997: 53 – Oliverio 1997: 86 – Bogi & Cauli 1998: 134 – Cachia 1999: 90 – Olabarria, Urgorri & Troncoso 1999: 89 – Avila et al. 2000: 154 – Borja & Muxika 2001: 76 – Cachia, Mifsud & Sammut 2001: 67, pl. 10, fig. 5; 2004: 254 – Martini et al. 2001: 193 – Landau et al. 2003: 271, pls. 45, fig. 8 – Rolán 2005: 172, fig. 789 – Repetto et al. 2005: 39, 218, fig. 900 – Peñas et al. 2006: 43 – Oliver Baldoví 2007: 39 – Vardala-Theodorou & Nicolaidou 2007: 65 – Soppelsa et al. 2007: 27 – Cecalupo et al. 2008: 32 – Pusateri & Giannuzzi-Savelli 2008: 124, figs 5, 7, 9, 14 – Robin 2008: 454, fig. 16 – Scaperrotta et al. 2010: 96 (figured) – Cossignani & Ardovini 2011: 31, 326 (figured) – Gofas, Moreno & Salas 2011: 339 (figured) – Rolán 2011: 230, fig. 76L, M – Manousis 2012: 179 [only upper figure, lower one is L. concinna ] – Oliver et al. 2012: 55; 2015: 109 – Tro - no & Macrì 2013: 35 – Ceulemans et al. 2018: 113 – Manousis et al. 2018: 21, fig. 14A-C – Trigo et al. 2018: 355 – Muñoz Ferrera de Castro et al. 2019: 162-163 (figured).

TYPE MATERIAL. — Pleurotoma leufroyi Michaud : Potential syntypes. Mediterranean Sea • 2 sh; Lyon Museum 45018065; Mediterranean ; 13.1 and 15 mm long, glued with the apex pointing downward, on a cardboard labelled by Michaud (“PleuR. leufroyi Mich. /Médit.”) and by Locard (“ Defrancia Leufroyi Mich /méditerranée/coll. Michaud ”) ( Boyer & D’Audibert 2007: 151).

Pleurotoma zonalis Delle Chiaje : not found, it should be searched in the general collection at Museo Zoologico di Napoli, currently not revised (Sergio Duraccio pers. comm.).

Pleurotoma cyrilli O.G. Costa : not found, it should be searched in the general collection at Museo Zoologico di Napoli, currently not revised (Sergio Duraccio pers. comm.).

Murex caudicula Nardo : not available; it should be in the Stefano Chiereghini collections (Museo di Storia Naturale,Torino), currently not accessible (Paolo Russo pers. comm.).

Clathurella mirabilis Locard, 1891 :

Syntype. France • 1 sh; MNHN-IM-2000-3104; Belle Isle ; 18.6 mm .

TYPE LOCALITY. — Pleurotoma leufroyi Michaud : Mediterranean, “Habite la Méditerranée, côte d’Agde (Hérault)” [Lives in the Mediterranean, coast of Agde (Hérault): Michaud 1828].

Pleurotoma zonalis Delle Chiaje : Napoli.

Pleurotoma cyrilli O.G. Costa : Taranto.

Murex caudicula Nardo : Venezia.

Clathurella leufroyi var. albida Bucquoy, Dautzenberg & Dollfus : côtes de Barbarie.

Clathurella mirabilis Locard : l’Océan.

MATERIAL EXAMINED. — NE Atlantic. Portugal • 1 sh; Algarve, Marinha Portimão ; 16- 18 m; coll. PUS .

Canary Islands • 1 sh; Gran Canaria , La Luz ; 109 m; coll. PUS.

Açores • 1 sh; São Miguel Island , Ponta Delgada ; coll. DSC.

Mediterranean. Spain • 2 sh; Alboran Sea ; coll. OLI • 1 sh; Cabo de Gata , coll. CRO • 1 sh; Malaga ; coll. TRI • 1 sh; La Herradura ; 10 m; coll. AGA • 3 sh; Baleares ; unprecised locality; SMNH 73167 • 2 sh; Formentera I., El Calo ; leg. Alf Josefson; SMNH 73166 D. France • 2 sh; Antibes ; coll. HOA • 6 sh; Provence; HUJ coll. Coen 8092B • 1 sh; St. Raphael; labelled by Monts . “ leufroyi var. parva ”; MCZR-M-17111 • 1 sh; St. Raphael; coll. HOA • 2 sh; St. Raphael, Le Dramont ; coll. HOA • 1 sh; Cassis; MNHN coll. Locard .

Corsica • 8 sh; Baie de Calvi ; legit A. Warén, SMNH 73171 L • 1 sh; Ile Rouge ; 40 m; coll. BAR • 1 sh; Alistro ; coll. MAR • 1 sh; Ajaccio ; coll. CRO • 4 lv; Corsica 2019; Cap Corse ; MNHN- IM-2019-4036, MNHN-IM-2019-4093, MNHN-IM-2019-5302, MNHN-IM-2019-5315.

Sardinia • 3 sh; no. 8092A; no precise locality; HUJ coll. Coen • 3 sh; no precise locality; MCZR-M-17111 • 2 sh; S. Teresa di Gallura; 6 m; coll. CRO • 1 sh; Olbia ; coll. CRO • 1 sh; Porto Conte ; coll. OLI • 2 sh; Porto Conte, Nettuno cave; 15 m; coll. OLI • 14 sh; Poetto ; coll. PIS • 1 lv; Villasimius ; 39°07’43”N, 9°32’17”E; BAU-1742. Sicily • 1 sh; Porticello ; coll. GIR • 1 sh; Ustica I., Punta Gavazzi ; 50 m; coll. AGA • 3 sh; Palermo ; sub nomine leufroyi var. minor ; MCZR-M-17111 • 3 sh; Palermo; sub nomine “ Leufroyia malatina” Monts. ms; MCZR-M-17111 • 2 sh; Palermo ; “ var. albescens ” Monts ; MCZR-M-17111 • 1 h; Palermo Gulf ; coll. GIR • 8 sh; Palermo Gulf ; coll. PUS • 4 sh; Isola delle Femmine ; coll. PUS • 5 sh; Isola delle Femmine; coll. SER • 2 sh; Isola delle Femmine ; coll. GIR • 2 sh; Isola delle Femmine ; coll. CRO • 1 sh; Trapani ; coll. OCC • 2 sh; Favignana I. ; coll. PUS • 1 sh; Sicily Channel ; coll. TRI • 1 h; Lampedusa I., Punta Cappellone ; 45 m; coll. CRO • 1 sh; SE Sicily; coll. ARD • 1 sh; unprecised locality; MRSNT no. 45335 • 2 sh; Punta Braccetto ; coll. OCC • 3 sh; Porto Palo di Siracusa • 1 sh; coll. GER • Brucoli ; coll. CRO • 2 sh; Isola delle Correnti ; 15 m; coll. PIE • 4 sh; Cannizzaro ; 45 m; coll. GER • 3 sh; Can- nizzaro; 45 m; coll. CRO • 2 sh; Catania, Acireale ; 15 m; coll. PAG • 18 sh; Acitrezza ; coll. GER • 4 sh; Acitrezza; coll. CRO • 1 sh; Acitrezza ; coll. PAG.

Italy • 1 sh; Gallinara I. ; 40 m; coll. REP • 5 sh; Bergeggi ; 38 m; coll. REP • 1 sh; Capo Noli ; 2 m; coll. REP • 1 sh; Camogli , S. Frut- tuoso; 42 m; coll. REP • 1 sh; Sestri Levante, Riva Trigoso ; 20 m; coll. REP • 6 sh; Gorgona I. ; 80 m; coll. BAL • 1 sh; Capraia I. ; 80 m; coll. MAR • 8 sh; Capraia I.; 40 m; coll. PAO • 1 sh; Elba I. ; coll. MAR • 1 sh; Elba I.; coll. GOR • 1 sh; Elba I. ; coll. BAR • 1 sh; Elba I. , Buccale ; coll. PAG • 1 sh; Elba I. , Manaregno ; coll.TRI • 6 sh; Grosseto, Punta Ala ; 5 m; coll. REP • 1 sh; Grosseto; coll. OCC • 5 sh; Porto Santo Stefano ; coll. PAG • 7 sh; Gulf of Baratti; 5 m; coll. PAO • 7 sh; Gulf of Baratti; 5 m; coll. BAL • 1 sh; Livorno, Bagni Fiume ; coll. PAG • 1 sh; Livorno, Bagni Fiume ; coll. MAR • 1 sh; Livorno, Bagni Fiume ; coll. PAG • 6 sh; Castiglioncello ; coll. MAR • 1 sh; Vada ; coll. PAG • 1 sh; Ladispoli, Torre Flavia ; MCZR; coll. Pizzini • 2 sh; Ostia ; coll. PIE • 1 sh; off Fiumicino ; 30 m; coll. PAG • 7 sh; Santa Marinella ; 27 m; coll. AGA • 1 sh; Zannone I. ; coll. FUM • 1 sh; Ponza I., Secca dei mattoni; 26 m; coll. CRO • 23 sh; Pro- cida I.; coll. CRO • 4 sh; Procida I., Punta Pioppeto ; 6 m; coll. CRO • 3 sh; Napoli; var albolineata Monts ms.; MCZR-M • 1 sh; Sorrento, Marina di Puolo ; coll. DUR • 3 sh; Capri I. ; coll. CRO • 1 sh; Ce- traro; coll. RON • 3 sh; Costa Viola ; 38 m; coll. PAO • 4 sh; Scilla ; coll. GER • 67 sh; Scilla; coll. VAZ • 1 sh; Scilla ; 50 m; coll. PAG • 14 sh; Scilla ; 50 m; coll. CRO • 1 sh; Scilla ; 50 m; coll. TIS • 3 sh; Lazzàro ; coll. VAZ • 15 sh; Isola di Capo Rizzuto ; 45- 60 m; coll. MEL • 4 sh; Maratea ; coll. CAR • 5 sh; Campomarino ; coll. DIN • 1 sh; Porto Cesareo ; coll. MAC • 1 sh; Lama ; coll. NOT • 1 sh; Novaglie ; coll. MAC • 1 sh; Marina di Ugento ; coll. MAC • 1 sh; Monopoli, Cala Corvino ; coll. DUR • 2 sh; Giovinazzo ; coll. MEL • 2 sh; San Nicola I. ; coll. AGA • 1 sh; Ortona ; 8 m; coll. PAG.

Morocco • 2 sh; Restinga ; MRSNT no. 45337.

Malta • 1 sh; no precise locality; coll. MIF.

Croatia • 2 sh; Korcula I. ; 5 m; coll. AGA • 1 sh; Dalmatia ; coll. PUS • 2 sh; Zadar ; SMNH 70482 -70483 • 2 sh; Pag I., Košljun bay; 2- 5 m; coll. PRK • 2 sh; Pag I., Ljubačka Vrata ; 35 m; coll. UGA • 2 sh; Molat I. ; amidst red coral colonies, 35- 50 m; coll. PRK • 10 sh; Sukošan ; 2- 7 m; coll. PRK • 25 sh; Dugi Otok I., Veli Garmenjak ; 32- 40 m; coll. PRK • 37 sh; Murter I., Kosirina bay; 1- 7 m; coll. PRK • 2 sh; Žirje I. ; amidst red coral colonies, 70- 100 m; coll. PRK • 2 sh; Primošten, Šparadići ; 2- 6 m; coll. PRK • 1 sh; Sevid ; 18 m; coll. UGA • 33 sh; Sevid; 2- 8 m; coll. PRK • 6 sh; Jabuka I. ; 15- 35 m; coll. UGA • 5 sh; Jabuka I.; 15- 35 m; coll. STA • 2 sh; Čiovo I., Fumija ; 10 m; coll. UGA • 2 sh; Šolta I. ; 5- 10 m; coll. STA • 23 sh; Split ; 1- 5 m; coll. PRK • 1 sh; Vis I., Komiža ; 35 m; coll. UGA • 1 sh; Brač I., Milna ; 10- 18 m; coll. PRK • 13 sh; Brač I., Milna ; 10- 18 m; coll. STA • 2 sh; Brač I., Maslinova bay; 3- 15 m; coll. PRK • 15 sh; Brač I., Maslinova bay; 3- 15 m; coll. STA • 3 sh; Omiš, Balića Rat ; 4- 8 m; coll. PRK • 20 sh; Lastovo I. ; amidst red coral colonies; 70- 100 m; coll. PRK • 14 sh; Lastovo I.; amidst red coral colonies; 70- 100 m; coll. STA • 1 sh; Palagruža I. ; 28 m; coll. UGA • 2 sh; Pelješac, Žuljana ; 1 m; coll. STA • 2 sh; Pelješac, Prapratno ; 2- 6 m; coll. PRK • 5 sh; Mljet I., Pod Škoji ; 1- 3 m; coll.PRK • 32 sh; Mljet I.; amidst red coral colonies, 70- 100 m; coll. PRK • 1 sh; Zadar; 6 m; coll. PAG • 1 lv; Sevid ; 43°28’46”N, 16°02’08”E; 2- 4 m; BAU-2240.1.

Greece • 1 sh; Athens , Varkiza ; coll.TRI • 1 sh; Astypalea I. ; coll.TRI • 1 sh; Antiparos I. ; coll. BIN • 1 sh; Achaia ; 5 m; coll. PAG • 1 sh; Corfù I. ; coll. BAR • 2 sh; Chalkidiki, Sani ; coll. CRO • 1 sh; Crete Island, Mallia ; coll. CRO.

Turkey • 1 sh; Bozcaada I. ; coll. PUS • 1 sh; Adana, Yumurtalik , coll. CGS • 1 sh; Bodrum, Salih I. ; 35 m; coll. CGS • 1 sh; Bod- rum; 80 m; coll. CGS • 1 sh; Güllük Bay ; 44 m; coll. OZT • 1 sh; Marmara Sea, Silivri ; 25 m; coll. OZT.

DISTRIBUTION. — Known with certainty from the Early Pliocene to Recent; records from the Middle Miocene (Langhian of the Loire Basin: Peyrot 1938, Glibert 1954; Langhian of the Paratethys: Hörnes 1854, Bałuk 2003, Friedberg 1912; Serravallian of Karaman Basin: Landau et al. 2003) and from the Late Miocene (Messinian of the Loire Basin: Brébion 1964) although morphologically broadly similar are probably not conspecific (B. Landau pers. comm.), and need to be reanalysed in the framework of a revision of the genus Leufroyia in the Miocene.

Early Pliocene: central Mediterranean, Italy ( Sacco 1904). Early-late Pliocene: central Mediterranean, Italy ( Cipolla 1914).

Early Pleistocene: central Mediterranean, Italy (Cerulli-Irelli 1910). [From Landau et al. 2003].

Recent: Northeastern Atlantic from the Atlantic coast of France (Belle Isle), south to Portugal, Açores, Madeira and Canary Islands. The entire Mediterranean and Marmara Sea ( Ostroumoff 1896: 59). Leufroyia leufroyi is relatively common on rocky bottoms from a depth of 1 m to more than 100 m, where it lives under stones or amidst algae, but also in sciaphylous habitats, including amidst red coral colonies where it is also 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. concinna even under the same stone or amidst the same alga, while sampling in sciaphilous habitats all 4 Leufroyia species can be found together.

ORIGINAL DESCRIPTION. — “P. Testa turrito-conica; luteo-fusca, longitudinaliter costata, striis transversis decussantibus exarata; apice obtusa [sic], anfractibus septenis; convexis, postremo duabus, fasciis spadiceis interruptis, superioribus una tantum, ornais; sutura excavata; apertura alba; labro incrassato; canali brevi, recto, extus eleganter plicato” ( Michaud 1828:121).

DESCRIPTION

Shell

Robust and fusiform, of large size for the genus. Height: 10.83- 22.93 mm (mean 15.26 mm, SD: 3.07); width: 5.07-9.59 mm (mean 6.76 mm, SD: 1.12); H/W: 2.0-2.62 (mean 2.27, SD: 0.13).

Protoconch

Multispiral with 2.6 to 3.0 (mean 2.84, SD: 0.14) convex whorls and small nucleus (d: 150 µm). Protoconch I of 0.9 whorls, diameter 242 µm, covered by dense cancellate sculpture; protoconch II with axial threads under suture and less dense and diagonally cancellate sculpture on rest of whorl. A keel at the end of last whorl of varying length. Protoconch-teleoconch boundary of flexuose, opisthocline growth lines. Colour always dark brown or blackish, with white or yellowish nucleus.

Teleoconch

Of 5.8-8.0 convex whorls (mean 6.87, SD: 0.58), with deep and distinct wavy sutures and prominent sculpture; suture area slightly thickened abapically and white coloured with brown dots or lines; whole external surface covered with relatively thick periostracum; dense and slightly pronounced growth lines present on ribs and interspaces, more pronounced on the subsutural ramp; surface glossy; microgranules absent on 1-2 adapical whorls, remaining ones covered with extremely fine and dense microgranulation.

Axial sculpture of 12-19 (mean 14.16, SD: 1.73) strong, equidistant, orthocline or slightly opisthocline ribs, broader than spiral cords, narrower than interspaces; ribs occasionally less elevated on body whorl of largest shells.

Spiral sculpture of 10-17 narrow cords (mean 13, SD: 1.75) 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, rarely one thin cordlet between all pairs of strong cords; spiral interspaces 1.5-3 broader than cords; on first adapical whorl 2 or 3 strong spiral cords and one thin subsutural cordlet.

Siphonal fasciole with 6-11 (mean 8.21, SD: 1.11) strong and slightly nodulose cords, usually stronger than cords of body-whorl.

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

A B D C

Subsutural ramp narrow and more or less 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 moderately long, posterior canal deep and moderately wide.

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

Height of aperture: 4.6-9.99 mm (mean 6.24 mm, SD: 1.40); A/H: 36.98-45.97% (mean 40.83%, SD: 2.46).

Coloration

Background colour variable but usually light, from whitish to pale brownish; darker blotches, light brown to dark brown-blackish, variable in number (rarely absent), size and colour, irregularly placed on upper half of whorls and below periphery, usually on axial ribs, occasionally also in interspaces; a broad light spiral band almost devoid of blotches at aproximately 1/3 to 1/2 of whorl height. Occasionally a thin brown or reddish-brown cordlet above the median ligth band. Cords in light coloured areas often with many brownish dots and short lines, more evident in living specimens. Cords on siphonal fasciole light, whitish or yellowish, often with some light brown or pale orange narrow bands in the interspaces between low tubercles. Aperture internally white, often with light or dark brown blotches visible by transparency.

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.

Colour translucent white or cream, with minute white speckles densely covering upper side of foot, scattered or absent on sole, head and along neck. Tentacles translucent white or cream, with white speckles only along broadest part. Siphon translucent yellowish-white, pale ochre or pale orange, with dense white speckles ( Figs 6 , 7 ).

REMARKS

The collection Michaud was in part donated to the Musée de Lyon (now Musée des Confluences), in part passed to the city of Brive by Michaud’s son Elysée, and in part bought by Locard ( Locard, 1891 b: 7). Only the Lyon material remains identifiable ( Boyer & D’Audibert 2007: 13; pl. 1, fig. 2), the part in Brive having been lost, and the part in Locard’s collection having lost the original labels. The possible syntypes of Pleurotoma leufroyi Michaud remaining in Lyon Museum (lot no. 45018065) consist of 2 shells, with handwritten labels by Michaud and Locard. This is the same material referred to by Locard (1891 b: 11). Michaud gave as size 20 x 9 mm, making the identification of the Lyon material as syntypical still not conclusive.

Leufroyia leufroyi was mistakenly confused by many authors with L. concinna . The first source seems to be that of Forbes & Hanley (1853: 468) who nonetheless pointed out that their English specimens of “ leufroyi ” (evidently L. concinna ) were different from the Mediterranean ones (clearly typical L. leufroyi ) by being broader, with more obsolete colouration and by having some spirals cords of brownish colour. Also Jeffreys (1867: 366) recorded “ leufroyi ” from the British coast but his description seems to refer to a mix of characters of L. leufroyi and L. concinna . Other more recent misidentifications are e.g. in Fretter & Graham (1985: 539) and Wigham & Graham (2018: 137, fig. 161).

The maximum height of examined shells is 23 mm, but specimens larger than 18 mm are rare. The height and maximum diameter of the protoconch depend on the number of whorls; in the specimen of Fig. 8 with 2.7 protoconch whorls the protoconch is 525 µm high and 484 µm wide. The diameters of nucleus and first whorl are small, similar to L. concinna , and smaller than in L. erronea and L. villaria . The colour of the protoconch is always dark brown or blackish, even in specimens with very light teleoconch. The sexes are separated but we did not find any evidence of bimodality in morphometrics, thus guessing no size dimorphism. Diagnostic features of L. leufroyi with all other extant Leufroyia spp. are the very dark protoconch and the dark brown blotches on a light background of the teleoconch (although shells tend to fade out after some years from collection). Some problematic specimens with small and very pale or even absent blotches on the teleoconch can be easily diagnosed by the dark protoconch (if retained). Some dark coloured shells of L. erronea may be misleading, but they can be easily identified by the lighter and broader protoconch, the teleoconch spiral sculpture, less slender outline and shorter siphonal canal.

According to HØisaeter (2016: 28) L. leufroyi is not present along the European coasts North of Brittany ( France) and all such records (like e.g. Lebour 1934: 553 from Plymouth, UK) should be in fact ascribed to L. concinna . As far as we know the northernmost confirmed record is Belle Isle (Bay of Biscay, Brittany, France) from the Locard coll. (MNHN- IM-2000-3104) under the name Clathurella mirabilis , whereas the southern limit should be at Canary Islands.

The records of L. leufroyi from southern Angola ( Rolán et al. 1998: 108) are based on L. villaria , according to the different pattern of the living animal (cream colour with small white spots, siphon clear orange, with many light coloured spots) and the slightly different protoconch. It is quite common in the whole Mediterranean, including Israel from where it was recorded by Barash & Danin (1992) from various localities. Marquet (1998 b) correctly separated the Pliocene North Sea Basin records as Raphitoma pseudoleufroyi ( Marquet, 1998) , a species with very similar teleoconch characters, but paucispiral protoconch (vs multispiral in L. leufroyi ).

Figures

FIG. 4. — Leufroyia leufroyi (Michaud,1828): A, original engravings; B, potential syntypes,MDCL (‘Méditerranée’) 13.1 and 15 mm; C, potential syntype,13.1 mm; D, original label by Locard; E, Murter Island (Croatia), 15.4 mm (H/D = 2.54); F, Murter Island (Croatia), 15.2 mm (H/D = 2.10); G, Gulf of Termini Imerese (Palermo - Italy), h. 22.71 mm (with 19 ribs); H, Isola delle Femmine (Palermo - Italy), h. 14 mm (specimen with 10 axial ribs on last whorl). Photo courtesy: B-D, Cédric Audibert; G-H, Ignazio Sparacio.

FIG. 5. — Leufroyia leufroyi (Michaud, 1828): A, Sevid (Croatia) 18 m, h. 17 mm; B, Murter Island (Croatia) 3 m, h. 18.2 mm; C, Murter Island (Croatia) 3 m, h. 15.5 mm; D, Saronic Gulf (Greece), h. 15.8 mm; E, Le Dramont (Saint Raphael, France), h. 15.8 mm; F, Secca del Frate, Palmaiola Island (Tuscan Archipelago), h.15.2 mm; G, Clathurella mirabilis Locard,1891,syntype (MNHN),Belle Isle (Brittany,France),h. 18.6 mm;H, Tuscan Archipelago,h. 20 mm;I, Tuscan Archipelago, h. 15.5 mm; J, Split (Croatia) 3 m, h. 9 mm; K, Mljet Island (Croatia),60-100 m h. 6.1 mm; L, Ponta Delgada (São Miguel Island,Açores), h: 2.3 mm; M, Elba Island (Italy), h. 0.8 mm. Photo courtesy: A. Pero Ugarković; D. Costas Kontadakis; E. Dominique Horst.

FIG. 6. — Photographs of two living animals of Leufroyia leufroyi (Michaud, 1828). A-D, Split, Croatia, 3 m, h. 10 mm; E-F, Čiovo Island, Croatia, 5 m, h. 6 mm. Photo courtesy: E-F, Pero Ugarković.

FIG. 7. — Underwater photographs of three living animals of Leufroyia leufroyi (Michaud, 1828). A-C, France, Cap d’Antibes, 13-17 m. Photo courtesy: Dominique Horst.

FIG. 8. — Leufroyia leufroyi (Michaud, 1828). Murter Island (Croatia), 3 m, h. 7.85 mm. A, Frontal view of shell; B-D, apical (B), frontal (D) and dorsal (C) view of protoconch. Scale bars: 2 mm (A), 300 µm (B-D).

FIG. 16. — Leufroyia erronea Monterosato, 1884. Žirje Island (Croatia), 70-100 m, amidst red coral colonies, h. 9.32 mm. A, Frontal view of shell. B-D, apical (B), frontal (D) and dorsal (C) view of protoconch. Scale bars: 2 mm (A), 300 µm (B-D).

FIG. 14. — Leufroyia erronea Monterosato,1884:A, Sardinia (Italy),h. 15.8 mm,with original labels by Tiberi and Monterosato;B, sine loco, h. 10.5 mm;C, Gorgona Island (Italy), h. 12.5 mm; D, Spain, h. 24 mm; E, Capraia Island (Italy) 120 m, h. 27 mm; F, Latium (Italy),h. 15.5 mm; G, Capo Corso (Italy), h. 8.5 mm; H, Mljet Island (Croatia) 70-100 m, h. 10.6 mm; I, Mljet Island (Croatia) 70-100 m, h. 15.7 mm. Photo courtesy:D, Peter Stahlschmidt, E, Enzo Campani, H, Morena Tisselli.