Leufroyia erronea Monterosato, 1884

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 : 454-458

Leufroyia erronea Monterosato, 1884

( Figs 14-16 )

Defrancia leufroyi var. coralligena Monterosato, 1872: 51 (nomen nudum).

? Defrancia convexa Jeffreys, 1882: 33 .

Leufroyia erronea Monterosato, 1884: 134 .

Clathurella erronea – Locard & Caziot 1899: 63 (? L. villaria ).

Comarmondia inflata – sensu Chirli 1997: 91, pl. 26, figs 5, 6 non De Cristofori & Jan, 1832.

Leufroyia erronea – Carus 1893: 428 [in synonymy with Clathurella inflata ( De Cristofori & Jan, 1832) ] – Cipolla 1914: 72 (176) [in synonymy with Peratotoma (Leufroyia) inflata sensu Cipolla 1914 non De Cristofori & Jan, 1832] – Pallary 1900: 257 – Van Aartsen 1988: 142.

Pleurotoma (Leufroyia) leufroyi erronea – Kobelt 1905: 366, no. 22. Pleurotomella demosia – sensu Bogi 1986: 27-28 non Dautzenberg & Fischer, 1896 – sensu Crocetta & Spanu 2008: 67, 72, fig 3A, B non Dautzenberg & Fischer, 1896 – sensu Giribet & Peñas 1997: 52, Figs 66-68 non Dautzenberg & Fischer, 1896 – sensu Negri & Corselli 2016: 69, fig. 15Q-T non Dautzenberg & Fischer, 1896 – sensu Scaperrotta et al. 2012: 99 non Dautzenberg & Fischer, 1896.

Raphitoma (Leufroyia) erronea – Piani 1980: 156 – Sabelli et al. 1990: 44, 216.

Raphitoma (Leufroyia) leufroyi erronea – Nordsieck 1968: 179.

Raphitoma (Leufroyia) sp. De Casa & Hallgass, 1979: 11 pl. 1 (3-4).

Raphitoma erronea – Poppe & Goto 1991: 44, 174 – Peñas & Giribet 2003: 181 – Repetto et al. 2005: 39, 218, fig. 895 (uncertain, very badly figured) – Pusateri & Giannuzzi-Savelli 2008: 124, fig. 15 – Cossignani & Ardovini 2011: 31, 325 (figured) – Appolloni et al. 2018: 60-61, 112, fig 22M, N – Ceulemans et al. 2018: 113 – Manousis et al. 2018: 21, fig.11A-E.

TYPE MATERIAL. — Leufroyia erronea Monterosato : Syntypes. Sardinia • 1 sh; Sardinia, unprecised locality; Tiberi’s handwriting label, “ Defrancia volutella Valenciennes Sardegna”, H: 15.8 mm, W: 7.5 mm, H/D = 2.1; MCZR-M-16704-L. Mediterranean • 1 sh; unprecised locality; Monterosato handwritten label “ Leufroya erronea, Monts. in Nom. gen. e sp. p. 126 Adr. (Stossich) Palermo (Monts)”, H: 10.2 mm, W: 4.9 mm. Defrancia convexa Jeffreys : type material lost ( Warén 1980: 33); MCZR-M-17340-L.

TYPE LOCALITY. — Leufroyia erronea Monterosato : Originally described from Corsica (France), Sardinia, Palermo to San Vito lo Capo (Sicily) and Dalmatia (Croatia). Defrancia convexa Jeffreys : West of Italy.

MATERIAL EXAMINED. — Atlantic. Great Britain • 7 sh; Shetland Islands; USNM Jeffreys coll. • 1 sh; Moray Firth; under leufroyi ; NMW 01065.

Spain • 1 sh; Galicia; coll. HOU.

Açores • 1 sh; São Miguel I., Ponta Delgada; 18 m, Josephine exp. 1869 • 1 sh; SMNH 70501; São Miguel; SMNHN 70499.

Madeira • 2 sh; unprecised locality; sub nomine Defrancia leufroyi with handwritten label by Watson; MCZR-M 16704.

Mediterranean. Spain • 1 sh; unprecised locality; coll. GUB • 1 sh; Malaga; coll. PUS.

Corsica • 1 sh; Capo Corso; coll. BAR.

Sardinia • 5 sh; Alghero; coll. SPM.

Sicily • 1 sh; Isola delle Femmine; coll. PUS.

Italy • 1 sh; Gorgona I.; coll. PAG • 1 sh; Capraia I.; coll. CAM • 1 sh; Livorno, Bagni Fiume; coll. PAG • 1 sh; off Fiumicino; 300 m; coll. PAG • 2 sh; Napoli; Tiberi label; MCZR-M-1730.

Croatia • 4 sh; Molat I.; 35-50 m; coll. PRK • 3 sh; Žirje I.; 70-100 m; coll. PRK • 78 sh; Lastovo I.; 70-100 m; coll. PRK • 43 sh; Lastovo I.; 70-100 m; coll. STA • 110 sh; Mljet I.; 70-100 m; coll. PRK • 6 sh; Mljet I.; 70-100 m; coll. TIS • 3 sh; Dugi Otok I., Veli Garmenjak; 32-40 m, rocky bottom; coll. PRK • 1 sh; Voluja; 40-60 m, rocky bottom; coll. PRK • 3 sh; Jabuka I.; 30 m, rocky bottom; coll. STA • 14 sh; off Dubrovnik; 300-380 m, soft bottom; coll. LET.

Greece • 1 sh; Saronic Gulf; coll. KON.

DISTRIBUTION. — North East Atlantic from Great Britain to Portugal, and the whole Mediterranean Sea. More frequently collected on rocky bottoms, from a depth of ca. 30 m to more than 100 m, than dredged from deep soft bottoms (100-400 m). In Croatia, L. erronea is the commonest raphitomid amidst the red coral colonies. Often found in sympatry/syntopy with other raphitomids, also with other Leufroyia spp.

ORIGINAL DESCRIPTION. — “= P. volutella , (non Valenc.) auct. =? P. fortis, Forbes — Rep. 1843, p. 196 — Reeve t. 19, f. 165 (Mar Egéo). Coralligena, rarissima in Corsica e Sardegna (Tiberi); Palermo a S. Vito (Monts.); Dalmazia (Brusina)” ( Monterosato 1884: 134). DESCRIPTION

Shell

Solid and suboval, of large size for the genus. Height: 9.04- 20.8 mm (mean 13.52 mm, SD: 3.35); width: 4.4-8.8 mm (mean 6.28 mm, SD: 1.36); H/W: 1.95-2.36 (mean 2.15, SD: 0.11). Protoconch

Multispiral with 2.55 to 3.1 (mean 2.82, SD: 0.15) convex whorls and large nucleus (d: 172 µm). Protoconch I of 1.15 whorls, diameter 315 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 light, whitish to brown, occasionally darker brown, never very dark or blackish; with or without white nucleus.

Teleoconch

Of 4.7-7.15 convex whorls (mean 5.88, SD: 0.68), with deep and distinct wavy sutures and prominent axial sculpture; suture area often slightly thickened abapically; periostracum not observed; whole surface covered with fine and dense axial growth lines and extremely fine microgranules, less dense or scattered on 1-2 adapical whorls and very dense on others.

Axial sculpture of 11-22 (mean 14.89, SD: 2.41) strong, equidistant, orthocline or slightly opisthocline ribs, broader than spiral cords, narrower than interspaces; ribs occasionally very weak and low on body whorl, sometimes also on penultimate whorl, in largest shells.

Spiral sculpture of 12-26 narrow cords (mean 18.18, SD: 3.77) above the aperture, of which 3-4 weak on subsutural ramp, remaining never equally strong due to variable number of thinner cords (number of thinner cords lower than or equal to that of strong ones); interspaces narrower, equal or slightly broader than stronger cords; on first adapical whorl 3 spiral cords equal in strength and one thinner subsutural cordlet.

Siphonal fasciole with 8-16 (mean 11.15, SD: 1.87) strong and nodulose cords, stronger than cords on body-whorl, thinner cords often present between strong ones.

Cancellation rectangular, with elongate and slightly elevated tubercles at the intersections. When the ribs are weak and low, tubercles are also very weak; tubercles usually stronger on first 3-4 whorls.

Subsutural ramp narrow and inclined, covered with dense growth marks of the anal sinus, crossed by 3-4 weak spiral cordlets; ramp on first whorls less inclined; ramps less inclined in shells with strong ribs.

Columella simple, straight medially and angled posteriorly. Siphonal canal very short and wide, posterior canal deep and wide.

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

Height of aperture: 3.8-8.4 mm (mean 5.65, SD: 1.35); A/H: 37.69-46.56% (mean 42.36%, SD: 2.02).

Coloration

Background colour variable but usually light, from whitish to light brown or reddish-brown, rarely dark brown, with areas or blotches of different colour, rarely uniformly coloured. One cord lighter than background, usually white-whitish, above suture, at approximately 1/3 to 1/2 of whorl height, the area above the cord often darker than the rest. Darker blotches and areas irregularly placed on axial ribs and interspaces on upper part of each whorl, sometimes below periphery. Cords on siphonal fasciole always same light colour as background, occasionally with some pale brownish lines in the interspaces. Aperture internally white or beige, often with some light brownish blotches visible by transparency.

Soft parts

Unknown.

REMARKS

We suspect that Defrancia convexa Jeffreys, 1882 may be this species but the type material is lost ( Warén 1980: 33). Description (based on a single specimen), size and colour fit relatively well a L. erronea with obsolete ribs. But it could be also a L. villaria for its “longer spire ... and the canal is not so short and abrupt” ( Jeffreys 1882: 33). Therefore, we consider for the moment Defrancia convexa as nomen dubium.

Leufroyia erronea was proposed by Monterosato (1884: 134) to distinguish a specimen received around 1878 by Tiberi, as Pleurotoma volutella Kiener, 1839 ex Valenciennes ms. ( Monterosato 1878: 106), considered to be a distinct species. Only in 1884, after the acquisition of Tiberi collection, Monterosato realized that misidentification and named erronea this species (see also Carus 1893: 428; Cipolla 1914: 72). The misidentification by Tiberi is quite strange as his specimen is very different from Kiener’s P. volutella (see Fig. 21D ).

The height and maximum diameter of the protoconch depend on the number of whorls; in the specimen of Fig. 16 , with 3 protoconch whorls, the protoconch is 715 µm high and 596 µm wide. The diameter of nucleus and first whorl are quite large and similar to L. villaria , larger than in L. leufroyi and L. concinna .

The maximum height of examined shells is 27 mm, but shells larger than 18 mm are rare. The largest shell (27 mm) found so far was dredged from a depth of 120 m at Capraia Island, Italy ( Fig. 14E ).

Diagnostic features of L. erronea with all other Leufroyia spp. are the suboval profile, the wide aperture, the very short siphonal canal and the stronger and less spaced spiral cords. It differs from L. concinna also in the lack of brown coloured cords and pinkish/purple areas, and in its larger protoconch. From L. leufroyi it differs in the lack of dark brown or blackish blotches and in its larger protoconch, which is never very dark or blackish. L. villaria has a more slender shell, more stepped whorls and thinner spiral cords. After some years from collection the coloration tends to fade out, with shells becoming lighter and almost uniformly coloured.

The specimen MCZR-M-16704 from Sardinia, labeled by Tiberi as Defrancia volutella , has been figured by Pusateri & Giannuzzi-Savelli (2008: 123, fig. 15, as “ holotype ” and with wrong measures) and by Appolloni et. al. (2018: 112, fig. 22M, N, with wrong catalog number).

Specimens of L. erronea have been frequently identified erroneously as Pleurotomella demosia (Dautzenberg & Fischer, 1896) an Atlantic species, the actual presence of which in the Mediterranean is very doubtful ( Fig. 15I ). Small specimens of L. erronea have strong axial and spiral sculpture ( Fig. 15 G-H), making confusion easier with similarly sized specimens of P. demosia : the two species have protoconch with similar ranges of whorls number (2.5-3, see Bouchet & Warén 1980: fig. 226) but P. demosia has a lower protoconch (H: 550 µm [Bouchet & Warén 1980] v. 650-720 µm in L. erronea ). In fact, several Mediterranean records of P. demosia (e.g. Bogi 1986; Giribet & Peñas 1997; Crocetta & Spanu 2008; Scaperrotta et al 2012; Negri & Corselli 2016) are based on misidentification with L. erronea . Adriatic and Aegean specimens generally are smaller and more vividly colored than those in the central-western basin.

Figures

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.

FIG. 15. — Leufroyia erronea Monterosato, 1884. A, Syntype, Shetland Islands (USNM 190581 ex Jeffreys coll.), h. 14.8 mm; B, Shetland Islands (USNM 190582 ex Jeffreys coll.), h. 14.5 mm;C, Shetland Islands (USNM 190578 ex Jeffreys coll.), h. 19.4 mm; D, Galicia (Spain),h. 16.2 mm;E, Mljet Island (Croatia),70-100 m, amidst red coral colonies, h. 9.1 mm; F, Mljet Island (Croatia), 70-100 m, amidst red coral colonies, h. 8.3 mm; G, Mljet Island (Croatia), 70-100 m, amidst red coral colonies, h. 5.3 mm; H, Saronic Gulf (Greece), h. 6.15 mm - scale bar 200 µm. I, Pleurotomella demosia (Dautzenberg & Fischer, 1896), Açores (MNHN), 1300 m, 38°33’21”N, 30°28’54” W, h. 7.3 mm. Photo courtesy: A-C, Jolanda Villacampa (USNM); H, Costas Kontadakis.

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. 21. — A, Pleurotoma inflata De Cristofori & Jan, 1832, Lectotype, (MCSNM i 4296) Tabiano (Parma) Piacentian (Pliocene), h. 15.3 mm (after Pinna & Spezia, 1978, pl. 40); B, P. inflata, Ficarazzi (Palermo) (Lower Pleistocene),h. 18.9 mm (NMW,12930), erroneously identified by Monterosato as Leufroyia volutella; C, P. inflata, Monte Pellegrino (Palermo) (Plio-Pleistocene) (MCZR-M-16706), h. 23.9 mm; D, Pleurotoma volutella Kiener,1839, original drawings by Kiener, h. 25.4 mm; E, P. volutella, Ficarazzi (Palermo) (Lower Pleistocene), h. 21.5 mm; F, Pleurotoma cf. volutella Monte Pellegrino (Palermo) (Pliocene), h. 25.8 mm.