Leufroya villaria (Pusateri & Giannuzzi-Savelli, 2008)

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 : 459-466

Leufroya villaria (Pusateri & Giannuzzi-Savelli, 2008)

( Figs 17-20 )

Raphitoma villaria Pusateri & Giannuzzi-Savelli, 2008: 119 , figs 1-4, 6, 8, 10-12, 18.

Peratotoma (Leufroyia) inflata – sensu Cipolla 1914: 72 non De Cristofori & Jan, 1832.

Philbertia leufroyi – sensu Knudsen 1952: 173, pl. 2, fig. 11; 1956: 527, pl. 2, figs 16, 17 non Michaud, 1828.

Pleurotoma volutella – sensu Aradas & Benoit 1876: 251 non Kiener, 1839 – var. B sensu Brugnone 1862: 27 non Kiener, 1839.

Pleurotomella sp. – Peñas et al. 2006: 126, figs 274-276.

Raphitoma (Leufroyia) erronea – sensu Mifsud 2003: 44, fig. 15 non Monterosato, 1884.

Raphitoma cf. erronea – sensu Cachia et al. 2001: 66, pl. 10, fig. 3 non Monterosato, 1884.

Raphitoma leufroyi – sensu Ardovini & Cossignani 2004: 38, 225 [only the fourth of first row] non Michaud, 1832 – sensu Rolán et al. 1998: 108, figs 26-28 non Michaud, 1828.

Raphitoma villaria – Cossignani & Ardovini 2011: 328 (figured) – Manousis et al. 2018: 24, fig. 18A-E.

TYPE MATERIAL. — Holotype. Sicily • 1 sh; Taormina ; 15 m; H: 17.3 mm, W: 6.8 mm; MNHN-IM-2000-27244 .

Paratypes. Malta • 1 sh; Ras il-Wahx; 80-100 m; paratype A, H: 11.5 mm, W: 4.6 mm; MNHN-IM-2000-23353 • 1 sh; Ras il- Wahx; 80-100 m; paratype B, H: 14.4 mm, W: 5.6 mm; coll. MIF . Italy • 1 sh; Capraia I.; 150 m; paratype C, H: 16.9 mm, W: 6.5 mm; coll. GOR .

Sicily • 1 sh; Carini Bay ; paratype D, H: 12 mm, W: 4.8 mm; coll. PUS .

Sardinia • 1 sh; unprecised locality; paratype F, H: 9.3 mm, W: 4.7 mm; sub nomine Pleurotoma (Defrancia) volutella with Tiberi’s handwritten label); SMNH 70490 .

Greece • 1 sh; Antiparos I.; paratype E, H: 8.6 mm, W: 3.9 mm; coll. BIN .

TYPE LOCALITY. — Taormina (Messina), bioclastic sediments.

MATERIAL EXAMINED. — Atlantic. Angola • 1 sh; Ilha de Luanda; 40-60 m; leg. S. Gofas, as R. leufroyi ; MNHN-IM-2010-11650 .

São Tomé & Príncipe • 1 sh; Príncipe I., S. Antonio beach; coll. Marche Marchad, as R. leufroyi ; MNHN-IM-2010-11655.

Ivory Coast • 11 sh; Region d’Abidjan; as R. leufroyi .; MNHN Paris IM-2010-11653, MNHN-IM-2010-11651.

Gambia • 1 sh; SW Cap de Bald ; 18 m, as R. leufroyi ; MNHN- IM-2010-11656 .

Senegal • 1 sh; Gorée Bay ; coll. Knudsen, as Philbertia leufroyi ; MNHN • 1 sh; Region de Dakar; 95 m, coll. Marche Marchad as R. leufroyi ; MNHN-IM-2010-11654 • 1 sh; Sud Ile Madeleine ; 48 m, as R. leufroyi ; MNHN-IM-2010-11653 .

Mediterranean. Spain • 1 sh; unprecised locality; coll. STH • 1 sh; Malaga; 80-150 m; coll. PUS.

Sicily • 1 sh; Marzamemi; coll. MAC • 1 sh; Cannizzaro; coll. BOG • 1 sh; Taormina; coll. PUS.

Italy • 2 sh; Capo Rizzuto; coll. ARD • 1 sh; Porto Cesareo; 70 m; coll. FIO.

Malta • 18 sh; Gnejna Bay ; 80-100 m; coll. MIF • 5 sh; Qammich ; 80 m; coll. MIF • 1 sh; Fomm ir-Riħ; 140 m; coll. PRK • 1 lv; Ras il-Wahx; 80-100 m; BAU 3590.1 .

Lybia • 1 sh; Misurata; 60-80 m; coll. MIF.

Turkey • 1 sh; Bozcaada I.; coll. BAR.

Greece • 1 sh; unprecised locality; coll. PUS • 6 sh; Saronic Gulf; coll. KON.

Croatia • 2 sh; Kvarnerić; 60-90 m; coll. UGA • 2 sh; Dugi Otok Island, Veli Garmenjak; 35-57 m; coll. PRK • 2 sh; Voluja; 40- 60 m; coll. PRK • 1 sh; Jabuka I.; 30 m; coll. STA • 1 sh; Hvar Channel; 80 m; coll. STA • 1 sh; Lastovo I.; 200 m; coll. UGA • 1 sh; Lastovo I.; coll. BAR • 2 sh; Lastovo I.; amidst red coral colonies, 70-100 m; coll. STA • 27 sh; Mljet I.; red coral colonies; 70-100 m; coll. PRK • 12 sh; Mljet I.; red coral colonies; 70-100 m; coll. LET • 2 sh; Mljet I.; red coral colonies; 70-100 m; coll. TIS • 2 sh; off Dubrovnik; 350-380 m; coll. LET.

DISTRIBUTION. — East Atlantic ( Senegal to Angola) and the Mediterranean Sea, from Spain to the Aegean Sea. More frequently collected by dredging on soft or mixed bottoms than on rocky bottoms, live specimens from ca. 20-30 m to 400 m. Most Croatian specimens were found amidst red coral colonies (70-100 m depth). It is frequently found in sympatry/syntopy with other raphitomids, also with other Leufroyia spp.

ORIGINAL DESCRIPTION. — “Protoconch multispiral, of 3.25 whorls [sic]. Protoconch I of 1.2, dome shaped whorls, with a sculpture of 8-9 spiral threads, covered by minute tubercles; tubercles tending to become oblong and anastomosing, giving rise to a somewhat clathrate pattern. Protoconch II of slightly more than 2 convex whorls, with a sculpture of only subsutural axial threads on the adapical third, tending to curve adaperturally. On the lower two thirds sculpture of diagonally crossing threads. Sinusigera outer lip at the protoconchteleoconch boundary.Teleoconch of 6-7 whorls, with stepped spire. Subsutural ramp narrow, suborizontal, tending to obliterate after the sixth whorl. Adsutural marks of the anal sinus visible on the ramp. Axial sculpture of 15-18 ribs, narrower than the interspaces, reaching the base. Spiral sculpture of numerous continuous cordlets, 13-16 above the aperture, regularly spaced, on the body whor; 10- 12 strong cords on the siphonal canal. Aperture suboval, tapering anteriorly. Outer lip, internally smooth. Anal sinus as deep as the interspace between two axial ribs. Inner lip smooth, arcuate posteriorly, straigth medially. Siphonal canal short, open. Background colour yellowish with a ligther narrow band on the lower third of the spire. The spiral cordlets within this band may be slightly stronger that the others. Animal with a short head and two short tentacles. Eyes placed on the external, thickened basal part of the tentacles, at about one third of their total height. Foot narrow and long with two anterior lateral triangular lobes and a pointed posterior end. Colour light yellow with a lighter coloured foot, darker tentacles and black eyes. Siphon much darker (nearly orange) in colour. Operculum absent.” (Pusateri & Giannuzzi-Savelli 2008: 120).

DESCRIPTION

Shell

Moderately solid, slender, fusiform, of large size for the genus. Height: 10.62-19 mm (mean 14.56 mm, SD: 2.86); width: 4.33-7.94 mm (mean 5.77 mm, SD: 1.02); H/W: 2.29- 2.89 (mean 2.51, SD: 0.14).

Protoconch

Multispiral with 3 to 3.25 (mean 3.1, SD: 0.03) convex whorls and large nucleus (d: 170 µm). Protoconch I of 1.1 whorls, diameter 302 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 or absent. Protoconch- teleoconch boundary of flexuose, opisthocline growth lines. Colour always light, from whitish to light brown, never dark brown or blackish; with or without white nucleus.

Teleoconch

Of 5.5-7.5 (mean 6.57, SD: 0.59) convex and more or less stepped whorls, with deep and distinct sutures and prominent axial sculpture; periostracum very thin, hardly visible and not persistent; whole surface covered with fine and dense axial growth lines and extremely fine microgranules, scattered on 1-2 adapical whorls and very dense on others.

A B D C

Axial sculpture of 12-17 (mean 14.5, SD: 1.54) strong, equidistant, orthocline or opisthocline ribs (often on the same shell), broader than spiral cords and usually narrower than interspaces.

Spiral sculpture of 13-24 (mean 16.55, SD: 2.33) very narrow and low cords above the aperture, of which 3-4 very weak on narrow subsutural ramp, the most adapical cordlet well spaced from suture and often slightly stronger than other cordlets. Remaining cords equally strong and regularly spaced, often with a variable number of thinner cordlets between pairs of stronger ones; interspaces 3-5 times broader than cords; on first adapical whorl 5 thin spiral cords, the most adapical thinner than others.

Siphonal fasciole with 9-12 (mean 10.35, SD: 1.0) moderately strong, well spaced and slightly nodulose cords.

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

Subsutural ramp narrow, subhorizontal on first whorls, concave and slightly inclined on lower ones, covered with dense growth marks of the anal sinus, crossed by 3-4 thin spiral cordlets.

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

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

Height of aperture: 4.3-7.03 mm (mean 5.92 mm, SD: 0.96); A/H: 35.93-43.46% (mean 39.61%, SD: 2.21).

Coloration

Background colour variable but always light, from whitish to light brown, with areas, blotches and lines of different colour, occasionally uniformly coloured; first two teleoconch whorls often whitish. One cord (occasionally two) lighter than background, usually white-whitish, above suture, at approximately 1/3 to 1/2 of whorl height, the area above the cord usually slightly darker than the rest. Darker blotches and areas irregularly placed on axial ribs and interspaces on upper part of each whorl, often also below periphery and on siphonal fasciole and on subsutural ramps. Spiral cords occasionally of same colour as background, more often entirely or dashed brown or reddish-brown. Aperture internally white or beige, sometimes with light brownish blotches and lines 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 narrower than basal. Operculum absent. Body translucent white with some parts yellowish or orange. Scattered white speckles on siphon, tentacles and upper side of foot. Posterior part of foot yellowish; tentacles translucent yellowish or pale orange; siphon pale yellowish in small specimens, orange in large ones ( Fig. 19 ).

REMARKS

The foregut anatomy of a specimen from Malta, Qummick, 80 m depth (vi.1988, C. Mifsud leg; 15.5 x 6.1 mm, protoconch tip missing, 1.5 protoconch whorls remaining, diagonally cancellate) was examined and described by Don Tippett (in litt., 2001). The specimen lacked a proboscis, poison apparatus, radula and salivary glands. The rhynchodaeum was very retracted and folded, with the posterior end bearing a tiny circular opening into the oesophagus.

This species is everywhere less common than the other 3 species of the genus. The first specimen of this species was submitted to our attention by Alberto Villari who identified it as Raphitoma volutella Kiener, 1839 . With the same name, additional specimens were present in some other private collections. Kiener (1839: 67) described P. volutella as recent from Sicily, yet it is known that sometimes he reported the fossil species as living (Palazzi, unpublished ms.). In fact, Pleurotoma volutella is relatively well known from the Plio-Pleistocene ( Seguenza 1880; Cerulli-Irelli 1910; Harmer 1915). Aradas & Benoit (1876: 251) reported the finding of a fresh specimen of Pleurotoma volutella from Acitrezza. Unfortunately, a search in the Aradas collection at the Museo di Storia Naturale di Milano (merged into that of Priolo), was unsuccessful. This specimen must be considered lost as subsequently confirmed by Leonardi (MSNM, pers. com.). The type material of P. volutella has also not been found, despite a meticulous search in the Delessert collection studied by Kiener himself (Y. Finet, MHNG, pers. com.). The comparison with fossil specimens from Monte Pellegrino (Plio-Pleistocene) and Ficarazzi (Low Pliestocene) from the Monterosato collection and that of Melvill-Tomlin (NMW, 12930, given by Monterosato) showed that L. villaria and P. volutella are different species. This last specimen ( Fig. 21B ) is labelled by Monterosato “ Leufroyia volutella Kiener , fossile di Ficarazzi, non vivente”. The second handwritten label reads: “Kiener a cru vivant un specimen ramassé par la mer”. However, the specimens identified by Monterosato as “ volutella ” are quite different from each other and do not seem to be conspecific. The specimen ( Fig. 21E ) from Ficarazzi is the most similar to Kiener’s drawings of P. volutella . According to Cipolla (1914: 73), Pleurotoma volutella falls within the chronological variation of Pleurotoma inflata De Cristofori & Jan, 1832 whereas for Cerulli-Irelli (1910: 63 (255)) the two entities are separated. We are inclined to share the opinion of Cerulli-Irelli and believe that P. inflata and P. volutella are two different species (both known only as Plio-Pleistocene fossils, Fig. 21 ), and that both of them are different from L. villaria . L. villaria is distinguished from L. inflata by having:

1 – different outline (with less arched spire);

2 – axial ribs weaker (less high and wide), opistocline vs orthocline or slightly prosocline;

3 – much thinner and less dense spiral cords, and alternat- ing cords and cordlets;

4 – interspaces between cords quite large vs very narrow;

5 – different opening shape (more oval vs more oblong);

6 – very short siphonal canal vs long one;

L. villaria is distinguished from L. volutella by having:

1 – more rounded outline;

2 – greater number of axial ribs which are flatter;

3 – more prominent, more numerous and less spaced spi- ral cords;

4 – different opening shape (more oval vs piriformis elon- gated);

5 – less wide and shorter siphonal canal.

The height and maximum diameter of protoconch depend on the number of whorls, in the photographed specimen ( Fig. 20 ) with PW = 3.04 these two features are: H = 735 µm and max. D = 593 µm. Diameter of nucleus and first whorl are quite large and similar to L. erronea , while in L. concinna and L. leufroyi these features are significantly smaller.

Maximum height of examined shells is 19 mm, those larger than 20 mm are very rare. Mifsud (2003, as Raphitoma erronea ) reported the maximum size of 25 mm.

The original description was based on light and uniformly coloured specimens, but at least in the Adriatic and Aegean Sea they are usually brightly coloured, with brown or reddishbrown spiral cords and brown areas and blotches. Peñas et al. (2006: 126, figs 274-276 as Pleurotomella sp.) mentioned the presence of some brown cords also on the shells examined by them from Alboran Sea.

After a few years from collection the shells always look significantly different from the fresh ones, since the darker colours fade out and the shells become much lighter in colour, so even the shells with brown spiral cords often look almost uniformly coloured.

Diagnostic features of L. villaria from all other living Leufroyia spp. are: somewhat thinner and generally more slender shell, very thin spiral cords and narrow axial ribs, narrower subsutural ramps and more stepped whorls, medially the largest protoconch, differently coloured tentacles and foot of the animal.

Its identification is usually relatively easy, especially of large specimens, due to their slender shell and very thin spiral cords. Smaller specimens have less slender shell and can be confused with some untypical small shells of other species.

It differs from L. concinna , which also have brown spiral cords, by absence of pinkish/purple areas on the teleoconch, by thinner spiral cords and much larger protoconch, also by much bigger maximum size of shell.

L. leufroyi sometimes has similar colour pattern of shell ( Fig. 5H ) to some colour forms of L. villaria , but even in such cases they can easily be differentiated by several features: small and dark vs large and light protoconch, wide and less spaced vs very thin and well spaced spiral cords; none or only one brown cord on each whorl vs often almost all.

L. erronea has much lower H/W ratio; wider aperture; shorter siphonal canal; much stronger thickening of outher lip; stronger, denser and never brown coloured spiral cords.

Figures

FIG. 17. — Leufroyia villaria (Pusateri & Giannuzzi-Savelli, 2008): A, holotype (MNHN-IM-2000-27244), Taormina (Italy) -15 m, h. 17.3 mm; B, paratype C (GOR), Capraia Island (Italy) 150 m, h. 16.9 mm; C, paratype A (MNHN-IM-2000-23353), Ras il-Wahx (Malta), 80-100 m, h. 11.5 mm; D, East Sicily (Italy), h. 12.4 mm; E, Porto Cesareo (Taranto - Italy),h. 14.2 mm; F, Fomm Ir-Riħ (Malta) 140 m,h. 10.76 mm;G, Ionian coasts of Calabria (Italy),h. 13.7 mm;H, Gorée (Senegal) (MNHN, Knudsen coll.),h. 11.4 mm;I, Mljet Island (Croatia),70-100 m, amidst red coral colonies,h.10.62 mm;J, Mljet Island (Croatia),70-100 m, amidst red coral colonies, h. 8.58 mm; K, Saronikos Gulf (Greece), h. 17.1 mm; L, Saronikos Gulf (Greece), h. 16.6 mm. Photo courtesy: D, G, Roberto Ardovini; K, L, Costas Kontadakis.

FIG. 18. — Leufroyia villaria (Pusateri & Giannuzzi-Savelli, 2008): A, Mljet Island (Croatia) -70-100 m, amidst red coral colonies, h. 12.22 mm; B, Saronic Gulf (Greece), h. 16.1 mm; C, Saronic Gulf (Greece), h. 18.3 mm; D, South Eastern Sicily (Italy), h. 13.1 mm; E, Mljet Island (Croatia) 70-100 m, amidst red coral colonies, h. 11.5 mm; F, Lastovo Island (Croatia) 200 m, h. 16.7 mm; G, Mljet Island (Croatia) 70-100 m, amidst red coral colonies, h. 8.1 mm. Photo courtesy: B, C, Costas Kontadakis; D, Roberto Ardovini; F, Pero Ugarković.

FIG. 19. — Photographs of two living animals of Leufroyia villaria (Pusateri & Giannuzzi-Savelli,2008). Dugi Otok Island (Croatia), 35-57 m, h. 7 mm (A), h. 7.5 mm (B-E). Photo courtesy: Alen Petani.

FIG. 20. — Leufroyia villaria (Pusateri & Giannuzzi-Savelli, 2008). Dugi Otok Island (Croatia), 35-57 m, h. 7.18 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.

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.