Scabricola, SWAINSON, 1840

Fedosov, Alexander, Puillandre, Nicolas, Herrmann, Manfred, Kantor, Yuri, Oliverio, Marco, Dgebuadze, Polina, Modica, Maria Vittoria & Bouchet, Philippe, 2018, The collapse of Mitra: molecular systematics and morphology of the Mitridae (Gastropoda: Neogastropoda), Zoological Journal of the Linnean Society 183, pp. 253-337 : 310

GENUS SCABRICOLA SWAINSON, 1840

( FIGS 28D–F, 29A–F)

Type species: Mitra serpentina Lamarck, 1811 (= Voluta variegata Gmelin, 1791 ); SD, Gray (1847: 141).

Diagnosis: see under ‘Remarks’.

Distribution: Indo-Pacific, upper subtidal depths, on sand.

Species included: Scabricola bicolor (Swainson, 1824) 3, S. desetangsii (Kiener, 1838) 1, S. olivaeformis (Swainson, 1821) 1, S. padangensis (Thiele, 1925) 2, S. variegata (Gmelin, 1791) 1,? Scabricola barrywilsoni (J. Cate, 1968) 3,? S. albina A. Adams , 18533,? S. caerulea (Reeve, 1844) 3,? S. condei Guillot de Suduiraut , 2 0 0 1 3, S. cor i acea (Reeve, 1 8 4 5) 2,? S. b a ck ae (Cernohorsky, 1973) 3,? S. dampierensis Salisbury & Heinicke , 19983,? S. dianneae (Salisbury & Guillot de Suduiraut, 2003) 3 comb. nov.,? S. eximia (A. Adams, 1853) 3,? S. geigeri Poppe, Tagaro & Salisbury , 20093,? S. gilbertsoni (Cate, 1968) 3 comb. nov.,? S. guttata (Swainson, 1824) 3 comb. nov.,? S. hayashii (Kira, 1959) 3,? S. incisa (A. Adams & Reeve, 1850) 3,? S. ivanmarrowi Marrow , 20163,? S. lavoisieri Guillot de Suduiraut , 20023,? S. lorenzi ( Poppe & Tagaro, 2006) 3 comb. nov.,? S. mariae (A. Adams, 1853) 3,? S. martini Poppe & Tagaro, 2006 3,? S. petiti Poppe & Tagaro , 20063,? S. parkinsoni Salisbury & Wolff, 2005 3,? S. potensis (Montrouzier, 1858) 3,? S. sowerbyi d’Orbigny , 18523,? S. splendidula Salisbury & Guillot de Suduiraut , 20033,? S. vicdani Cernohorsky , 19813,? S. yaekoae (Habe & Kosuge, 1966) 3.

Remarks: The genus Scabricola as commonly understood is extremely heterogeneous. It combines such dissimilar forms as S. yaekoae , which in shell outline is close to Mitra mitra , and S.potensis (Montrouzier, 1858) , with its ovate shell resembling species of Pterygia . The range of sculpture patterns is equally considerable: whereas most species of Scabricola are characterized by wide and low cords or punctate grooves, some (e.g. S. padangensis ) have a ‘ Subcancilla -type’ sculpture pattern ( Cernohorsky, 1991) of narrow, high cords, and still others, like S. coriacea , show a distinctly crenulated sculpture. In the treatment of Cernohorsky, the more slender species of Scabricola , with weaker sculpture, were separated in the subgenus Scabricola (Swainsonia) . It should be noted, though, that the boundaries of the two subgenera appear blurred, primarily because S. variegata (the type species of Scabricola ) is conchologically closer to Swainsonia than to many species of Scabricola (Scabricola) . One of the important diagnostic characters of Scabricola is the morphology of the radula lateral teeth, unique for the Mitridae , ‘with 3–5 strong cusps on the inward facing angular plate’ ( Cernohorsky, 1991: 115) ( Fig. 28D – S . variegata). This radular morphology is shared by seven species of Scabricola (Scabricola) and Scabricola (Swainsonia) studied by Cernohorsky (1970), whereas radulae with divergent morphologies are found in S. desetangsii ( Fig. 28E) and S. coriacea ( Cernohorsky, 1970: fig. 120): the former has unicuspidate laterals, the latter lacks the major cusp on laterals which, therefore, have a somewhat intermediate morphology between typical Scabricola and Neocancilla .

Because the boundaries of the genus remain poorly understood, we are only confident about the placement of those species for which molecular and radular morphology data are available, and the remaining species are placed in Scabricola with a question mark. Given this major uncertainty on its contents, we refrain from giving a formal diagnosis of the genus.