Drilus, OLIVIER, 1790: 3

GENUS DRILUS OLIVIER, 1790: 3

( FIG. 9E, K, U)

Type species: Drilus flavescens (Geoffroy, 1785) ; by monotypy.

Species included: Altogether 45 described and several undescribed species: Drilus adustus Chevrolat, 1854 , D.akbesianus (Fairmaire, 1895) , D. amabilis Schaufuss, 1867 , D. attenuatus Pic, 1914 , D. badius Kobieluszova & Kundrata, 2015 , D. baenai Kundrata et al., 2015 , D. bleusei ( Olivier, 1913) , D. bicolor Schaufuss, 1867 , D. concolor Ahrens, 1812 , D. creticus Pic, 1905 , D.distincticollis Pic, 1907 , D.flavescens (Geoffroy, 1785) , D. frontalis Schaufuss, 1867 , D. fulvicollis Audouin, 1824 , D. fulvicornis Kiesenwetter, 1859 , D. fulvitarsis Baudi di Selve, 1872 , D. funebris Reitter, 1884 , D. horasfakionus Kundrata et al. 2015 , D. huijbregtsi Kobieluszova & Kundrata, 2015 , D.humeralis Pic, 1931 , D. iljini Barovskij, 1922 , D. iranicus Wittmer, 1967 , D. latithorax Pic, 1902 , D. longulus Kiesenwetter, 1859 , D. mauritanicus Lucas, 1842 , D. mertliki Kobieluszova & Kundrata, 2015 , D. nemethi Kundrata et al., 2014 , D. novoathonius Sumakow, 1903 , D. obscuricornis Pic, 1899 , D. posticus Schaufuss, 1867 , D. rectus Schaufuss, 1867 , D. rittneri Petrzelkova & Kundrata, 2015 , D. robustus Kobieluszova & Kundrata, 2015 , D. rufipes (Baudi di Selve, 1872) , D. sanliurfensis Kobieluszova & Kundrata, 2015 , D. schwarzi Reitter, 1891 , D. subparallelus Pic, 1934 , D. testaceipes Pic, 1933 , D. teunisseni Kobieluszova & Kundrata, 2015 , D. turcicus Kobieluszova & Kundrata, 2015 . The following five species described from the Afrotropical Region have different morphology from the here defined Drilus and will be revised in the near future: D. atripennis Pic, 1934 , D. basilewskyi Wittmer, 1962 , D. impressiceps Pic, 1913 , D. ramosus Fairmaire, 1883 , D. testaceipennis Pic, 1918 .

Diagnosis: Drilus is recognized by the following combination of characters: dorsal body surface uneven, usually matt; eyes relatively small to medium-sized, their frontal separation 1.60–3.00 times eye diameter; antenna slightly serrate to weakly flabellate; pronotum transverse, 1.20–1.70 times as wide as long, with lateral sides sinuate or rounded, lateral carina almost complete, and posterior margin usually only shallowly arcuately emarginate; prosternum ( Fig. 9E) with reduced prosternal process; mesoventrite ( Fig. 9K) V-shaped, with only indistinctly defined or shallow mesoventral cavity; surface of elytra usually without distinct striae or lines of punctures; abdominal ventrites free ( Fig. 9U), sternite IX and tergite X not apparently elongate, with more or less rounded sides, usually not more than 1.80 times as long as wide; paramere without latero-apical projection.

Relationships: The phylogenetic relationships among the Drilus lineages are only partly resolved based on the six-marker phylogeny ( Fig. 1; Supporting Information, Fig. S1). The presence of the monophyletic group informally known as the ‘Greek clade’, which is formed by the elongate, usually tiny species with the serrate antennae that are specialized predators of Albinaria snails and are distributed in the Peloponnese, the Ionian islands and Crete, is consistent with the results of the cox1 analysis by Kundrata et al. (2015a). Its statistically strongly supported sister group relationship with D. rufipes (Baudi di Selve) from Cyprus agrees with the results of the three-marker analysis by Sormova et al. (2018). The remaining species, which include usually more robust and hairy representatives with weakly flabellate antennae and a strongly transverse pronotum, formed a gradual branching in previous studies ( Kundrata et al., 2015a; Sormova et al., 2018), but they are found to be monophyletic in this study, although with low support ( Fig. 1; Supporting Information, Fig. S1). This clade included two branches, i.e. D. fulvicornis + ( D. flavescens + D. mauritanicus ) and D. concolor + ( D. rectus + D. cf. mertliki ). More species in the analysis in combination with a detailed morphological investigation of this genus are needed for a better understanding of Drilus phylogenetic interrelationships.

Distribution: Northern Africa ( Algeria, Morocco), Europe, Asia Minor, Levant, Caucasus, Iran. Species from tropical Africa do not belong to the genus (see comment above).

GENUS MALACODRILUS KUNDRATA & BOCAK ,