Bolognaia Ruiz, Garcia-Paris , Sanchez-Vialas & Recuero, subgen. nov.

Sanchez-Vialas, Alberto, Ruiz, Jose L., Recuero, Ernesto, Gutierrez-Perez, Felipe & Garcia-Paris, Mario, 2022, A new systematic arrangement for the blister beetle genus Eurymeloe (Meloini, Meloidae, Coleoptera) with the description of a new species from Spain, ZooKeys 1109, pp. 17-48 : 17

Bolognaia Ruiz, Garcia-Paris, Sanchez-Vialas & Recuero, subgen. nov.

Type species.

Meloe mediterraneus Müller, 1925, by present designation.

Description (adult).

Size small or medium to large (8-36 mm). Body integument black, dull grey or dark brown, occasionally sandy brown [ E. pallidicolor ( Martínez de la Escalera, 1909)], with an opaque, silky or bright appearance, never bluish or metallic (Fig. 2C-J ). Body pubescence quite distinct, black, yellowish, whitish or golden, short or very short. Head rounded, temples usually forming a regular arc, except in E. murinus (Brandt and Erichson 1832) and E. affinis (Lucas, 1847), which have strongly enlarged temples; occiput usually weakly concave. Medium-sized eyes [smaller in E. affinis and E. apivorus (Reitter, 1895)], subreniform, moderately protruding, without a longitudinal depression behind them. Antennae moniliform, normally slender, not thickened towards the apex; long or medium in length, usually reaching the posterior margin of the pronotum or exceeding it; unmodified in males, straight. Antennomeres IV-IX subcylindrical, always longer than wide. Clypeus transverse, approximately twice as wide as long. Labrum wide, fore margin broadly emarginate. Mandibles robust, regularly curved along the outer margins. Pronotum slightly convex, transverse, mainly subhexagonal or subtrapezoidal, wider than long, usually 1.4-2.1 × as wide as long [in E. fernandezi , 1.2-1.3 × as wide as long], sides not parallel, converging backward, posterior margin broadly emarginated, posterior corners rounded. Pronotum surface variable, with or without a depressed area or groove in the middle, frequently with two depressed or smooth areas, diffuse, on both sides of the disc. Head and pronotum punctation fine to coarse, of variable density, always with pubescence. Posterior margin of mesonotum straight or weakly arcuate. Mesepisterna usually not meeting at the midline of the body. Elytra short and dehiscent, smooth to strongly rough, usually rugulose. Legs normal, usually slender, pubescent. Tarsomeres without hair pads on the inferior side, though some species [e.g., E. nanus (Lucas, 1847), E. baudueri (Grenier, 1863), E. gomari ] have fairly dense setose pubescence that appears as small and short brushes. Last abdominal ventrite broadly and deeply emarginated at the hind margin in males. Male genitalia: Gonostyli usually elongate, distal regions narrow with their apices acuminate or digitiform in lateral view; gonocoxal plate long, usually narrow and slightly widened at the middle in dorsal view; aedeagus usually elongate, equally long as or longer than gonoforceps.

Etymology.

The name Bolognaia , formed by the noun “Bologna” plus the Italian suffix “-aia” derived from the Latin “-aria” (used, in this case, to form a word meaning an animal associated with the specified noun Bologna), is in honour of Marco A. Bologna, a distinguished Italian entomologist specialising in the Meloidae and a friend who, among other excellent works, was able to clarify, for the first time, the complex taxonomy of the small-sized species of Eurymeloe related to E. rugosus for which the new subgenus is here erected.

Taxonomic remarks.

We selected M. mediterraneus as the type species of Bolognaia because it is a morphologically well-characterised species, with low morphological or genetic intraspecific geographic differentiation ( Bologna 1988, 1991; Sánchez-Vialas et al. 2021), and without nomenclatural or identity problems associated to synonyms ( García-París et al. 2010; Bologna 2020a). Bolognaia , a monophyletic subgenus, largely corresponds to the E. rugosus species group of Eurymeloe defined by Bologna (1988, 1991). It includes species whose adults are characterised mainly by the following traits: small or medium body size; black, dull grey, or dark brown body colour; a distinctive black or pale-coloured (yellowish, whitish, or golden) pilosity; elongated and subcylindrical antennomeres that are longer than wide; and generally marked punctation and rugosity.

Based on molecular data (this work) and adult morphology (see Reitter 1895; Martínez de la Escalera 1909; Pliginskij 1910; Pardo Alcaide 1951; Kaszab 1958, 1983; Bologna 1988, 1991, 1994a, 1994b; Ruiz and García-París 2009, 2015; García-París and Ruiz 2011; Di Giulio et al. 2013), we include within the subgenus Eurymeloe Bolognaia the following species: Eurymeloe (Bolognaia) affinis (Lucas, 1847), E. (B.) apivorus (Reitter, 1895), E. (B.) apenninicus (Bologna, 1988), E. (B.) baamarani ( Ruiz and García-París 2015), E. (B.) baudii (Leoni, 1907), E. (B.) baudueri (Grenier, 1863), E. (B.) fernandezi , E. (B.) flavicomus (Wollaston, 1854), E. (B.) ganglbaueri , E. (B.) glazunovi (Pliginskij, 1910), E. (B.) gomari , E. (B.) kandaharicus (Kaszab, 1958), E. (B.) mediterraneus , E. (B.) murinus , E. (B.) nanus , E. (B.) omanicus (Kaszab, 1983), E. (B.) pallidicolor , and E. (B.) rugosus .

According to our molecular analyses (Fig. 1 ), three sublineages can be recognised within Bolognaia : two generally corresponding to Bologna’s (1988) subgroups A and B, defined by presenting, respectively, an entirely black body pilosity (in our analyses, E. mediterraneus , E. apivorus , and E. glazunovi ) or a pale-coloured (whitish, yellowish, reddish yellow, or golden) pilosity over the entire body or parts of it [in our analyses, E. ganglbaueri , E. murinus , E. nanus , E. gomari , E. rugosus , and Eurymeloe orobates sp. nov. from central Spain]. Notably, based on our molecular analyses, E. rugosus , which was included in Bologna’s (1988) subgroup A, appears to be genetically more related to the species included in his subgroup B. In this regard, following a detailed examination of some specimens belonging to E. rugosus , we observed that several have inconspicuous brownish and reddish to yellowish setae (but not tufts) on their abdominal tergites, similar to those observed on the morphologically related species E. apenninicus (JLR, pers. obs.). In fact, Escherich (1890) and Bologna (1988, 1991) pointed out that some specimens of E. rugosus show yellowish brown setae on the last abdominal tergites; these correspond to the named var. Eurymeloe rugosus abdominalis ( Escherich 1890) (which has even been confused with E. ganglbaueri , see Bologna 1988: 247). Likewise, E. ganglbaueri , which presents a golden-yellow pilosity on a part of the body, resolved as genetically more related to species in subgroup B. The third sublineage diverged from its sister group, the A and B sublineages, during the Middle Miocene ( Sánchez-Vialas et al. 2021). This sublineage is composed of only one species, E. fernandezi , an endemic of the Canary Islands that is morphologically singular and isolated within the subgenus ( Pardo Alcaide 1951; Bologna 1988, 1991, 1994a; Ruiz and García-París 2015).

For practical purposes, but also supported by our analyses and some morphological traits (mainly, pilosity colour, integument aspect, pronotum punctation, and elytra rugosity; see Bologna 1988; Ruiz and García-París 2009, 2015), we redefine the specific composition of the subgroups established by Bologna (1988) within Bolognaia (defined as the E. rugosus species group by Bologna 1988) as follows:

(1) group A or E. mediterraneus group (now renamed), characterised mainly by a dark body pilosity (black or dark brown) and a black body integument that is usually glossy, semi-glossy, or silky in appearance [exceptionally, it is matte as in E. (B.) baamarani ]. This group integrates the following species: E. (B.) affinis , E. (B.) apivorus , E. (B.) baamarani , E. (B.) baudii , E. (B.) glazunovi , and E. (B.) mediterraneus . In some specimens of E. (B.) mediterraneus , particularly those from Sardinia ( Bologna 1988, 1991), the pilosity of the temples, pronotum and, sometimes, abdomen, is brown. The unstudied E. (B.) affinis setosus Escherich, 1890 from Algeria, which differs from the typical form of the species by the presence of isolated yellowish setae along the abdominal tergites, among other traits (e.g., smaller size, constant frontal furrow, and different elytral sculpture) ( Escherich 1890; Bologna 1988, 1991; Di Giulio et al. 2013), possibly constitutes a distinct species, as mentioned by Di Giulio et al. (2013) and previously suggested by Escherich (1890) and Peyerimhoff (in Cros 1934: 90).

(2) group B or E. murinus group, characterised mainly by a pale-coloured (reddish, golden, brownish, yellowish, or whitish) body pilosity, either over the entire body or parts of it, and a body integument that is usually greyish, greyish black, or dark brown and opaque; exceptionally, it is glossy black as in E. (B.) apenninicus and E. (B.) rugosus . This group comprises the following species: E. (B.) apenninicus , E. (B.) baudueri , E. (B.) flavicomus , E. (B.) ganglbaueri , E. (B.) gomari , E. (B.) kandaharicus , E. (B.) murinus , E. (B.) nanus , E. (B.) omanicus , E. (B.) pallidicolor , and E. (B.) rugosus . Within this group, E. (B.) apenninicus and E. (B.) rugosus can be clearly differentiated from the others by having a glossy black body integument and dark reddish brown (sometimes almost black) body setation, with scattered and sparse yellowish brown short setae on the abdominal tergites that are often barely noticeable.

(3) group C, composed by only E. (B.) fernandezi , well characterised morphologically within Bolognaia (see Pardo Alcaide 1951; Ruiz and García-París 2015).

Bologna (1988, 1990) integrated M. saharensis (= M. otini Peyerimhoff, 1949, M. marianii Kaszab, 1983; see Ruiz et al. 2010; Bologna 2020a) and M. vignai in the E. rugosus species group. Ruiz et al. (2010) considered the closely related E. saharensis and E. vignai as morphologically isolated and proposed a new group for them. As neither E. saharensis nor E. vignai have been studied at the molecular level, we have tentatively ascribed them to Bolognaia as Eurymeloe (Bolognaia) saharensis (Chobaut, 1898) and E. (B.) vignai (Bologna, 1990).

Sánchez-Vialas et al. (2021) considered the six Asian species that Bologna (1988) included in the E. rugosus species group as belonging to Eurymeloe . These species are Eurymeloe heptapotamicus (Pliginski, 1910), E. primaeveris (Kaszab, 1958), E. punjabensis (Kaszab, 1958), E. schmidi (Kaszab, 1978), E. scutellatus (Reitter, 1895), and E. subsetosus (Reitter, 1895). However, the available information on these taxa is currently insufficient to assign them to the subgenus Eurymeloe Bolognaia ; therefore, they require further study at both the morphological and the molecular levels.

Figures

Figure 1. Bayesian phylogeny of the genus Eurymeloe based on the concatenated matrix of mitochondrial and nuclear genes (COI, 16 S, Wg, 18 S). The three subgeneric taxonomic categories are indicated as follows: Eurymeloe (in green), Coelomeloe (in violet) and Bolognaia (in blue). Numbers on the nodes represent the posterior probabilities of the clades. Letters in parentheses (A-J) correspond to the species portrayed in Fig. 2. Phylogeny in lower left shows the outgroups (black) and the lineage of Eurymeloe (orange). Additional information for each specimen is provided in Table 1.

Figure 2. Habitus of some living adult specimens of the current subgenera of Eurymeloe A subgenus Eurymeloe Eurymeloe, represented by a specimen of Eurymeloe (Eurymeloe) ibericus from Puebla de la Sierra, northern Madrid, Spain B subgenus Eurymeloe Coelomeloe, represented by a specimen of Eurymeloe (Coelomeloe) tuccia found 3 km east of Celin, Sierra de Gador, Almeria, Spain C-I subgenus Eurymeloe Bolognaia C Eurymeloe (Bolognaia) affinis from the surroundings of Bab Taza, Rif Mountains, Morocco D E. (Bolognaia) mediterraneus (type species of the subgenus Eurymeloe Bolognaia) from Puebla de la Sierra, northern Madrid, Spain E E. (Bolognaia) glazunovi from Istria, Romania F E. (Bolognaia) gomari from the surroundings of Bab Taza, Rif Mountains, Morocco G E. (Bolognaia) nanus from Alcazar de San Juan, Ciudad Real, Spain H E. (Bolognaia) fernandezi from Los Sauces, La Palma, Canary Islands I Eurymeloe orobates sp. nov., from Puerto de la Quesera, Guadalajara, Spain J Eurymeloe (Bolognaia) murinus from Tizi n'Tichka, High Atlas, Morocco. Photographs: A-C, F, G, I (ASV) E, H, J (MGP) D (J. Aznar Gonzalez de Rueda).