Research Article |
Corresponding author: Alberto Sánchez-Vialas ( alberto.alytes@gmail.com ) Academic editor: Aaron Smith
© 2022 Alberto Sánchez-Vialas, José L. Ruiz, Ernesto Recuero, Felipe Gutiérrez-Pérez, Mario García-París.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Sánchez-Vialas A, Ruiz JL, Recuero E, Gutiérrez-Pérez F, García-París M (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: 17-48. https://doi.org/10.3897/zookeys.1109.83863
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The taxonomic status and subgeneric arrangement of the genus Eurymeloe have been debated for decades. In this work, the internal taxonomy of Eurymeloe is redefined by recognising three subgenera: Eurymeloe for the former Eurymeloe brevicollis species group, Coelomeloe for Eurymeloe tuccia, and Bolognaia Ruiz, García-París, Sánchez-Vialas & Recuero, subgen. nov., to accommodate the species of the formerly recognised Eurymeloe rugosus species group. Additionally, a new species of the newly described subgenus Bolognaia is described from the Iberian Peninsula based on molecular and morphological traits. The new species, Eurymeloe (Bolognaia) orobates sp. nov., can be distinguished from all other species of Eurymeloe by the following combination of morphological traits: dispersed brownish setae over the body that are arranged in small tufts on the abdominal terga; a small, very transverse pronotum that presents a unique macrosculpture; a deeply and densely punctured integument of the head and pronotum; and the very rugose elytra. The characters displayed by E. orobates suggest that the species groups that were previously defined and recognised for Eurymeloe, and that are now integrated within the newly erected subgenus Bolognaia, are non-monophyletic.
Bolognaia subgen. nov., Coelomeloe, Iberian Peninsula, new species, phylogeny, systematics
The taxonomy of Eurymeloe Reitter, 1911, originally described as a subgenus of Meloe Linnaeus, 1758, has been controversial due to both its differential use at the genus (
The Eurymeloe rugosus species group was comprehensively revised by
Recently,
We recently collected specimens representing a new distinctive species of the E. rugosus species group from central Spain. The new species, which can be easily diagnosed by both conspicuous morphological characters and molecular data, is characterised by a combination of morphological traits present in either one or the other of
In this study, we (1) describe the new species of Eurymeloe and analyse its phylogenetic relationships by using the Meloini molecular framework of
We studied the external morphology of a total of 326 specimens belonging to 17 species of Eurymeloe. All specimens are listed in either
The description of the new species of Eurymeloe is based on a total of five specimens (one male, dried preserved, and four females, ethanol-preserved), all belonging to the type series. These specimens were collected from the mountains of central Spain, at Puerto de la Quesera (Province of Guadalajara, Autonomous Community of Castilla-La Mancha). The type series is held at
For the morphological study, dry-mounted specimens were examined under a stereomicroscope. The male specimen was rehydrated in water before the extraction of the genital structures, which was subsequently mounted on a piece of cardboard using dimethylhydantoin formaldehyde (DMHF) resin and pinned adjacent to the specimen. Measurements were taken using a micrometre that was coupled to one of the microscope eyepieces. Digital images of live, dry-mounted specimens, and of male and female genital structures, were taken with a reflex camera (Canon 77D) fitted with a macro-lens (Sigma 105 mm F2.8) and two external flashes. We used the terminology suggested by
For the molecular analyses, we used sequences of Eurymeloe available from GenBank and those of four new specimens that we had collected and preserved in ethanol (now housed at the
Specimen identity, collection locality, voucher number, and GenBank accession numbers of the new and previously published sequences analysed in this work.
Taxon | Locality | Voucher number | GenBank # CoxI | GenBank # 16S | GenBank # Wg | GenBank # 18S |
---|---|---|---|---|---|---|
Eurymeloe apivorus | Morocco: Fès-Meknès: Ifrane, 2 km South of Cedro Gouran, Middle Atlas | mel 81009 | MW158218 | MW158046 | MW157964 | MW158119 |
Eurymeloe apivorus | Morocco: Fès-Meknès: Ifrane, 2 km South of Cedro Gouran, Middle Atlas | mel 81013 | MW158220 | MW158048 | MW157966 | MW158121 |
Eurymeloe apivorus | Morocco: Fès-Meknès: Ifrane, 2 km South of Cedro Gouran, Middle Atlas | mel 81054 | MW158219 | MW158047 | MW157965 | MW158120 |
Eurymeloe brevicollis | Spain: Cantabria: Brañavieja, Pico Tres Mares | mel 04107 | MW158305 | MW158088 | MW157987 | MW158142 |
Eurymeloe brevicollis | Andorra: Arinsal | mel 07092 | MW158306 | MW158089 | MW157988 | MW158143 |
Eurymeloe corvinus | Japan: Niigata, Sado, Kitaushima | 9060 | LC583106.1 | |||
Eurymeloe corvinus | Japan: Saga, Karatsu, Hirose | 11881 | LC583105.1 | |||
Eurymeloe corvinus | Japan: Nagano, Iriyamabe | 10521 | LC583104.1 | |||
Eurymeloe fernandezi | Spain: Islas Canarias: La Palma, Los Sauces | mel 07045 | MW158266 | MW158068 | MW157972 | MW158127 |
Eurymeloe fernandezi | Spain: Islas Canarias: La Palma, Los Sauces | mel 07048 | MW158267 | MW158069 | MW157973 | MW158128 |
Eurymeloe ganglbaueri | Italy: Lazio: Tarquinia | mel 81064 | MW158268 | MW158070 | MW157974 | MW158129 |
Eurymeloe ganglbaueri | Italy: Lazio: Tarquinia | mel 81065 | MW158269 | MW158071 | MW157975 | MW158130 |
Eurymeloe ganglbaueri | Italy: Sardinia: 4 km North-West of Orgosolo | ASV19011 | OM918705 | OM925566 | ||
Eurymeloe glazunovi | Romania: Dobruja: Istria | mel 07001 | MW158265 | MW158067 | MW157971 | MW158126 |
Eurymeloe gomari | Morocco: 9 km South-West of Moulay Abdeselam | ASV18019 | OM936883 | OM918704 | OM925565 | |
Eurymeloe gomari | Morocco: Tangier-Tetouan: Chaouen, Talassemtane National Park | mel 81063 | MW158275 | MW158076 | MW157981 | MW158136 |
Eurymeloe ibericus | Spain: Ávila: Villanueva del Campillo, Puerto de Villatoro | mel 06039 | MW158307 | MW158090 | MW157989 | MW158144 |
Eurymeloe ibericus | Spain: Ávila: Hoyos del Espino, Plataforma de Gredos | mel 81039 | MW158308 | MW158091 | MW157990 | MW158145 |
Eurymeloe mediterraneus | Spain: Cádiz: Puerto Real | mel 04255 | MW158221 | MW158049 | MW157967 | MW158122 |
Eurymeloe mediterraneus | Morocco: Moulay Abdelsalam | mel 07010 | MW158222 | MW158050 | MW157968 | MW158123 |
Eurymeloe mediterraneus | Spain: Cuenca: Saelices | mel 07147 | MW158224 | MW158052 | MW157970 | MW158125 |
Eurymeloe mediterraneus | Morocco: Tetouan: Agnan, Sierra del Haus | mel 81066 | MW158223 | MW158051 | MW157969 | MW158124 |
Eurymeloe murinus | Morocco: Marrakesh-Safi: 2 km North of Aguelmouse, Tizi n’Tichka, High Atlas | mel 81018 | MW158270 | MW158072 | MW157976 | MW158131 |
Eurymeloe murinus | Spain: Madrid: Colmenar Viejo | mel 81053 | MW158271 | MW157977 | MW158132 | |
Eurymeloe nanus | Spain: Madrid: 7 km South of Tielmes | mel 01028 | MW158273 | MW158074 | MW157979 | MW158134 |
Eurymeloe nanus | Spain: Madrid: Tielmes | mel 81042 | MW158274 | MW158075 | MW157980 | MW158135 |
Eurymeloe nanus | Spain: Toledo: Villacañas, Sierra del Romeral | mel 05001 | MW158272 | MW158073 | MW157978 | MW158133 |
Eurymeloe orobates sp. nov. | Spain: Guadalajara: Puerto de la Quesera | ASV18002 | OM936884 | OM925567 | ||
Eurymeloe orobates sp. nov. | Spain: Guadalajara: Puerto de la Quesera | ASV18003 | OM936885 | OM925568 | ||
Eurymeloe rugosus | Germany: Saxony-Anhalt | ZFMK-TIS-2003300 | KU918912.1 | |||
Eurymeloe tuccia | Spain: Menorca: 2 km South of Binimella | mel 06034 | MW158276 | MW158077 | MW157982 | MW158137 |
Eurymeloe tuccia | Spain: Islas canarias: La Palma: Don Pedro | mel 07058 | MW158277 | MW158078 | MW157983 | MW158138 |
Eurymeloe tuccia | Spain: Almería: La Mela | mel 81001 | MW158278 | MW158079 | MW157984 | MW158139 |
Eurymeloe tuccia | Portugal: Algarve: Sagres, Praia do Martinhal | mel 81002 | MW158279 | MW158080 | MW157985 | MW158140 |
Eurymeloe tuccia | Morocco: Larache: Lixus | mel 81006 | MW158280 | MW158081 | MW157986 | MW158141 |
Lampromeloe aff. variegatus | Morocco: Marrakesh-Safi: 5.5 km North-East of Aguelmouse, Tizi n’Tichka, High Atlas | mel 81010 | MW158202 | MW158033 | MW157953 | MW158108 |
Lampromeloe cavensis | Morocco: Casablanca-Settat: Ouled Bahmad | mel 06011 | MW158201 | MW158032 | MW157952 | MW158107 |
Lampromeloe variegatus | Spain: Salamanca: 5 km West of Palencia de Negrilla | mel 05015 | MW158203 | MW158034 | MW157954 | MW158109 |
Lampromeloe variegatus | Hungary: Komárom-Esztergom: Vèrtesszölös | mel 81068 | MW158204 | MW158035 | MW157955 | MW158110 |
Meloe (Anchomeloe) autumnalis | Spain: Guadalajara: Villanueva de la Torre | mel 04246 | MW158189 | MW158025 | MW157949 | MW158104 |
Meloe (Anchomeloe) autumnalis | Spain: Zaragoza: El Frago | mel 10070a | MW158191 | MW158027 | MW157951 | MW158106 |
Meloe (Anchomeloe) autumnalis | Morocco: Tangier-Tetouan: Chaouen, Djebel Tissouka | mel 81071 | MW158190 | MW158026 | MW157950 | MW158105 |
Meloe (Meloe) proscarabaeus | Spain: Menorca: 3.5 km South-West of Fornells | mel 06026 | MW158148 | MW157993 | MW157939 | MW158094 |
Meloe (Meloe) proscarabaeus | Morocco: Souss-Massa: 2 km South of Chafarni | mel 81007 | MW158150 | MW157995 | MW157941 | MW158096 |
Meloe (Meloe) proscarabaeus | Hungary: Tolna: Bátaapáti | mel 06004 | MW158151 | MW157996 | MW157942 | MW158097 |
Meloe (Meloe) proscarabaeus | Italy: Tuscany: Alberese | mel 81082 | MW158149 | MW157994 | MW157940 | MW158095 |
Meloe (Meloe) tropicus | Guatemala: El Quiché: 9 km North-East of Santa Cruz Quiché | mel 81075 | MW158188 | MW158024 | MW157948 | MW158103 |
Meloe (Meloe) cf. violaceus | Hungary: Tolna: Mőcsény | mel 07033 | MW158186 | MW158022 | MW157946 | MW158101 |
Meloe (Meloe) cf. violaceus | Hungary: Vas: Csákánydoroszló | mel 07036 | MW158187 | MW158023 | MW157947 | MW158102 |
Meloe (Meloe) violaceus | Spain: Ávila: Hoyos del Espino | mel 05024 | MW158183 | MW158019 | MW157943 | MW158098 |
Meloe (Meloe) violaceus | Spain: León: Correcillas: Pico Polvareda | mel 81051 | MW158185 | MW158021 | MW157945 | MW158100 |
Meloe (Meloe) violaceus | Spain: León: Correcillas: Pico Polvareda | mel 81052 | MW158184 | MW158020 | MW157944 | MW158099 |
Meloegonius cicatricosus | Hungary: Pest: Tatárszentzgyörgy | mel 06002 | MW158208 | MW158038 | MW157959 | MW158114 |
Meloegonius cicatricosus | Hungary: Komárom-Esztergom: Vèrtesszölös | mel 81069 | MW158209 | MW158039 | MW157960 | MW158115 |
Mesomeloe coelatus | Morocco: Guelmine-Smara: Reg Labyad | Mcoelatus_labyad | MW805179 | |||
Mesomeloe coelatus | Morocco: Guelmine-Smara: Jbel Ouarkziz | Mcoelatus_ouarkziz | MW805180 | |||
Mesomeloe ottomerkli | Qatar: 1.8 km West of Al Marrawnah | mel Qatar1 | HG003653 | MW158044 | MW157962 | MW158117 |
Mesomeloe ottomerkli | Qatar: 1.8 km West of Al Marrawnah | mel Qatar2 | HG003654 | MW158045 | MW157963 | MW158118 |
Physomeloe corallifer | Spain: Madrid: 5 km South-East of Agustín de Guadalix, 618 m asl | mel 09051 | MW158210 | MW158040 | MW157961 | MW158116 |
Taphromeloe erythrocnemus | Morocco: 2.5 km North-West of Douar Azerzou | ASV18007 | MW158309 | MW158092 | MW157991 | MW158146 |
Treiodous gracilicornis | Guatemala: El Quiché: 3.4 km North of Uspantán | mel 81077 | MW158205 | MW158036 | MW157956 | MW158111 |
Treiodous gracilicornis | Mexico: Guerrero: 5 km West of Carrizal de Bravo | mel 874 | MW158206 | MW157957 | MW158112 | |
Treiodous laevis | Mexico: Guanajuato: Dolores Hidalgo | mel 08094 | MW158207 | MW158037 | MW157958 | MW158113 |
Lytta vessicatoria | Spain: Ourense: A Acea | mel 05073 | MW158147 | MW157992 | MW158093 | |
Phodaga alticeps | Mexico: Baja California Norte: Ejido Luchadores del Desierto, Northwest of Laguna Salada | KRN14 | MK024506 | MK024642 | MK024601 | |
Cordylospasta fulleri | USA: California: 4.8 km North-East of Big Pine | KRN23 | MK024478 | MK024619 | MK024572 |
The molecular data set consisted of two mitochondrial (COI and 16S rRNA) and two nuclear (Wg and 18S rRNA) gene fragments from 66 specimens (including the four new ones). All sequences were compiled and revised using Sequencer v. 4.9 and aligned with MAFFT (
To delimit species, we adopted the evolutionary species concept (
Our Bayesian phylogenetic tree, which has a topology similar to the one presented by
Bayesian phylogeny of the genus Eurymeloe based on the concatenated matrix of mitochondrial and nuclear genes (COI, 16S, Wg, 18S). 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.
With respect to the internal taxonomic structure of Eurymeloe (sensu
Bolognaia is comprised of three main lineages: one represented by E. fernandezi, an endemic of the Canary Islands, which resolved as the sister taxon to the other two lineages formed, respectively, by E. mediterraneus (Müller, 1925) and its closely related species and by E. murinus (Brandt and Erichson 1832) and its related species, including the new species of Eurymeloe from central Spain. The sequences of the new samples from Spain resolved as a distinctive lineage that is the sister taxon to the clade formed by E. murinus and E. ganglbaueri. This new lineage is not morphologically congruent with any other described species of Eurymeloe (Fig.
Habitus of some living adult specimens of the current subgenera of Eurymeloe A subgenus Eurymeloe, represented by a specimen of Eurymeloe (Eurymeloe) ibericus from Puebla de la Sierra, northern Madrid, Spain B subgenus Coelomeloe, represented by a specimen of Eurymeloe (Coelomeloe) tuccia found 3 km east of Celín, Sierra de Gádor, Almería, Spain C–I subgenus Bolognaia C Eurymeloe (Bolognaia) affinis from the surroundings of Bab Taza, Rif Mountains, Morocco D E. (Bolognaia) mediterraneus (type species of the subgenus 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 Alcázar 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 González de Rueda).
Lastly, our molecular phylogenetic analysis confirmed the species identification of the other two newly sequenced specimens as E. gomari and E. ganglbaueri, and their relationship with the other specimens. As a result, the northern limit of the distribution range of the Moroccan endemic E. gomari, which was previously known only from its type locality, has been expanded by approximately 37 km to the northwest along the Rif Mountains. The new specimen of E. ganglbaueri from Sardinia resolved, as expected, within the same clade as the conspecific Italian continental samples.
Given the new systematic arrangement proposed here for Eurymeloe as three subgenera (Eurymeloe, Coelomeloe, and Bolognaia subgen. nov.), we consider it necessary to present a revised diagnosis of Eurymeloe based on the morphology of adult specimens.
Meloe brevicollis Panzer, 1793, by subsequent designation of
(adult). Size small to medium (6–36 mm), with diverse appearance, ranging from very robust to comparatively slender. Body integument colour variable, black, dull grey or dark brown (exceptionally sandy brown) to moderately metallic blue, opaque, bright, silky or sometimes with an oily shininess (Fig.
Larva. The morphological traits of previously known first instar larvae of Eurymeloe (triungulines), including Coelomeloe, and the descriptions of the triungulines of several additional species of Eurymeloe, have been synthesised and studied by
The adult instar of species of the genus Eurymeloe is morphologically diverse, and the three subgenera can be recognised based on this diversity. Adults of the subgenera Eurymeloe, Coelomeloe, and the newly described Bolognaia are distinguishable particularly by the shape of the antennae and the pronotum, the macrosculpture of the pronotum, body integument and pilosity, and the punctation of the head, pronotum, and elytra, among other traits.
Meloe mediterraneus Müller, 1925, by present designation.
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.
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.
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 (
Based on molecular data (this work) and adult morphology (see
According to our molecular analyses (Fig.
For practical purposes, but also supported by our analyses and some morphological traits (mainly, pilosity colour, integument aspect, pronotum punctation, and elytra rugosity; see
(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 (
(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
Meloe brevicollis Panzer, 1793 (by subsequent designation of Pinto and Selander, 1970).
Size small or medium (6–30 mm), usually robust in appearance. Body integument colour black to moderately metallic blue, bright, silky, or with an oily shininess (Fig.
According to the present definition of the subgenus Eurymeloe, it is correlated with the E. brevicollis species group defined by
On the basis of the molecular and morphological data (
Another 13 species [from Palaearctic Asia, except E. aleuticus (Borchmann, 1942), from the Aleutian Islands] were provisionally assigned by
Regarding other species of Eurymeloe,
1 | Body entirely black and opaque. Body pubescence absent dorsally. Pronotum flat, subrectangular, transverse, depressed in middle of the base, with sides straight, parallel. Punctation of the head and pronotum very broad, dense, subcontiguous (less dense in Sicilian and southern Italian populations) and deep, clearly foveate in appearance (Fig. |
Coelomeloe |
– | Body black, dull grey or dark brown (exceptionally sandy brown) to moderately metallic blue, bright, silky or more seldom opaque in appearance, sometimes with an oily shininess. Body pubescence quite distinct, or very short, recumbent, often almost imperceptible. Pronotum slightly to moderately convex, wider than long, with sides not parallel, more or less converging backward, and posterior angles usually broadly rounded. Head and pronotum punctation from fine and scattered, sometimes almost absent, to somewhat deep and dense, but never foveolate (Fig. |
2 |
2 | Body colour black to moderately metallic blue, bright or silky. Overall appearance robust. Body pubescence very short, recumbent, almost imperceptible or even absent on the head and pronotum. Antennae compact, robust, sometimes smoothly thickened towards the apex, short or medium in length, not reaching the posterior margin of the pronotum. Antennomeres subcylindrical or subconical, V to VII (in some species IV to IX) wider than long or, at most, as wide as long | Eurymeloe |
– | Body integument black, dull grey or dark brown, exceptionally sandy brown, with an opaque, silky or bright appearance, never bluish or metallic. Body pubescence quite distinct. Overall appearance more graceful, sometimes moderately robust. Antennae normally slender, not thickened towards the apex, long or medium in length, usually reaching the posterior margin of the pronotum or exceeding it. Antennomeres IV to IX subcylindrical, always longer than wide | Bolognaia |
Our molecular results revealed a distinctive lineage of Eurymeloe nested within the clade comprising E. rugosus, E. murinus, and E. ganglbaueri. This lineage, morphologically distinguishable from all its congeneric species, represents a new species that we herein describe.
adult male (Fig.
four adult females, labelled: two females: “Puerto de la Quesera, Guadalajara, Spain 41°11'30.11"N, 3°24'27.55"W, 1625 m, 22-V-2016, M. García París, A. Fernández Liger, A. Corral Lou leg. [white label, printed]; ASV 18002 and ASV 18003, respectively [white label, handwritten]; MNCN_Ent 325407 and MNCN_Ent 325408, respectively [white label, printed]. One adult female (Fig.
Total length (frons to apex of the tergite VIII): 11.05 mm. Length from the frons to the posterior margin of elytra: 6.55 mm. Maximum width (located slightly anterior to the apex of the elytra): 6.81 mm. Body relatively robust, with slender appendages (Fig.
Head voluminous, broadly rounded and clearly wider than the pronotum, weakly truncated on the posterior margin of the temples, with integument black, silky in appearance, finely microreticulated, and without longitudinal depressions behind the eyes (Fig.
Antennae with 11 antennomeres, moniliform, slender and long, surpassing the base of the pronotum when extended backward (Fig.
Pronotum black, silky in appearance (Fig.
Elytra relatively short (length: 4.05 mm), strongly dehiscent and weakly convex, imbricated basally (the right over the left), divergent backwards and reaching the middle area of the fourth tergite, covering the first tergite, almost completely covering the second, and covering the lateral areas of the third (of which, only the central plate is clearly visible), and lateral basal portions on both sides of the fourth; elytral surface strongly rugose, corrugated, with marked wavy foveoles (Fig.
Abdomen black, voluminous (Fig.
Legs relatively slender (Fig.
Male genitalia with gonoforceps dark brown, hairless, moderately elongated, slender in dorsal, ventral, and lateral views (Fig.
Female similar to the male (Fig.
Eurymeloe (B.) orobates is characterised morphologically, with respect to all the other species of the Bolognaia, by the following combination of diagnostic traits: (1) body size small or medium (total length: 10–14 mm); (2) body integument entirely black, semi-glossy in appearance; (3) setation of the head, pronotum and elytra, short and decumbent, reddish brown, moderately dark, sometimes very dark (almost black) ventrally and on the legs; (4) setation of the central plates of the abdominal tergites yellowish red (some almost golden), longer and forming inconspicuous tufts; (5) antennae slender and long, surpassing the base of the pronotum when extended backwards; (6) head broadly rounded, with a weak and relatively short longitudinal median groove; (7) pronotum small, very transverse (more than 1.5 × wider than long), sub-hexagonal; (8) pronotal surface showing a weak and narrow depressed longitudinal-middle area, but without a marked groove; (9) punctation of the head and pronotum dense, forming rounded and markedly deep punctures; (10) elytral surface strongly rugose, corrugated, with marked foveoles; and (11) male genitalia with long gonostyli with no excavated or depressed areas in the distal regions and a narrow and long gonocoxal plate.
The species most similar to E. orobates are E. rugosus and E. apenninicus (both belonging to the group B or E. murinus group). Both species present dark (black or dark brown) pilosity all over the body and, on the abdominal tergites, some inconspicuous (usually barely perceptible) yellowish brown or yellow setae, but not tufts. In addition to the colour pattern of the body setation, E. rugosus and E. apenninicus differ from E. orobates by the shape of their pronotum, which is longer, less transverse, and flatter (less convex), and has a strong median longitudinal groove (absent in E. orobates). The punctation of the head and pronotum are also markedly larger, deeper, and denser in E. rugosus and E. apenninicus (see
Within group B (E. murinus group), in which E. (B.) orobates is integrated, the new species can be readily distinguished from E. (B.) baudueri (southern France, Iberian Peninsula, and northern Morocco), E. (B.) flavicomus (Canary Islands), E. (B.) ganglbaueri (mainland Italy, Sardinia, Corsica, Sicily, Greece, Albania, Bulgaria, Bosnia and Herzegovina, Montenegro, Turkey, Syria, Spain, and southern France), E. (B.) gomari (northern Morocco), E. (B.) kandaharicus (Iran and Afghanistan), E. (B.) murinus (Iberian Peninsula, Sicily, Sardinia, Corse, Crete, Maghreb, and Libya), E. (B.) nanus (Iberian Peninsula, North Africa, and Middle East), E. (B.) omanicus (eastern Arabian peninsula), and E. (B.) pallidicolor (western Morocco). For instance, in contrast to E. (B.) orobates, all these species present, among other specific traits, a body integument that is dull grey or dark brown, occasionally reddish brown or, rarely, sandy brown (E. pallidicolor) or almost black (E. ganglbaueri). In addition, the body integument is generally opaque or matte in appearance or, at most, silky (but never glossy or semi-glossy as in E. orobates). The setation of these species is also quite distinct from that of E. orobates: it is yellowish, whitish, or golden all over the body and usually longer, and on the abdominal tergites, the tufts of setae, when present, are highly conspicuous (see Fig.
Eurymeloe (B.) orobates differs from the species of group A (E. mediterraneus group, composed of, at least, E. affinis from the Maghreb and Libya; E. apivorus and E. baamarani, which are restricted to Morocco; E. baudii from the Italian Peninsula, Sicily, and Croatia; E. glazunovi from Eastern Europe and Central Asia; and E. mediterraneus, which is widely distributed throughout Europe, the Mediterranean basin, the Canary Islands, and the Middle East) by presenting reddish brown body setation and abdominal tergites with small tufts of reddish yellow setae, among other diagnostic characters (see above). By contrast, the body pilosity, including on the abdominal tergites, of species of the E. mediterraneus group is black (see
The only species in group C is E. (B.) fernandezi (endemic to the Canary Islands). In comparison with E. (B.) orobates, this species presents, among other distinctive characters, an entirely black body setation; a clearly longer, not transverse pronotum with sinuous margins; an integument surface with wrinkles and parallel ridges that form eddies; and an elytral sculpture consisting of a fine zig-zag roughness (see
Eurymeloe orobates is only known from a single locality, Puerto de la Quesera (in the province of Guadalajara, Spain) in the Iberian Peninsula (Fig.
The specific epithet orobates is derived from the Greek word “oros”, meaning mountain, and “bates”, meaning walker. This name alludes to the mountainous environment where the specimens of the new species were found, sometimes, wandering on mountain pastures and trails (Fig.
In light of previous morphological data and recent phylogenetic analyses (
Morphological traits of larvae have been traditionally considered relevant in the systematics of the group, sometimes even more informative than adult characters for phylogenetic studies (
Some conspicuous adult traits are also shared between the subgenera Eurymeloe and Coelomeloe, including antennae that are submoniliform, robust, short or medium in length, and which do not usually not reach the posterior margin of the pronotum; antennomeres V to VII that are wider than long or, at most, as wide as long; and very short or not [e.g., E. (E.) brevicollis, E. (E.) ibericus, and E. (C.) tuccia] body pubescence. These character states differ from those of the Bolognaia, which usually presents antennae that are moniliform, normally slender, long or medium in length, and which usually reach or exceed the posterior margin of the pronotum; antennomeres IV-IX that are subcylindrical, always longer than wide; and distinctive short or very short (black, yellowish, whitish, or golden) body pubescence. Therefore, the close relationship between Eurymeloe s. str. and Coelomeloe is supported by both our molecular analysis (BPP = 0.9) and morphology.
Our results confirm that E. (B.) rugosus, a species previously assigned to
With the addition of the new species, eight species of the Bolognaia are known from the Iberian Peninsula: E. (B.) baudueri, E. (B.) ganglbaueri, E. (B.) mediterraneus, E. (B.) murinus, E. (B.) nanus, E. (B.) orobates, E. (B.) rugosus, and E. (B.) saharensis (
We are very grateful for help during the field work to E. Karen López-Estrada, Concha Cano, Andrea Corral Lou, Andrea Fernández Liger, Javier Aznar González de Rueda and Arlo Hinckley. We thank Melinda Modrell for the thorough language revision. We are also grateful to Mercedes París and Antonio Melic for providing us with the opportunity to revise the MNCN entomological collection and the Maynar Collection of the Asociación Entomológica Aragonesa, respectively. We thank Antonio Recalde for allowing us to examine an Iberian specimen of E. rugosus. This work was funded by the Spanish Government, project grant PID2019-110243GB-100 /AEI/10.13039/501100011033 (Ministerio de Ciencia, Innovación y Universidades, Spain) (PI: M. García-París).