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Hycleus hayki Serri & Bologna n. sp. Figs 6, 18, 63–69 Hycleus pirata, Serri, 2015: 235, nec Bologna & Turco, 2007. Type material. Holotype ♂ (HMIM) " [Iran] Dogonbadan / 15. V.68/ [leg.] Zairi, Ayat ". 4 ♂ 2 ♀ Paratypes (HMIM) with the same label of holotype; 3 ♂ 1 ♀ paratypes (HMIM) "Dogonbadan/ 11. V.68 [leg.] Ayat., Zairi"; 3 ♂ 2♀ paratypes (HMIM) 1 ♂ (MABC); "Dogonbadan/ 9. V.67/ [leg.] Zairi, Ayat'; 1♂ paratype (HMIM): Behbahan/ 1. V.69/[leg.] Zairi, Ayat"; 1 ♂ paratype (HMIM): Behbahan/ 3. VI.69/ [leg.] Zairi, Ayat". Type locality: " Iran, Dogonbadan " (hoc opus). Records from literature. Iran. Khuzestan, Kuhgiloyeh & Boyerahmad (Serri, 2015 as pirata; Ghahari & Campos-Soldini 2019 as pirata). Description. Body black, elytra testaceous with black pattern; setation black, short, sparser shorter and obliquely recumbent on elytra. Golden dense pubescence on ventral side of fore tibiae and tarsi of male, only on tibiae of female. Body length 8–14 mm. Head subquadrate (Fig. 18); with maximum width at level of eye; punctures relatively wide, deep and irregular, surface among punctures relatively shiny; eye globose, with antero-dorsal margin slightly sinuate, just behind antennal insertion. Temples parallel, only slightly curved posteriad and shorter than longitudinal diameter of eye; frons almost flat; clypeus transverse, convex, with slightly rounded anterior and lateral margins, fronto-clypeal suture clearly visible; labrum only slightly shorter than clypeus, anterior margin slightly sinuate, weakly depressed in middle; mandibles robust, curved, longer than clypeus and labrum together; maxillary palpomeres subcylindrical, IV flattened; antennae (male Fig. 63, female Fig. 64) not extending to posterior margin of pronotum, antennomere III more than 1/3 time longer than IV, IV–VII progressively longer and widened, VIII as long as IX, IX slightly narrower than VIII, X narrower than IX, IX widest before middle and narrowed to apex, with a transverse depressed line right in middle of curved part; in female last antennomere short. Pronotum (Fig. 18) convex, slightly longer than wide, bulging on sides and slightly narrowing anteriad, fore portion without depression, posterior margin slightly depressed just in front of mesonotum; with one middle longitudinal furrow, punctation deep, interstices shiny, mesosternun as in Fig. 68. Elytral pattern as in Fig. 69, each elytron with two black spots on anterior third, middle fascia with sinuate posterior margin, and a wide black fascia on apical part with straight fore margin. Last male ventrite convex, posterior margin with a deep middle emargination; aedeagus (Fig. 67) with endophallic hook slightly narrow, distal aedeagal hook scarcely curved, far from proximal one and almost at apex; gonoforceps (Figs 65–66) in ventral view regularly narrowed, in lateral view sinuate in last third but not curved. Taxonomic remarks. This new species is very similar to H. sexmaculatus and in some ways to H. pirata, but can be distinguished by the shape of last antennomere of male which has a transverse depressed line in the middle of the curved part, by the shape of the pronotum which is slightly longer than wide, by the elytral pattern which middle fascia is wide and has always sinuate hind margin, and by the shape of male genitalia. We failed to find fresh material for this species for DNA extraction, but we hope future molecular studies may reveal the phylogenetic relationships among these species. Distribution. Hycleus hayki sp. n. is hitherto known from southwestern Iran. Etymology. The species is named after Hayk Mirzayans, orthopterist and founder of Hayk Mirzayans Insect Museum (HMIM), the largest insect collection of Iran. He was the first to draw up the blister beetles of the HMIM coleoptera collection and to publish a contribution to the knowledge of Meloidae of Iran.

Hycleus pirata Bologna & Turco, 2007 Figs 8, 20, 77–83 Hycleus pirata Bologna & Turco, 2007: 16. Type material. Holotype ♂ and 2 ♀ paratypes:" United Arab Emirates: Abu Dhabi Emirate, EA4, Al Bateen dunes, 4 Km W Al Ain on the road to Abu Dhabi, 195 m a.s.l. / 24, 20491° N 55, 61263° E, 15.III.2005, leg. M. Bologna & F. Turco (MABC)"; 4 ♂ 4 ♀ paratypes (MABC), 1 ♂ paratype (HMIM): "United Arab Emirate, Abu Dhabi Emirate, EA10, Al Aïn, 3–4 Km E/ Al Selimat on road Al Aïn-Abu Dhabi, 24, 19263° N 55, 58861° E, 220 m, 12–14.III.2005, leg. M. Bologna & F. Turco ". Type locality: " United Arab Emirates: Abu Dhabi Emirate, EA4, Al Bateen dunes, 4 Km W Al Ain on the road to Abu Dhabi " (Bologna & Turco 2007). Additional material examined. 1 ♀ Oman, Wadi Air, 7. V.1957, leg. Popov & Wood (MABC). Records from literature. Oman. Oman (Koçak & Kemal 2015). UAE. UAE (Gillet & Gillet, 2005 as bipunctata; Koçak & Kemal 2015; Ghahari & Campos-Soldini 2019); Abu Dhabi Emirate, EA4, Al Bateen dunes, 4 Km W Al Aïn on the road to Abu Dhabi, 24, 20491° N 55, 61263° E (Bologna & Turco 2007, type locality); Abu Dhabi Emirate, EA10, Al Aïn, 3–4 Km E Al Selimat on road Al Aïn-Abu Dhabi, 24, 19263° N 55, 58861° E (Bologna & Turco 2007); Jebel Hatif (Gillet & Howarth 2004 as bipunctata); Al Aïn area (Gillet & Gillet 1996 as bipunctata); Wadi Bih; Wadi Shawkah (Batelka & Geistardt 2009 as bipunctatus). Description. Body length 12–16.5 mm. Body black, elytra yellowish bright brown, sometimes red, with black pattern. Head subquadrate; male antennomere III (Fig. 77) less than half longer than IV, V–VIII transverse and widened, IX–X narrower than VIII, male XI narrower and longer than X; in female (Fig. 78) shorter than X. Pronotum (Fig. 20) longer than wide, with anterior depression; mesosternum as in Fig. 82. Each elytron with two black spots on anterior third, one sinuate middle fascia, and one black oblique fascia on apical part (Fig. 83). Male last ventrite regularly concave and depressed, posterior margin with a wide and deep middle emargination; aedeagus (Fig. 81) with endophallic hook narrow, distal aedeagal hook curved, far from proximal one and close to apex, proximal hook slightly curved; gonoforceps (Figs 79–80) narrow and elongate both in ventral and lateral view (see Bologna & Turco 2007 for a detailed description). Taxonomic remarks. This species resembles H. apicipennis but it can be distinguished by the elongate shape of antennomere IV, which is subequal in length to V, the shape of last male antennomere, which is strikingly curved, and by the male genitalia. The shape of last antennomere is also very similar to that of H. hayki and of H. sexmaculatus, but H. pirata differs because of the pronotum and aedeagus shape. Distribution. This species seems to have restricted distribution in the United Arab Emirates and Oman (Bologna & Turco 2007). The record of this species from south Iran (Serri 2015) refers to the new species H. hayki., Published as part of Serri, Sayeh, Bologna, Marco A. & Riccieri, Alessandra, 2020, Revision of the Hycleus sexmaculatus species group (Coleoptera: Meloidae Mylabrini), pp. 1-42 in Zootaxa 4790 (1) on pages 23-24, DOI: 10.11646/zootaxa.4790.1.1, http://zenodo.org/record/3887775, {"references":["Bologna, M. A. & Turco, F. (2007) The Meloidae (Coleoptera) of the United Arab Emirates with an updated Arabian checklist. Zootaxa, 1625 (1), 1 - 33. https: // doi. org / 10.11646 / zootaxa. 1625.1.1","Kocak, A. O. & Kemal, M. (2015) Initial list of the Pterygota species of Malatya Prov., based upon the info-system of the Cesa (East Turkey). Cesa News, 115, 1 - 150.","Gillet, M. P. T. & Gillet, C. (2005) Insects and other arthropods. In: Hellyer, P. & Aspinall, S. (Eds.), The Emirates. A Natural History. Trident Press Limited, Abu Dhabi, pp. 169 - 196.","Ghahari, H. & Campos-Soldini, M. P. (2019) An annotated catalogue of blister-beetles (Coleoptera: Tenebrionoidea: Meloidae) of Iran. Entomofauna, 40, 59 - 138.","Gillet, M. P. T. & Howarth, B. (2004) Insects. In: Aspinall, S. & Hellyer, P. (Eds.), Jebel Hafit, a natural history. Emirates Natural History Group, Abu Dhabi, pp. 94 - 143.","Gillet, M. P. T. & Gillet, J. (1996) A large blister beetle Mylabris maculiventris Klug, 1845 from Al Ain and neighbouring Oman (Coleoptera: Meloidae). Tribulus, 6, 20 - 21.","Serri, S. (2015) Blister beetles (Coleoptera: Meloidae) of Southwest Iran. Proceedings of the 1 st Iranian International Congress of Entomology, 2015, 232 - 237."]}

Hycleus is the most speciose genus of the beetle family Meloidae, with almost 430 species widespread in Africa and Eurasia. It belongs to the tribe Mylabrini, subfamily Meloinae. The systematics of the genus Hycleus has never been investigated but, according to the mesosternal morphology, it has been divided by Pardo Alcaide into three main “sections”: Mesogorbatus, with almost 70 species; Mesotaeniatus, with about 50 species and Mesoscutatus with almost 300 species. Within these “sections”, different lineages have been identified according to other morphological characters. Furthermore, previous and ongoing studies on the phylogeny of this family pointed out that the genera Paractenodia and Ceroctis are strictly related to the genus Hycleus and should probably be included within it. Paractenodia includes five species endemic to South Africa and Namibia, and Ceroctis 59 species distributed through the tropical Africa and the Saharan-Arabic deserts. In order to understand the relationships among some of the species belonging to these genera, about one hundred species were molecularly tested with mitochondrial (COI, 16S) and nuclear (CAD, ARGK, Wingless) markers. The sequences obtained were analysed under the assumptions of Maximum parsimony, Maximum likelihood and Bayesian inferences and a first set of phylogenetic trees is proposed. Preliminary results seem to corroborate the validity of some lineages already defined according to morphological characters, moreover both nuclear and mitochondrial markers confirm that both Ceroctis and Paractenodia belong to the genus Hycleus, in agreement with previous preliminary molecular results. Further analyses to increment the number of tested species are already in progress.

Nemognathinae is the most widespread subfamily of Meloidae, with ~600 species, and includes the only blister beetles distributed in Australia and on islands of the western Pacific. Four tribes are recognised based on morphology: Stenoderini, Palaestrini, Horiini and Nemognathini. Using two mitochondrial (16S, COI) and three nuclear markers (CAD, 28S, ITS2), and both maximum likelihood and Bayesian approaches, this study describes the evolutionary history of Nemognathinae based on molecular data for the first time. We provided a fossil-calibrated phylogeny that unravels the phylogenetic relationships among the tribes and among most of the genera, and a reconstruction of the biogeographic history using a parametric approach. Our results recognised the four tribes that were described previously based on morphology and revealed the presence of another well-differentiated clade corresponding to the genus Zoltanzonitis. Phylogenetic relationships among the tribes are well supported, with Stenoderini as the most ancient lineage, followed by Zoltanzonitini, Palaestrini, Horiini and Nemognathini. A few long-standing genera within Nemognathini (Nemognatha, Zonitis, Stenoria) and the nominate subgenus Stenodera (Stenodera) were recovered as polyphyletic. In addition, biogeographic analyses revealed the origin of the subfamily in the Old World during the Eocene, and the associated diversification into the five tribes astride the Eocene and Oligocene between 46 and 30 Ma. Based on these results we propose the new tribe Zoltanzonitini, and the elevation of the subgenus Pronemognatha to genus level, new status. In addition, Zonitoschema breveapicalis new comb., Z. curticeps new comb. and Z. pulchella new status are proposed. ZooBank: urn:lsid:zoobank.org:pub:72EECC6D-36A6-4DD7-B4DB-D0692034E775.

Sex is an important factor affecting bacterial diversity in insects (Xu et al. 2016, J. Insect Phy, 95: 110–117). Divergence in some life-history traits between males and females could provide differential opportunities for bacterial colonization (Tang et al. 2012, FEMS Microb Ecol 80: 659–670). Blister beetles include almost 3000 species (Riccieri et al. 2022, Syst Ent 1-12) and are characterized by sexual dimorphism in body content of the toxic terpene cantharidin (CA). CA is produced and exuded by blister beetles in yellowish oily hemolymph droplets from leg and antennal joints as a defensive strategy (Gisondi et al. 2019, Entomol Sci 22: 258-263). Although the latest steps of CA production are still elusive (Fratini et al. 2021, BMC Genomics 22: 808), males are known to produce more CA than females and accumulate large amounts of this terpene in their reproductive organs (Carrel et al. 1993, Experientia 49: 171–174). Females are especially attracted by males with high amount of CA, which is transferred to females upon mating for protecting eggs from predators or parasites (Sierra et al. 1976, Experientia 32: 142-144). Through 16S metabarcoding we compared the bacterial composition between males and females in five species of blister beetles (Hycleus polymorphus, Mylabris variabilis, Lydus trimaculatus, Zonitis flava, Meloe proscarabeus). Gammaproteobacteria, Bacilli, Actinobacteria and Alphaproteobacteria were the most prevalent classes of bacteria in all species. Overall, we observed a sexual dimorphism in the microbial component in all examined species, with males harboring lower values of microbial alpha-diversity, as in other insects (Wan et al. 2020, Animals 10: 2352). Plausible roles of sex-specific microbes are hypothesized and, as in other terpene-producing beetles (i.e., Paederus fuscipennis, Song et al. 2022, Insect Mol Biol 1-14), the potential involvement of bacteria in the de novo biogenesis of CA in blister beetles is discussed.

Protease inhibitors are widely studied since the unrestricted activity of proteases can cause extensive organ lesions. In particular, elastase activity is involved in the pathophysiology of acute lung injury, for example during SARS-CoV-2 infection, while serine proteases and thrombin-like proteases are involved in the development and/or pathology of the nervous system. Natural protease inhibitors have the advantage to be reversible and with few side effects and thus are increasingly considered as new drugs. Kunitz-type protease inhibitors (KTPIs), reported in the venom of various organisms, such as wasps, spiders, scorpions, and snakes, have been studied for their potent anticoagulant activity and widespread protease inhibitor activity. Putative KTPI anticoagulants have been identified in transcriptomic resources obtained for two blister beetle species, Lydus trimaculatus and Mylabris variabilis. The KTPIs of L. trimaculatus and M. variabilis were characterized by combined transcriptomic and bioinformatics methodologies. The full-length mRNA sequences were divided on the base of the sequence of the active sites of the putative proteins. In silico protein structure analyses of each group of translational products show the biochemical features of the active sites and the potential protease targets. Validation of these genes is the first step for considering these molecules as new drugs for use in medicine.

In this study, the blister beetles belonging to the Hycleus sexmaculatus species group are revised. Morphological characters are described and figured for each species and keys to both male and female are provided. Hycleus hayki new species from Southwest Iran is described and the following new synonymies are proposed: Mylabris triangulifera ab. reitterioides Mader 1929 = Hycleus bipunctatus (Olivier, 1811) new synonymy ; Mylabris triangulifera ab. bushirensis Mader 1929 = Hycleus bipunctatus (Olivier, 1811) new synonymy ; Mylabris javeti Marseul, 1870 = Hycleus colligatus (Redtenbacher, 1850) new synonymy ; Mylabris javeti umbilicatus Kaszab, 1958 = Hycleus colligatus (Redtenbacher, 1850) new synonymy ; Hycleus amrishi Makhan, 2012 = Hycleus cingulatus (Faldermann, 1837) new synonymy . Additional remarks which rectify previous errors in the identification of the species are provided. The taxonomic positions of Zonabris soumacovi Pic, 1930 and Zonabris sialanus Pic, 1929 are also discussed. Phylogenetic relationships of the species, based on a subset of available species inferred from molecular data are proposed, and some morphologically defined subgroups of species are distinguished with the molecular support. The geographic and ecological distributions of all the species are defined, and a detailed list of localities is provided. The ecological information including phenology, elevation and host plants of the species are summarized in a table. Based on literature records, collections and recently collected data, the distribution of the sexmaculatus species and their biogeographic characteristics are discussed.

Many continental species distributed in the Eurasian steppe occur as relict populations in the mountains of Western Europe. Their biogeographical responses to Quaternary climate changes have been poorly studied; however, they could have responded as cold-adapted species. We investigated the biogeographic history of a steppe beetle, Hycleus polymorphus, using mitochondrial and nuclear DNA sequences (COI, CAD, ITS2), and species distribution modelling (SDM) under present and past bioclimatic envelopes. We first performed a phylogenetic assessment to define species boundaries within the H. polymorphus species group. Specimens previously treated as Hycleus humerosus on morphological grounds are assigned to H. polymorphus, and those identified as Hycleus zebraeus assigned to Hycleus atratus. ITS2 data analyses revealed a strong phylogeographical structure of H. polymorphus populations, with four haplogroups corresponding to the (i) Italian Alps, (ii) French Alps and Pyrenees, (iii) South Balkan and Pontic mountains, and (iv) North Dinaric Alps. Based on these analyses and the SDM, we propose that during a glacial period, following the spread of steppic habitat, H. polymorphus underwent a range expansion from Asia to South-West Europe. Within the Mediterranean area, during the last interglacial the climatic suitability for the species was limited to mountains that acted as refugia and prompted allopatric divergence into four main lineages.

The genus Hycleus is a hyper-diverse genus of blister beetles including ~500 species with a wide geographic distribution in different biogeographic regions in the Old World, and with the highest diversity occurring in the Afrotropical Region. The phylogenetic relationships among the species and the biogeographic processes related to their diversification have never been investigated. In this study, to provide a biogeographic hypothesis that allow explaining the current diversity and the observed distribution patterns of Hycleus, we built a robust time-calibrated phylogenetic tree using mitochondrial and nuclear DNA obtained from 125 species. Phylogenetic results were subsequently used for biogeographic inference carried out with the R package BioGeoBEARS. Four main lineages were detected in our tree: A. including only Afrotropical species; B. including only Saharo-Sindian species; C. comprising Afrotropical, Palaearctic and Saharo-Sindian species; and D. comprising Afrotropical and Oriental species. According to our results, the genus Hycleus likely originated in the Afrotropical Region during the Early Miocene (~20 Mya), and subsequently spread in the Saharo-Sindian Transitional Region and in the Palaearctic Region. In the Late Miocene (~7.5 Mya) a second Saharo-Sindian group branched off from the Palaearctic lineage, whereas the Oriental Region was colonized (~7.8 Mya) following a dispersal event through the Arabian Peninsula from the Afrotropical Region.

Three non-phoretic blister-beetle species (Mylabris schreibersi, Actenodia distincta and Cabalia segetum) were investigated to trace their Siculo-Maghrebian biogeographic origins and to understand how Tertiary and Quaternary paleogeographic events (e.g. the ‘Messinian Salinity Crisis’ and the cyclic Quaternary glacio-eustatic marine oscillations) shaped their phylogeographic patterns. Data from CAD and RpP01 nuclear genes highlighted a clear lineage sorting between Sicilian and Maghrebian populations of M. schreibersi, explainable by a vicariance event likely occurred after the ‘Messinian Salinity Crisis’. A less marked – but still noticeable – genetic distinction among Sicilian and Maghrebian populations of A. distincta and C. segetum was observed, which could be more referable to moderately old (during Pliocene or late Quaternary) dispersal events. In addition, the low genetic variability and lack of population structure found in Sicily for the three blister-beetle species suggest some drastic reduction in size of Sicilian populations during Pleistocene glacial phases. In general, our results on Meloidae beetles are consistent with published literature, indicating that vicariance or dispersal scenarios can be both plausible explanations of the Siculo-Maghrebian distribution currently observed in a number of animal and plant species.

Several animal and plant species show a West Mediterranean distribution restricted to Maghreb and Sicily. According to geological evidences, these lands became separated after the opening of the straits of Gibraltar at the beginning of Pliocene (5.33 million years ago). During Pleistocene glaciations, apparently, no land connections occurred between Sicily and Tunisian coasts, even if these lands were greatly approached. Therefore, the presence of species with a fragmented distribution at both shores of the Sicilian channel stimulated two zoogeographic hypotheses: (a) a pre-Pliocene origin of these species, whose range was subsequently fragmented by a vicariance event at the beginning of Pliocene (which implies a quite old species age); (b) Pleistocene dispersal events, during glacial stages, across the Sicilian channel. In order to test these hypotheses, we performed molecular analysis in two blister beetle species (subfamily Meloinae) having a Sicilian-Maghrebian distribution, but belonging to distinct phylogenetic lineages (BOLOGNA, 1991; BOLOGNA and PINTO, 2002): Mylabris schreibersi Reiche, 1866 (tribe Mylabrini), referred to a Palaearctic, widely distributed and speciose genus, and Cabalia segetum (Fabricius, 1792) (tribe Lyttini) belonging to a small Afrotropical genus distributed in eastern Africa, SW Arabian peninsula and Sahara, with the single C. segetum exclusive of Maghreb and Sicily. We sequenced one mitochondrial (COI) and three nuclear (CAD, Wingless, RpS9b) gene fragments in 9 Sicilian and 6-8 Maghrebian (from Morocco and Tunisia) populations of both species. Preliminary results (mainly based on CAD, the most informative gene) show that both species represent cohesive genetic units with a shallow divergence between Maghrebian and Sicilian populations supporting a Pleistocene dispersal scenario. North-African populations show higher genetic variability than Sicilian populations, suggesting an older history of these species in the Maghreb. Additional data and statistical analyses will allow to better understand the phylogeography of M. schreibersi and C. segetum, to estimate timing of population divergence, as well as to infer colonization routes between Maghreb and Sicily.