A parasitic coevolution since the Miocene revealed by phase-contrast synchrotron X-ray microtomography and the study of natural history collections.

The discovery of a new fossil species of the Caribbeo-Mexican genus Proptomaphaginus (Coleoptera, Leiodidae, Cholevinae) from Dominican amber, associated with a new fossil parasitic fungus in the genus Columnomyces (Ascomycota, Laboulbeniales), triggered an investigation of extant species of Proptomaphaginus and revealed the long-enduring parasitic association between these two genera. This effort resulted in the description of the fossil species †Proptomaphaginus alleni sp. nov., and one fossil and two extant species of Columnomyces, selectively associated with species of Proptomaphaginus: †Columnomyces electri sp. nov. associated with the fossil †Proptomaphaginus alleni in Dominican amber, Columnomyces hispaniolensis sp. nov. with the extant Proptomaphaginus hispaniolensis (endemic of Hispaniola), and Columnomyces peckii sp. nov. with the extant Proptomaphaginus puertoricensis (endemic of Puerto Rico). Based on biogeography, our current understanding is that the Caribbean species of Proptomaphaginus and their parasitic species of Columnomyces have coevolved since the Miocene. This is the first occurrence of such a coevolution between a genus of parasitic fungus and a genus of Coleoptera. The phylogenetic relations among Proptomaphaginus species are also addressed based on a parsimony analysis. Fossil specimens were observed by propagation phase-contrast synchrotron X-ray microtomography (PPC-SRµCT) and extant specimens were obtained through the study of preserved dried, pinned insects, attesting for the importance of (i) technological advancement and (ii) natural history collections in the study of microparasitic relationships.

Sound vs. light: wing-based communication in Carboniferous insects.

Acoustic communication is well-known in insects since the Mesozoic, but earlier evidence of this behavior is rare. Titanoptera, an 'orthopteroid' Permian-Triassic order, is one of the few candidates for Paleozoic intersex calling interactions: some specimens had highly specialized broadened zones on the forewings, which are currently considered-despite inconclusive evidence-as 'resonators' of a stridulatory apparatus. Here we argue that the stridulatory apparatus hypothesis is unlikely because the Titanoptera lack a stridulatory file on their bodies, legs or wings. Instead, comparing these broadened zones with similar structures in extant locusts, flies, and fossil damselflies, we find evidence that the Titanoptera used their wings to produce flashes of light and/or crepitated sounds. Moreover, we describe the first Carboniferous (~310 Mya) Titanoptera, which exhibits such specialized zones, thus corresponding to the oldest record of wing communication in insects. Whether these communication systems were used to attract sexual partners and/or escape predators remain to be demonstrated.

Archaeocerus uenoi n. gen. n. sp. (Coleoptera Leiodidae Catopocerinae) from Albian/Cenomanian age amber of Myanmar.

The first fossil Catopocerinae: Archaeocerus uenoi n. gen. n. sp. (Coleoptera Leiodidae) is described and figured. It comes from the upper Cretaceous (Albian/Cenomanian) amber deposit of the Hukawng valley, in northern Myanmar. Archaeocerus has developed eyes and flight wings, contrast to all the other genera of the subfamily which are anophthalm and apterous.

Phylogeny of Bathysciotina Guéorguiev, 1974, based on morphology with a special emphasis to Italian genera and with the description of a new species of Halbherria (Coleoptera Leiodidae Cholevinae Leptodirini).

The discovery of the new species Halbherria daffneri Perreau, n. sp., a troglobitic species with pholeuonid body shape, in the high altitude karst system of Piani Eterni in southern Dolomites (northern Italy) is taken as an opportunity for a morphological investigation of the phylogeny of Bathysciotina with a special emphasis to the Italian genera. The monophyly of the set of Italian genera, (Aphaotus, Halbherria, Lessiniella, Neobathyscia, Sinuicollia) is confidently supported, with Neobathysica as the sister group of the four other genera. However the monophyly of the subtribe Bathysciotina is not significantly supported and the genus Sphaerobathyscia should clearly be excluded from the subtribe. The subgenus Albanella of the genus Speonesiotes is raised to genus rank.

Ptomaphaginus troglodytes sp. n., the first anophthalmic species of Ptomaphaginina from China (Coleoptera, Leiodidae, Cholevinae, Ptomaphagini).

Ptomaphaginus troglodytessp. n., the first anophthalmic species of Ptomaphaginus Portevin, 1914 is described from two close caves in Libo Karst, south Guizhou Province, China.

A review of the Cholevinae from the island of Borneo (Coleoptera, Leiodidae).

The available knowledge of the round fungus beetle subfamily Cholevinae (Leiodidae) from the island of Borneo is reviewed, and the results of newly studied material presented. The currently known 30 species (of which 14 are newly described herein) represent the genera Micronemadus (one species), Catops (one species), Baryodirus (one species), Ptomaphaginus (14 species), and Ptomaphaminus (13 species). The following new species are described: Micronemadussondaicus Schilthuizen & Perreau, sp. n., Ptomaphaginusgrandis Schilthuizen & Perreau, sp. n., P.louis Schilthuizen & Perreau, sp. n., P.muluensis Schilthuizen & Perreau, sp. n., and P.isabellarossellini Schilthuizen, Njunjic & Perreau, sp. n., and Ptomaphaminuskinabatanganensis Njunjic, Schilthuizen & Perreau, sp. n., P.testaceus Schilthuizen & Perreau, sp. n., P.nanus Schilthuizen & Perreau, sp. n., P.marshalli Schilthuizen & Perreau, sp. n., P.hanskii Schilthuizen & Perreau, sp. n., P.sarawacensis Schilthuizen & Perreau, sp. n., P.layangensis Schilthuizen & Perreau, sp. n., P.microphallus Schilthuizen & Perreau, sp. n., and P.alabensis Schilthuizen & Perreau, sp. n. It is expected that the cholevine biodiversity of Borneo is still far from completely known. Nonetheless, provisional identification keys to all species known so far are presented.

Comprehensive evolutionary analysis of the Anthroherpon radiation (Coleoptera, Leiodidae, Leptodirini).

The genus Anthroherpon Reitter, 1889 exhibits the most pronounced troglomorphic characters among Coleoptera, and represents one of the most spectacular radiations of subterranean beetles. However, radiation, diversification, and biogeography of this genus have never been studied in a phylogenetic context. This study provides a comprehensive evolutionary analysis of the Anthroherpon radiation, using a dated molecular phylogeny as a framework for understanding Anthroherpon diversification, reconstructing the ancestral range, and exploring troglomorphic diversity. Based on 16 species and 22 subspecies, i.e. the majority of Anthroherpon diversity, we reconstructed the phylogeny using Bayesian analysis of six loci, both mitochondrial and nuclear, comprising a total of 4143 nucleotides. In parallel, a morphometric analysis was carried out with 79 landmarks on the body that were subjected to geometric morphometrics. We optimized morphometric features to phylogeny, in order to recognize the way troglomorphy was expressed in different clades of the tree, and did character evolution analyses. Finally, we reconstructed the ancestral range of the genus using BioGeoBEARS. Besides further elucidating the suprageneric classification of the East-Mediterranean Leptodirini, our main findings also show that Anthroherpon dates back to the Early Miocene (ca. 22 MYA) and that the genus diversified entirely underground. Biogeographic reconstruction of the ancestral range shows the origin of the genus in the area comprising three high mountains in western Montenegro, which is in the accordance with the available data on the paleogeography of the Balkan Peninsula. Character evolution analysis indicates that troglomorphic morphometric traits in Anthroherpon mostly evolve neutrally but may diverge adaptively under syntopic competition.

Revision of the genus Ptomaphagus Hellwig from eastern Asia (Coleoptera, Leiodidae, Cholevinae).

The species belonging to the genus Ptomaphagus Hellwig, 1795 (Coleoptera, Leiodidae, Cholevinae, Ptomaphagini) from eastern Asia are assigned to three species groups. Group yasutoshii has a single species: P. (s. str.) yasutoshii Nishikawa, 1993 from Taiwan, China. Group nepalensis with three species: P. (s. str.) nepalensis Perreau, 1988 from Nepal and P. (s. str.) masumotoi Nishikawa, 2011 from Thailand are redescribed, and P. (s. str.) piccoloi Wang, Ruzicka, Nishikawa, Perreau & Hayashi, 2016 is recorded for the first time from China (Zhejiang). Group sibiricus with seven species, including two newly described Chinese ones P. (s. str.) funiu sp. n. from Henan, and P. (s. str.) haba sp. n. from Yunnan, and five known species: P. (s. str.) chenggongi Wang, Nishikawa, Perreau, Ruzicka & Hayashi, 2016, P. (s. str.) hayashii Wang, Ruzicka, Perreau, Nishikawa & Park, 2016, P. (s. str.) kuntzeni Sokolowski, 1957 (distribution records from Myanmar excluded), P. (s. str.) sibiricus Jeannel, 1934 and P. (s. str.) tingtingtae Wang, Nishikawa, Perreau, Ruzicka & Hayashi, 2016. Specimens of other undescribed species of the group sibiricus are also recorded, revealing a high diversity of this genus in eastern Asia, especially in central and north Sichuan, China, which essentially remains to be investigated. Relevant morphological characters of the examined species are illustrated with colour plates, and their known distributions are mapped. A key to species of Ptomaphagus from eastern Asia is provided.

Revision of the genus Ptomaphagus Hellwig (Coleoptera, Leiodidae, Cholevinae) from Taiwan Island.

Ptomaphagus (s. str.) chenggongi sp. n. and Ptomaphagus (s. str.) tingtingae sp. n. (Coleoptera, Leiodidae, Cholevinae, Ptomaphagini) are described from Taiwan Island. In addition, a new subjective synonym is proposed, Ptomaphagus (s. str.) yasutoshii Nishikawa, 1993 = Ptomaphagus (s. str.) smetanai Perreau, 1996, syn. n. Relevant morphological characters of the examined Ptomaphagus species are illustrated with colour plates, and their known distributions are mapped.

Revision of the genus Ptomaphagus Hellwig (Coleoptera, Leiodidae, Cholevinae) from Japan.

After examining Japanese material of Ptomaphagus Hellwig from various collections, a new species is described, Ptomaphagus (s. str.) piccoloi sp. n., and a new subjective synonym proposed, Ptomaphagus (s. str.) kuntzeni Sokolowski, 1957 = Ptomaphagus (s. str.) amamianus Nakane, 1963, syn. n., in this paper. Relevant morphological characters of examined species of Ptomaphagus are illustrated with colour plates, and known distributions are mapped.

Revision of the genus Ptomaphagus Hellwig (Coleoptera, Leiodidae, Cholevinae) from the Russian Far East and the Korean Peninsula.

The conundrum of Ptomaphagus (s. str.) sibiricus Jeannel, 1934 (Coleoptera, Leiodidae, Cholevinae, Ptomaphagini) is solved, and it is redescribed and newly recorded in South Korea. A new species is also described from the Russian Far East: Ptomaphagus (s. str.) hayashii sp. n. Relevant morphological characters of the concerned species are illustrated with colour plates, and their known distributions are mapped.

One new species of Ptomaphaminus Perreau, 2000 (Coleoptera: Leiodidae: Cholevinae: Ptomaphagini) from a granitic subterranean environment in Vietnam.

Ptomaphaminus granophilus n. sp. from Vietnam is described and illustrated. The special subterranean biotope of this species, deep interstices in a granitic blockfield, is emphasized.

Two new species of the genus Anthroherpon Reitter, 1889 from northern Montenegro with notes on the "A. ganglbaueri" species group (Coleoptera: Leiodidae: Leptodirini).

Two new species of Anthroherpon belonging to the "ganglbaueri" species group are described from northern Montenegro: Anthroherpon sinjajevina n. sp. and Anthroherpon cecai n. sp. Based on morphological investigations of other species of this group, we elevate A. udrzali Giachino & Vailati from subspecies to species rank, and suggest the synonymy of A. brckoensis Giachino & Guéorguiev with A ganglbaueri alticola Knirsch. An identification key of the ganglbaueri species group (sensu Guéorguiev, 1990) is given and the distributions of species and subspecies are mapped and discussed.