Diversification of the phytophagous lineages of true bugs (Insecta: Hemiptera: Heteroptera) shortly after that of the flowering plants.

More than 95% of phytophagous true bug (Hemiptera: Heteroptera) species belong to four superfamilies: Miroidea (Cimicomorpha), Pentatomoidea, Coreoidea, and Lygaeoidea (all Pentatomomorpha). These iconic groups of highly diverse, overwhelmingly phytophagous insects include several economically prominent agricultural and silvicultural pest species, though their evolutionary history has not yet been well resolved. In particular, superfamily- and family-level phylogenetic relationships of these four lineages have remained controversial, and the divergence times of some crucial nodes for phytophagous true bugs have hitherto been little known, which hampers a better understanding of the evolutionary processes and patterns of phytophagous insects. In the present study, we used 150 species and concatenated nuclear and mitochondrial protein-coding genes and rRNA genes to infer the phylogenetic relationships within the Terheteroptera (Cimicomorpha + Pentatomomorpha) and estimated their divergence times. Our results support the monophyly of Cimicomorpha, Pentatomomorpha, Miroidea, Pentatomoidea, Pyrrhocoroidea, Coreoidea, and Lygaeoidea. The phylogenetic relationships across phytophagous lineages are largely congruent at deep nodes across the analyses based on different datasets and tree-reconstructing methods with just a few exceptions. Estimated divergence times and ancestral state reconstructions for feeding habit indicate that phytophagous true bugs explosively radiated in the Early Cretaceous-shortly after the angiosperm radiation-with the subsequent diversification of the most speciose clades (Mirinae, Pentatomidae, Coreinae, and Rhyparochromidae) in the Late Cretaceous.

Taxonomic corrections for Asian Fingulus (Hemiptera: Heteroptera: Miridae).

The identities of two East and Southeast Asian species of the plant bug genus Fingulus Distant, 1904 (Hemiptera: Heteroptera: Miridae: Deraeocorinae: Deraeocorini) are clarified based on their type specimens. The following new subjective synonymies are proposed: F. longicornis Miyamoto, 1965 = F. brevirostris Ren, 1983, syn. nov.; and F. ruficeps Hsiao Ren, 1983 = F. henrytomi Nakatani Yasunaga, 2018, syn. nov. Previous records of F. umbonatus Stonedahl Cassis, 1991 from Hainan, China, and F. collaris Miyamoto, 1965 from Zhejiang and Yunnan, China, are regarded as misidentifications of F. ruficeps and F. inflatus Stonedahl Cassis, 1991, respectively.

Taxonomic corrections for East and Southeast Asian Reduviidae (Hemiptera: Heteroptera).

The identities of certain East and Southeast Asian genera and species of Reduviidae (Hemiptera: Heteroptera) are clarified based on their type materials and taxonomic conclusions are drawn. The following new subjective synonymies and new combinations are proposed: Harpactorinae: Cutocoris Stål, 1859 = Paracydnocoris Miller, 1954, syn. nov.; Cutocoris distinctus (Miller, 1954), comb. nov. (transferred from Paracydnocoris) = Cydnocoris ventralis Hsiao, 1979, syn. nov.; Cutocoris macgillavryi (Miller, 1954), comb. nov. (transferred from Paracydnocoris); Henricohahnia wangi Ren, 2001 = H. obscara Cai Li, 2003, syn. nov.; Pahabengkakia Miller, 1941 = Stalireduvius Tomokuni Cai, 2004, syn. nov.; Pahabengkakia piliceps Miller, 1941 = S. nodipes Tomokuni Cai, 2004, syn. nov.; Reduviinae: Reduvius xantusi (Horváth, 1879), comb. nov. (transferred from Velitra) = R. decliviceps Hsiao, 1976, syn. nov. Pahabengkakia piliceps is recorded from Laos for the first time.

Aphelonotus schuhi sp. nov., the first s--pecies of the family Pachynomidae from Puerto Rico (Hemiptera: Heteroptera).

Aphelonotus schuhi sp. nov. (Hemiptera: Heteroptera: Pachynomidae: Aphelonotinae) is described from Puerto Rico based on a single male collected by flight interception trap. The new species is morphologically similar to A. taino Schuh, Weirauch Grillo, 2015 and A. xenos Schuh, Weirauch Grillo, 2015, being distinguished from them by the combination of the presence of a stout subapical spine on the protibia and the characteristic shape of the parameres. An informal A. xenos species-group is proposed to accommodate these three species.

From lanceolate to plate-like: Gross morphology, terminology, and evolutionary trends of the trichophoran ovipositor.

The ovipositor morphology of Trichophora (Hemiptera: Heteroptera) is revisited. Skeletomuscular structure of the ovipositor of selected species and outgroups is documented. Homologies of the structures are established, different homology hypotheses of previous authors are discussed and rejected. The groundplan of the trichophoran ovipositor is reconstructed, apomorphic conditions of each part of the ovipositor are documented. A standard nomenclature is proposed for muscles associated with the ovipositor and extrinsic muscles of the female inner ectodermal genital tracts. Character transformations of the ovipositor are reconstructed via cladistic analysis. The concepts of "lanceolate ovipositor" and "plate-like ovipositor", frequently used in the literature, are discussed; it is concluded that no unambiguous definition of them is possible on a morphological basis, these terms refer merely to evolutionary grades rather than strictly distinct character states. "Plate-like ovipositors" evolved at least four times within Trichophora; they exhibit considerable differences among and within these clades. It is demonstrated that the "M- or W-shaped sclerites" of Pyrrhocoridae and Urostylididae are not homologous: in Pyrrhocoridae they are expansions of the mesal face of the posterior portion of valvifers IX, whilst the superficially similar structures in Urostylididae are infoldings of the ventral rim of the mesal portion of laterotergites IX.

When did the ancestor of true bugs become stinky? Disentangling the phylogenomics of Hemiptera-Heteroptera.

The phylogeny of true bugs (Hemiptera: Heteroptera), one of the most diverse insect groups in terms of morphology and ecology, has been the focus of attention for decades with respect to several deep nodes between the suborders of Hemiptera and the infraorders of Heteroptera. Here, we assembled a phylogenomic data set of 53 taxa and 3102 orthologous genes to investigate the phylogeny of Hemiptera-Heteroptera, and both concatenation and coalescent methods were used. A binode-control approach for data filtering was introduced to reduce the incongruence between different genes, which can improve the performance of phylogenetic reconstruction. Both hypotheses (Coleorrhyncha + Heteroptera) and (Coleorrhyncha + Auchenorrhyncha) received support from various analyses, in which the former is more consistent with the morphological evidence. Based on a divergence time estimation performed on genes with a strong phylogenetic signal, the origin of true bugs was dated to 290-268 Ma in the Permian, the time in Earth's history with the highest concentration of atmospheric oxygen. During this time interval, at least 1007 apomorphic amino acids were retained in the common ancestor of the extant true bugs. These molecular apomorphies are located in 553 orthologous genes, which suggests the common ancestor of the extant true bugs may have experienced large-scale evolution at the genome level.

A new synonymy in East Asian Urostylididae (Hemiptera: Heteroptera).

Urostylididae is a moderately species-rich family of phytophagous, mainly arboricolous pentatomoid true bugs (Hemiptera: Heteroptera) currently comprising 8 genera and 172 species (Rider et al. 2018). All described species occur in Asia, but undescribed taxa are present in Madagascar (P. Stys, pers. comm.). The present short paper was prompted by a recent contribution by Kim et al. (2018), who reviewed the species of the genus Urostylis Westwood, 1837, occurring in Korea, including the description of a purportedly new species, U. koreana Kim Jung, 2018. The identity of the latter species is clarified below, with proposal of a new subjective synonymy.

A review of the species of the tribe Chorosomatini of China (Hemiptera: Heteroptera: Rhopalidae).

Species of the tribe Chorosomatini (Hemiptera: Heteroptera: Rhopalidae: Rhopalinae) occurring in China are reviewed. Type materials of three species described from China, so far of uncertain identities, are re-examined and documented, and taxonomic changes are accordingly proposed. The following synonymies, suspected but not explicitly proposed by previous authors, are confirmed and explicitly proposed: Agraphopus lethierryi Stål, 1872 = A. yunnanus Hsiao, 1965; Leptoceraea viridis Jakovlev, 1873 = L. granulosa Hsiao, 1965; Chorosoma macilentum Stål, 1858 = Ch. brevicolle Hsiao, 1964. Faunistic records of other species of the tribe are provided, and several misidentifications in the previous literature are corrected; Ch. gracile Josifov, 1968, and Ch. schillingii (Schilling, 1829) are recorded for the first time from China. A checklist of the species and subspecies of Chorosomatini occurring in China, with a review of their distribution, and a key for their identification, are provided.

The genus Alphocoris in the Indomalaya (Hemiptera: Heteroptera: Scutelleridae).

The Indomalayan species of the genus Alphocoris Germar, 1839 (Hemiptera: Heteroptera: Scutelleridae: Odontotarsinae: Odontotarsini) are revised. Three species, A. caudatus Rédei, Tsai Jindra, sp. nov. (India: Maharashtra), A. naso Rédei Tsai, sp. nov. (India: Goa), and A. asper Rédei, Tsai Jindra, sp. nov. (India: Goa and Maharashtra), are recognized. The type material of A. lixoides Germar, 1839 (type locality: Senegal) is documented; previous records of this species from India and Pakistan are considered as based on misidentifications, the species is restricted to the Afrotropical Region.

A revision of Graptophara (Hemiptera: Heteroptera: Scutelleridae).

The Indomalayan genus Graptophara Stål, 1865 (Hemiptera: Heteroptera: Scutelleridae: Scutellerinae) is redescribed and revised, and its systematic position is discussed. Two species, G. reynaudii (Guérin-Méneville, 1834) (the type species, distributed in the Sundaic Region) and G. anomala sp. nov. (southern India) are recognized, redescribed or described, and illustrated. A lectotype is designated for Callidea pulchra Westwood, 1837, now a junior synonym of G. reynaudii.

Phylogeny and the colourful history of jewel bugs (Insecta: Hemiptera: Scutelleridae).

Members of the family Scutelleridae (Heteroptera: Pentatomomorpha: Pentatomoidea) are also called shield bugs because of the greatly enlarged scutellum, or jewel bugs because of the brilliant colours of many species. All scutellerids are phytophagous, feeding on various parts of their host plants. Due to lack of obvious synapomorphies and the failure to apply rigorous phylogenetic methods, the higher classification of Scutelleridae has been disputed for more than 150 years. Here we reconstructed a phylogeny of Scutelleridae based on complete sequences of 18S and 28S nuclear rDNAs and all 13 protein-coding genes of the mitochondrial genome, with the sampled taxa covering all of the currently recognized subfamilies. The monophyly of Scutelleridae was confirmed by the congruence of the results of analyses conducted using Bayesian inference, maximum likelihood and maximum parsimony. The phylogenetic relationships among subfamilies were well resolved for the first time. Furthermore, time-divergence studies estimated that the time of origin of Scutelleridae was in the Early Cretaceous (142.1-122.8 Ma), after the origin of the angiosperms. The diversification between the extant subfamilies of Scutelleridae and within the subfamilies occurred from the late Palaeocene to the late Miocene, simultaneously with the rise of the major groups of angiosperms and other phytophagous insects.

Taxonomic corrections on Cronion (Hemiptera: Heteroptera: Plataspidae).

Vollenhoven (1864) presented a preliminary report and subsequently (Vollenhoven 1865) a detailed description of a new genus of pentatomoid true bugs (Hemiptera: Heteroptera), Poseidon Vollenhoven, 1864, with a single included species, P. malayanus Vollenhoven, 1864; the species is therefore the type species of the genus by monotypy. The genus was placed into Scutelleridae by Vollenhoven (1864, 1865), but the latter taxon was defined broadly, in a way much different from the current concept of this family (see e.g. Rider et al. 2017); Stål (1876) transferred it into Plataspidae defined in the present sense. Bergroth (1891) showed that Poseidon was preoccupied by Poseidon Girard, 1825, a valid genus of ribbon worms (Nemertea), and therefore he proposed a replacement name, Cronion Bergroth, 1891, for it. Miller (1955) described another new plataspid genus, Triodocoris Miller, 1955, and designated the newly described T. cinctus Miller, 1955, as its type species. Both of these genera have remained monotypic, and no author have presented new data on them since their original descriptions.

Physoderinae Miller, 1954 (Hemiptera: Heteroptera: Reduviidae): a junior synonym of Epiroderinae Distant, 1904, and a junior homonym of Physoderina Chaudoir, 1877 (Coleoptera: Carabidae).

The purpose of the present paper is to review and resolve nomenclatural problems with the family-group name Physoderinae Miller, 1954 (type genus: Physoderes Westwood, 1846), a name being in prevailing usage for a subfamily of assassin bugs (Hemiptera: Heteroptera: Reduviidae). It is demonstrated that this name is a junior synonym of Epiroderinae Distant, 1904 (type genus: Epirodera Westwood, 1847, a junior objective synonym of Physoderes) and permanently invalid as a junior homonym of the family-group name Physoderina Chaudoir, 1877 (type genus: Physodera Eschscholtz, 1829) (Coleoptera: Carabidae: Harpalinae: Lebiini), therefore it is replaced by its senior synonym, Epiroderinae.

Acrosternum heegeri Fieber, 1861 (Hemiptera: Heteroptera: Pentatomidae), another Mediterranean bug expanding to the north.

Acrosternum heegeri Fieber, 1861 (Hemiptera: Heteropera: Pentatomidae: Pentatominae: Pentatomini), a Holomediterranean-Turanian shield bug species, is recorded for the first time from Hungary, representing the first known occurrence of the species in Central Europe. The species is illustrated, its geographic distribution is reviewed. The occurrence of A. heegeri in two localities in Budapest, regular records from one of the localities in the years 2015 and 2016, and the finding of an overwintering individual indicate the presence of a stable population in the city; the species is probably already established and it is locally abundant. Its further spread in Europe is expected.

The identity of Equatobursa, with proposal of new genus and species level synonymies (Hemiptera: Heteroptera: Heterogastridae).

The identities of the genus Equatobursa Zou, 1985 and its single included species, E. nigra Zou, 1985 (Hemiptera: Heteroptera: Lygaeoidea: Heterogastridae) are clarified based on the re-examination of the type material of the species. The following new subjective synonymies are proposed: Sadoletus Distant, 1903 = Equatobursa Zou, 1985, syn. nov.; Sadoletus izzardi Hidaka, 1959 = Equatobursa nigra Zou, 1985, syn. nov.

Fossil record of stem groups employed in evaluating the chronogram of insects (Arthropoda: Hexapoda).

Insecta s. str. (=Ectognatha), comprise the largest and most diversified group of living organisms, accounting for roughly half of the biodiversity on Earth. Understanding insect relationships and the specific time intervals for their episodes of radiation and extinction are critical to any comprehensive perspective on evolutionary events. Although some deeper nodes have been resolved congruently, the complete evolution of insects has remained obscure due to the lack of direct fossil evidence. Besides, various evolutionary phases of insects and the corresponding driving forces of diversification remain to be recognized. In this study, a comprehensive sample of all insect orders was used to reconstruct their phylogenetic relationships and estimate deep divergences. The phylogenetic relationships of insect orders were congruently recovered by Bayesian inference and maximum likelihood analyses. A complete timescale of divergences based on an uncorrelated log-normal relaxed clock model was established among all lineages of winged insects. The inferred timescale for various nodes are congruent with major historical events including the increase of atmospheric oxygen in the Late Silurian and earliest Devonian, the radiation of vascular plants in the Devonian, and with the available fossil record of the stem groups to various insect lineages in the Devonian and Carboniferous.

Resolving the taxonomy and nomenclature of Metochus abbreviatus (Hemiptera: Heteroptera: Rhyparochromidae).

The identity of Metochus abbreviatus Scott, 1874 (Hemiptera: Heteroptera: Rhyparochromidae: Rhyparochrominae: Rhyparochromini) is clarified based on reexamination of the lectotype. The subjective synonymy of Dieuches kreyenbergi Breddin, 1906 with M. abbreviatus is confirmed. The following new subjective synonymies are proposed: Rhyparochromus erosus Walker, 1872 = Metochus abbreviatus Scott, 1874, syn. nov. = Metochus holsti Distant, 1918, syn. nov. Bibliographies of the names are provided and it is demonstrated that the name M. abbreviatus is in prevailing usage. Reversion to the senior name Rh. erosus is considered to be undesirable, therefore an application has simultaneously been submitted to the International Commission on Zoological Nomenclature to give the specific name abbreviatus precedence over the specific name erosus. The diagnostic characters, bionomics and distribution of the species are briefly reviewed.

A new synonymy in the genus Macrocixius (Hemiptera: Fulgoromorpha: Cixiidae).

Until recently the genus Macrocixius Matsumura, 1914 (Hemiptera: Fulgoromorpha: Cixiidae) contained two species: M. giganteus Matsumura, 1914 (the type species) and M. grossus Tsaur & Hsu, 1991. Two recent papers revised the genus within a short interval of time. Both Zhang & Chen (2013) and Orosz (2013) provided redescriptions of the genus and its two previously included species; Zhang & Chen (2013) described two new species from China; Orosz (2013) described five new species from Nepal, Taiwan, Vietnam, and Indonesia.

Lectotype designations and taxonomic corrections on Neotropical Scutelleridae described by G. Breddin (Hemiptera: Heteroptera).

Lectotypes are designated and documented for the following species and infrasubspecific taxa: Dystus villosus Breddin, 1904; Lobothyreus breviceps Breddin, 1914; Pachycoris torridus (Scopoli, 1772) var. laetissimus Breddin, 1906 (originally proposed as var. laetissima); P. torridus var. moestissimus Breddin, 1906 (originally proposed as var. moestissima); Polytes speculiger Breddin, 1914. The identities of the taxa in concern are clarified. The following new junior subjective synonyms are proposed: Pachycoris torridus (Scopoli, 1772) = P. torridus var. laetissimus Breddin, 1906, syn. nov., = P. torridus var. moestissimus Breddin, 1906, syn. nov.

The identity of the Brachyplatys species recently introduced to Panama, with a review of bionomics (Hemiptera: Heteroptera: Plataspidae).

A recent report of a population of Brachyplatys vahlii (Fabricius, 1787) (Hemiptera: Heteroptera: Plataspidae) introduced to Panama is considered as misidentification, the species in concern is recognized as B. subaeneus (Westwood, 1837). Syntypes of B. subaeneus and diagnostic characters of the species are illustrated, published information on its distribution, bionomics and economic importance is reviewed. Syntypes of B. vahlii are illustrated, taxonomic problems in connection with the species are highlighted.