The earliest known holometabolous insects.

The Eumetabola (Endopterygota (also known as Holometabola) plus Paraneoptera) have the highest number of species of any clade, and greatly contribute to animal species biodiversity. The palaeoecological circumstances that favoured their emergence and success remain an intriguing question. Recent molecular phylogenetic analyses have suggested a wide range of dates for the initial appearance of the Holometabola, from the Middle Devonian epoch (391 million years (Myr) ago) to the Late Pennsylvanian epoch (311 Myr ago), and Hemiptera (310 Myr ago). Palaeoenvironments greatly changed over these periods, with global cooling and increasing complexity of green forests. The Pennsylvanian-period crown-eumetabolan fossil record remains notably incomplete, particularly as several fossils have been erroneously considered to be stem Holometabola (Supplementary Information); the earliest definitive beetles are from the start of the Permian period. The emergence of the hymenopterids, sister group to other Holometabola, is dated between 350 and 309 Myr ago, incongruent with their current earliest record (Middle Triassic epoch). Here we describe five fossils--a Gzhelian-age stem coleopterid, a holometabolous larva of uncertain ordinal affinity, a stem hymenopterid, and early Hemiptera and Psocodea, all from the Moscovian age--and reveal a notable penecontemporaneous breadth of early eumetabolan insects. These discoveries are more congruent with current hypotheses of clade divergence. Eumetabola experienced episodes of diversification during the Bashkirian-Moscovian and the Kasimovian-Gzhelian ages. This cladogenetic activity is perhaps related to notable episodes of drying resulting from glaciations, leading to the eventual demise in Euramerica of coal-swamp ecosystems, evidenced by floral turnover during this interval. These ancient species were of very small size, living in the shadow of Palaeozoic-era 'giant' insects. Although these discoveries reveal unexpected Pennsylvanian eumetabolan diversity, the lineage radiated more successfully only after the mass extinctions at the end of the Permian period, giving rise to the familiar crown groups of their respective clades.

Laying the foundations of evolutionary and systematic studies in crickets (Insecta, Orthoptera): a multilocus phylogenetic analysis.

Orthoptera have been used for decades for numerous evolutionary questions but several of its constituent groups, notably crickets, still suffer from a lack of a robust phylogenetic hypothesis. We propose the first phylogenetic hypothesis for the evolution of crickets sensu lato, based on analysis of 205 species, representing 88% of the subfamilies and 71% tribes currently listed in the database Orthoptera Species File (OSF). We reconstructed parsimony, maximum likelihood and Bayesian phylogenies using fragments of 18S, 28SA, 28SD, H3, 12S, 16S, and cytb (~3600 bp). Our results support the monophyly of the cricket clade, and its subdivision into two clades: mole crickets and ant-loving crickets on the one hand, and all the other crickets on the other (i.e. crickets sensu stricto). Crickets sensu stricto form seven monophyletic clades, which support part of the OSF families, "subfamily groups", or subfamilies: the mole crickets (OSF Gryllotalpidae), the scaly crickets (OSF Mogoplistidae), and the true crickets (OSF Gryllidae) are recovered as monophyletic. Among the 22 sampled subfamilies, only six are monophyletic: Gryllotalpinae, Trigonidiinae, Pteroplistinae, Euscyrtinae, Oecanthinae, and Phaloriinae. Most of the 37 tribes sampled are para- or polyphyletic. We propose the best-supported clades as backbones for future definitions of familial groups, validating some taxonomic hypotheses proposed in the past. These clades fit variously with the morphological characters used today to identify crickets. Our study emphasizes the utility of a classificatory system that accommodates diagnostic characters and monophyletic units of evolution. Moreover, the phylogenetic hypotheses proposed by the present study open new perspectives for further evolutionary research, especially on acoustic communication and biogeography.

300 million years of diversification: elucidating the patterns of orthopteran evolution based on comprehensive taxon and gene sampling.

Orthoptera is the most diverse order among the polyneopteran groups and includes familiar insects, such as grasshoppers, crickets, katydids, and their kin. Due to a long history of conflicting classification schemes based on different interpretations of morphological characters, the phylogenetic relationships within Orthoptera are poorly understood and its higher classification has remained unstable. In this study, we establish a robust phylogeny of Orthoptera including 36 of 40 families representing all 15 currently recognized superfamilies and based on complete mitochondrial genomes and four nuclear loci, in order to test previous phylogenetic hypotheses and to provide a framework for a natural classification and a reference for studying the pattern of divergence and diversification. We find strong support for monophyletic suborders (Ensifera and Caelifera) as well as major superfamilies. Our results corroborate most of the higher-level relationships previously proposed for Caelifera, but suggest some novel relationships for Ensifera. Using fossil calibrations, we provide divergence time estimates for major orthopteran lineages and show that the current diversity has been shaped by dynamic shifts of diversification rates at different geological times across different lineages. We also show that mitochondrial tRNA gene orders have been relatively stable throughout the evolutionary history of Orthoptera, but a major tRNA gene rearrangement occurred in the common ancestor of Tetrigoidea and Acridomorpha, thereby representing a robust molecular synapomorphy, which has persisted for 250 Myr.

Inside the Melanoplinae: new molecular evidence for the evolutionary history of the Eurasian Podismini (Orthoptera: Acrididae).

The Podismini are melanopline grasshoppers with a Holarctic distribution and well represented in the Eurasian fauna. To investigate their controversial taxonomy and evolutionary history, we studied 86%, 78% and 33% respectively of the Eurasian, European and Asian Palaearctic genera (Otte, 1995; Eades et al., 2013). We reconstructed parsimony, maximum likelihood and Bayesian phylogenies using fragments of four genes (ITS1, 16S, 12S, CO2). We applied a Bayesian molecular clock to estimate the times of species divergence, and the event-based parsimony method to depict the biogeographic framework of the diversification. Our results suggest that the selected Eurasian Podismini constitute a monophyletic group inside the Melanoplinae, provided it includes the North American genus Phaulotettix. The clades proposed by the present study inside the Podismini do not fit the older morphological or cytological classifications, but are in agreement with more recent proposals. Furthermore, our results can be explained by a plausible biogeographic history in which the present geographical distribution of the Eurasian Podismini resulted from known changes, to the Cenozoic climate and vegetation, induced by major geological events including the genesis of high mountain chains (e.g., Himalayas, Altay, Alps) and large deserts (e.g., Gobi, Karakoum, Taklamakan), and the opening of marginal seas (e.g., Bering, Japanese and Yellow Seas).

Revision of the genus Pteroplistes in India, with the description of two new species Pteroplistes kervasae Jaiswara, n. sp. and Pteroplistes masinagudi Jaiswara, n. sp. (Orthoptera, Grylloidea, Pteroplistinae).

Pteroplistes Brunner von Wattenwyl, 1873 is an Indo-Malaysian cricket genus with only one species, P. platycleis Bolivar, 1899(1900) known from India. Here, we redescribe the genus Pteroplistes and P. platycleis and describe two new Indian species, Pteroplistes kervasae Jaiswara, n. sp. and Pteroplistes masinagudi Jaiswara, n. sp. from the Western Ghats, using morphology and genitalia. Identification keys to separate the Indian species of Pteroplistes are provided and the distribution of the genus in India is discussed.

Zebragryllus Desutter-Grandcolas & Cadena-Casteñada, n.gen. a new Gryllinae genus from Eastern and Western Amazonia, South America (Orthoptera, Grylloidea, Gryllidae).

We describe a new genus of grylline cricket, Zebragryllus Desutter-Grandcolas & Cadena-Casteñada n. gen., from the Neotropical Region, using characters of morphology and male genitalia; genitalic characters clearly show that Zebragryllus n. gen. is closely related to Anurogryllus Saussure, 1878. Six species are described as new to science, originating from western (Peru, Colombia) and eastern (French Guiana) Amazonia: Zebragryllus fuscus Desutter-Grandcolas, n. sp., Z. guianensis Desutter-Grandcolas, n. sp., Z. intermedius Desutter-Grandcolas, n. sp., Zebragryllus nauta Desutter-Grandcolas, n. sp., Zebragryllus nouragui Desutter-Grandcolas, n. sp., and Zebragryllus wittoto Desutter-Grandcolas and Cadena-Casteñada, n. sp., type species of the genus. They are characterized by their size, coloration (shining black, most often with white patterns of coloration, hence the genus name), and male and female genitalia. The calling songs of Z. guianensis Desutter-Grandcolas, n. sp., Z. intermedius Desutter-Grandcolas, n. sp., Z. nouragui Desutter-Grandcolas, n. sp., and Z. wittoto Desutter-Grandcolas and Cadena-Casteñada, n. sp. are described. An identification key is proposed for both males and females.

Mechanisms of high-frequency song generation in brachypterous crickets and the role of ghost frequencies.

Sound production in crickets relies on stridulation, the well-understood rubbing together of a pair of specialised wings. As the file of one wing slides over the scraper of the other, a series of rhythmic impacts causes harmonic oscillations, usually resulting in the radiation of pure tones delivered at low frequencies (2-8 kHz). In the short-winged crickets of the Lebinthini tribe, acoustic communication relies on signals with remarkably high frequencies (>8 kHz) and rich harmonic content. Using several species of the subfamily Eneopterinae, we characterised the morphological and mechanical specialisations supporting the production of high frequencies, and demonstrated that higher harmonics are exploited as dominant frequencies. These specialisations affect the structure of the stridulatory file, the motor control of stridulation and the resonance of the sound radiator. We placed these specialisations in a phylogenetic framework and show that they serve to exploit high-frequency vibrational modes pre-existing in the phylogenetic ancestor. In Eneopterinae, the lower frequency components are harmonically related to the dominant peak, suggesting they are relicts of ancestral carrier frequencies. Yet, such ghost frequencies still occur in the wings' free resonances, highlighting the fundamental mechanical constraints of sound radiation. These results support the hypothesis that such high-frequency songs evolved stepwise, by a form of punctuated evolution that could be related to functional constraints, rather than by only the progressive increase of the ancestral fundamental frequency.

Stalacris Desutter-Grandcolas n. gen., an amazing cricket from South Africa (Orthoptera, Grylloidea, Phalangopsidae).

Stalacris n. gen. (Insecta, Grylloidea, Phalangopsidae) is described from South Africa using characters of morphology and male genitalia. This taxon, known from only two species, Stalacris meridionalis n. gen., n. sp. and Stalacris sp, is characterized by the unique structure of its forewings. These are prolonged distally as long, acute and more or less articulated processes, forming a kind of forceps when forewings are opened.

New pseudophyllinae from the Lesser Antilles (Orthoptera: Ensifera: Tettigoniidae).

Two new Cocconitini Brunner von Wattenwyl, 1895 species belonging to Nesonotus Beier, 1960 are described from the Lesser Antilles: Nesonotus caeruloglobus Hugel, n. sp. from Dominica, and Nesonotus vulneratus Hugel, n. sp. from Martinique. The songs of both species are described and elements of biology are given. The taxonomic status of species close to Nesonotus tricornis (Thunberg, 1815) is discussed.

Nomenclatural and taxonomic problems related to the electronic publication of new nomina and nomenclatural acts in zoology, with brief comments on optical discs and on the situation in botany.

In zoological nomenclature, to be potentially valid, nomenclatural novelties (i.e., new nomina and nomenclatural acts) need first to be made available, that is, published in works qualifying as publications as defined by the International Code of zoological Nomenclature ("the Code"). In September 2012, the Code was amended in order to allow the recognition of works electronically published online after 2011 as publications available for the purpose of zoological nomenclature, provided they meet several conditions, notably a preregistration of the work in ZooBank. Despite these new Rules, several of the long-discussed problems concerning the electronic publication of new nomina and nomenclatural acts have not been resolved. The publication of this amendment provides an opportunity to discuss some of these in detail. It is important to note that: (1) all works published only online before 2012 are nomenclaturally unavailable; (2) printed copies of the PDFs of works which do not have their own ISSN or ISBN, and which are not obtainable free of charge or by purchase, do not qualify as publications but must be seen as facsimiles of unavailable works and are unable to provide nomenclatural availability to any nomenclatural novelties they may contain; (3) prepublications online of later released online publications are unavailable, i.e., they do not advance the date of publication; (4) the publication dates of works for which online prepublications had been released are not those of these prepublications and it is critical that the real release date of such works appear on the actual final electronic publication, but this is not currently the case in electronic periodicals that distribute such online prepublications and which still indicate on their websites and PDFs the date of release of prepublication as that of publication of the work; (5) supplementary online materials and subsequent formal corrections of either paper or electronic publications distributed only online are nomenclaturally unavailable; (6) nomenclatural information provided on online websites that do not have a fixed content and format, with ISSN or ISBN, is unavailable. We give precise examples of many of these nomenclatural problems. Several of them, when they arise, are due to the fact that the availability of nomenclatural novelties now depends on information that will have to be sought not from the work itself but from extrinsic evidence. As shown by several examples discussed here, an electronic document can be modified while keeping the same DOI and publication date, which is not compatible with the requirements of zoological nomenclature. Therefore, another system of registration of electronic documents as permanent and inalterable will have to be devised. ZooBank also clearly needs to be improved in several respects. Mention in a work of its registration number (LSID) in ZooBank would seem to be possible only if this registration has occurred previously, but some works that have purportedly been registered in ZooBank are in fact missing on this web application. In conclusion, we offer recommendations to authors, referees, editors, publishers, libraries and the International Commission on Zoological Nomenclature, in the hope that such problems can be limited along with the potential chaos in zoological nomenclature that could result, if careful attention is not paid to the problems we highlight here, from a somewhat misplaced, and perhaps now widespread, understanding that electronic publication of nomenclatural novelties is now allowed and straightforward. We suggest that, as long as the problematic points linked to the new amendment and to electronic publication as a whole are not resolved, nomenclatural novelties continue to be published in paper-printed journals that have so far shown editorial competence regarding taxonomy and nomenclature, which is not the case of several recent electronic-only published journals.

New intertidal crickets from Comoros and Mascarene islands (Orthoptera: Trigonidiidae: Nemobiinae: Burcini).

Nemobiinae crickets of the tribe Burcini Gorochov, 1986 are described for the first time from the shores of South Western Indian Ocean islands. The new genus Makalapobius n. gen. is proposed to include M. aigrettensis n. gen. n. sp. from Mauritius and M. masihu n. gen. n. sp. from Grande Comore, and the new genus Gabusibius n. gen. to include G. ndzilu n. gen. n. sp. from Anjouan, G. mosi n. gen. n. sp., from Mohéli, and G. dzindzanu n. gen. n. sp. from Mayotte. The species Speonemobius littoreus Vannini Chelazzi, 1978 from Somalia coast is tentatively placed in the genus Gabusibius n. gen. as G. ? litoreus (Vannini Chelazzi, 1978) n. gen. n. comb. The songs of G. mosi n. gen. n. sp. and M. aigrettensis n. gen. n. sp. are described. The threats to SWIO Burcini and endemism of Orthoptera from SWIO coastal areas are discussed.

Taxonomic revision of Teleogryllus mitratus (Burmeister, 1838) and T. occipitalis (Serville, 1838) in India, with the description of Teleogryllus rohinae Jaiswara amp; Jain sp. nov. and a key for Teleogryllus species from India (Orthoptera: Gryllidae).

The genus Teleogryllus is known from 52 species distributed worldwide, of which India is home to 11 species. We update Teleogryllus diversity by describing a new species, T. rohinae Jaiswara Jain sp. nov., from Kerala, India. In addition, we revise the description of the two widespread Teleogryllus species, i.e., T. mitratus (Burmeister, 1838) and T. occipitalis (Serville, 1838) and describe their female genital structures for the first time. Finally, we provide updated identification keys for all Indian Teleogryllus species.

The Phalangopsidae crickets (Orthoptera, Grylloidea) of the Seychelles Archipelago: Taxonomy of an ecological radiation.

The Phalangopsidae crickets (Grylloidea) of the Seychelles are examined following extensive field sampling on several main islands of the archipelago (Mah, Silhouette, Praslin, La Digue). Despite the small area of these islands, six genera (12 species) are documented, including one new genus and five new species. The type species of the genus Seychellesia Bolivar, 1912 is transferred to the genus Paragryllodes Karny, 1909 as Paragryllodes nitidula (Bolivar, 1912) n. comb. The other species described in Seychellesia are transferred to the genus Seselia Hugel Desutter-Grandcolas, n. gen., as Seselia longicercata (Bolivar, 1912) n. comb. and Seselia patellifera (Bolivar, 1912) n. comb. Two new species are also described in the genus Seselia Hugel Desutter-Grandcolas, n. gen., Seselia coccofessei Hugel Desutter-Grandcolas, n. gen., n. sp. (type species of the genus) and Seselia matyoti Hugel Desutter-Grandcolas, n. gen., n. sp. The genera Phaeogryllus Bolivar, 1912 and Phalangacris Bolivar, 1895 are redescribed, including Phalangacris ferlegro Hugel Desutter-Grandcolas, n. sp. and Phalangacris sotsote Hugel Desutter-Grandcolas, n. sp. that are new to science. The genus Gryllapterus Bolivar, 1912 is redescribed and transferred from the Landrevinae (Gryllidae) to the Cachoplistinae (Phalangopsidae). New tribes are defined for the genus Paragryllodes (Paragryllodini Hugel Desutter-Grandcolas, n. tribe) on the one hand, and for Seselia Hugel Desutter-Grandcolas, n. gen., Phalangacris, Phaeogryllus and Gryllapterus (Seselini Hugel Desutter-Grandcolas, n. tribe) on the other, using morphological characters and the results of molecular phylogenetic studies (Warren et al. 2019). Phaloria (Papuloria) insularis (Bolivar, 1912) (Phaloriinae) is redescribed and restricted to Mah, and its calling song is documented for the first time, while Phaloria (Papuloria) bolivari Hugel Desutter-Grandcolas, n. sp. is newly described from Silhouette. Identification keys are proposed for the genera of Seselini Hugel Desutter-Grandcolas, n. tribe, and for the species of Seselia Hugel Desutter-Grandcolas, n. gen. and Phalangacris. The confusion between the Mogoplistidae Ornebius succineus Bolivar, 1912 and the Phalangopsidae Heterotrypus succineus Bolivar, 1910 is discussed, and the name Subtiloria succineus (Bolivar, 1912) considered a nomen nudum.

The genus Arachnomimus/ Saussure, 1897 (Orthoptera: Phalangopsidae) in India, with description of a new subgenus and species from a cave in Chhattisgarh, India.

The genus Arachnomimus Saussure, 1897 is one of the 13 genera of Phalangopsidae present in India. A total of 11 species belongs to this genus, of which, 9 are grouped in the subgenus Arachnomimus Saussure, 1897 with their distribution on the Indian subcontinent. One species occurs in Brazil and another in Euarachnomimus Gorochov, 1996 known from Southeast Asia. India hosts two species, Arachnomimus (Arachnomimus) lepidus Chopard, 1969 and Arachnomimus (Arachnomimus) maindroni (Chopard, 1969). In the present paper, we describe a new subgenus, Indimimus Desutter-Grandcolas Jaiswara and a new species Indimimus jayanti Jaiswara under genus Arachnomimus. Identification keys for all the three subgenera and three species from India are proposed.

Colonization of different biomes drove the diversification of the Neotropical Eidmanacris crickets (Insecta: Orthoptera: Grylloidea: Phalangopsidae).

The phylogeny of the cricket genus Eidmanacris is used to analyse its historical distribution and diversification in three South American biomes: Atlantic Forest, Cerrado, and Chiquitano Dry Forest. A morphological phylogeny with all the 29 species of Eidmanacris and the Geographically explicit Event Model (GEM) is used to explain their colonization and diversification through three different biomes and their ancestral habitats and distributional areas. We analysed ecologically-significant characters, such as body size and metanotal characters, to test whether if morphology, habitat, or behaviour are connected. The relations of these features with the colonisation of wetter or drier biomes based on the distributional area, phylogeny and diversity of the genus were also tested. The results show that the ancestral distribution of the genus was the Atlantic Forest, and that biome occupancy, habitat, size, and mating behaviour evolved congruently through the phylogeny, drawing a coherent pattern of changes through Eidmanacris evolution toward the colonisation of drier biomes. Our results indicate that gallery forests could play a key role in the distribution and diversification of Eidmanacris species.

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.

A new genus and species of cricket from eastern Cuba: the first Antillean true Eneopterinae (Orthoptera: Gryllidae: Eneopterinae).

The occurrence of true Eneopterinae in the Antilles is recorded herein for the first time, with the description of a new genus and species herein described from eastern Cuba (Greater Antilles): Antillobinthus inexpectatus Yong Desutter-Grandcolas, n. gen. n. sp. It is described and illustrated in detail, including color photographs of habitus, morphologically diagnostic characters and habitat. The present finding revives a biogeographical debate, as no Eneopterinae was up-to-now known from the Antilles.

First Record of Dericorys albidula Serville, 1838 (Orthoptera: Dericorythidae) in Tunisia and Libya.

The humpbacked grasshopper Dericorys albidula (Serville, 1838) (Orthoptera: Dericorythidae: Dericorythinae) is reported here for the first time from Southern Tunisia (Tozeur province) and northwestern Libya. Details on the ecological occurrence of D. albidula are given and compared to available data from Central Asia. A lectotype is designated for D. albidula and its type locality identified as the Sinaï desert in Egypt.

A new genus and five new species of Anostostomatidae from the Lesser Antilles (Orthoptera: Ensifera).

Most high volcanic islands of Lesser Antilles harbor one single genus of Anostostomatidae: Rhumosa n. gen: Rhumosa bolognei n. gen. n. sp. in Guadeloupe, Rhumosa macoucheriei n. gen. n. sp. in Dominica, Rhumosa depazei n. gen. n. sp. in Martinique, Rhumosa admiralrodneyei n. gen. n. sp. in Saint Lucia, Rhumosa captainblighei n. gen. n. sp., in Saint Vincent. These species are restricted to well preserved rainforests; species from northern islands apparently occurring at higher elevation than species of southern islands. The distribution and generic position of Rhumosa n. gen. species is discussed, as well as the generic position of Lutosa cubaensis (Haan, 1843).

New insights on basivenal sclerites using 3D tools and homology of wing veins in Odonatoptera (Insecta).

Being implied in flight, mimetism, communication, and protection, the insect wings were crucial organs for the mega diversification of this clade. Despite several attempts, the problem of wing evolution remains unresolved because the basal parts of the veins essential for vein identification are hidden in the basivenal sclerites. The homologies between wing characters thus cannot be accurately verified, while they are of primary importance to solve long-standing problems, such as the monophyly of the Palaeoptera, viz. Odonatoptera, Panephemeroptera, and Palaeozoic Palaeodictyopterida mainly known by their wings. Hitherto the tools to homologize venation were suffering several cases of exceptions, rendering them unreliable. Here we reconstruct the odonatopteran venation using fossils and a new 3D imaging tool, resulting congruent with the concept of Riek and Kukalová-Peck, with important novelties, viz. median anterior vein fused to radius and radius posterior nearly as convex as radius anterior (putative synapomorphies of Odonatoptera); subcostal anterior (ScA) fused to costal vein and most basal primary antenodal crossvein being a modified posterior branch of ScA (putative synapomorphies of Palaeoptera). These findings may reveal critical for future analyses of the relationships between fossil and extant Palaeoptera, helping to solve the evolutionary history of the insects as a whole.