Tightening the requirements for species diagnoses would help integrate DNA-based descriptions in taxonomic practice.

Modern advances in DNA sequencing hold the promise of facilitating descriptions of new organisms at ever finer precision but have come with challenges as the major Codes of bionomenclature contain poorly defined requirements for species and subspecies diagnoses (henceforth, species diagnoses), which is particularly problematic for DNA-based taxonomy. We, the commissioners of the International Commission on Zoological Nomenclature, advocate a tightening of the definition of "species diagnosis" in future editions of Codes of bionomenclature, for example, through the introduction of requirements for specific information on the character states of differentiating traits in comparison with similar species. Such new provisions would enhance taxonomic standards and ensure that all diagnoses, including DNA-based ones, contain adequate taxonomic context. Our recommendations are intended to spur discussion among biologists, as broad community consensus is critical ahead of the implementation of new editions of the International Code of Zoological Nomenclature and other Codes of bionomenclature.

Phylogeny and diversification of planthoppers (Hemiptera: Fulgoromorpha) based on a comprehensive molecular dataset and large taxon sampling.

Our understanding of the evolution of Fulgoromorpha (Insects, Hemiptera) has relied on molecular studies that have only considered either a limited number of taxa where all the families were not represented simultaneously, or a reduced number of genes.The absence of a global analysis comparing all the available data has thus led to significant biases in the analyzes, as evidenced by the incongruence of the results reported for planthopper phylogeny. Here we provide a phylogenetic and dating analysis of the Fulgoromorpha with a large sampling of 531 ingroup taxa, representing about 80% of the currently described suprageneric taxonomic diversity in this group. This study is based on most of the molecular sequences available to date and duly verified, for a set of nuclear and mitochondrial genes from a taxonomic sampling as complete as possible. The most significant results of our study are: (1) the unexpected paraphyly of Delphacidae whose Protodelphacida seem more related to Cixiidae than to other Delphacidae;(2) the group Meenoplidae-Kinnaridae recovered sister to the remaining Fulgoroidea families; (3) the early branching node of Tettigometridae sister of all the other families;(4) the Achilidae-Derbidae clade with Achilidae Plectoderini including Achilixiidae recovered as monophyletic as well as theFulgoridae-Dictyopharidae clade; and (5) the Tropiduchidae placed sister to the other so called 'higher' families (sec. Shcherbakov, 2006).Our divergence times analysis, calibrated with a set of duly verified fossils, suggests that the first diversification of planthoppers occurred in the Early Triassic around 240 Mya and those of the superfamilies Delphacoidea and Fulgoroidea in the Middle-Late Triassic around 210 Mya and 230 Mya, respectively. By the end of the Jurassic, all major planthopper lineages were originated, and all families, around 125 Mya, might havebeen driven in their distribution and evolution (in their first subfamilial divisions) by the geographical constraints of the Gondwanan break-up.Rapid evolutionary radiations occurred particularly in Fulgoridae around 125-130 Mya. Our results stress the importance of the good quality of the sequences used in the molecular analyzes and the primordial importance of a large sampling when analyzing the phylogeny of the group.

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.

Planthopper family Issidae (Insecta: Hemiptera: Fulgoromorpha): linking molecular phylogeny with classification.

A molecular phylogeny of the planthopper family Issidae (Hemiptera, Fulgoroidea) is provided using both Maximum likelihood and Bayesian inference analyses. The phylogeny is based on 18S, two parts of 28S, COXI and Cytb genes from 50 genera and 79 ingroup species (including 8 species recently excluded from Issidae). As with the only few previous studies, an important taxonomic impediment is observed with the sampling; however for the first time, all analyses depict several fully supported lineages, which challenge the recent proposed "modern classification" of the family. It also highlights a strong coherence between these lineages and their respective geographical distribution. All previously excluded taxa are confirmed as not being part of the Issidae as recently defined which monophyly is confirmed. Accordingly, a new classification of the family is proposed with 3 subfamilies and 7 tribes as follows. Neotropical issid Thioniini in Thioniinae stat. rev. is re-established as an independent lineage sister to all other Issidae. Palaearctic Issidae are weakly supported as a monophyletic lineage, Issinae stat. nov., including 2 tribes: Issini stat. nov. (genera Issus and Latissus) and Hysteropterini stat. rev. (all other Palaearctic genera). Oriental Issidae form a strongly supported monophyletic subfamily group Hemisphaeriinae stat. rev. including 4 tribes: Kodaianellini trib. nov., Sarimini trib. nov., Parahiraciini Cheng & Yang, 1991, and Hemisphaeriini Melichar, 1906, the latter including 2 subtribes: Mongolianina s.trib. nov., and Hemisphaeriina Melichar, 1906. A Neotropical lineage including the genus Picumna is provisionally placed in incertae sedis within the Hemisphaeriinae stat. nov.

New Cretaceous Fossil Achilidae Taxa (Insecta, Hemiptera, Fulgoromorpha) from Burmese Ambers with Description of Niryasaburniini Trib. Nov.

A new species Niryasaburnia nigrutomia sp. nov. of the planthopper family Achilidae from Burmese amber collected from Hukawng Valley (Tanai) of northern Myanmar, is described, notably based on forewing pattern coloration and metatibiotarsal teeth conformation. A new fossil genus with its type species Sinuovenaxius kachinensis gen. et sp. nov. is also described. The tribe Niryasaburniini trib. nov. is established to include Niryasaburnia Szwedo, 2004, and Sinuovenaxius gen. nov., based on a unique combination of characters, of which the following states are particularly notable: head with compound eyes around half the length of pronotum, late forking of ScP+R and CuA after the fusion of Pcu+A1 on the forewing, apical teeth of metatarsomeres I and II both with subapical platellar sensilla, and a unique hindwing pattern with simple RP and biforked MP, CuA with two terminals only, and with A2 simple, reaching the posterior wing margin. The hindwing venation of this new tribe with RP with only one terminal and both MP and CuA with two terminals is unique in Achilidae.

Two New Species of Pentacarinus from the Acrotiarini Tribe in Burmese Amber (Hemiptera, Fulgoromorpha, Cixiidae).

Two new species, Pentacarinus maculosus sp. nov. and Pentacarinus tenebrosus sp. nov., from Burmese amber are described. Alongside the type species P. kachinensis already described, they are easily distinguishable by the tegmina patterns. The diagnosis of the genus Pentacarinus is amended, notably with fusion of Pcu + A1 distad of forking CuA, the fork ScP + R approximately close to basal 1/5 of tegmen, basad of forking CuA, and only one transverse veinlet ir between RP and RA on forewings. Additionally, a key to these three species of Pentacarinus is provided.

Disparity of metatibial and metatarsal cuticular and sensory structures in Cixiidae (Hemiptera: Fulgoromorpha) with a metatibiotarsal diagnosis for the tribes.

A review of the cuticular and sensory metatibial and metatarsal structures in cixiid planthoppers (Hemiptera: Fulgoromorpha) is proposed, depicting both their strong disparity and the great diversity of the patterns observed. Terminology and definitions for these structures are provided. The so-called lateral teeth of the metatibia in fact are particular styloconic sensory structures, called spiniform sensilla in Cixiidae. The apical metatibiotarsal teeth are non-sensory cuticular expansions, often bearing one or several chaetic sensilla ventrally, generally distributed in an internal and an external group of three teeth each, in some instances separated by a diastema; innermost and outermost teeth are generally larger. On the first tarsomere seven to eight teeth generally occur in one row, although two rows are observed in Brixidiini. A strong diversity of conformations and patterns is observed in the second metatarsomere. A specific subdorsal sensillum, of platellar type, may be present in the first metatarsomere teeth for a few taxa. It is generally present in the second metatarsomere, either as a narrow-based acutellar sensillum or as a broad-based platellar sensillum according to the taxon. Scanning electron microscope (SEM) analyses of 54 species of cixiids from all described tribes of the family, supplemented by data from the literature, are used to provide a metatibiotarsal diagnosis for each of the tribes of Cixiidae. In the state of our knowledge of the sufficiently precise observations of metatibiotarsal structures in the Cixiidae which are available, and of the phylogeny of the group as a frame of reference for their interpretations, we note that the observed patterns are probably the result of multiple and independent convergences and evolutionary regressions. These occurred at all levels of cixiid classification. Although these patterns can be useful in the identification of taxa at a low taxonomic level, they would be less useful for phylogenetic approaches.

Taxonomic and nomenclatural notes on some Andes species (Hemiptera, Fulgoromorpha, Cixiidae) with two synonymies and one homonymy.

Two African Cixiidae species names, Andes goniodes and Andes synavei proposed by Emeljanov in 2001 to correct two homonymies published by Synave in 1953, are synonymized respectively with Andes muiri and Andes spinifer, which were already proposed as replacement names by Synave in 1970. Additionally, the name Andes synafenni is proposed as new for the Vietnamese species Andes truncatus Fennah, 1978 preoccupied by the African species Andes truncatus Synave, 1953.

Bioinspire-Explore: Taxonomy-Driven Exploration of Biodiversity Data for Bioinspired Innovation.

Successful bioinspired design depends on practitioners' access to biological data in a relevant form. Although multiple open-access biodiversity databases exist, their presentation is often adapted to life scientists, rather than bioinspired designers. In this paper, we present a new tool, "Bioinspire-Explore", for navigating biodiversity data in order to uncover biological systems of interest for a range of sectors. Bioinspire-Explore allows users to search for inspiring biological models via taxa (species, genera, etc.) as an entry point. It provides information on a taxon's position in the "tree of life", its distribution and climatic niche, as well as its appearance. Bioinspire-Explore also shows users connections in the bioinspiration literature between their taxon of interest and associated biological processes, habitats, and physical measurements by way of their semantic proximity. We believe Bioinspire-Explore has the potential to become an indispensable resource for both biologists and bioinspired designers in different fields.

Toward a new classification of planthoppers Hemiptera Fulgoromorpha: 2. Higher taxa, their names and their composition.

A synthesis of planthopper suprafamilial and familial classification is provided and discussed, solving some nomenclatural issues for the group, and adopting an enriched presentation with a clear distinction between the taxon name and its concept.

Spotted beautyGedanochila museisucini gen. et sp. nov.a new Achilidae from Eocene Baltic amber (Hemiptera: Fulgoromorpha) and its relation to Achilini.

Gedanochila museisucini gen. et sp. nov. is described, based on inclusions in the Eocene Baltic amber. A morphological phylogenetic analysis supports the placement of Gedanochila gen. nov. into the tribe Achilini. Definition, content and subdivisions of the tribe as well as position of extinct taxa placed within are briefly discussed.

A New Cixiid Fossil Genus of the Tribe Acrotiarini from Mid-Cretaceous Burmese Amber (Insecta, Hemiptera, Fulgoromorpha).

A new Burmese amber genus Maculixiusgen. nov. with its type species Maculixius jiewenaesp. nov. is described in the planthopper family Cixiidae. This new genus is unique in Burmese Cixiidae by its forewing venation, with Pcu and A1 merging in the middle of clavus, the late bifurcation of ScP+R slightly after this level, and the early CuA forking well before this level. Although the head capsule is missing, it belongs to the recently described tribe Acrotiarini, based on the pentacarinated mesonotum and the distinctly arched RA on forewing with cell C1 wider submedially than apically. Morphological characteristics of Acrotiarini are discussed, and a key to identification of Acrotiarini genera is provided. The new taxon broadens the knowledge of the tribe, and it underlines the already great diversity of the family in the Cretaceous.

Distribution patterns of Chinese Cixiidae (Hemiptera, Fulgoroidea), highlight their high endemic diversity.

BACKGROUND: Cixiidae are small strictly phytophagous hemipteran insects worldwide distributed. Ecology and systematics of Chinese fauna remains poorly investigated. For instance, does their distribution follows the patterns of biogeogaphical distribution established for their host plants or other related-taxa because they are all obligatory phytophagous taxa? Do they follow the usual distributional Chinese realms and boundaries already recognized? Which zoogeographical Chinese regions and connections between them do they depict. To investigate these issues, we provide here a referenced and comprehensive checklist of the 250 cixiid species currently reported from China (77 new records), with their precise distribution at the regional level. In the 8 Chinese main zoogeographical regions usually recognized and 2 adjacent areas, we analyzed further their diversity at the tribal, generic, and specific levels using a non-metric multidimensional scaling and an unweighted pairwise group analysis using an arithmetic mean cluster analyses. The observed distribution patterns shown that an intercalary Sino-Japanese realm is recognisable between the Palaearctic and Oriental realms. At the regional level, the South China region clusters more closely with the Southwest, Central and North China regions. Taiwan, clearly separated from the South China region and mainland China, is more closely related to the Qinghai-Tibet region and Indochina countries. Although Central and South China regions remain close to each other, the Qinghai-Tibet region appears singularly different. NEW INFORMATION: An updated checklist of the 250 Cixiidae species, known to occur in China and counting for 10% of the Chinese planthopper fauna, is presented based on literature, recent collections, and museum records. More than 400 records distributed among the 28 provinces and 8 regions in China are extensively provided, including 77 new records. Of these, more than 80% of the species (205 species, 82%) have been only reported from China, and most of them are endemic species, which could reflects the great diversity degree of the Chinese regions and local biotypes highlights the uniqueness of this fauna. These species are found in 8 Chinese zoogeographical regions: The Taiwan region is the most diversified with 161 species and the highest rate of endemic species (69.57%), followed by South China (78 species, 17.95%), Central China (60 species, 33.33%), Southwest China (43 species, 39.53%), North China (29 species, 34.48%), Qinghai-Tibet region (10 species, 20%), Northeast China (8 species, 12.5%), and 5 species found in the Inner Mongolia-Xinjiang region that are not endemic ones. Endemism was analyzed for each region and repeated for species distribution patterns across them, 9 being bi-regionally and tri-regionally distributed. The South China-Taiwan pattern is the most richest one, followed by the Central-South China-Taiwan pattern. Semonini and Pentastirini tribes are widespread among all the zoological regions, representing respectively 21.20% and 17.20% of all the species, while Cixiini being is the most common tribe with 45.20%, remains absent from the North-Eastern China region. Andini with only 5.20% of the species is distributed in the Sino-Japanese - Oriental Region; Eucarpini (6.40%) and Borysthenini (2.00%) are mainly concentrated in the south of the Qingling Mountain-Huai River. The remaining four tribes, Bennini (0.40%), Briixini (0.80%), Oecleini (1.20%) and Stenophlepsiini (0.40%) are relatively rare and restricted to Taiwan. At the generic level, Kuvera (7.2%) is the most widely distributed genus in China while Cixius, Betacixius, Kuvera, Oecleopsis and Andes are the more diversified. One genus (Oliparisca) is distributed only in the Tibet region, while 10 genera are distributed only in the Taiwan region. In addition, nearly half of the genera (16 genera, 48.48%) are distributed south of the Palearctic/Oriental boundary. A non-metric multidimensional scaling and an unweighted pairwise group method analysis using arithmetic mean clustering based on the Jaccard similarity coefficient matrix support a Palaearctic/Sino-Japanese boundary and a South China region closer to the Southwest, Central and North China regions. The Taiwan region appears clearly separated from the South China region and to mainland China, and more closely related to the Qinghai-Tibet region and Indochina countries. The Central and South China regions appear close to each other, but the Qinghai-Tibet region is singularly isolated.

Four new species of the planthopper genus Metanigrus Tsaur, Yang & Wilson from China (Hemiptera, Fulgoromorpha, Meenoplidae).

Four new species of the genus Metanigrus Tsaur, Yang & Wilson, 1986, M. chromus Lv & Chen, sp. nov., M. guttatus Lv & Chen, sp. nov., M. gremius Lv & Chen, sp. nov., and M. spinatus Lv, Chen & Bourgoin, sp. nov. from China (Hubei, Guizhou and Yunnan), are described and illustrated, giving the genus six species in total. A key to all known species of Metanigrus is provided, as well as a map of their geographic distributions.

Two new species of the genus Saigona Matsumura (Hemiptera, Fulgoromorpha, Dictyopharidae) from China.

Two new species of the genus Saigona Matsumura, 1910, S.baiseensis Zheng & Chen sp. nov. and S.maculata Zheng & Chen sp. nov., from China (Guanxi) are described and illustrated. A revised identification key to the 16 species of Saigona is provided. 15 species of the genus are known from China only.

Complete mitochondrial genome of the planthopper Orthopagus splendens (Germar, 1830) (Hemiptera: Fulgoromorpha: Dictyopharidae).

The first complete mitochondrial genome of a dictyopharid planthopper, Orthopagus splendens (Germar, 1830) (Hemiptera: Fulgoroidea: Dictyopharidae) is sequenced. The 15,349 bp long complete mitogenome contains 13 protein-coding genes (PCGs), 22 tRNA genes, two rRNA genes, and 1 A + T-rich region with an arrangement identical to that observed in most insect mitogenomes (GenBankNo. MW441850). All PCGs start with ATN, and end with TAN or single T (nad1, nad5, and atp6). A phylogenetic analysis places O. splendens as sister to Fulgoridae confirming a sister relationship between Dictyopharidae and Fulgoridae.

A new genus of the tribe Sarimini (Fulgoromorpha, Issidae) from the Guangxi Province of China.

A new genus with a new species Eusarimissus hezhouensis gen. nov. et sp. nov. from Guangxi Province of China are described in the tribe Sarimini of the family Issidae. Molecular sequences of 18S, 28S and COXI genes are provided for the new taxon. Phylogenetic analysis places this taxon sister to a previously sequenced but not yet described Sarimini genus 'Eusarima sp. 4'. Taxonomic notes are provided for the genus Eusarima Yang, 1994. The species Eusarima (Nepalius) iranica Gnezdilov & Mozaffarian, 2011 is transferred to the genus Sarima Melichar 1903.

Parahiraciini (Hemiptera, Fulgoromorpha, Issidae): one new genus, two new species and three new subtribes.

A new genus Pusulissus gen. nov. with two new species P. phiaoacensis sp. nov. and P. coronomensis sp. nov. are described respectively from Vietnam and China in the tribe Parahiraciini of the family Issidae. A molecular phylogeny using combined genes (18S, 28S, COX1 and Cytb) confirms its placement in the tribe Parahiraciini. The tribe is divided into three subtribes: Scantiniinasubtribe nov. with the type genus Scantinius Stål, 1866, and Vindilisinasubtribe nov. with type genus Vindilis Stål, 1870 plus Nisoprincessa Gnezdilov, 2017, and the nominal subtribe Parahiraciina Cheng & Yang, 1991 subtribe nov. The characteristics of these subtribes are provided, with a key to identification. Genus Folifemurum Che, Zhang & Wang, 2013 is transferred to HemisphaeriiniMongolianina, and genus Gelastyrella Yang, 1994 is maintained as a valid name.

Notes on genera of Sarimini with description of two new species in the genera Microsarimodes and Tetrichina (Hemiptera: Fulgoromorpha: Issidae).

Two new species of the tribe Sarimini in the planthopper family Issidae: Microsarimodes flavomaculata sp. nov. and Tetrichina fuscovinclum sp. nov. are described from China. The genera Duplexissus Wang, Zhang Bourgoin, 2019 and Sarimissus Wang, Zhang Bourgoin, 2019 are reestablished as valid genera based on both morphological grounds and molecular analyses.