Revisionary notes on the genus Adelungia Melichar, 1902 (Hemiptera: Cicadellidae, Megophthalminae), with the description of a new genus.

Leafhopper genus Adelungia Melichar, 1902 is reviewed. Spathicerula Gnezdilov gen. nov. is described with Adelungia calligoni Oshanin, 1908 as the type species, and a new combination-Spathicerula calligoni (Oshanin, 1908), comb. nov., is proposed. Adelungia tridigitata Dlabola, 1984 is placed in synonymy under A. calligoni. Lectotypes are designated for Adelungia elegans Melichar, 1902 and A. breviceps Kusnezov, 1931 to stabilize the nomenclature in the group according to the International Code of Zoological Nomenclature.

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.

A new genus and species of giant mealybugs (Hemiptera: Coccomorpha: Monophlebidae) from eastern China.

A new genus of giant mealybugs (Hemiptera: Coccomorpha; Monophlebidae), Coronaproctus Li, Xu & Wu gen. nov., is established for the species Coronaproctus castanopsis Li, Xu & Wu sp. nov., collected from Zhejiang province, China on the twigs of Castanopsis eyrei (Fagaceae). The adult female and male are described and illustrated. The new genus is characterized by the adult female possessing quadrilocular disc pores, 13 cicatrices on the venter, and a U-shaped opening of the marsupium, but lacking large bilocular pores.

A new species of planthopper from Costa Rica in the genus Herpis (Hemiptera: Derbidae) from palms in the cloud forest.

Recent survey efforts in Costa Rica have documented many new species of planthoppers, primarily in the families Derbidae and Cixiidae, on palms. Recently, a specimen was collected sweeping palms in the Los Angeles cloud forest in Costa Rica and was identified as belonging to the genus Herpis (Derbidae). It was subsequently determined to represent a previously undescribed species. Herein, the new species, Herpis circumsoros Bahder & Bartlett sp. n. is described with supplemental molecular data for the cytochrome c oxidase subunit I (COI) gene and 18S rRNA gene to support placement of the new species in the genus Herpis.

A new genus, Duffelsa gen. n., with descriptions of three new species and one new combination (Hemiptera: Cicadidae).

One new cicada genus, Duffelsa gen. n., is erected in the tribe Leptopsaltriini Moulton, 1923, with the designation of Pomponia orientalis (Distant, 1912) as the type species. Three new species, D. parvula sp. n., D. grandia sp. n. and D. rubida sp. n., are described from China. A key to species of Duffelsa gen. n. is provided. The relationship of this new genus to related taxa is discussed.

Review of Cyphotes Burmeister, 1835 (Hemiptera: Membracidae) with the description of a related new genus.

The Neotropical treehopper genus Cyphotes Burmeister, 1835 (= Aspona Stal, 1862 syn. nov.) is redefined. Cyphotes contains only two species, the type species Cyphotes nodosa Burmeister, 1835 (= Aspona bullata Stl, 1862 syn. nov.) and Cyphotes quadrinodosa (Fonseca & Diringshofen, 1969) reinstated. comb. Allocyphotes gen. nov. (type species Cyphotes insolita Goding, 1929) is proposed to accommodate two other species previously placed in Cyphotes, Allocyphotes pompanoni (Boulard, 2011) comb. nov. and Allocyphotes colombiensis (Gonzlez-Mozo, 2017) comb. nov., and three new species from Ecuador: A. flavus sp. nov., A. waoraniorum sp. nov and A. robertoi sp. nov. for a total of six species. Illustrations, including genitalia images, new locality records and keys to genera and species are provided.

A new species of planthopper in the genus Anchimothon (Hemiptera: Auchenorrhyncha: Derbidae) on palms from Costa Rica.

Anchimothon is a small genus of derbid planthoppers known from Mesoamerica and northern South America. Recent survey efforts in the Caribbean basin have documented many new taxa of fulgoroids from palms. During this survey, a novel taxon identified as Anchimothon was collected from palm seedlings at La Selva Biological Station in Costa Rica. Here, the novel taxon is described as A. myriei sp. n. with molecular data for the cytochrome c oxidase subunit I (COI) and18S rRNA providing support for placement of the new species in Anchimothon. An updated key is provided for the current species of Anchimothon.

Microbiome analyses of 12 psyllid species of the family Psyllidae identified various bacteria including Fukatsuia and Serratia symbiotica, known as secondary symbionts of aphids.

BACKGROUND: Psyllids (Hemiptera: Psylloidea) comprise a group of plant sap-sucking insects that includes important agricultural pests. They have close associations not only with plant pathogens, but also with various microbes, including obligate mutualists and facultative symbionts. Recent studies are revealing that interactions among such bacterial populations are important for psyllid biology and host plant pathology. In the present study, to obtain further insight into the ecological and evolutionary behaviors of bacteria in Psylloidea, we analyzed the microbiomes of 12 psyllid species belonging to the family Psyllidae (11 from Psyllinae and one from Macrocorsinae), using high-throughput amplicon sequencing of the 16S rRNA gene. RESULTS: The analysis showed that all 12 psyllids have the primary symbiont, Candidatus Carsonella ruddii (Gammaproteobacteria: Oceanospirillales), and at least one secondary symbiont. The majority of the secondary symbionts were gammaproteobacteria, especially those of the family Enterobacteriaceae (order: Enterobacteriales). Among them, symbionts belonging to "endosymbionts3", which is a genus-level monophyletic group assigned by the SILVA rRNA database, were the most prevalent and were found in 9 of 11 Psyllinae species. Ca. Fukatsuia symbiotica and Serratia symbiotica, which were recognized only as secondary symbionts of aphids, were also identified. In addition to other Enterobacteriaceae bacteria, including Arsenophonus, Sodalis, and "endosymbionts2", which is another genus-level clade, Pseudomonas (Pseudomonadales: Pseudomonadaceae) and Diplorickettsia (Diplorickettsiales: Diplorickettsiaceae) were identified. Regarding Alphaproteobacteria, the potential plant pathogen Ca. Liberibacter europaeus (Rhizobiales: Rhizobiaceae) was detected for the first time in Anomoneura mori (Psyllinae), a mulberry pest. Wolbachia (Rickettsiales: Anaplasmataceae) and Rickettsia (Rickettsiales: Rickettsiaceae), plausible host reproduction manipulators that are potential tools to control pest insects, were also detected. CONCLUSIONS: The present study identified various bacterial symbionts including previously unexpected lineages in psyllids, suggesting considerable interspecific transfer of arthropod symbionts. The findings provide deeper insights into the evolution of interactions among insects, bacteria, and plants, which may be exploited to facilitate the control of pest psyllids in the future.

New genera and species of felt scales (Hemiptera: Coccomorpha: Eriococcidae), with descriptions of new species and immature instars of described species.

In this work, emphasis is placed on the Eriococcidae (Hemiptera: Coccomorpha) of the Nearctic Region including the immature instars of many species not previously described, two new genera (Carphococcus Miller and Stocks and Ypofloiococcus Miller and Stocks), and 11 new species (Acanthococcus pennyae Miller and Stocks, Carph. apodus Miller and Stocks, Carpochloroides eugeniae Miller and Stocks, Ovaticoccus betsyae Miller and Stocks, Ov. gordoni Miller and Stocks, Ov. haigi Miller and Stocks, Ov. maryfoleybensonae Miller and Stocks, Ov. telotrichus Miller and Stocks, Ov. tuttlei Miller and Stocks, Ov. villanuevorum Miller and Stocks and Y. libeauae Miller and Stocks). In addition to the new taxa, descriptions and illustrations are provided for adult females, adult males and/or immature instars of Ac. arenosus (Cockerell), Ac. coccineus (Cockerell), Ac. dennoi Miller and Miller, Ac. gallicolus (Cockerell & Rohwer), Ac. hoyi Miller and Miller, Ac. kemptoni (Parrott), Apezococcus idiastes Ferris, Carpo. mexicanus Ferris, Cryptococcus fagisuga Lindinger, Cr. williamsi Kosztarab and Hale, Hypericicoccus hyperici (Ferris), Ov. agavacearum Pellizzari and Kozár, Ov. agavium (Douglas), Ov. cornutus (Ferris), Ov. densus (Miller), Ov. eriogoni (Miller), Ov. exoticus Pellizzari and Kozár, Ov. neglectus (Cockerell), Ov. peruvianus (Granara de Willink & Díaz), Ov. salviae Miller, Ov. sanguineus (Miller), Ov. tippinsi Miller and Miller, Ov. variabilis Miller, Spiroporococcus braggi (Cockerell & Robinson), S. yuccae (Ferris) and Xerococcus foquieriae Ferris. Lectotype designations are made for Ap. idiastes. Carpo, mexicanus and H. hyperici. New collection data are provided for Ov. californicus McKenzie, Ov. johnsoni (Miller), Ov. mackenziei Miller, Ov. parvispina (Chaffin) and Ov. viscosa (Kondo). Dichotomous keys are given for: determination of most eriococcid instars of both sexes; Nearctic genera of the Eriococcidae based on adult females; adult females, second-instar females, second-instar males, and first-instar nymphs of Ovaticoccus Kloet 1944; and the seven species of Acanthococcus Signoret 1875 treated here, based on first-instar nymphs. Oregmopyga Hoy 1963 and Cornoculus Ferris 1955 are treated as junior subjective synonyms of Ovaticoccus. Atriplicia Cockerell and Rohwer 1909 has been placed previously as a junior synonym of Eriococcus but here is regarded as a junior subjective synonym of Acanthococcus. We treat 44 species; new descriptions are provided for 101 treated instars, and detailed line drawings are given for all but three of them. The following species, listed in ScaleNet as occurring in the Nearctic Region, are transferred to Acanthococcus from Eriococcus Targioni Tozzetti 1868: Acanthococcus actius Miller and Miller 1993 rev. comb.; Ac. arenariae Miller and Miller 1993 rev. comb.; Ac. barri Miller 1991 rev. comb.; Ac. beshearae Miller and Miller 1993 rev. comb.; Ac. dennoi Miller and Miller 1993 rev. comb.; Ac. droserae Miller, Liu and Howell 1992 rev. comb.; Ac. epacrotrichus Miller and Miller 1992 rev. comb.; Ac. froebeae Miller 1991 rev. comb.; Ac. gallicolus (Cockerell & Rohwer 1909) comb. n.; Ac. hoyi Miller and Miller 1992 rev. comb.; Ac. mackenziei Miller and Miller 1992 rev. comb.; Ac. macrobactrus Miller and Miller 1992 rev. comb.; Ac. megaporus Miller and Miller 1993 rev. comb.; A. mesotrichus Miller and Miller 1993 rev. comb.; Ac. microtrichus Miller and Miller 1992 rev. comb.; Ac. monotrichus Miller and Miller 1993 rev. comb.; Ac. ophius Miller and Miller 1993 rev. comb.; Ac. stauroporus Miller and Miller 1992 rev. comb.; Ac. washingtonensis Miller and Miller 1992 rev. comb.; and Ac. whiteheadi Miller 1991 rev. comb. In addition, two species are transferred to Acanthococcus from Rhizococcus Signoret: Ac. coccineus (Cockerell 1894a) rev. comb. and Ac. kemptoni (Parrott 1900) rev. comb. Three images of water color paintings by Mary Foley Benson are included.

Five new Brazilian species of Fuminana Freytag (Hemiptera: Cicadellidae: Gyponini) with new records and key to males.

Five new Brazilian species of Fuminana Freytag, 1989 are described and illustrated: F. diaphana sp. nov. from states of Maranhão and Piauí, F. guaricica sp. nov. from Paraná State, F. incurvata sp. nov. and F. retorta sp. nov. from Rondônia State, and F. niluga sp. nov. from states of Acre, Pará, Rondônia and Mato Grosso. The female genitalia is described and illustrated for the first time and a revised genus description is presented. New distribution records are given and seven species are newly recorded from Brazil: F. astra (DeLong & Freytag, 1969), F. conspicua (DeLong & Freytag, 1969), Fuminana curra (DeLong & Freytag, 1969), F. extata Freytag, 1989, F. jaculum (DeLong & Freytag, 1969), F. signa (DeLong & Freytag, 1969) and F. superba (DeLong & Freytag, 1969). A key to males is provided as well as some taxonomic notes about the genus.

Complete mitochondrial genome sequencing of Oxycarenus laetus (Hemiptera: Lygaeidae) from two geographically distinct regions of India.

Oxycarenus laetus is a seed-sap sucking pest affecting a variety of crops, including cotton plants. Rising incidence and pesticide resistance by O. laetus have been reported from India and neighbouring countries. In this study, O. laetus samples were collected from Bhatinda and Coimbatore (India). Pure mtDNA was isolated and sequenced using Illumina MiSeq. Both the samples were found to be identical species (99.9%), and the complete genome was circular (15,672 bp), consisting of 13 PCGs, 2 rRNA, 23 tRNA genes, and a 962 bp control region. The mitogenome is 74.1% AT-rich, 0.11 AT, and - 0.19 GC skewed. All the genes had ATN as the start codon except cox1 (TTG), and an additional trnT was predicted. Nearly all tRNAs folded into the clover-leaf structure, except trnS1 and trnV. The intergenic space between trnH and nad4, considered as a synapomorphy of Lygaeoidea, was displaced. Two 5 bp motifs AATGA and ACCTA, two tandem repeats, and a few microsatellite sequences, were also found. The phylogenetic tree was constructed using 36 mitogenomes from 7 super-families of Hemiptera by employing rigorous bootstrapping and ML. Ours is the first study to sequence the complete mitogenome of O. laetus or any Oxycarenus species. The findings from this study would further help in the evolutionary studies of Lygaeidae.

Taxonomic study of the leafhopper genus Anufrievia Dworakowska (Hemiptera: Cicadellidae: Typhlocybinae: Erythroneurini) from China, with descriptions of two new species.

The genus Anufrievia Dworakowska is reviewed with two new species from China: A. anchoroides sp. nov., and A. drepanoides sp. nov.. Detailed morphological descriptions and illustrations of these two new species and a key to all known species (males) from China are provided.

Morphological reassessment of the movable calcar of delphacid planthoppers (Hemiptera: Fulgoromorpha: Delphacidae).

This study presents the morphology of calcar in adult Delphacidae based on representatives of the genera Ugyops Guérin-Meneville, 1834, Notuchus Fennah, 1969 (Ugyopini), Asiraca Latreille, 1798 (Asiracini), Kelisia Fieber, 1866, (Kelisini), Stenocranus Fieber, 1866 (Stenocranini), Chloriona Fieber, 1866, Megadelphax Wagner, 1963, Muellerianella Wagner, 1963, Javesella Fennah, 1963, Conomelus Fieber, 1866, Euconomelus Haupt, 1929, Hyledelphax Vilbaste, 1968, Stiroma Fieber, 1866, Struebingianella Wagner, 1963 and Xanthodelphax Wagner, 1963 (Delphacini). We used SEM electron microscopy, to define seven types of calcar structure (Types 1, 2, 5, 6, 7, 8, and 9) based on combinations of characters including shape, number of teeth and differentiation of sensory structures in species from fifteen genera. Additionally, two other types (Types 3 and 4) were determined based on the calcar descriptions from previous studies. Similarities and differences in calcar structure and function were discussed and emerging relationships between planthopper species and their particular habitats were indicated.

Cytotype Affects the Capability of the Whitefly Bemisia tabaci MED Species To Feed and Oviposit on an Unfavorable Host Plant.

The acquisition of nutritional obligate primary endosymbionts (P-symbionts) allowed phloemo-phageous insects to feed on plant sap and thus colonize novel ecological niches. P-symbionts often coexist with facultative secondary endosymbionts (S-symbionts), which may also influence their hosts' niche utilization ability. The whitefly Bemisia tabaci is a highly diversified species complex harboring, in addition to the P-symbiont "Candidatus Portiera aleyrodidarum," seven S-symbionts whose roles remain poorly understood. Here, we compare the phenotypic and metabolic responses of three B. tabaci lines differing in their S-symbiont community, reared on three different host plants, hibiscus, tobacco, or lantana, and address whether and how S-symbionts influence insect capacity to feed and produce offspring on those plants. We first show that hibiscus, tobacco, and lantana differ in their free amino acid composition. Insects' performance, as well as free amino acid profile and symbiotic load, were shown to be plant dependent, suggesting a critical role for the plant nutritional properties. Insect fecundity was significantly lower on lantana, indicating that it is the least favorable plant. Remarkably, insects reared on this plant show a specific amino acid profile and a higher symbiont density compared to the two other plants. In addition, this plant was the only one for which fecundity differences were observed between lines. Using genetically homogeneous hybrids, we demonstrate that cytotype (mitochondria and symbionts), and not genotype, is a major determinant of females' fecundity and amino acid profile on lantana. As cytotypes differ in their S-symbiont community, we propose that these symbionts may mediate their hosts' suitable plant range. IMPORTANCE Microbial symbionts are universal in eukaryotes, and it is now recognized that symbiotic associations represent major evolutionary driving forces. However, the extent to which symbionts contribute to their hosts' ecological adaptation and subsequent diversification is far from being fully elucidated. The whitefly Bemisia tabaci is a sap feeder associated with multiple coinfecting intracellular facultative symbionts. Here, we show that plant species simultaneously affect whiteflies' performance, amino acid profile, and symbiotic density, which could be partially explained by differences in plant nutritional properties. We also demonstrate that, on lantana, the least favorable plant used in our study, whiteflies' performance is determined by their cytotype. We propose that the host plant utilization in B. tabaci is influenced by its facultative symbiont community composition, possibly through its impact on the host dietary requirements. Altogether, our data provide new insights into the impact of intracellular microorganisms on their animal hosts' ecological niche range and diversification.

Complete Mitochondrial Genomes of Metcalfa pruinosa and Salurnis marginella (Hemiptera: Flatidae): Genomic Comparison and Phylogenetic Inference in Fulgoroidea.

The complete mitochondrial genomes (mitogenomes) of two DNA barcode-defined haplotypes of Metcalfa pruinosa and one of Salurnis marginella (Hemiptera: Flatidae) were sequenced and compared to those of other Fulgoroidea species. Furthermore, the mitogenome sequences were used to reconstruct phylogenetic relationships among fulgoroid families. The three mitogenomes, including that of the available species of Flatidae, commonly possessed distinctive structures in the 1702-1836 bp A+T-rich region, such as two repeat regions at each end and a large centered nonrepeat region. All members of the superfamily Fulgoroidea, including the Flatidae, consistently possessed a motiflike sequence (TAGTA) at the ND1 and trnS2 junction. The phylogenetic analyses consistently recovered the familial relationships of (((((Ricaniidae + Issidae) + Flatidae) + Fulgoridae) + Achilidae) + Derbidae) in the amino acid-based analysis, with the placement of Cixiidae and Delphacidae as the earliest-derived lineages of fulgoroid families, whereas the monophyly of Delphacidae was not congruent between tree-constructing algorithms.

Mitochondrial Genomes from Two Specialized Subfamilies of Reduviidae (Insecta: Hemiptera) Reveal Novel Gene Rearrangements of True Bugs.

Reduviidae, a hyper-diverse family, comprise 25 subfamilies with nearly 7000 species and include many natural enemies of crop pests and vectors of human disease. To date, 75 mitochondrial genomes (mitogenomes) of assassin bugs from only 11 subfamilies have been reported. The limited sampling of mitogenome at higher categories hinders a deep understanding of mitogenome evolution and reduviid phylogeny. In this study, the first mitogenomes of Holoptilinae (Ptilocnemus lemur) and Emesinae (Ischnobaenella hainana) were sequenced. Two novel gene orders were detected in the newly sequenced mitogenomes. Combined 421 heteropteran mitogenomes, we identified 21 different gene orders and six gene rearrangement units located in three gene blocks. Comparative analyses of the diversity of gene order for each unit reveal that the tRNA gene cluster trnI-trnQ-trnM is the hotspot of heteropteran gene rearrangement. Furthermore, combined analyses of the gene rearrangement richness of each unit and the whole mitogenome among heteropteran lineages confirm Reduviidae as a 'hot-spot group' of gene rearrangement in Heteroptera. The phylogenetic analyses corroborate the current view of phylogenetic relationships between basal groups of Reduviidae with high support values. Our study provides deeper insights into the evolution of mitochondrial gene arrangement in Heteroptera and the early divergence of reduviids.

Characterization, Comparative Analysis and Phylogenetic Implications of Mitogenomes of Fulgoridae (Hemiptera: Fulgoromorpha).

The complete mitogenomes of nine fulgorid species were sequenced and annotated to explore their mitogenome diversity and the phylogenetics of Fulgoridae. All species are from China and belong to five genera: Dichoptera Spinola, 1839 (Dichoptera sp.); Neoalcathous Wang and Huang, 1989 (Neoalcathous huangshanana Wang and Huang, 1989); Limois Stål, 1863 (Limois sp.); Penthicodes Blanchard, 1840 (Penthicodes atomaria (Weber, 1801), Penthicodes caja (Walker, 1851), Penthicodes variegata (Guérin-Méneville, 1829)); Pyrops Spinola, 1839 (Pyrops clavatus (Westwood, 1839), Pyrops lathburii (Kirby, 1818), Pyrops spinolae (Westwood, 1842)). The nine mitogenomes were 15,803 to 16,510 bp in length with 13 protein-coding genes (PCGs), 22 transfer RNA genes (tRNAs), 2 ribosomal RNA genes (rRNAs) and a control region (A + T-rich region). Combined with previously reported fulgorid mitogenomes, all PCGs initiate with either the standard start codon of ATN or the nonstandard GTG. The TAA codon was used for termination more often than the TAG codon and the incomplete T codon. The nad1 and nad4 genes varied in length within the same genus. A high percentage of F residues were found in the nad4 and nad5 genes of all fulgorid mitogenomes. The DHU stem of trnV was absent in the mitogenomes of all fulgorids sequenced except Dichoptera sp. Moreover, in most fulgorid mitogenomes, the trnL2, trnR, and trnT genes had an unpaired base in the aminoacyl stem and trnS1 had an unpaired base in the anticodon stem. The similar tandem repeat regions of the control region were found in the same genus. Phylogenetic analyses were conducted based on 13 PCGs and two rRNA genes from 53 species of Fulgoroidea and seven outgroups. The Bayesian inference and maximum likelihood trees had a similar topological structure. The major results show that Fulgoroidea was divided into two groups: Delphacidae and ((Achilidae + (Lophopidae + (Issidae + (Flatidae + Ricaniidae)))) + Fulgoridae). Furthermore, the monophyly of Fulgoridae was robustly supported, and Aphaeninae was divided into Aphaenini and Pyropsini, which includes Neoalcathous, Pyrops, Datua Schmidt, 1911, and Saiva Distant, 1906. The genus Limois is recovered in the Aphaeninae, and the Limoisini needs further confirmation; Dichoptera sp. was the earliest branch in the Fulgoridae.

A new planthopper species in the genus Omolicna (Hemiptera: Auchenorrhyncha: Derbidae) from the Reserva Privada el Silencio de Los Angeles Cloud Forest in Costa Rica.

Recent survey work on palms in Costa Rica has resulted in the discovery of several new species of Derbidae, especially in the Cenchreini. During a recent expedition, specimens collected by light trapping at the Hotel Villa Blanca (cloud forest) were determined to be a novel species of Omolicna Fennah. Sequence data was generated for the novel taxon for the cytochrome c oxidase subunit I (COI) and 18S loci. Pairwise analyses and phylogenetic analyses support placement of the novel taxon in Omolicna.

Taxonomy of the leafhopper genus Nirvanguina (Hemiptera: Cicadellidae: Deltocephalinae: Drabescini) with description of a new species from China.

The leafhopper genus Nirvanguina Zhang Webb (Cicadellidae: Deltocephalinae: Drabescini: Paraboloponina) is reviewed, including description of a new species, Nirvanguina bifurcatus Xu Zhang sp. nov., from China. An updated checklist along with distribution and a key to males of this genus are also provided. Type specimens of this new species are deposited in the Entomological Museum, Northwest A F University, Yangling, Shaanxi, China (NWAFU).

Fuscmacula, a new leafhopper genus from China (Hemiptera: Cicadellidae: Deltocephalinae).

A new leafhopper genus of Athysanini from China, Fuscmacula gen. nov. with F. biprocessa sp. nov. as type species is described and illustrated. This species was collected from grass. The type specimens of the new species are deposited in the Institute of Entomology, Guizhou University, Guiyang, China (GUGC).