Phylogeny and historical biogeography of leafhopper subfamily Iassinae (Hemiptera: Cicadellidae) with a revised tribal classification based on morphological and molecular data

Phylogenetic relationships among major lineages of the leafhopper subfamily Iassinae were explored by analysing a dataset of 91 discrete morphological characters and DNA sequence data from nuclear 28S rDNA and histone H3 genes and mitochondrial 12S rDNA. Bayesian, maximum‐likelihood and maximum parsimony analyses yielded similar tree topologies that were well resolved with strong branch support except at the base of the tree, resulting in equivocal support for inclusion of Bythoniini as a tribe of Iassinae but strong support for the monophyly of Iassinae (excluding Bythoniini) and most previously recognized iassine tribes. Divergence times for recovered nodes were estimated using a Bayesian relaxed clock method with two fossil calibration points. The results suggest that the deepest divergences coincided with Gondwanan vicariant events but that more recent divergences resulted from long‐range dispersal and colonization. Biogeographical analyses suggest that the group most likely has a Neotropical origin. The following changes to the taxonomic classification are proposed: establishment of three new tribes, Batracomorphini trib.n. (based on type genus Batracomorphus Lewis), Hoplojassini trib.n. (based on type genus Hoplojassus Dietrich and including one other South American genus), Lipokrisnini trib.n. (based on type genus Lipokrisna Freytag and including two other endemic Caribbean genera); Krisnini is redefined to include only the Old World genera Krisna and Gessius; Iassini is redefined to include only the type genus and four endemic Afrotropical genera; Bascarrhinus Fowler and Platyhynna Berg, recently treated as genera incertae sedis, are placed in Hyalojassini; Thalattoscopus Kirkaldy is added to the previously monobasic tribe Trocnadini. Iassinae now includes 12 tribes, all of which appear to be monophyletic. Revised morphological diagnoses of the subfamily and each of the included tribes are provided and a key to tribes is also given. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:41295B68‐2DAB‐4C4F‐B260‐F7C054922173 .     

Phylogeny and classification of the leafhopper subfamily Eurymelinae (Hemiptera: Cicadellidae) inferred from molecules and morphology

Phylogenetic analysis of the globally distributed arboreal leafhopper subfamily Eurymelinae was conducted based on DNA sequence data from three nuclear and two mitochondrial genes in addition to 86 discrete morphological characters. The analysis included 89 species representing 61 genera from all major biogeographic regions including six species from outgroups, Megophthalminae and Ulopinae. Trees resulting from partitioned Bayesian and maximum likelihood analyses of the combined data were well resolved and largely congruent, differing mainly in the relationships among the earliest diverging lineages. The results are consistent with an expanded concept of Eurymelinae, including tribes Austroagalloidini and Macropsini. Six monophyletic groups are recognized as new tribes, Balocerini, Chiasmodolini, Chileanoscopini, Idioceroidini, Kopamerrini and Nesocerini, tribe n. , and the previously recognized tribes Eurymelini, Idiocerini and Megipocerini are redefined. A new synonym, Busonini Zhang & Li, 2015 syn.n. is proposed here for Megipocerini Isaev, 1988. Molecular divergence time estimates were calibrated using two fossil taxa and suggested that the earliest divergences occurred in the Lower Cretaceous and that most major lineages of this group arose during the Cretaceous. Reconstruction of ancestral areas revealed considerable continental-scale biogeographical structure. The place of origin of Eurymelinae is equivocal but major lineages arose in the Neotropical, Australian and Afrotropical regions. A key to tribes and a checklist of genera showing current tribal placements are provided.     

巴基斯坦角顶叶蝉亚科两新纪录属（半翅目：叶蝉科）

报道巴基斯坦角顶叶蝉亚科1新纪录属Changwhania Kwon和1新纪录亚属Limotettix(Scleroracus)Van Duzee,并记述2新纪录种Changwhania ceylonensis Baker,1925 n.rec.和Limotettix(Scleroracus)cacheolus n.rec.。     

Phylogeny of the leafhopper subfamily Evacanthinae with a review of Neotropical species and notes on related groups (Hemiptera: Membracoidea: Cicadellidae)

Abstract. The phylogenetic analysis of ninety‐two adult morphological characters supports the treatment of Nirvaninae as a junior synonym of Evacanthinae and the redefinition of Evacanthinae to include the tribes Nirvanini, Balbillini, Evacanthini and Pagaroniini. The analysis indicates that Nirvaninae, as previously defined, is polyphyletic. A key to tribes and Neotropical genera is provided and diagnostic features for these taxa are reviewed. Jassoqualus Kramer, Neonirvana Oman, Synogonia Melichar (=Jassopronus Nielson & Godoy, syn.n.) and Tahura Melichar are retained within Nirvanini and two new Neotropical genera of this tribe are described and illustrated: Antillonirvana, gen.n., based on two new species from the Dominican Republic and one from Cuba; and Chibchanirvana, gen.n., based on two new species from Colombia. Pentoffia Kramer and Evanirvana Hill are treated as incertae sedis within Evacanthinae. Six new species of Pentoffia, a new species of Synogonia, a new species of Jassoqualus, two new species of Neonirvana and eleven new species of Tahura, all from South America, are also described and illustrated. The following taxa included previously in Nirvaninae are excluded from Evacanthinae, sensu lato: Tungurahuala Kramer to Cicadellinae; Columbonirvana Linnavuori to Typhlocybinae; Macroceratogoniini to Coelidiinae; Occinirvanini Evans to Deltocephalinae. Omaranus Distant, placed previously in Occinirvanini, is transferred to Doraturini (Deltocephalinae).     

Review of the South American leafhopper genus Parandanus (Hemiptera,Cicadellidae, Deltocephalinae)

The South American leafhopper genus Parandanus Linnavuori & DeLong (Deltocephalinae: Deltocephalini) is reviewed and four of its six species are illustrated and male genital characters are provided. Three new species from Peru, Parandanus longistylus Duan, sp. n., Parandanus nigricephalus Duan, sp. n. and Parandanus paracruciatus Duan, sp. n. are described. A key to species is also provided.     

Phylogeny and biogeography of Neotropical spittlebugs (Hemiptera: Cercopidae: Ischnorhininae): revised tribal classification based on morphological data

The spittlebug family Cercopidae is currently divided into two subfamilies: the paraphyletic Old World Cercopinae and the monophyletic New World Ischnorhininae. The most recent classification scheme proposed by Fennah in 1968 divided the New World Cercopidae into four tribes: Tomaspidini, Ischnorhinini, Hyboscartini and Neaenini. Herein we present a phylogenetic analysis of Ischnorhininae using 108 morphological characters and including 53 of the 59 recognized genera, to evaluate the tribal‐level classification and understand the processes underlying the current distributional patterns of these genera. We found significant support for the monophyly of many Neotropical genera, but Fennah's tribal classification is revised because tribes Neaenini, Ischnorhinini and Tomaspidini were recovered as polyphyletic. Hyboscartini was synonymized with Tomaspidini. A taxonomic key to tribes and genera of Neotropical spittlebugs is provided based mostly on recovered apomorphies. The biogeographical analysis suggests a Neotropical origin of ischnorhinines, more specifically in northwestern South America. This was possibly coincident spatially and temporally with the origin of grasses, with ancestral range expansions southward to the Amazonian and Paraná regions, and posterior vicariant events, possibly related to the expansion of forests in the Chacoan region, the South America diagonal of open formations. Dispersals to the Chacoan region and to the Nearctic region are hypothesized to have occurred only within genera. In the Chacoan region it is associated with more recent events, such as the diversification of C4 grasses and establishment of the savannas and seasonally dry forests.     

A taxonomic study on the leafhopper genus Scaphoidella vilbaste (Hemiptera: Cicadellidae: Deltocephalinae) from China

All six species of the leafhopper genus Scaphoidella Vilbaste from China are reviewed. Among them, two new species, S. undosa sp. nov. and S. acaudata sp. nov., are described; two new combinations, Scaphoide. unihamata (Li et Kuoh) comb. nov. and Scaphoidella wideaedeaga (Wang et Li) comb. nov. are proposed; and two new synonyms are revealed, Scaphoideus inermis Cai et He (2001) syn. nov. as a junior synonym of Scaphoidella unihamata (Li et Kuoh 1993) and Scaphoideus multipunctus Li et Dai (2004) syn. nov. as a junior synonym of Scaphoidella stenopaea Anufriev (1977). A key is given to separate all six Chinese species, and illustrations of genitalia are provided.     

A new deltocephaline leafhopper genus,Melanetettix, from Melanesia (Hemiptera: Cicadellidae: Deltocephalinae)

Melanetettix gen. nov. is described from Melanesia for 22 new species of leafhoppers. The new species fall into four groups. New species described are M. bicolor, M. bifidus, M. capitatus, M. curvatus, M. delmoi, M. elongatus, M. maai, M. marginatus, M. ocellatus, M. pallidus, M. rhamphodes, M. roseus, M. semeraroae, M. sinuatus, M. aculeus, M. bifurcatus, M. truncatus, M. clavatus, M. ensiferus, M. nielsoni, M. rotundatus and M. alatus. An identification key to the new species is provided along with discussion of the affinities of Melanetettix, which is close to Scaphoideus Uhler.     

基于16S rDNA和28S rDNA D2基因序列与形态特征联合分析的中国角顶叶蝉亚科系统发育研究（半翅目： 叶蝉科）

首次在国内利用28S rDNA D2区段和16S rDNA基因序列,结合50个形态特征对角顶叶蝉亚科(Deltocephalinae)［半翅目（Hemiptera）: 叶蝉科（Cicadellidae）］19个属进行系统发育分析研究。从无水乙醇浸泡保存的标本中提取基因组DNA并扩增了19个内群和1种外群Typhlocybinae［半翅目(Hemiptera): 叶蝉科(Cicadellidae)］种类的28S rDNA D2基因片段并测序,同时扩增了16S rDNA基因片段并测序11条,采用了GenBank中1个种类的16S rDNA同源序列。采用PAUP*4.0和MrBayes3.0两个分析软件和3种建树方法,利用同源28S D2 rDNA和16S rDNA两个基因序列与形态特征结合进行系统发育分析研究。分析结果表明,二叉叶蝉族Macrostelini是一个单系,并在角顶叶蝉亚科的系统发育中处于基部的位置,是内群中最原始的族;角顶叶蝉族Deltocephalini中除了纹翅叶蝉属Nakaharanus,其余各属构成单系;殃叶蝉族Euscelini内属的归属比较混乱,可能是一个并系群,属间差异有待进一步研究。隆额叶蝉族Paralimnini与顶带叶蝉族Athysanini是姐妹群。带叶蝉属Scaphoideus与纹翅叶蝉属Nakaharanus是姐妹群,二者与木叶蝉属Phlogotettix的关系最近,三者构成一个单系,建议将三者归为带叶蝉族Scaphoideini。研究结果还表明,小眼叶蝉族Xestocephalini和Balcluthini的系统发育位置不明,有待进一步研究。     

Biology and natural enemies of the leafhopper Balclutha incisa (Matsumura) (Hemiptera: Cicadellidae: Deltocephalinae) in south-western Australia

Abstract  Balclutha incisa is a widely distributed leafhopper common on grasses. In Perth, Western Australia, crab grass ( Digitaria sanguinalis ) and veldt grass ( Ehrharta longiflora ) were identified as host plants. Mating and subsequent oviposition occurred in late summer to early winter (August) on the panicles and stems of crab grass. We describe the biology and phenology of B. incisa within a restricted habitat over a 2-year period and we comment on oviposition sites, eggs and emergence of B. incisa . We describe the behaviour of two main predators, the Pacific damsel bug, Nabis kinbergii Reuter and muscid fly belonging to the genus Coenosia Meigen , particularly in regard to their response to visual and auditory signals produced by the leafhoppers.     

Phylogeny of the tribe Empoascini (Hemiptera: Cicadellidae: Typhlocybinae) based on morphological characteristics,with reclassification of the Empoasca generic group

A morphology‐based phylogenetic analysis of the tribe Empoascini (Hemiptera: Cicadellidae: Typhlocybinae) is presented for 58 of 83 formerly recognized genera based on 99 morphological characters of adults. The results support excluding the New World Beamerana generic group from Empoascini. The remaining genera of Empoascini were recovered as a monophyletic sister group of Dikraneurini. Previously recognized tribes Jorumini and Helionini are derived from within Empoascini and are considered synonyms of the latter tribe. Three previously recognized informal generic groups, the Empoasca group, Alebroides group and Usharia group were paraphyletic but the Ficiana group was recovered as monophyletic based on five synapomorphies. Genera previously placed in the Alebroides group represent at least six independent lineages, indicating that the hind wing character separating this group from the Empoasca group (CuA and MP veins free) is highly homoplasious. Empoasca (sensu lato) is also paraphyletic. Thus, twelve previously recognized subgenera of Empoasca are elevated to genus status and five species groups of Empoasca from the New World are recognized as separate new genera. Sikkimasca Dworakowska, 1993 is treated as synonym of Marolda Dworakowska, 1977 based on the phylogeny. Biogeographic analysis suggests that Empoascini most likely first evolved in the Oriental region and spread to other biogeographic realms more recently by multiple independent invasions.     

Review of the oriental leafhopper genus Lampridius (Hemiptera: Cicadellidae: Deltocephalinae), with description of a related new genus

The leafhopper genus Lampridius Distant, 1918 (type species: L. spectabilis Distant, 1918 ) is redescribed, and a second species, L. cuspidatus sp. nov. (Thailand: Loei), is described. A related new genus Paralampridius gen. nov. and four new species are described: P. mimicus sp. nov. (China: Guangdong, Hainan), P. rotundatus sp. nov. (Thailand: Loei), P. sinuatus sp. nov. (Thailand: Loei, Suphanburi, Petchaburi), and P. truncatus sp. nov. (Thailand: Chaiyaphum, Ubon Ratchathani). Both genera are tentatively included in the tribe Opsiini, although only Lampridius has paired aedeagal shafts with separate gonopores. All six species are illustrated, and a key is provided for their identification. Problems with the current tribal classification are discussed in light of the present discovery of closely related species that show variation in characters previously assumed to be stable at the tribal level in the subfamily Deltocephalinae.     

亚草叶蝉属分类订正（半翅目：叶蝉科：角顶叶蝉亚科：隆脊叶蝉族）

对亚草叶蝉属进行系统研究并首次报道其在中国分布,共记述5种：E. acarifer (Lethierry),E. hilaris (Melichar),E. magnus Mitjaev,E. medius (Mulsant & Rey) 和E. pratensis Emeljanov,其中E. hilaris (Melichar) 和E. medius (Mulsant & Rey) 为中国新记录种。文中提供了分种检索表和雄性外生殖器特征图,并首次记述和提供雌性生殖器特征。     

Nomenclatural changes in the grass-feeding Mexican leafhopper genus Cocrassana Blocker & Larsen (Hemiptera: Cicadellidae: Deltocephalinae: Athysanini)

Two nomenclatural changes are proposed in the tribe Athysanini (Cicadellidae: Deltocephalinae). Chlorotettix sexvarus DeLong, 1959, is newly placed in the genus Cocrassana Blocker & Larsen and is a senior synonym of the type species of Cocrassana, C. riepmai Blocker & Larsen, 1991, syn. nov. A revised diagnosis and illustration of the species are provided.     

A review of the morphologically diverse leafhopper subfamily Stegelytrinae (Hemiptera: Cicadellidae) with description of new taxa

Abstract. The morphologically diverse leafhopper subfamily Stegelytrinae is reviewed and its biology and biogeography are discussed as well as its classification, monophyly and relationship to Deltocephalinae s.l. In addition, the first stegelytrine is recorded east of Wallace's and Weber's lines, extending the distribution of the subfamily outside the Oriental and Palaearctic regions. A checklist and key to the genera of the subfamily are given. The following new genera and species are described: Honguchia rubrofascia Wei & Webb, gen.n & sp.n. from Sabah; H. superba Wei & Webb, sp.n. from Sumatra; Yaontogonia flavopicta Wei & Webb, gen.n & sp.n. from Brunei and Sabah; Y. darjeelingensis Wei & Webb, sp. n. from West Bengal; Sychentia breviata Wei & Webb, gen.n & sp.n. from Malaysia; Louangana stellata Wei & Webb, gen.n & sp.n. from Laos; Shangonia sarawakensis Wei & Webb, gen.n & sp.n. from Sarawak; Quiontugia fuscomaculata Wei & Zhang, gen.n. & sp.n. from Hainan; and Neophansia wallacei Wei & Webb, gen.n & sp.n. from New Guinea.     

Copulation, the dynamics of sperm transfer and female refractoriness in the leafhopper Balclutha incisa (Hemiptera: Cicadellidae: Deltocephalinae)

Abstract.  Mature sperm of the leafhopper Balclutha incisa (Matsumara) (Cicadellidae: Auchenorrhyncha: Hemiptera) are stored as a series of sperm bundles within seminal vesicles prior to ejaculation. During transfer, sperm are pumped from the vesicles into the ejaculatory duct to the complex aedeagus. Sperm transfer is marked by a c . 30-fold expansion of the spermatheca to accommodate both sperm and seminal fluid. Sperm number increases exponentially with male age, reaching a maximum of 700 000 after 14 days, while the number of sperm available on days 2–5 is between 70 000 and 100 000. During mating, maximum sperm transfer occurs after 7 min and mating is complete after about 10 min. Ejaculate size, defined by both sperm and associated accessory gland fluid, is influenced by male mating status and the interval since the previous mating. There is a positive correlation between duration of copulation and both ejaculate and the time to subsequent mating. Sperm are more likely to be retained in the testes during mating by males of 2–5 days post-emergence than older males. The number of sperm received by the female can be manipulated experimentally by mating males once (medium ejaculate) or twice (small ejaculate) immediately after their first mating. Females that receive small ejaculates from sperm-depleted males have a far shorter refractory period than females receiving medium to large ejaculates. Both ejaculate size and the time after males have mated influence the female post-mating refractory period as measured by the female's responsiveness to male sexual signalling.