Endogena , two new genera of the Alebroides genus group ( Hemiptera : Cicadellidae : Typhlocybinae ) from the Oriental Region with notes on the phylogeny of Empoascini

Two new microleafhopper genera of Empoascini within the subfamily Typhlocybinae (Hemiptera: Cicadellidae), Condensella Xu, Dietrich & Qin gen. n., based on the type species C. fi lamenta Xu, Dietrich & Qin sp. n., and Endogena Xu, Dietrich & Qin gen. n., based on the type species E. fl ava Xu, Dietrich & Qin sp. n., are described from southern China and Thailand. Male habitus photos and illustrations of male genitalia of the two new species are provided. Comparative notes on related genera are provided. Phylogenetic relationships and the status of genus groups within the tribe are also discussed. ZooBank Article LSID: C08BD6DF-8E3A-474F-8402-51EDC1EEAB53 * Corresponding authors, e-mails: chdietri@illinois.edu, qindaozh0426@aliyun.com INTRODUCTION Empoascini is a tribe of the microleafhopper subfamily Typhlocybinae. It comprises 89 previously described genera and more than 1000 species worldwide (Xu et al., 2017), including widespread, polyphagous agricultural pests such as the Potato Leafhopper, Empoasca fabae (Harris) (Chasen et al., 2014) and Cotton Leafhopper Amrasca biguttula biguttula (Ishida) (Saeed et al., 2015), as well as pests that appear to be monophagous on one particular crop, such as the Tea Green Leafhopper, Empoasca onukii Matsuda (Qin et al., 2015). Knowledge of the world fauna of Empoascini remains incomplete and new genera and species continue to be discovered at a rapid pace. Members of this tribe have been assigned to three informal generic groups (Qin et al., 2011; Xu et al., 2015, 2016), one of which, the Alebroides group, is characterized by the vein CuA in the hind wing branched preapically. The Alebroides group comprises 158 species in 27 genera so far, widely distributed in the Old World (Xu et al., 2017). The Oriental Region harbors the richest biodiversity of the Alebroides group, with 23 genera reported so far, including 16 genera known in the Chinese fauna. Despite recently published papers treating the Alebroides genus Eur. J. Entomol. 114: 462–469, 2017 doi: 10.14411/eje.2017.059


INTRODUCTION
Empoascini is a tribe of the microleafhopper subfamily Typhlocybinae.It comprises 89 previously described genera and more than 1000 species worldwide (Xu et al., 2017), including widespread, polyphagous agricultural pests such as the Potato Leafhopper, Empoasca fabae (Harris) (Chasen et al., 2014) and Cotton Leafhopper Amrasca biguttula biguttula (Ishida) (Saeed et al., 2015), as well as pests that appear to be monophagous on one particular crop, such as the Tea Green Leafhopper, Empoasca onukii Matsuda (Qin et al., 2015).Knowledge of the world fauna of Empoascini remains incomplete and new genera and species continue to be discovered at a rapid pace.Members of this tribe have been assigned to three informal generic groups (Qin et al., 2011;Xu et al., 2015Xu et al., , 2016)), one of which, the Alebroides group, is characterized by the vein CuA in the hind wing branched preapically.The Alebroides group comprises 158 species in 27 genera so far, widely distributed in the Old World (Xu et al., 2017).
The Oriental Region harbors the richest biodiversity of the Alebroides group, with 23 genera reported so far, including 16 genera known in the Chinese fauna.Despite recently published papers treating the Alebroides genus 2).Crown short and broad, rounded anteriorly, anterior and posterior margins nearly parallel, middle length shorter than half width between eyes (Figs 1, 3); coronal suture long, passing to face and terminating at level of antennal bases (Figs 1,3,4).Ocelli distinct, on face, well separated from eyes (Fig. 4).Face broad, lateral frontal sutures extended ventromesad of ocelli, anteclypeus slightly convex, not expanded (Fig .4).Pronotum relatively short (Figs 1, 3).Forewing narrow, rounded apically, apical cells occupying nearly one-third of total length, c and r cells nearly equal in width, both narrower than m and cua cells; vein RP arising from r cell and MP' and MP''+CuA' from m cell, MP' and MP''+CuA' almost parallel throughout their length (Fig. 9).Hind wing with CuA branched, branching point distad of coalescence of CuA with MP'' (Fig. 10).Fro nt femur seta AM1 stout, well developed, situated near ventral margin; intercalary row with one large basal seta tion Suite (LAS) V3.7 and edited using Adobe Photoshop CS 8.0 (Adobe Systems).Body measurements (in mm) are from apex of vertex to tip of forewing.
Etymology.The generic name is derived from the Latin word "condensus" (dense), referring to the densely grouped setal group A of the subgenital plate.Gender: feminine.
Remarks.Among previously described genera of the Alebroides group, this new genus is most similar to Apheliona Kirkaldy, 1907 andZnana Dworakowska, 1994a.It differs from both in having vein MP' in the forewing aris- ing from the m cell (Fig. 9), in having a caudoventral process on the pygofer in addition to a ventral appendage (Figs 5,11,12) and having a small central lobe of the connective (Fig. 17).This new genus is also similar to Schizandrasca Anufriev, 1972 in the slender and sinuate aedeagus, but differs from the latter in having a long coronal suture (Figs 3,4), the dorsal habitus of head rounded anteriorly, the crown broader than the pronotum (Figs 1, 3), a ventral pygofer appendage (Figs 5,11) and setal group C of the subgenital plate uniseriate near the base (Figs 11,19).
Etymology.The specifi c epithet is derived from Latin adjective "fi lamentus" (fi laments), which refers to the shape of the aedeagus.
Distribution.China (Guangxi, Yunnan); Thailand (Kanchanaburi).Description.Head including eyes almost as wide as pronotum in dorsal aspect (Figs 22,24).Crown short and narrow, rounded anteriorly, anterior and posterior margins nearly parallel, middle length shorter than width between eyes, coronal suture distinct (Figs 22,24); posteromedial corner of eye separated from posterior margin of head (Fig. 24).Head in profi le only slightly upturned relative to pronotum, transition of crown to face rounded (Fig. 23).Ocelli distinct (Figs 22,24,25), on crown margin well separated from eyes.Face broad and distinctly convex in profi le, lateral frontal suture extended ventromesad of ocelli, anteclypeus strongly infl ated in male (Fig. 25).Pronotum large (Figs 22,24).Forewing narrow, rounded apically, apical cells occupying more than one-fourth total length, c, r and m cells nearly equal in width, all narrower than cua cell; veins RP and MP' arising from r cell and MP''+CuA' from m cell (Fig. 29).Hind wing with CuA branched, branching point distad of coalescence of CuA with MP'' (Fig. 30).Front femur seta AM1 stout, situated near ventral margin; intercalary row with one large basal seta and seven to eight smaller setae near tip of femur.Hind femur with macrosetal formula 2+1+1.Hind-tibia row AV with ~13 preapical macrosetae.
Etymology.The generic name is derived from the Latin word "endogenus" (internal), referring to the small mesoapical spine near caudo-dorsal angle of the pygofer lobe.Gender: feminine.
Etymology.The specifi c epithet is derived from the Latin adjective "fl avus" (yellow), which refers to the body color of the new species.

DISCUSSION
Although Empoascini is now widely accepted as a valid tribe of subfamily Typhlocybinae, its phylogenetic position has still not been adequately elucidated.The relationship between Empoascini and other tribes was fi rst discussed by Mahmood & Ahmed (1968), who proposed splitting Typhlocybini (sensu Young 1952) into two tribes (Empoascini and Typhlocybini) but suggested that Empoascini is "closely related to Typhlocybini."Later, Zhang (1990) suggested that Empoascini has a closer relationship with Dikraneurini based on the presence of a well-developed submarginal vein in the hind wing of both groups.
Like other typhlocybine tribes, Empoascini has traditionally been characterized by the wing venation.Species of this tribe lack an appendix on the forewing, and have the hind wing submarginal vein present apically between the jugal lobe and MP' or RP+MP' but not extended around the wing apex along the costal margin.Other diagnostic traits include the presence of well developed ocelli, the characteristically curved distal segment of forewing vein MP''+CuA', presence of a longitudinal row or band of numerous macrosetae on the male subgenital plate, and absence of a well-developed preapical lobe on the paramere.These latter four features are shared with Alebrini and are presumably symplesiomorphic.The presence of welldeveloped paired processes at the base of the anal tube is possibly a synapomorphy of the tribe, although it is present in a few genera in other tribes [e.g., Aphanalebra McAtee, 1926 (Alebrini), Tautoneura Anufriev, 1969 (Erythroneurini)].The venation of the hind wing may also be synapomorphic, although a similar pattern was recently reported in two Oriental genera of Dikraneurini (Dietrich, 2013), providing additional support for Zhang's (1990) hypothesis of a relationship between Dikraneurini and Empoascini.
Three informal generic groups have been explicitly recognized within Empoascini.One of them is the Alebroides group characterized by a branched vein CuA in the hind wing and including 27 genera, widely distributed in the Oriental, Palaearctic, Afrotropical and Australian Regions but absent in the New World (Xu et al., 2017).The second is the Ficiana group comprising seven genera (Ficiana Ghauri, 1963;Ishiharella Dworakowska, 1970b;Dialecticopteryx Kirkaldy, 1907;Mahmoodia Dwora kowska, 1970a;Nimabanana Dworakowska, 1994b;Kotwaria Dworakowska, 1984 andDaluana Ram akrishnan, 1982) defi ned by having hind wing vein CuA unbranched, the male subgenital plates fused, the ventral pygofer appendage lacking and three apical veins in the fore wing arising from cell m (Xu et al., 2015).Species in the Ficiana group are confi ned to the Oriental, Palaearctic and Australian Regions.In addition to these two groups, our recent work suggests that a third genus group (the Usharia group) is also recognizable and includes another seven genera in the Empoasca-complex (hind wing CuA unbranched): Baguoidea Mahmood, 1967;Dayus Mahmood, 1967;Goifa Dworakowska, 1977;Homa Distant, 1908;Ifugoa Dworakowska & Pawar, 1974;Treufalka Qin &Zhang, 2008 andUsharia Dworakowska, 1977.Species in the Usharia group all have the connective fused with the aedeagus and are distributed in the Oriental and Australian Regions.Among the genus groups mentioned above, it seems reasonable to hypothesize that the Ficiana and Usharia groups are monophyletic, given the unique features of their male genitalia.The Alebroides group lacks such features and its monophyly is therefore doubtful.The remaining genera of Empoascini, including the type genus Empoasca Walsh, have hind wing vein CuA unbranched, which is presumably a derived trait within the tribe, but it is doubtful that these genera form a monophyletic group because they are apparently united only by the absence of the diagnostic traits present in other generic groups.A formal phylogenetic analysis will be needed to elucidate the status and relationships of all groups currently recognized within Empoascini and the relationship of this tribe to other Typhlocybinae.