South American leafhoppers of the tribe Typhlocybini (Hemiptera: Cicadellidae: Typhlocybinae)

The fauna of Typhlocybini (sensu stricto, excluding Empoascini) endemic to South America is reviewed and comprises seven closely related genera, five described herein as new, and 55 species, 52 here described as new. The genera and species are described and keys and illustrations are provided to aid in their identification. Columbonirvana Linnavuori comprises 17 species, 16 new. Eualebra Baker comprises 19 species, 17 new. Euzyginella gen. nov., comprises four new species. Neozyginella gen. nov., comprises six new species. Pseudhadina gen. nov., comprises one new species. Pseudozyginella gen. nov., comprises three new species. Tahurella gen. nov., comprises five new species. One new synonym is recognized: Eualebra smithii Baker, 1899 equals Dikraneura (Hyloidea) reticulata Osborn, 1928, syn. nov. One previously described species placed in Eualebra belongs in tribe Dikraneurini; thus, a new combination is proposed: Alconeura (Hyloidea) rubra (Van Duzee), comb. nov. Most of the specimens available for this study were from Malaise trap and canopy fogging samples obtained at very few localities in Bolivia, Brazil, Colombia, Ecuador, and Peru, suggesting that further sampling in South America, particularly in the Amazonian rainforest and eastern foothills of the Andes Mountains, will reveal large numbers of additional species.

The leafhopper subfamily Typhlocybinae comprises ~6,000 described species of mostly small, delicate leafhoppers that feed preferentially on the contents of leaf parenchyma cells of their host plants.Based on described species, this is currently the second largest leafhopper subfamily (after Deltocephalinae, ZAHNISER & DIETRICH 2010), but recent sampling of tropical faunas indicates that the extant, but mostly undescribed, fauna of Typhlocybinae is far larger than that of any other leafhopper subfamily (DIETRICH & WALLNER 2002).The Neotropical fauna, in particular, is highly diverse but appears to remain largely undocumented (DIETRICH & DMITRIEV 2006, 2008).
Study of canopy fogging samples from lowland rainforests in eastern Peru and Ecuador (DIETRICH & WALLNER 2002) indicates the presence of an enormously diverse typhlocybine fauna.Composition of this fauna contrasts strikingly with that of the somewhat better known, but still poorly documented faunas of the Old World tropics.Tribes Dikraneurini, Empoascini (= Jorumini) and Alebrini, in decreasing order of species richness, are well represented in the samples from Amazonia; but Erythroneurini and Typhlocybini, so abundant in north temperate forests and in the Old World tropics, are relatively rare.
Malaise trap samples from the eastern Andean foothills above 500 m elevation differ from those obtained from the canopy of lowland rainforest (unpublished data) in that they include larger numbers of Typhlocybini (sensu stricto).These species belong to a group of apparently closely related, endemic South American genera that includes Eualebra Baker.The group also includes Columbonirvana Linnavuori, a genus previously placed in Nirvaninae (LINNAVUORI 1959), but transferred to Typhlocybinae by DIETRICH (2004) based on phylogenetic analysis of morphological characters.These two genera include a total of three previously described species (plus one new synonym, designated below).Recent sampling indicates that the South American fauna of Typhlocybini is much richer in both species and genera.In this paper, I review the endemic South American genera and species of Typhlocybini and discuss their possible relationships to the fauna of this tribe in other parts of the world.Considering that the 52 new species of Typhlocybini described herein were collected from only eight localities in Bolivia, Brazil, Colombia, Ecuador and Peru, further sampling in South America, especially in cloud forests of the eastern Andean foothills, will likely reveal the presence of a much more speciose assemblage belonging to this tribe.

MATERIAL AND METHODS
Morphological terminology follows that of YOUNG (1952) as modified by DIETRICH (2005) except wing veins are labeled according to the simplified system illustrated in figures 3i, j.Drawings of genitalia were made by tracing over photographs taken using a digital camera mounted on a compound microscope.In drawings of the male genital capsule, the entire anal complex (segments X-XI) is shown in lateral view but only the well sclerotized and pigmented parts of segment X are shown South American leafhoppers of the tribe Typhlocybini ZOOLOGIA 30 (5): 519-568, October, 2013 Typhlocybini, but varies intraspecifically, and occasionally within the same individual, for features traditionally diagnostic of tribes Typhlocybini,Eupterygini,and Zyginellini (Figs 82 and 83).Hind wing veins RP and MA may be separate and connected by a crossvein (as in Eupterygini) or confluent (as in Typhlocybini, s.s.).Also, the hind wing submarginal vein may be strongly curved and colinear with CuA preapically (as in "Zyginellini") or straight and connected to CuA by a crossvein (as in Typhlocybini, s.s.).Presence of such variation within a single species provides support for AHMED's (1983) treatment of Eupterygini and Zyginellini as junior synonyms of Typhlocybini.
The presence of a submarginal vein at the hind wing apex and relatively numerous macrosetae on the male subgenital plates (as in Alebrini and Empoascini) suggests that Eualebra and related South American genera represent a plesiomorphic lineage of Typhlocybini not represented in the Old World.Other Typhlocybini, including genera and species endemic to the Old World and North America, lack a submarginal vein in the hind wing and almost always have two or fewer (usually 0-1) macrosetae on the male subgenital plate.
As in some Old World Typhlocybini, the lower part of the face in most South American genera is sexually dimorphic: females have the clypellus and lorum flattened and well delimited, but males have the lorum fused to the clypellus and both structures are markedly inflated (Fig. 59).
Based on the morphological similarities mentioned above, the South American fauna of Typhlocybini appear to be related to some Old World genera previously included in "Zyginellini" (sensu DWORAKOWSKA 1979) as well as to the "Eupteryx-complex" (sensu DWORAKOWSKA 1969) of genera that are most diverse in the Indomalayan region.These relationships need to be elucidated by phylogenetic analysis.
The endemic South American fauna of Typhlocybini comprises seven genera: Eualebra, placed by YOUNG (1952) in Typhlocybini based on the branched vannal vein and absence of a submarginal vein along the costal margin in the hind wing; Columbonirvana Linnavuori, previously placed in Nirvaninae presumably based on its depressed form and produced head, but transferred to Typhlocybinae by DIETRICH (2004); and five new genera described below, Euzyginella, Neozyginella, Pseudhadina, Pseudozyginella, and Tahurella.
Two Palearctic species of Typhlocybini known to be established in Argentina and Chile, Edwardsiana froggatti (Baker) and Ribautiana tenerrima (Herrich-Schaffer), have been well described elsewhere (CHRISTENSEN 1942, CHRISTIAN 1953) and are not treated here.
Remarks.The type species of this genus was described based on a single female from Colombia (illustrated by LINNAVUORI 1959 andDIETRICH 2004).No additional specimens matching the type specimen of C. aurea Linnavuori in structure or color pattern have been found, but numerous congeneric male specimens representing 16 additional species were collected by Malaise trap in cloud forests in the Andes Mountains of Peru.No additional female representatives of this ge-nus have yet been found.DIETRICH (2004) transferred Columbonirvana from Evacanthinae (= Nirvaninae) to Typhlocybinae but considered its tribal placement uncertain.The genus is here placed in the tribe Typhlocybini based on the concept of the tribe embodied in the above diagnosis, which is narrower than that of YOUNG (1952) but broader than that of OMAN et al. (1990).It is similar to Tahurella gen.nov.but differs in the smaller size, darker overall coloration, lack of transverse black band on the anterior margin of the head, and rounded forewing apex.
Columbonirvana ameliae sp.nov.Figs 2,[91][92][93][94]152 Description.Length of male 4.2 mm.Dorsum (Fig. 2) dark reddish brown with symmetrical pale yellow markings; anterior margin of head with pair of thin red transverse bands between eyes; crown yellow bordered anteriorly with brown; pronotum yellow medially with anterior and lateral margins brown and with brown transverse submarginal band posteriorly; mesonotum yellow medially, anterolateral triangles and apex of scutellum brown; forewing clavus with small white transcommisural spot bordered posteriorly with larger crimson area, veins orange bordered with brown in distal half; venter of thorax pale yellow except mesosternum orange brown.Head as wide as pronotum, crown nearly twice as long medially as next to eye.

Columbonirvana aurea
Columbonirvana craigi sp.nov.Figs 4,[95][96][97][98]153 Description.Length of male 4.0 mm.Dorsum (Fig. 4) brown with extensive symmetrical pale markings; anterior margin of head with pair of thin red transverse bands between eyes; crown brown medially with broad white anterior and posterior margins; pronotum with white median crescent, posterior margin broadly white; mesonotum and scutellum white except midline anterad of scutellar suture and basolateral triangles brown; forewing with extensive large symmetrical white markings.Head subequal to pronotum in width, crown with median length much less than twice length next to eye.
Etymology.This species is named in honor of my brother, Craig B. Dietrich.
Columbonirvana davidi sp.nov.Figs 5,[99][100][101][102]154 Description.Length of male 3.4 mm.Dorsum (Fig. 5) dark brown with large symmetrical yellow markings.Anterior margin of head with two narrow transverse red bands; crown with posterior margin infused with yellow medially.Pronotum with large transverse yellow crescent.Mesonotum brown with large yellow median spot.Forewing clavus with large symmetrical transcommisural marking; veins mostly orange; anteapical and apical cells each with medial area hyaline; brachial and discal cell each with hyaline area near apex.Head as wide as pronotum, crown twice as long medially as next to eye.
Male 2S apodemes (Fig. 155) broad, parallel sided, extended nearly to posterior margin of sternite IV.Pygofer emargination quadrate (Fig. 104), tergite nearly 1/2 length of lobe; dorsal sclerotized area of lobe with digitiform distal section bent ventrad at slightly less than 90° angle, extended slightly beyond margin; ventral appendage gradually curved dorsad, apex curved slightly mesad (Figs 103 and 104 Etymology.This species is named in honor of my father, Edgar H. Dietrich.Description.Length of male 3.8 mm.Dorsum (Fig. 7) brown, without distinct pale markings; head anterior margin with pair of thin red transverse bands between eyes; forewing veins yellow distally, apical and anteapical cells hyaline in middle, discal and brachial cell each with hyaline spot near apex.Head subequal to pronotum in width, crown less than twice as long medially as next to eyes.
Columbonirvana tuberculata sp.nov.Figs 14,[135][136][137][138]163 Description.Length of male 4.3 mm.Dorsum (Fig. 14) mostly brown, venter pale yellow except medial part of frontoclypeus, gena below eye, and mesepisternum brown.Head anterior margin with pair of transverse red bands, crown, pronotum, mesonotum and scutellum brown infused with stramineous; forewing with indistinct stramineous band extended from base of clavus across mesonotum and another larger transcommisural band near midlength, veins mostly orange, cells hyaline medially, costal margin with orange pigment in distal third.Head subequal to pronotum in width, crown only slightly longer medially than next to eye.
Etymology.The species name refers to the small dorsal preapical tooth (tubercle) on the male pygofer lobe.
Type species: E. smithii Baker 1899.Diagnosis.Small to medium sized, strongly depressed, somewhat ovoid leafhoppers  broadest across midlength of resting forewings in dorsal view.Color pale yellow to dark brown with symmetrical white, yellow, orange, brown, and/or black markings dorsally.
Head  narrower than pronotum, depressed, produced, anterior margin parabolic in dorsal view; crown slightly convex, declivous, glabrous, coronal suture restricted to posterior third; face horizontal (Figs 62-63); rostrum not extended beyond front trochanters; lower part of face sexually dimorphic, male anteclypeus inflated and expanded laterad, parallel-sided and weakly convex in female; lorum very narrow; frontoclypeus flat or concave medially; lateral frontal sutures not extended dorsad of antennal pits; antennal ledges curved, nearly vertical; antennae slightly longer than head width; ocelli vestigial, on crown margin approximately midway between eye and midline.Pronotum  with lateral margins long, divergent in dorsal view, distinctly carinate, carina even with posterior corner of eye.Front femur with AM1 enlarged and situated on ventral margin; intercalary row with few fine setae, basal seta larger than others; PV1 well developed; tibia somewhat flattened dorsally, AD and PD without preapical macrosetae.Hind femur macrosetae 2+1+1; tibia row AV with 3-4 macrosetae near apex.Forewing (Figs 71 and 73) with RA slightly to strongly reflexed, RP and MA separate and connected by crossvein or confluent for short distance preapically.Hind wing (Figs 72 and 74) with anterior branch of R absent, veins RP and MA separate, joined by crossvein; MP and CuA separate, joined by oblique m-cu crossvein; submarginal vein extended from apex of RP to jugal lobe.
Male 2S apodemes (Fig. 242-260) well developed, usually joined at base by transverse bridge, variable interspecifically in length and orientation.Pygofer (Figs 167 and 168) slightly to strongly emarginate dorsally; tergite variable interspecifically in length and shape; posterior lobe with or without dorsal process, side not or only weakly produced posteriorly, ventral appendage elongate, curved dorsad, usually areolate distally.Anal tube and appendages highly variable interspecifically, append-age usually well developed, arising from base of anal tube (segment X), articulated to posterodorsal margin of pygofer laterally and to dorsal connective medially, usually with at least one separate projection extended mesad and/or ventrad.Sclerotized dorsal connective present (Fig. 214), usually U-shaped (rarely platelike) and articulated between dorsal apodeme of aedeagus and paired basal processes of anal tube.Valve very short, rectangular, fused to pygofer.Subgenital plate (Fig. 167) depressed, ovoid in ventral view, distal half tapered with 5 or more macrosetae arranged in lateral band, fine setae short and sparse on dorsal surface, apex rounded or subtruncate.Style (Figs 169 and 170) apodeme short, broad; apophysis elongate, slender, apex simple, tapered and slightly hooked, without conspicuous setae.Connective (Fig. 170) U-shaped with pair of posterodorsal lobes articulated to aedeagus.Aedeagus (Figs 169 and 170) with preatrium well developed; dorsal apodeme short, lobelike; shaft short, tubular, with or without processes; gonopore apical.
Remarks.DIETRICH ( 2004) noted that Eualebra and Columbonirvana share a unique hind wing venational pattern different from that of other Typhlocybinae and therefore treated both genera as unplaced to tribe within Typhlocybinae.Based on the revised tribal diagnosis given above, both of these genera are now included in Typhlocybini (also see Remarks under Columbonirvana).Species of Eualebra resemble species of the Old World genus Eurhadina Haupt in their ovoid shape in dorsal view and strongly depressed body form.However Eurhadina differs in having the structure of the lower part of the face similar in males and females, the forewing narrow distally, the male subgenital plates usually with a single subbasal macroseta, and the pygofer lacking an elongate ventral process.Also, like all other known Old World Typhlocybini, Eurhadina lacks a submarginal vein at the apex of the hind wing.Species of Eualebra occur in both lowland Neotropical rainforest and premontane cloud forests.
Eualebra previously included five species.YOUNG (1952) excluded E. notata Baker from Guatemala, noting that the hind wing venation of the female holotype is consistent with that of tribe Dikraneurini, but did not suggest an appropriate generic placement for the species so it remains incertae sedis.Examination of the holotype of Eualebra rubra Van Duzee from Jamaica, deposited in the California Academy of Sciences collection, indicates that it belongs in the subgenus Alconeura (Hyloidea) of tribe Dikraneurini.Therefore, A. (H.) rubra (Van Duzee) is proposed as a new combination.As a result, only three taxa previously included, E. smithii Baker, E. reticulata Osborn, and E. rufoornata (Stål), all described from Brazil, are retained in Eualebra.Seventeen new species, described below, are also included.Based on the species known at present, the genus appears to be restricted to South America.
Eualebra, as presently defined, is somewhat heterogeneous morphologically, comprising species that range in size from 2.7 to 4.7 mm and exhibit a variety of color patterns, shapes, and male genitalia configurations.Given the still poor state of knowledge of the genus, and particularly considering that females are known for only a few species, I have opted not to recognize additional genera or subgenera but, rather, place the known species into five informal species groups that may represent distinct lineages.Recognition of some of these groups as separate genera may be warranted once the fauna becomes better known.In particular, the ovipositors of the few female specimens available for study exhibit considerable variation and may provide genus-level characters.

Eualebra barbarae species group
Diagnosis.Species of this group may be distinguished from other Eualebra in their very small size (<3 mm), relatively elongate crown (Figs 18 and 19), mottled brown coloration, cruciform or trident-shaped pronotal marking, basally quadrate third apical cell of the forewing, platelike dorsal connective, and absence of paired processes on the aedeagus (Fig. 170).
Eualebra barbarae sp.nov.(Figs 18,(167)(168)(169)(170)242) Description.Length of male 2.7 mm. Color (Fig. 18) mostly dark brown, marked with white and reddish orange; crown mostly brown with midline and lateral margins pale, face brown except for broad pale anterior band; pronotum brown with pale midline and symmetrical pattern of pale markings anteriorly; scutellum and mesonotal midline pale; forewing mostly mottled brown, clavus with two transcommisural pale areas outlined with dark brown, corium with large reddish orange submedial spot; veins yellow except near wing margin; venter dark brown, tibiae somewhat paler.Body relatively short and broad.Forewing with RP and MA separate, connected by crossvein; CuA connected to M basad of its fork.Hind tibia row AV with 3 macrosetae.
ZOOLOGIA 30 (5): 519-568, October, 2013 brown, pygofer and medial area of male subgenital plates dark brown.Body relatively broad.Forewing with RP and MA separate, connected by crossvein; CuA connected to M at or basad of its branching point.Hind tibia row AV with 4 macrosetae.
Eualebra jessicae sp.nov.Figs 21,[179][180][181][182]245 Description.Length of male 4.1 mm.Dorsal coloration (Fig. 21) mostly pale brown with symmetrical white, orange, and dark brown markings; venter dull stramineous except mesepisternum dark brown.Crown and pronotum orange, darkening to brown posteriorly with three white longitudinal stripes; mesonotum with anterolateral triangles and pair of small medial spots dark brown, medial area white tinged with orange; scutellum white with transverse brown band; forewing mostly brown with complex color pattern consisting of two transcommisural markings concentrically outlined with dark brown and white, numerous small brown spots and lines, and large orange brochosome field, veins mostly bright orange.Body robust, ovoid.Forewing with RP and MA separate, connected by crossvein; CuA connected to M basad of its branching point.Hind tibia row AV with 3 macrosetae.
Material examined.Holotype male, ECUADOR: Orellana, Res.Etnica Waorani, Transect Ent.Etymology.This species is named in memory of my brother-in-law, Charles Craft.

Eualebra dorisae species group
Diagnosis.Species of this group may be distinguished from other Eualebra in their moderate size, the color pattern  which includes a pair of broad orange or brown longitudinal stripes extended from the crown onto the pronotum, the basally quadrate third apical cell of the forewing, and the elongate aedeagal atrium (Fig. 177).
Eualebra dorisae sp.nov.Figs 20,[175][176][177][178]244 Description.Length of male 4.2 mm.Coloration of dorsum (Fig. 20) mostly brownish orange with paler orange and white markings; crown with midline and lateral margins white, with pair of irregular orange longitudinal submedial stripes; pronotum orange brown with medial and pair of submedial white stripes not extended to posterior margin; mesonotal triangles brown, scutellum white with brown preapical band; forewing mostly brown, mottled distally, with two angulate transcommisural marks on clavus, two smaller white spots on corium adjacent to clavus, brochosome field pale orange; abdominal terga brown; venter of head and thorax white, infused with brown except mesepisternum dark brown; abdominal terga Male abdomen (Fig. 245) with 2S apodemes robust, slightly divergent, extended slightly beyond midlength of sternite IV.Pygofer posterodorsal emargination V-shaped, forming right angle (Fig. 180); dorsal process absent; lobe weakly developed; ventral appendage robust, relatively short, evenly curved posterodorsad.Anal hook (Fig. 179) moderately short and broad, extended ventrad, apex footlike, articulated with broadly Wshaped dorsal connective.Subgenital plate (Fig. 179 Etymology.This species is named in honor of my niece, Jessica C. Bailey. Eualebra patriciae sp.nov.Figs 22,[183][184][185][186]246 Description.Length of male 4.0 mm.Dorsal coloration (Fig. 22) pale yellow with symmetrical white, orange, and brown markings; venter uniformly dull yellow; head with crown margins and midline white, with pair of orange spots along anterior margin, pair of orange S-shaped lines submedially, and pair of large orange rectangles posteriorly; pronotum with paired orange spots anteromedially and at posterolateral angles, large orange marking posteromedially, divided along midline anteromedially; mesonotal triangles orange, pair of small brown spots present anterad of scutellar suture; scutellum with transverse brown band; forewing mostly yellow orange with veins white, clavus with two transcommisural white markings, brachial cell with two irregular white spots, brochosome field white infused with bright yellow, apex with diffuse brown markings.Body robust, ovoid.Forewing with RP and MA separate, connected by crossvein; CuA connected to M at or basad of its branching point.Hind tibia row AV with 3 macrosetae.
Etymology.This species is named in honor of my sisterin-law, Patricia B. Schulz.
Remarks.When dissected, the holotype specimen had a late-instar dryinid (Hymenoptera: Dryinidae) larva in the abdominal cavity.Parasitism by dryinids sometimes causes abnormal development of the abdominal apodemes and genitalia in leafhoppers.The genitalia of the holotype appear to be fully developed, but the relatively small size of the 2S apodemes may be attributable to presence of the parasite.

Eualebra gingerae species group
Diagnosis.Species of this group may be distinguished from other Eualebra by their moderately small size, brown overall coloration (Fig. 23-30), and the presence of large pale transcommisural markings on the forewing and, usually, large pale symmetrical areas on the crown and pronotum (absent in E. jenniferae).
Eualebra gingerae sp.nov.[187][188][189][190]247,367 Description.Length of male 3.3 mm, female 3.0 mm.Coloration of dorsum (Figs 23 and 24) mostly dark brown with symmetrical yellow, orange and white markings; crown and anterior half of pronotum with white medial area bearing 4 lateral extensions, 2 on crown, one on margin of crown and pronotum, and one medially on pronotum, latter with 2 smaller white spots more laterad, posterior margin of pronotum narrowly yellow; mesonotum and base of forewing with inverted pentagonal white mark bearing 2 small black spots on mesonotum and arcuate black band across clavus and scutellum; clavus with slender white transcommisural mark near midlength and larger triangular mark near apex; corium with small yellow spot near base and large oblique orange macula extended from near base of costal margin across brochosome field and into medial cell; veins orange in distal half, bordered with brown; venter of head, thorax and abdomen brown.Body moderately broad.Forewing with RP and MA separate, connected by crossvein; CuA connected to MP. Hind tibia row AV with 3 macrosetae.
Female sternite VII (Fig. 367) with short, acute median posterior lobe and pair of shorter lateral lobes.Second valvulae evenly curved and moderately narrow, right blade with ~13 teeth.
Material examined.Holotype male, BRAZIL: Goias, Uruaçu, Serra da Mesa Survey 14°17.Etymology.This species is named in honor of my sisterin-law, Ginger P. Dietrich.Remarks.The association between male specimens from Goias, Brazil and the female from Orellana, Ecuador, based on the nearly identical color pattern, must be considered tentative.The Ecuadorean specimen is excluded from the type series because it is somewhat smaller than the males from Brazil and differs in some small details of coloration, e.g., the specimen from Ecuador lacks the pair of small black submedial spots on the mesonotum and has the medial pale area of the pronotum relatively large.
Eualebra marilynae sp.nov.Figs 25,[191][192][193][194]248,[363][364][365][366] Description.Length of male 3.0 mm, female 3.0 mm.Coloration of dorsum (Fig. 25) mostly tan to medium brown with symmetrical white and black markings; crown white with pair of black marginal spots at apex and another just anterad of eyes; pronotum with large white anteromedial marking bearing pair of rounded posterolateral lobes; mesonotum with brown M-shaped medial marking; forewing with large white transcommisural marking at base of clavus continuous with scutellum and smaller marking at apex of clavus; veins pale, bordered with brown; venter of head, thorax and abdomen tan, legs dull stramineous, macrosetae black.Body relatively narrow.Forewing with RP and MA separate, connected by crossvein; CuA connected to MP. Hind tibia row AV with 3 macrosetae.
Remarks.The female specimen from Tiputini is very similar to the specimens included in the type series in coloration but has a pair of black spots near the posterior margin of the crown, lacks the bright yellow mark on the costal margin of the forewing, and has the pregenital sternite of the abdomen more strongly produced.It may represent a distinct species or a variant of E. susanae.
Remarks.This species is similar to E. rufoornata (Stål), but differs in its much larger size, brown ventral coloration, more shallowly emarginate tergite IX, and bifid anal hook.
Eualebra peggyae sp.nov.Figs 29,[207][208][209][210]252 Description.Length of male 3.9 mm.Dorsal coloration (Fig. 29) mostly medium brown with symmetrical pale yellow, white, and orange markings; venter dull yellow except dark brown mesepisternum and meso-and metathoracic femora.Head and pronotum with large median white wedge-shaped marking extended beyond midlength of pronotum; posterior margin of pronotum narrowly yellow.Mesonotum brown anteriorly, yellow posteriorly, area anterad of scutellar suture with pair of brown spots; scutellum yellow with brown transverse band.Forewing clavus with two transcommisural diamond-shaped markings, one near base an another smaller one near apex; corium with round spots near midlength in discal cell midlength; costal area with brochosome field orange; veins mostly orange; outer two apical cells hyaline distally.Body robust, ovoid.Forewing with RP and MA separate, connected by crossvein; CuA connected to MP. Hind tibia row AV with 3-4 macrosetae.
Etymology.This species is named in memory of my mother-in-law, Margarete (Peggy) D. Braxton.
Remarks.This species is very similar to E. rufoornata (Stål) in size and coloration, but differs in its short, broad anal hook and elongate basal aedeagal processes.

Eualebra rufoornata (Stål)
Typhlocyba rufoornata Stål 1860: 55.Remarks.Specimens were not examined for the present study but the species was redescribed by Linnavuori (1954), who examined the male holotype from "Brazil" and illustrated the dorsal color pattern and male genitalia.Based on Linnavuori's drawings, the color pattern resembles that of E. peggyae sp.nov.(Fig. 29) in having a wedgelike white marking extended from the crown onto the pronotum, the posterior portion of the mesonotum and scutellum pale, and two diamond-shaped transcommisural markings on the forewing, but the aedeagus of E. rufoornata has a pair of short distal processes that are perpendicular to the shaft rather than the long, sinuate basal processes present in E. peggyae.
Etymology.This species is named in honor of my sisterin-law, Jennifer L. Dietrich.

Eualebra smithii species group
Diagnosis.Species of this group, which includes the type species of the genus, may be distinguished from other Eualebra by the pale overall coloration , color pattern consisting of numerous irregular tiny brown spots and larger creamcolored spots, the absence of paired processes on the aedeagus (Figs 218 and 222), and the broad, straight ovipositor which lacks teeth on the second valvulae (Fig. 374).Description.Length of male 3.3-3.4mm, female 3.8 mm.Color (Figs 31 and 32) mostly pale yellow to white, marked with white, orange and brown; crown and pronotum with few indistinct paired orange maculae; antennal ledge dark brown; pronotum, mesonotum and forewing with sparse brown stipples posteriorly; forewing with brown maculae at midlength of costal margin and in third and fourth apical cells, clavus and distal half of corium stippled with brown, median area of wing with ca.15-20 conspicuous opaque white spots of various sizes; abdominal terga brown medially; venter pale except anteclypeus and femorotibial joint of mesothoracic leg dark brown.Body relatively short and broad.Forewing with RP and MA separate, connected by crossvein; CuA connected to branching point of M. Hind tibia row AV with 3 macrosetae.

Eualebra smithii Baker
Male abdomen (Fig. 254) with 2S apodemes moderately broad, slightly divergent and expanded distally, extended nearly to posterior margin of sternite IV.Pygofer with posterodorsal emargination broadly V-shaped (Fig. 216); posterodorsal appendage absent but margin thickened and heavily sclerotized; ventral appendage arising near midheight of posterior margin, extended ventromesad then curved posterodorsad, apical half areolate (Figs 215 and 216).Anal hook (Fig. 215) with short acute ventral projection and slender posterior branch extended mesad to dorsal connective, slightly broadened distally.Dorsal connective broadly U-shaped.Subgenital plate (Fig. 215) relatively broad at base, slightly broadened to near midlength, then tapered to broadly rounded apex, with ~13-16 macrosetae in lateral band on distal half.Style (Figs 217 and 218) apodeme short and slender, apophysis moderately long and slender, nearly straight, bent slightly mesad at apex, apex surpassing midlength of subgenital plate.Connective (Fig. 218 Remarks.Young (1952) illustrated and described the male genitalia of E. smithii, presumably based on study of the type specimen.Comparisons of specimens from Peru and Bolivia to Young's illustrations of E. smithii and to the female holotype of Dikraneura (Hyloidea) reticulata indicate that these two species are synonyms.The specimen from Bolivia has the connective proportionately larger than those of the specimens from Peru, but the genitalia appear otherwise to be identical.Further collecting is needed to determine whether such variation is indeed intraspecific.
Male abdomen (Fig. 255) with 2S apodemes broad, slightly divergent, extended to midlength of sternite IV.Pygofer with posterodorsal emargination broadly V-shaped (Fig. 220); dorsal appendage absent but dorsal margin with strongly sclerotized ridge; with two paired ventral appendages, one arising near mid-height of posterior margin and another more ventrad, both elongate and slender, extended ventromesad then strongly curved dorsad, ventral pair twice as long as dorsal and contorted distally (Figs 219 and 220).Anal hook (Fig. 219) massive, extended mesad, broad distally, apex broadly and shallowly bifid, with ventral lobe articulated to dorsal connective; segment X without additional lobes or processes (Figs 219 and 220).Dorsal connective very small, platelike.Subgenital plate (Fig. 219 Etymology.This species is named in honor of my mother, Margaret Anne B. Dietrich.

Eualebra michaelorum species group
Diagnosis.Members of this group differ from other Eualebra as follows: size relatively large; crown relatively short with anterior margin in profile relatively thick; forewing veins RP and MA partially confluent preapically, forming petiolate apical cell (Fig. 73); pygofer dorsomedial emargination short and relatively narrow (Fig. 224); pygofer lobe with semicircular unpigmented area posterodorsally.
Etymology.This species is named in memory of my grandfather, Albert E. Dietrich Jr.
Eualebra dorothyae sp.nov.Figs 35,[227][228][229][230]257 Description.Length of male 4.7 mm.Dorsal coloration (Fig. 35) mostly yellow; head with diffuse brown spot at apex, pronotum with small brown medial stripe extended slightly onto mesonotum; forewing with narrow brown maculae at base and apex of brochosome field, and brown macula at base of inner apical cell.Body relatively long and narrow; forewing veins RP and MA partially confluent preapically, CuA connected to M basad of its branch; hind tibial row AV with 3 macrosetae.
Etymology.This species is named in memory of my grandfather, Michael Baka, Sr., and in honor of my uncle, and cousin, Michael Baka, Jr., and III.

Eualebra leoni species group
Diagnosis.Species of this group may be distinguished from other Eualebra by their small size, relatively weakly produced head, brown overall coloration (Figs 37 and 38), white posterior pronotal margin, partially confluent forewing veins RP and MA, and relatively shallow, broad emargination of male abdominal tergite IX (Fig. 236).
Eualebra leoni sp.nov.Figs 37,[235][236][237]298 Description.Length of male 3.1 mm.Overall color medium brown (Fig. 37).Face uniformly brown; band across head apex stramineous, complete; crown midline dark brown in distal half, coronal suture black.Pronotum with pair of submedian ovoid white spots, and two smaller sublateral spots; posterior margin narrowly white, bordered anteriorly with dark brown.Mesonotum and scutellum black except paired white posterolateral spots.Scutellum black except white apex.Forewing clavus with three diamond-shaped white transcommisural markings, progressively larger posteriorly; corium with two small white spots along claval suture, one in inner anteapical cell; brochosome field white tinged with orange; costal margin with three brown false veinlets preapically.Legs stramineous except middle and hind femora mostly black, hind tibia with macrosetae in rows AD, PD and AV black.Forewing R and M confluent for considerable distance.
Remarks.This species closely resembles E. moralesi but is most readily distinguished by the external color pattern, the slender, needlelike pygofer appendage, and the shorter aedeagal shaft relative to the preatrium.
Eualebra moralesi sp.nov.Figs 38,[238][239][240][241]260 Description.Length of male 3.4 mm.Overall color medium brown (Fig. 38).Face with lateral areas dull stramineous; ocellar vestiges and indistinct broken band across head apex stramineous; crown midline stramineous in distal half, coronal suture black.Pronotum with teardrop-shaped median white spot, pair of large submedian ovoid white spots, and two smaller sublateral spots; posterior margin narrowly white, bordered anteriorly with dark brown.Mesonotal triangles dark brown, posterolateral angles white.Scutellum black except white anteromedially and at apex.Forewing clavus with two white transcommisural markings, anterior one thin and arcuate, posterior one pentagonal; corium with two small white spots along claval suture and large ovoid white spot tinged with orange encompassing brochosome field; costal margin with three brown false veinlets preapically.Legs stramineous except front femur and tibia pale orange, middle femur mostly black, hind tibia with macrosetae in basal half of rows AD and PD and at apex of AV black.Forewing with RP and MA confluent for short distance.
Male 2S apodemes (Fig. 260) widely separated, moderately wide and parallel-sided, extended to posterior margin of sternite IV.Pygofer in dorsal view with posterior margin broadly and shallowly concave (Fig. 239), ventral appendage slender, areolate distally, curved evenly dorsad in lateral view, nearly straight in ventral view (Figs 238 and 239).Anal hook (Fig. Etymology.The species is named in honor of Juan Evo Morales Ayma, President of the Republic of Bolivia.
Remarks.This species differs from E. leoni in its color pattern, longer aedeagal shaft, and thicker, areolate ventral pygofer appendage.
Type species: Euzyginella marki sp.nov.Diagnosis. Small,.Coloration pale yellow, orange, or reddish brown, dorsum with symmetrical white, orange, black and/or red markings.Head  wider than pronotum, depressed, anterior margin angulately produced; face nearly horizontal (Fig. 64); rostrum not extended beyond front trochanters; lower part of face sexually dimorphic, male anteclypeus inflated and expanded laterad, lorum very narrow; female anteclypeus relatively narrow, flat, and tapered, lorum relatively broad; frontoclypeus convex; lateral frontal sutures not extended dorsad of antennal pits; antennal ledges weakly developed; antennae as long as head width; ocelli absent; crown convex, coronal suture restricted to posterior third of crown.Pronotum (Figs 39-42) with lateral margins moderately long, slightly divergent posteriorly in dorsal view, distinctly carinate, carina even with posterior margin of eye.Front femur with AM1 enlarged and situated on ventral margin; intercalary row with few fine setae, basal seta larger than others; PV1 well developed; tibia somewhat flattened dorsally, AD and PD without preapical macrosetae.Hind femur macrosetae 2+1+1; tibia row AV with three macrosetae near apex.Forewing (Fig. 75) with RA reflexed, RP confluent with MA for short distance, apical cell 3 triangular, inner apical cell trapezoidal; hind wing (Fig. 76) venation somewhat variable interand intraspecifically, submarginal vein absent between CuA and RP, RP and MA separate, connected by crossvein or confluent; distal segment of CuA (beyond connection with submarginal vein) present.
Etymology.The genus name, a feminine noun, was chosen to indicate its morphological similarity to Zyginella and Eualebra.
Remarks.This genus is similar to Pseudozyginella (and most other Typhlocybini) in having the hind wing submarginal vein not extended to MP (i.e., the hind wing lacks closed apical cells), but differs in in having the row of macrosetae on the subgenital plate continuous and the connective Y-shaped, and in lacking basolateral apodemes on the anal ring.The species are also somewhat broader and more strongly depressed than in Pseudozyginella.The four known species, described below, were collected by fogging the canopy of terra firme forest at two localities in Orellana Province, Ecuador.
Material examined.Holotype male, ECUADOR: Orellana, Res.Etnica Waorani, Transect Ent.Etymology.This species is named in honor of my nephew, Mark Schultz.
Etymology.This species is named in honor of my cousin, Albert E. Dietrich, IV.
Euzyginella teralberti sp.nov.Figs 41,[273][274][275][276]339 Description.Length of male 2.8 mm.Crown (Fig. 41) white with narrow transverse red-orange band between anterior corners of eyes; pronotum red-orange with white band across posterior margin; mesonotum red-orange; scutellum white; forewing clavus red-orange with small white mark along anal margin, large transcommisural diamond-shaped mark near midlength, and white mark at extreme apex; corium red-orange basally, brochosome field white bordered with brown, distal third smoky hyaline with darker mark in inner apical cell and veins bordered with fuscous; venter dull stramineous with apex of anteclypeus and front and middle legs infused with fuscous.
Etymology.This species is named in honor of my uncle, Albert E. Dietrich, III.
Type species: N. baileyi sp.nov.Diagnosis. Small,.Coloration pale yellow, dorsum with symmetrical bright yellow, orange, black and/or red markings; head with two pairs of red spots, small pair on margin of crown near apex, larger pair just anteromesad of eyes.Head wider than pronotum, depressed, anterior margin angulately produced; face nearly horizontal (Fig. 65); rostrum not extended beyond front trochanters; lower part of face sexually dimorphic, male anteclypeus inflated and expanded laterad, lorum very narrow; female anteclypeus relatively narrow, flat and tapered, lorum relatively broad; frontoclypeus convex; lateral frontal sutures not extended dorsad of antennal pits; antennal ledges weakly developed; antennae as long as head width; ocelli absent; crown convex, coronal suture restricted to posterior third of crown.Pronotum  with lateral margins moderately long, slightly divergent in dorsal view, distinctly carinate, carina even with posterior margin of eye.Front femur with AM1 enlarged and situated on ventral margin; intercalary row with few fine setae, basal seta larger than others; PV1 well developed; tibia somewhat flattened dorsally, AD and PD without preapical macrosetae.Hind femur macrosetae 2+1+1; tibia row AV with three macrosetae near apex.Forewing (Fig. 77) with RA reflexed, RP confluent with MA for short distance, apical cell 2 triangular, inner apical cell trapezoidal; hind wing (Fig. 78) veins RP and MA separate, connected by crossvein, submarginal vein incomplete, extended from jugal lobe to point slightly beyond MP, forming closed cell between MP and CuA.
Male abdomen (Figs 341-346) with 2S apodemes well developed, connected by sclerotized bridge at base but widely separated distally, more or less parallel, usually extended beyond posterior margin of sternite III.Pygofer (Figs 277 and 278) with dorsal emargination well developed; tergite well developed but short; dorsal part of lobe heavily sclerotized or not, with or without digitiform distal process; ventral appendage well developed, elongate; one or more additional posterior pygofer spines also present in several species.Valve short, emarginate posteriorly, fused to pygofer.Segment X (Fig. 277) largely membranous with sclerotized basal ring, lateral ends of ring articulated to internal surface of pygofer lobe or to separately articulated anal hook.Subgenital plate (Fig. 277) narrow basally, broadened near middle, with 3-5 macrosetae in sublateral row near apex; apex weakly to strongly compressed.Connective (Fig. 280) U-or Yshaped, articulated to aedeagus, without prominent posterodorsal lobes.Style (Figs 279 and 280) apodeme short, apophysis elongate, with few fine setae preapically, apex tapered and curved.Aedeagus (Figs 279 and 280) with preatrium well developed, dorsal apodeme weakly to strongly developed, shaft longer than preatrium, with or without paired processes.
Etymology.The genus name, a feminine noun, combines Neo-(new) with Zyginella, the name of an apparently related Old World genus.
Remarks.This genus shares a hind wing venational pattern with Pseudhadina gen.nov., in which the submarginal vein connects the apices of CuA and MP to form a closed apical cell but does not extend to MA.This pattern is unknown among other Typhlocybini.Neozyginella is easily distinguished from other Typhlocybini by the unique color pattern.The known species are recorded from terra firme forests in western Amazonia and premontane forests of the eastern Andean foothills.
Neozyginella baileyae sp.nov.Figs 43,[277][278][279][280]341 Description.Length of male 3.3-3.5 mm.Color (Fig. 43) off white, heavily infused with yellow; head with posterior pair of red spots relatively large; pronotum, base of clavus, and apex of scutellum without brown pigment; pronotum with extensive posteromedial orange area extended posterolaterally onto mesonotum; scutellum apex orange; irregular orange band extended length of clavus onto corium, ending just short of apical cells; dark brown maculae present on costal margin at base and apex of brochosome field.
Neozyginella bethae sp.nov.Figs 44,[281][282][283][284]342 Description.Length of male 3.3 mm.Color (Fig. 44) off white, infused with yellow; head with posterior pair of red spots relatively large; pronotum with large round medial black spot; apex of scutellum black; forewing clavus with black macula at base and another near midlength of commisural margin, corium with black macula at base of brochosome well developed.
Etymology.This species is named in honor of my cousin, Marilyn Beth D. Lindberg.
Etymology.This species is named in honor of my cousin, Bradley Dietrich.Neozyginella braxtoni sp.nov.Figs 46,[289][290][291][292]344 Description.Length of male 3.0 mm.Color (Fig. 46) off white, heavily infused with yellow; head with posterior pair of red spots relatively large; small brown spot present medially on pronotum, one at base of forewing clavus, and one at base of costal brochosome field; apex of scutellum black.
Male abdomen (Fig. 345) with 2S apodemes slender, bowed laterad, extended to posterior margin of sternite IV.Pygofer posterodorsal emargination U-shaped, moderately broad (Fig. 294); dorsal process absent; posterior lobe rounded, posteromedial appendage present just dorsad of and closely appressed to ventral appendage; both appendages slender, acuminate and curved dorsomesad (Figs 293 and 294).Anal hook (Fig. 293) robust, moderately long, curved ventrad, apex connected to basal ring of anal tube by sclerotized bar.Subgenital plate (Fig. 293) with four preapical macrosetae.Style apophysis (Figs 295 and 296) slender and sinuate in lateral view, broadened near midlength in ventral view, apex blunt, straight.Connective (Fig. 296) broadly U-shaped.Aedeagus (Figs 295 and 296) with preatrium moderately long, moderately broad in ventral view; dorsal apodeme well developed with compressed anterior lobe and posterior transverse plate; shaft in lateral view broadened toward midlength with prominent dorsal lobe, then abruptly narrowed and slender to slightly expanded apex, broad and parallel sided in posterior view, with pair of rounded lateral lobes at apex and pair of slender strongly recurved lateroapical processes, basal processes absent; gonopore apical.Female as described for genus.
Etymology.This species is named in memory of my niece, Jennifer Craft.
Etymology.The name, a feminine noun, was formed by combining "pseud-", meaning false, with Eurhadina, a superficially similar Old World genus.
Remarks.This genus is based on a single species from the eastern Ecuadorean rainforest canopy.It appears to be morphologically intermediate between Neozyginella and Columbonirvana.It is readily distinguished from other South American Typhlocybini by the narrow, flattened (not sexually dimorphic) male anteclypeus.It resembles Eualebra in shape and proportions but differs in having the coronal suture extended nearly to the apex of the crown and the face in profile oblique rather than horizontal.
Etymology.The genus name, a feminine noun, combines "pseudo-" meaning "false" with Zyginella, and reflects the morphological similarity to the latter genus.
Remarks.This is the only New World typhlocybine genus known to include species with hind wing venation resembling that of the Old World genus Zyginella, although the Zyginella-like features (i.e., absence of a distal segment of CuA extended beyond the submarginal vein) are not consistently present and, in some cases, vary between the right and left wings of a single specimen.Thus, for members of Pseudozyginella, at least, the criteria previously used by some previous authors to justify recognition of Zyginellini as a separate tribe are unstable, providing support for the present treatment of Zyginellini as a junior synonym of Typhlocybini.The new genus differs from Zyginella and other genera placed by Dworakowska (1979) in "Zyginellini" in having a well-developed, elongate ventral pygofer process similar to that of Columbonirvana.It also resembles Columbonirvana in having a well-developed anal hook articulated to the pygofer and anal tube, a slender apically setose style, and subgenital plates with a reduced number of macrosetae.The genus includes three species, all collected by fogging terra firme rainforest canopy at a single locality in eastern Ecuador.
Male abdomen (Fig. 349) with 2S apodemes vestigial (possibly due to parasitism of only available male specimen).Pygofer (Figs 309 and 310) with dorsal appendage robust, tapered, evenly curved posteroventrad and extended slightly beyond margin of lobe; ventral appendage relatively slender, curved slightly dorsad through most of length, apex curved slightly laterad.Subgenital plate (Fig. 309) with 1-2 macrosetae near midlength and 1 at apex.Style apophysis (Figs 311 and 312) slightly tapered through most of length, nearly straight, apex bent ventrad and curved slightly mesad, acuminate.Connective (Fig. 312) broadly U-shaped, arms slender.Aedeagus (Figs 311 and 312) with preatrium slightly shorter than shaft, relatively slender and constricted near midlength in ventral view; dorsal apodeme weakly developed; shaft slender, tubular, curved slightly dorsad, with basal processes relatively large, evenly divergent from shaft in ventral view, extended well beyond shaft midlength; with pair of slender retrorse lateral processes at apex, extended anteroventrad.Female as described for genus.
Remarks.This genus is structurally similar to Columbonirvana but differs in the larger body size, paler dorsal coloration, emarginate forewing apex, and Y-shaped connective.The five known species were collected in Malaise traps in cloud forests at two localities in Colombia and Peru.Tahurella josephi sp.nov.Figs 53,[317][318][319][320]351 Description.Length of male 4.8 mm.Dorsum (Fig. 53) pale yellow with symmetrical red-orange and dark brown markings; head anterior margin with narrow dark brown band between eyes, discontinuous with similar band extended across thoracic pleuron; pronotum with anterior margin red-orange, posterior margin with convex red-orange marking medially; mesonotum orange anteriorly; scutellum orange at apex.Forewing with series of 7 oblique brown lines distributed along costal margin; basal third of corium and clavus hyaline with narrow red-orange transcommisural arcuate band; distal 2/3 of forewing smoky hyaline with paler areas in apical cells; veins orange; distal lobe well developed.Thoracic venter pale yellow; abdomen pale yellow ventrally except sternite VIII and valve brown.
Tahurella katherinae sp.nov.Figs 54,[321][322][323][324]352 Description.Length of male 5.0 mm.Dorsum (Fig. 54) pale yellow with symmetrical red-orange and dark brown markings; head anterior margin with narrow dark brown band between eyes, discontinuous with similar band extended across thoracic pleuron; pronotum with anterior margin red-orange, posterior margin with convex red-orange marking medially; mesonotum orange anteriorly; scutellum orange at apex.Forewing with series of 7 oblique brown lines distributed along costal margin; basal third of corium and clavus hyaline with narrow red-orange transcommisural arcuate band; distal 2/3 of forewing smoky hyaline with paler areas in apical cells; veins orange; apical lobe well developed.Thoracic venter pale yellow; abdomen pale yellow ventrally except sternite VIII and valve brown.Head slightly narrower than pronotum, crown twice as long medially as next to eye; forewing apical cell 2 quadrate; hind tibia row AV with 4 macrosetae.Tahurella lynnae sp.nov.Figs 55,68,84,[325][326][327][328]353 Description.Length of male 5.3 mm.Dorsum (Fig. 55) yellow to white, heavily marked with dark brown and orange; head anterior margin with pair of thin brown transverse bands between eyes, crown yellow; pronotum white with anterior margin red-orange and incomplete posterior submarginal orange transverse band; mesonotum and scutellum orange; forewing basal half with alternating white, red-orange, white, and dark brown transcommisural arcuate bands; brochosome field bright yellow, costal margin with 6 oblique brown bands, veins yellow, cells with large fuscous areas; thorax and abdomen brown ventrally.Head subequal to pronotum in width, crown slightly less than twice as long medially as next to eyes.Forewing with apical cell 2 triangular, apical lobe short (Fig. 84).Hind tibia row AV with 4 macrosetae.Male 2S apodemes (Fig. 353) weakly divergent, extended to posterior margin of sternite IV.Pygofer with dorsal emargination broad, parabolic (Fig. 326); tergite ca.1/3 length of distal lobe; apical process long, slender, extended ventrad; ventral appendage weakly curved dorsomesad (Figs 325 and 326).Anal hook (Fig. 325) small, triangular.Subgenital plate (Fig. 325   Etymology.This species is named in honor of my niece Lynn Dietrich. Tahurella clairae sp.nov.Figs 56,[329][330][331][332]354 Description.Length of male 4.7 mm.Dorsum (Fig. 56) yellow to white, heavily marked with dark brown and orange; head anterior margin with pair of thin brown transverse bands between eyes, crown yellow; pronotum white with anterior margin red-orange and incomplete posterior submarginal orange transverse band; mesonotum and scutellum orange; forewing basal half with alternating white, red-orange, white, and dark brown transcommisural arcuate bands; brochosome field bright yellow, costal margin with 6 oblique brown bands, veins yellow, cells with large fuscous areas; thorax and abdomen brown ventrally; apical lobe weakly developed.Head subequal to pronotum in width, crown slightly less than twice as long medially as next to eyes.Forewing with apical cell 2 triangular, apical margin concave.Hind tibia row AV with 4 macrosetae.
Etymology.The species is named in honor of my cousin, Heather B. Denne.
brown anteriorly, yellow posteriorly, area anterad of scutellar suture brown; scutellum yellow with brown transverse band.Forewing clavus with two triangular transcommisural markings, one near base an another smaller one near apex, basal triangle orange medially; corium with two round spots, one near base and another near midlength; costal area with brochosome field bright red; veins mostly orange; outer two apical cells hyaline distally.Body robust, ovoid.Forewing with RP and MA separate, connected by crossvein; CuA connected to MP. Hind tibia row AV with 3 macrosetae.