South American Leafhoppers of the Tribe Typhlocybini (Hemiptera: Cicadellidae: Typhlocybinae)

ZOOLOGIA 30 (5): 519–568, October, 2013 http://dx.doi.org/10.1590/S1984-46702013000500008 South American leafhoppers of the tribe Typhlocybini (Hemiptera: Cicadellidae: Typhlocybinae)

Christopher H. Dietrich

Illinois Natural History Survey, Prairie Research Institute, University of Illinois, 1816 S. Oak St., Champaign, IL 61820, USA. E-mail: [email protected]

ABSTRACT. 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, sug- gesting 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. KEY WORDS. Distribution; Homoptera; morphology; Neotropical; taxonomy.

The leafhopper subfamily Typhlocybinae comprises ~6,000 also includes Columbonirvana Linnavuori, a genus previously described species of mostly small, delicate leafhoppers that feed placed in Nirvaninae (LINNAVUORI 1959), but transferred to preferentially on the contents of leaf parenchyma cells of their Typhlocybinae by DIETRICH (2004) based on phylogenetic analy- host plants. Based on described species, this is currently the sec- sis of morphological characters. These two genera include a ond largest leafhopper subfamily (after Deltocephalinae, ZAHNISER total of three previously described species (plus one new syn- & DIETRICH 2010), but recent sampling of tropical faunas indi- onym, designated below). Recent sampling indicates that the cates that the extant, but mostly undescribed, fauna of South American fauna of Typhlocybini is much richer in both Typhlocybinae is far larger than that of any other leafhopper species and genera. In this paper, I review the endemic South subfamily (DIETRICH & WALLNER 2002). The Neotropical fauna, in American genera and species of Typhlocybini and discuss their particular, is highly diverse but appears to remain largely un- possible relationships to the fauna of this tribe in other parts documented (DIETRICH & DMITRIEV 2006, 2008). of the world. Considering that the 52 new species of Study of canopy fogging samples from lowland rainforests Typhlocybini described herein were collected from only eight in eastern Peru and Ecuador (DIETRICH & WALLNER 2002) indi- localities in Bolivia, Brazil, Colombia, Ecuador and Peru, fur- cates the presence of an enormously diverse typhlocybine fauna. ther sampling in South America, especially in cloud forests of Composition of this fauna contrasts strikingly with that of the the eastern Andean foothills, will likely reveal the presence of somewhat better known, but still poorly documented faunas of a much more speciose assemblage belonging to this tribe. the Old World tropics. Tribes Dikraneurini, Empoascini (= Jorumini) and Alebrini, in decreasing order of species rich- MATERIAL AND METHODS ness, are well represented in the samples from Amazonia; but Erythroneurini and Typhlocybini, so abundant in north tem- Morphological terminology follows that of YOUNG (1952) perate forests and in the Old World tropics, are relatively rare. as modified by DIETRICH (2005) except wing veins are labeled Malaise trap samples from the eastern Andean foothills according to the simplified system illustrated in figures 3i, j. above 500 m elevation differ from those obtained from the Drawings of genitalia were made by tracing over photographs canopy of lowland rainforest (unpublished data) in that they taken using a digital camera mounted on a compound micro- include larger numbers of Typhlocybini (sensu stricto). These scope. In drawings of the male genital capsule, the entire anal species belong to a group of apparently closely related, endemic complex (segments X-XI) is shown in lateral view but only the South American genera that includes Eualebra Baker. The group well sclerotized and pigmented parts of segment X are shown

© 2013 Sociedade Brasileira de Zoologia | www.sbzoologia.org.br | All rights reserved. 520 C.H. Dietrich in dorsal view. Material examined is deposited in the follow- 2’. Hind wing submarginal vein incomplete, either extended ing institutions: Humboldt Institute, Villa de Leyva, Colombia along costal margin but not connecting vein apices along (HIC); Carnegie Museum of Natural History (ICCM); Illinois apical margin, or connecting vein apices along apical Natural History Survey, Champaign (INHS); Universidad de San margin but not extended along costal margin (terminated Marcos, Lima (USML), United States National Museum (USNM). at apex of vein RP or R+M)...... 3 3. Forewing with inner apical cell parallel-sided, hind wing TAXONOMY submarginal vein absent along apical margin, vannal vein unbranched ...... Erythroneurini Typhlocybinae Kirschbaum, 1868 3’. Forewing with inner apical cell usually distinctly tapered The tribal classification of Typhlocybinae has long been through most of length (often short and oblique), hind unstable, with some authors (e.g., YOUNG 1952) having recog- wing submarginal vein, if present, not extended along costal nized as few as four tribes and others (e.g., RUPPEL 1987) as many margin basad of RP or R+M, vannal vein branched ...... 4 as ten. In his review of Western Hemisphere Typhlocybinae, 4. Ocelli absent or vestigial, hind wing submarginal vein absent YOUNG (1952) employed a relatively broad concept of between apices of veins MA or RP+MA and MP (Fig. 78) or, Typhlocybini, including as synonyms the tribes Eupterygini if present (Fig. 70), body strongly depressed with face nearly Kirkaldy, 1906, Jorumini McAtee, 1926, and Empoascini Dis- horizontal ...... Typhlocybini tant, 1908, despite differences in the hind wing venation of 4’. Ocelli well developed (except in Paulomanus), hind wing these groups. Dworakowska, the principal worker on the Old submarginal vein present between apices of veins MA or World typhlocybine fauna for the past four decades, recognized RP+MA and MP, body not strongly depressed, face in profile tribes Typhlocybini, with synonym Eupterygini; Empoascini, oblique, not nearly horizontal ...... Empoascini with synonym Jorumini; and Zyginellini (DWORAKOWSKA 1979). The latter tribe was erected to comprise species having the hind Typhlocybini Kirschbaum wing submarginal vein apparently connected directly to CuA Typhlocybini Kirschbaum, 1868: 16. rather than being joined by a crossvein (alternatively inter- Eupteryginae Kirkaldy, 1906: 296. preted as the absence of the segment of vein CuA distad of its Zyginellini Dworakowska, 1979: 299. junction with the submarginal vein). Despite the unique hind Diagnosis. Species of Typhlocybini (sensu AHMED 1983) wing venation, Zyginellini, as defined by DWORAKOWSKA (1979), differ from other Typhlocybinae in having the following com- is a rather heterogeneous assemblage. Most genera placed in bination of morphological features: ocelli absent or vestigial; this tribe strongly resemble many Typhlocybini (sensu stricto), forewing appendix absent, inner apical cell short and oblique, particularly in the structure of the male genital capsule (e.g., not extended to apical wing margin; hind wing vannal vein presence of a single macroseta near the base of the plate) and branched, submarginal vein present or absent at wing apex; aedeagus (e.g., presence of elongate paired apical processes). male pygofer with few or no macrosetae (except Eualebra). Moreover, some of the Neotropical species described herein Remarks. The endemic South American taxa treated vary intraspecifically (occasionally intra-individually) for the herein fit AHMED’s (1983) concept of Typhlocybini, but form a features used to distinguish Zyginellini from Typhlocybini. distinctive group within this tribe. Members of this group are AHMED (1983) considered Zyginellini to be an artificial group readily distinguished by their depressed form, produced head, and treated it as a synonym of Typhlocybini. Until phyloge- and presence of two or more macrosetae on the male subgenital netic analyses can clarify the status and relationships of the plate (most Old World and Nearctic Typhlocybini have 0-1). various tribes proposed for groups of typhlocybine genera, it Some of the genera included here have hind wing vena- seems reasonable to adopt the classification of AHMED (1983), tional patterns not known to occur among Typhlocybini in which recognizes five tribes in the subfamily: Alebrini, other parts of the world. One pattern (Figs 72 and 74), exem- Dikraneurini (= Forcipatini), Empoascini (= Jorumini), plified by the genus Eualebra, has the submarginal vein present Erythroneurini, and Typhlocybini (= Eupterygini, Zyginellini). and connecting the apices of veins RP, MA, MP and CuA at the Key to tribes of Typhlocybinae wing apex, but absent along the costal margin; veins RP and MA are free, connected by a crossvein. 1. Forewing appendix present ...... Alebrini The second pattern (Figs 78 and 80), exemplified by new 1’. Forewing appendix absent ...... 2 genera Neozyginella and Pseudhadina, has the submarginal vein 2. Hind wing with submarginal vein complete, extended along extended distad along the anal margin from the jugal lobe and costal margin and connecting apices of all veins (except crossing the tips of veins CuA and MP but not extended to MA. Typhlocybella: submarginal vein absent, vannal vein unbran- A third group of South American genera, exemplified by ched, forewing inner apical cell tapered distally) ...... Euzyginella gen. nov. and Pseudozyginella gen. nov., has the hind ...... Dikraneurini wing submarginal vein absent at the wing apex as in Old World

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Typhlocybini, but varies intraspecifically, and occasionally from MA apically ...... 5 within the same individual, for features traditionally diagnostic 2. Hind wing submarginal vein extended across apices of CuA of tribes Typhlocybini, Eupterygini, and Zyginellini (Figs 82 and and MP, forming closed cell (Figs 78, 80) ...... 4 83). Hind wing veins RP and MA may be separate and connected 2’. Hind wing submarginal vein not extended to apex of MP, by a crossvein (as in Eupterygini) or confluent (as in either forming continuous line with CuA (Fig. 82) or Typhlocybini, s.s.). Also, the hind wing submarginal vein may terminating slightly beyond CuA apex (Fig. 83) ...... 3 be strongly curved and colinear with CuA preapically (as in “Zyginellini”) or straight and connected to CuA by a crossvein 3. Male connective Y-shaped, stem as long as or longer than (as in Typhlocybini, s.s.). Presence of such variation within a arms (Fig. 264); sclerotized base of segment X, in dorsal single species provides support for AHMED’s (1983) treatment of view, U-shaped, without anterolateral projections (Fig. 262) Eupterygini and Zyginellini as junior synonyms of Typhlocybini...... Euzyginella gen. nov. The presence of a submarginal vein at the hind wing 3’. Male connective U-shaped, stem not developed (Fig. 308); apex and relatively numerous macrosetae on the male sclerotized base of segment X, in dorsal view, with pair of subgenital plates (as in Alebrini and Empoascini) suggests that sclerotized anterolateral projections articulated to pygofer Eualebra and related South American genera represent a (Fig. 306) ...... Pseudozyginella gen. nov. plesiomorphic lineage of Typhlocybini not represented in the 4. Head with coronal suture not extended beyond midlength Old World. Other Typhlocybini, including genera and species of crown, crown with paired red spots (Fig. 43) ...... endemic to the Old World and North America, lack a submar- ...... Neozyginella gen. nov. ginal vein in the hind wing and almost always have two or 4’. Head with coronal suture extended nearly to apex of crown; fewer (usually 0-1) macrosetae on the male subgenital plate. crown without spots (Fig. 49) ...... Pseudhadina gen. nov. As in some Old World Typhlocybini, the lower part of the face in most South American genera is sexually dimorphic: 5. Form broad and strongly depressed (Figs 18-38), face hori- females have the clypellus and lorum flattened and well de- zontal and flat or concave in lateral view (Figs 62, 63); male limited, but males have the lorum fused to the clypellus and subgenital plate with numerous macrosetae in irregular both structures are markedly inflated (Fig. 59). longitudinal band (Fig. 8a) ...... Eualebra Baker Based on the morphological similarities mentioned above, 5’. Form slender and less depressed (Figs 1-17), face oblique and the South American fauna of Typhlocybini appear to be related usually convex in lateral view (Figs 60, 61, 68); male subgenital to some Old World genera previously included in “Zyginellini” plate with 3-4 macrosetae in single row (Fig. 87) ...... 6 (sensu DWORAKOWSKA 1979) as well as to the “Eupteryx-complex” 6. Dorsal coloration mostly pale yellow with arcuate red and (sensu DWORAKOWSKA 1969) of genera that are most diverse in the brown transverse bands; forewing apex emarginate or with Indomalayan region. These relationships need to be elucidated distinct projection (Figs 84, 85); male subgenital plate with by phylogenetic analysis. small stout setae at apex in addition to row of macrosetae The endemic South American fauna of Typhlocybini near midlength (Fig. 317) ...... Tahurella gen. nov. comprises seven genera: Eualebra, placed by YOUNG (1952) in 6’. Dorsal coloration mostly brown with pale spots (Figs 1-17); Typhlocybini based on the branched vannal vein and absence of a submarginal vein along the costal margin in the hind wing; forewing apex rounded (Fig. 69); male subgenital plate Columbonirvana Linnavuori, previously placed in Nirvaninae without stout apical setae (Fig. 87) ...... Columbonirvana Linnavouri presumably based on its depressed form and produced head, but transferred to Typhlocybinae by DIETRICH (2004); and five new genera described below, Euzyginella, Neozyginella, Columbonirvana Linnavuori, new placement Pseudhadina, Pseudozyginella, and Tahurella. Columbonirvana Linnavuori, 1959: 34. Two Palearctic species of Typhlocybini known to be es- Type species: C. aurea Linnavuori, 1959. tablished in Argentina and Chile, Edwardsiana froggatti (Baker) Diagnosis. Small to medium-sized, depressed, slender and Ribautiana tenerrima (Herrich-Schaffer), have been well de- leafhoppers. Dorsal coloration (Figs 1-17) orange to dark brown scribed elsewhere (CHRISTENSEN 1942, CHRISTIAN 1953) and are with symmetrical yellow, brown or black markings; forewing not treated here. apex usually with false eye spot, costal margin with oblique Key to endemic genera of South American Typhlocybini dark brown false veins; ventral coloration including legs yellow except mesepisternum brown. 1. Hind wing with submarginal vein obsolete apically (Figs 76, Head (Figs 1-17) subequal to or narrower than pronotum, 78, 80, 82, 83); vein RP weak or appearing confluent with depressed, anterior margin angulately produced; face nearly MA ...... 2 horizontal; rostrum extended slightly beyond front trochan- 1’. Hind wing with submarginal vein well developed apically ters; lower part of face sexually dimorphic, male anteclypeus (Figs 70, 72, 74, 86); vein RP well developed and separated inflated and expanded laterad, lorum very narrow; female

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anteclypeus narrow, flat, tapered distally, lorum broader than nus have yet been found. DIETRICH (2004) transferred in male; frontoclypeus convex; lateral frontal sutures not ex- Columbonirvana from Evacanthinae (= Nirvaninae) to tended dorsad of antennal pits; antennal ledges weakly devel- Typhlocybinae but considered its tribal placement uncertain. oped; antennae as long as head width; ocelli absent; crown The genus is here placed in the tribe Typhlocybini based on flat or slightly convex, coronal suture extended to or near apex the concept of the tribe embodied in the above diagnosis, which of crown. Pronotum (Figs 1-17) with lateral margins moder- is narrower than that of YOUNG (1952) but broader than that of ately long, slightly divergent in dorsal view, distinctly carinate, OMAN et al. (1990). It is similar to Tahurella gen. nov. but dif- carina even with posterior margin of eye. Front femur with fers in the smaller size, darker overall coloration, lack of trans- AM1 enlarged and situated on ventral margin; intercalary row verse black band on the anterior margin of the head, and with few fine setae, basal seta larger than others; PV1 absent; rounded forewing apex. tibia rounded dorsally, AD and PD without preapical The Southeast Asian Lowata Dworakowska resembles some macrosetae. Hind femur macrosetae 2+1+1; tibia row AV with species of Columbonirvana in having a pair of narrow transverse 4 macrosetae near apex. Forewing (Fig. 69) with apex rounded, red stripes on the anterior margin of the head but differs in RA reflexed, RP confluent with MA for short distance, apical having the hind wing submarginal vein absent at the wing apex. cell 2 triangular or petiolate; CuA joining M basad of its fork, Key to species of Columbonirvana inner apical cell trapezoidal; apical margin convex. Hind wing (Fig. 70) with RP and MA separate, connected by crossvein; 1. Pronotum with large white or yellow crescent medially (Fig. submarginal vein extended from apex of RP to jugal lobe. 2) ...... 2 Male 2S apodemes (Figs 151-166) well developed, joined 1’. Pronotum uniformly brown or with pair of small white spots at base by transverse bridge, subparallel, extended beyond pos- (Figs 1, 7) ...... 8 terior margin of sternite III. Pygofer (Figs 87 and 88) strongly 2. Posterior margin of pronotum yellow or white (Fig. 2), emarginate dorsally; tergite variable in length and shape among aedeagus with pair of elongate processes arising near base species; posterior lobe with strongly sclerotized acuminate dor- and extended distad along shaft (Fig. 93) ...... 3 sal section variably clothed with microtrichia, unpigmented ventral section bearing longitidinal row or band of short fine setae; 2’. Posterior margin of pronotum brown (Fig. 5); aedeagus ventral appendage slender, elongate, arising near base of ventral without spinelike processes (Fig. 102) ...... 5 margin and curved posterodorsad, usually areolate distally. Anal 3. Aedeagus with ventral processes closely appressed to shaft tube (Figs 87 and 88) depressed, sclerotized dorsal band short, in lateral view (Fig. 93) ...... ameliae sp. nov. with pair of basolateral hooks. Subgenital plate (Figs 87 and 88) 3’. Aedeagus with ventral processes distinctly divergent from constricted near base, broadest near midlength, with row of 3-4 shaft in lateral view (Fig. 105) ...... 4 macrosetae near lateral margin, apical macrosetae absent; fine dorsal setae divided into medial and distal groups; apical lobe 4. Dorsal sclerotized part of pygofer apex broad and only slightly compressed, parallel sided and darkly pigmented. Style (Figs 89 curved ventrad (Fig. 95) ...... craigi sp. nov. and 90) apodeme short; apophysis elongate, without preapical 4’. Dorsal sclerotized part of pygofer apex slender and sharply lobe, with fine setae preapically. Connective (Figs 90 and 94) U- bent ventrad (Fig. 103) ...... edgari sp. nov. or V-shaped, articulated to aedeagus. Aedeagus (Figs 89 and 90) 5. Aedeagal shaft distinctly denticulate preapically (Fig. 129) .... 6 with preatrium elongate, dorsal apodeme weakly developed; shaft 5’. Aedeagal shaft without denticuli (Fig. 133) ...... 7 elongate and nearly straight, with or without paired ventral processes; apical processes absent; gonopore apical. 6. Aedeagus with denticuli restricted to margins of preapical Female (based only on type species) sternite VII with pos- flange (Fig. 150) ...... vatia sp. nov. terior margin angulately produced; ovipositor not extended 6’. Aedeagus with denticuli scattered over preapical surface of beyond pygofer; first valvulae with strigate sculpturing dorsally shaft (Fig. 133) ...... thomasi sp. nov. near apex; second valvulae (Fig. 357) elongate, slender, curved 7. Pygofer with ventral appendage gradually curved dorsad in dorsad through most of length, slightly sinuate preapically, right lateral view (Fig. 127); aedeagal shaft widest at apex in la- blade with 9-10 small, rounded, close-set teeth near apex. teral view (Fig. 129) ...... schulzi sp. nov. Remarks. The type species of this genus was described 7’. Pygofer with ventral appendage abruptly bent dorsad near based on a single female from Colombia (illustrated by midlength (Fig. 99); aedeagal shaft widest near base in la- LINNAVUORI 1959 and DIETRICH 2004). No additional specimens teral view (Fig. 101) ...... davidi sp. nov. matching the type specimen of C. aurea Linnavuori in structure or color pattern have been found, but numerous conge- 8. Anterior margin of head with pair of transverse parallel red neric male specimens representing 16 additional species were stripes (Fig. 61) ...... 9 collected by Malaise trap in cloud forests in the Andes Moun- 8’. Anterior margin of head white, bordered with light brown, tains of Peru. No additional female representatives of this ge- without red stripes (Fig. 11) ...... noahi sp. nov.

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9. Forewing apical cells with dark brown markings extended Male 2S apodemes (Fig. 151) robust, parallel sided, weakly to margin (Fig. 61) ...... 10 divergent, extended to posterior margin of sternite IV. Pygofer 9’. Forewing apical cells pale to light brown, third apical cell dorsal emargination (Fig. 88) with lateral margins convexly with dark brown medial spot not extended to margin (Fig. rounded, broadest distally; tergite produced medially, 1/3 69) ...... 16 length of distal lobe; dorsal sclerotized area of lobe weakly falcate but not produced beyond margin; ventral appendage 10. Aedeagus with paired ventral processes (Fig. 109) ...... 11 strongly curved dorsad, apex areolate, curved posterolaterad 10’. Aedeagus without processes (Fig. 117) ...... 14 (Fig. 87). Anal hook (Fig. 87) elongate and slender, weakly sinu- 11. Aedeagal processes arising at or near base of shaft (Fig. 109) ate, extended ventrad. Subgenital plate (Fig. 87) with 3-4 ...... 12 macrosetae near midlength, without stout apical setae, apex 11’. Aedeagal processes arising at or near midlength of shaft abruptly curved dorsad. Style apophysis broadened preapically (Fig. 137) ...... 13 in ventral view (Fig. 87), apex tapered, curved ventrolaterad. Connective U-shaped (Fig. 90). Aedeagus (Figs 89 and 90) with 12. Pygofer with dorsal process gently curved ventrad (Fig. 107); preatrium distinctly shorter than shaft, broadened basally in aedeagal processes straight in lateral view (Fig. 109) ...... ventral view; shaft slender, tubular, curved dorsad, with pair ...... ernae sp. nov. of ventral processes arising near base, closely paralleling shaft, 12’. Pygofer with dorsal process bent ventrad at right angle and extended to or slightly beyond shaft apex; shaft apex slen- (Fig. 111); aedeagal processes distinctly curved in lateral der, compressed. view (Fig. 113) ...... joshuai sp. nov. Material examined. Holotype male, PERU: Pasco, Yanachaga- 13. Aedeagal processes very short, arising from ventrolateral Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 Octo- flange, shaft apex not expanded in lateral view (Fig. 141)... ber 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio ...... tumida sp. nov. San Alberto [USML]. Paratypes: 7 males, same data [INHS]. 13’. Aedeagal processes long, arising from tubular shaft, shaft Etymology. This species is named in honor of my nephew, apex expanded in lateral view (Fig. 137) ...... Aidan Dietrich...... tuberculata sp. nov. Columbonirvana ameliae sp. nov. 14. Aedeagal shaft in lateral view with apex narrower than base Figs 2, 91-94, 152 (Fig. 117) ...... margaretae sp. nov. 14’. Aedeagal shaft in lateral view with apex wider than base Description. Length of male 4.2 mm. Dorsum (Fig. 2) (Fig. 121) ...... 15 dark reddish brown with symmetrical pale yellow markings; anterior margin of head with pair of thin red transverse bands 15. Aedeagal shaft with angulate preapical ventral projection between eyes; crown yellow bordered anteriorly with brown; (Fig. 121) ...... matthewi sp. nov. pronotum yellow medially with anterior and lateral margins 15’. Aedeagal shaft with ventral margin broadly rounded brown and with brown transverse submarginal band posteri- preapically (Fig. 145) ...... urodolobrata sp. nov. orly; mesonotum yellow medially, anterolateral triangles and 16. Forewing with round white spot near midlength of clavus apex of scutellum brown; forewing clavus with small white and another on adjacent area of corium (Fig. 1) ...... transcommisural spot bordered posteriorly with larger crim- ...... aidani sp. nov. son area, veins orange bordered with brown in distal half; venter of thorax pale yellow except mesosternum orange brown. 16’. Forewing with oblique white band on clavus extended onto Head as wide as pronotum, crown nearly twice as long medi- corium (Fig. 60)...... aurea Linnavuori ally as next to eye. Columbonirvana aidani sp. nov. Male 2S apodemes (Fig. 152) robust, subparallel, extended to posterior margin of sternite IV. Pygofer dorsal emargination Figs 1, 69-70, 87-90, 151 (Fig. 92) parabolic; tergite 1/3 length of distal lobe; dorsal scle- Description. Length of male 4.3 mm. Dorsum (Fig. 1) rotized area of lobe falcate, sinuate preapically, not produced mottled orange brown with symmetrical white spots; head beyond apical margin; ventral appendage gradually curved anterior margin with pair of thin red transverse bands between dorsad, apex curved posterolaterad (Fig. 91). Anal hook (Fig. eyes; pronotum with two pairs of posterolateral white spots; 91) small and triangular. Subgenital plate (Fig. 91) with 3 macro- forewing clavus with large round white medial spot, brachial setae near midlength and few short, stout preapical setae, apex cell with smaller white spot, veins orange bordered with dark extended posterad. Style apophysis (Figs 93 and 94) parallel brown distally; venter of thorax pale yellow except mesoster- sided through most of length, apex tapered, curved ventro- num dark brown; abdomen brown ventrally except near laterad. Connective (Fig. 94) V-shaped. Aedeagus (Figs 93 and midlength of subgenital plates. Head as wide as pronotum, 94) with preatrium distinctly shorter than shaft, slightly broad- crown slightly longer medially than next to eye. ened basally in ventral view; shaft slender, tubular, nearly

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3233 34 35 36 37 38 Figures 1-38. Dorsal habitus. (1-17) Columbonirvana: (1) C. aidani sp. nov.; (2) C. ameliae sp. nov.; (3) C. aurea Linnavuori; (4) C. craigi sp. nov.; (5) C. davidi sp. nov.; (6) C. edgari sp. nov.; (7) C. ernae sp. nov.; (8) C. joshuai sp. nov.; (9) C. margaretae sp. nov.; (10) C. matthewi sp. nov.; (11) C. noahi sp. nov.; (12) C. schulzi sp. nov.; (13) C. thomasi sp. nov.; (14) C. tuberculata sp. nov.; (15) C. tumida sp. nov.; (16) C. urodolobrata sp. nov.; (17) C. vatia sp. nov.; (18-38) Eualebra: (18) E. barbarae sp. nov.; (19) E. charlesi sp. nov.; (20) E. dorisae sp. nov.; (21) E. jessicae sp. nov.; (22) E. patriciae sp. nov.; (23) E. gingerae sp. nov., male holotype; (24) same, female specimen from Ecuador; (25) E. marilynae sp. nov.; (26) E. susanae sp. nov.; (27) E. helenae sp. nov.; (28) E. kathyae sp. nov.; (29) E. peggyae sp. nov.; (30) E. jenniferae sp. nov.; (31) E. smithii Baker, male specimen from Bolivia; (32) E. smithii Baker, female holotype of D. (H.) reticulata Osborn from Brazil; (33) E. margaretannae sp. nov.; (34) E. alberti sp. nov.; (35) E. dorothyae sp. nov.; (36) E. michaelorum sp. nov.; (37) E. leoni sp. nov.; (38) E. moralesi sp. nov. Scale bars = 1 mm.

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60 61 62

65 63 64

66 67 68 Figures 39-68. Dorsal habitus, lateral habitus, and face. (39-42) Euzyginella: (39) E. marki sp. nov.; (40) E. quartalberti sp. nov.; (41) E. teralberti sp. nov.; (42) E. theari sp. nov. (43-48) Neozyginella: (43) N. baileyae sp. nov.; (44) N. bethae sp. nov.; (45) N. bradleyi sp. nov.; (46) N. braxtoni sp. nov.; (47) N. dougi sp. nov.; (48) N. jenncraftae sp. nov.; (49) Pseudhadina amazonica sp. nov.; (50-52) Pseudozyginella: (50) P. grahami sp. nov.; (51) P. davei sp. nov.; (52) P. dougbraxtoni sp. nov. (53-57) Tahurella: (53) T. josephi sp. nov.; (54) T. katherinae sp. nov.; (55) T. lynnae sp. nov.; (56) T. clairae sp. nov.; (57) T. heatherae sp. nov. (58-69) anteroventral view of head: (58) Pseudhadina amazonica, male; (59) Columbonirvana noahi, male. (60-68) lateral habitus: (60) Columbonirvana aurea, holotype female; (61) C. matthewi sp. nov.; (62) Eualebra margaretannae sp. nov.; (63) E. michaelorum sp. nov.; (64) Euzyginella quartalberti sp. nov.; (65) Neozyginella baileyi sp. nov.; (66) Pseudhadina amazonica sp. nov.; (67) Pseudozyginella grahami sp. nov.) (68) Tahurella lynnae sp. nov. Scale bars = 1 mm.

ZOOLOGIA 30 (5): 519–568, October, 2013 526 C.H. Dietrich

straight, with pair of ventral processes arising near base, closely Material examined. Holotype male, PERU: Pasco, Villa paralleling shaft, and extended 2/3 distance to shaft apex; shaft Rica, 1400m, 10°43’21"S 75°15’43"W, 21 October 2002, C. H. apex slightly compressed, rounded in lateral view, weakly emar- Dietrich, vacuum, 2-28-1 [USML]. ginate in ventral view. Etymology. This species is named in honor of my brother, Material examined. Holotype male, PERU: Pasco, Yanachaga- Craig B. Dietrich. Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 Octo- ber 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio Columbonirvana davidi sp. nov. San Alberto [USML]. Paratypes: 4 males, same data [INHS]. Figs 5, 99-102, 154 Etymology. This species is named in honor of my niece, Amelia Dietrich. Description. Length of male 3.4 mm. Dorsum (Fig. 5) dark brown with large symmetrical yellow markings. Anterior Columbonirvana aurea Linnavuori, 1959 margin of head with two narrow transverse red bands; crown with posterior margin infused with yellow medially. Pronotum Figs 3, 60, 18a-b with large transverse yellow crescent. Mesonotum brown with Columbonirvana aurea Linnavuori, 1959: 34. large yellow median spot. Forewing clavus with large symmetri- Remarks. This species, the type species of the genus, was cal transcommisural marking; veins mostly orange; anteapical described based on a single female from “Sierra San Lorenzo”, and apical cells each with medial area hyaline; brachial and Colombia, by LINNAVUORI (1959). DIETRICH (2004) examined the discal cell each with hyaline area near apex. Head as wide as holotype from the Hungarian National Museum, Budapest, and pronotum, crown twice as long medially as next to eye. illustrated the ovipositor. No male specimens that could be Male 2S apodemes (Fig. 154) slender, bowed laterad, ex- considered conspecific have been found. tended slightly beyond posterior margin of sternite IV. Pygofer emargination parallel sided (Fig. 100); tergite weakly developed Columbonirvana craigi sp. nov. and poorly sclerotized; dorsal sclerotized area of lobe gradu- Figs 4, 95-98, 153 ally tapered distad, apex strongly hooked ventrad but not extended beyond apical margin; ventral appendage gradually Description. Length of male 4.0 mm. Dorsum (Fig. 4) curved dorsad, apex extended posterolaterad (Fig. 99). Anal brown with extensive symmetrical pale markings; anterior hook (Fig. 99) digitiform, extended ventrad, apex hooked me- margin of head with pair of thin red transverse bands between sad. Subgenital plate (Fig. 99) with 3 macrosetae near mid- eyes; crown brown medially with broad white anterior and length, without stout apical setae, apex gradually curved posterior margins; pronotum with white median crescent, pos- posterodorsad. Style apophysis (Figs 101 and 102) slightly terior margin broadly white; mesonotum and scutellum white broadened preapically, apex tapered, curved ventrolaterad. except midline anterad of scutellar suture and basolateral tri- Connective (Fig. 102) U-shaped. Aedeagus (Figs 101 and 102) angles brown; forewing with extensive large symmetrical white with preatrium distinctly shorter than shaft, slightly broad- markings. Head subequal to pronotum in width, crown with ened basally in ventral view; shaft tubular, somewhat com- median length much less than twice length next to eye. pressed basally, tapering toward apex in lateral view; without Male 2S apodemes (Fig. 153) slender, divergent, extended paired ventral processes, with indistinct ventrolateral flange beyond posterior margin of sternite IV. Pygofer dorsal emar- extended to apex; apex tubular, subtruncate. gination (Fig. 96) quadrate; tergite 1/3 length of distal lobe; Material examined. Holotype male, PERU: Pasco, Yanachaga- dorsal sclerotized area of lobe relatively broad, weakly falcate Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 Octo- with angulate dorsal preapical projection, not produced be- ber 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio yond apical margin; ventral appendage gradually curved dor- San Alberto [USML]. Paratypes: 4 males, same data [INHS]. sad, apex curved posteromesad (Figs 95 and 96). Anal hook Etymology. This species is named in honor of my brother, (Fig. 95) well developed, triangular. Subgenital plate (Fig. 95) David A. Dietrich. with 3 macrosetae near midlength and few short, stout preapical setae, apex bent posterodorsad. Style apophysis (Figs 97 and Columbonirvana edgari sp. nov. 98) parallel sided through most of length, apex blunt, curved ventrolaterad, extended slightly mesad in ventral view. Con- Figs 6, 103-106, 155 nective (Fig. 98) broadly V-shaped. Aedeagus (Figs 97 and 98) Description. Length of male 4.0 mm. Dorsum (Fig. 6) with preatrium distinctly shorter than shaft, nearly parallel medium brown with extensive symmetrical pale markings; sided in ventral view; shaft slender, tubular, nearly straight, anterior margin of head with pair of thin red transverse bands curved slightly dorsad near apex; with pair of slender ventral between eyes; crown brown with anterior and posterior mar- processes arising near base and extended nearly to apex of shaft, gins yellow; pronotum with large white median spot, poste- only slightly divergent from shaft; shaft apex slightly com- rior margin broadly white; mesonotum and scutellum yellow pressed, rounded in lateral view, truncate in ventral view. except midline and basolateral triangles brown; forewing with

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 527 extensive large symmetrical white markings. Head subequal to preapical setae, apex bent posterodorsad. Style apophysis (Figs pronotum in width, crown with median length much less than 105 and 106) acuminate, apex curved ventrolaterad. Connec- twice length next to eye. tive (Fig. 106) V-shaped. Aedeagus (Figs 105 and 106) with Male 2S apodemes (Fig. 155) broad, parallel sided, ex- preatrium nearly as long as shaft, parallel sided in ventral view; tended nearly to posterior margin of sternite IV. Pygofer emar- shaft tubular, very slender, slightly sinuate in lateral view; with gination quadrate (Fig. 104), tergite nearly 1/2 length of lobe; pair of acuminate ventral processes arising near midlength, dorsal sclerotized area of lobe with digitiform distal section bent curved away from, then toward shaft in ventral and lateral views; ventrad at slightly less than 90° angle, extended slightly beyond apex obliquely rounded in lateral view, blunt in ventral view. margin; ventral appendage gradually curved dorsad, apex curved Material examined. Holotype male, PERU: Pasco, Villa slightly mesad (Figs 103 and 104). Anal hook (Fig. 103) slender, Rica, 1400m, 10°43’21"S 75°15’43"W, 21 October 2002, C.H. bifurcate, with slender dorsomedial branch and longer, broader Dietrich, vacuum, 2-28-1[USML]. ventral branch curved dorsad in lateral view. Subgenital plate Etymology. This species is named in honor of my father, (Fig. 103) with 3 macrosetae near midlength and few short, stout Edgar H. Dietrich.

69 71 73

70 72 74

77 75 79

76 78 80

85 81 83

86 82 84

Figures 69-86. Wings, photographed with transmitted light to highlight venation. (69-70) Columbonirvana aidani sp. nov.) forewing and hind wing; (71-72) Eualebra jenniferae sp. nov., same; (73-74) Eualebra michaelorum sp. nov., same; (75-76) Euzyginella marki sp. nov., same, absence of distal part of vein R in hind wing is aberrant; (77-78) Neozyginella jenncraftae sp. nov., same; (79-80) Pseudhadina amazonica sp. nov., same; (81-82) Pseudozyginella grahami sp. nov., same; (83) P. davei sp. nov., hind wing; (84) Tahurella lynnae sp. nov., forewing; (85-86) Tahurella heatherae sp. nov.) forewing and hind wing. (iac) Inner apical cell, (smv) submarginal vein.

ZOOLOGIA 30 (5): 519–568, October, 2013 528 C.H. Dietrich

87 88 90

91 92 94

98 95

101

102 99 100

105

106 103 104

Figure 87-106. Columbonirvana, male genitalia. (87-90) C. aidani sp. nov.: (87) genital capsule) lateral view; (88) pygofer and basal sclerites of anal tube) dorsal view; (89) genitalia) lateral view; (90) genitalia, ventral view, only right style shown. (91-94) C. ameliae sp. nov., same. (95-98) C. craigi sp. nov., same. (99-102) C. davidi sp. nov., same; (103-106) C. edgari sp. nov., same.

Columbonirvana ernae sp. nov. apex. Head subequal to pronotum in width, crown less than twice as long medially as next to eyes. Figs 7, 107-110, 156 Male 2S apodemes (Fig. 156) long, slightly divergent, par- Description. Length of male 3.8 mm. Dorsum (Fig. 7) allel sided, extended nearly to posterior margin of sternite V. brown, without distinct pale markings; head anterior margin Pygofer emargination with sides divergent posteriorly (Fig. 108), with pair of thin red transverse bands between eyes; forewing base truncate; tergite ca. 1/3 length of distal lobe; dorsal sclero- veins yellow distally, apical and anteapical cells hyaline in tized area of lobe moderately broad, gradually tapered distad, apex middle, discal and brachial cell each with hyaline spot near hooked ventrad and extended slightly beyond margin; ventral

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 529 appendage acuminate, gradually curved dorsad, bowed slightly Etymology. This species is named in honor of my nephew, mediad (Figs 107 and 108). Anal hook (Fig. 107) short, tapered, Joshua Dietrich. extended ventrad, apex weakly bidentate in dorsal view (Fig. 108). Subgenital plate (Fig. 107) with 3 macrosetae near midlength and Columbonirvana margaretae sp. nov. few smaller stout setae distally, apex gradually curved dorsad. Figs 9, 115-118, 158 Style (Figs 109 and 110) somewhat expanded beyond midlength, Description. Length of male 3.9 mm. Dorsum (Fig. 9) apex hooked ventrolaterad. Connective (Fig. 5d) V-shaped, stem mostly dark brown; head anterior margin with pair of thin red poorly developed. Aedeagus (Figs 109 and 110) with preatrium transverse bands between eyes; forewing with oblique pale or- nearly as long as shaft; shaft very slender, sinuate in lateral view, ange mark along posterior part of brochosome field and smaller with pair of long slender straight processes arising near base and orange areas more distad along costal margin, veins orange, extended distad slightly more than half distance to shaft apex, bordered with brown, central areas of distal cells hyaline; ven- divergent from shaft in lateral view, parallel in ventral view; shaft ter of head and thorax stramineous except mesosternum dark apex obliquely truncate in lateral view. brown, abdomen brown ventrally except submedial areas of Material examined. Holotype male, PERU: Pasco, Villa sternite III, posterior margins of pregenital sternites and basal Rica, 1400m, 10°43’21"S 75°15’43"W, 21 October 2002, C. H. 2/3 of subgenital plate stramineous. Head narrower than Dietrich, vacuum, 2-28-1 [USML]. Paratype: 1 male, PERU: Pasco, pronotum, crown less than twice as long medially as next to Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, eyes. 10-13 October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise Male 2S apodemes (Fig. 158) weakly divergent, extended trap across Rio San Alberto [INHS]. to posterior margin of sternite IV. Pygofer emargination paral- Etymology. This species is named in honor of my niece, lel sided, base irregularly rounded (Fig. 116); tergite ca. 1/4 Erna Dietrich. length of distal lobe; dorsal sclerotized area of lobe slender, Columbonirvana joshuai sp. nov. parallel sided through most of length, apex tapered and decurved, extended slightly beyond apical margin; ventral ap- Figs 8, 111-114, 157 pendage gradually curved dorsad, apex directed posteromesad Description. Length of male 3.8 mm. Dorsum (Fig. 8) (Figs 115 and 116). Anal hook (Fig. 115) with short antero- brown, without distinct pale markings; head anterior margin ventral process, longer irregularly sinuate ventral extension with pair of thin red transverse bands between eyes; forewing hooked mesad. Subgenital plate (Fig. 115) with 3-4 macrosetae veins yellow distally, apical and anteapical cells hyaline in near midlength. Style (Figs 117 and 118) apophysis with slight middle, discal and brachial cell each with hyaline spot near preapical gibbosity, apex tapered blunt, weakly hooked apex. Head subequal to pronotum in width, crown approxi- ventrolaterad. Connective (Fig. 118) U-shaped with arms mately twice as long medially as next to eyes. strongly bent mesad distally. Aedeagus (Figs 117 and 118) with Male 2S apodemes (Fig. 157) parallel sided, parallel to preatrium much shorter than shaft, narrowest at base in ven- each other, extended nearly to posterior margin of sternite IV. tral view; shaft weakly compressed, nearly straight, somewhat Pygofer emargination with sides divergent posteriorly (Fig. 112), broadened medially, with indistinct angulate dorsolateral flange base truncate; tergite ca. 1/2 length of distal lobe; dorsal scle- preapically without paired ventral processes, apex expanded rotized area of lobe moderately broad, gradually tapered dis- in lateral view, compressed and subtruncate. tad, apex abruptly hooked ventrad and extended slightly Material examined. Holotype male, PERU: Pasco, Yanacha- beyond margin; ventral appendage acuminate, evenly curved ga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 dorsad, apex curved slightly mesad (Figs 111 and 112). Anal October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across hook (Fig. 111) relatively long, tapered, extended ventrad, apex Rio San Alberto [USML]. hooked mesad in dorsal view. Subgenital plate (Fig. 111) with Etymology. This species is named in honor of my grand- 3 macrosetae near midlength, apex gradually curved dorsad. mother, Margaret R. Baka. Style (Figs 113 and 114) somewhat expanded beyond midlength, apex hooked ventrolaterad. Connective (Fig. 114) Columbonirvana matthewi sp. nov. V-shaped, stem absent. Aedeagus (Figs 113 and 114) with Figs 10, 61, 119-122, 159 preatrium nearly as long as shaft; shaft very slender, sinuate in Description. Length of male 3.7-4.0 mm. Dorsum (Fig. lateral view, with pair of long tapered evenly curved processes 10) mostly brown, venter pale yellow except medial part of arising near base and extended distad half distance to shaft frontoclypeus, gena below eye, and mesepisternum brown. apex, divergent from shaft in lateral view, parallel in ventral Head anterior margin with pair of transverse red bands, crown, view; shaft apex obliquely truncate in lateral view. pronotum, mesonotum and scutellum brown infused with Material examined. Holotype male, PERU: Pasco, Villa stramineous; forewing with indistinct stramineous band ex- Rica, 1400m, 10°43’21"S 75°15’43"W, 21 October 2002, C. H. tended from base of clavus across mesonotum and another Dietrich, vacuum, 2-28-1 [USML].

ZOOLOGIA 30 (5): 519–568, October, 2013 530 C.H. Dietrich larger transcommisural band near midlength, veins mostly near midlength, without stout apical setae, apex gradually orange, cells hyaline medially, costal margin with orange pig- curved dorsad. Style (Figs 125 and 126) apex somewhat ex- ment in distal third. Head subequal to pronotum in width, panded near apex, apex hooked ventrolaterad. Connective (Fig. crown only slightly longer medially than next to eye. 126) broadly V-shaped. Aedeagus (Figs 125 and 126) with Male 2S apodemes (Fig. 159) weakly divergent, extended preatrium nearly as long as shaft; shaft compressed, relatively to near midlength of sternite IV. Pygofer emargination with broad in lateral view, without paired ventral processes, with lateral margins concave, widest posteriorly (Fig. 120); tergite weak ventrolateral flange; apex tubular and subtruncate. 1/5 length of distal lobe; dorsal sclerotized area of lobe slen- Material examined. Holotype male, PERU: Pasco, der, tapered distally, apex curved ventrad and extended slightly Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, beyond margin; ventral appendage with weakly sclerotized 10-13 October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise jointlike area near midlength, distal portion strongly curved trap across Rio San Alberto [USML]. Paratype: 1 male, same dorsad (Figs 119 and 120). Anal hook (Fig. 119) somewhat elon- data [INHS]. gate, divided preapically into digitiform posterolateral and flat- Etymology. This species is named in honor of my nephew, tened anteromedial process. Subgenital plate (Fig. 119) with Noah Dietrich. 3-4 macrosetae near midlength, apex gently curved dorsad. Style (Figs 121 and 122) apophysis parallel sided, apex tapered, blunt, Columbonirvana schulzi sp. nov. weakly hooked ventrolaterad. Connective (Fig. 122) V-shaped Figs 12, 127-130, 161 with tiny digitiform medial lobe. Aedeagus (Figs 121 and 122) Description. Length of male 4.6 mm. Dorsum (Fig. 12) with preatrium much shorter than shaft, only slightly expanded dark brown with large symmetrical yellow markings. Anterior basally in ventral view; shaft tubular, slender, with indistinct margin of head with two narrow transverse red bands; crown ventrolateral flange ending 1/4 distance from apex, without with posterior margin infused with yellow medially. Pronotum paired ventral processes, apex compressed with broad triangu- with large transverse yellow crescent. Mesonotum yellow ex- lar ventral keel. cept dark brown anterolateral triangles; scutellum with ante- Material examined. Holotype male, PERU: Pasco, Yanachaga- rior half yellow. Forewing clavus with large diamond-shaped Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 Octo- transcommisural marking; veins mostly orange; anteapical and ber 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio apical cells each with medial area hyaline; brachial and discal San Alberto [USML]. Paratypes: 3 males, same data [INHS]. cell each with hyaline area near apex. Head as wide as Etymology. This species is named in honor of my brother, pronotum, crown twice as long medially as next to eye. Matthew M. Dietrich. Male 2S apodemes (Fig. 161) broad, tapered, not ex- Columbonirvana noahi sp. nov. tended beyond posterior margin of sternite III. Pygofer emargination nearly parallel sided (Fig. 128); tergite weakly Figs 11, 59, 123-126, 160 developed and poorly sclerotized; dorsal sclerotized area of Description. Length of male 4.0-4.1 mm. Dorsum (Fig. lobe abruptly tapered distad, apex hooked ventrad at 90° angle 11) brown marked with white and orange; head anterior mar- but not extended beyond apical margin; ventral appendage gin white; pronotum with anterior margin orange; mesonotum areolate distally, gradually curved dorsad, apex extended and scutellum orange; forewing base brown infused with or- posterolaterad well beyond pygofer apex (Fig. 127). Anal hook ange, costal area with two large white oblique markings; veins (Fig. 127) short, weakly bidentate, extended anteromesad. orange, cells mostly dark brown distally; venter mostly tan, Subgenital plate (Fig. 127) with 4 macrosetae near midlength, mesosternum dark brown, abdomen tan ventrally, medial and without stout apical setae, apex gradually curved posterodor- anterior parts of sternites and apex of subgenital plate brown. sad. Style (Figs 129 and 130) apophysis abruptly narrowed Head subequal to pronotum in width, crown less than twice as preapically, apex tapered, curved ventrolaterad. Connective long medially as next to eyes. (Fig. 130) Y-shaped; stem broad, slightly shorter than arms. Male 2S apodemes (Fig. 160) straight, slightly divergent, Aedeagus (Figs 129 and 130) with preatrium shorter than shaft, extended slightly beyond posterior margin of sternite IV. parallel sided basally in ventral view; shaft strongly com- Pygofer emargination parallel sided, base broadly V-shaped (Fig. pressed, in lateral view nearly straight, parallel sided through 124); tergite short, ca. 1/6 length of distal lobe; dorsal sclero- most of length, apex broadened and subtruncate; without pro- tized area of lobe broad, gradually tapered distad, apex strongly cesses or flanges. hooked ventrad but not extended beyond apical margin; ven- Material examined. Holotype male, PERU: Pasco, tral appendage broadened medially and abruptly narrowed and Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, somewhat twisted preapically, curved dorsad, apex bent 10-13 October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise posterad (Figs 123 and 124). Anal hook (Fig. 123) bifid with trap across Rio San Alberto [USML]. two processes digitiform and subequal in length, extended Etymology. This species is named in honor of my brother- ventromesad. Subgenital plate (Fig. 123) with 3-4 macrosetae in-law, David Schulz.

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 531

109

107 108 110

113

111 114 112

117

118 115 116

121

122 119 120 125

126 123

124 Figure 107-126. Columbonirvana, male genitalia. (107-110) C. ernae sp. nov.: (107) genital capsule) lateral view; (108) pygofer and basal sclerites of anal tube, dorsal view; (109) genitalia, lateral view; (110) genitalia, ventral view, only right style shown; (111-114) C. joshuai sp. nov., same; (115-118) C. margaretae sp. nov., same; (119-122) C. matthewi sp. nov., same; (123-126) C. noahi sp. nov., same.

Columbonirvana thomasi sp. nov. line; brachial and discal cell each with hyaline area near apex. Head as wide as pronotum, crown twice as long medially as Figs 13, 131-134, 162 next to eye. Description. Length of male 3.4-3.9 mm. Dorsum (Fig. Male 2S apodemes (Fig. 162) robust, parallel-sided, ex- 13) dark brown with large symmetrical yellow markings. Ante- tended beyond posterior margin of sternite IV. Pygofer emar- rior margin of head with two narrow transverse red bands; gination nearly parallel sided (Fig. 132); tergite weakly developed crown with posterior margin infused with yellow medially. and poorly sclerotized; dorsal sclerotized area of lobe broadend Pronotum with large transverse yellow crescent. Mesonotum preapically, abruptly bent ventrad, apex tapered, not extended and scutellum without pale markings. Forewing clavus with beyond apical margin; ventral appendage areolate distally, large pentagonal transcommisural marking; veins mostly or- abruptly bent dorsad near midlength, apex not extended be- ange; anteapical and apical cells each with medial area hya- yond pygofer apex (Fig. 131). Anal hook (Fig. 131) short, sinu-

ZOOLOGIA 30 (5): 519–568, October, 2013 532 C.H. Dietrich ate in lateral view, hooked mesad. Subgenital plate (Fig. 131) Etymology. The species name refers to the small dorsal with 3 macrosetae near midlength, without stout apical setae, preapical tooth (tubercle) on the male pygofer lobe. apex gradually curved posterodorsad. Style (Figs 133 and 134) apophysis narrowed preapically, apex tapered, curved Columbonirvana tumida sp. nov. ventrolaterad. Connective (Fig. 134) V-shaped. Aedeagus (Figs Figs 15, 139-142, 164 133 and 134) with preatrium shorter than shaft, slightly broadened basally in ventral view; shaft strongly compressed, in lat- Description. Length of male 4.2 mm. Dorsum (Fig. 15) mostly brown, venter pale yellow except mesepisternum brown. eral view nearly straight, gradually tapered through most of Head anterior margin with pair of transverse red bands, crown, length, apex broadened and obliquely truncate; with pair of preapical ventrolateral flanges and numerous preapical denticuli. pronotum, mesonotum and scutellum brown infused with stramineous; forewing with indistinct stramineous band ex- Material examined. Holotype male, PERU: Pasco, Yanachaga- tended from base of clavus across mesonotum and another Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 Octo- ber 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio larger transcommisural band near midlength, veins mostly orange, cells hyaline medially, costal margin with orange pig- San Alberto [USML]. Paratypes: 4 males, same data [INHS]. ment in distal third. Head subequal to pronotum in width, Etymology. This species is named in honor of my brother- in-law, Thomas Braxton. crown only slightly longer medially than next to eye. Male 2S apodemes (Fig. 164) relatively slender, parallel- Columbonirvana tuberculata sp. nov. sided, extended slightly beyond sternite IV. Pygofer emargination with lateral margins slightly divergent posteriorly (Fig. Figs 14, 135-138, 163 140); tergite weakly sclerotized; dorsal sclerotized area of lobe Description. Length of male 4.3 mm. Dorsum (Fig. 14) constricted preapically, apex curved ventrad and extended mostly brown, venter pale yellow except medial part of slightly beyond margin; ventral appendage areolate distally, frontoclypeus, gena below eye, and mesepisternum brown. abruptly bent dorsad near midlength and extended postero- Head anterior margin with pair of transverse red bands, crown, dorsad slightly beyond dorsal pygofer margin (Figs 139 and 140). pronotum, mesonotum and scutellum brown infused with Anal hook (Figs 139 and 140) short, sinuate in lateral view, stramineous; forewing with indistinct stramineous band ex- hooked mesad. Subgenital plate (Fig. 139) with 3-4 macrosetae tended from base of clavus across mesonotum and another near midlength, apex gently curved dorsad. Style apophysis (Figs larger transcommisural band near midlength, veins mostly 141 and 142) parallel sided, apex tapered, blunt, weakly hooked orange, cells hyaline medially, costal margin with orange pig- ventrolaterad. Connective (Fig. 142) V-shaped. Aedeagus (Figs ment in distal third. Head subequal to pronotum in width, 141 and 142) with preatrium short, slightly expanded basally in crown only slightly longer medially than next to eye. ventral view; shaft slender, somewhat compressed distally, apex Male 2S apodemes (Fig. 163) robust, parallel sided, ex- slightly broadened and obliquely rounded in lateral view; with tended to posterior margin of sternite IV. Pygofer emargination pair of ventrolateral flanges broadening distad in ventral view with lateral margins slightly divergent posteriorly (Fig. 136); and each giving rise to spine extended distad and slightly diver- tergite weakly sclerotized; dorsal sclerotized area of lobe with gent from shaft, not reaching shaft apex. dorsal tooth adjacent to anal hook, apex curved ventrad and Material examined. Holotype male, PERU: Pasco, Yanacha- extended slightly beyond margin; ventral appendage areolate ga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 distally, abruptly bent dorsad near midlength and extended October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across posterodorsad slightly beyond dorsal pygofer margin (Figs 135 Rio San Alberto [USML]. and 136). Anal hook (Fig. 135) short, weakly bidentate, curved Etymology. The species name refers to the tumid (swol- anteromesad. Subgenital plate (Fig. 135) with 3-4 macrosetae len) medial section of the aedeagal shaft. near midlength, apex gently curved dorsad. Style apophysis (Figs 137 and 138) parallel sided, apex tapered, blunt, weakly hooked Columbonirvana urodolobrata sp. nov. ventrolaterad. Connective (Fig. 138) V-shaped, stem not devel- Figs 16, 143-146, 165 oped. Aedeagus (Figs 137 and 138) with preatrium much shorter than shaft, slightly expanded basally in ventral view; shaft tu- Description. Length of male 4.7 mm. Dorsum (Fig. 16) bular, slender, somewhat compressed distally, apex slightly mostly dull orange-brown, venter pale yellow except broadened and obliquely truncate in lateral view; with pair of mesepisternum brown. Head anterior margin with pair of trans- slender ventral processes arising near midlength, extended dis- verse red bands, crown, pronotum, mesonotum and scutellum tad and slightly divergent from shaft, not reaching shaft apex. brown with few diffuse white markings; forewing with narrow Material examined. Holotype male, PERU: Pasco, Yanacha- transcommisural white band near midlength of clavus, veins ga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 mostly orange, cells hyaline medially, costal margin with or- October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across ange pigment in distal third. Head subequal to pronotum in Rio San Alberto [USML]. width, crown only slightly longer medially than next to eye.

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 533

129

130 128

127 133

134 132

131 137

136 138

135

141

142 140

139 145

146

144 143

149

150 148 147

Figures 127-150. Columbonirvana, male genitalia. (127-130) C. schulzi sp. nov.: (127) genital capsule) lateral view; (128) pygofer and basal sclerites of anal tube, dorsal view; (129) genitalia, lateral view; (130) genitalia, ventral view, only right style shown; (131-134) C. thomasi sp. nov., same; (135-138) C. tuberculata sp. nov., same; (139-142) C. tumida sp. nov., same; (143-146) C. urodolobrata sp. nov., same; (147-150) C. vatia sp. nov., same.

ZOOLOGIA 30 (5): 519–568, October, 2013 534 C.H. Dietrich

Male 2S apodemes (Fig. 165) robust, short, tapered, ex- expanded basally in ventral view; shaft slender, tubular, nearly tended slightly beyond sternite III. Pygofer emargination with straight, with pair of rounded, finely serrate ventrolateral lateral margins slightly divergent posteriorly(Fig. 144); tergite flanges arising near apex; shaft apex compressed, slightly ex- sclerotized in basal fifth; dorsal sclerotized area of lobe very panded, obliquely rounded in lateral view. slender, apex abruptly curved ventrad and extended slightly Material examined. Holotype male, PERU: Pasco, Yanachaga- beyond margin; ventral appendage areolate distally, abruptly Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 Octo- bent dorsad near midlength, bowed mesad and extended ber 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio posterodorsad slightly beyond dorsal pygofer margin (Figs 143 San Alberto [USML]. Paratypes: 2 males, same data [INHS]. and 144). Anal hook (Fig. 143) short, sinuate in lateral view, Etymology. The species name is derived from the Latin abruptly constricted preapically, hooked mesad. Subgenital word vatius, meaning “bent”, and refers to the sharply bent plate (Fig. 143) with 3 macrosetae near midlength, apex gently pygofer appendages. curved dorsad. Style apophysis (Figs 145 and 146) parallel sided, apex tapered, blunt, weakly hooked ventrolaterad. Connective Eualebra Baker (Fig. 146) V-shaped. Aedeagus (Figs 145 and 146) with preatrium Eualebra Baker 1899: 402. shorter than shaft, tapered basally in ventral view; shaft strongly Type species: E. smithii Baker 1899. compressed, nearly straight, slightly tapered in lateral view; Diagnosis. Small to medium sized, strongly depressed, apex strongly expanded with acute posterodorsal angle and somewhat ovoid leafhoppers (Figs 18-38) broadest across rounded posteroventral lobe; without processes. midlength of resting forewings in dorsal view. Color pale yel- Material examined. Holotype male, PERU: Pasco, Yanacha- low to dark brown with symmetrical white, yellow, orange, ga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 brown, and/or black markings dorsally. October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Head (Figs 18-38) narrower than pronotum, depressed, Rio San Alberto [USML]. produced, anterior margin parabolic in dorsal view; crown Etymology. The species name combines uro, meaning slightly convex, declivous, glabrous, coronal suture restricted “tail”, with dolobrata, meaning “hatchet”, referring to the to posterior third; face horizontal (Figs 62-63); rostrum not hatchetlike apex of the aedeagus. extended beyond front trochanters; lower part of face sexually Columbonirvana vatia sp. nov. dimorphic, male anteclypeus inflated and expanded laterad, parallel-sided and weakly convex in female; lorum very nar- Figs 17, 147-150, 166 row; frontoclypeus flat or concave medially; lateral frontal su- Description. Length of male 3.4-3.9 mm. Dorsum (Fig. tures not extended dorsad of antennal pits; antennal ledges 17) dark brown with symmetrical white markings; anterior curved, nearly vertical; antennae slightly longer than head margin of head with pair of thin red transverse bands between width; ocelli vestigial, on crown margin approximately mid- eyes; crown white bordered anteriorly with brown; pronotum way between eye and midline. Pronotum (Figs 18-38) with lat- white medially with anterior and lateral margins brown and eral margins long, divergent in dorsal view, distinctly carinate, with brown transverse submarginal band posteriorly; carina even with posterior corner of eye. Front femur with AM1 mesonotum white medially, anterolateral triangles and apex enlarged and situated on ventral margin; intercalary row with of scutellum brown; forewing clavus with large white few fine setae, basal seta larger than others; PV1 well devel- transcommisural spot, veins orange bordered with brown in oped; tibia somewhat flattened dorsally, AD and PD without distal half; venter of thorax pale yellow except mesosternum preapical macrosetae. Hind femur macrosetae 2+1+1; tibia row orange brown. Head as wide as pronotum, crown slightly longer AV with 3-4 macrosetae near apex. Forewing (Figs 71 and 73) medially than next to eye. with RA slightly to strongly reflexed, RP and MA separate and Male 2S apodemes (Fig. 166) robust, subparallel, extended connected by crossvein or confluent for short distance to posterior margin of sternite IV. Pygofer dorsal emargination preapically. Hind wing (Figs 72 and 74) with anterior branch quadrate, lateral margins constricted medially (Fig. 148); terg- of R absent, veins RP and MA separate, joined by crossvein; ite 1/3 length of distal lobe; dorsal sclerotized area of lobe fal- MP and CuA separate, joined by oblique m-cu crossvein; sub- cate, produced slightly beyond apical margin; ventral marginal vein extended from apex of RP to jugal lobe. appendage gradually curved dorsad, apex curved dorsolaterad Male 2S apodemes (Fig. 242-260) well developed, usu- (Figs 147 and 148). Anal hook (Fig. 147) long, tapered, sinu- ally joined at base by transverse bridge, variable interspecifically ate, apex hooked mesad. Subgenital plate (Fig. 147) with 3 in length and orientation. Pygofer (Figs 167 and 168) slightly macrosetae near midlength and few short, stout preapical se- to strongly emarginate dorsally; tergite variable interspecifically tae, apex bent dorsad. Style apophysis (Figs 149 and 150) in length and shape; posterior lobe with or without dorsal pro- slightly sinuate, apex tapered, curved ventrolaterad then cess, side not or only weakly produced posteriorly, ventral ap- hooked mesad. Connective (Fig. 150) U-shaped. Aedeagus (Figs pendage elongate, curved dorsad, usually areolate distally. Anal 149 and 150) with preatrium much shorter than shaft, not tube and appendages highly variable interspecifically, append-

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 535

151 152 154 153

155 158 157 156

159 160 162

161

163

164 165

Figures 151-166. Columbonirvana, base of male abdomen, ventral view. (151) C. aidani sp. nov.; (152) C. ameliae sp. nov.; (153) C. craigi sp. nov.; (154) C. davidi sp. nov.; (155) C. edgari sp. nov.; (156) C. ernae sp. nov.; (157) C. joshuai sp. nov.; (158) C. margaretae sp. nov.; (159) C. matthewi sp. nov.; (160) C. noahi sp. nov.; (161) C. schulzi sp. nov.; (162) C. thomasi sp. nov.; (163) C. tuberculata sp. nov., right apodeme broken; (164) C. tumida sp. nov.; (165) C. urodolobrata sp. nov.; (166) C. vatia sp. nov.

age usually well developed, arising from base of anal tube (seg- turing strigate; second valvulae (Figs 362, 370, and 374) vari- ment X), articulated to posterodorsal margin of pygofer later- able interspecifically in shape and dentition. ally and to dorsal connective medially, usually with at least Remarks. DIETRICH (2004) noted that Eualebra and one separate projection extended mesad and/or ventrad. Scle- Columbonirvana share a unique hind wing venational pattern rotized dorsal connective present (Fig. 214), usually U-shaped different from that of other Typhlocybinae and therefore (rarely platelike) and articulated between dorsal apodeme of treated both genera as unplaced to tribe within Typhlocybinae. aedeagus and paired basal processes of anal tube. Valve very Based on the revised tribal diagnosis given above, both of these short, rectangular, fused to pygofer. Subgenital plate (Fig. 167) genera are now included in Typhlocybini (also see Remarks depressed, ovoid in ventral view, distal half tapered with 5 or under Columbonirvana). Species of Eualebra resemble species of more macrosetae arranged in lateral band, fine setae short and the Old World genus Eurhadina Haupt in their ovoid shape in sparse on dorsal surface, apex rounded or subtruncate. Style dorsal view and strongly depressed body form. However (Figs 169 and 170) apodeme short, broad; apophysis elongate, Eurhadina differs in having the structure of the lower part of slender, apex simple, tapered and slightly hooked, without con- the face similar in males and females, the forewing narrow dis- spicuous setae. Connective (Fig. 170) U-shaped with pair of tally, the male subgenital plates usually with a single subbasal posterodorsal lobes articulated to aedeagus. Aedeagus (Figs 169 macroseta, and the pygofer lacking an elongate ventral pro- and 170) with preatrium well developed; dorsal apodeme short, cess. Also, like all other known Old World Typhlocybini, lobelike; shaft short, tubular, with or without processes; go- Eurhadina lacks a submarginal vein at the apex of the hind nopore apical. wing. Species of Eualebra occur in both lowland Neotropical Female with posterior margin of sternite VII (Fig. 361) rainforest and premontane cloud forests. angulately produced medially; ovipositor not extended beyond Eualebra previously included five species. YOUNG (1952) pygofer; first valvulae Figs 359, 364, and 369) with dorsal sculp- excluded E. notata Baker from Guatemala, noting that the hind

ZOOLOGIA 30 (5): 519–568, October, 2013 536 C.H. Dietrich wing venation of the female holotype is consistent with that 5. Forewing clavus with small round white spots (Fig. 33); of tribe Dikraneurini, but did not suggest an appropriate ge- aedeagus with paired lateral flanges extended full length neric placement for the species so it remains incertae sedis. Ex- of shaft (Fig. 10t); female sternite VII with posterior margin amination of the holotype of Eualebra rubra Van Duzee from broadly but shallowly produced (Fig. 18p) ...... Jamaica, deposited in the California Academy of Sciences col- ...... magaretannae sp. nov. lection, indicates that it belongs in the subgenus Alconeura 5’. Forewing clavus with irregular white vermiform areas (Fig. (Hyloidea) of tribe Dikraneurini. Therefore, A. (H.) rubra (Van 31); aedeagus without paired lateral flanges (Fig. 10p); Duzee) is proposed as a new combination. As a result, only female sternite VII with short median lobe . smithii Baker three taxa previously included, E. smithii Baker, E. reticulata 6. Crown and pronotum with pair of orange or brown longitu- Osborn, and E. rufoornata (Stål), all described from Brazil, are dinal stripes extended onto pronotum (Fig. 20); length >4.0 retained in Eualebra. Seventeen new species, described below, mm (dorisae species group) ...... 7 are also included. Based on the species known at present, the genus appears to be restricted to South America. 6’. Crown and pronotum without pair of longitudinal stripes Eualebra, as presently defined, is somewhat heteroge- or, if stripes present, length <3.0mm ...... 9 neous morphologically, comprising species that range in size 7. Dorsum predominantly bright yellow (Fig. 22); male tergite from 2.7 to 4.7 mm and exhibit a variety of color patterns, IX with narrow, parallel-sided posterior emargination (Fig. shapes, and male genitalia configurations. Given the still poor 184); aedeagus without processes (Fig. 185) ...... state of knowledge of the genus, and particularly considering ...... patriciae sp. nov. that females are known for only a few species, I have opted 7’. Dorsum predominantly brown; male tergite IX with broad, not to recognize additional genera or subgenera but, rather, obtusely angulate emargination (Fig. 8j); aedeagus with pair place the known species into five informal species groups that of processes (Fig. 178) ...... 8 may represent distinct lineages. Recognition of some of these 8. Crown width between eyes much less than 2X median length groups as separate genera may be warranted once the fauna (Fig. 21); aedeagus with paired processes arising near apex becomes better known. In particular, the ovipositors of the (Fig. 182) ...... jessicae sp. nov. few female specimens available for study exhibit considerable variation and may provide genus-level characters. 8’. Crown width between eyes ~2X median length (Fig. 20); aedeagus with paired processes arising near base (Fig. 178) Key to species of Eualebra ...... dorisae sp. nov. 1. Body length >4.5 mm, head in profile with anterior margin 9. Pronotum with narrow white median cruciform or trident- thick (Fig. 63), hind tibia row AV with three macrosetae shaped mark surrounded by brown (Fig. 18); length <3 mm (michaelorum species group) ...... 2 (barbarae species group) ...... 10 1’. Body length < 4.5 mm, head in profile with anterior margin 9’. Pronotum with white marking, if present, broad and not thin (Fig. 62), hind tibia row AV usually with four cruciform or trident-shaped; length >3.0 mm ...... 11 macrosetae ...... 4 10. Male pygofer with elongate dorsal spine and short, bifurcate 2. Dorsal coloration mostly pale yellow with few brown or black ventral process (Fig. 167); aedeagus strongly curved in la- markings (Fig. 34) ...... 3 teral view (Fig. 169) ...... barbarae sp. nov. 2’. Dorsal coloration mostly brown with smaller pale markings 10’. Male pygofer with dorsal spine absent, ventral process long (Fig. 36) ...... michaelorum sp. nov. and acuminate (Fig. 8e); aedeagus nearly straight in lateral 3. Pronotum with narrow median posterior macula; forewing view (Fig. 173) ...... charlesi sp. nov. clavus unmarked (Fig. 35) ...... dorothyae sp. nov. 11. Crown brown with pale marking, if present, restricted to 3’. Pronotum with broad transverse macula on posterior narrow area near apex (Fig. 38) ...... 12 margin; forewing clavus with distinct transverse brown 11’. Crown with large medial lobed or wedge-shaped pale area macula at midlength (Fig. 34) ...... alberti sp. nov. (Fig. 23, 29) (gingerae species group, part) ...... 14 4. Dorsum color pale stramineous marked with numerous 12. Pronotum uniformly brown, without white spots (Fig. 30); minute brown spots (Fig. 31), forewing with irregular cream- forewing clavus with pair of triangular yellow transcommi- colored spots, crown unmarked or with small symmetrically sural markings (gingerae species group, part) ...... arranged pale orange markings (smithii species group) .. 5 ...... jenniferae sp. nov. 4’. Dorsal color pattern not consisting of small brown spots on 12’. Pronotum brown with narrow white posterior band and a stramineous background, forewing with cream-colored pair of white submedial spots; forewing clavus with spots absent or large and symmetrically arranged, crown transcommisural markings white (Fig. 37) ...... with orange markings, if present, large ...... 6 ...... (leoni species group) ... 13

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 537

13. Forewing clavus with three small white transcommisural crown mostly brown with midline and lateral margins pale, spots (Fig. 37) ...... leoni sp. nov. face brown except for broad pale anterior band; pronotum 13’. Forewing clavus with large white arcuate transcommisural brown with pale midline and symmetrical pattern of pale mark- marking near midlength and pentagonal white transcom- ings anteriorly; scutellum and mesonotal midline pale; forew- misural marking near apex (Fig. 38) ...... moralesi sp. nov. ing mostly mottled brown, clavus with two transcommisural pale areas outlined with dark brown, corium with large red- 14. Crown with pair of oblique claviform posterior maculae dish orange submedial spot; veins yellow except near wing mesad of eyes; pronotum with distinct white transverse margin; venter dark brown, tibiae somewhat paler. Body rela- band on posterior margin (Fig. 23) ...... gingerae sp. nov. tively short and broad. Forewing with RP and MA separate, 14’. Crown without pair of oblique claviform posterior maculae connected by crossvein; CuA connected to M basad of its fork. mesad of eyes; pronotum with posterior margin brown or Hind tibia row AV with 3 macrosetae. with very narrow indistinct white posterior band ...... 15 Male abdomen (Fig. 242) with 2S apodemes broad, 15. Mesonotum with pair of black spots just anterad of scutellar slightly divergent, extended to midlength of sternite IV. Pygofer suture (Fig. 29) ...... 16 with posterodorsal emargination narrow and elliptical (Fig. 168); dorsal process elongate, curved posterolaterad and slightly 15’. Mesonotum without pair of black spots just anterad of ventrad; ventral appendage short, slender, divided into two scutellar suture ...... 17 slender acuminate processes, processes weakly divergent and 16. Forewing with transcommisural pale marking at base of curved dorsad (Figs 167 and 168). Anal hook (Fig. 167) slen- clavus much larger than marking at apex of clavus (Fig. der, without ventrolateral arm, extended ventromesad and 28); aedeagus with pair of distal processes shorter than shaft united with dorsal connective, apex acuminate, extended ven- (Fig. 206) ...... kathyae sp. nov. trad; segment X without additional posterior lobes or processes. 16’. Forewing with transcommisural pale markings of clavus Dorsal connective narrow, platelike. Subgenital plate (Fig. 167) subequal in size (Fig. 27); aedeagus with pair of basal pro- strongly constricted at base, widest near midlength, apical half cesses much longer than shaft (Fig. 210) .... peggyae sp. nov. somewhat roundly tapered, with ~12 macrosetae in lateral band 17. Crown with pale areas extended to lateral margin ...... 18 on distal half. Style apodeme (Figs 169 and 170) short, apophysis straight, extended posterad to midlength of plate. Con- 17’. Crown with pale areas not extended to lateral margin, nective (Fig. 170) U-shaped, moderately broad, with pair of margin brown throughout length ...... 19 prominent posterodorsal lobes. Aedeagus (Figs 169 and 170) 18. Pronotum with pair of small white spots laterad of large with preatrium shorter than shaft; dorsal apodeme well devel- median pale marking (Fig. 25) ...... marilynae sp. nov. oped; shaft slender, curved dorsad, apex weakly spatulate, atrial 18’. Pronotum with pair of small white spots posterolaterad of processes absent. Female unknown. large median pale marking (Fig. 26) ...... susanae sp. nov. Material examined. Holotype male. ECUADOR: Orellana, 19. Margin of median pale area of crown lobed laterally (Fig. Tiputini Biodiversity Sta. nr. Yasuni Nat. Pk. 220-250m 27); forewing with single pale transcommisural marking; 00°37’55"S, 076°08’39"W, 6 February 1999, T. L. Erwin et al., aedeagus without apical processes (Fig. 202) ...... lot 2073, transect T/, fogging terra firme forest [USNM]...... helenae, sp. nov. Paratypes: 2 males, same data [USNM, INHS]. Etymology. This species is named in memory of my sister-in-law, Barbara B. Craft. 19’. Margin of median pale area of crown entire; forewing with two pale transcommisural markings; aedeagus with pair of Eualebra charlesi sp. nov. apical processes ...... rufoornata (Stål) Figs 19, 171-174, 243, 358-360 Eualebra barbarae species group Description. Length of male 2.8 mm, female 3.3 mm. Color (Fig. 19) mostly dark brown, marked with white and red- Diagnosis. Species of this group may be distinguished dish orange; crown mostly brown with midline and lateral from other Eualebra in their very small size (<3 mm), relatively margins pale, face brown except for broad pale anterior band; elongate crown (Figs 18 and 19), mottled brown coloration, pronotum brown with pale midline and symmetrical pattern cruciform or trident-shaped pronotal marking, basally quad- of pale markings anteriorly; scutellum and mesonotal midline rate third apical cell of the forewing, platelike dorsal connec- pale; forewing mostly mottled brown, clavus with two tive, and absence of paired processes on the aedeagus (Fig. 170). transcommisural pale areas outlined with dark brown, corium Eualebra barbarae sp. nov. with large reddish orange submedial spot; veins yellow except near wing margin; venter dark brown, tibiae somewhat paler. (Figs 18, 167-170, 242) Body relatively short and broad. Forewing with RP and MA Description. Length of male 2.7 mm. Color (Fig. 18) separate, connected by crossvein; CuA connected to M basad mostly dark brown, marked with white and reddish orange; of its fork. Hind tibia row AV with 3 macrosetae.

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Male abdomen (Fig. 243) with 2S apodemes broad, slightly brown, pygofer and medial area of male subgenital plates dark divergent, extended to midlength of sternite IV. Pygofer with brown. Body relatively broad. Forewing with RP and MA sepa- posterodorsal emargination poorly delimited, shallow (Fig. 172); rate, connected by crossvein; CuA connected to M at or basad dorsal process absent but margin strongly sclerotized with broad of its branching point. Hind tibia row AV with 4 macrosetae. posterodorsal lobe; ventral appendage arising near base of ven- Male 2S apodemes (Fig. 244) moderately broad, slightly tral margin, relatively robust, acuminate, curved dorsomesad divergent, extended to midlength of sternite IV. Pygofer with (Figs 171 and 172). Anal hook (Fig. 171) extended postero- posterodorsal emargination broadly V-shaped (Fig. 176), dor- ventrad, apex footlike with heel oriented posterodorsad and toe sal process short, acuminate, extended posteromesad; ventral extended anteroventrad, united medially with dorsal connec- appendage shifted dorsad, partly fused to pygofer lobe, elon- tive; segment X without additional posterior lobes or processes. gate, extended ventromesad then curved posterodorsad, apex Dorsal connective narrow, platelike. Subgenital plate (Fig. 171) contorted (Figs 175 and 176). Anal hook (Fig. 175) short, fal- constricted at base, widest near midlength, apical half evenly cate, extended ventrad, united medially with dorsal connec- tapered to bluntly rounded apex, with ~12 macrosetae in lateral tive; segment X without additional posterior lobes or processes. band on distal half. Style apodeme (Figs 173 and 174) short, Dorsal connective broadly U-shaped. Subgenital plate (Fig. 175) apophysis straight, extended posterad to distal 1/3 of plate. narrowest at base, convexly broadened to midlength and nar- Connective (Fig. 174) U-shaped, moderately broad, with pair of rowed distally to rounded apex, with ~13 macrosetae in lateral prominent posterodorsal lobes. Aedeagus (Figs 173 and 174) with band on distal half. Connective (Fig. 178) compact, U-shaped, preatrium slightly longer than shaft; shaft slender, weakly sinu- with prominent posterodorsal lobes. Style apodeme (Figs 177 ate and slightly tapered in lateral view, apex blunt. and 178) moderately long; apophysis relatively straight, ta- Female sternite VII (Fig. 358) posterior margin with acute pered, apex blunt, curved ventrad. Aedeagus (Figs 177 and 178) median projection longer than lateral angles; second valvulae with preatrium slightly longer than shaft; dorsal apodeme (Fig. 360) moderately slender, evenly curved dorsad, apical 1/4 weakly developed; shaft short, tapered in lateral view, curved of dorsal margin with ~12 teeth decreasing in size distally. dorsad, with pair of long, slender, somewhat asymmetrical lat- Material examined. Holotype male, ECUADOR: Orellana, eral processes arising near base, extended laterad, curved me- Res. Etnica Waorani, Transect Ent. 1 km S Onkonegare Camp, sad and extended nearly to midline below and distad of shaft 00°39’10"S, 076°26’00"W, 220 m, 26 June 1996, T. L. Erwin et apex; shaft also with pair of depressed lateral flanges extended al., lot 1589, fogging terra firme forest [USNM]. Paratype: 1 to apex, apex acute. Female unknown. female, same data except, 23 February 1995, lot 1059 [INHS]. Material examined. Holotype male, PERU: Pasco, Yanacha- Etymology. This species is named in memory of my ga-Chemillén N.P., Huampal Stn. 10°11’9"S 75°34’27"W 6-9 X 2002, brother-in-law, Charles Craft. D. Takiya, C. Peña, R. Rakitov, Malaise trap acr. R. Huancabamba [USML]. Paratypes: 2 males, same data [INHS]; 7 males, PERU: Pasco, Eualebra dorisae species group Yanachaga-Chemillén N.P., Puesto de Control Huampal, 10°11’9"S Diagnosis. Species of this group may be distinguished 75°34’27"W, 1050m, 6 X 2002, D.M. Takiya [INHS]. from other Eualebra in their moderate size, the color pattern Etymology. This species is named in honor of my aunt, (Figs 20-22) which includes a pair of broad orange or brown Doris A. Dietrich. longitudinal stripes extended from the crown onto the pronotum, the basally quadrate third apical cell of the forew- Eualebra jessicae sp. nov. ing, and the elongate aedeagal atrium (Fig. 177). Figs 21, 179-182, 245 Description. Length of male 4.1 mm. Dorsal coloration Eualebra dorisae sp. nov. (Fig. 21) mostly pale brown with symmetrical white, orange, Figs 20, 175-178, 244 and dark brown markings; venter dull stramineous except Description. Length of male 4.2 mm. Coloration of dor- mesepisternum dark brown. Crown and pronotum orange, sum (Fig. 20) mostly brownish orange with paler orange and darkening to brown posteriorly with three white longitudinal white markings; crown with midline and lateral margins white, stripes; mesonotum with anterolateral triangles and pair of with pair of irregular orange longitudinal submedial stripes; small medial spots dark brown, medial area white tinged with pronotum orange brown with medial and pair of submedial orange; scutellum white with transverse brown band; forew- white stripes not extended to posterior margin; mesonotal tri- ing mostly brown with complex color pattern consisting of angles brown, scutellum white with brown preapical band; forew- two transcommisural markings concentrically outlined with ing mostly brown, mottled distally, with two angulate dark brown and white, numerous small brown spots and lines, transcommisural marks on clavus, two smaller white spots on and large orange brochosome field, veins mostly bright orange. corium adjacent to clavus, brochosome field pale orange; ab- Body robust, ovoid. Forewing with RP and MA separate, con- dominal terga brown; venter of head and thorax white, infused nected by crossvein; CuA connected to M basad of its branch- with brown except mesepisternum dark brown; abdominal terga ing point. Hind tibia row AV with 3 macrosetae.

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 539

169

170 168 167

173

174 171 172

177

176 178 175

181

182

180

179

185

184 186 183 Figures 167-186. Eualebra, male genitalia. (167-170) E. barbarae sp. nov.: (167) genital capsule, lateral view; (168) pygofer and basal sclerites of anal tube, dorsal view; (169) genitalia, lateral view; (170) genitalia, ventral view, only right style shown; (171-174) E. charlesi sp. nov., same; (175-178) E. dorisae sp. nov., same; (179-182) E. jessicae sp. nov., same; (183-186) E. patriciae sp. nov., same.

ZOOLOGIA 30 (5): 519–568, October, 2013 540 C.H. Dietrich

Male abdomen (Fig. 245) with 2S apodemes robust, slightly half, apex broadly rounded. Style (Figs 185 and 186) apodeme divergent, extended slightly beyond midlength of sternite IV. short, apophysis very long, nearly straight, slightly arcuate in Pygofer posterodorsal emargination V-shaped, forming right lateral view, extended to apical 1/4 of subgenital plate. Con- angle (Fig. 180); dorsal process absent; lobe weakly developed; nective (Fig. 186) small, slender, broadly U-shaped. Aedeagus ventral appendage robust, relatively short, evenly curved (Figs 185 and 186) short, preatrium longer than shaft; shaft posterodorsad. Anal hook (Fig. 179) moderately short and broad, tubular, tapered, without processes. Female unknown. extended ventrad, apex footlike, articulated with broadly W- Material examined. Holotype male, PERU: Pasco, Yanacha- shaped dorsal connective. Subgenital plate (Fig. 179) in ventral ga-Chemillén N.P., Huampal Stn. 10°11’9"S 75°34’27"W 6-9 X view widest near midlength, lateral margin evenly convex 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap acr. R. Huanca- throughout length, apex rounded. Style apodeme (Figs 181 and bamba [USML]. 182) relatively long, apophysis sinuate in lateral view, extended Etymology. This species is named in honor of my sister- to apical 1/3 of subgenital plate. Connective (Fig. 182) robust, in-law, Patricia B. Schulz. U-shaped. Aedeagus (Figs 181 and 182) curved dorsad; preatrium Remarks. When dissected, the holotype specimen had a longer than shaft; shaft tubular, tapered, extended posterodorsad, late-instar dryinid (Hymenoptera: Dryinidae) larva in the ab- with pair of slender subapical lateral processes curved and form- dominal cavity. Parasitism by dryinids sometimes causes ab- ing semicircle in ventral view. Female unknown. normal development of the abdominal apodemes and genitalia Material examined. Holotype male, PERU: Pasco, Yanacha- in leafhoppers. The genitalia of the holotype appear to be fully ga-Chemillén N.P., Puesto de Control Huampal, 10°11’9"S developed, but the relatively small size of the 2S apodemes 75°34’27"W, 1050m, 6 X 2002, D.M. Takiya [USML]. Paratypes: may be attributable to presence of the parasite. two males, same data [INHS]. Etymology. This species is named in honor of my niece, Eualebra gingerae species group Jessica C. Bailey. Diagnosis. Species of this group may be distinguished from other Eualebra by their moderately small size, brown over- Eualebra patriciae sp. nov. all coloration (Fig. 23-30), and the presence of large pale Figs 22, 183-186; 246 transcommisural markings on the forewing and, usually, large Description. Length of male 4.0 mm. Dorsal coloration pale symmetrical areas on the crown and pronotum (absent in (Fig. 22) pale yellow with symmetrical white, orange, and brown E. jenniferae). markings; venter uniformly dull yellow; head with crown mar- Eualebra gingerae sp. nov. gins and midline white, with pair of orange spots along anterior margin, pair of orange S-shaped lines submedially, and Figs 23-24, 187-190, 247, 367 pair of large orange rectangles posteriorly; pronotum with Description. Length of male 3.3 mm, female 3.0 mm. paired orange spots anteromedially and at posterolateral angles, Coloration of dorsum (Figs 23 and 24) mostly dark brown with large orange marking posteromedially, divided along midline symmetrical yellow, orange and white markings; crown and anteromedially; mesonotal triangles orange, pair of small brown anterior half of pronotum with white medial area bearing 4 spots present anterad of scutellar suture; scutellum with trans- lateral extensions, 2 on crown, one on margin of crown and verse brown band; forewing mostly yellow orange with veins pronotum, and one medially on pronotum, latter with 2 smaller white, clavus with two transcommisural white markings, bra- white spots more laterad, posterior margin of pronotum nar- chial cell with two irregular white spots, brochosome field white rowly yellow; mesonotum and base of forewing with inverted infused with bright yellow, apex with diffuse brown markings. pentagonal white mark bearing 2 small black spots on Body robust, ovoid. Forewing with RP and MA separate, con- mesonotum and arcuate black band across clavus and scutel- nected by crossvein; CuA connected to M at or basad of its lum; clavus with slender white transcommisural mark near branching point. Hind tibia row AV with 3 macrosetae. midlength and larger triangular mark near apex; corium with Male abdomen (Fig. 246) with 2S apodemes slender, par- small yellow spot near base and large oblique orange macula allel, widely separated, extended to nearly to posterior margin extended from near base of costal margin across brochosome of sternite III. Pygofer posterodorsal emargination very nar- field and into medial cell; veins orange in distal half, bordered row, forming deep acute angle (Fig. 184); dorsal process ab- with brown; venter of head, thorax and abdomen brown. Body sent; lobe weakly developed, ventral appendage slender, evenly moderately broad. Forewing with RP and MA separate, con- curved ventromesad then dorsolaterad, somewhat arcuate in nected by crossvein; CuA connected to MP. Hind tibia row AV lateral view; apex tapered and irregularly sinuate (Figs 183 and with 3 macrosetae. 184). Anal hook (Fig. 183) moderately broad, short, sinuate, Male abdomen (Fig. 247) with 2S apodemes relatively apex curved mesad, articulated with broadly W-shaped dorsal narrow, slightly divergent, extended to midlength of sternite connective; segment X weakly sclerotized. Subgenital plate (Fig. IV. Pygofer with posterodorsal emargination broadly V-shaped 183) in ventral view widest near midlength, tapered in distal (Fig. 188); dorsal appendage absent but posterodorsal margin

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 541 thickened and darkly pigmented; ventral appendage shifted scutellum and smaller marking at apex of clavus; veins pale, dorsad, articulated to pygofer lobe, broad basally with dorsal bordered with brown; venter of head, thorax and abdomen lobe, strongly tapered and attenuate distally, extended tan, legs dull stramineous, macrosetae black. Body relatively anteroventrad then curved dorsolaterad (Figs 187 and 188). narrow. Forewing with RP and MA separate, connected by Anal hook (fig. 187) without ventrolateral arm, mesal arm ex- crossvein; CuA connected to MP. Hind tibia row AV with 3 tended to dorsal connective, with digitiform distal process; macrosetae. segment X with separate pair of slender spinelike processes Male abdomen (Fig. 248) with 2S apodemes short, broad, arising from ringlike sclerite near midlength. Dorsal connec- well separated at base, convergently extended dorsomesad, not tive broadly U-shaped. Subgenital plate (Fig. 187) narrowest at exceeding posterior margin of sternite III. Pygofer with base, slightly broadened, then convexly tapered to blunt apex, posterodordal emargination V-shaped (Fig. 192); dorsal pro- with ~9 macrosetae in lateral band on distal half. Connective cess absent; ventral appendage slender, elongate, arising ven- (Fig. 190) broadly U-shaped with prominent posterodorsal trally anddd curved dorsad and slightly laterad with apical half lobes. Style (Figs 189 and 190) apodeme short, apophysis ta- contorted. Anal hook short, robust, extended posteromesad pered, sinuate, apex curved posteromesad, not surpassing and terminating ventrally in short process, apex obliquely ta- midlength of subgenital plate. Aedeagus (Figs 189 and 190) pered, with basomesal arm united with dorsal connective. with preatrium longer than shaft, dorsal apodeme well devel- Dorsal connective broadly U-shaped. Subgenital plate (Fig. 191) oped with compressed anterior lobe and posterior transverse narrowest at base, slightly broadened, then evenly tapered to plate; shaft compressed basally, tubular distally, curved dor- blunt apex, with ~12 macrosetae in lateral double row on dis- sad; without atrial processes, apex with pair of long slender tal 2/3. Style apodeme (Figs 193 and 194) short, apophysis ex- laterally directed processes. tended mesad, then gradually curved posterad, extended to Female sternite VII (Fig. 367) with short, acute median distal 1/3 of subgenital plate. Connective (Fig. 194) broadly U- posterior lobe and pair of shorter lateral lobes. Second valvu- shaped. Aedeagus (Figs 193 and 194) with preatrium longer lae evenly curved and moderately narrow, right blade with ~13 than shaft; shaft very short, slender, straight, tubular; atrium teeth. with pair of elongate ventrolateral processes recurved in lat- Material examined. Holotype male, BRAZIL: Goias, eral view, bowed laterad in ventral view with apices curved Uruaçu, Serra da Mesa Survey 14°17.1’S, 48°55.6’W, 26-31 May slightly laterad, > 3X longer than shaft. 1996, Malaise trap, cerrado [MZSP]. Paratype: 1 male, same data Female sternite VII (Fig. 363) with posterior margin pro- [INHS]. Other material: 1 female, ECUADOR: Orellana, Tiputini duced and forming narrowly obtuse angle; second valvulae (Figs Biodiversity Sta. nr. Yasuni Nat. Pk. 220-250m 00°37’55"S, 365 and 366) slender, evenly curved dorsad, right blade with 076°08’39"W, 7 February 1999, T.L. Erwin et al., lot 2058, ~13 close-set teeth decreasing in size distally. transect T/6, fogging terra firme forest [USNM]. Material examined. Holotype male, ECUADOR: Orellana, Etymology. This species is named in honor of my sister- Tiputini Biodiversity Sta. nr. Yasuni Nat. Pk. 220-250m in-law, Ginger P. Dietrich. 00°37’55"S, 076°08’39"W, 8 February 1999, T. L. Erwin et al., Remarks. The association between male specimens from lot 2031, transect T/4, fogging terra firme forest [USNM]. Goias, Brazil and the female from Orellana, Ecuador, based on Paratype: 1 female, same data except transect T/3, 8 February the nearly identical color pattern, must be considered tenta- 1999, lot 2028 [INHS]. tive. The Ecuadorean specimen is excluded from the type se- Etymology. This species is named in honor of my aunt, ries because it is somewhat smaller than the males from Brazil Marilyn B. Dietrich. and differs in some small details of coloration, e.g., the specimen from Ecuador lacks the pair of small black submedial spots Eualebra susanae sp. nov. on the mesonotum and has the medial pale area of the Figs 26, 195-198, 249, 368-370 pronotum relatively large. Description. Length of male 2.8 mm; female 2.6 mm. Eualebra marilynae sp. nov. Coloration of dorsum (Fig. 26) brown with large symmetrical cream-white areas bordered in dark brown; crown mostly white Figs 25, 191-194, 248, 363-366 with margins anterad of eyes discontinuously dark brown; Description. Length of male 3.0 mm, female 3.0 mm. pronotum with large white medial area and pair of smaller Coloration of dorsum (Fig. 25) mostly tan to medium brown posterolateral white spots; forewing clavus with two large with symmetrical white and black markings; crown white with transcommisural white spots extended onto corium, anterior pair of black marginal spots at apex and another just anterad spot continuous with white area on mesonotum and scutel- of eyes; pronotum with large white anteromedial marking bear- lum; brochosome field with small basal macula and large dis- ing pair of rounded posterolateral lobes; mesonotum with tal macula dark brown, costal margin more distad with brown M-shaped medial marking; forewing with large white triangular bright yellow marking bordered with dark brown, transcommisural marking at base of clavus continuous with veins pale, cells darker brown; venter stramineous with apex

ZOOLOGIA 30 (5): 519–568, October, 2013 542 C.H. Dietrich of anteclypeus dark brown; apical portion of femur and base Eualebra helenae sp. nov. of tibia of mid- and hind legs dark brown, basal 2 setae of hind Figs 27, 199-202, 250, 361-362 tibial row AD black. Body relatively slender. Forewing with RP and MA confluent preapically, outer apical cell relatively large; Description. Length of male 3.4 mm. Color mostly dark CuA connected to MP. Hind tibia row AV with 3 macrosetae. brown (Fig. 27); dorsum with three conspicuous opaque white Male abdomen (Fig. 249) with 2S apodemes relatively narrow, medial markings, one on crown triangular with lateral invagi- slightly divergent, extended to midlength of sternite IV. Pygofer nation, one on pronotum winglike, one at base of clavus and with posterodorsal emargination V-shaped with basal constric- extended onto scutellum and mesonotum semicircular anteri- tion (Fig. 196), dorsal appendage absent; ventral appendage orly, tapered posteriorly; forewing costal brochosome field arising posteroventrad, rigidly attached to pygofer lobe, slen- bright red. Body short and broad. Forewing with RP and MA der and moderately long, curved dorsolaterad, apex hooked separate, connected by crossvein; CuA connected to MP. Hind laterad (Figs 195 and 196). Anal hook (Fig. 195 and 196) con- tibia row AV with 3 macrosetae. sisting of quadrate sclerotized plate articulated to pygofer side Male abdomen (Fig. 250) with 2S apodemes long, nar- and extended dorsomesad to midlength of segment X, with row, slightly divergent, joined at base, extended to midlength acute ventromesal extension and ventrolateral articulation to of sternite V. Pygofer with posterodorsal emargination broadly dorsal connective, posterior of anal ring present, not extended U-shaped (Fig. 200); dorsal appendage absent; lobe produced ventrad of segment X margin. Dorsal connective broadly U- posterodorsad; ventral appendage arising from base of dorsal shaped. Subgenital plate (Fig. 195) narrowest at base, slightly lobe, elongate, slender, extended posteroventrad then curved broadened, then convexly tapered to blunt apex, with ~10 dorsad, enclosed laterally by unpigmented acuminate poste- macrosetae in lateral band on distal half. Connective small H- rior sheath (Figs 199 and 200). Anal hook (Fig. 199) extended shaped with posterodorsal lobes larger than anterior arms. Style ventromesad, with short digitiform apical process extended apodeme (Figs 197 and 198) short, apophysis tapered, sinuate, ventrad from articulation with U-shaped dorsal connective; apex curved ventrolaterad, blunt, not surpassing midlength of segment X without additional posterior lobes or processes. subgenital plate. Aedeagus (Figs 197 and 198) with preatrium Subgenital plate (Fig. 199) narrowest at base, slightly broad- longer than shaft, dorsal apodeme well developed with com- ened in basal 1/3, distal 2/3 of lateral margin concavely ta- pressed anterior lobe and posterior transverse plate; shaft very pered to blunt apex, with ~10 macrosetae in irregular lateral short and slender, tubular, straight; with pair of slender pro- row on distal 2/3. Style apodeme (Figs 201 and 202) short, apo- cesses arising from atrium and extended ventrad of shaft be- physis straight, extended posteromesad, not surpassing yond shaft apex, bent slightly dorsad in lateral view near apex, midlength of subgenital plate. Connective (Fig. 202) broadly slightly divergent in ventral view. W-shaped. Aedeagus (Figs 201 and 202) with preatrium as long Female sternite VII (Fig. 368) convexly produced poste- as shaft; dorsal apodeme well developed; shaft narrow, tubu- riorly with slight median emargination; ovipositor evenly lar, tapered, curved dorsad, without processes. curved posterodorsad; second valvulae (Fig. 370) slightly broad- Female sternite VII (Fig. 361) strongly produced ened from base to apex, with ~13 irregular teeth. posteromedially, concealing basal half of ovipositor. Second valvulae (362) moderately broad, evenly curved, right blade Material examined. Holotype male, ECUADOR: Orellana, Res. Etnica Waorani, Transect Ent. 1 km S Onkonegare Camp, with ~13 close-set teeth grouped near apex. 00°39’10"S, 076°26’00"W, 220 m, 9 February 1995, T.L. Erwin Material examined. Holotype male, PERU: Madre de Dios; et al., lot 986, fogging terra firme forest [USNM]. Paratypes: 1 Rio Tambopata Res; 30 air km. SW Pto. Maldonado, 290m. 16- male, same data except lot 977; 1 female, same data except 30 20 XI 1979 J.B. Heppner, subtropical moist forest [USNM]. September 1996, lot 1680; 1 female, same data except lot 1677 Paratype: 1 female, same data [INHS]. [USNM, INHS]. Other material: 1 female, ECUADOR: Orellana, Etymology. This species is named in memory of my aunt, Tiputini Biodiversity Sta. nr. Yasuni Nat. Pk. 220-250m Helen R. Huchko. 00°37’55"S, 076°08’39"W, 9 February 1999, T. L. Erwin et al., Eualebra kathyae sp. nov. lot 2035, transect T/3, fogging terra firme forest [USNM]. Etymology. This species is named in honor of my wife, Figs 28, 203-206, 251 Susan M. Braxton. Description. Length of male 4.4 mm. Coloration (Fig. Remarks. The female specimen from Tiputini is very simi- 28) mostly medium brown with symmetrical pale yellow, white, lar to the specimens included in the type series in coloration and red markings. Head brown, gena, anterior margin, and but has a pair of black spots near the posterior margin of the median wedge-shaped area of crown yellow, crown with paired crown, lacks the bright yellow mark on the costal margin of darker brown maculae mesad of eyes and along anterior mar- the forewing, and has the pregenital sternite of the abdomen gin. Pronotum with large median white marking consisting of more strongly produced. It may represent a distinct species or pair of small anteromedial lobes extended to anterior margin a variant of E. susanae. and pair of larger posterolateral quadrate lobes. Mesonotum

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 543

189

188 187 190

192 194 191

193

197

196

195 198

201

199 200 202

Figures 187-202. Eualebra, male genitalia. (187-190) E. gingerae sp. nov.: (187) genital capsule, lateral view; (188) pygofer and basal sclerites of anal tube, dorsal view; (189) genitalia, lateral view; (190) genitalia, ventral view, only right style shown. (191-194) E. marilynae sp. nov., same. (195-198) E. susanae sp. nov., same. (199-202) E. helenae sp. nov., same.

brown anteriorly, yellow posteriorly, area anterad of scutellar near base and another near midlength; costal area with suture brown; scutellum yellow with brown transverse band. brochosome field bright red; veins mostly orange; outer two Forewing clavus with two triangular transcommisural mark- apical cells hyaline distally. Body robust, ovoid. Forewing with ings, one near base an another smaller one near apex, basal RP and MA separate, connected by crossvein; CuA connected triangle orange medially; corium with two round spots, one to MP. Hind tibia row AV with 3 macrosetae.

ZOOLOGIA 30 (5): 519–568, October, 2013 544 C.H. Dietrich

Male abdomen (Fig. 251) with 2S apodemes robust, par- then dorsolaterad; apex tapered and irregularly sinuate (Figs allel, extended to midlength of sternite IV. Pygofer postero- 207 and 208). Anal hook (Fig. 207) very short, broad, with dorsal emargination broadly V-shaped, forming shallow obtuse ventromedial toothlike process, articulated with broadly V- angle (Fig. 204); dorsal process absent; lobe weakly developed; shaped dorsal connective; segment X weakly sclerotized. ventral appendage robust, nearly straight, somewhat expanded Subgenital plate (Fig. 207) in ventral view widest near near midlength, tapered and irregularly sinuate distally, ex- midlength, tapered in distal half, with lateral margin concave, tended posteromesad (Figs 203 and 204). Anal hook (Fig. 203) apex narrowly rounded. Style apodeme (Figs 209 and 210) short, short, extended ventrad, bidentate apically with teeth widely apophysis long, nearly straight, extended to apical 1/3 of separated, articulated with broadly W-shaped dorsal connec- subgenital plate. Connective (Fig. 210) robust, U-shaped. tive; segment X with pair of denticulate ventrolateral lobes Aedeagus (Figs 209 and 210) short, preatrium subequal in length distally. Subgenital plate (Fig. 203) in ventral view widest near to shaft; shaft tubular, extended posterodorsad, apex depressed, midlength, tapered in distal half, apex rounded. Style apodeme rounded in posterior view; atrium expanded with pair of elon- (Figs 205 and 206) short, apophysis sinuate, extended to api- gate slender lateral processes curved posterodorsad then cal 1/3 of subgenital plate. Connective (Fig. 206) relatively large, strongly hooked ventrolaterad, extended well beyond shaft U-shaped with weak anteromedial tooth. Aedeagus (Figs 205 apex. Female unknown. and 206) nearly straight; preatrium subequal in length to shaft; Material examined. Holotype male, PERU: Pasco, Yanacha- shaft tubular, tapered, extended posterodorsad, with pair of ga-Chemillén N.P., Huampal Stn. 10°11’9"S 75°34’27"W 6-9 X slender subapical lateral processes oriented approximately per- 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap acr. R. Huanca- pendicular to shaft. Female unknown. bamba [USML]. Material examined. Holotype male, PERU: Pasco, Yanacha- Etymology. This species is named in memory of my ga-Chemillén N.P., Huampal Stn. 10°11’9"S 75°34’27"W 6-9 X mother-in-law, Margarete (Peggy) D. Braxton. 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap acr. R. Huanca- Remarks. This species is very similar to E. rufoornata (Stål) bamba [USML]. in size and coloration, but differs in its short, broad anal hook Etymology. This species is named in honor of my sister- and elongate basal aedeagal processes. in-law, Katherine (Kathy) E. Dietrich. Remarks. This species is similar to E. rufoornata (Stål), Eualebra rufoornata (Stål) but differs in its much larger size, brown ventral coloration, Typhlocyba rufoornata Stål 1860: 55. more shallowly emarginate tergite IX, and bifid anal hook. Remarks. Specimens were not examined for the present Eualebra peggyae sp. nov. study but the species was redescribed by Linnavuori (1954), Figs 29, 207-210, 252 who examined the male holotype from “Brazil” and illustrated the dorsal color pattern and male genitalia. Based on Description. Length of male 3.9 mm. Dorsal coloration Linnavuori’s drawings, the color pattern resembles that of E. (Fig. 29) mostly medium brown with symmetrical pale yellow, peggyae sp. nov. (Fig. 29) in having a wedgelike white marking white, and orange markings; venter dull yellow except dark extended from the crown onto the pronotum, the posterior brown mesepisternum and meso- and metathoracic femora. Head portion of the mesonotum and scutellum pale, and two dia- and pronotum with large median white wedge-shaped marking mond-shaped transcommisural markings on the forewing, but extended beyond midlength of pronotum; posterior margin of the aedeagus of E. rufoornata has a pair of short distal processes pronotum narrowly yellow. Mesonotum brown anteriorly, yel- that are perpendicular to the shaft rather than the long, sinu- low posteriorly, area anterad of scutellar suture with pair of brown ate basal processes present in E. peggyae. spots; scutellum yellow with brown transverse band. Forewing clavus with two transcommisural diamond-shaped markings, Eualebra jenniferae sp. nov. one near base an another smaller one near apex; corium with Figs 30, 71-72, 211-214, 253 round spots near midlength in discal cell midlength; costal area with brochosome field orange; veins mostly orange; outer two Description. Length of male 3.4 mm. Color mostly dark apical cells hyaline distally. Body robust, ovoid. Forewing with brown dorsally (Fig. 30), apex of crown paler, scutellum white RP and MA separate, connected by crossvein; CuA connected to with brown transverse preapical stripe; forewing clavus with MP. Hind tibia row AV with 3-4 macrosetae. pair of triangular transcommisural yellow areas, large triangu- Male abdomen (Fig. 252) with 2S apodemes robust, par- lar area on costal margin distad of brochosome field, three small allel, extended to midlength of sternite IV. Pygofer posterodor- yellow marks more basad; venter dull yellow, heavily infused sal emargination narrowly V-shaped, forming deep acute angle with brown. Body relatively narrow. Forewing with RP and MA (Fig. 208); dorsal process absent; lobe weakly developed, with separate, connected by crossvein (Fig. 3c), or confluent for short hyaline membranous cleft near midheight of posterior mar- distance; CuA connected to MP. Hind tibia row AV with 4 gin; ventral appendage slender, evenly curved ventromesad macrosetae.

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 545

205

206 204

203 209

210 208

207

214

213 212

211

217

218 216 215

221

222

219

220

Figures 203-222. Eualebra, male genitalia. (203-206) E. kathyae sp. nov.; (203) genital capsule, lateral view; (204) pygofer and basal sclerites of anal tube, dorsal view; (205) genitalia, lateral view; (206) genitalia, ventral view, only right style shown; (207-210) E. peggyae sp. nov., same; (211-214) E. jenniferae sp. nov., same; (215-218) E. smithii, same, style not shown in ventral view; (219-222) E. margaretannae sp. nov., same. (dc) Dorsal connective.

ZOOLOGIA 30 (5): 519–568, October, 2013 546 C.H. Dietrich

Male abdomen (Fig. 253) with 2S apodemes slender, and distal half of corium stippled with brown, median area of slightly divergent, extended nearly to posterior margin of stern- wing with ca. 15-20 conspicuous opaque white spots of vari- ite IV. Pygofer posterodorsal emargination U-shaped, moderately ous sizes; abdominal terga brown medially; venter pale except broad (Fig. 212); ventral appendage shifted to near dorsal mar- anteclypeus and femorotibial joint of mesothoracic leg dark gin, articulated to pygofer lobe, moderately long, slender, par- brown. Body relatively short and broad. Forewing with RP and tially enclosed by posteroventral lobe, extended ventromesad MA separate, connected by crossvein; CuA connected to branch- then curved posterodorsad (Figs 211 and 212). Anal hook (Fig. ing point of M. Hind tibia row AV with 3 macrosetae. 211) robust, extended ventromesad, articulated with dorsal con- Male abdomen (Fig. 254) with 2S apodemes moderately nective, apex with short digitiform process; segment X with pair broad, slightly divergent and expanded distally, extended nearly of short, slender posteriorly directed posteroventral spines. Dor- to posterior margin of sternite IV. Pygofer with posterodorsal sal connective relatively narrow, slightly U-shaped, platelike (Fig. emargination broadly V-shaped (Fig. 216); posterodorsal append- 214). Subgenital plate (Fig. 211) only slightly constricted at base, age absent but margin thickened and heavily sclerotized; ven- lateral margin gradually tapered to more strongly tapered apical tral appendage arising near midheight of posterior margin, 1/4, with ~15 macrosetae in lateral double row on distal 2/3. extended ventromesad then curved posterodorsad, apical half Style apodeme (Figs 213 and 214) short, apophysis extended areolate (Figs 215 and 216). Anal hook (Fig. 215) with short acute posterad and curved slightly mesad, apex blunt, reaching distal ventral projection and slender posterior branch extended me- 1/3 of plate. Connective (Fig. 214) very broad and short, U- sad to dorsal connective, slightly broadened distally. Dorsal con- shaped with prominent posterodorsal lobes. Aedeagus (Figs 213 nective broadly U-shaped. Subgenital plate (Fig. 215) relatively and 214) with preatrium as long as shaft; dorsal apodeme well broad at base, slightly broadened to near midlength, then ta- developed; shaft depressed, apex broken off in only known speci- pered to broadly rounded apex, with ~13-16 macrosetae in lat- men, with pair of slender lateral processes arising near base and eral band on distal half. Style (Figs 217 and 218) apodeme short extended along sides of shaft. Female unknown. and slender, apophysis moderately long and slender, nearly Material examined. Holotype male, ECUADOR: Orellana, straight, bent slightly mesad at apex, apex surpassing midlength Res. Etnica Waorani, Transect Ent. 1 km S Onkonegare Camp, of subgenital plate. Connective (Fig. 218) U-shaped, very short 00°39’10"S, 076°26’00"W, 220 m, 4 October 1996, T.L. Erwin et and broad. Aedeagus (Figs 217 and 218) with preatrium very al., lot 1754, fogging terra firme forest [USNM]. Paratypes: 1 short; dorsal apodeme weakly developed; shaft short and slen- male, same data as holotype except 22 June 1996, lot 1561; 1 der, tubular, with depressed rounded lateral flanges near base; male, same data except 6 July 1995, lot 1113; 1 male, same apex acute. Female sternite VII slightly produced posteromedially. data except 2 October 1996, lot 1704. Material examined. 2 males, BOLIVIA: Cochabamba, Villa Etymology. This species is named in honor of my sister- Tunari, Hotel Los Tucanes 16°58.302’S, 65°23.793’W, 320m 5- in-law, Jennifer L. Dietrich. IX-2000, M. Hauser, S. Gaimari, D. Yeates [INHS]; 4 males, 4 females, PERU: Madre de Dios, Rio Tambopata Eualebra smithii species group Res. 30 km (air) SW Pto. Maldonado, 290 m, 12°50’S, 069°17’W, Smithsonian Institution Canopy Fogging Project, T.L. Erwin et Diagnosis. Species of this group, which includes the type al. colls. 02 May 1984, 02/03/58 [USNM. Female holotype, species of the genus, may be distinguished from other Eualebra Dikraneura (Hyloidea) reticulata Osborn labeled: “Janoria/Minca/ by the pale overall coloration (Figs 31-33), color pattern con- Brazil; Dec 17/1907; Carn. Mus./Acc. 3702; 5045; TYPE” [ICCM]. sisting of numerous irregular tiny brown spots and larger cream- Remarks. Young (1952) illustrated and described the male colored spots, the absence of paired processes on the aedeagus genitalia of E. smithii, presumably based on study of the type (Figs 218 and 222), and the broad, straight ovipositor which specimen. Comparisons of specimens from Peru and Bolivia lacks teeth on the second valvulae (Fig. 374). to Young’s illustrations of E. smithii and to the female holo- Eualebra smithii Baker type of Dikraneura (Hyloidea) reticulata indicate that these two species are synonyms. The specimen from Bolivia has the con- Figs 31-32, 215-218, 254 nective proportionately larger than those of the specimens from Eualebra smithii Baker 1899b: 402. Peru, but the genitalia appear otherwise to be identical. Fur- Dikraneura (Hyloidea) reticulata Osborn, 1928a: 279, syn. nov. ther collecting is needed to determine whether such variation Description. Length of male 3.3-3.4 mm, female 3.8 mm. is indeed intraspecific. Color (Figs 31 and 32) mostly pale yellow to white, marked with white, orange and brown; crown and pronotum with few Eualebra margaretannae sp. nov. indistinct paired orange maculae; antennal ledge dark brown; Figs 33, 62, 219-222, 255, 371-374 pronotum, mesonotum and forewing with sparse brown Diagnosis. Length of male 3.5-3.8 mm, female 3.8 mm. stipples posteriorly; forewing with brown maculae at midlength Color (Fig. 33) mostly pale yellow to white, marked with white, of costal margin and in third and fourth apical cells, clavus orange and brown; crown and pronotum with few indistinct

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 547 paired orange maculae; antennal ledge dark brown; pronotum and relatively narrow (Fig. 224); pygofer lobe with semicircu- with sparse brown stipples posteriorly; scutellum apex dark lar unpigmented area posterodorsally. brown; forewing with brown maculae at midlength of costal margin and in third and fourth apical cells, clavus and distal Eualebra alberti sp. nov. half of corium stippled with brown, median area of wing with Figs 34, 223-226, 256 ca. 15-20 conspicuous opaque white spots of various sizes; ab- Description. Length of male 4.7 mm. Dorsal coloration dominal terga brown medially; venter pale except femorotibial (Fig. 34) yellow, crown with pair of oblique brown maculae joint of mesothoracic leg dark brown. Body relatively short betwen eyes extended toward apex; pronotum with large pos- and broad. Forewing with RP and MA separate, connected by teromedial black semicircular marking; mesonotum with crossvein; CuA connected to MP. Hind tibia row AV with 3 basolateral triangles brown; forewing clavus with oblique black macrosetae. transcommisural macula near midlength, costal margin with Male abdomen (Fig. 255) with 2S apodemes broad, narrow black maculae at base and apex of brochosome field, slightly divergent, extended to midlength of sternite IV. Pygofer apex with diffuse brown markings. Body relatively long and with posterodorsal emargination broadly V-shaped (Fig. 220); narrow; forewing veins RP and MA partially confluent dorsal appendage absent but dorsal margin with strongly scle- preapically, CuA connected to M basad of its branch; hind tibial rotized ridge; with two paired ventral appendages, one arising row AV with 3 macrosetae. near mid-height of posterior margin and another more ven- Male 2S apodemes (Fig. 256) short and tapered. Pygofer trad, both elongate and slender, extended ventromesad then with tergite deeply and narrowly emarginate posteriorly in strongly curved dorsad, ventral pair twice as long as dorsal and dorsal view (Fig. 224); with small rigid posterodorsal process; contorted distally (Figs 219 and 220). Anal hook (Fig. 219) ventral appendage elongate, slender, curved dorsad (Figs 223 massive, extended mesad, broad distally, apex broadly and shal- and 224). Anal hook (Fig. 223) present, articulated to pygofer, lowly bifid, with ventral lobe articulated to dorsal connective; base of anal tube, and dorsal connective. Dorsal connective segment X without additional lobes or processes (Figs 219 and broadly U-shaped in posterior aspect. Subgenital plate (Fig. 223) 220). Dorsal connective very small, platelike. Subgenital plate with base slender, broadened to midlength then gradually ta- (Fig. 219) narrowest at base, broadened to near midlength, then pered toward apex; with 10 or more macrosetae. Style apodeme tapered to blunt apex, with ~13 macrosetae in lateral band on (Figs 225 and 226) short; apophysis elongate, without preapical distal half. Style (Figs 221 and 222) apodeme short, apophysis lobe, with few fine preapical setae, apex hooked. Connective extended mesad then curved posterad, apex surpassing (Fig. 226) weakly U-shaped with pair of posterodorsal lobes midlength of subgenital plate. Connective (Fig. 222) U-shaped, articulated to aedeagus. Aedeagus (Figs 225 and 226) with moderately broad. Aedeagus (Figs 221 and 222) with preatrium preatrium elongate and cleft in dorsal view with margins closely as long as shaft, arcuate; dorsal apodeme well developed; shaft appressed in basal half and divergent in distal half; dorsal tapered and curved dorsad in lateral view; with broad depressed apodeme weakly developed; shaft tubular, broad, curved dor- lateral flanges extended over most of length but terminated sad, without processes. Female unknown. slightly basad of apex; apex acute. Female sternite VII (Fig. 371) Material examined. Holotype male, PERU: Pasco, Yanacha- broadly and shallowly rounded posteriorly. ga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, D. Takiya, Material examined. Holotype male, ECUADOR: Orellana, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML]. Tiputini Biodiversity Sta. nr. Yasuni Nat. Pk. 220-250m Paratypes: 3 males, same data [INHS]. 00°37’55"S, 076°08’39"W, 8 February 1999, T.L. Erwin et al., lot Etymology. This species is named in memory of my 2035, transect T/4, fogging terra firme forest [USNM]. Paratypes: grandfather, Albert E. Dietrich Jr. 5 males and 4 females, same data [USNM, INHS]. Other material: 1 male, 2 females, ECUADOR: Orellana, Res. Etnica Waorani, Transect Ent. 1 km S Onkonegare Camp, 00°39’10"S, Eualebra dorothyae sp. nov. 076°26’00"W, 220 m, 4 February 1996, T. L. Erwin et al., lot Figs 35, 227-230, 257 1413, fogging terra firme forest [USNM, INHS]. Description. Length of male 4.7 mm. Dorsal coloration Etymology. This species is named in honor of my mother, (Fig. 35) mostly yellow; head with diffuse brown spot at apex, Margaret Anne B. Dietrich. pronotum with small brown medial stripe extended slightly onto mesonotum; forewing with narrow brown maculae at base Eualebra michaelorum species group and apex of brochosome field, and brown macula at base of Diagnosis. Members of this group differ from other inner apical cell. Body relatively long and narrow; forewing Eualebra as follows: size relatively large; crown relatively short veins RP and MA partially confluent preapically, CuA connected with anterior margin in profile relatively thick; forewing veins to M basad of its branch; hind tibial row AV with 3 macrosetae. RP and MA partially confluent preapically, forming petiolate Male 2S apodemes (Fig. 257) small, digitiform, not ex- apical cell (Fig. 73); pygofer dorsomedial emargination short tended to posterior margin of sternite III. Pygofer with tergite

ZOOLOGIA 30 (5): 519–568, October, 2013 548 C.H. Dietrich

deeply and narrowly emarginate posteriorly in dorsal view (Fig. Material examined. Holotype male, PERU: Pasco, 228); with small rigid posterodorsal process; ventral append- Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, age elongate, slender, curved dorsad (Figs 227 and 228). Anal D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San hook (Fig. 227) present, articulated to pygofer, base of anal Alberto [USML]. Paratypes: 7 males, same data [INHS]. tube, and dorsal connective. Dorsal connective broadly U- Etymology. This species is named in memory of my shaped in posterior aspect. Subgenital plate (Fig. 227) with base grandfather, Michael Baka, Sr., and in honor of my uncle, and slender, broadened to midlength then gradually tapered to- cousin, Michael Baka, Jr., and III. ward apex; with 10 or more macrosetae. Style apodeme (Figs 229 and 230) short; apophysis elongate, without preapical lobe, Eualebra leoni species group with few fine preapical setae, apex hooked. Connective (Fig. Diagnosis. Species of this group may be distinguished 230) U-shaped with posterodorsal lobes weak. Aedeagus (Figs from other Eualebra by their small size, relatively weakly pro- 229 and 230) with preatrium elongate and cleft in dorsal view duced head, brown overall coloration (Figs 37 and 38), white with margins parallel-sided in basal half, constricted posterior pronotal margin, partially confluent forewing veins preapically; dorsal apodeme weakly developed; shaft tubular, RP and MA, and relatively shallow, broad emargination of male broad, curved dorsad, without processes. abdominal tergite IX (Fig. 236). Material examined. Holotype male, PERU: Pasco, Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Eualebra leoni sp. nov. Alberto [USML]. Paratype: 1 male, same data [INHS]. Figs 37, 235-237, 298 Etymology. This species is named in memory of my Description. Length of male 3.1 mm. Overall color me- grandmother, Dorothy H. Dietrich. dium brown (Fig. 37). Face uniformly brown; band across head apex stramineous, complete; crown midline dark brown in dis- Eualebra michaelorum sp. nov. tal half, coronal suture black. Pronotum with pair of subme- Figs 36, 73-74, 231-234, 258 dian ovoid white spots, and two smaller sublateral spots; Description. Length of male 4.7 mm. Ground color (Fig. posterior margin narrowly white, bordered anteriorly with dark 36) stramineous, heavily marked with brown and black. Crown brown. Mesonotum and scutellum black except paired white midline black bordered with diffuse brown pigment; pronotum posterolateral spots. Scutellum black except white apex. Forew- with diffuse brown median longitudinal stripe marked with ing clavus with three diamond-shaped white transcommisural white anteromedially, pair of dark brown oblique maculae markings, progressively larger posteriorly; corium with two posterolaterally; mesonotum with basal triangles and medial small white spots along claval suture, one in inner anteapical area dark brown; forewing mostly brown with two white cell; brochosome field white tinged with orange; costal mar- transcommisural markings, brochosome field yellow. Body rela- gin with three brown false veinlets preapically. Legs stramineous tively long and narrow; forewing veins RP and MA partially except middle and hind femora mostly black, hind tibia with confluent preapically (Fig. 73), CuA connected to M basad of macrosetae in rows AD, PD and AV black. Forewing R and M its branch; hind tibial row AV with 3 macrosetae. confluent for considerable distance. Male 2S apodemes (Fig. 258) widely separated, weakly Male 2S apodemes (Fig. 258) widely separated, moderately hooked mesad, extended to midlength of sternite IV. Pygofer wide and parallel sided, extended to posterior margin of sternite with tergite deeply and narrowly emarginate posteriorly in IV. Pygofer in dorsal view with posterior margin broadly V-shaped dorsal view (Fig. 232); with small rigid posterodorsal process; (Fig. 236); side evenly sclerotized; ventral appendage very slen- ventral appendage elongate, slender, curved dorsad (Figs 231 der, needlelike, smooth, curved evenly dorsad in lateral view, and 232). Anal hook (Fig. 231) present, articulated to pygofer, bowed laterad in ventral view (Figs 235 and 236). Anal hook (Fig. base of anal tube, and dorsal connective. Dorsal connective 235) short and slender, apex acuminate, extended ventromesad. broadly U-shaped in posterior aspect. Subgenital plate (Fig. 231) Dorsal connective broadly U-shaped in posterior view. Subgenital with base slender, broadened to midlength then gradually ta- plate (Fig. 235) with 7-8 macrosetae; dorsolateral band of fine pered toward apex; with 10 or more macrosetae. Style apodeme setae with longer setae interspersed with shorter ones. Style apo- (Figs 233 and 234) short; apophysis elongate, without preapical physis acuminate, curved dorsad in lateral view, apex bent ven- lobe, with few fine preapical setae, apex hooked. Connective trad. Connective stout, narrowly U-shaped. Aedeagus (Fig. 237) (Fig. 234) H-shaped with posterodorsal lobes as long as ante- with shaft very short, tubular, straight in lateral view, lateral pro- rior arms. Aedeagus (Figs 233 and 234) with preatrium elon- cesses extended posterolaterad and surpassing shaft apex, curved gate and cleft in dorsal view with margins tapered in basal dorsolaterad apically. Female unknown. two-thirds and divergent distally; dorsal apodeme weakly de- Material examined. Holotype male, BOLIVIA: Cochabamba, veloped; shaft tubular, broad, curved dorsad, without processes. Villa Tunari, Hotel Los Tucanes, 16°58.302’S, 65°23.793’W, Female unknown. 320m 5-IX-2000, M. Hauser, S. Gaimari, D. Yeates [INHS].

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 549

224 225

223 226

228 229

227

230

233

232 231 234

237 235

236 240

238 241

239

Figures 223-241. Eualebra, male genitalia. (223-226). E. alberti sp. nov.: (223) genital capsule, lateral view; (224) pygofer and basal sclerites of anal tube, dorsal view; (225) genitalia, lateral view; (226) genitalia, ventral view, only right style shown; (227-230) E. dorothyae sp. nov., same; (231-234) E. michaelorum sp. nov., same; (235-237) E. leoni sp. nov.: (235-236) lateral and dorsal view of genital capsule; (237) ventral view of aedeagus; (238-241) E. moralesi sp. nov.: (238-239) lateral and dorsal view of genital capsule; (240-241) lateral and ventral view of genitalia, only right style shown.

ZOOLOGIA 30 (5): 519–568, October, 2013 550 C.H. Dietrich

Etymology. This species is named in honor of my father- posterolateral angles white. Scutellum black except white in-law, Leon E. Braxton. anteromedially and at apex. Forewing clavus with two white Remarks. This species closely resembles E. moralesi but is transcommisural markings, anterior one thin and arcuate, pos- most readily distinguished by the external color pattern, the terior one pentagonal; corium with two small white spots along slender, needlelike pygofer appendage, and the shorter aedeagal claval suture and large ovoid white spot tinged with orange shaft relative to the preatrium. encompassing brochosome field; costal margin with three brown false veinlets preapically. Legs stramineous except front Eualebra moralesi sp. nov. femur and tibia pale orange, middle femur mostly black, hind Figs 38, 238-241, 260 tibia with macrosetae in basal half of rows AD and PD and at Description. Length of male 3.4 mm. Overall color me- apex of AV black. Forewing with RP and MA confluent for short dium brown (Fig. 38). Face with lateral areas dull stramineous; distance. ocellar vestiges and indistinct broken band across head apex Male 2S apodemes (Fig. 260) widely separated, moder- stramineous; crown midline stramineous in distal half, coro- ately wide and parallel-sided, extended to posterior margin of nal suture black. Pronotum with teardrop-shaped median white sternite IV. Pygofer in dorsal view with posterior margin broadly spot, pair of large submedian ovoid white spots, and two smaller and shallowly concave (Fig. 239), ventral appendage slender, sublateral spots; posterior margin narrowly white, bordered areolate distally, curved evenly dorsad in lateral view, nearly anteriorly with dark brown. Mesonotal triangles dark brown, straight in ventral view (Figs 238 and 239). Anal hook (Fig.

242 243 244 245

247 248 246 249

251 252 253 250

256 257 254 255

260 258 259 Figures 242-260. Eualebra, base of male abdomen, ventral view. a242 Eualebra barbarae sp. nov.; (243) E. charlesi sp. nov.; (244) E. dorisae sp. nov.; (245) E. jessicae sp. nov.; (246) E. patriciae sp. nov.; (247) E. gingerae sp. nov.; (248) E. marilynae sp. nov.; (249) E. susanae sp. nov.; (250) E. helenae sp. nov.; (251) E. kathyae sp. nov.; (252) E. peggyae sp. nov.; (253) E. jenniferae sp. nov.; (254) E. smithii; (255) E. margaretannae sp. nov.; (256) E. alberti sp. nov.; (257) E. dorothyae sp. nov.; (258) E. michaelorum sp. nov.; (259) E. leoni sp. nov.; (260) E. moralesi sp. nov.

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 551

238) short and broad, dorsal margin with subbasal tooth, apex yond posterior margin of sternite III. Pygofer (Figs 261 and acute, extended posteromesad. Dorsal connective broadly U- 262) with dorsal emargination well developed; tergite well de- shaped in posterior view. Subgenital plate (Fig. 238) with 5-6 veloped but short; posterior lobe acutely produced, dorsal part macrosetae not reaching apex of plate; dorsolateral band of heavily sclerotized, with falcate distal process; ventral append- fine setae with longer setae toward apex. Style apophysis (Figs age well developed, elongate. Valve short, emarginate posteri- 240 and 241) acuminate, curved dorsad in lateral view, apex orly, fused to pygofer. Segment X (Fig. 261) largely membranous bent ventrad. Connective (Fig. 241) stout, broadly U-shaped. with sclerotized basal ring, lateral ends of ring produced and Aedeagus (Figs 240 and 241) with preatrium longer than shaft; free posteriorly. Dorsal connective absent. Subgenital plate (Fig. shaft slightly arcuate in lateral view, with lateral processes ex- 261) narrow basally, slightly broadened near middle, with 3-5 tended posterolaterad and slightly surpassing shaft apex, curved macrosetae arranged in continuous row along lateral margin dorsad apically. Female unknown. distally; apex weakly to strongly compressed and slightly up- Material examined. Holotype male, BOLIVIA: Cochabamba, turned. Connective (Fig. 264) Y- shaped, articulated to aedeagus, Villa Tunari, Hotel Los Tucanes, 16°58.302’S, 65°23.793’W, without prominent posterodorsal lobes. Style (Figs 263 and 264) 320m 5-IX-2000, M. Hauser, S. Gaimari, D. Yeates [INHS]. apodeme short, apophysis elongate, with few fine setae Etymology. The species is named in honor of Juan Evo preapically, apex tapered and curved. Aedeagus (Figs 263 and Morales Ayma, President of the Republic of Bolivia. 264) with preatrium well developed, dorsal apodeme weakly Remarks. This species differs from E. leoni in its color to strongly developed, shaft with or without paired processes, pattern, longer aedeagal shaft, and thicker, areolate ventral gonopore apical. pygofer appendage. Female sternite VII (Fig. 375) parabolically produced posteriorly; ovipositor (Fig. 377) elongate, slender, slightly Euzyginella gen. nov. sigmoid, first valvulae (Fig. 376) with strigate dorsal and ventral sculpturing near apex, second valvula (Fig. 377 and 378) Type species: Euzyginella marki sp. nov. right blade with ~13 small close-set teeth near apex of dorsal Diagnosis. Small, depressed leafhoppers (Figs 39-42). margin. Coloration pale yellow, orange, or reddish brown, dorsum Etymology. The genus name, a feminine noun, was cho- with symmetrical white, orange, black and/or red markings. sen to indicate its morphological similarity to Zyginella and Head (Figs 39-42) wider than pronotum, depressed, anterior Eualebra. margin angulately produced; face nearly horizontal (Fig. 64); Remarks. This genus is similar to Pseudozyginella (and rostrum not extended beyond front trochanters; lower part most other Typhlocybini) in having the hind wing submar- of face sexually dimorphic, male anteclypeus inflated and ginal vein not extended to MP (i.e., the hind wing lacks closed expanded laterad, lorum very narrow; female anteclypeus rela- apical cells), but differs in in having the row of macrosetae on tively narrow, flat, and tapered, lorum relatively broad; the subgenital plate continuous and the connective Y-shaped, frontoclypeus convex; lateral frontal sutures not extended and in lacking basolateral apodemes on the anal ring. The spe- dorsad of antennal pits; antennal ledges weakly developed; cies are also somewhat broader and more strongly depressed antennae as long as head width; ocelli absent; crown convex, than in Pseudozyginella. The four known species, described be- coronal suture restricted to posterior third of crown. low, were collected by fogging the canopy of terra firme forest Pronotum (Figs 39-42) with lateral margins moderately long, at two localities in Orellana Province, Ecuador. slightly divergent posteriorly in dorsal view, distinctly carinate, carina even with posterior margin of eye. Front femur Key to species of Euzyginella with AM1 enlarged and situated on ventral margin; interca- 1. Aedeagus with pair of preapical processes (Fig. 272, 276) ... 2 lary row with few fine setae, basal seta larger than others; PV1 well developed; tibia somewhat flattened dorsally, AD 1’. Aedeagus without processes (Fig. 264, 268) ...... 3 and PD without preapical macrosetae. Hind femur macrosetae 2. Crown mostly orange with narrow white band along poste- 2+1+1; tibia row AV with three macrosetae near apex. Forew- rior margin (Fig. 40); anal hook slender, unbranched (Fig. ing (Fig. 75) with RA reflexed, RP confluent with MA for short 269) ...... quartalberti, sp. nov. distance, apical cell 3 triangular, inner apical cell trapezoi- 2’. Crown mostly white with narrow transverse red band dal; hind wing (Fig. 76) venation somewhat variable inter- between anterior margins of eyes (Fig. 41); anal hook with and intraspecifically, submarginal vein absent between CuA three acute distal branches (Fig. 273) ... teralberti, sp. nov. and RP, RP and MA separate, connected by crossvein or confluent; distal segment of CuA (beyond connection with 3. Anal hook short, blunt (Fig. 265); aedeagal shaft evenly submarginal vein) present. curved dorsad in lateral view (Fig. 13g) ..... marki, sp. nov. Male abdomen (Figs 337-340) with 2S apodemes well 3’. Anal hook long and slender (Fig. 261); aedeagal shaft nearly developed, connected by sclerotized bridge at base but widely straight with ventral margin slightly sinuate in lateral view separated distally, more or less parallel, usually extended be- (Fig. 263) ...... theari sp. nov.

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Euzyginella marki sp. nov. hooked ventrad, acuminate. Connective (Fig. 272) consisting of triangular basal plate with well-developed stem. Aedeagus (Figs Figs 39, 75-76, 265-268, 337, 375-378 271 and 272) with preatrium shorter than shaft; dorsal apodeme Description. Length of male 3.1 mm. Color (Fig. 39) weakly developed; shaft slender, compressed, slightly sinuate, dull stramineous, crown and pronotum each with arcuate with pair of basolateral spines extended less than half shaft transverse orange band near midlength. Forewing with brown length ventrad of shaft; shaft apex slightly broadened and trun- maculae at base and apex of costal brochosome field, clavus cate in ventral view. Female unknown. with three large milky spots; remainder of wing in basal 2/3 Material examined. Holotype male, ECUADOR: Orellana, milky. Tiputini Biodiversity Sta. nr. Yasuni Nat. Pk. 220-250m Male abdomen (Fig. 337) with 2S apodemes robust, nearly 00°37’55"S, 076°08’39"W, 26 October 1998, T. L. Erwin et al., parallel sided, extended slightly beyond posterior margin of lot 1940, transect T/5, fogging terra firme forest [USNM]. sternite IV. Pygofer with posterodorsal emargination extended Etymology. This species is named in honor of my cousin, to base, broadly U-shaped (Fig. 266); dorsal appendage robust, Albert E. Dietrich, IV. extended posterad, abruptly tapered and decurved apically; ventral appendage slender, straight, extended posterodorsad Euzyginella teralberti sp. nov. along pygofer margin, expanded preapically in ventral view Figs 41, 273-276, 339 apex acuminate and curved laterad (Figs 265 and 266). Anal Description. Length of male 2.8 mm. Crown (Fig. 41) hook (Fig. 265) short, slender, spinelike. Subgenital plate (Fig. white with narrow transverse red-orange band between ante- 265) with 3-4 preapical macrosetae. Style (Fig. 267 and 268) rior corners of eyes; pronotum red-orange with white band apophysis sinuate, apex hooked mesad, acute. Connective (Fig. across posterior margin; mesonotum red-orange; scutellum 268) Y-shaped, stem as long as arms. Aedeagus (Fig. 267 and white; forewing clavus red-orange with small white mark along 268) with preatrium slightly shorter than shaft, slender in ven- anal margin, large transcommisural diamond-shaped mark near tral view; dorsal apodeme weakly developed; shaft tubular, midlength, and white mark at extreme apex; corium red-or- curved evenly dorsad throughout length, slightly tapered in ange basally, brochosome field white bordered with brown, lateral and ventral views, without processes, apex rounded. distal third smoky hyaline with darker mark in inner apical Female as described for genus. cell and veins bordered with fuscous; venter dull stramineous Material examined. Holotype male, ECUADOR: Orellana, with apex of anteclypeus and front and middle legs infused Res. Etnica Waorani, Transect Ent. 1 km S Onkonegare Camp, with fuscous. 00°39’10"S, 076°26’00"W, 220 m, 7 February 1996, T. L. Erwin Male abdomen (Fig. 339) with 2S apodemes slightly ta- et al., lot 1440, fogging terra firme forest [USNM]. Paratypes: 1 pered and divergent, extended to posterior margin of sternite male, 2 females, same data as holotype; 4 males, 4 females, IV. Pygofer with posterodorsal emargination quadrate, ex- same data except 21 June 1996, lot #1560 [USNM, INHS] tended to base of segment (Fig. 274); dorsal appendage ab- Etymology. This species is named in honor of my nephew, sent; ventral appendage slender, not strongly curved, Mark Schultz. irregularly sinuate through most of length (Figs 273 and 274). Euzyginella quartalberti sp. nov. Anal hook (Fig. 273) long, extended posterad, slightly arcuate, with 3 terminal spines. Subgenital plate (Fig. 273) with 2 Figs 40, 64, 269-272, 338 preapical macrosetae. Style apophysis (Figs 275 and 276) Description. Length of male 3.0 mm. Dorsum (Fig. 40) straight, slender, apex hooked slightly ventrad, acute. Con- mostly red-orange; crown with white anterior and posterior nective (Fig. 276) Y-shaped with stem short and broad, ante- bands; pronotum with posterior margin white; mesonotal tri- rior margin semicircular. Aedeagus (Figs 275 and 276) with angles white; clavus with white basal, medial and apical spots; preatrium slightly longer than shaft; dorsal apodeme com- corium with white medial spot, brochosome field white in- pressed anteriorly with transverse posterior plate; shaft slen- fused with orange and bordered with dark brown on each end; der, tubular, curved dorsad near base, with pair of slender venter stramineous infused with fuscous. lateral processes arising near midlength and extended distad Male abdomen (Fig. 338) with 2S apodemes parallel-sided, along shaft but not reaching shaft apex; shaft apex slightly slightly divergent, extended to midlength of sternite IV. Pygofer broadened, rounded in lateral view and compressed; gonop- with posterodorsal emargination subquadrate (Fig. 270); dorsal ore apical. Female unknown. appendage robust, evenly tapered, apex hooked ventrad; ven- Material examined. Holotype male, ECUADOR: Orellana, tral appendage robust, swollen preapically, apex hooked slightly Tiputini Biodiversity Sta. nr. Yasuni Nat. Pk. 220-250m mesad (Figs 269 and 270). Anal hook (Fig. 269) moderately long, 00°37’55"S, 076°08’39"W, 7 February 1999, T. L. Erwin et al., slender, needlelike, curved slightly dorsad. Subgenital plate (Fig. lot 2040, transect T/5, fogging terra firme forest [USNM]. 269) with 3 preapical macrosetae. Style apophysis (Figs 271 and Etymology. This species is named in honor of my uncle, 272) nearly straight, somewhat broadened preapically, apex Albert E. Dietrich, III.

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 553

Euzyginella theari sp. nov. Male abdomen (Fig. 340) with 2S apodemes robust, parallel sided, extended to posterior margin of sternite IV. Pygofer Figs 42, 261-264, 340 with posterodorsal emargination parabolic (Fig. 262); dorsal Description. Length of male 2.8 mm. Dorsum (Fig. 42) process slender, falcate; ventral appendage slender, somewhat mostly pale-orange; crown with white anterior and posterior arcuate, apex curved slightly laterad (Figs 261 and 262). Anal bands; pronotum with posterior margin white; mesonotal tri- ring with elongate slender spine extended posterad from each angles white; clavus with white basal, medial and apical spots; side (Fig. 261). Subgenital plate (Fig. 261) with 3 preapical corium with white medial spot, brochosome field white in- macrosetae. Style apophysis (Figs 263 and 264) straight, slen- fused with orange and bordered with dark brown on each end; der, apex hooked slightly ventrad, acute. Connective (Fig. 264) venter stramineous infused with fuscous. with stem slender, arms very short, anterior margin slightly

263

264 261 262

267

268 266 265

271

270 272 269

275

276 273 274

Figures 261-276. Euzyginella, male genitalia. (261-264) E. theari sp. nov.: (261) genital capsule) lateral view; (262) pygofer and basal sclerites of anal tube, dorsal view; (263) genitalia, lateral view; (264) genitalia, ventral view, only right style shown; (265-268) E. marki sp. nov., same; (269-272) E. quartalberti sp. nov., same; (273-276) E. teralberti sp. nov., same, anal tube not shown.

ZOOLOGIA 30 (5): 519–568, October, 2013 554 C.H. Dietrich concave. Aedeagus (Figs 263 and 264) with preatrium slightly surface of pygofer lobe or to separately articulated anal hook. shorter than shaft; dorsal apodeme compressed anteriorly with Subgenital plate (Fig. 277) narrow basally, broadened near transverse posterior plate; shaft long, slender, tubular, some- middle, with 3-5 macrosetae in sublateral row near apex; apex what sinuate in lateral view, distal half keeled dorsally, with- weakly to strongly compressed. Connective (Fig. 280) U- or Y- out processes, apex blunt. Female unknown. shaped, articulated to aedeagus, without prominent posterodor- Material examined. Holotype male, ECUADOR: Orellana, sal lobes. Style (Figs 279 and 280) apodeme short, apophysis Tiputini Biodiversity Sta. nr. Yasuni Nat. Pk. 220-250m elongate, with few fine setae preapically, apex tapered and 00°37’55"S, 076°08’39"W, 26 October 1998, T. L. Erwin et al., curved. Aedeagus (Figs 279 and 280) with preatrium well devel- transect T/5, lot 1945, fogging terra firme forest [USNM]. oped, dorsal apodeme weakly to strongly developed, shaft longer Etymology. This species is named in honor of my nephew, than preatrium, with or without paired processes. T.R. Braxton; the name is a phonetic spelling of his first two Female sternite VII (Fig. 379) with posterior margin con- initials. vex; ovipositor elongate, slender, somewhat sigmoid, first valvulae (Fig. 380) with strigate dorsal and ventral sculpturing, Neozyginella gen. nov. second valvulae (Fig. 380) with right blade bearing ~13 small, Type species: N. baileyi sp. nov. close-set teeth near apex of dorsal margin. Diagnosis. Small, depressed leafhoppers (Figs 43-48). Etymology. The genus name, a feminine noun, combines Coloration pale yellow, dorsum with symmetrical bright yel- Neo- (new) with Zyginella, the name of an apparently related low, orange, black and/or red markings; head with two pairs of Old World genus. red spots, small pair on margin of crown near apex, larger pair Remarks. This genus shares a hind wing venational pat- just anteromesad of eyes. Head wider than pronotum, de- tern with Pseudhadina gen. nov., in which the submarginal vein pressed, anterior margin angulately produced; face nearly hori- connects the apices of CuA and MP to form a closed apical cell zontal (Fig. 65); rostrum not extended beyond front but does not extend to MA. This pattern is unknown among trochanters; lower part of face sexually dimorphic, male other Typhlocybini. Neozyginella is easily distinguished from anteclypeus inflated and expanded laterad, lorum very nar- other Typhlocybini by the unique color pattern. The known row; female anteclypeus relatively narrow, flat and tapered, species are recorded from terra firme forests in western Amazonia lorum relatively broad; frontoclypeus convex; lateral frontal and premontane forests of the eastern Andean foothills. sutures not extended dorsad of antennal pits; antennal ledges Key to species of Neozyginella weakly developed; antennae as long as head width; ocelli absent; crown convex, coronal suture restricted to posterior third 1. Male pygofer lobe with three posterior processes (Fig. 285) of crown. Pronotum (Figs 43-48) with lateral margins moder- ...... 3 ately long, slightly divergent in dorsal view, distinctly carinate, 1’. Male pygofer lobe with two posterior processes (Fig. 293).. carina even with posterior margin of eye. Front femur with ...... 2 AM1 enlarged and situated on ventral margin; intercalary row 1". Male pygofer lobe with only one posterior processes, arising with few fine setae, basal seta larger than others; PV1 well de- ventrally (Fig. 281)...... bethae, sp. nov. veloped; tibia somewhat flattened dorsally, AD and PD without preapical macrosetae. Hind femur macrosetae 2+1+1; tibia 2. Male pygofer lobe with processes well separated from each row AV with three macrosetae near apex. Forewing (Fig. 77) other, dorsal process much shorter than ventral process (Fig. with RA reflexed, RP confluent with MA for short distance, 297); aedeagus with pair of massive ventrolateral processes apical cell 2 triangular, inner apical cell trapezoidal; hind wing arising from base, distal processes very short (Fig. 300) ... (Fig. 78) veins RP and MA separate, connected by crossvein, ...... jenncraftae sp. nov. submarginal vein incomplete, extended from jugal lobe to point 2’. Male pygofer lobe with processes closely adjacent to each slightly beyond MP, forming closed cell between MP and CuA. other, subequal in length (Fig. 293); aedeagus without basal Male abdomen (Figs 341-346) with 2S apodemes well de- processes, apex with pair of slender, curved lateral proces- veloped, connected by sclerotized bridge at base but widely sepa- ses (Fig. 296) ...... dougi, sp. nov. rated distally, more or less parallel, usually extended beyond 3. Aedeagus with pair of short basal spines arising below shaft posterior margin of sternite III. Pygofer (Figs 277 and 278) with (Fig. 291) ...... braxtoni, sp. nov. dorsal emargination well developed; tergite well developed but 3’. Aedeagus without pair of basal spines (Fig. 287) ...... 4 short; dorsal part of lobe heavily sclerotized or not, with or without digitiform distal process; ventral appendage well developed, 4. Pygofer lobe with three processes subequal in size; anal hook elongate; one or more additional posterior pygofer spines also acuminate (Fig. 285) ...... bradleyi, sp. nov. present in several species. Valve short, emarginate posteriorly, 4. Pygofer lobe with dorsal process much shorter than medial fused to pygofer. Segment X (Fig. 277) largely membranous with spine and both much shorter than ventral process; anal sclerotized basal ring, lateral ends of ring articulated to internal hook short and blunt (Fig. 277) ...... baileyae, sp. nov.

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 555

Neozyginella baileyae sp. nov. acuminate but weakly sclerotized; ventral appendage elongate, slender, sinuate, apex curved slightly mesad (Figs 281 and 282). Figs 43, 277-280, 341 Anal hook (Fig. 281) absent but anal tube with ventrolateral Description. Length of male 3.3-3.5 mm. Color (Fig. 43) extensions of basal ring well developed. Subgenital plate (Fig. off white, heavily infused with yellow; head with posterior pair 281) with 3 preapical macrosetae. Style apophysis (Figs 283 of red spots relatively large; pronotum, base of clavus, and apex and 284) sinuate, apex bent abruptly mesad, acuminate. Con- of scutellum without brown pigment; pronotum with exten- nective (Fig. 284) robust, V-shaped. Aedeagus (Figs 283 and sive posteromedial orange area extended posterolaterally onto 284) with preatrium moderately long, moderately wide in ven- mesonotum; scutellum apex orange; irregular orange band tral view; dorsal apodeme well developed with compressed extended length of clavus onto corium, ending just short of anterior lobe and transverse posterior plate; shaft slender, tu- apical cells; dark brown maculae present on costal margin at bular, tapered, curved dorsad, apex with pair of short lateral base and apex of brochosome field. spines, basal processes absent; gonopore apical. Female un- Male abdomen (Fig. 341) with 2S apodemes moderately known. wide, bowed laterad, extended to posterior margin of sternite Material examined. Holotype male, PERU: Huanuco, 5km IV. Pygofer posterodorsal emargination quadrate (Fig. 278); N Tingo Maria, 600m, 9°14’49"S, 75°58’59"W, 24 Oct 2002, C.H. posterodorsal spine well developed, extended posterodorsad, Dietrich, merc. vapor light, 02-38-1 [USML]. Paratype: 1 male, with row of fine setae along posteroventral margin; postero- same data [INHS]. mesal spine short, slender, straight, extended posterodorsad Etymology. This species is named in honor of my cousin, along pygofer margin; ventral appendage elongate, slender, Marilyn Beth D. Lindberg. curved dorsad, areolate and curved slightly mesad distally (Figs 277 and 278). Anal hook (Fig. 277) robust, curved posteroven- Neozyginella bradleyi sp. nov. trad, connected by membrane with basal ring of anal tube. Figs 45, 285-288, 343 Subgenital plate (Fig. 277) with 4-5 preapical macrosetae. Style Description. Length of male 3.0 mm. Color (Fig. 45) off apophysis (Figs 279 and 280) slightly sinuate, slightly expanded white, infused with yellow; head with posterior pair if red spots beyond midlength, apex blunt in lateral view, curved laterad relatively small; pronotum with small brown medial spot; then hooked mesad and acute in ventral view. Connective (Fig. forewing clavus with small brown basal spot and diffuse brown 280) broadly U-shaped. Aedeagus (Figs 279 and 280) with maculae more distally, corium with brown spots at base and preatrium short, broad in ventral view; dorsal apodeme well apex of brochosome field small. developed, with compressed anterior lobe and depressed Male abdomen (Fig. 343) with 2S apodemes moderately posterodorsal extension; shaft slender, tubular, sinuate, apex robust, convexly rounded laterally, extended to posterior mar- with pair of small lateral spines, with pair of short basal spines gin of sternite IV. Pygofer posterodorsal emargination V-shaped, arising on atrium below shaft; gonopore apical. Female un- moderately broad (Fig. 286); pygofer lobe strongly attenuated known. into two posteromedial spines appressed to each other and to Material examined. Holotype male, PERU: San Martín, ventral appendage basally but separate and curved dorsad dis- 35km N Tarapoto, Bosque Prot. Cord. Escalera, 900m, 6°27.46’S, tally; ventral appendage long, moderately broad, acuminate, 76°17.24’W, 6-16 Mar 2005, ME Irwin, JD Vasquez, Malaise, nearly straight throughout length (Figs 285 and 286). Anal hook humid montane forest 12-01 [INHS]. Paratypes: 3 males, same (Fig. 285) well developed, curved ventrad, connected by mem- data [INHS] brane to basal ring of anal tube. Subgenital plate (Fig. 285) Etymology. This species is named in honor of my niece, with four preapical macrosetae. Style apophysis (Figs 287 and Bailey Schultz. 288) slender and sinuate in lateral view, broadened near Neozyginella bethae sp. nov. midlength in ventral view, apex blunt. Connective (Fig. 288) broadly U-shaped. Aedeagus (Fig. 287 and 288) with preatrium Figs 44, 281-284, 342 short; dorsal apodeme well developed with large compressed Description. Length of male 3.3 mm. Color (Fig. 44) off anterior lobe and transverse posterior plate; shaft very slender white, infused with yellow; head with posterior pair of red spots and elongate, curved slightly dorsad, apex in ventral view with relatively large; pronotum with large round medial black spot; pair of short, slender spines extended laterad, basal processes apex of scutellum black; forewing clavus with black macula at absent; gonopore apical. Female unknown. base and another near midlength of commisural margin, co- Material examined. Holotype male, BOLIVIA: Cochabamba, rium with black macula at base of brochosome well developed. Villa Tunari, Hotel Los Tucanes, 16°58.302’S, 65°23.793’W, Male abdomen (Fig. 342) with 2S apodemes short and 320m 5-IX-2000, M. Hauser, S. Gaimari, D. Yeates [INHS]. slender, slightly tapered, slightly surpassing posterior margin Paratypes: 2 males, same data [INHS]. of sternite III. Pygofer with posterodorsal emargination quad- Etymology. This species is named in honor of my cousin, rate (Fig. 282); posterodorsal spine absent; pygofer lobe strongly Bradley Dietrich.

ZOOLOGIA 30 (5): 519–568, October, 2013 556 C.H. Dietrich

279

280 277 278

283

281

282 284

287

285

288 286

291 290

289 292

295

294 293 296

299

297 300 298

Figures 277-300. Neozyginella, male genitalia. (277-280) N. baileyae sp. nov.: (277) genital capsule) lateral view; (278) pygofer and basal sclerites of anal tube, dorsal view; (279) genitalia, lateral view; (280) genitalia, ventral view, only right style shown; (281-284) N. bethae sp. nov., same; (285-288) N. bradleyi sp. nov., same; (289-292) N. braxtoni sp. nov., same; (293-296) N. dougi sp. nov., same; (297-300) N. jenncraftae sp. nov., same.

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 557

Neozyginella braxtoni sp. nov. broadened near midlength in ventral view, apex blunt, straight. Connective (Fig. 296) broadly U-shaped. Aedeagus (Figs 295 Figs 46, 289-292, 344 and 296) with preatrium moderately long, moderately broad Description. Length of male 3.0 mm. Color (Fig. 46) off in ventral view; dorsal apodeme well developed with com- white, heavily infused with yellow; head with posterior pair of pressed anterior lobe and posterior transverse plate; shaft in red spots relatively large; small brown spot present medially lateral view broadened toward midlength with prominent dor- on pronotum, one at base of forewing clavus, and one at base sal lobe, then abruptly narrowed and slender to slightly ex- of costal brochosome field; apex of scutellum black. panded apex, broad and parallel sided in posterior view, with Male abdomen (Fig. 344) with 2S apodemes robust, pair of rounded lateral lobes at apex and pair of slender strongly roundly tapered laterally, extended to posterior margin of ster- recurved lateroapical processes, basal processes absent; gonop- nite IV. Pygofer posterodorsal emargination V-shaped (Fig. 290); ore apical. Female as described for genus. posterodorsal spine well developed, extended dorsad, with row Material examined. Holotype male, ECUADOR: Orellana, of fine setae along posteroventral margin; pair of elongate Res. Etnica Waorani, Transect Ent. 1 km S Onkonegare Camp, posterior spines of equal length extended posterad, closely 00°39’10"S, 076°26’00"W, 220 m, 10 February 1996, T. L. Erwin appressed through most of length (Figs 289 and 290). Anal et al., lot 1485, fogging terra firme forest [USNM]. Paratypes: 4 hook (Fig. 289) robust, moderately long, with angulate ventral males and 1 female, same data [USNM, INHS]. projection near midlength, apex acuminate, curved mesad. Etymology. This species is named in honor of my cousin, Subgenital plate (Fig. 289) with four preapical macrosetae. Style Douglas Dietrich. apophysis (Figs 291 and 292) moderately robust, slightly sinuate, apex blunt in lateral view, hooked mesad and acute in ven- Neozyginella jenncraftae sp. nov. tral view. Connective (Fig. 292) broadly U-shaped. Aedeagus Figs 48, 77-78, 297-300, 346 (Figs 291 and 292) with preatrium shorter than shaft, broad in Description. Length of male 3.8 mm, female 3.8 mm. Color ventral view; dorsal apodeme well developed, with compressed (Fig. 48) pale yellow, head with posterior pair of red spots rela- anterior lobe and depressed posterodorsal extension; shaft slen- tively large; small brown spot present medially on pronotum; der, tubular, slightly sinuate in lateral view, without apical pro- apex of scutellum brown; base of forewing clavus with small cesses, base with pair of short posteriorly directed ventrolateral brown spot; costal margin with three black maculae, one near spines; gonopore apical. Female unknown. base, one near midlength and one near apex, wing margin with Material examined. Holotype male, PERU: San Martín, several additional diffuse brown markings; venter pale yellow. 35km N Tarapoto, Bosque Prot. Cord. Escalera, 900m, 6°27.46’S, Male abdomen (Fig. 346) with 2S apodemes small, tapered, 76°17.24’W, 6-16 Mar 2005, ME Irwin, JD Vasquez, Malaise, extended to midlength of sternite III. Pygofer posterodorsal humid montane forest 12-01 [INHS]. emargination deep, quadrate (Fig. 298); dorsal appendage mov- Etymology. This species is named in honor of my nephew, ably articulated with margin and fused to anal hook arising from Braxton Schultz. ringlike basal sclerite of segment X and terminating in long Neozyginella dougi sp. nov. posteroventrally directed spine; posterodorsal process short, tapered to acute angle, extended posterodorsad, with row of fine Figs 47, 293-296, 345, 379-380 setae along posteroventral margin; ventral process elongate, slen- Description. Length of male 3.0 mm. Color (Fig. 47) off der, extended posterad to apex of subgenital plate, weakly curved white, infused with yellow; head with posterior pair of red spots dorsomesad (Figs 297 and 298). Subgenital plate (Fig. 297) with relatively large; brown spot present medially on pronotum, at five preapical macrosetae. Style apophysis (Figs 299 and 300) base of forewing clavus, and at base of costal brochosome field; moderately robust, slightly sinuate in lateral view, apex tapered, apex of scutellum, male subgenital plate, and female third val- hooked mesad and acute in ventral view. Connective (Fig. 300) vulae also brown. broadly U-shaped. Aedeagus (Figs 299 and 300) with preatrium Male abdomen (Fig. 345) with 2S apodemes slender, very short; dorsal apodeme well developed, columnlike; shaft bowed laterad, extended to posterior margin of sternite IV. very slender, tubular, slightly sinuate in lateral view, with pair Pygofer posterodorsal emargination U-shaped, moderately of short retrorse lateral spines apically, base with pair of robust broad (Fig. 294); dorsal process absent; posterior lobe rounded, posteriorly directed ventrolateral spines as long as shaft; gonop- posteromedial appendage present just dorsad of and closely ore apical. Female as described for genus. appressed to ventral appendage; both appendages slender, Material examined. Holotype male, PERU: Pasco, acuminate and curved dorsomesad (Figs 293 and 294). Anal Yanachaga-Chemillén N.P., Huampal Stn. 10°11’9"S 75°34’27"W hook (Fig. 293) robust, moderately long, curved ventrad, apex 6-9 X 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap acr. R. connected to basal ring of anal tube by sclerotized bar. Huancabamba [USML]. Paratype: 1 female, same data [INHS]. Subgenital plate (Fig. 293) with four preapical macrosetae. Style Etymology. This species is named in memory of my niece, apophysis (Figs 295 and 296) slender and sinuate in lateral view, Jennifer Craft.

ZOOLOGIA 30 (5): 519–568, October, 2013 558 C.H. Dietrich

Pseudhadina gen. nov. Remarks. This genus is based on a single species from the eastern Ecuadorean rainforest canopy. It appears to be Type species P. amazonica sp. nov. morphologically intermediate between Neozyginella and Diagnosis. Small, depressed leafhoppers (Fig. 49). Over- Columbonirvana. It is readily distinguished from other South all color stramineous, dorsum with symmetrical dark brown American Typhlocybini by the narrow, flattened (not sexually markings; forewing apical cell 2 with false eyespot, costal mar- dimorphic) male anteclypeus. It resembles Eualebra in shape gin with 2 oblique dark brown lines. Head (Fig. 49) wider than and proportions but differs in having the coronal suture ex- pronotum, depressed, anterior margin angulately produced; tended nearly to the apex of the crown and the face in profile face nearly horizontal (Fig. 66); rostrum not extended beyond oblique rather than horizontal. front trochanters; male anteclypeus not inflated (Fig. 58), flattened and tapered; lorum long and narrow; frontoclypeus flat- Pseudhadina amazonica sp. nov. tened; lateral frontal sutures not extended dorsad of antennal Figs 49, 58, 66, 79-80, 301-304, 347, 381-383 pits; antennal ledges weakly developed; antennae shorter than head width; ocelli absent; crown flat, coronal suture extended Description. Length of male 3.4 mm. Crown unmarked to apex of crown. Pronotum (Fig. 49) with lateral margins short, (Fig. 49); pronotum with narrow lateral longitudinal stripe, slightly divergent in dorsal view, distinctly carinate, carina even two transverse posterior lines, and posterior margin dark brown; with posterior margin of eye. Front femur with AM1 enlarged mesonotum with 4 small brown spots in transverse row; forew- and situated on ventral margin; intercalary row with few fine ing clavus with narrow brown arch extended posteriorly onto setae, basal seta much larger than others; PV1 stout; tibia corium, veins in distal half yellow bordered with dark brown; rounded dorsally, AD and PD without preapical macrosetae. venter of head and thorax brown, legs stramineous. Hind femur macrosetae 2+1+1; tibia row AV with 4 macrosetae Male abdomen (Fig. 347) with 2S apodemes moderately near apex. Forewing (Fig. 79) with RA reflexed, RP confluent robust, falcate, extended to midlength of sternite IV. Pygofer with MA for short distance, apical cell 2 triangular, inner api- dorsal emargination rounded, tergite 1/3 total length of pygofer cal cell trapezoidal; hind wing (Fig. 80) with RP and MA sepa- (Fig. 302); dorsal process short, bent mesad; lobe concave rate, connected by crossvein; submarginal vein absent between posterodorsally, rounded apically; ventral appendage curved RP and MP+CuA.Male abdomen (Fig. 347) with 2S apodemes dorsad in lateral view, weakly contorted and curved mesad dis- well developed, connected by sclerotized bridge at base but tally in dorsal view (Figs 301 and 302). Subgenital plate (Fig. widely separated distally, more or less parallel. Pygofer (Fig. 301) with 4 macrosetae. Style apophysis (Figs 303 and 304) 301-302) with dorsal emargination well developed; tergite well abruptly narrowed preapically, apex curved ventrolaterad, developed; lobe evenly sclerotized, with dorsal process; ven- blunt. Aedeagus (Figs 303 and 304) with preatrium very nar- tral appendage well developed, elongate. Valve short, emar- row at base in ventral view, atrium broad with two pairs of ginate posteriorly, fused to pygofer. Segment X (Fig. 301) largely lateral lobes; shaft strongly depressed, slender and slightly ar- membranous with sclerotized basal ring, lateral ends of ring cuate in lateral view, broad and nearly parallel sided with articulated to internal surface of pygofer lobe and extended slightly expanded, rounded apex in ventral view. Female ventrad into anal hook. Subgenital plate (Fig. 301) narrow ba- terminalia as described for genus. sally, broadened near middle, with 4 macrosetae in sublateral Material examined. Holotype male, ECUADOR: Orellana, row near apex; apex weakly compressed, with preapical lateral Tiputini Biodiversity Sta. nr. Yasuni Nat. Pk. 220-250m row of long fine setae and apical group of short rigid setae. 00°37’55"S, 076°08’39"W, 8 February 1999, T. L. Erwin et al., Connective (Fig. 304) U-shaped, articulated to aedeagus, with lot# 2028, transect T/3, fogging terra firme forest [USNM]. small posterodorsal lobes. Style apodeme short, apophysis elon- Paratypes: 1 female, same data except 26 October 1998, transect gate, with few fine setae preapically, apex tapered and curved. T/6, lot# 1950; 1 male, ECUADOR: Orellana, Res. Etnica Aedeagus (Figs 303 and 304) with preatrium very short, dorsal Waorani, Transect Ent. 1 km S Onkonegare Camp, 00°39’10"S, apodeme absent, shaft depressed, with paired lateral flanges; 076°26’00"W, 220 m, 4 October 1995, T. L. Erwin et al., lot gonopore apical. 1188, fogging terra firme forest; 1 male, same data except 8 Female abdominal sternite VII (Fig. 381) ~1.5X longer October 1995, lot# 1263 [USNM, INHS]. than sternite VI, posterior margin truncate with small median notch and paired lateral lobes; first valvulae with strigate sculp- Pseudozyginella gen. nov. turing; second valvulae (Figs 382 and 383) relatively elongate Type species: P. grahami sp. nov. and slender, evenly curved dorsad through most of length, apex Diagnosis. Small, depressed leafhoppers (Figs 50-52). Col- slightly sigmoid, dorsal margin of apex with ~12 small, close- oration mostly brown with large, symmetrical white markings set teeth. dorsally. Head (Figs 50-52) wider than pronotum, depressed, Etymology. The name, a feminine noun, was formed by anterior margin angulately produced; face nearly horizontal combining “pseud-”, meaning false, with Eurhadina, a superfi- (Fig. 67); rostrum not extended beyond front trochanters; lower cially similar Old World genus. part of face sexually dimorphic, male anteclypeus inflated and

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 559 expanded laterad, lorum very narrow; female anteclypeus rela- Remarks. This is the only New World typhlocybine ge- tively narrow, tapered and flat, lorum relatively broad; nus known to include species with hind wing venation resem- frontoclypeus convex; lateral frontal sutures not extended dor- bling that of the Old World genus Zyginella, although the sad of antennal pits; antennal ledges weakly developed; an- Zyginella-like features (i.e., absence of a distal segment of CuA tennae as long as head width; ocelli absent; crown convex, extended beyond the submarginal vein) are not consistently coronal suture restricted to posterior third of crown. Pronotum present and, in some cases, vary between the right and left (Figs 50-52) with lateral margins moderately long, slightly di- wings of a single specimen. Thus, for members of Pseudozygi- vergent in dorsal view, distinctly carinate, carina even with nella, at least, the criteria previously used by some previous posterior margin of eye. Front femur with AM1 enlarged and authors to justify recognition of Zyginellini as a separate tribe situated on ventral margin; intercalary row with few fine se- are unstable, providing support for the present treatment of tae, basal seta larger than others; PV1 well developed; tibia Zyginellini as a junior synonym of Typhlocybini. The new ge- somewhat flattened dorsally, AD and PD without preapical nus differs from Zyginella and other genera placed by macrosetae. Hind femur macrosetae 2+1+1; tibia row AV with Dworakowska (1979) in “Zyginellini” in having a well-devel- three macrosetae near apex. Wing venation variable intra- and oped, elongate ventral pygofer process similar to that of interspecifically; forewing (Fig. 81) inner apical cell trapezoi- Columbonirvana. It also resembles Columbonirvana in having a dal; hind wing (Figs 82 and 83) submarginal vein absent be- well-developed anal hook articulated to the pygofer and anal tween CuA and RP, RP and MA separate, connected by crossvein tube, a slender apically setose style, and subgenital plates with or confluent; distal segment of CuA, beyond connection with a reduced number of macrosetae. The genus includes three submarginal vein, present or (more rarely) absent. species, all collected by fogging terra firme rainforest canopy Male abdomen (Figs 348-350) with 2S apodemes vari- at a single locality in eastern Ecuador. ably developed, connected by sclerotized bridge at base but Key to species of Pseudozyginella widely separated distally, more or less parallel, usually extended beyond posterior margin of sternite III. Pygofer (Figs 305 and 1. Male subgenital plate without one or more macrosetae at 306) with dorsal emargination well developed, broadly U- apex (Fig. 305); aedeagus without pair of apical processe... shaped; tergite well developed but short; lateral lobe tapered ...... P. grahami sp. nov. from base to apex, dorsal part heavily sclerotized, with 1’. Male subgenital plate with one or more macrosetae at apex posteroventrally directed distal process; ventral appendage well (Fig. 313); aedeagus with pair of apical processes (Fig. 312) developed, elongate, extended posterodorsad along inner sur- ...... 2 face of lobe. Segment X (Fig. 305) largely membranous but with 2. Forewing mostly brown with two white spots near midlength, well sclerotized and darkly pigmented dorsal ring basally bear- one along costal margin and one in clavus (Fig. 52); ing falcate posterolateral process and anterolateral projection aedeagus in ventral view with preatrium evenly tapered articulated to dorsal margin of pygofer lobe. Dorsal connec- toward base (Fig. 316) ...... P. dougbraxtoni sp. nov. tive absent. Valve short, emarginate posteriorly, fused to 2’. Forewing white in basal half, brown distally (Fig. 51); pygofer. Subgenital plate (Fig. 305) very narrow basally, slightly aedeagus in ventral view with preatrium constricted near broadened near middle, tapered distally, with 1-2 macrosetae midlength (Fig. 312) ...... P. davei sp. nov. slightly beyond midlength and, usually 1-2 macrosetae at apex, apex compressed. Connective (Fig. 308) U-shaped, stem absent, articulated to aedeagus, without prominent posterodorsal Pseudozyginella grahami sp. nov. lobes. Style (Figs 307 and 308) apodeme short, apophysis elon- Figs 50, 67, 81-82, 305-308, 348, 384-387 gate, with few fine setae preapically, apex tapered and curved. Description. Length of male 2.5-2.6 mm, female 2.6 mm. Aedeagus (Figs 307 and 308) with preatrium well developed, Color (Fig. 50) off white, heavily infused with brown, crown dorsal apodeme weakly to strongly developed, shaft longer than uniformly pale, pronotum and basal triangles of mesonotum preatrium, with pair of distally directed spines arising near base brown; forewing base and apex brown, interrupted medially of shaft, with or without apical processes, gonopore apical. by large transcommisural pale area containing irregular areas Female abdominal sternite VII (Fig. 384) produced pos- of orange in some specimens, widest medially and narrowest teriorly; ovipositor moderately elongate, curved dorsad, some- at costal margin; corresponding areas of abdomen pale with what expanded preapically, first valvulae (Fig. 385 and 386) brown areas at base and apex including pygofer. with strigate dorsal and ventral sculpturing, second valvulae Male abdomen (Fig. 348) with 2S apodemes robust, nearly (Fig. 387) with right blade bearing ~10 large, close-set, quad- parallel sided, extended to midlength of sternite V. Pygofer (Figs rate teeth near apex of dorsal margin. 305 and 306) with dorsal appendage slender, acuminate, bent Etymology. The genus name, a feminine noun, combines posteroventrad and extended well beyond ventral margin of “pseudo-” meaning “false” with Zyginella, and reflects the mor- lobe; ventral appendage robust, weakly convoluted through phological similarity to the latter genus. most of length, apex bent laterad then hooked slightly mesad.

ZOOLOGIA 30 (5): 519–568, October, 2013 560 C.H. Dietrich

Anal hook (Fig. 305) moderately long, slender, acuminate, 00°39’10"S, 076°26’00"W, 220 m, 2 July 1995, T. L. Erwin et al., curved posteroventrad. Subgenital plate (Fig. 305) with 2 lot 1068, fogging terra firme forest [USNM]. Paratypes: 2 fe- preapical macrosetae. Style apophysis (Figs 307 and 308) males, same data; 1 female, same data except 10 February 1996, slightly tapered through most of length, nearly straight, apex lot# 1496 [USNM, INHS]. hooked strongly mesad, acuminate. Connective (Fig. 308) ro- Etymology. This species is named in honor of my nephew, bust, V-shaped. Aedeagus (Fig. 307 and 308) with preatrium David Braxton. shorter than shaft, parallel sided in ventral view; dorsal apodeme weakly developed; shaft slender, tubular, curved dor- Pseudozyginella dougbraxtoni sp. nov. sad, with pair of short slender spines arising near midlength, Figs 52, 313-316, 350 only slightly divergent from shaft, and extended slightly beyond shaft midlength; shaft apex slightly expanded and ob- Description. Length of male 2.4-2.5mm. Color (Fig. 52) liquely truncate in lateral view, without processes. Female mostly brown, crown uniformly pale, pronotum and basal tri- abdominal sternite VII (Fig. 384) with posterior margin ob- angles of mesonotum brown; forewing with brochosome field tusely angulate; ovipositor as described for genus. white and small transcommisural white spot relatively small Material examined. Holotype male, ECUADOR: Orellana, and restricted to clavus; abdomen brown. Res. Etnica Waorani, Transect Ent. 1 km S Onkonegare Camp, Male abdomen (Fig. 350) with 2S apodemes robust, nearly 00°39’10"S, 076°26’00"W, 220 m, 7 October 1995, T. L. Erwin parallel sided, extended to midlength of sternite V. Pygofer (Figs et al., lot 1246, fogging terra firme forest [USNM]. Paratypes: 1 313 and 314) with dorsal appendage robust, tapered, evenly male, 1 female, same data except 10 February 1996, lot#1496; curved posteroventrad and extended slightly beyond margin 1 male, same data except 21 June 1996, lot# 1549 [USNM, of lobe; ventral appendage robust, weakly convoluted through INHS]. most of length, apex curved slightly laterad. Subgenital plate Etymology. This species is named in honor of my nephew, (Fig. 313) with 2 macrosetae near midlength and 2 at apex. Graham Dietrich. Style apophysis (Figs 315 and 316) slightly tapered through most of length, nearly straight, apex bent ventrad and curved Pseudozyginella davei sp. nov. slightly mesad, acuminate. Connective (Fig. 316) broadly U- Figs 51, 309-312, 349 shaped, arms stout. Aedeagus (Fig. 315 and 316) with preatrium slightly shorter than shaft, evenly tapered toward base in ven- Description. Length of male 2.4 mm, female 2.5 mm. tral view; dorsal apodeme weakly developed; shaft slender, tu- Color (Fig. 51) off white, heavily infused with brown, crown bular, curved slightly dorsad, with basolateral processes uniformly pale, pronotum and basal triangles of mesonotum relatively large, evenly divergent from shaft in ventral view, brown; forewing with basal half white, distal half brown; ab- extended well beyond shaft midlength; with pair of slender domen pale dorsally except near apex. Forewing veins RP and retrorse lateral processes at apex, extended anteroventrad. Fe- MA confluent for short distance distally. male unknown. Male abdomen (Fig. 349) with 2S apodemes vestigial (pos- Material examined. Holotype male, ECUADOR: Orellana, sibly due to parasitism of only available male specimen). Res. Etnica Waorani, Transect Ent. 1 km S Onkonegare Camp, Pygofer (Figs 309 and 310) with dorsal appendage robust, ta- 00°39’10"S, 076°26’00"W, 220 m, 2 July 1995, T. L. Erwin et al., pered, evenly curved posteroventrad and extended slightly lot# 1069, fogging terra firme forest [USNM]. Paratypes: 2 males, beyond margin of lobe; ventral appendage relatively slender, same data except 8 October 1995, lot# 1266; 1 male, same data curved slightly dorsad through most of length, apex curved except 12 February 1995, lot# 1239 [USNM, INHS]. slightly laterad. Subgenital plate (Fig. 309) with 1-2 macrosetae Etymology. This species is named in honor of my brother- near midlength and 1 at apex. Style apophysis (Figs 311 and in-law, Douglas Braxton. 312) slightly tapered through most of length, nearly straight, apex bent ventrad and curved slightly mesad, acuminate. Con- Tahurella gen. nov. nective (Fig. 312) broadly U-shaped, arms slender. Aedeagus (Figs 311 and 312) with preatrium slightly shorter than shaft, Type species: T. lynnae, sp. nov. relatively slender and constricted near midlength in ventral Diagnosis. Medium-sized, depressed leafhoppers (Figs 53- view; dorsal apodeme weakly developed; shaft slender, tubu- 57). Dorsal coloration pale with symmetrical arcuate lar, curved slightly dorsad, with basal processes relatively large, transcommisural bands across forewings; forewing apex with evenly divergent from shaft in ventral view, extended well be- false eyespot, costal margin with 6 or more oblique dark brown yond shaft midlength; with pair of slender retrorse lateral pro- false veins. Head (Figs 53-57) narrower than pronotum, de- cesses at apex, extended anteroventrad. Female as described pressed, anterior margin angulately produced; face nearly hori- for genus. zontal (Fig. 68); rostrum extended slightly beyond front Material examined. Holotype male, ECUADOR: Orellana, trochanters; male anteclypeus inflated and expanded laterad Res. Etnica Waorani, Transect Ent. 1 km S Onkonegare Camp, (Fig. 59); lorum very narrow; frontoclypeus convex; lateral fron-

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 561

303

302

301

304

306 307

305 308

309 311 310

312

314 315

313 316

Figures 301-316. (301-304) Pseudhadina amazonica sp. nov., male genitalia: (301) genital capsule, lateral view; (302) pygofer and basal sclerites of anal tube, dorsal view; (303) genitalia, lateral view; (304) genitalia, ventral view, only right style shown. (305-308) same, Pseudozyginella grahami sp. nov.; (309-312) same P. davei sp. nov.; (313-316) same, P. dougbraxtoni sp. nov. tal sutures not extended dorsad of antennal pits; antennal separate and joined by crossvein or confluent for short dis- ledges weakly developed; antennae as long as head width; ocelli tance; CuA joining M basad of its fork; inner apical cell trap- absent; crown flat, coronal suture extended to crown apex. ezoidal; apical margin with distinct concavity. Hind wing (Fig. Pronotum (Figs 53-57) with lateral margins moderately long, 86) venation as in Eualebra. slightly divergent in dorsal view, distinctly carinate, carina even Male 2S apodemes (Figs 351-355) well developed, joined with posterior margin of eye. Front femur with AM1 enlarged at base by transverse bridge, variable interspecifically in shape. and situated on ventral margin; intercalary row with few fine Pygofer (Figs 317 and 318) strongly emarginate dorsally; terg- setae, basal seta larger than others; PV1 absent; tibia rounded ite variable in length and shape among species; posterior lobe dorsally, AD and PD without preapical macrosetae. Hind fe- with strongly sclerotized, distally falcate dorsal section vari- mur macrosetae 2+1+1; tibia row AV with 4 macrosetae near ably clothed with microtrichia, unpigmented ventral section apex. Forewing (Figs 84 and 85) with RA reflexed, RP and MA bearing longitidinal row or band of short fine setae; ventral

ZOOLOGIA 30 (5): 519–568, October, 2013 562 C.H. Dietrich appendage slender, acuminate, arising near base of ventral thoracic pleuron; pronotum with anterior margin red-orange, margin and curved posterodorsad, usually areolate distally. Anal posterior margin with convex red-orange marking medially; tube (Figs 317 and 318) depressed, sclerotized dorsal band short, mesonotum orange anteriorly; scutellum orange at apex. Forew- with pair of basolateral hooks. Dorsal connective absent. ing with series of 7 oblique brown lines distributed along costal Subgenital plate (Fig. 317) constricted near base, broadest near margin; basal third of corium and clavus hyaline with narrow midlength, with row of 3-4 macrosetae near lateral margin and red-orange transcommisural arcuate band; distal 2/3 of forew- row of smaller setae ventrally near apex; apical lobe compressed, ing smoky hyaline with paler areas in apical cells; veins orange; parallel sided and darkly pigmented. Style (Figs 319 and 320) distal lobe well developed. Thoracic venter pale yellow; abdo- apodeme short; apophysis elongate, without preapical lobe, men pale yellow ventrally except sternite VIII and valve brown. with several fine setae preapically. Connective (Fig. 320) Y- Male 2S apodemes (Fig. 351) robust, close together at shaped with stem short and broad, articulated to aedeagus. base, evenly tapered toward apex, extended to midlength of Aedeagus (Figs 319 and 320) with preatrium elongate, dorsal sternite IV. Pygofer with dorsal emargination broad, quadrate apodeme weakly developed; shaft elongate and nearly straight, (Fig. 318); tergite ca. 1/4 length of distal lobe; dorsal sclero- usually with paired ventral preapical spines or processes; api- tized area of lobe broad, truncate apically and terminating ven- cal processes absent; gonopore apical. Female unknown. trally in short ventrally directed spine; ventral appendage Etymology. The genus name, a feminine noun, is a di- gradually curved dorsad in lateral view, apex bowed laterad in minutive form of Tahura, a genus of Nirvanini that is superfi- dorsal view (Figs 317 and 318). Anal hook (Fig. 317) small, cially similar in having distal projections on the forewings. triangular. Subgenital plate (Fig. 317) with 2-4 macrosetae near Remarks. This genus is structurally similar to Columbonir- midlength and another 5 at apex. Style apophysis (Figs 319 vana but differs in the larger body size, paler dorsal coloration, and 320) gradually tapered distally, apex curved ventrolaterad, emarginate forewing apex, and Y-shaped connective. The five blunt. Connective (Fig. 320) stem short and broad. Aedeagus known species were collected in Malaise traps in cloud forests (Figs 319 and 320) with preatrium relatively short; narrow at at two localities in Colombia and Peru. base; shaft broad basally and slightly sinuate in lateral view, with pair of slender ventral processes separated from but par- Key to species of Tahurella alleling shaft and extended 3/4 distance to shaft apex, diver- 1. Anterior margin of head with two parallel transverse black gent in ventral view; apex compressed with ventral keel, bands (Fig. 68); male pygofer with distal spine elongate rounded in lateral view. (Fig. 325); aedeagal processes not diverging from shaft in Material examined. Holotype male, COLOMBIA: Nariño, RN ventral view (Fig. 328) ...... 2 La Planada Parcela Olga, 1°15’N 78°15’W 1850m, Malaise 16.vii- 1’. Anterior margin of head with one transverse black band; 2.viii.2001, G. Oliva leg. M.2402 [HIC]. Paratype: 1 male, COLOM- male pygofer with distal spine short (Fig. 317); aedeagal BIA: Nariño, RN La Planada Parcela Permanente 1°15’N 78°15’W processes diverging from shaft in ventral view (Fig. 320) or 1885m Malaise 2-16.vii.2001, G. Oliva leg. M.2404 [INHS] absent (Fig. 336)...... 3 Etymology. This species is named in honor of my nephew, Joseph Dietrich. 2. Pygofer lobe distinctly emarginate apically (Fig. 329) ...... T. clairae sp. nov. Tahurella katherinae sp. nov. 2’. Pygofer lobe entire apically (Fig. 325) ....T. lynnae sp. nov. Figs 54, 321-324, 352 3. Aedeagus without processes or spines (Fig. 335) ...... Description. Length of male 5.0 mm. Dorsum (Fig. 54) ...... T. heatherae sp. nov. pale yellow with symmetrical red-orange and dark brown mark- 3’ Aedeagus with paired ventral processes or spines (Fig. 319). ings; head anterior margin with narrow dark brown band be- ...... 4 tween eyes, discontinuous with similar band extended across 4. Aedeagal processes very short, spinelike (Fig. 323) ...... thoracic pleuron; pronotum with anterior margin red-orange, ...... T. katherinae sp. nov. posterior margin with convex red-orange marking medially; 4’ Aedeagal processes elongate, extended nearly to shaft apex mesonotum orange anteriorly; scutellum orange at apex. Forew- (Fig. 319) ...... T. josephi sp. nov. ing with series of 7 oblique brown lines distributed along costal margin; basal third of corium and clavus hyaline with Tahurella josephi sp. nov. narrow red-orange transcommisural arcuate band; distal 2/3 of forewing smoky hyaline with paler areas in apical cells; veins Figs 53, 317-320, 351 orange; apical lobe well developed. Thoracic venter pale yel- Description. Length of male 4.8 mm. Dorsum (Fig. 53) low; abdomen pale yellow ventrally except sternite VIII and pale yellow with symmetrical red-orange and dark brown mark- valve brown. Head slightly narrower than pronotum, crown ings; head anterior margin with narrow dark brown band be- twice as long medially as next to eye; forewing apical cell 2 tween eyes, discontinuous with similar band extended across quadrate; hind tibia row AV with 4 macrosetae.

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 563

Male 2S apodemes (Fig. 352) robust, bowed laterad, ex- preatrium relatively short, not broadened basally in ventral view; tended to midlength of sternite IV. Pygofer with dorsal emar- shaft slender, compressed, nearly straight, with two pairs of small gination parallel sided, base broadly V-shaped (Fig. 322); tergite ventral spines, one near midlength and one near apex; apex ca. 1/3 length of distal lobe; apical process short, toothlike; ven- compressed, broadened in lateral view and obliquely rounded. tral appendage weakly curved dorsomesad (Figs 321 and 322). Material examined. Holotype male, PERU: Pasco, Anal hook (Fig. 321) small, triangular. Subgenital plate (Fig. 321) Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, with 3-4 macrosetae near midlength and 0-2 stout apical setae. 10-13 October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise Style apophysis (Figs 323 and 324) slightly broadened medially, trap across Rio San Alberto [USML]. apex curved ventrolaterad and bluntly rounded. Connective (Fig. Etymology. This species is named in honor of my niece, 324) stem short and broad. Aedeagus (Figs 323 and 324) with Katherine Dietrich.

319

317 318 320

323

324 321 322

327

326

328 325

331

332 329 330

335

336

333 334 Figures 317-336. Tahurella, male genitalia. (317-320) T. josephi sp. nov.: (317) genital capsule) lateral view; (318) pygofer and basal sclerites of anal tube, dorsal view; (319) genitalia, lateral view; (320) genitalia, ventral view, only right style shown. (321-324) T. katherinae sp. nov., same. (325-328) T. lynnae sp. nov., same. (329-332) T. clairae sp. nov., same. (333-336) T. heatherae sp. nov., same.

ZOOLOGIA 30 (5): 519–568, October, 2013 564 C.H. Dietrich

Tahurella lynnae sp. nov. Male 2S apodemes (Fig. 353) weakly divergent, extended to posterior margin of sternite IV. Pygofer with dorsal emar- Figs 55, 68, 84, 325-328, 353 gination broad, parabolic (Fig. 326); tergite ca. 1/3 length of Description. Length of male 5.3 mm. Dorsum (Fig. 55) distal lobe; apical process long, slender, extended ventrad; yellow to white, heavily marked with dark brown and orange; ventral appendage weakly curved dorsomesad (Figs 325 and head anterior margin with pair of thin brown transverse bands 326). Anal hook (Fig. 325) small, triangular. Subgenital plate between eyes, crown yellow; pronotum white with anterior (Fig. 325) with 2-4 macrosetae near midlength and two small margin red-orange and incomplete posterior submarginal or- stout apical setae. Style apophysis (Figs 327 and 328) parallel ange transverse band; mesonotum and scutellum orange; forew- sided through most of length, apex curved laterad then bent ing basal half with alternating white, red-orange, white, and mesad, blunt. Connective (Fig. 328) Y-shaped, stem short and dark brown transcommisural arcuate bands; brochosome field broad. Aedeagus (Figs 327 and 328) with preatrium relatively bright yellow, costal margin with 6 oblique brown bands, veins short, slightly broadened basally in ventral view; shaft very yellow, cells with large fuscous areas; thorax and abdomen slender, slightly sinuate, with pair of long slender processes brown ventrally. Head subequal to pronotum in width, crown arising near base, closely appressed to each other, separated slightly less than twice as long medially as next to eyes. Forew- from shaft through most of length, contacting shaft apically; ing with apical cell 2 triangular, apical lobe short (Fig. 84). apex compressed, expanded in lateral view, obliquely trun- Hind tibia row AV with 4 macrosetae. cate.

338 339 337 340

342 343 344 341

347 346 345 348

349 351 350 352

355 353 354 Figures 337-355. Euzyginella, Neozyginella, Pseudhadina, and Tahurella, base of male abdomen, ventral view. (337) Euzyginella marki sp. nov.; (338) E. quartalberti sp. nov.; (339) E. teralberti sp. nov.; (340) E. theari sp. nov.; (341) Neozyginella baileyae sp. nov.; (342) N. bethae sp. nov.; (343) N. bradleyi sp. nov.; (344) N. braxtoni sp. nov.; (345) N. dougi sp. nov.; (346) N. jenncraftae sp. nov., left apodeme broken; (347) Pseudhadina amazonica sp. nov.; (348) Pseudozyginella grahami sp. nov.; (349) P. davei sp. nov.; (350) P. dougbraxtoni sp. nov.; (351) Tahurella josephi sp. nov.; (352) T. katherinae sp. nov.; (353) T. lynnae sp. nov.; (354) T. clairae sp. nov.; (355) T. heatherae sp. nov.

ZOOLOGIA 30 (5): 519–568, October, 2013 South American leafhoppers of the tribe Typhlocybini 565

358

356

357

359

362 360

361

364

363

366

369 365

367 370

373

368

372

374 371

Figures 356-374. Female seventh sternite and ovipositor. (356) Columbonirvana aurea, holotype, first valvula; (357) same, second valvula, ventral part of apex broken. Eualebra charlesi sp. nov.: (358) sternite VII; (359) first valvula; (360) second valvulae. E. helenae sp. nov.: (361) sternite VII; (362) ovipositor. E. marilynae sp. nov.: (363) sternite VII; (364) first valvula; (365) second and third valvulae; (366) detail of second valvulae apex. E. gingerae sp. nov.: (367) sternite VII. E. susanae sp. nov.: (368) sternite VII; (369) first valvula; (370) second valvulae. E. margaretannae sp. nov.: (371) sternite VII; (372) first valvula; (373) same, detail of apex; (374) second and third valvulae.

ZOOLOGIA 30 (5): 519–568, October, 2013 566 C.H. Dietrich

376

375

377 378

379 383 380

381 382 386

385

384 387

Figure 375-387. Female seventh sternite and ovipositor. Euzyginella marki sp. nov.: (375) sternite VII; (376) apex of first valvula; (377) second and third valvulae; (378) apex of second valvula. Neozyginella dougi sp. nov.: (379) sternite VII; (380) apex of first and second valvulae. Pseudhadina amazonica sp. nov.: (381) sternite VII; (382) second and third valvulae; (383) apex of second valvula. Pseudozyginella grahami sp. nov.: (384) sternite VII; (385) first valvula; (386) same, apex; (387) second and third valvulae.

Material examined. Holotype male, PERU: Pasco, Yanacha- bright yellow, costal margin with 6 oblique brown bands, veins ga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 yellow, cells with large fuscous areas; thorax and abdomen brown October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across ventrally; apical lobe weakly developed. Head subequal to Rio San Alberto [USML]. Paratypes: 4 males, same data [INHS]. pronotum in width, crown slightly less than twice as long me- Etymology. This species is named in honor of my niece dially as next to eyes. Forewing with apical cell 2 triangular, Lynn Dietrich. apical margin concave. Hind tibia row AV with 4 macrosetae. Male 2S apodemes (Fig. 354) weakly divergent, extended Tahurella clairae sp. nov. nearly to posterior margin of sternite IV. Pygofer with dorsal Figs 56, 329-332, 354 emargination relatively narrow, parabolic (Fig. 330); tergite ca. Description. Length of male 4.7 mm. Dorsum (Fig. 56) 1/3 length of distal lobe; lobe with distinct apical emargin- yellow to white, heavily marked with dark brown and orange; ation above distal spine, spine nearly straight; ventral append- head anterior margin with pair of thin brown transverse bands age curved dorsad, extended into emargination of pygofer lobe between eyes, crown yellow; pronotum white with anterior (Figs 329 and 330). Anal hook (Fig. 329) short, blunt. Subgenital margin red-orange and incomplete posterior submarginal or- plate (Fig. 329) with 3 macrosetae near midlength, without ange transverse band; mesonotum and scutellum orange; forew- stout preapical setae. Style apophysis (Figs 331 and 332) paral- ing basal half with alternating white, red-orange, white, and lel-sided through most of length, apex curved ventrolaterad, dark brown transcommisural arcuate bands; brochosome field blunt. Connective (Fig. 332) V-shaped, stem poorly developed.

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Aedeagus (Fig. 331 and 332) with preatrium nearly as long as ACKNOWLEDGMENTS shaft, slightly broadened basally in ventral view; shaft slender, compressed distally, constricted preapically in lateral view, I am indebted to Carlos Peña, Pedro Lozada and Gerardo slightly sinuate, with pair of long slender processes arising near Lamas for helping facilitate fieldwork in Peru, to Terry Erwin for midlength and extended ventrally along shaft nearly to apex, providing access to his canopy fogging samples from Western closely parallel to each other in ventral view; apex obliquely Amazonia, to Paul Freytag and Michael Sharkey for providing rounded in lateral view. access to Malaise trap samples from Colombia, and to Mike Irwin Material examined. Holotype male, PERU: Pasco, for donating Malaise trap samples from Bolivia and Peru. I am Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, also grateful to Norman Penny (California Academy of Sciences) 10-13 October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise for providing photos of type material, to Chen Young (Carnegie trap across Rio San Alberto [USML]. Museum) and Stuart McKamey (U.S. National Museum) for lend- Etymology. This species is named in honor of my daugh- ing other types and to Gabriel Mejdalani and two anonymous ter, Claire E. Dietrich. referees for constructive criticism of the manuscript. This work was funded in part by NSF grants DEB 0089671 and DEB-0529679. Tahurella heatherae sp. nov. Figs 57, 85-86, 333-336, 355 LITERATURE CITED Description. Length of male 4.8 mm. Dorsum (Fig. 57) pale yellow with symmetrical red-orange and dark brown mark- AHMED, M. 1983. Biotaxonomy of typhlocybine leafhoppers of ings; head anterior margin with narrow dark brown band be- Pakistan, p. 179-183. In: W.J. KNIGHT; N.C. PANT; T.S. ROBERTSON tween eyes, discontinuous with similar band extended across & M.R. WILSON (Eds) Proceedings of the 1st International thoracic pleuron; pronotum with anterior margin red-orange, Workshop on Biotaxonomy, Classification and Biology posterior margin with convex red-orange marking medially; of Leafhoppers and Planthoppers (Auchenorrhyncha) of mesonotum orange anteriorly; scutellum orange at apex. Forew- Economic Importance. London, Commonwealth Institute ing with series of 7 oblique brown lines distributed along cos- of Entomology. tal margin; basal third of corium and clavus hyaline with CHRISTENSEN, J.R. 1942. Algunos cicadellidos de la Argentina y narrow red-orange transcommisural arcuate band; distal 2/3 Bolivia. Revista de la Sociedad Entomológica Argentina of forewing smoky hyaline with paler areas in apical cells; veins 11: 336-339. orange; apical lobe well developed (Fig. 85). Thoracic venter CHRISTIAN, P.G. 1953. A revision of the North American species pale yellow; abdomen pale yellow ventrally except sternite VIII of Typhlocyba and its allies (Homoptera, Cicadellidae). and valve brown. University of Kansas Science Bulletin 35: 1103-1277. Male 2S apodemes (Fig. 355) parallel sided, slightly di- DIETRICH, C.H. 2004. Phylogeny of the leafhopper subfamily vergent, extended to midlength of sternite IV. Pygofer with Evacanthinae with a review of Neotropical species and dorsal emargination parabolic (Fig. 334); tergite ca. 1/3 length Remarks on related groups (Hemiptera: Membracoidea: of distal lobe; dorsal sclerotized area of lobe gradually tapered, Cicadellidae). Systematic Entomlogy 29: 455-487. doi: apex sharply hooked ventrad and extended slightly beyond 10.1111/j.0307-6970.2004.00250.x margin; ventral appendage nearly straight in lateral and dor- DIETRICH, C.H. 2005. Keys to the families of Cicadomorpha and sal view (Figs 333 and 334). Anal hook (Fig. 333) short, apex subfamilies and tribes of Cicadellidae (Hemiptera: truncate. Subgenital plate (Fig. 333) with 3-4 macrosetae near Auchenorrhyncha). Florida Entomologist 88: 502-517. doi: midlength and several shorter stout setae preapically. Style 10.1653/0015-4040(2005)88[502:KTTFOC]2.0.CO;2 apophysis (Figs 335 and 336) nearly straight and parallel sided DIETRICH, C.H. & D.A. DMITRIEV. 2006. Review of the New World through most of length, apex curved ventolaterad, truncate. genera of the leafhopper tribe Erythroneurini (Hemiptera: Connective (Fig. 336) Y-shaped with stem as long as arms. Cicadellidae: Typhlocybinae). Bulletin of the Illinois Na- Aedeagus (Figs 335 and 336) with preatrium distinctly shorter tural History Survey 37: 119-190. than shaft, slightly broadened basally in ventral view; shaft DIETRICH, C.H. & D.A. DMITRIEV. 2008. Review of the New World slender, compressed distally, constricted preapically in lateral Erythroneurini (Hemiptera: Cicadellidae: Typhlocybinae). view, slightly sinuate, without spines or processes, apex ob- II. Genus Zyginama. Bulletin of the Illinois Natural History liquely rounded in lateral view. Survey 38: 129-175. Material examined. Holotype male, PERU: Pasco, DIETRICH, C.H. & A.M. WALLNER. 2002. Diversity and taxonomic Yanachaga-Chemillén N.P., Puesto de Control Huampal composition of Cicadellidae in the Amazonian rainforest 10°11’9"S 75°24’27"W, 2300m, 6 October 2002, D. Takiya canopy (Hemiptera, Cicadomorpha, Membracoidea), p. 18. [USML]. In: H. HOCH; M. ASCHE; C. HÖMBERG & P. KESSLING (Eds.). 11th Etymology. The species is named in honor of my cousin, International Auchenorrhyncha Congress, 5-9 August Heather B. Denne. 2002. Berlin, Museum für Naturkunde.

ZOOLOGIA 30 (5): 519–568, October, 2013 568 C.H. Dietrich

DWORAKOWSKA, I. 1969. Revision of the Palaearctic and Oriental OMAN, P. W., W. J. KNIGHT & M. W. NIELSON. 1990. Leafhoppers species of the genus Eurhadina Hpt. (Homoptera, Cicadellidae, (Cicadellidae): a bibliography, generic check-list and Typhlocybinae). Annales Zoologici 27: 67-88. index to the world literature 1956-1985. Wallingford, CAB DWORAKOWSKA, I. 1979. On the leafhopper tribe Zyginellini International Institute of Entomology, 368p. (Homoptera: Auchenorrhyncha, Cicadellidae, Typhlocybinae). RUPPEL, R.F. 1987. A summary of the tribes proposed in Typhlocybinae Revue de Zoologie Africaine 93: 288-331. (Hemiptera, Cicadellidae). Michigan Academician 19: 29-35. LINNAVUORI, R. 1954. Contributions to the neotropical leafhopper YOUNG, D.A. 1952. A reclassification of Western Hemisphere fauna of the family Cicadellidae II. A revision of some of Typhlocybinae (Homoptera, Cicadellidae). University of Stål’s and Osborn’s neotropical leafhopper species. Annales Kansas Science Bulletin 35: 1-217. Entomologica Fennica 20: 124-145. ZAHNISER, J.N. & C.H. Dietrich. 2010. Phylogeny of the leafhopper LINNAVUORI, R. 1959. Revision of the Neotropical Deltocephalinae subfamily Deltocephalinae (Hemiptera: Cicadellidae) based and some related subfamilies (Homoptera). Suomalaisen on molecular and morphological data with a revised family- Elain-ja Kasvitieteellisen Seuran Vanamon Elaintieteellisia group classification. Systematic Entomology 35: 489-511. Julkaisuja 20: 1-370. doi: 10.1111/j.1365-3113.2010.00522.x

Submitted: 04.IV.2013; Accepted: 07.VII.2013. Editorial responsibility: Gabriel L.F. Mejdalani

ZOOLOGIA 30 (5): 519–568, October, 2013