Subtropical and tropical zones of the New World have a rich but still largely undescribed leafhopper fauna (Dietrich & Wallner 2002;Dietrich & Rakitov 2002). Deltocephalinae is the most diverse subfamily of Cicadellidae, including >1,070 genera in 39 tribes, of which Athysanini Van Duzee is the largest. However, this tribe is poorly defined and not monophyletic. A recent comprehensive phylogenetic analysis of the subfamily did not include a sufficient number of athysanine genera to resolve relationships confidently or support dividing the tribe into smaller monophyletic groups of genera (Zahniser & Dietrich 2013). Many endemic Neotropical genera of Deltocephalinae are enigmatic due to bizarre morphological features that obscure their affinities and often give them a superficial resemblance to other leafhopper subfamilies (Linnavuori 1959;Linnavuori & DeLong 1976, 1978;Dietrich & Rakitov 2002). Nevertheless, many of the unusual forest-dwelling Neotropical Athysanini included in the phylogenetic analysis of Zahniser & Dietrich (2013) grouped together in a clade with the endemic New World tribes Pendarini and Scaphytopiini. A more recent analysis including more taxa and gene regions (Lin Lu, Northwest A&F University, pers. comm.) also placed the endemic Neotropical tribe Bahitini in this large New World clade. This suggests that, despite the high morphological diversity found among the Neotropical Athysanini, they are closely related to other Deltocephalinae inhabiting the forests of Central and South America.
Among the unusual Neotropical Athysanini are the genera Brazosa Oman and Zabroza Oman, which have an arched crown/pronotum and bright orange dorsal spots resembling those found in many Idiocerini (Eurymelinae). These two genera appear to belong to a larger group of Neotropical Athysanini that includes species with bold dorsal markings including paired black spots and orange lines or patches, a relatively short, broad crown rounded to the face, unmodified wing venation, and relatively simple male genitalia (pygofer and subgenital plates with reduced setation, segment 10th without processes and usually unsclerotized, aedeagus U-shaped and usually without large processes, style without or with week preapical lobe and apex digitiform).
In the present work, study of samples from the Amazon Basin, collected either at lights or by fogging rainforest canopies, revealed the presence of several species belonging to three new genera from Ecuador, Peru, Brazil, and Bolivia. The new taxa are notable in showing affinities to previously described Neotropical genera, particularly in overall structure and dorsal color pattern. They are described and illustrated below and a table comparing the morphology of the new genera to that of some similar Neotropical genera is provided.
General descriptive morphological terminology herein follows Dietrich (2005). Wing venation follows Anufriev & Emeljanov (1988) and leg chaetotaxy follows Rakitov (1998). Abdomens of males and females studied were cleared in hot 10% KOH solution, rinsed in distilled water and stored in glycerin. Holotypes and paratypes on loan were relaxed in a humid chamber then dissected and remounted. Images were taken using a Big Kahuna camera coupled to stereo microscope (http://www.duninc.com/bk-lab-system.html). Zerene Stacker software was used to combine images captured at different focal planes and Adobe Illustrator and Photoshop were used to process final images. Body length was measured using an electronic vernier. In quoted label text, different labels are separated by a reversed virgule (\) starting with the top label; bracket ([ ]) for additional information that is not written on the labels. Body color is described based on pinned dry specimens.
The material studied here is deposited at the following institutions: 1).
External morphology. Robust small to medium-sized leafhoppers (3.9-5.2 mm), overall dorsal coloration grey, venter variously marked with black. Crown with transverse orange submarginal band between eyes and median apical black spot; pronotum with four oblong orange spots; mesonotum, scutellum and claval veins also variably marked with orange, basolateral triangles of mesonotum black; forewing veins and cells without distinct pigmentation except in clavus. Face with midline of frontoclypeus yellowish with various amounts of dark pigmentation laterally, ocellocular region and area bordering ventrolateral part of frontoclypeus black. Head subequal to or wider than pronotum. Crown short, arched above eyes, anterior margin bluntly angulate in dorsal view, transition to face forming blunt obtuse angle in lateral view, crown width between eyes approximately twice eye width. Face mostly yellowish with darkish marks (except in appendiculata, mostly dark with a central orange line). Ocellocular area parallel sided; lateral frontal sutures complete, extended to ocelli. Ocelli on anterior margin of head, near eyes. Frontoclypeus longer than wide, slightly convex; erect fine seta present laterad of frontal suture. Antennal ledge represented by indistinct carina, antennal base situated slightly below midheight of eye, antenna length 1.2x width of head. Anteclypeus widened apically. Lorum narrowly separated from lower margin of face, wider than or subequal to anteclypeus in width near base. Gena slightly incised below eye. Rostrum short, tapered, slightly surpassing front trochanters.
Pronotum strongly convex, anterior margin parabolic, lateral margin carinate, shorter than eye width. Scutellum protuberant in lateral view. Forewing macropterous, translucent, apex rounded, appendix restricted to anal margin, without extra crossveins, with three anteapical cells, outer apical cell large with apex delimited by crossvein, inner anteapical cell open (without m-cu2 crossvein), apical cells subequal in width, Pcu connected to claval suture by crossvein; Pcu and A1 free basally but usually confluent through most of length distally, rarely separate with crossvein present. Hind wing venation fully developed, without pigmentation, RP-MA and MP-CuA separated by crossvein. Front femur with long AM1 at mid-height, IC setae long and thin (10-25), row AV with short stout setae (8-25), AV1 poorly differentiated from IC setae. Front tibia dorsal macrosetal formula 1+4 (AD+PD), PV without or with macrosetae. Mesotrochanter with one stout setae. Mesofemur row AV with short stout setae. Hind femur macrosetal formula 2+2+1, without extra setae basad of usual set. Metatarsomere I as long as II and III combined, not expanded apically, plantar setae simple, pecten with 4 platellae and inner apical seta platelliform.
Male genitalia. Apodemes of sternite II inconspicuous. Basolaterad pygofer apodemes absent. Pygofer in dorsal view with moderately long dorsal sclerotized section, lobes in lateral view more or less triangular, longest along dorsal margin, well-sclerotized, without processes, short ventral process sometimes present, macrosetae sparse, present near posterior margin, basolateral cleft present. Anal tube membranous. Valve and subgenital plates free from each other, articulated with pygofer. Valve relatively short, posterior margin weakly produced medially. Subgenital plate triangular, apex narrowly rounded, macrosetae uniseriate sublaterally or somewhat irregularly arranged near margin. Connective somewhat H-shaped, with arms subparallel and stem emarginate. Style broadly bilobed basally; median lobe short, not extended dorsad, preapical lobe short and rounded; apophysis somewhat elongated, extended posteromesad then strongly curved laterad with apex pointed or rounded, with few fine lateral setae, without obvious microsclupture or teeth. Aedeagus with atrium slightly broader than shaft; preatrium absent or very short; shaft curved dorsad, usually with pair of lateral flanges with or without teeth, rarely with distal processes, apex of aedeagus blunt, bifid or pointed; gonoduct not sclerotized, narrower than base of shaft width; gonopore subapical on dorsal surface; phragma weakly sclerotized or membranous, with or without conspicuous setae.
Female genitalia. Pygofer conical with numerous macrosetae concentrated on posterior portion. Ovipositor not protruding far beyond pygofer apex. First valvula, ramus convex, dorsal sculpturing imbricate with overlapping scales, reaching dorsal margin, more elongated medially than apically, ventral sculpturing restricted to apical portion. Second valvula abruptly broadened medially with medial dorsal tooth, distal teeth small restricted to apical portion and serrated. Third valvula with one row of distinct setae. Sternite VII truncate. Eighth sternite membranous.
Distribution. Brazil, Bolivia and Peru Male genitalia. Pygofer much longer than tall, macrosetae mixed with microsetae near posterior margin, ventrolateral cleft bordered with fine setae. Anal tube as long as wide, broad apically, same length as pygofer. Valve short, flat, weakly produced posteromesad. Subgenital plate relatively short, with submarginal row of 4 somewhat fine macrosetae in distal half. Connective, stem shorter and narrower than arms. Style, preapical lobe very short and broad, apophysis relatively long and strongly curved, forming less than 90º angle laterally. Aedeagus curved dorsocaudad forming less than half circle, shaft robust basally in lateral view; distal flange without teeth, projected anterad, in posterior view rounded laterally and not extended to apex.
Female genitalia. Unknown. Type material. Holotype ♂ (MZSP)-BRASIL, Goiás: Campinaçu, Serra da Mesa Survey, 13°52.1´S 48°23. 2´W, 19-23.ii. 1996, Malaise trap. A. Sharkov and F. Ejchel Colls. Distribution. Brazil Measurements. Body length, ♂ 4.6 mm, ♀ unknown.
Etymology. The species name, a feminine noun, refers to the apical-shape of the aedeagus, using a Spanish prefix "espatula" meaning "spatula".
Brazosa negra Pinedo-Escatel & Dietrich, sp. nov. (Figs. 1 D and L,2 D,6) urn:lsid:zoobank.org:act:A3C6D75C-770E-46C0-9708-6DE092A17A43 Description. External morphology. Frontoclypeus with center yellowish and lateral suture with a broad black band. Anteclypeus yellowish. Lorum black. Gena black next to frontoclypeal suture emerging below antennal pit running to lorum, yellowish distally. Anterior margin of pronotum ivory and posterior half dark-greenish. Forewings light brownish. Ventral sclerites yellowish and black. Front tibia and tarsomeres orange.
Male genitalia. Pygofer as long as tall, posterior margin rounded, macrosetae restricted to apical portion, fine setae scattered along posterior margin, broadly rounded ventrocaudally, teeth on inner margin. Anal tube more than 2x longer than wide, parallel, longer than pygofer. Valve broader than long, projected posteromesad. Subgenital plate longer than width, with uniseriate row of 9-10 somewhat stout macrosetae. Connective, stem shorter and wider than arms. Style, preapical lobe short and rounded, apophysis short and slightly curved, less than a 90º angle laterally. Aedeagus curved dorsad, shaft slender in lateral view; apex emarginate; apical flanges with teeth, projected anterad.
Female genitalia. Unknown. The species name, a feminine noun, refers to color of the lorum and gena, using the Spanish "negra" meaning black.
Remarks. This is species is similar to B. beni in coloration, but the different aedeagal apex separates these species.
Brazosa encrustada Pinedo-Escatel & Dietrich, sp. nov. (Figs. 1 E and M,7) urn:lsid:zoobank.org:act:BDCBB0B7-98DC-4099-B63C-BC5A072C0EC3 Description. External morphology. Frontoclypeus yellowish with or without two median longitudinal lines on midline. Ocellocular area with upper half black and lower yellowish. Anteclypeus with a center yellowish line bordered with black. Lorum black with a broad black band reaching ocelli. Gena yellowish with a gold line surround- Male genitalia. Pygofer as long as tall, slightly pointed, macrosetae near posterior margin with fine setae intercalated, rounded caudoventrally. Anal tube as long as wide, parallel, shorter than pygofer. Valve, short, weakly produced posteromesad. Subgenital plate long as pygofer, strongly tapered to apex, with subapical row of macrosetae slightly submarginal. Connective short, stem shorter than arms, apex truncate. Style, preapical lobe distinct and rounded, apophysis relatively long and curved. Aedeagus curved dorsocaudad, shaft slender in lateral view; apex produced, distal flange in posterior view rounded laterally with small teeth.
Female genitalia. Unknown. Type material. Holotype ♂ (MZSP)-BRASIL, Goiás: Campinaçu, Serra da Mesa Survey, 13°52.1´S 48°23. 2´W, 19-23.ii. 1996, Malaise trap. A. Sharkov and F. Ejchel Colls.; Paratypes, 1♂ (MZSP)-BRASIL, Goiás: Campinaçu, Serra da Mesa Survey, 13°52.1´S 48°23. 2´W, 19-23.ii. 1996, Malaise trap. A. Sharkov and F. Ejchel Colls; 1♂ (MZSP)-BRASIL, Goiás: Campinaçu, Serra da Mesa Survey, 13°52.1´S 48°23. 2´W, 19-23.ii. 1996, Malaise trap. A. Sharkov and F. Ejchel Colls. Distribution. Brazil Measurements. Body length, ♂ 5.1 mm, ♀ unknown Etymology. The species name, a feminine noun, refers to the variable color of the face, using a Spanish word "encrustada" meaning "imbedded".
Remarks. This species is similar externally to B. campinacu, but it differs in the coloration of the face and the structure of the male genitalia.
Brazosa mildredireanae Pinedo-Escatel, sp. nov. (Figs. 1 F and N,2 G,8) urn:lsid:zoobank.org:act:3AC21553-2F38-4180-A002-8FC27CDB141C Description. External morphology. Frontoclypeus whitish flanked by a dark band on basolateral suture. Anteclypeus and lorum whitish. Gena whitish with center yellowish. Pronotum greenish with two paired long broad bands, inner pair shorter than lateral. Forewings reddish. Prothorax blackish and abdomen white. Legs white.
Male genitalia. Pygofer longer than tall in lateral view, long and stout macrosetae in more or less a row subapically, fine setae through of posterior margin, strongly rounded ventrally. Anal tube 1.5x longer than wide, broader basally, slightly beyond pygofer. Valve flat, short, weakly produced posteromesad. Subgenital plate as long as pygofer, gradually tapering to apex, macrosetae uniseriate subapically. Connective, stem broader and longer than arms. Style, preapical lobe strongly rounded, apophysis long and curved, forming a 90º angle laterally. Aedeagus strongly curved dorsad, dorsal margin forming a distinct half circle, shaft slender in lateral view; apex emarginate medially; lateral flanges with teeth, projected anterad.
Female genitalia. Brazosa campinacu Pinedo-Escatel & Dietrich, sp. nov. (Figs. 1 G and O,2 H,9) urn:lsid:zoobank.org:act:9A079306-92E3-4D74-B363-416B2CEB2905 Description. External morphology. Frontoclypeus yellowish with two broad median longitudinal lines on midline connected to lorum and anteclypeus. Anteclypeus yellowish basally then black. Lorum black. Gena black, yellowish distally. Pronotum dark-greenish. Forewings smoky. Ventral sclerites black. Front coxa, trochanter and femur marked with black.
Male genitalia. Pygofer as long as tall, posterior margin pointed, scattered macrosetae and several fine setae on posterior margin, weakly rounded ventrally, ventrolateral cleft bordered with long and fine setae. Anal tube 1.2x longer than wide, tapered apically, shorter than pygofer. Valve short, produced posteromesad, weakly inflated in lateral view. Subgenital plate long as pygofer, gradually tapering to apex, macrosetae uniseriate subapically. Connective, stem shorter than arms, apex slightly emarginate. Style, preapical lobe distinct and rounded, apophysis long and strongly curved. Aedeagus curved dorsad, shaft slender in lateral view; apex flange-like, emarginate medially, membranous subapically in posterior view.
Female genitalia. Unknown Type material. Holotype ♂ (MZSP)-BRASIL, Goiás: Campinaçu, Serra da Mesa Survey, 13°52.1´S 48°23. 2´W, 19-23.ii. 1996, Malaise trap. A. Sharkov and F. Ejchel Colls. Distribution. Brazil Measurements. Body length, ♂ 5.1 mm, ♀ unknown Etymology. The species name refers to the type locality, Campinaçu in Serra da Mesa, Brazil. The species name is a noun in apposition.
Remarks. This species is similar in coloration to B. encrustada, but differs in the apex of aedeagus. Remarks. This species is superficially similar to B. negra, but it can be easily distinguished by the apex of the aedeagus.
Spaltumtettix Pinedo-Escatel & Dietrich, gen. nov. 12 A and C,(14)(15)(16)(17)(18)31 B) urn:lsid:zoobank.org:act:6AC34C29-B662-4FDD-AEBC-36FC8C826B64 Type species. Brazosa caesarea Linnavouri & Heller, 1961 Description. Diagnosis. Dorsal coloration yellowish-ochraceous and forewings with orange bands. Crown short, arched above eyes. Pronotum convex with strong lateral carina. Forewings without extra crossveins; anal veins fused; Cs-Pcu with 1 crossvein; Pcu-A1 crossveins absent. Front tibia formula 1+5; AM1 position at middle. Pygofer without or posterad processes; valve fused with subgenital plate: aedeagus without or with apical processes; first valvula dorsal sculpturing strigate (Table 1).
External morphology. Moderately robust and large leafhoppers (8.8-10.2 mm), general coloration yellowish-ochraceous in female to greenish in male; live specimens green with orange markings. Male crown, pronotum, mesaepisternum, and scutellum with orange markings (faded in some specimens); crown with pair of submedial orange spots, pronotum with two pairs of orange spots, forewing translucent with indistinct pigmentation except vein CuA and fused claval veins bright orange; venter stramineous. Head wider than pronotum. Crown short, relatively flat, anterior and posterior margins parallel, interocular width less than twice eye width, transition of crown to face bluntly angled, texture shagreen. Ocellocular area parallel; lateral frontal sutures subparallel, extended to ocelli.
Ocelli on anterior margin of head, separated from eyes by one ocellar diameter. Antennal ledge obsolete, antennal base close to lower corner of eye, antenna length 1.1x width of head. Frontoclypeus moderately narrow, not tumid in lateral view, parallel sided, fine erect seta present next to frontoclypeal suture. Anteclypeus widened apically with fine microsetae, apex slightly surpassing natural curve of gena. Lorum wider than anteclypeus near base, extended to lower margin of face. Gena slightly incised laterad, with fine microsetae basad of lorum and anteclypeus. Rostrum short, tapered, not extended beyond front trochanters.
Pronotum convex, anterior margin evenly rounded, posterior margin only slightly concave, lateral margin carinate, shorter than half eye width. Scutellum not protuberant. Forewing macropterous, translucent, appendix restricted to anal margin, apex rounded, without extra crossveins, three anteapical cells present, outer apical cell moderately large with apex delimited by crossvein, apical cells subequal in width, inner anteapical cell open (without m-cu2 crossvein), vein CuA very close to claval suture (brachial cell unusually narrow); clavus with crossvein connecting Pcu to claval suture, Pcu and A1 completely confluent throughout length, combined vein distinctly sinuate. Hind wing venation fully developed, without pigmentation, RP-MA and MP-CuA each separated by crossvein. Front femur with long AM1 at mid-height, IC setae long and thin (15-21), row AV with short stout setae (18-26), AV1 poorly differentiated from IC row. Front tibia dorsal macrosetal formula 1+5 (AD+PD), PV with multiple macrosetae. Mesotrochanter with more than one stout seta apically. Mesofemur row AV with short stout setae. Hind femur macrosetal formula 2+2+1, without extra setae basad of usual set. Metatarsomere I longer than metatarsomeres II and III combined, not expanded apically, plantar setae simple, pecten with 3 platellae and inner apical setae tapered.
Male genitalia. Apodemes of sternite II inconspicuous. Basolateral pygofer apodemes absent. Pygofer moderately long, triangular, longest along dorsal margin with short, well-sclerotized band dorsally at base, broadly incised below apex or not, without macrosetae, only fine setae concentrated on incised posterior margin, basolateral cleft present, with well sclerotized process or thickened area on lobe posterodorsally. Anal tube very long and narrow, fully sclerotized dorsally. Valve fused to subgenital plates, articulated with pygofer, with strong basolateral constriction delimiting point of fusion with plates. Subgenital plates very long, slender, fused to each other basally, free distally, extended beyond pygofer, slightly compressed, rounded apically, with several stout setae in single sublateral row preapically. Connective Y-shaped, anterior arms only slightly separated anteriorly, stem long, extended dorsad. Style broadly bilobed basally, median basal lobe strongly sinuate and elongated dorsad, preapical lobe reduced, apophysis elongate, curved mesad then hooked laterad at apex, apex pointed or rounded. Aedeagus long, atrium well developed but not substantially wider than shaft, preatrium poorly developed; shaft tubular, curved dorsad, with or without distal processes, gonoduct not sclerotized, gonophore apical.
Female genitalia. Pygofer conical, posterior margin rounded, long macrosetae distributed near posterior margin, stout setae concentrated near midlength. Ovipositor slightly protruding beyond pygofer apex. First valvula, ramus convex, dorsal sculpturing pattern strigate, reaching dorsal margin, ventroapical sculpturing restricted to apical portion. Second valvula, slender and evenly curved throughout length, with numerous subquadrate dorsal teeth along distal third, teeth serrated. Third valvula, with basal half narrow and apical half abruptly expanded, apex broad with few small setae on apical portion. Sternite VII greatly enlarged, longer than wide, compressed and directed downwards, tip with very prominent ventral projection preapically in lateral view, apex slightly notched. Eighth sternite well-sclerotized, attached at midlength of sternite VII.
Habitat. Amazonian rainforest Etymology. The genus name refers to its extravagant genitalia, using the combination of Latin words "spatha" (spatula) and "multum" (long), ending with "tettix", a common suffix of leafhopper names. Gender: masculine.
Remarks. This genus is similar to Brazosa, which it resembles externally in having a very short crown and prominent orange dorsal markings but is larger and more elongate and differs in the male genitalia including the large and partly exposed aedeagus. Etymology. The specific epithet refers to the external color pattern, is a Latinization of the Spanish adjective "coloreada", meaning "colored".
Spaltumtettix caesareus (Linnavouri & Heller), comb. nov. (Figs. 2 K,11 B,12 C,(17)(18) Brazosa caesarea Linnavouri & Heller, 1961: 11 Redescription. External morphology. Male external morphology as in coloradus. Pronotum greenish marked with four large orange spots. Ocellocular area only in male orange. Female, crown and scutellum ochraceous, pronotum almost fully ochraceous with minute pair of orange stripes on each laterobasal margin (rudimentary coloration matching that of male present but faded). Thorax and abdomen yellowish ventrad.
Male genitalia. Pygofer well sclerotized, without weakly sclerotized dorsoapical lobe, posterior margin with more strongly sclerotized area posteroventrally near apex, process absent. Subgenital plates more than twice pygofer length. Style apophysis greatly elongate and broadly curved following curve of subgenital plate. Aedeagus strongly and evenly curved dorsad, with pair of short lateral spines flanking gonopore apically.
Female genitalia. Pygofer conical with long setae posterad. First valvula dorsal sculpturing pattern strigate. Second valvula serrate with numerous tooth, ducts connected between each other and directed to apical portion. Third valvula punctate. Sternite VII long, broad, posterior margin slightly notched.
Distribution. Peru and Ecuador nov. rec. 1).
External morphology. Medium-sized (5.7-7.0 mm) moderately robust leafhoppers. Overall coloration yellowish-orange with bright orange markings; crown mostly bright orange, face with pair of black spots mesad of ocelli connected by more or less distinct arcuate black line that extends slightly onto crown; pronotum with pair of oblique lateral orange bands; mesonotum and scutellum mostly orange with basolateral triangles and median triangular marking black. Head wider than pronotum. Crown convex, declivous, relatively short, anterior margin broadly rounded, more or less parallel to posterior margin, width between eyes more than twice eye width, transition of crown to face bluntly angulate, shagreen. Ocellocular area parallel-sided and short, lateral frontal sutures extended to ocelli. Ocelli on anterior margin of head, separated from eyes by one ocellar diameter. Frontoclypeus, narrow, not tumid, parallel-sided through most of length, erect fine seta present laterad of frontal suture. Antennal ledges weakly developed, without carina. Anteclypeus constricted medially, widening apically, not tumid, apex slightly surpassing natural curve of gena. Lorum subequal to anteclypeus in width near base, extended nearly to lower margin of face. Gena slightly incised below eye. Rostrum short, narrow, extended slightly beyond front trochanters.
Pronotum with anterior margin parabolic; strongly convex and declivous anteriorly in lateral view; lateral margin carinate, shorter than half eye width; posterior margin slightly concave. Scutellum not protuberant. Forewing macropterous, transparent, appendix restricted to anal margin, apex rounded, without extra crossveins, with three anteapical cells, outer apical cell large with apex delimited by crossvein, apical cells subequal in width, inner anteapical cell open (without m-cu2 crossvein). Clavus without crossvein connecting Pcu to claval suture, Pcu and A1 free throughout length, crossvein absent. Hindwing venation complete, RP-MA and MP-CuA separated by crossvein. Front femur with short AM1 on ventral margin, IC setae long and thin (12-20), row AV without setae, AV1 poorly differentiated from IC row. Front tibia with dorsal macrosetal formula 1+4 (AD+PD), PV without setae. Mesotrochanter without setae. Mesofemur row AV with short, stout setae. Hind femur macrosetal formula 2+2+1, without extra setae basad of usual set. Metatarsomere I as long as II and III combined, not expanded apically, plantar setae simple, pecten with 3 platellae and inner apical setae tapered.
Male genitalia. Apodemes of sternite II inconspicuous. Basolateral pygofer apodemes absent. Pygofer elongate, incised nearly to base dorsally, posterior margin of lobe rounded, macrosetae concentrated near posterior margin, basolateral cleft longer than half of pygofer length, basolateral section ventrad of cleft extended anteroventrad into digitiform process. Anal tube short, narrow, sclerotized dorsally. Valve and subgenital plates free, articulated to pygofer. Valve with posterior margin rounded. Subgenital plate strongly constricted at basal third and gradually tapered distally, depressed, apex rounded, macrosetae in distal two thirds, uniseriate and well separated from margin; fine setae sparsely distributed laterally and dorsally. Connective elongate, Y-shaped, 3x style length; stem much longer than arms, emarginate apically. Style broadly bilobed basally, median lobe short and broad, preapical lobe absent, apophysis relatively short and strongly curved mesad, with group of fine setae dorsally near apex; apex abruptly narrowed and hooked laterad to sharply pointed tip. Aedeagus with atrium moderately enlarged relative to shaft, dorsum of atrium poorly developed, preatrium absent; shaft slender, tapered, curved dorsad, with or with-out pair of weakly sclerotized basal processes arising dorsally at base of shaft and extended distad, apex of shaft pointed, gonoduct not sclerotized, gonopore short, preapical on ventral surface.
Female genitalia. Pygofer long, produced posterad, posterior margin rounded, with macrosetae grouped near middle of posteroventral margin. Ovipositor extended far beyond pygofer apex. First valvula ramus not strongly convex, dorsal sculpturing granulose, narrowly separated from dorsal margin through most of length, ventroapical sculpturing restricted to apical portion. Second valvula gradually broadened distad, without dorsal medial tooth, distal teeth absent. Third valvula with basal half narrow and apical half abruptly expanded, apex with setae on apical portion. Sternite VII much longer than wide, narrowing towards posterior margin, apex pointed. Eighth sternite long, well-sclerotized.
Distribution. Peru, Ecuador, and Brazil.
Napo bellus Zanol, 2006 Measurements. Body length, ♂ 6.5 mm, ♀ 6.9 mm Remarks. Male specimens examined from MEPN, INHS and OSUC vary in the length and orientation of the aedeagal processes (Fig. 24 A-D), also the style and apex of the apophysis differ notably in size (Fig. 24 E-G).
Napo bellus Zanol, 2006: 65 Remarks. Zanol (2006) noted that N. bellus is easily separated from N. brazosellus due to the absence of processes on the aedeagus. Aside from this reported difference in the male genitalia, this taxon appears to be identical to the type species of the genus. Zanol's specimens were not available for study but need to be re-examined because the basal aedeagal processes of N. brazosellus are weakly sclerotized and often not easily visible in the specimens examined. Thus, it is possible that the Brazilian specimens of N. bellus examined by Zanol 1). External morphology. Medium-sized (5.9-7.1 mm) moderately robust leafhoppers. Overall coloration yellowish-ochraceous with bright orange dorsal markings; crown mostly orange (faded in some individuals); face with pair of black spots ventromesad of ocelli, with or without dark marking between spots; pronotum uniformly greenish brown; mesonotum and scutellum orange with basolateral triangles and median triangular area black. Head wider than pronotum. Crown convex, declivous, relatively short, anterior margin broadly rounded, more or less parallel to posterior margin, width between eyes more 1.7 times eye width, transition of crown to face bluntly angulate, shagreen. Ocellocular area parallel-sided, short, lateral frontal reaching ocelli. Ocelli on anterior margin of head, separated from eyes by 1.2 times ocellar diameter. Frontoclypeus, narrow and not tumid, parallel-sided through most of length, erect fine seta present laterad of frontal suture. Antennal ledges weakly developed, without carina. Anteclypeus narrowed medially, widening apically, not tumid, apex slightly surpassing natural curve of gena. Lorum subequal to anteclypeus in width near base, extended nearly to lower margin of face. Gena weakly incised below eye.
Pronotum with anterior margin parabolic; convex and declivous anteriorly in lateral view; lateral margin carinate, shorter than half eye width; posterior margin slightly concave. Scutellum not protuberant. Forewing macropterous, transparent, appendix restricted to anal margin, apex rounded, without extra crossveins, with three anteapical cells, outer apical cell large with apex delimited by crossvein, apical cells subequal in width, inner anteapical cell open (without m-cu2 crossvein). Clavus without crossvein between Pcu and claval suture, Pcu and A1 free throughout length, crossvein absent. Hindwing venation complete, RP-MA and MP-CuA separated by crossvein. Front femur with short AM1 on ventral margin, IC setae long and thin (14-17), row AV without stout setae, AV1 poorly differentiated from IC row. Front tibia with dorsal macrosetal formula 1+4 (AD+PD), PV without setae. Mesotrochanter with more than one stout setae. Mesofemur row AV with short and stout setae. Hind femur macrosetal formula 2+2+1, without extra setae basad of usual set. Metatarsomere I as long as II and III combined, not expanded apically, plantar setae simple, pecten with 3 platellae and inner apical setae tapered.
Male genitalia. Apodemes of sternite II inconspicuous. Basolateral pygofer apodemes absent. Pygofer moderately longer than tall, incised to near base dorsally, with processes well-sclerotized dorsally, posterior margin projected, lobe with dorsal preapical process extended posterodorsad, macrosetae reduced and concentrated nearly to processes and posterior margin, basolateral cleft shorter than half of pygofer length. Anal tube long as half-length of pygofer, broad, fully sclerotized dorsally. Valve and subgenital plates free, articulated to pygofer. Valve wider than long, posterior margin strongly rounded, articulated to pygofer. Subgenital plate flat, slender throughout its length, macrosetae uniseriate and well or not separated from margin; fine setae distributed basally. Connective short, Y-shaped, shorter than style, arms well separated from each other, stem short and truncate. Style bilobed basally, median lobe inconspicuous or short; apophysis short, straight or strongly curved mesad and hooked with or without setae, apex blunt or pointed. Aedeagus with atrium not enlarged, atrium dorsally longer than half length of shaft, preatrium absent; shaft broad at base then slender, tapering to apex, strongly curved dorsad, without basal processes; apex of shaft pointed with or without processes, gonoduct not sclerotized, gonopore broad as shaft width subapically on ventral surface.
Female genitalia. Pygofer long, posterior margin squarish, macrosetae distributed near posteroventral margin. Ovipositor extended far beyond pygofer apex. First valvula, ramus slightly curved, dorsal and ventroapical sculpturing granulose, reaching dorsal margin, restricted to apical portion. Second valvula gradually broadened toward apex, without dorsal medial tooth, inconspicuous teeth near apex and distant teeth absent. Third valvula, basal half narrow and apical half abruptly expanded, apex with several setae on apical portion. Sternite VII much longer than wide, apex produced and rounded, medially ornate and carinate. Eighth sternite weakly sclerotized.
Distribution. Ecuador and Peru.
Etymology. The genus name is a feminine noun, a combination of prefix "pseudo" meaning "false" with "Napo", due to the morphological similarity to the latter genus.
Remarks. Pseudonapo is very similar to Napo Linnavuori & DeLong externally based on color pattern and overall structure. The male genital capsule is distinctly different from that of Napo in having the pygofer incised near base dorsally with dorsal processes and the aedeagus strongly curved dorsad with or witout apical processes. Redescription. Diagnosis. Overall coloration yellowish. Crown produced, arched above eyes. Pronotum convex with strong lateral carina. Forewings without extra crossveins; anal veins free; Cs-Pcu without or with 1 crossvein; Pcu-A1 without crossvein. Front tibia formula 3+4; AM1 position at middle. Pygofer with ventral processes; valve free; aedeagus without processes; first valvula dorsal sculpturing granulose or strigate (Table 1).
External morphology. Robust, medium-sized (5.9-7.4 mm) leafhoppers, overall coloration yellowish with symmetrical black marks on crown, face and pronotum; face with pair of oblong black spots mesad of ocelli; crown with one or two pairs of spots; pronotum with pair of sublateral spots; forewing smoky hyaline without distinct markings. Head wider than pronotum. Crown short, strongly arched above eyes and anterior margin of pronotum, slightly produced medially, posterior and anterior margins subparallel, transition to face bluntly angled, width between eyes than twice eye width. Face with ocellocular area parallel sided. Ocelli on anterior margin of head, separated from eyes by one ocellar diameter. Frontoclypeus more or less evenly tapered from base to apex, not tumid, fine seta present just laterad of frontal suture. Antennal ledges indistinct, without carina; antennal base close to lower corner of eye, antenna length 1.1x width of head. Anteclypeus strongly widened apically, extended slightly beyond normal curve of gena. Lorum wider than anteclypeus near base, extended to ventral margin of face. Gena slightly incised below eye, rostrum short, tapered, extended slightly beyond front trochanters.
Pronotum moderately convex, anterior margin parabolic, posterior margin slightly concave, lateral margin shorter than half width of eye, carinate. Scutellum not protuberant. Forewing macropterous, translucent, appendix restricted to anal margin, apex rounded, without extra crossveins, with three anteapical cells, outer apical cell large with apex delimited by crossvein, apical cells subequal in width, inner anteapical cell open (without m-cu2 crossvein), Claval veins separate throughout length, with or without crossvein connecting Pcu to claval suture, Pcu-A1 crossvein absent. Hindwing venation fully developed, RP-MA and MP-CuA separated by crossvein. Front femur with short AM1 at mid-height, IC setae long and thin (7-9), row AV with short stout setae (17-20), AV1 large and distinct from IC row. Front tibia dorsal macrosetal formula 3+4 (AD+PD), PV with multiple macrosetae. Mesotrochanter with one stout seta. Mesofemur row AV with numerous stout setae and 1 longer apical seta. Hind femur macrosetal formula 2+2+1, without extra setae basad of usual set. Metatarsomere I longer than metatarsomeres II and III combined, not expanded apically, plantar setae simple, pecten with 4 platellae and inner apical seta platelliform.
Male genitalia. Apodemes of sternite I inconspicuous, apodemes of sternite II present but not particularly large. Basolateral pygofer apodemes absent. Pygofer relatively short and broad, squarish in lateral view, with ventral process, with few scattered macrosetae preapically, basolaterad cleft present. Anal tube membranous or with thin dorsal sclerotized band distally. Valve and subgenital plates free from each other, articulated with pygofer. Valve very short with only slight median posterior extension. Subgenital plate large, triangluar, lateral margin slightly convex, apex rounded, macroseate irregularly arranged near lateral margin. Conective Y-shaped, arms short, stem very long. Style broadly bilobed basally, median lobe not extended dorsad, preapical lobe short and rounded, apophysis short and smoothly curved with apex pointed or rounded. Aedeagus with atrium only slightly wider than shaft base, shaft curved dorsad, without processes, without processes, symmetrical, apex in posterior view with distinct medial notch, gonoduct not sclerotized, gonopore on dorsal surface near midlength of shaft, phragma membranous, with or without conspicuous setae.
Female genitalia. Pygofer short, conical with numerous macrosetae concentrated on posterior portion. Ovipositor not protruding far beyond pygofer apex. First valvula, ramus convex or straight, dorsal sculpturing pattern imbricate, reaching dorsal margin, dorsal sculpturing more elongated near middle than near apex, ventroapical sculpturing restricted to apical portion. Second valvula broadened medially without medial dorsal tooth, teeth small and serrated with inconspicuous tooth, ducts globose. Third valvula, wide, stout setae restricted to apex or a row of stout setae along length. Sternite VII or emarginate medially and carinate ventrad. Eighth sternite fully or weakly sclerotized.
Distribution. Bolivia, Ecuador, Brazil, Paraguay, Argentina, Mexico and USA
Zabrosa amazonensis (Osborn, 1923) 1). External morphology. Robust, medium-sized (5.0-5.7 mm) leafhoppers, overall coloration stramineous to brownish with symmetrical black spots and lines and diffuse brown markings; face with pair of round black spots above antennal pits, crown with paired black maculae, mesonotum with basolateral triangles black; forewing translucent brown with unpigmented areas in some cells, veins pale bordered with brown. Head wider than pronotum. Crown very short, arched above eyes and anterior part of pronotum, anterior margin rounded and subparallel to posterior margin, transition to face rounded, width between eyes less than twice eye width. Ocellocular area at tip narrower than base. Ocelli slightly below anterior margin of head and not visible in dorsal view, separated from eyes by twice ocellar diameter. Frontoclypeus with lateral margins weakly convex, surface not tumid, erect fine seta close to lateral margin. Antennal ledges not carinate, antennal base close to lower corner of eye, antenna length 1.0x width of head. Anteclypeus strongly widening apically and extended slightly beyond normal curve of face margin. Lorum subequal to anteclypeus in width near base, extended to ventral margin of face. Gena slightly incised below eye. Rostrum short, tapered, extended slightly beyond front trochanters.
Pronotum strongly convex, punctate, anterior margin parabolic, posterior margin slightly concave; lateral margin carinate, shorter than half eye width. Scutellum not protuberant. Forewing macropterous, appendix restricted to anal margin, apex rounded, with one extra costal crossvein, with three anteapical cells, outer anteapical cell short with apex delimited by crossvein, apical cells subequal in width, inner anteapical cell open (without m-cu2 crossvein), Claval veins separate throughout length, Pcu connected to claval suture by crossvein, Pcu-A1 crossvein present. Hindwing scheme fully developed, with smoky pigmentation, RP-MA and MP-CuA separated by crossvein. Front femur with long AM1 on ventral margin, IC setae very long and thin (10-14), row AV with short stout setae (8-15), AV1 poorly differentiated from IC row. Front tibia dorsal macrosetal formula 1+4 (AD+PD), PV without macrosetae. Mesotrochanter with one stout setae. Mesofemur row AV without short setae. Hind femur macrosetal formula 2+2+1, without extra setae basad of usual set. Metatarsomere I longer than metatarsomeres II and III combined, not expanded apically, plantar setae simple, pecten with 3 platellae and inner apical seta tapered. Male genitalia. Apodemes of sternite II inconspicuous. Basolateral pygofer apodemes weakly developed. Pygofer moderately long, well-sclerotized dorsally, without processes, macrosetae grouped near posterior margin, basolateral cleft present, with large membranous area ventrally near base, posteroventral margin serrate with several evenly spaced teeth. Anal tube short and weakly sclerotized dorsally. Valve partially fused to subgenital plates medially, articulated with pygofer. Subgenital plate large, triangular, lateral margin convex, apex rounded, macrosetae uniseriate laterad with some additional irregular setae near margin. Connective U-shaped, stem poorly developed. Style broadly bilobed basally, medial lobe not extended dorsad, preapical lobe obsolete, apophysis elongated and smoothly curved mesad then laterad with apex slightly expanded and blunt. Aedeagus with atrium poorly developed, shaft strongly recurved, with many lateral denticles arranged somewhat irregularly, with small apical flange and pair of slender apical processes, gonoduct not sclerotized, gonopore medial on ventral surface, phragma with conspicuous setae.
Female genitalia. Unknown Distribution. Brazil, Bolivia, Paraguay and Argentina Remarks. The genus is closely similar to Goiattus, which it resembles in the shape of the head and pronotum but with major differences in the male genitalia such as pygofer with large membranous area with posteroventral margin serrate and the aedeagus slender with lateral denticles.
Pseudalaca multipunctata (Osborn) (Figs. 13 B,E and I,29,31 H) Agallia multipunctata Osborn, 1923: 10 Alaca multipunctata Oman, 1938: 386 Pseudalaca multipunctata Linnavuori, 1959: 236 Redescription. External morphology. Crown yellowish with stripes and minute black spots centrally. Frontoclypeus with transversal brownish stripes plus a longitudinal yellowish midline. A pair of circle black spots in ocelluclar area. Anteclypeus and lorum brownish. Gena yellowish marked with brownish marks. Pronotum brownish with longitudinal black, brown and yellowish stripes. Forewing brownish. Venter yellowish. Front trochanter and femur marked with black.
Male genitalia. Pygofer 2x longer than tall, macrosetae grouped at caudal apex. Anal tube wide apically. 1). External morphology. Robust medium-sized (5.3 mm) leafhopper, overall coloration brownish with stramineous regions; with symmetrical and asymmetrical black spots. Head wide as pronotum. Crown very short, arched above eyes and anterior part of pronotum, anterior margin rounded, transition to face rounded, width between eyes less than twice eye width, texture shagreen. Ocellocular area parallel sided. Ocelli slightly below anterior margin of head, not visible in dorsal view, distance to adjacent eye approximately 3x ocellar diameter. Frontoclypeus with lateral margins somewhat parallel and reaching ocelli, weakly tumid medially, erect fine seta close to lateral margin. Antennal ledge absent, antennal base close to lower corner of eye, antenna length 1.0x width of head. Anteclypeus paralleled, weakly tumid in lateral view, extended slightly more than normal curve of face margin. Lorum shorter than anteclypeus in width near base. Gena slightly incised below eye.
Pronotum strongly convex, anterior margin parabolic, posterior margin concave; lateral margin weakly carinate, shorter than half eye width. Scutellum slightly protuberant, broad. Forewing macropterous, appendix reduced, apex rounded, without extra crossveins, with three anteapical cells, outer apical cell shorter than adjacent cell with apex delimited by crossvein, apical cells subequal in width, inner anteapical cell open (without m-cu2 crossvein), Claval veins separate throughout length, Pcu unconnected to claval suture, Pcu-A1 crossvein present. Hindwing scheme fully developed, without pigmentation, RP-MA and MP-CuA separated by crossvein. Front femur with long AM1 at mid-height, IC setae very long and thin (12-19), row AV with hair-like setae (8-15), AV1 poorly differentiated from IC row. Front tibia dorsal macrosetal formula 1+4 (AD+PD), PV without macrosetae. Mesotrochanter with one stout setae apically. Mesofemur row AV without setae. Hind femur macrosetal formula 2+2+1, without extra setae basad of usual set. Metatarsomere I twice as long as II and III combined, not expanded apically, plantar setae simple, pecten with 5 platellae and inner apical setae tapered.
Male genitalia. Apodemes of sternite II inconspicuous. Basolateral pygofer apodemes absent. Pygofer moderately broad, strongly pointed, macrosetae grouped posterad, well-sclerotized dorsally, caudolateral margin striate, without processes, basolateral cleft present. Anal tube short and broad, sclerotized dorsally. Valve free and articulated with subgenital plates and pygofer. Subgenital plate triangular, slightly shorter than pygofer length, free from each other, apex rounded, macrosetae uniseriate distant from lateral margin. Connective Y-shaped, stem longer than arms. Style bilobed basad, medial lobe not extended dorsad, preapical lobe absent, apophysis short and straight, apex blunt. Aedeagus with atrium weakly developed, curved dorsad; shaft in lateral view broad at base then slender in distal portion, without basal process, without denticles, lateral flanges developed; apex with pair of retrorse processes, curved basad. Gonoduct not sclerotized, gonopore subapical on dorsal surface, phragma membranous.
Female genitalia. Unknown. Distribution. Brazil Etymology. The genus name is masculine and refers to the type locality of the type species, Goiás state, Brazil.
Remarks. This genus is similar to Brazosa, Zabroza and Pseudalaca in external and internal features. The latter genus, Pseudalaca, appears closely related, but Goiattus differs from it in having the pygofer striated and corrugated, and the aedeagus not dentate but robust with retrorse distal processes (see also Table 1).
Goiattus reyesi Pinedo-Escatel, sp. nov. (Figs. 13 C,F and J,30,31 G) urn:lsid:zoobank.org:act:9C100174-1431-4A38-900A-68344D22904B Description. External morphology. Crown stramineous with minute pair of black spots on posterior margin next to midline, big black circular spot on anterior margin. Face with black spot above and below ocelli. Frontoclypeus with a broad black median longitudinal line. Anteclypeus black. Lorum stramineous with lateral suture black. Gena stramineous marked below antennal pit with black. Pronotum brownish with irregular black band on anterior margin behind eyes. Scutellum posterior margin stramineous and anterior brownish. Forewing brownish. Venter stramineous with black. Front femur black basally. Metafemur black.
Male genitalia. Pygofer pointed and striate anteroventrally, macrosetae concentrated at apex. Anal tube black. Connective, stem longer than arms. Style, apophysis straight with blunt apex. Aedeagus curved dorsad, shaft inflated basally and slender apically with lateral flanges and apical processes.
The Neotropical taxa described above appear to form a morphologically distinct group within the poorly defined and polyphyletic tribe Athysanini. Most have the crown of the head relatively short and broad in dorsal view, rounded to the face, and often more or less arched, and the pronotum in some cases is declivous anteriorly and more convex than usual. All except Spaltumtettix also have paired black spots or lines on the head and most also have bright orange dorsal markings. In the most recent key to Neotropical genera of this tribe (Linnavuori 1959, as "Euscelini") some of the included genera will run to Brazosa or Zabrosa. Goiattus and Spaltumtettix are similar to Brazosa in having a short and rounded head and are also similar in body size and the male genital capsule and genitalia also have numerous features in common (Table 1). Dietrich & Rakitov (2002) noted that Neotropical athysanine leafhoppers are important because several exhibit morphological affinities to different deltocephaline tribes and may represent intermediate stages in the evolutionary diversification of cicadellids. Several genera of Athysanini from South America seems to lack affinities to North American groups and, therefore, presumably represent deltocephaline lineages that originated in South America (Dietrich & Rakitov 2002). Other New World genera of this tribe present in subtropical regions (e.g., in southern Mexico) show similarities to both Nearctic and Neotropical genera (Pinedo-Escatel et al. 2016).
The new genera described above appear to be endemic to South America. The morphological limits of some of these genera are not clear and more detailed taxonomic study and further phylogenetic research are needed to clarify their status and relationships to other genera in the tribe. More detailed studies including phylogenetic analyses are needed to better understand hotspots in diversification and evolution of Neotropical athysanine lineages. The comprehensive phylogeny of Zahniser & Dietrich (2013) included only two of the genera treated here (Brazosa and Napo) and they were part of a large and almost exclusively New World clade including Athysanini (in part), Pendarini, Bahitini and Scaphytopiini. This previous analysis indicates that many of the deltocephaline genera endemic to the Neotropical region, including those treated here, are related to each other and diversified in isolation from the deltocephaline faunas of other parts of the world.