Pandara DeLong & Freytag, 1972 stat. nov.

(Figs 1–6)

Type-species: Gyponana (Pandara) eleganta DeLong & Freytag, 1972

Diagnosis. Medium sized and fusiform leafhoppers (Figs 6A–M); dull yellow, crown (Figs 1A, 4A) with two pairs of brown maculae, one pair anterior to ocelli and one pair adjacent to posterior margin of the crown; pronotum (Figs 1A, 3A) with dark punctures; forewing (Figs 1D, 4D) mottled with brown maculae. Head (Figs 1A, 2A) in dorsal view, moderately produced; median length of crown approximately half of interocular width. Ocelli (Figs 2A, 4A) slightly closer to anterior margin and equidistant from median line and inner margin of eyes. Clypeus (Figs 1B, 4B) with lateral margins slightly converging to apex. Frontogenal suture (Figs 2B, 3B) very close to inner margin of the eyes. Crown-face transition (Figs 1C, 2C) distinct, thick and striated. Forewing (Figs 2D, 3D) with venation not reticulated; appendix well developed. Male sternite VIII (Figs 1E, 4E) wider than long, subrectangular. Male pygofer (Figs 1H, 4H) with dorsal margin with small preapical process. Connective (Figs 1J, 3J) Y-shaped. Style (Figs 2I, 4J) with outer lobe well developed, subtriangular; blade (Figs 1K, 4K) elongated, apex expanded with dorsal tooth. Aedeagus (Figs 1L–N, 3L–M) with shaft long curved anterodorsally, with one or two pair of subapical processes, dorsal pair (DPP), when present, short, spiniform; ventral pair (VPP) usually bifid. Ovipositor with second valvula (Figs 5K–L) with several small dorsal teeth at apical third.

Coloration. Dull yellow (Figs 6A–M). Crown (Figs 1A, 3A, 5A) with two pairs of brown maculae, one pair extending from ocelli to anterior margin of crown, a second pair behind ocelli, adjacent to posterior margin of the crown. Ocelli (Fig 4A) light red. Pronotum (Figs 1A, 3A) with irregular brown maculae near anterior margin, surface with several dark punctures on disc and posterior third; lateral lobe (Figs 1C, 5C) with a dark brown band below dorsopleural carina. Mesonotum (Figs 2A, 4A) with a dark brown longitudinal band at lateral corners. Forewing (Figs 2D, 5D) dull yellow subhyaline at basal half and light yellow hyaline at apical half; costal area with a dark brown band extending from base to RM vein bifurcation and a large dark brown macula at middle third, between costal margin and R vein; membrane with several small brown and dark brown maculae distributed over the entire length of wing, being more numerous at apical half; veins with apex dark brown; appendix fuscous at basal and apical portion. Legs dull yellow with brown areas. Metatibia with basis of cucullate setae dark brown.

Redescription. Head (Figs 1A, 4A), in dorsal view, with crown moderately produced, median length about half length of interocular width, anterior margin broadly rounded, median portion longer than region adjacent to eyes, surface slightly concave near ocelli, texture with fine transverse parallel striations, more evident near apex; ocelli slightly closer to anterior margin and equidistant from median line and eyes. Head (Figs 1B, 3B), in ventral view, with face slightly wider than high; frontogenal suture very close to inner margin of the eyes, extending dorsally above antennal ledge to anterior margin of crown; antennal ledge carinated, obliquely downwards in relation to frons and extending over frons; frons approximately as high as wide; epistomal suture distinct, but incomplete, evanescent medially; clypeus approximately as wide as high; lateral margins slightly converging to apex; rectilinear apex; not inflated; maxillary plate very narrow, produced ventrally as far as clypeus apex; gena with ventrolateral margins slightly convex near eyes. Head (Figs 1C, 3C) with crown-face transition defined and thick; with five to seven distinct transverse striae; frons slightly swollen and concave below anterior margin of crown.

Pronotum (Figs 2A, 4A), in dorsal view, with superficial transverse striae on disc and posterior third; lateral margins convergent anterad; in lateral view (Figs 2C, 3C), rounded and moderately declivous; head and pronotum in continuous slope. Exposed part of mesonotum (Figs 1A, 3A) wider than long; in lateral view (Figs 2C, 4C), scutellum slightly inflated. Forewing (Figs 3D, 5D) with venation evident, without extra crossveins; cross vein mcu 1 positioned anterior to cross vein r-m 1; appendix well developed and involving first and second apical cells, as wide as maximum width of first apical cell; apex rounded. Profemur four times longer than high; AV row absent, and PV row about four-five setae spaced; AD, PD, and AM row absent except to AD 1, PD 1, and AM 1; IC row formed by a lightly arched comb of fine setae at distal half of femur extending to AM 1; protibia approximately triangular in cross section; AV row formed approximately by 13 setae increasing in size towards apex; PD row formed by four long setae spaced along entire length of tibia; PV row developed, with four setae; metafemur with setal formula 2:2:1; metatibia AD row with three intercalary setae between macrosetae; PV row with setae of apical half long and heterogeneous in length and thickness; first tarsomere with two rows of setae not cuculate, the outer is remarkably smaller, apex with three-four platellae; second tarsomere apex with three apical platellae.

Male terminalia. Sternite VIII (Figs 1E, 3E) subrectangular, wider than long; posterior margin rounded, slightly emarginated medially. Valve (Figs 1F, 4F) wider than long; integument thickening on anterior margin. Pygofer (Figs 1G, 3G, 4G) subtriangular, high at base and tapering towards the apex; ventral margin broadly rounded; numerous macrosetae distributed throughout apical half; dorsal margin concave preapically, with small preapical or apical serrated process (Figs 2G, 4H). Subgenital plate (Figs 2F, 4G) in lateral view, extending to apical third of pygofer; in ventral view (Figs 1I, 4I), elongated; usually widened at the base and gradually narrowing towards the apex Connective (Figs 1J, 3J) Y-shaped. Style (Figs 2I, 4J) in dorsal view, with outer lobe strongly produced, approximately subtriangular; in lateral view (Figs 1K, 4K), apophysis elongated and enlarged towards apex; apex blunt with dorsal tooth. Aedeagus (Figs 1L–N, 3L–M, 4L–M) without atrial processes; preatrium reduced; shaft dorsoventrally flattened, sometimes cylindrical, elongated and curved anterodorsally; with one or two pairs of preapical processes, dorsal pair (DPP) short or elongate and spiniform, ventral pair (VPP), when present, bifid.

Female terminalia. Ovipositor (Figs 5E–F) not surpassing apex of pygofer. Internal sternite VIII membranous. Second valvula (Figs 5K–L) with dorsal protuberance at basal half; apical third with several small dorsal teeth.

Distribution. Bolivia (Santa Cruz); Brazil (Amazonas); French Guiana; and Peru (Amazonas, Cusco).

Notes. Pandara was originally described as a subgenus of Gyponana . However, differences between the Pandara species and the other Gyponana are notable. Firstly, they differ in geographic distribution. While most Gyponana species are distributed in Central and North America (except for Zerana), Pandara species are restricted to South America (Fig. 7). Secondly, Gyponana species are mostly completely green (yellow when preserved) without distinct spots or some species with orange longitudinal stripes on pronotum. While in Pandara species have a constant color pattern, with two dark pairs of maculae on the crown, dark punctures on the pronotum and mottled brown maculae on forewings. In Gyponana, the forewing frequently has reticulate venation and a reduced appendix, while in Pandara venation is not reticulate and the appendix is well developed. In relation to the male genitalia, there are also some differences: in Gyponana the pygofer does not have processes on the dorsal margin, the styles have reduced or poorly projected lateral lobes and the apex usually foot-shaped, and the aedeagus generally has an apical pair of unbranched processes, each not bifurcated. In Pandara, the pygofer bears a lobiform process on the dorsal margin, the style has an expanded apical portion with a blunt apex, and the aedeagus bears subapical processes, each generally bifid.

Furthermore, in an unpublished phylogeny of Gyponini based in based on 182 morphological characters combined with sequence data (28S and 16S rDNA and cytochrome oxidase subunit I), recovered Pandara as monophyletic with moderate support, separated from other Gyponana species and forming a clade with representatives of the genera Nullana DeLong, 1980, Regalana DeLong & Freytag, 1975, and Alapona DeLong, 1980 (Gonçalves 2016). In fact, these genera have morphological characteristics similar to Pandara species, for example: crown with parallel striations; crown-face transition distinct with five or more striae, forewing with non-reticulated venation, membrane covered by mottled dark maculae and well-developed appendix; connective Y-shaped; and style with well-produced lateral lobes, except in Nullana .

Considering the phylogenetic results and clear morphological differences between species of Pandara and species of the other subgenera of Gyponana, we propose herein to elevate Pandara to genus rank.

Species of Pandara stat. nov. (Fig. 7)

Pandara eleganta DeLong & Freytag, 1972: 160 . Bolivia (Santa Cruz).

Pandara inca sp. nov. Peru (Amazonas).

Pandara indivisa sp. nov. Bolivia (Santa Cruz); Peru (Cusco).

Pandara parallela sp. nov. Brazil (Amazonas); French Guiana.

Pandara species occur in very different biomes in South America (Fig. 7), such as the Cerrado savannah in Central Bolivia; moist tropical forests, such as the Amazonian rainforest and Guianan Eastern Lowland; and cloud forests such North-Central Moist Andes.