A new genus and five new species of Neocoelidiinae leafhoppers from Brazil (Insecta: Hemiptera: Cicadellidae) with a key to males

A new Brazilian leafhopper genus of the subfamily Neocoelidiinae is proposed: Takiyaella gen. nov. based on T. sexguttata (Chiamolera & Cavichioli, 2003) gen. et comb. nov. and five new species: T. anomala gen. et sp. nov., T. coelhomarquesae gen. et sp. nov., and T. daniela gen. et sp. nov. from the state of Rio de Janeiro; T. cavichiolii gen. et sp. nov. from the state of Paraná, and T. mejdalanii gen. et sp. nov. from the state of Minas Gerais. A distribution map and identification key to Takiyaella species are provided.


Introduction
includes leafhoppers restricted to the New World, most of which are endemic to the Neotropical region and only four genera occur in the Nearctic region (Nielson & Knight 2000). Currently, the subfamily is divided into two tribes: Neocoelidiini Oman, 1943 with 28 genera and 175 species and Krocodonini Marques-Costa & Cavichioli, 2012 with 14 species allocated in four living genera and one fossil (Dietrich 2003;Marques-Costa & Cavichioli 2012;Coelho-Marques & Ale-Rocha 2016Gonçalves & Mejdalani 2017). In the phylogenomics of Membracoidea Rafi nesque, 1815 (Dietrich et al. 2017), using 388 loci and more than 99,000 aligned nucleotide positions, Neocoelidiinae was recovered as a sister group of Portanini Linnavuori, 1959 (Aphrodinae), both as a sister group of Deltocephalinae Dallas, 1870. Neocoelidiines are small to large leafhoppers, measuring 3-14 mm, commonly having coloration in shades of green, yellow or orange, and are distinguished from other subfamilies by the following combination of characters: (1) crown generally elevated and fl at between the eyes; (2) ocelli in the transition between the crown and frons or close to it, rarely located on the crown or face; (3) frons with muscle impressions inconspicuous or slightly visible; (4) very long antenna, often exceeding the total body length; (5) pronotum with posterior margin usually emarginate V-shaped; (6) mesothorax frequently well developed, pleurae extending to the sternum, giving individuals a swollen aspect ventrally; (7) forewing with venation usually indistinct, except apically and (8) with transverse vein r-m 1 absent; (9) hind wing with R 4+5 and M 1+2 veins usually confl uent preapically, fused at the apex, forming a single vein; and (10) valve of the male terminalia generally entirely fused to the subgenital plates (Kramer 1964;Dietrich 2003;Marques-Costa & Cavichioli 2012).

Etymology
The generic name Takiyaella (feminine noun) is a tribute to Prof. Dr Daniela Maeda Takiya in recognition of her remarkable contribution to our knowledge of the Brazilian Auchenorrhyncha.
Description HEAD AND THORAX. Head, in dorsal view (Figs 1A,7A), slightly produced anteriorly, median length of crown approximately equal or slightly less than interocular width; transocular width about 4 /5 humeral width of pronotum; crown subrectangular, anterior margin rounded, surface fl at, texture shagreen; ocellus medium-sized, on anterior margin of head, distant from eye margin, visible in dorsal view; coronal maculae distinct at basal half, between midline and eyes; coronal suture distinct in basal third, evanescent anteriorly. Head, in frontal view (Figs 1B,10B), with face slightly higher than wide; frontogenal suture extending to antennal ledge but not reaching ocellus; antennal ledge narrow, slightly prominent, oblique and weakly carinated; frons approximately 1.8 × as long as wide; muscle impressions indistinct; epistomal suture distinct but incomplete, evanescent medially; clypeus 1.5 × as long as maximum width, lateral margins parallel, apex slightly emarginated; maxillary plate produced ventrally, reaching clypeus apex; lorum ellipse-shaped, apical margin not reaching apex of clypeus; gena incompletely covering episternum. Head, in lateral view (Figs 1C,3C), with crown-face transition rounded, without carina; lateral margins of crown, adjacent to eyes, elevated and not carinated; antennal pits at same level as an imaginary line tangent to anteroventral angles of eyes; antenna with long fl agellum, exceeding half-length of forewing; frons convex. Pronotum (Figs 1A,5A) with inconspicuous transverse striae on disc; lateral margins rounded, convergent anterad, and slightly shorter than eye length; posterior margin roundly excavated; in lateral view (Figs 1C,8C), slightly declivous; dorsopleural carina present and complete. Mesonotum (Fig. 8A) as long as wide. Forewing (Figs 1D, 7D) mostly semi-hyaline, except for apical cells and apex of anteapical cells hyaline, 3.5 × as long as maximum width; venation indistinct, except apically; two anteapical cells, crossvein sectorial ʻsʼ missing; four apical cells: fi rst to third approximately rectangular, third enlarged apically, fourth rhomboid, base of second more proximal than base of third; appendix narrow; apex rounded. Hind wing with vein R 4+5 and M 1+2 preapically convergent, fused to each other at apex, forming single vein. Profemur with AD, AM, and PD rows reduced and poorly defi ned, with exception of apical setae AD 1 , AM 1 , and PD 1 , respectively; AV and PV rows absent; IC row formed by slightly arched comb of fi ne setae, beginning at distal half of femur and extending to apex. Protibia, in cross-section, semi-circular; AV row formed by approximately 20 setae, slightly longer and thicker towards apex; AD and PD rows without differentiated setae; PV row with 2-3 widely spaced setae. Metafemur with setal formula 2:2:1, with inner setae of second pair reduced in size. Metatibial AD row with 2-4 intercalary setae between macrosetae; PD, AD, and PV rows with 13-15, 10-12, and 28-32 macrosetae, respectively; AV row with approximately 11 macrosetae distributed along most of tibia, except in the proximal and distal portions. Metatarsomere I longer than combined length of two distal tarsomeres; plantar surface with two rows of setae, external row with longer and robust setae than inner row; pecten with fi ve platellae. Metatarsomere II pecten with three platellae.

Distribution
Brazil (states of Minas Gerais, Paraná and Rio de Janeiro). Species of Takiyaella gen. nov. occur in southern and southeastern Brazil, in the Atlantic Rainforest biome (Fig. 13).
However, the new genus can be easily separated from Paraphysiana due to hind wing with veins R 4+5 and M 1+2 preapically convergent, fused at apex, forming single vein, whereas in Paraphysiana R 4+5 and M 1+2 are convergent, fused to each other, but they are divergent apically; pygofer without inner ventral processes (present in Paraphysiana); subgenital plates without dorsal tooth (present in Paraphysiana); style with preapical lobe moderately developed, whereas in Paraphysiana it is strongly developed and produced caudally; paraphysis absent (present in Paraphysiana); and the aedeagus with atrial processes (absent in Paraphysiana).

Material
COLORATION. Pale yellow ( Fig. 12A-B). Forewing ( Fig. 1D) yellow, translucent, with three dark maculae: fi rst on claval comissure between anal veins, second on apex of clavus and extending to brachial cell, and third on fourth apical cell.
HEAD AND THORAX. External morphological characters as in generic description. MALE TERMINALIA. Pygofer, in lateral view (Fig. 1E), 1.6 × as long as maximum height, not expanded apically, basal portion slightly higher than apical portion; dorsal margin slightly convex; ventral tooth  prominent and rounded; caudal margin truncated; few small setae scattered on middle third. Subgenital plate, in lateral view (Fig. 1F), 2.9 × as long as maximum height, expanded medially; dorsal margin slightly convex; ventral margin oblique in the apical half; apical third strongly narrowed; in ventral view (Fig. 1G), subtriangular, lateral margins parallel on basal half and strongly converging on apical half; ventral surface with pair of uniseriate rows of 5-6 macrosetae; apex acute. Connective (Fig. 1H) Y-shaped, total length about 3 /5 length of style; stem approximately as long as arms. Style (Fig. 1H-I) with dorsal margin of apodeme broadly rounded, ventral margin acutely produced anteroventrally; apophysis elongated, robust. Aedeagus (Fig. 1J-K) with pair of slender atrial processes, each process with approximately half length of shaft, directed posterad and strongly divergent in ventral view, apical third with three spiniform projections, one longer on dorsal margin, and two smaller on apex; shaft fl attened laterally, curved dorsally at basal third, with pair of small subapical ventral lamellae. Anal tube ( Fig. 1E) with sternite X long and tubular, without basiventral process.
COLORATION. Pale yellow (Fig. 12C-D). Forewing (Fig. 3D) translucent, with three small dark maculae on claval comissure: fi rst, smaller, between base and second anal vein, second, between anal veins, and third on apex of clavus and extending to brachial cell; in some specimens the fi rst pair is absent, and in one specimen, fi rst and second pairs are absent.
HEAD AND THORAX. External morphological characters as in generic description. MALE TERMINALIA. Pygofer, in lateral view (Fig. 3E), 1.5 × as long as maximum height, expanded apically, apical portion higher than basal portion; dorsal margin with prominent preapical subtriagular lobe; ventral tooth broadly rounded; caudal margin with ventral rounded lobe; few small setae scattered on apical half; apical portion with inner integument thickening. Subgenital plate, in lateral view (Fig. 3F), subtriangular, 3.3 × as long as maximum height, slightly expanded medially; in ventral view (Fig. 3G), subrhomboid, lateral margins converging anterad on basal one-fi fth and converging towards apex on apical four-fi fths; ventral surface with one or two pairs of apical macrosetae; apex slightly truncated. Connective (Fig. 3H) V-shaped, total length approximately ⅔ length of style; stem very short. Style ( Fig. 3H-I) with apophysis very elongated and slender; ventral margin distinctly rounded near median third. Aedeagus (Fig. 3J-L) with pair of atrial processes parallel and following curvature of shaft, each process approximately ½ length of shaft, broad at base and tapering towards acute apex; shaft slightly fl attened dorsoventrally, apical portion expanded laterally, spatulate, lateral margins with some small spiniform projections. Anal tube (Fig. 3M) with sternite X long and tubular, with triangular basiventral process with denticles. FEMALE TERMINALIA. Sternite VII (Fig. 4A-C) as long as wide; posterior margin slightly excavated medially. First valvula of ovipositor ( Fig. 4F) with dorsal sculptured area areolate-strigate. Other characters as in generic description.

Remarks
Takiyaella cavichioli gen. et sp. nov. is most similar to T. sexguttata (Chiamolera & Cavichioli, 2003) gen. et comb. nov. by the shape of male pygofer with two lobes, one dorsal and one apical (Figs 3E,10D) and the shape of the aedeagal shaft expanded apically, with lateral margins rounded, bearing several small spiniform projections (Figs 3J-L, 10I-J). However, T. cavichioli gen. et sp. nov. differs in having the style slender and the aedeagus with atrial processes parallel to the shaft and following its curvature (Fig. 3J), while in T. sexguttata gen. et comb. nov. the style is wider, and the atrial processes are shorter, not parallel to the shaft (Fig. 10I).

Diagnosis
Male pygofer (Fig. 5E) expanded apically, with apical lobe. Aedeagus (Fig. 5J-K) with two pairs of slender atrial processes anterior to shaft curvature, one pair curved dorsally, and the second directed ventrally; shaft with apical portion expanded laterally forming a small process on each side; anal tube ( Fig. 5L) with basiventral process truncated with few denticles.
COLORATION. Pale yellow (Fig. 12E-F). Forewing (Fig. 5D) translucent, with three small dark maculae on claval comissure: fi rst, smaller, between base and second anal vein, second, between anal veins, and third on apex of clavus and extending to brachial cell. HEAD AND THORAX. External morphological characters as in generic description. MALE TERMINALIA. Pygofer, in lateral view (Fig. 5E), 1.7 × as long as maximum height, expanded apically, apical portion higher than basal portion; dorsal margin slightly concave medially; ventral tooth slightly rounded; caudal margin truncated with small ventral rounded lobe; few small setae scattered near ventral margin and apical half; apical portion with inner integument thickening. Subgenital plate, in lateral view (Fig. 5F), subtriangular, 3.3 × as long as maximum height; in ventral view (Fig. 5G), subovoid, lateral margins converging towards apex; ventral surface with pair of uniseriate rows of 5-6 macrosetae; apex acutely rounded. Connective (Fig. 5H) V-shaped, total length approximately ⅔ length of style; stem very short. Style (Fig. 5H-I) with dorsal margin of apodeme broadly rounded; apophysis elongated and robust. Aedeagus (Fig. 5J-K) with two pairs of slender atrial processes with acute apexes: fi rst more dorsal in comparison to second, longer, approximately ⅓ ength of shaft, curved dorsally, anterior to shaft curvature; second pair, shorter, approximately ⅓ length of fi rst, directed ventrally; shaft tubular, slightly fl attened dorsoventrally at apex, ventral surface with longitudinal row of denticles on each side, extending from base to apex, apical portion expanded laterally forming small process on each side. Anal tube (Fig. 5E, L) with sternite X long and tubular, with basiventral process truncated, with few denticles.

Remarks
Takiyaella coelhomarquesae gen. et sp. nov. is most similar to T. daniela gen. et sp. nov. by the shape of apex of aedeagus expanded laterally forming a small process on each side (Figs 5K,7K). However, T. coelhomarquesae gen. et sp. nov. differs by the dorsal margin of the male pygofer without distinct lobe (Fig. 5E); aedeagus with two pairs of atrial processes (Fig. 5J) and anal tube with basiventral process (Fig. 5L). While T. daniela gen. et sp. nov. have the dorsal margin of the male pygofer with distinct lobe (Fig. 7E); aedeagus with a pair of bifi d atrial processes (Fig. 7J) and anal tube without basiventral process (Fig. 7E).

Etymology
The specifi c epithet is a tribute to Prof. Dr Daniela Maeda Takiya in recognition of her remarkable contribution to our knowledge of the Brazilian Auchenorrhyncha, and also in gratitude for her great friendship and for bringing the fi rst author to the study of leafhoppers. The species epithet is treated as a noun in apposition. Description MEASUREMENTS (mm). Holotype (♂): total length 6.5. Paratypes (♂): 6.3-6.6 (n = 2).

Holotype
COLORATION. Pale yellow (Fig. 12G-H). Forewing (Fig. 7D) translucent, with three small dark maculae on claval comissure: fi rst, between base and second anal vein; second between anal veins and third on apex of clavus and extending to brachial cell. HEAD AND THORAX. External morphological characters as in generic description. MALE TERMINALIA. Pygofer, in lateral view (Fig. 7E), 1.8 × as long as maximum height, expanded apically; dorsal margin with prominent preapical lobe; caudal margin with ventral subacute lobe; ventral tooth broadly rounded; apical portion with inner integument thickening. Subgenital plate, in lateral view (Fig. 7F), subtriangular, 2.9 × as long as maximum height; in ventral view (Fig. 7G), subovoid, lateral margins converging towards apex; ventral surface with pair of uniseriate rows of 3-4 macrosetae; apex acute. Connective (Fig. 7H) V-shaped, total length approximately ⅔ length of style; stem very short. Style (Fig. 7H-I) with apophysis elongated and robust. Aedeagus (Fig. 7J-K) with pair of bifi d atrial process: dorsal ramus about ⅓ length of shaft, directed dorsally, anterior to curvature of shaft; ventral ramus approximately ⅓ length of dorsal ramus, directed posteriorly, posterior to curvature of shaft; shaft tubular, slightly fl attened dorsoventrally at apex, ventral surface with longitudinal row of denticles on each side, extending from base to apex, apical portion expanded laterally forming small process on each side. Anal tube (Fig. 7E) with sternite X long and tubular, without basiventral process.

Remarks
Xenocoelidia sexguttata was described by Chiamolera & Cavichioli (2003) based on two males and one female from Rio de Janeiro, Brazil. Our study of the X. sexguttata types (Figs 10-11, 12K-L) indicated the following morphological incongruities with the other species of Xenocoelidia: (1) male pygofer with caudal margin with small lobe without robust setae (Fig. 10D), whereas in other species of Xenocoelidia the caudal margin of pygofer is broadly rounded bearing several small and robust setae; (2) subgenital plates fused to each other in the basal two-thirds, with only the apical third being free (Fig. 10F), while in other species the subgenital plates are fused only at base, being free for most of its length; (3) connective with stem short (Fig. 10G), while in other species the connective is Y-shaped, with stem long; (4) style with preapical lobe moderately developed and slightly produced posterad (Fig. 10H), whereas other species of Xenocoelidia have the preapical lobe strongly developed and expanded laterally; and (5) aedeagus with shaft moderately long, wider at base, bearing atrial processes (Fig. 10I), while in other species of Xenocoelidia the aedeagus lacks atrial process, and the aedeagal shaft is very long and narrow in almost all its length. Based on these differences, we propose the transfer of X. sexguttata to Takiyaella gen nov.
Takiyaella gen nov. differs from the other mentioned genera by the forewing (Figs 1D,7D) with two or three dark maculae on clavus, crossvein ʻsʼ missing; hind wing with veins R 4+5 and M 1+2 convergent preapically and fused to each other at apex, forming a single vein; male pygofer (Figs 5E,8E) with a sclerotized and dentated ventral tooth on apical portion and caudal margin usually with ventral rounded lobe; valve (Figs 3G,8G) without indication of separation with subgenital plates; subgenital plates (Figs 7G,10F) fused to each other in the basal two-thirds; connective (Figs 3H, 7H) usually with stem reduced; style (Figs 1H,7H) with preapical lobe moderately developed; and the aedeagus (Figs 5J,7J,8J) with one or two pairs of atrial processes. In addition, all species known for Takiyaella gen. nov. have distribution restricted to the Atlantic Rainforest biome (Fig. 13), unlike what occurs with the species of the other genera mentioned above, which mostly are distributed in the Amazon Rainforest biome, except for Chinaia bidentata Chiamolera & Cavichioli, 2002 that is found in the Atlantic Rainforest.