Three new genera from South America and some taxonomic changes in Gyponini (Insecta: Hemiptera: Cicadellidae)

. Three new South American leafhopper genera of the tribe Gyponini are proposed: Beltrana gen. nov. based on Beltrana reticulata gen. et sp. nov. from French Guiana, Fulana gen. nov. based on Fulana brasiliensis gen. et sp. nov. from Brazil, and Sicrana gen. nov. based on Sicrana plana gen. et sp. nov. from Brazil and Ecuador. Diagnoses, detailed descriptions, and illustrations are provided for each taxon, as well as comparisons with closely related genera. In addition, the following synonyms are proposed: Freytagana DeLong, 1975 as a junior synonym of Marganana DeLong, 1948 and F. gibsoni DeLong, 1975 as a junior synonym of M. ( Marganana ) mexicana DeLong & Freytag, 1963. Chilella DeLong & Freytag, 1967 is transferred from Gyponini to Selenomorphini Evans, 1974. Three new genera from South America and some taxonomic changes in Gyponini (Insecta: Hemiptera: Cicadellidae). European 750: as dorsoapical apex

could include more than 500 species. Currently, considering the recently described taxa, the tribe comprises 1405 species in 71 genera Freytag 2021). Based on the large number of undescribed species deposited in Brazilian collections, we believe that the diversity of Gyponini will exceed 2000 species. Specimens can be collected using sweeping nets and sometimes yellow plates; however, light and Malaise traps are the most effective methods for collecting these leafhoppers (Domahovski & Cavichioli 2018). This paper proposes the establishment of three new Gyponini genera, with very peculiar morphological features: Beltrana gen. nov. from French Guiana, Fulana gen. nov. from Brazil, and Sicrana gen. nov. from Brazil and Ecuador. The names "Fulana, Beltrana, and Sicrana" form a Portuguese expression equivalent to the English expression "Tom, Dick, and Harry". This expression is used to refer to multiple unspecified people. The names were chosen because this expression is very popular in Brazil and, also, they end in '-ana' which is a common suffix in most generic names of Gyponini.

Type species
Beltrana reticulata gen. et sp. nov., by present designation and monotypy.

Etymology
The generic name is feminine. Beltrana, together with Fulana and Sicrana, forms a very popular expression in Brazil ("Fulana, Sicrana e Beltrana"), which refers to unspecified people or people whose actual names should not be mentioned.

Description
Head and tHorax. Head in dorsal view ( Fig. 1A) moderately produced anteriorly, median length of crown slightly less than interocular width; transocular width about six-sevenths humeral width of pronotum; crown with anterior margin broadly rounded, surface mostly flat, slightly concave medially, with subtle oblique striae between ocelli and adjacent to anterior margin; ocellus medium-sized, equidistant between median line and eye margin, closer to posterior than anterior margin of crown; coronal suture distinct only on basal portion of crown. Head in ventral view ( Fig. 1B) with face slightly higher than wide; frontogenal suture distant from eye margin by slightly less than maximum width of clypeus and extending to anterior margin of crown; antennal ledge carinated and adjacent to anterior margin of crown, obliquely ascending and not extending over frons; frons approximately 1.7 times as long as wide; epistomal suture indistinct; clypeus approximately 1.3 times as long as maximum width, lateral margins parallel, apex slightly emarginated; maxillary plate produced ventrally, not reaching clypeus apex; gena excavated just below eye, ventrolateral margin slightly excavated. Head in lateral view ( Fig. 1C) with crown-face transition distinct and foliaceous, with two carinae; frons concave below anterior margin of crown. Pronotum ( Fig. 1A) with transverse striae on disc and posterior half; lateral margins straight, convergent anterad, and approximately as long as eye length; posterior margin excavated; in lateral view (Fig. 1C (2021) very narrow. Profemur with AD, AM, and PD rows reduced and poorly defined, with exception of apical setae AD 1 , AM 1 , and PD 1 , respectively; AV and PV rows formed by 4-5 setae; IC row formed by slightly arched comb of fine setae, beginning at distal third of femur and extending to AM 1 . Protibia, in cross-section, semi-circular; AV row formed by long setae, slightly longer and thicker towards apex; AD and PD rows without differentiated setae; PV row with 5 setae. Metafemur with setal formula 2:2:1. Metatibial AD row without intercalary setae between macrosetae; PD, AD, and AV rows with 26-28, 13-14, and 19-20 macrosetae, respectively. Metatarsomere I with outer setal row indistinct, apex with 5 platellae. Metatarsomere II apex with 4 platellae.

Distribution
French Guiana.
Head and tHorax. External morphological characters as in generic description.

Type species
Fulana brasiliensis gen. et sp. nov., by present designation and monotypy.

Diagnosis
Medium-sized leafhoppers ( Fig. 6C-F). Head in dorsal view ( Fig. 2A) short, median length of crown about 2.5 times smaller than interocular width; crown transversely striated; ocellus large, closer to anterior margin of crown; in lateral view (

Etymology
The generic name is feminine. Fulana, together with Beltrana and Sicrana, forms a very popular expression in Brazil ("Fulana, Sicrana e Beltrana"), which refers to unspecified people or people whose actual names should not be mentioned.

Description
Head and tHorax. Head in dorsal view ( Fig. 2A) short; median length of crown about 2.5 times smaller than interocular width; crown with anterior margin slightly sinuous; surface adjacent to ocelli slightly concave between ocellus and eye; texture transversely striated; transocular width of head six-sevenths of humeral width of pronotum; ocellus large, close to anterior margin of crown and slightly closer to eye than to median line; coronal suture distinct, extending to near anterior margin of crown. Head in ventral view (Figs 2B, 3A) with face wider than high; frontogenal suture distant from eye margin by basal width of clypeus and surpassing antennal ledge, extending to anterior margin of crown; oriented obliquely downward in relation to frons; frons approximately as wide as long, lateral margins strongly convergent ventrally; epistomal suture distinct; clypeus approximately 1.2 times as long as maximum width, lateral margins divergent apically, apex slightly emarginated; maxillary plate produced ventrally as far as clypeus apex; gena with ventrolateral margin straight on mid-length and slightly excavated below eye margin. Head in lateral view ( 2D) translucent, covered by numerous small brown spots; long and narrow, approximately 3.3 times as long as wide; venation distinct; appendix very narrow and bordering first to second apical cells. Profemur with AD, AM, and PD rows reduced and poorly defined, with exception of apical setae AD 1 , AM 1 , and PD 1 , respectively; AV and PV rows formed by 2-3 setae; IC row formed by slightly arched comb of fine setae, beginning at distal half of femur and extending to AM 1 . Protibia in cross-section semi-circular; AV row formed by long setae, slightly longer and thicker towards apex; AD row without differentiated setae; PD row with 7-8 setae; PV row with 6-7 setae. Metafemur with setal formula 2:2:1. Metatibia PD, AD, and AV rows with 26-31, 13-14, and 16-19 macrosetae, respectively; AD row with 1-4 intercalary setae between macrosetae; PV row with setae of apical half formed by sequence of a thicker and 2-4 thinner setae. Metatarsomere I with two rows of setae, inner row consisting of larger and more robust setae than those of outer row; apex with 6 platellae; metatarsomere II apex with 2-3 platellae. FeMale terMinalia. Second valvula of ovipositor ( Fig. 3G) elongated, slightly higher along apical fourth; apical portion ( Fig. 3H) with dorsal margin bearing minute and uniform denticles.

Remarks
An unpublished study based on 182 morphological characters combined with sequence data (28S and 16S rDNA and cytochrome oxidase subunit I) recovered Fulana gen. nov. as the sister group to all other gyponines except Chilenana DeLong & Freytag, 1967(Gonçalves 2016. In fact, the genera Fulana European Journal of Taxonomy 750: 70-93 (2021) gen. nov. and Chilenana share plesiomorphic characteristics with other tribes of Iassinae, which are absent in other Gyponini genera: (1) male pygofer with lateral lobe partly separated from base by flexible membranous cleft; and (2) presence of basiventral inner process on the male pygofer.
The new genus resembles Alapona DeLong, 1980, Ponana Ball, 1920, and Dumorpha DeLong & Freytag, 1975 because of its slightly anteriorly produced and transversely striated crown; ocellus closest to the anterior margin of the crown; forewing with dark spots; and metatibia with intercalary setae between macrosetae of the AD row. However, Fulana gen. nov. differs from Alapona by the lack of flaps in the aedeagus (Fig. 2K-L), from Ponana and Alapona by its thinner crown-face transition (Fig. 2C), and from Ponana and Dumorpha by its aedeagus without apodemal processes (Fig. 2K).

Etymology
The species name is in reference to the country of origin of the studied specimens.     Coloration. Pale yellow to yellow (Fig. 6C-F). Crown ( Fig. 2A) pale yellow, with circular black spot behind each ocellus, adjacent to posterior margin; anterior margin with three dark maculae, one medially and two in front of ocelli. Face (Figs 2B, 3A) pale yellow to yellow; frons with about ten pairs of dark bands over muscular impressions, dorsal portion with small dark punctuations; epistomal and frontogenal sutures black; lorum with dark subtriangular macula adjacent to lateral margin of clypeus; clypeus with central dark macula and apical margin black; gena with three black maculae that may be interconnected, forming large spot in some specimens: one adjacent to antennal pit, one adjacent to inner corner of compound eye, and one adjacent to lorum base. Pronotum ( Fig. 2A) with small dark punctures except near anterior margin, behind eyes. Mesonotum ( Fig. 2A) with ringed black maculae near lateral corners; in lateral view (Fig. 2C), lateral lobe of pronotum with large black spot, anepisternum with small black spot. Forewing (Fig. 2D) pale yellow, translucent, with yellow-cream areas on basal twothirds; numerous small black spots throughout surface and several larger black spots on apices of anal veins, along costal margin, over crossveins, and apical margin. Legs (Fig. 6D, F) pale yellow with scattered dark maculae; tarsus of pro-and mesothoracic legs black.
Head and tHorax. External morphological characters as in generic description.
Male terMinalia. Sternite VIII (Fig. 2E) strongly produced posteriorly, about 1.6 times as long as wide; lateral margins converging towards apex, posterior margin broadly rounded. Valve subrectangular, three times as wide as long; posterior margin shallowly excavated medially. Pygofer in lateral view (Fig. 2F) elongate, about 2.1 times as long as maximum height; apex rounded; macrosetae distributed only on apical half; basiventral inner process (Fig. 2F) strongly sclerotized, long and thin, almost reaching pygofer apex and contiguous to ventral margin, apical fourth curved dorsally; in ventral view (Fig. 2G), process broadened preapically, with short preapical tooth on external side, apex acute. Subgenital plate European Journal of Taxonomy 750: 70-93 (2021) in lateral view (Fig. 2F) not produced posteriorly as far as pygofer apex; in ventral view (Fig. 2H) long and narrow, about 6.4 times longer than wide; lateral margins parallel; apex abruptly rounded. Connective (Fig. 2I) approximately one-fourth of length of style; stalk longer than wide; arms short, slightly wider than stalk; dorsal keel developed. Style in dorsal view (Fig. 2I) with outer lobe developed and rounded; in lateral view (Fig. 2J), broadened on apical two-thirds and tapering towards apex; ventral margin with few denticles at median third; apex acute and curved dorsally. Aedeagus (Fig. 2K-L) with preatrium long; dorsal apodeme projected dorsally; shaft elongated and tubular, curved posterodorsally, expanded apically; apex bilobed, with pair of small apical spines.

Remarks
Male specimens collected from different localities show a variation in the shape of the apical portion of the ventral process of pygofer, which may be more curved or straighter, more dilated or slightly thinner. In addition, some specimens have two conspicuous and rounded black spots over crossveins of discal cells of forewing. However, remaining structures of the male terminalia and shape of the female sternite VII are invariable and we thus consider these variations as intraspecific.
Specimens of Fulana brasiliensis gen. et sp. nov. were mainly collected in areas of Atlantic Forest, at altitudes that vary from 50 to 1960 m above sea level. A putative second species of the new genus, known only from female specimens from the Brazilian Amazon Rain Forest, State of Amazonas, Manaus, was studied but will not be described until male specimens become available. This undescribed species is similar to F. brasiliensis gen. et sp. nov. in all aspects of the external morphology and female terminalia described herein. However, it can be distinguished by its larger size (15.5-15.8 mm); head in ventral view with fewer black maculae on face (lorum with small spot adjacent to lateral margin of clypeus, frons with punctuations and a semi-circular macula near anterior margin of crown, and antennal pits black); and sternite VII more deeply excavated medially.

Type species
Sicrana plana gen. et sp. nov., by present designation and monotypy.

Diagnosis
Large-sized, flattened leafhoppers ( Fig. 6G-J). Head in dorsal view (Fig. 4A) with rugose surface, strongly produced anteriorly, anterior margin parabolic; ocellus closer to median line than to adjacent eye and closer to posterior than to anterior margin of crown; in lateral view (Fig. 4C) 4D) with appendix absent. Aedeagus (Fig. 4J-K) with pair of apodemal processes. Second valvula of ovipositor with apical portion (Fig. 5G-H) with large preapical tooth.

Etymology
The generic name is feminine. Sicrana, together with Beltrana and Fulana, forms a very popular expression in Brazil ("Fulana, Sicrana e Beltrana'), which refers to unspecified people or people whose actual names should not be mentioned.

Description
Head and tHorax. Head in dorsal view (Fig. 4A) moderately produced anteriorly, median length of crown almost as long as interocular width; transocular width three-fourths of humeral width of pronotum; crown with anterior margin parabolic; surface flat; texture rugose; ocellus medium-sized, closer to median line than to adjacent eye and closer to posterior than to anterior margin of crown; coronal suture distinct along basal two-thirds of crown. Head in lateral view ( Fig. 4C) with crown-face transition distinct and strongly foliaceous, with two very close carinae; frons tumid. Head in ventral view (Figs 4B, 5A) approximately as wide as high; frontogenal suture strongly sinuous, reaching antennal ledge and distant from eye margin by twice maximum width of clypeus; antennal ledge carinated, strongly arched, adjacent to anterior margin of crown and not extending over frons; frons narrow, approximately two times longer than wide; epistomal suture indistinct; clypeus approximately 1.4 times as long as maximum width, lateral margins parallel, apex slightly emarginated; maxillary plate very narrow, not reaching clypeus apex; gena with ventrolateral margin slightly excavated, texture with several oblique striations parallel to ventrolateral margin. Pronotum (Fig. 4A) rugose, except disc and posterior third with transverse parallel striae; anterior margin almost straight; lateral margins convergent anterad, about two times as long as eye length, rounded, carinated and foliaceous, expanded laterally; posterior margin slightly excavated; in lateral view (Fig. 4C), pronotal surface declivous; head and pronotum in continuous slope. Mesonotum (Fig. 4A) as long as wide; scutellum (Fig. 4C) flat. Forewing (Fig. 4D) long and narrow, approximately 3.2 times as long as wide; venation with some additional crossveins located mainly apically; appendix absent; apex subacute. Profemur with AD, AM, and PD rows reduced and poorly defined, with exception of apical setae AD 1 , AM 1 , and PD 1 , respectively; AV and PV rows formed by 5-6 very short and thin setae; IC row formed by slightly arched comb of fine setae, beginning at distal half of femur and extending to AM 1 . Protibia in cross-section semi-circular; AV row formed by short setae, slightly longer and thicker towards apex, setae of apical portion shorter than diameter of tibia; AD row without differentiated setae; PD row with three small setae and undifferentiated intercalary setae; PV row with 4-5 small setae on apical half and undifferentiated intercalary setae. Metafemur with setal formula 2:2:1. Metatibia PD, AD, and AV rows with 24-25, 12, and 16 macrosetae, respectively; metatibia AD row without intercalary setae between macrosetae. Metatarsomere I with two rows of cucullate setae, inner row formed by 5-7 setae, outer row reduced, with 0-2 median setae, apex with 5 platellae. Metatarsomere II apex with 2-3 apical platellae.

Etymology
The species name refers to its flattened body (Fig. 6H, J).

GONÇALVES C.C. et al., Three new genera of Gyponini from South America
Male terMinalia. Sternite VIII (Fig. 4E) subrectangular, about 1.3 times as wide as long; lateral margins parallel, posterior margin slightly rounded. Valve subrectangular, 2.3 times as wide as long; posterior margin convex. Pygofer in lateral view (Fig. 4F) (2021) times as long as maximum height, without processes; dorsal margin strongly excavated; less than 10 macrosetae forming a longitudinal row, near dorsal margin of apical half; apex tapered and rounded. Subgenital plate, in lateral view (Fig. 4F), produced posteriorly as far as pygofer apex; in ventral view (Fig. 4G), ligulate, about 3.6 times as long as wide; ventral surface covered by small setae; dorsal surface with tuft of moderately long filiform setae near apex; lateral margins divergent towards apex; apex abruptly tapered and rounded. Connective (Fig. 4H) with anterior portion membranous, posterior portion sclerotized; approximately one-seventh length of style; arms narrow, strongly divergent. Style in dorsal view (Fig. 4H) with outer lobe poorly developed; in lateral view (Fig. 4I), with margins parallel; ventral margin smooth; apex acute, more sclerotized and curved outwards. Aedeagus (Fig. 4J-K) with preatrium absent; atrium expanded posteriorly; dorsal apodeme well developed; apodemal processes claviform, extending to apex of shaft, subapical region of each process rounded and with preapical excavation dorsally; shaft elongated and tubular, curved dorsally; apex with pair of processes parallel to shaft, extending anteroventrally, half-length of shaft, each process with inner margin crenulate and apex acute.
The monotypic genus Freytagana was erected by DeLong (1975) to accomodate the type species, Freytagana gibsoni DeLong, 1975, from Mexico. According to that author, Freytagana differs from other gyponines by the following set of characteristics: (1) crown short and broad, with anterior and posterior margins almost parallel; (2) ocellus closer to median line than to adjacent eye; (3) transition crown-face distinct and thick; and (4) forewing long and narrow, with venation reticulated, and slight appendix.
Based on the original descriptions of the genera and a detailed study of the holotypes of the two included species, we conclude that there are no characters that distinguish Freytagana from Marganana. Therefore, we suggest that Freytagana be treated as a junior synonym of Marganana. In addition, considering the distribution data (the holotype of F. gibsoni and several paratypes of M. mexicana are from the same locality -Mexico City), forewing venation, and male terminalia ( Fig. 7A-H, I-P), we concluded that holotypes of F. gibsoni and M. mexicana are conspecific, so that the former should be treated as a junior synonym of the latter. We note that the holotype of F. gibsoni has its aedeagus shaft damaged and twisted, besides being strongly bent ventrally at apical half. This may be the reason why DeLong did not treat them as the same species.

Remarks
The monotypic genus Chilella was erected by DeLong & Freytag (1967) based on a single female specimen of Chilella rugella DeLong & Freytag, 1967 from Chile. According to the authors, Chilella differs from other genera of Gyponini by the following set of characteristics: (1) crown short and broad; (2) transition crown-face rounded, without defined margin; (3) ocelli distant from eyes, located on anterior portion of crown; (4) forewing rugose, corrugated and without an appendix; and (5) foreleg with metatibia intercalary setae between macrosetae.
Our study of the holotype of Chilella rugella (Fig. 8A-E) indicated that the position of the ocelli, which are located in the crown-face transition, is incongruent with the diagnoses proposed for Gyponini by DeLong (1942), Oman (1949), Linnavuori (1959), Dietrich (2005), and Krishnankutty et al. (2016). According to those authors, the ocelli in Gyponini are located on the crown, away from the anterior margin. Moreover, according to the key to the tribes of Iassinae proposed by Krishnankutty et al. (2016), Chilella rugella is classified as Selenomorphini Evans, 1974, based on the following characteristics: GONÇALVES C.C. et al., Three new genera of Gyponini from South America (1) head with crown poorly delimited, rounded to face; (2) hind wing veins R4+5 and M1+2 separated distally, not confluent at apex; and (3) body pigmentation primarily pale green or stramineous.
However, morphological characteristics of C. rugella are not congruent with any genera currently included in Selenomorphini. Chilella differs from Selenomorphus Evans, 1974 andLinnavuoria Dai &Dietrich, 2015 in having the crown-face transition indistinct, whereas in the last two genera the crownface transition is distinctly angulated in profile. Also, Chilella differs from Pachyopsis Uhler, 1877 and Uhleriana Domahovski, 2019 in having the metatibiae with intercalary setae. In addition, it differs from Parapachyopsis Domahovski, 2019 in having the head narrower than the pronotum and from Scaroidana Osborn, 1938 by its smaller size (9.0 mm) and position of the ocelli mesad to the antennal pits (Fig. 8C), whereas in Scaroidana they are closer to the eyes and located above the antennal pits, in ventral view.
Unfortunately, the holotype of C. rugella lost its forewings, which could have provided useful information about its relationships with other genera of Selenomorphini. However, according to DeLong & Freytag (1967), the forewings are rugose and corrugated, two features not found in any other genus of the tribe. We propose the transfer of Chilella to Selenomorphini, maintaining it as a valid genus. However, the discovery of male specimens as well as phylogenetic studies on the tribe are necessary to confirm our proposal.

Discussion
In this study, three new species of Gyponini from South America were described and illustrated. These species have morphological characteristics that are clearly not in accordance with the diagnoses of previously established genera of the tribe. Therefore, three new genera were erected to allocate them. Holotypes of Marganana (Marganana) mexicana DeLong &Freytag, 1963 andFreytagana gibsoni DeLong, 1975 were studied and we concluded that they are conspecific. Therefore, F. gibsoni is regarded as a junior synonym of M. mexicana, and the monotypic genus Freytagana DeLong, 1975 is regarded as a junior synonym of Marganana DeLong, 1948. In addition, our study of the holotype of Chilella rugella DeLong & Freytag, 1967, the only species so far described for the genus, showed that the position of the ocelli, which are located at the crown-face transition, is incongruent with the condition observed in the Gyponini; in the latter group, the ocelli are consistently located on the crown. This feature, added to the shape of the crown, the venation of the hind wings, and body color, supports the transfer of Chilella to Selenomorphini. With the new genera described here and the proposed taxonomic changes, the Gyponini now comprise 72 genera and 1406 valid species. Many species from the Neotropical region remain undescribed, especially from poorly known and threatened biomes such as the Amazon Forest.
Of the 72 genera currently recognized in the Gyponini, 20 are monotypic. Approximately 70% of the species of the tribe are included in only six genera, viz., Gypona Germar, 1821, Curtara DeLong & Freytag, 1972, Polana DeLong, 1942, Ponana Ball, 1920, Gyponana Ball, 1920, and Darma Walker, 1858. The present work contributes to our knowledge of the morphological diversity within this interesting leafhopper tribe.