Eight new species of Costanana DeLong & Freytag (Hemiptera: Cicadellidae: Gyponini): taxonomic changes, key to males, and description of Metacostana gen. nov

Costanana DeLong & Freytag, 1972 is a Neotropical genus of leafhoppers currently comprising 13 species with a wide distribution from Mexico to Argentina. In this study, eight new species of Costanana are described, C. xenomorpha sp. nov. from Department of Cuzco, Peru, and seven Brazilian species: C. alata sp. nov. from Minas Gerais State; C. bifida sp. nov. from Paraná and Mato Grosso states; C. cifi sp. nov., C. obtusa sp. nov. and C. piraquarensis from Paraná State; C. luzi sp. nov. from Distrito Federal; and C. rubromarginata sp. nov. from Mato Grosso and Pará states. We propose that Costanana cella DeLong & Freytag, 1972 is a junior synonym of C. praecellens (Stål, 1862) and C. asymmetrica DeLong & Freytag, 1972 is a junior synonym of Gypona viridans DeLong & Martinson, 1972. Costanana costata DeLong & Freytag, 1972 and C. flavicosta (Stål, 1862) are transferred to Gypona Germar, 1821, C. minuta (Spångberg, 1878) is transferred to Acuponana DeLong & Freytag, 1970 and Gypona nupera Van Duzee, 1907 is transferred to Costanana. Gypona costata comb. nov. and Costanana santana DeLong & Wolda, 1983 are newly recorded from Pernambuco State, and Gypona flavicosta comb. nov. is recorded from Espírito Santo and Paraná states, Brazil. The female genitalia of Costanana flavina DeLong & Freytag, 1972, C. praecellens and Gypona flavicosta comb. nov. are described and illustrated for the first time and a brief note on feeding behavior is given for C. flavina and C. praecellens. A key to males and a new diagnosis are provided for Costanana. We also describe and illustrate Metacostana cornuta gen. et sp. nov., a new Brazilian genus for Gyponini sharing characteristics with Costanana, Acuthana Domahovski & Cavichioli, 2018, Domahovana Silva et al., 2022 and Dumorpha DeLong & Freytag, 1975.

Coloration. Dorsal portion of head and thorax reddish-brown (Figs 1A, C, 11A); ventral portion yellow 11B). Crown (Fig. 1A) with black maculae surrounding ocelli; pair of rounded black spots behind ocelli, near posterior margin; median line and posterior margin light brown; eyes (in life) and ocelli red. Pronotum (Fig. 1A) with black irregular maculae and light brown spots near anterior and lateral margins; lateral margin yellow. Mesonotum with pair of dark brown maculae near lateral angles and medially; scutellum yellowish, with brown areas adjacent to scutoscutellar suture. Face (Fig. 1B) with frons slightly brownish dorsally. Forewing (Fig. 1D) with narrow light yellow macula on anterior third of costal margin and a depigmented (white) macula on costal margin at level of outer anteapical cell, and apex of third and fourth apical cells; veins outlined by dark brown. Legs (Fig. 11B) yellow, apex of metatibia dark brown.
StruCture. Head in dorsal view (Fig. 1A): median length of crown as long as half interocular width; transocular width of head eight tenths humeral width of pronotum; in lateral view (Fig. 1C), with anterior margin moderately thick, with 5-6 well developed carinae. Forewing (Fig. 1D) with inner discal cell very short, one-third as long as the outer discal cell; apex slightly tapered. Profemur AV row with 3-4 setae restricted to basal half and PV row with 2 setae restricted to distal half. Protibia PD row with 3 long setae and apical PD 1 seta developed; PV row developed, with 8 setae increasing in thickness and length towards apex. Metatibia rows PD, AD, and AV with 21, 12, and 12 macrosetae, respectively. Metatarsomere I apex with 3 platellae. Metatarsomere II apex with 2 apical platellae. Other characteristics as in the generic description.
Male terMinalia. Sternite VIII (Fig. 1E) 1.2× as wide as long; lateral margins excavated at base, forming a triangular projection and then converging towards apex, narrow and rounded. Valve (Fig. 1F) 3.1× as wide as long; posterior margin almost straight. Pygofer (Fig. 1G), in lateral view, 2.3× as long as maximum height; ventral margin with a small protrusion near base; posterodorsal and ventral margins rounded; external surface near ventral margin with short filiform setae; macrosetae distributed on apical half; apex slightly tapered and rounded. Subgenital plate (Fig. 1G), in lateral view, short, surpassing half length of pygofer; in ventral view (Fig. 1H), 5× as long as wide; inner margin straight; external margin rounded on basal third; ventral surface and external margin with long filiform setae; apex strongly tapered and weakly sclerotized. Connective (Fig. 1I) Y-shaped; arms weakly produced laterally; stalk about as long as maximum width of arms; median keel weakly developed. Style (Fig. 1J), in dorsal view, with outer lobe small and rounded; inner margin with a small dentiform process near midlength of blade; in lateral view (Fig. 1K), ventral margin not serrated, with dentiform process near base of blade; apex truncated, forming a pair of acute processes directed dorsally and ventrally.  preatrium not developed; dorsal apodeme strongly elongated laterally; shaft short, nearly tubular, with pair of ventral wing-shaped processes near base, curved dorsally with divergent and acute apices; dorsal margin of shaft with slender basodorsal process; ventral margin with pair of minute spurs subapically.

Female
Unknown.

Remarks
Costanana alata sp. nov. is most similar to C. flavina in having the same color pattern (Figs 11A-B. 13A-B) and the aedeagal shaft short with a pair of ventral processes near the base, not contiguous to shaft and following its curvature. The new species can easily be separated from C. flavina by having the apex of the style truncated (Fig. 1K) and the longer and winged ventral processes of the aedeagal shaft ( Fig. 1L-M).

Diagnosis
Coloration (Fig. 11C) of head and pronotum yellowish-brown with small black maculae, forewing costal margin with broad light yellow macula on anterior third and large depigmented (white) macula on apical third. Style (Fig. 2K) without dentiform ventral process near base of blade; apical portion bifid, with slender process at apical third, directed posterad and adjacent to dorsal apex.  with pair of lateral processes at base, strongly curved dorsally; shaft short, strongly compressed laterally, without apical processes.

Etymology
The species epithet 'bifida' refers to the branched blade of the style.
Coloration. Dorsal portion of head and thorax yellowish-brown (Figs 2A,C,11C); ventral portion yellow (Figs 2B-C, 11D). Crown ( Fig. 2A) with transverse narrow black stripe interrupted medially over median line; pair of rounded black spots behind ocelli, near posterior margin; eyes (in life) and ocelli red. Pronotum ( Fig. 2A) with small black irregular maculae near anterior and lateral margins. Mesonotum ( Fig. 2A) with pair of rounded black spots medially and pair of elongated black macula submedially (paratypes). Face (Fig. 2B) without maculae. Forewing (Fig. 2D) with large light yellow macula on anterior half of costal margin and a large rounded depigmented (white) macula at level of outer anteapical cell, remaining portion of costal cell, apex of apical cells, apex of clavus, apex of anal veins and appendix dark brown. Legs (Fig. 11D) yellow, without black markings.
StruCture. Head in dorsal view ( Fig. 2A): median length of crown slightly shorter than interocular width; transocular width of head 8.5 tenths humeral width of pronotum; in lateral view (Fig. 2C), with anterior margin thick, with 6-7 carinae. Forewing (Fig. 2D) inner discal cell open, m-cu 2 crossvein absent; apex rounded. Profemur AV row with 3-4 setae restricted to basal half and PV row with 1 apical seta. Protibia PD row with 3 long setae and apical PD 1 seta developed; PV row developed, with 8 setae increasing in thickness and length towards apex. Metatibia rows PD, AD, and AV with 20-21, 12, and 12-13 macrosetae, respectively. Metatarsomere I apex with 3 platellae. Metatarsomere II apex with 2 platellae. Other characteristics as in the generic description.   Male terMinalia. Sternite VIII (Fig. 2E) 1.1× as wide as long; lateral margins excavated at base, forming a triangular projection and then converging towards apex, narrow and rounded. Valve (Fig. 2F) 2.5× as wide as long; posterior margin slightly rounded. Pygofer (Fig. 2G), in lateral view, 2.2× as long as maximum height; ventral margin straight; posterodorsal margin slightly rounded; posteroventral margin straight; external surface near ventral margin with short filiform setae; macrosetae distributed on apical half; apex truncated. Subgenital plate (Fig. 2G), in lateral view, short, surpassing half length of pygofer; in ventral view (Fig. 2H), 3.7× as long as wide; inner margin approximately straight; external margin rounded on basal half; ventral surface and external margin with long filiform setae; apex strongly tapered and weakly sclerotized. Connective (Fig. 2I) Y-shaped; arms weakly produced laterally; stalk about twice as long as maximum width of arms; median keel weakly developed. Style (Fig. 2J), in dorsal view, with outer lobe small and rounded; in lateral view (Fig. 2K), blade evenly curved dorsally; ventral margin not serrated, with a slender process at apical third, directed posterad and adjacent to dorsal apex; apex broad and rounded, forming small tip directed anterad. Aedeagus ( Fig. 2L-M) preatrium not developed; dorsal apodeme produced laterally and broadly rounded; pair of lateral processes at base, strongly curved dorsally, wide at base and slender distally, apex acute; shaft short, wide, strongly compressed laterally; apical portion enlarged in posterior view and without processes.

Remarks
Costanana bifida sp. nov. is similar to C. helvacosta, C. nana, C. santana and C. rubromarginata sp. nov. in having the aedeagus with a pair of processes arising near the base of the shaft and the apical portion of the shaft without processes ( Fig. 2L-M). However, the new species is easily separated from these former species by having the apex of the style branched (Fig. K) and the aedeagus with the shaft short, wide and strongly compressed laterally ( Fig. 2L-M). The crown-face transition being weakly defined (Fig. 2C) is similar to in species of Polana DeLong, 1942; however, the morphology of the female ovipositor (Fig. 3D, F)
Male terMinalia. Sternite VIII (Fig. 4E) 1.2× as wide as long; lateral margins excavated at base, forming a triangular projection; apex broadly rounded. Valve (Fig. 4F) 2× as wide as long; posterior margin rounded, slightly emarginated medially. Pygofer (Fig. 4G), in lateral view, 2.2× as long as maximum height; ventral margin slightly rounded; posterodorsal margin rounded; posteroventral margin straight; external surface near ventral margin with short filiform setae; macrosetae distributed on apical half; apex subtruncated. Subgenital plate (Fig. 4G), in lateral view, short, slightly surpassing half length of pygofer; in ventral view (Fig. 4H), 4.5× as long as wide; inner margin straight; external margin broadly rounded; ventral surface and external margin with long filiform setae; apex tapered and weakly sclerotized. Connective (Fig. 4I) Y-shaped; arms weakly produced laterally; stalk narrow, about as long as maximum width of arms; median keel weakly developed. Style (Fig. 4J), in dorsal view, with outer lobe reduced; in lateral view (Fig. 4K), blade strongly elongated, almost straight; ventral margin not serrated, with short slender process at apical third, directed posterad, weakly curved dorsally and overlapping dorsal apex; apex tapered and curved dorsally. Aedeagus ( Fig. 4L-M) preatrium developed; dorsal apodeme strongly produced, with divergent arms; shaft elongated curved dorsally; dorsal margin with subapical spiniform  process; pair of long subapical processes almost as long as shaft length, directed dorsally and curved anterad, with apex tapered and sharply acute.

Remarks
Costanana cifi sp. nov. is similar in coloration to C. nupera comb. nov. in having the costal margin of the forewing with a small and slender yellow macula on the anterior fifth followed by a sequence of small yellowish spots and a larger white macula at the apex of the outer apical cell (Figs 4D,20A). Costanana cifi can be separated from the other species of Costanana by the unusual shape of the aedeagus, with a short subapical spiniform process on the dorsal margin and a pair of long subapical processes directed dorsally and curved anterad (Fig. 4L).  11G) of head and pronotum reddish-brown without black maculae, forewing costal margin with narrow light yellow macula on anterior third, depigmented (white) macula on apical third absent. Style ( Fig. 6K) with dentiform process near base of blade; dorsal margin strongly produced medially, forming a triangular process. Aedeagus ( Fig. 6L-M) without processes near base; shaft compressed, with pair of long subapical processes directed basally, branched near base, with ventral ramus half as long as dorsal ramus; apex rounded with pair of short spiniform processes directed dorsally.

Etymology
The new species name is in honor to Prof. Dr José Roberto Pujol Luz of the Universidade de Brasília who collected the holotype.

Male
MeaSureMentS. Holotype male: total length 7.1 mm. StruCture. Head in dorsal view ( Fig. 6A): median length of crown as long as half interocular width; transocular width of head 7.5 tenths of humeral width of pronotum; in lateral view (Fig. 6C), with anterior margin moderately thin, with 5-6 well developed carinae. Forewing ( Fig. 6D) with inner discal cell very short, one-third as long as outer discal cell; apex slightly tapered. Profemur AV row with 4-5 setae restricted to basal half and PV row with 1 apical seta. Protibia PD row with 3 long setae and apical PD 1 seta developed; PV row developed, with 8 setae increasing in thickness and length towards apex. Metatibia rows PD, AD, and AV with 18-19, 13, and 12 macrosetae, respectively. Metatarsomere I apex with 3 platellae. Metatarsomere II apex with 2 apical platellae. Other characteristics as in the generic description.
Male terMinalia. Sternite VIII ( Fig. 6E) 1.3× as wide as long; lateral margins excavated at base, forming a triangular projection; apex broadly rounded. Valve (Fig. 6F) 3.1× as wide as long; posterior margin almost straight. Pygofer (Fig. 6G), in lateral view, 1.7× as long as maximum height; ventral margin with rounded protrusion near midlength; posterodorsal and ventral margins rounded; macrosetae distributed on dorsoapical quadrant; apex slightly tapered and rounded. Subgenital plate (Fig. 6G), in lateral view, surpassing half length of pygofer; in ventral view (Fig. 6H), 3.5× as long as wide; inner margin straight; external margin rounded on basal half; external margin with long filiform setae; apex strongly tapered and weakly sclerotized. Connective (Fig. 6I) Y-shaped; arms produced laterally; stalk shorter than maximum width of arms, widened apically; median keel weakly developed. Style (Fig. 6J), in dorsal view, with outer lobe reduced and rounded; apical portion directed outward; in lateral view ( Fig. 6K), ventral margin not serrated, with dentiform process near base of blade; dorsal margin strongly produced on basal half, forming a triangular process; apical portion slender, curved dorsally; apex acute. Aedeagus ( Fig. 6L-M) preatrium not developed; dorsal apodeme strongly elongated laterally, forming divergent and truncated apices; shaft compressed, curved dorsally; pair of long subapical processes directed basally, branched near base, ventral ramus half as long as dorsal ramus; apex rounded with pair of short spiniform processes directed dorsally.

Female
Unknown.

Remarks
Costanana luzi sp. nov. is most similar to C. praecellens in having the same color pattern (Figs 11G-H, 13C-D) and the aedeagal shaft compressed and bearing a pair of subapical processes branched near the base (Fig. 6L), and the apex of the aedeagus rounded with pair of short spiniform processes directed dorsally (Fig. 6M). The new species can easily be separated from C. praecellens in having the style with a process on the dorsal margin of the blade and the distal portion longer and distinctly more curved (Fig. 6K), and the branched subapical process of the aedeagus with the ventral ramus half as long as the dorsal ramus (

Etymology
The species epithet 'obtusa' refers to the unusual apex of the style being truncated.

Description Male
MeaSureMentS. Holotype male: total length 6.8 mm.
Coloration. Dorsal portion of head and thorax brown ( (Fig. 7G), in lateral view, 1.8× as long as maximum height; basodorsal process present but weakly developed; ventral margin with rounded protrusion near midlength; posterodorsal and ventral margins rounded; external surface near ventral margin with short filiform setae; macrosetae distributed on posterior half; apex broadly rounded. Subgenital plate ( Fig. 7G), in lateral view, slightly surpassing half length of pygofer; in ventral view ( Fig. 7H), 3.5× as long as wide; inner margin straight; external margin slightly rounded on basal half; ventral surface and external margin with long filiform setae; apex tapered and weakly sclerotized. Connective (Fig. 7I) approximately linear; arms not produced laterally; stalk longer than maximum width of arms, widened apically; median keel weakly developed. Style (Fig. 7J), in dorsal view, with outer lobe reduced and subtruncated; in lateral view (Fig. 7K), blade broadened medially; ventral margin not serrated, with dentiform process near base of blade; apex truncated. Aedeagus ( Fig. 7L-M) preatrium not developed; dorsal apodeme strongly elongated laterally, forming divergent and truncated apices; shaft slender, curved dorsally; pair of long filiform subapical processes directed basally, with one or two spurs near base; apex spatulate and rounded.

Female
Unknown.

Remarks
Costanana obtusa sp. nov. is most similar to C. piraquarensis sp. nov. in having a similar color pattern ( Fig.12A-D), especially the pair of large maculae on the costal margin of the forewing (Figs 7D, 8D)  (5) subapical pair of processes having one or two spurs near the base (Fig. 7M). Unfortunately the legs of the holotype are missing but, due to the distinctly different genitalia and the lack of importance for species delimitation of the morphology of the legs, we suppose this does not prejudge the description or future recognition of this species.

Etymology
The new species name refers to the municipality of Piraquara, where the holotype was collected. StruCture. Head in dorsal view (Fig. 8A): median length of crown slightly shorter than interocular width; transocular width of head eight tenths humeral width of pronotum; in lateral view (Fig. 8C), with anterior margin thick, with 5-6 well developed carinae. Forewing (Fig. 8D) inner discal cell open, m-cu 2 crossvein absent; apex broadly rounded. Profemur AV row with 3-4 setae restricted to basal half and PV row with 1 apical seta. Protibia PD row with 4 long setae and apical PD 1 seta developed; PV row developed, with 8 setae increasing in thickness and length towards apex. Metatibia rows PD, AD, and AV with 24-25, 14, and 15 macrosetae, respectively. Metatarsomere I apex with 4 platellae. Metatarsomere II apex with 2 apical platellae. Other characteristics as in the generic description. Male terMinalia. Sternite VIII (Fig. 8E) 1.1× as wide as long; lateral margins excavated at base, forming a triangular projection and then converging to rounded apex. Valve (Fig. 8F) 2.7× as wide as long; posterior margin almost straight. Pygofer (Fig. 8G), in lateral view, 1.8× as long as maximum height; basodorsal process present but weakly developed; ventral margin with rounded protrusion near basal third; posterodorsal and ventral margins straight; external surface near ventral margin with short filiform setae; macrosetae distributed on posterodorsal quadrant; apex broad and truncate. Subgenital plate (Fig. 8G), in lateral view, slightly surpassing half length of pygofer; in ventral view (Fig. 8H), 3.2× as long as wide; inner margin straight; external margin slightly rounded; ventral surface and external margin with long filiform setae; apex tapered and weakly sclerotized. Connective (Fig. 8I) linear; arms not produced laterally; stalk very broad, slightly longer than maximum width of arms; median keel weakly developed. Style (Fig. 8J), in dorsal view, with outer lobe reduced and rounded; in lateral view (Fig. 8K), blade broadened medially; ventral margin not serrated, with dentiform process near base of blade; apex abruptly tapered, slightly directed dorsally, acute. Aedeagus (Fig. 8L-M) preatrium not developed; dorsal apodeme elongated laterally, forming divergent and truncated apices; shaft slender, curved dorsally; pair of long subapical filiform processes directed basally, almost as long as shaft length; apex compressed, subacute.

Female
Unknown.

Remarks
Costanana piraquarensis sp. nov. is separated from C. obtusa sp. nov. in having the following differences: (1) apex of pygofer truncated (

Etymology
The species epithet refers to the anterior margin of head being red.
StruCture. Head in dorsal view (Fig. 9A): median length of crown as long as half interocular width; transocular width of head 8.5 tenths humeral width of pronotum; in lateral view (Fig. 9C), with anterior margin moderately thick, with 5-6 well developed carinae. Forewing (Fig. 9D) with inner discal cell half long as outer discal cell; apex rounded. Profemur AV row with 4-5 setae restricted to basal half and PV row with 1 apical seta. Protibia PD row with 3 long setae and apical PD 1 seta developed; PV row developed, with 8 setae increasing in thickness and length towards apex. Metatibia rows PD, AD, and AV with 20-23, 12, and 13 macrosetae, respectively. Metatarsomere I apex with 3 platellae. Metatarsomere II apex with 2 apical platellae. Other characteristics as in the generic description.
Male terMinalia. Sternite VIII (Fig. 9E) 1.1× as wide as long; lateral margins excavated at base, forming a triangular projection and then converging to rounded apex. Valve (Fig. 9F) 1.6× as wide as long; posterior margin rounded. Pygofer (Fig. 9G), in lateral view, 2× as long as maximum height; ventral margin broadly rounded; posterodorsal margin approximately straight; external surface near ventral margin with short filiform setae; macrosetae distributed on posterodorsal quadrant; apex slightly tapered and almost truncate. Subgenital plate (Fig. 9G), in lateral view, surpassing half length of pygofer; in ventral view (Fig. 9H), 4× as long as wide; lateral margins slightly rounded; ventral surface and external margin with long filiform setae; apex tapered and weakly sclerotized. Connective (Fig. 9I) T-shaped; arms weakly produced laterally; stalk narrow, slightly shorter than maximum width or arms; median keel weakly developed. Style (Fig. 9J), in dorsal view, with outer lobe well developed, rounded; in lateral view (Fig. 9K), blade slender, evenly curved dorsally, slightly broadened subapically; ventral margin not serrated; apex tapered, curved dorsally and acute. Aedeagus (Fig. 9L-M) preatrium not developed; dorsal apodeme weakly produced laterally, dorsal margin straight; shaft long and slender, cylindrical, curved anterad; pair of long and slender processes arising from basal third, parallel to shaft and slightly surpassing apex, enlarged at midlength and thin and acute apically; shaft without apical or subapical processes.

Female
Unknown.

Remarks
Costanana rubromarginata sp. nov. is similar to C. helvacosta, C. nana and C. santana in having the aedeagus with the shaft long and slender, with the pair of processes arising near the base of the shaft and without processes distally (Fig. 9L-M). The shape of the apex of the style, slightly expanded subapically (Fig. 9K), is most similar to that of C. nana, but the new species is separated in having the pair of lateral processes abruptly expanded medially and not parallel to the shaft along the entire length as in C. nana.  (Fig. 12G) of head and pronotum brown with small black maculae, forewing costal margin with broad light yellow macula on anterior third and large depigmented (white) macula on apical third. Style (Fig. 10K) blade without processes near base or apical portion, ventral margin serrated; apex abruptly curved dorsally. Aedeagus (Fig. 10L-M) without processes near base; apical portion with four pairs of lateral processes and an unpaired subapical process on ventral surface.

Etymology
The new species name comes from the Greek 'xeno' ('strange', 'unusual') + 'morpha' ('form', 'shape'). It refers to the unusual shape of the male genital structures. It also alludes to the fictional extraterrestrial "xenomorphs" from the classic film "Alien", due to the aedeagus, in posterior view, resembling the creature named "face hugger". StruCture. Head in dorsal view (Fig. 10A): median length shorter than interocular width; transocular width of head eight tenths humeral width of pronotum; in lateral view (Fig. 10C), with anterior margin thick, with 7-8 carinae. Forewing (Fig. 10D) with inner discal cell not distinctly shorter than outer discal cell; apex rounded. Profemur AV row with 6 setae restricted to basal half and PV row with about 10 setae distributed from base to apex. Protibia PD row with 4 long setae and apical PD 1 seta developed; PV row developed, with 6-7 setae increasing in thickness and length towards apex. Metatibia rows PD, AD, and AV with 23, 13, and 15 macrosetae, respectively. Metatarsomere I apex with 3 platellae. Metatarsomere II apex with 2 apical platellae. Other characteristics as in the generic description.

Holotype
Male terMinalia. Sternite VIII (Fig. 10E) 1.3× as wide as long; lateral margins straight and parallel at basal fourth and rounded distally; posterolateral angles rounded; posterior margin almost straight, weakly produced medially. Valve (Fig. 10F) 2× as wide as long; posterior margin rounded, weakly emarginated medially. Pygofer (Fig. 10G), in lateral view, 2× as long as maximum height; basodorsal process present but weakly developed; ventral margin broadly rounded; posterodorsal margin excavated; external surface near ventral margin with a few short setae; macrosetae distributed on posterodorsal quadrant; apex produced and rounded, curved inwards, inner surface with short filiform setae and a small dentiform process. Anal tube (segment X) sclerotized, without processes. Subgenital plate (Fig. 10G), in lateral view, not surpassing apex of pygofer; in ventral view (Fig. 10H), 3.1× as long as wide, strongly broadened medially; lateral margins rounded; external margin with minute setae; apex rounded, well sclerotized. Connective (Fig. 10I) T-shaped; arms well produced laterally; stalk narrow, short, one-third as long as maximum width of arms; median keel weakly developed. Style (Fig. 10J), in dorsal view, with outer lobe well developed, rounded; in lateral view (Fig. 10K), blade slender, sinuous; ventral margin serrated; apex abruptly curved dorsally and acute. Aedeagus (Fig. 10L-M) preatrium weakly developed; dorsal apodeme produced dorsally but not laterally, dorsal margin straight; shaft long and slender, almost cylindrical, sigmoid; apical portion with four pairs of processes; two pairs of lateral processes adjacent to each other, directed laterally; long pair almost apical, thicker and longer than preceding ones, directed ventrolaterally; and a short apical claw-like pair directed dorsally; posterior surface with a unpaired subapical process curved ventrally.

Female
Unknown.

Remarks
The male genitalia of C. xenomorpha sp. nov. differ in several aspects from those of the other species of Costanana as in the larger size of body ( Fig. 12G-H); the shape of the sternite VIII, not tapered apically (Fig. 10E); the pygofer with apex curved inwards, bearing a spiniform process (Fig. 10G); the anal tube well sclerotized (Fig. 10G); the subgenital plate broadened subapically, well sclerotized distally, and lacking filiform setae (Fig. 10H); and the connective T-shaped with arms long (Fig. 10I). The shape of the subgenital plate and style is most similar to that of C. dunda. The new species can easily be recognized by the apex of the aedeagus with four pairs of lateral processes and a single subapical process on the ventral surface (Fig. 10M).

Behavior note
In a locality of the municipality of São José dos Pinhais (Paraná State, Brazil, 25°36′18″ S, 49°11′37″ W) C. praecellens and C. flavina are abundant species among the approximately 55 species of Gyponini found there. Both species are apparently polyphagous, feeding on shrubs and trees at a fragment of Araucaria moist forest. Both species were found presenting a feeding behavior in a curious upside down position, but we were able to record images only for a female specimen of C. praecellens (Fig. 17). The pro-and mesothoracic legs are extended, forming a cross, while the frons almost touches the leaf surface and the abdomen and metathoracic legs are directed almost vertically (Fig. 17A-B). In the figures 17B and C it is possible to observe the mouthparts inserted on the leaf parenchyma. We observed the specimen in this position for an interval of approximately 10 minutes in which the leafhopper also showed grooming behavior (Fig. 17C), but without releases of drops of excreta or brochosome fluid from the anus. We have never observed this type of behavior in species of the other Gyponini genera and perhaps this is characteristic of Costanana.

Remarks
Gypona praecellens Stål, 1862 was described based on a single female specimen labeled as "Brasil" (Fig. 15C-F). DeLong and Freytag (1972b) examined an additional female specimen from Santa Catarina State (Southern Brazil) deposited at NHRS and transferred this species to Costanana. In the same work, the authors described Costanana cella based on a single male specimen from Santa Catarina State (Fig. 15A-B) and commented that this specimen probably is a male of C. praecellens. We examined 65 pinned males and 21 females from Paraná State (Southern Brazil) (Figs 13C-D, 16-17), that fit perfectly with the descriptions of these species and therefore we propose that C. cella is a junior synonym of C. praecellens. DeLong & Wolda, 1983 Fig

Remarks
Costanana helvacosta closely resembles C. nana, C. santana and C. rubromarginata sp. nov. and was described based on a single male from Colombia.

Remarks
This species is known based on a single female specimen from Jamaica ( Fig. 19A-D) and was one of the 16 species not studied in the revision of the genus Gypona (DeLong & Freytag 1964). The external morphology and general coloration of Gypona nupera is very similar to that of Costanana cifi sp. nov. (Fig. 11E-F) -see remarks on this species -and because of that we decided to transfer this species to Costanana. Genus Acuponana DeLong & Freytag, 1970 Acuponana minuta (Spångberg, 1878) comb. nov. Fig. 20 Gypona minuta Spångberg, 1878: 37.

Remarks
This species is only known based on a female specimen from Colombia ( Fig. 20A-D). The coloration of the forewing differs from all the other species of Costanana by the entire length of the costal margin being yellow and the veins black, strongly contrasting with the translucent yellowish membrane. The crown with transverse striae, the anterior margin moderately thick and the alar appendix well developed are similar features to those of species of Costanana. We have in our possession four undescribed new species from the states of Amazonas, Mato Grosso and Pará, Brazil, with exactly the same color pattern on the forewings but that differ from C. minuta in the coloration of the pronotum, having two or three pairs of well defined round black spots. The male genitalia of these new species are most similar to those of Acuponana DeLong & Freytag, 1970or Marganana (Declivana) DeLong & Freytag, 1963 in having the male sternite VIII short, the subgenital plates exposed, without filiform setae, the male pygofer with short process on the dorsoapical margin and the few macrosetae grouped near the apex, similar to A. consesa DeLong & Freytag, 1970, A. fera DeLong & Freytag, 1970  1970, and the connective T-shaped. In our understanding, these species must belong to Acuponana and, by comparison of the external morphology and general coloration, this also applies to Costanana minuta.

Remarks
DeLong & Martinson (1972) described Gypona viridans based on a single male specimen from Santa Catarina State, Southtern Brazil (Fig. 21A-B). The holotype of Costanana asymmetrica was collected by Heinrich Wilhelm Schott, Austrian botanist taxonomist of Araceae Juss., who spent four years in Brazil (1817-1821) studying the flora in locations in Rio de Janeiro State such as Cabo Frio, Rio Paraíba, Rio Paraibuna, Cantagalo and Cachoeira de Macacu. This species was described by DeLong & Freytag (1972b) based on this single male specimen (Fig. 21C-E) that strongly contrasts the other species of Costanana in having the coloration uniformly yellow, the crown surface with oblique-rugose striae (like in figure 25D), and the alar appendix undeveloped. We also studied two male specimens from Minas Gerais State, Brazil ( Fig. 21F-G). Curiously, one of the specimens has the branched process at the left side of the aedeagus as in G. viridans and the other at the right side as in C. asymmetrica. After comparing these two specimens with the illustrations and descriptions of G. viridans and C. asymmetrica, we concluded that are no differences that can separate these two species and therefore they should be considered as synonyms.

Remarks
This species was described based on a single male from Rio de Janeiro State, Brazil (Fig. 22A-D). We studied an additional male specimen from Pernambuco State (Fig. 22E-F) that leaves no doubt that this species should be transferred to Gypona because of its having the crown with oblique striations between the ocelli (Fig. 25C) and the genitalia and coloration similar to those of several species of Gypona.

Remarks
Gypona flavicosta is remarkably similar in coloration and external morphology to C. asymmetrica and was described based on a single male from Rio de Janeiro State, Brazil ( Fig. 23A-D). DeLong & Freytag (1972b) transferred this species to Costanana. We studied four males and two females from the states of Paraná and Espírito Santo, Brazil.

Key to males of Costanana
Except Costanana nupera only known for the female. 15. Aedeagus with two pairs of apical processes equal in length (DeLong & Wolda 1983: 467, fig. 6

Etymology
The new genus name is feminine and combines the prefix 'meta' ('different') + part of the name Costanana, a closely related genus of Gyponini.

Description
StruCture. Head in dorsal view (Fig. 26A): slightly produced anteriorly, median length of crown less than half interocular width; crown surface with fine parallel transverse striations; anterior margin broadly rounded and parallel to anterior margin of pronotum, slightly extending over eye margin; transocular width of head narrower than humeral width of pronotum; ocelli equidistant between median line and eyes and slightly closer to anterior than posterior margin of crown; in frontal view (Fig. 26B), frons longer than wide, flat, not excavated below anterior margin of crown and with a few thin striations, texture shagreen; frontogenal suture distant from eye margins by maximum width of clypeus, not surpassing antennal ledge; antennal ledge carinated, obliquely downwards in relation to frons and extending over frons by short distance; epistomal suture indistinct medially; gena with ventrolateral margins slightly convex at midlength and weakly excavated near eye; maxillary plate produced ventrally as far as clypeus apex; clypeus not inflated, ca 1.5× as long as wide, lateral margins weakly divergent apically, apex emarginated; in lateral view (Fig. 26C), crown-face transition thin, with three transverse carinae. Pronotum in dorsal view (Fig. 26A): transversely striated except near anterior margin; lateral margins convergent anterad; in lateral view (Fig. 26C), moderately declivous; head and pronotum in continuous slope. Scutellum (Fig. 26C), not inflated. Forewing (Fig. 26A) without extra crossveins; venation distinct; M vein with segment after divergence between R+M and before cross vein m-cu 1 , as long as length of m-cu 1 ; outer anteapical cell short, almost half length of central and inner anteapical cells; five apical cells (R1 vein present); appendix well developed and involving first and second apical cells; apex slightly tapered. Profemur elongated, approximately 3.5× as long as wide; AD, AM, and PD rows reduced and poorly defined, with exception of apical setae AD 1 , AM 1 and PD 1 respectively; AV row with 3 short setae; PV row with only an apical seta present. Protibia, in cross-section, more or less cylindrical, with longitudinal carina adjacent to PD row weakly developed; AV row formed by long setae, gradually increasing in thickness and length towards apex; AD formed by many small undifferentiated setae; PD row with 3 long setae and intercalary undifferentiated setae; dorsal surface with apical setae AD 1 and PD 1 developed; PV row with 4 setae. Metafemur with setal formula 2:2:1. Metatibia rows PD, AD, and AV with 22-23, 12-13, and 14-15 macrosetae, respectively; AD row with intercalary setae between macrosetae; PV row with setae of apical half formed by sequence of 1 thicker and 3-4 thinner setae, ending with a thick seta. Metatarsomere I inner row of plantar surface with 5-6 setae, outer row very reduced in size; apex with 3 patellae flanked by 1 tapered lateral seta on inner and 1 on external corner. Metarsomere II pecten with 2 platellae flanked by 2 tapered lateral setae on inner and 1 on external corner.

Distribution
Southern Brazil.

Remarks
The crown of Metacostana cornuta gen. et sp. nov. transversely striated and the anterior margin defined are features present in the genera Acuthana, Costanana, Domahovana, Dumorpha, Delongiana Domahovski et al., 2020, Nullana DeLong, 1976, Ponana Ball, 1920, Regalana DeLong & Freytag, 1975, and some species of Polana DeLong, 1942. Among these genera, the presence of apodemal processes on the aedeagus is shared only with Dumorpha, Ponana and some subgenera of Polana, and the male sternite VIII being well produced posterad, often hiding the subgenital plates, is shared with Acuthana, Costanana, Domahovana, Dumorpha and some species of Polana. Probably the unusual ventrally curved shape of the ovipositor will place Metacostana gen. nov. as sister group of Acuthana, which was the only known genus of Gyponini with this feature. The new genus can be differentiated from Acuthana by the color, with black and yellow maculae (uniformly yellowishbrown in Acuthana), the absence of punctuations and reticulations on the forewings, the male pygofer with process on the ventroapical margin, the subgenital plate not tapered apically, the connective liner, the presence of apodemal processes on the aedeagus, the most strongly curved ovipositor, with smaller dorsal protuberance on the second valvulae, and the apex without denticles on the ventral margin. Unfortunately the ovipositor of females of Dumorpha is unknown, but the new genus can be separated by the color (pale yellow without black markings or only a few in Dumorpha); the ocelli smaller and equidistant between eye and midline (closer to the eye margin in Dumorpha), the presence of a process on the ventroapical margin of the male pygofer, the connective linear shape, and the atrial processes of the aedeagus being broader. Compared to Costanana, the new genus can be differentiated due to the metatibia row AD having intercalary setae between the macrosetae (also present in Dumorpha), the presence of apodemal processes on the aedeagus and the different morphology of the female ovipositor.  (Fig. 26G), with robust process on ventroposterior margin, extending posterad and surpassing pygofer apex. Style (Fig. 26K), in lateral view, blade slender, evenly curved dorsally, with uniform height to near apex. Aedeagus (Fig. 26L-M) apodemal processes broad at base, curved dorsally, apex tapered, not surpassing apex of aedeagus; shaft with pair of short subapical processes extending laterally and abruptly curving anterad.

Etymology
The new species epithet comes from de Latin word 'cornutus' meaning 'horned'. It alludes to the hornshaped process of the male pygofer. Other material BRAZIL • 1 ♂, 1 ♀ (stored in ethanol); same collection data as for holotype; DZUP.
Coloration. Head and thorax (Figs 26A-C, 28) yellowish-brown. Crown (Fig. 26A) with pair of round black spots behind ocelli, near posterior margin; eyes and ocelli red. Pronotum (Fig. 26A) with pair of small round black spots behind eyes, near anterior margin. Face (Fig. 26B) without maculae. Proepimeron (Fig. 26C) without maculae. Forewing (Fig. 26D) costal margin with pair of black maculae near base and midlength, yellow macula between black maculae, and white macula at level of outer anteapical cell and third and fourth apical cells; veins outlined by black; cross veins of inner discal cell black; apex of anal veins, first to third apical cells and appendix smoky; appendix with whitish macula on basal third. Legs yellowish-brown with basal half of mesotibia black.  Male terMinalia. Sternite VIII (Fig. 26E) 1.1× as long as wide; lateral margins with triangular projection near base; apex broad, truncated, and weakly emarginated. Valve (Fig. 26F) 1.8× as wide as long; integument thickening present only on anterior margin; posterior margin rounded. Pygofer (Fig. 26G), in lateral view, 1.7× as long as maximum height; basodorsal process absent; ventral margin broadly rounded; posterodorsal margin rounded; robust process on ventroposterior margin, extending posterad and surpassing pygofer apex, with transverse thin rugae, apex abruptly tapered and acute; macrosetae distributed on posterodorsal quadrant; apex subtruncated. Anal tube (segment X) membranous, without processes. Subgenital plate (Fig. 26G), in lateral view, not surpassing apex of pygofer; in ventral view (Fig. 26H), 4× as long as wide, wider near base than apically; inner margin straight; outer margin rounded near base; apex abruptly tapered, subacute, well sclerotized. Connective (Fig. 26I) linear-shaped; arms not produced laterally; stalk almost as wide as width of arms, long, 2× as long as maximum width of arms; median keel well developed. Style (Fig. 26J), in dorsal view, with outer lobe strongly developed, parallel to blade, rounded; in lateral view (Fig. 26K), blade slender, evenly curved dorsally, with uniform height to near apex; ventral margin not serrated; apex tapered and subacute. Aedeagus ( Fig. 26L-M) preatrium not developed; dorsal apodeme produced dorsally but not laterally, dorsal margin excavated, dorsolateral corners rounded; apodemal processes broad at base, curved dorsally, apex tapered, not surpassing apex of aedeagus; shaft cylindrical, curved dorsally, gradually tapering to apex; apical portion with pair of short subapical processes extending laterally and abruptly curving anterad.

Discussion
Among the previously described species of Costanana we have not examined the holotypes of C. apicata and C. circumaga (OSUC), but based on the original descriptions and illustrations it is unlikely that these species belong to Costanana. The holotype of Costanana nana (BMNH) was also not studied, but this species closely resembles other species of Costanana. It would be interesting to carry out a phylogenetic study to test the monophyly of Costanana mainly due to the differences between the type species C. dunda in relation to most species of the genus, as for example, the different coloration (similar to G. costata), the valve deeply excavated medially, which is a characteristic feature not found in any other species of Costanana studied herein, but present in some species of Acuponana, and the male pygofer with a few macrosetae grouped near the apex, and having a dorsoapical process, similar to some species of Acuponana and Gypona. The shape and length of the male sternite VII represent diagnostic features for the generic recognition of Costanana, but they are unknown in C. dunda. We suspect that C. dunda may be a species of Acuponana, a poorly know genus that lacks strong diagnostic characters and need a taxonomic revision. As Acuponana is the oldest genus, then another species would have to be chosen as the type species under a new name for Costanana. Also, images of C. apicata and C. circumaga still need to be studied to determine the correct position for them.
With the taxonomic changes and new species described, Costanana now comprises 17 species with records in nine Brazilian States: Bahia, Mato Grosso, Minas Gerais, Pará, Paraná, Pernambuco, Rio Grande do Sul, Rio de Janeiro, Santa Catarina, and the Distrito Federal. Gyponini now comprises approximately 1471 species in 80 genera.