Two new species of Hyalojassini (Hemiptera: Cicadellidae: Iassinae) to the genera Comanopa Blocker and Gehundra Blocker, description of the male of G. sordida (Baker) and key to species

Comanopa Blocker, 1979 and Gehundra Blocker, 1976 are small and poorly known genera of leafhoppers, previously comprising four and five species, respectively. In this study, two new species are proposed: Comanopa mananciensis sp. nov. from the state of Paraná, Southern Brazil, and Gehundra cristalinensis sp. nov. from the state of Mato Grosso, Centralwestern Brazil. The male of Gehundra sordida (Baker, 1900) is studied by the first time. Detailed descriptions and illustrations of males and females are provided and keys to males of Comanopa and Gehundra are given.


Introduction
Cicadellidae Latreille, 1825 is the largest family of Hemiptera Linnaeus, 1758 and one of the most successful radiations of plant-feeding insects known as leafhoppers, currently classified into 25 subfamilies (Krishnankutty et al. 2016). The Neotropical region has the richest fauna of leafhoppers, with more than 5000 described species (Freytag & Sharkey 2002).
Hyalojassini comprises small and ovate leafhoppers, between 3.5 to 8.0 mm in length, which can be recognized mainly by the head usually rounded in profile, without distinctly delimited crown; the forewing with inner apical cell usually more membranous than adjacent cells; the hind wing with veins R4+5 and M1+2 confluent distally; male subgenital plates usually very short or vestigial and completely concealed by pregenital sternite; and the female genitalia with first valvula with dorsal sculpturing usually areolate and second valvula with few widely spaced dorsal teeth (Krishnankutty et al. 2016). Hyalojassini is distributed in SE Asia (9 genera and 32 species) and the New World (29 genera and 117 species), representing an interesting biogeographical disjunction Krishnankutty et al. 2016;Wang et al. 2018;Domahovski & Cavichioli 2019c;Domahovski 2020). Although most of the of hyalojassine species are described from the Neotropical region (more than 70% of the known species), the real diversity of the tribe remains poorly known, especially in Brazil, where only 12 genera and 21 species were previously recorded (Defea & Paradell 2019;Domahovski & Cavichioli 2019b, 2019cDomahovski 2020).  described the genus Gehundra with three new species from Peru, Gehundra tricosa (type species), G. avulsa and G. galbina, each known from a single male specimen, and provided a key to the species. Blocker (1979) transferred Bythoscopus pallidus Osborn, 1924 and Macropsis sordidus Baker, 1900 to Gehundra, both species known from a single female specimen from Brazil.
Comanopa was described by Blocker (1979) with Stragania puertoricensis Caldwell, 1952 from Puerto Rico (type species), Stragania fasciata Linnavuori, 1956 from Brazil (only know by female specimens) and one species describe as new, Comanopa adelpha from Panamá and Mexico. Blocker (1982) described the last species, Comanopa hamiltoni from Jamaica.
This study aims to contribute to the knowledge of Hyalojassini by describing two new species belonging to the genera Comanopa and Gehundra. Additionally, one male and four females of Gehundra sordida (Baker, 1900), from the state of Mato Grosso do Sul, Centralwestern Brazil, are studied and the male and female genitalia are described for the first time.

Material and methods
The descriptive terminology adopted herein follows mainly Young (1968Young ( , 1977, except for head features (Hamilton 1981;Mejdalani 1998), wings (Dietrich 2005) and leg chaetotaxy (Rakitov 1997). The term gonoplac follows Mejdalani (1998) and pseudostyle (thickened portion along the inner margin of the subgenital plate) follows Kramer (1963) and Blocker (1979). The techniques used for dissection of male and female terminalia follow Oman (1949), with the few modifications described by Cavichioli & Takiya (2012).The dissected parts are stored in microvials with glycerin which are pinned bellow the specimen.
Body. Small and ovate leafhoppers, not depressed dorsoventrally.
Head (Figs 1-3). Dorsal view: very short, transocular width nine-tenths humeral width of pronotum; crown not developed, nearly vertical, median length 1/10 of interocular width, surface with transverse striae; anterior and posterior margins parallel and broadly rounded; ocelli not visible dorsally; lateral view: crown-face transition broadly rounded with transverse striae; frons slightly inflated; clypeus almost flat; ventral view: face less than two times wider than high; ocelli small, on anterior margin, mesad of antennal pits and distant from eyes; frons with transverse striae, lateral margins strongly convergent ventrally; frontogenal suture extending to antennal ledges; antennal ledge carinate and slightly oblique, not concealing antennal base; antenna length ca ⅓ width of head; gena broad, with small setae; ventrolateral margin broadly rounded, completely concealing proepisternum; maxillary plate produced ventrally as far as clypeus apex; lorum broad, with short setae, maximum width approximately equal to that of clypeus, subgenal suture rounded, extended nearly to midlength of lateral margin of frons; clypeus slightly longer than wide, epistomal suture complete and straight, lateral margins parallel, apex carinated and straight.
legs. Profemur moderately elongated, 2.7 times longer than high; PD, AD and AM rows formed by undifferentiated setae, PD1 and AD1 setae short and AM1 absent; IC row formed by double row of fine setae, continuous with AV row; AV and PV rows formed by several fine setae. Protibia, in crosssection, almost circular; AV row formed by short, thin setae in the basal half and slightly more robust setae distally; AD row consisting of only undifferentiated setae; PD row with 3 long setae intercaled by undifferentiated setae; PV row with 3-4 long setae intercaled by undifferentiated setae. Hind leg with femoral setal formula 2:2:1; PD2 seta reduced in size; metatibia PD, AD and AV rows with 15-16, 9-10 and 9-10 macrosetae respectively; AD row without intercalary setae between macrosetae; PV row with setae of apical half intercalating 1 longer and thicker seta and 3-4 shorter and thinner setae, ending with 2 very short, thin setae; first tarsomere without enlarged dorsoapical seta; ventral surface with two longitudinal rows of simple (non-cucullate) thin setae; pecten with 4 platellae flanked by tapered lateral setae; second tarsomere pecten with 2 platellae flanked by tapered lateral setae. terMinalia. Sternite VIII (Fig. 4) convex, moderately produced posterad, 1.2 times wider than long, shorter than sternites VII and VI combined; ventral surface with many small setae; lateral angles  rounded; posterior margin straight. Valve (Figs 5-7) broadly fused laterally to pygofer; ventral margin excavated laterad of small median lobe. Pygofer (Figs 5-7) with pair of processes arising near base of plates, short and rounded apically, ventral surface with short setae. In lateral view, pygofer ( Fig. 6) higher on apical half; dorsal margin widely notched on basal ⅔; dorsoposterior margin fused to pygofer lobes; ventral margin strongly produced ventrad, with short setae forming two longitudinal lines, internal process fused to pygofer lobe, arising near apex and bordering ventral margin, forming a small dentiform protrusion near apex of subgenital plate; lateral surface with longitudinal carina on basal ⅔; apex very wide and truncated; few macrosetae dispersed near apex. Anal tube (Fig. 5) membranous, without processes. Subgenital plate (Figs 5-8) short, not overlapping pygofer laterally, produced posteriorly as far as half length of pygofer; ventral surface without setae; anterior portion elongated and curved dorsally; apical portion expanded and ovate; apex rounded. Connective (Fig. 9), thin, slightly arched bar. Style (Figs 9-10), short, not surpassing apex of subgenital plates; apex curved ventrad in lateral view; apex subacute, slightly curved outward in dorsal view. In lateral view, aedeagus ( Fig. 11) with preatrium moderately developed, produced anterad; dorsal apodeme developed, anterior margin forming pair of flattened divergent arms truncated apically, directed dorsally, posterior margin produced ventrally; shaft directed ventrally, slightly curved posterad and tapered apically. In anterior view, aedeagus ( Fig. 12) with shaft deeply split apically, whit pair of subapical processes long, directed laterally; gonopore subapical.

Female
Unknown.
Body. Small and ovate leafhoppers, not depressed dorsoventrally.
Head (Figs 13-15). Dorsal view: very short, transocular width nine-tenths humeral width of pronotum; crown not developed, vertical, surface whit transverse striae; anterior and posterior margins parallel and broadly rounded; ocelli not visible dorsally; lateral view: crown-face transition broadly rounded with transverse striae; frons and clypeus slightly inflated; ventral view: face more than two times wider than high; ocelli small, on anterior margin, mesad of antennal pits and distant from eyes; frons with transverse striae, lateral margins strongly convergent ventrally; frontogenal suture surpassing antennal ledge, extending to anterior margin of crown; antennal ledge carinate and slightly oblique, not concealing antennal base; antenna length ca ⅓ width of head; gena broad, with small setae; ventrolateral margin broadly rounded, completely concealing proepisternum; maxillary plate produced ventrally as far as clypeus apex; lorum with few short setae, narrower than clypeus width, subgenal suture rounded, extended nearly to midlength of lateral margin of frons; clypeus slightly shorter than basal width, epistomal suture complete and rounded, lateral margins excavated medially, apex slightly broadened, rounded and carinated.
legs. Profemur moderately elongated, 2.9 times longer than high; PD, AD and AM rows formed by undifferentiated setae, PD1 and AD1 setae short and AM1 absent; IC row formed by double row of fine setae, continuous with AV row; AV and PV rows formed by several fine setae. Protibia, in cross-section, almost circular, without defined longitudinal carina adjacent to PD row; AV row formed by short, thin setae in the basal half and slightly more robust setae distally; AD row consisting of only undifferentiated setae; PD row formed by with 3-4 long setae intercaled by undifferentiated setae; PV row with 6-7 short setae. Hind leg with femoral setal formula 2:2:1 or 2:2:1:1; PD2 seta reduced in size; metatibia PD, AD and AV rows with 14-19, 9-11 and 10-12 macrosetae respectively; AD row without intercalary setae between macrosetae; PV row with setae of apical half intercalating 1 longer and thicker seta and 3-4 shorter and thinner setae, ending with 3 thin setae; first tarsomere without enlarged dorsoapical seta; ventral surface with two longitudinal rows of simple (non-cucullate) thin setae; pecten with 4 platellae flanked by tapered lateral setae; second tarsomere pecten with 2 platellae flanked by tapered lateral setae.
coloration. Head and thorax (Figs 13-15) brownish-yellow. Face (Fig. 15) with muscle impressions of frons brown; lorum with ventral half brown. Pronotum (Fig. 13) with irregular brown markings near anterior margin. Mesonotum (Fig. 13) with lateral angles dark brown. Forewing (Figs 13-14) with brown setae and groups of black setae at midlength of anal margin, apex of clavus and apex of inner discal cell (variable in intensity between the specimens).
terMinalia. Sternite VIII (Fig. 16) convex, strongly produced posterad, two times longer than wide, longer than sternites VII and VI combined; ventral surface with many small setae, except basal portion and along median line; lateral margins tapered toward apex, with small rounded lobe at basal third; apex subacute. Valve (Figs 17-18) longer than wide, broadly fused laterally to pygofer; lateral margins convergent toward ventral margin, with strong integument thickening; ventral margin rounded. In lateral view, pygofer (Figs 17-18) without processes, higher near base; dorsal margin deeply notched after half its length; dorsoposterior margin fused to pygofer lobes, straight and oblique; ventral margin approximately straight, with short setae near base and long thin setae on apical half; lateral surface with longitudinal carina near ventral margin of basal ⅔; apex very wide and truncated; without macrosetae, but with few short setae near apex. In ventral view, pygofer (Fig. 19) with ventral margin folded inward, from base to apex, with short setae on inner surface. Anal tube (Fig. 17) membranous, without processes. Subgenital plate (Figs 17-20) short, with well developed pseudostyle; inner margin nearly straight; external margin rounded, almost circular, overlapping pygofer laterally; apex deeply notched, pseudostyle acute, with few short setae. Connective amorphous. Style (Figs 17, 21) long, approximately straight, apodeme and apophysis long, subequal in length; apex slightly tapered and curved dorsally. Aedeagus (Figs 22-23), with preatrium not developed; dorsal apodeme developed, with lateral margins extended laterad and posterior margin extending to almost apex of shaft; shaft slightly curved dorsally;  with many thin setae; ventral margin broadly rounded with group of thick setae at base; apex obliquely truncated. Ovipositor slender, evenly curved dorsad, not surpassing pygofer apex. First valvifer (Fig. 27) trapezoid, approximately as long as wide; anterior margins of both valvifers connected by sclerotized membrane. First valvula (Fig. 27) with dorsal and ventral margins almost parallel, slightly convergent apically, ca 7 times longer than high at base; ventral interlocking device long, extending over basal ⅔; dorsal sculptured area areolate starting before the midlength; apical portion ( Fig. 28) with ventral sculpture present only near apex, continuous with the dorsal sculpture, apex moderately tapered and acute. Second valvifer (Fig. 31) ca three times higher than long. Second valvula (Fig. 29) ca 10 times longer than high; dorsal and ventral margins parallel; dorsal margin with three distinct subapical teeth widely spaced; apical portion ( Fig. 30) of dorsal margin with two rounded subapical notches forming one rounded tooth between apex and first subapical tooth; ventral margin without denticles and with one subapical tooth. Gonoplac (Fig. 31) four times longer than high; dorsoapical margin long, half length of gonoplac; external surface without dentiform cuticular projections; ventral margin broadly rounded, with few spaced macrosetae near ventral margin; apex rounded.

Remarks
Gehundra cristalinensis sp. nov. is most similar to G. avulsa in having the male pygofer with ventral margin bearing long setae on apical half and the subgenital plate deeply notched apically. However, the new species can easily by separated by the subgenital plate ( Fig. 20) with external margin rounded, almost circular (less expanded laterally and not circular in G. avulsa) and the style (Fig. 21) approximately straight, with apex slightly tapered (style with apophysis more curved, with apex footshaped in G. avulsa).
terMinalia. Sternite VIII (Fig. 35) convex, strongly produced posterad, 1.5 times longer than wide, longer than sternites VII and VI combined; ventral surface with many small setae, except basal portion and along median line; lateral margins with small rounded lobe at basal third; apex abruptly tapered and rounded. Valve (Figs 36-37) longer than wide, broadly fused laterally to pygofer; lateral margins convergent toward ventral margin, with strong integument thickening; ventral margin rounded. In lateral view, pygofer (Figs 36-37) without processes, higher near basal third; dorsal margin notched after half its length; dorsoposterior margin fused to pygofer lobes, straight and oblique; ventral margin excavated basally, with rounded lobe at basal third, straight on posterior ⅔, with short, thick setae grouped in line near midlength; lateral surface without macrosetae, but with few short setae near apex. In ventral view, pygofer ( Fig. 38) with ventral margin folded inward, from base to apex, with short setae on inner surface. Anal tube membranous, without processes. Subgenital plate (Figs 36-39) short, with well developed pseudostyle; inner margin nearly straight; external margin with rounded lobe on basal half, overlapping pygofer laterally; apex of pseudostyle subacute, with thin setae along external margin and group of thicker setae on apex. Connective amorphous. Style (Figs 36, 40) long, approximately straight, apodeme and apophysis long, sub equal in length; apex rounded, curved dorsally. Aedeagus (Figs 41-42), with preatrium not developed; dorsal apodeme developed, lateral margins extended laterad and posterior margin extending to almost apex of shaft; shaft slightly curved dorsally, with height approximately constant from base to apex. In ventral view, aedeagus (Fig. 42) . 45) with dorsal and ventral margins, slightly convergent apically, ca 8 times longer than high at base; ventral interlocking device long, extending over basal ⅔; dorsal sculpturing area areolate starting before the midlength; apical portion ( Fig. 46) with ventral sculpture present only near apex, continuous with the dorsal sculpture, apex gradually tapered and acute. Second valvifer (Fig. 49) ca three times higher than long. Second valvula (Fig. 47) ca 10 times longer than high; dorsal and ventral margins parallel; dorsal margin with three distinct subapical teeth widely spaced; apical portion ( Fig. 48) dorsal margin with two rounded notches forming one rounded tooth between apex and first subapical tooth; ventral margin without denticles and with one subapical tooth. Gonoplac (Fig. 49) four times longer than high; dorsoapical margin long with half length of gonoplac; external surface without dentiform cuticular projections; ventral margin broadly rounded, with few spaced macrosetae near ventral margin; apex rounded.

Remarks
Gehundra sordida is similar to G. galbina and G. cristalinensis sp. nov. in lacking long setae on ventral margin of pygofer, and is similar to G. cristalinensis sp. nov. in having the apex of style not foot-shaped. However, G. sordida can be easily separated by the subgenital plate with external margin with lobe limited to the basal half of pseudostyle (external margin with lobe occupying more than half length of pseudostyle in G. avulsa and G. cristalinensis sp. nov., and occupying entire length of pseudostyle in G. tricosa and G. galbina. Key to males of Gehundra (G. pallida only known from the female) 4. Pygofer with long setae on ventral margin (Fig. 18). Subgenital plate with lobe of the external margin occupying more than half length of pseudostyle (Fig. 20) (Fig. 37). Subgenital plate with lobe of the external margin limited to the basal half of pseudostyle (Fig. 39) ..............G. sordida (Baker, 1900) (Brazil)

Discussion
The genus Comanopa can be recognized by the forewing with vein separating appendix and first apical cell incomplete (evanescent apically); the pygofer with process (pygofer hooks) arising near apex, extending anterad, similar to Penestragania Beamer & Lawson, 1945; the subgenital plates simple, slightly reduced in size, similar to Momoria Blocker, 1979; the style reduced in length; and the connective like a narrow semicircular band (Blocker 1979). In the new species of Comanopa proposed herein, the process of pygofer differs from the process present in species of Penestragania, because it is entirely fused to the lateral lobe of pygofer, resembling more an integument thickening than a process, whereas in species of Penestragania this process is separated from the lateral lobe by most of its length. However, the remaining species of Comanopa need to be studied to determine if this characteristic found in C. manancienis sp. nov. is unique for this species or is shared with other species of the genus.
The genus Gehundra can mainly be recognized by the forewing with vein separating appendix and first apical cell complete; the male sternite VIII very long, reaching the apex of pygofer, in repose; the subgenital plates with external margin lobed, overlapping the lateral portion of pygofer and with well developed pseudostyles; and the connective amorphous . Only two species of Gehundra were previously known from Brazil, G. pallida and G. sordida, both described based on a single female specimen. According to the descriptions and illustrations of Blocker (1979), G. sordida has the posterior margin of sternite VII with a narrow and deep notch medially (Fig. 43;Blocker 1979: 22, fig. 24) and G. pallida has the posterior margin evenly excavated, almost rounded (Blocker 1979: 22, fig. 23). Gehundra cristalinensis sp. nov. can be differentiated from G. pallida and G. sordida in having the posterior margin of the sternite VII with a broad and acute notch medially (Fig. 24).
The diversity and distribution of leafhoppers of the tribe Hyalojassini remain poorly known. The majority of species described from the Neotropical region don't have the male and female associated and are known based on a few specimens, commonly only by the holotype. Almost all studies which described neotropical species currently placed in this tribe are anterior to the 1980s and considering only a few generic revisions are available and many species remain undescribed in Brazilian collections, this is a wide field to be explored.