Pintalia Stål, 1862

SANTOS, JÚLIO CÉSAR DO CARMO VAZ, HOCH, HANNELORE, BARTLETT, CHARLES R. & FERREIRA, RODRIGO LOPES, 2025, Revision of the type series of the original species placed in the genus Pintalia Stål, 1862 (Hemiptera: Fulgoromorpha: Cixiidae), and description of 12 new species from Brazilian caves, Zootaxa 5678 (1), pp. 1-96 : 7-17

Genus Pintalia Stål, 1862

Type species: Pintalia lateralis Stål, 1862 by original designation.

Amended diagnosis. (Adapted after Muir 1934, Kramer 1983.) Vertex trapezoidal ( Fig. 1 ), with two compartments (apical and basal) and two carinae (apical transverse and subapical carina), apical transverse carina usually visible in dorsal view; basal compartment larger than apical compartment. Frons ( Fig. 2A ), with median carina present. Rostrum reaching or surpassing the hind coxae. Lateral ocelli, and frontal ocellus present well developed. Pronotum short ( Fig. 1 ), in dorsal view narrower medially. Mesonotum with three carinae ( Fig. 1 ), planate in lateral view, usually distally inclined in lateral view. Tegmina ( Fig. 12 ) tectiform, in resting position usually surpassing approx. 1/3 of body length with distal margins usually touching each other, and apical cells and veins usually elongated and parallel (between RA and icu veins), usually with 12–13 distal veins, with RP quadrifid, sometimes with punctual anastomosis between RA + RP1 ; rm-1 and mcu-1 usually near the first MP fork; fork of MP1 +2 usually trifid and MP3 +4 usually bifid; ScP+ RA and RP fork starting more proximally to the basal cell, than the fork in Cua1 and Cua2 that occurs more distally, in some species they occur at the same level; C1 and C5 cells usually more elongated; C1 cell always larger than C5 cell. Hind wing ( Fig. 12B–D ) with RP bifid, MP usually trifid (petiolate anastomosis in MP3 +4 + CuA1 distally forked – Y type) or bifid (complete anastomosis with distal fusion MP3 +4 + CuA1 – I type). Posterior legs, hind tibia in most species with lateral spines, in some without any (see genus description below), usually with 6 apical teeth-grouped as follows ( Fig. 3B ): 1 larger lateral (distal), 2 smaller together (medial), 1 larger (medial), and 2 (medium size) (internal).

Morphology. Body length: ♀ 2.8–6.0 mm (n=40). ♂ 2.6–5.3 mm (n=42).

Head ( Figure 1 ). Vertex trapezoidal, apical transverse carina and subapical carina approx. the same size usually concave and convex ( P. lateralis Stål, 1862 ) or straight, almost straight, straight bent forward; apical transverse carina distinct in some species, elongated medially and touching the median carina of the frons (Y-shape, in frontal view uniting forward of the eyes) ( P. ustulata Stål, 1862 ; P. inornata Stål, 1862 ); angle formed by the caudal margin of the vertex usually concave (e.g., P. lateralis ) or triangular (e.g., P. fasciatipennis Stål, 1862 ); lateral carinae elevated (except P. fasciatipennis ); interspecific variation observed in the basal emargination (median carina), absent or inconspicuous ( P. lateralis ; P. obscuripennis Stål, 1862 ; P. pictipennis Stål, 1862 ; P. fraterna Stål, 1862 ), weakly developed ( P. inornata ), or well developed ( P. fasciatipennis ; P. consobrina Stål, 1862 ; P. ustulata ). Frons, longer than wide; up to 2.0 times wider medially than apically (e.g., P. lateralis ); trapezoidal apically, with apical region usually above the compound eyes (except P. ustulata ; P. inornata ) and straight or concave apically (apical transverse carina frontally); median carina not reaching or rarely reaching the frontal ocellus (except P. consobrina ), short in some species, touching to curved apical transverse carina medially (Y-shape in frontal view); lateral carinae of the frons usually elevated (except P. fasciatipennis ). Antenna with short scape and semi-globular pedicel, slightly surpassing the margins of the frons when in frontal view.

Thorax ( Figure 1 ). Pronotum tricarinate, with median carina absent or inconspicuous; submedian carinae moderately developed (e.g., P. lateralis ), often not continuous dorsally (weakly developed); dorsally shorter than length in the middle between the eyes (e.g., P. lateralis ); caudal margin rounded or straight/obtuse (e.g., P. lateralis ). Mesonotum dorsally with three carinae (e.g., P. lateralis ), which are usually narrow near the pronotum and wide near the abdomen (except P. inornata ); in some species region between carinae lighter than on the lateral area near the tegulae ( P. lateralis and little in P. consobrina ). Tegmina (Forewings) with RP vein quadrifid (e.g., P. lateralis ), in some species with punctual anastomosis between RA+RP1 ( P. consobrina ; P. pictipennis ); tubercles occur throughout all veins of the tegmina, often more evident in some species; can be evanescent ( P. lateralis ; P. pictipennis ; P. fraterna ), double ( P. consobrina ), simple ( P. fasciatipennis ; P. obscuripennis ; P. proxima Stål, 1862 and P. ustulata ), or yet absent or inconspicuous ( P. inornata ); distal margin totally curved (except P. ustulata ); Cells C1b, C2, C3a, C3 and C4 (central prenodal cells), stacked together right behind the center of the tegmina (e.g., P. lateralis ); C1 and C5 elongated (lateral prenodal cells) (e.g., P. lateralis ), C1 a little closer to bc than C5 (except, P. ustulata ), or C5 very short ( P. inornata ) or approx. the same size as C1 ( P. ustulata ); usually 12 apical cells (nodal) and 7 subapical cells (prenodal) (except P. fasciatipennis and P. obscuripennis ). Hind wing with RP bifid (e.g., P. lateralis ), MP trifid usually with petiolate anastomosis in MP3+4 + CuA1 distally forked, Y – type ( P. lateralis , P. ustulata , P. fraterna ) or with complete anastomosis with distal MP3+4 + CuA1 fusion, I-type (possibly P. pictipennis ).

Legs. Hind tibia with 6 apical teeth; usually with 1–7 lateral spines, tiny near base, or unarmed in some species ( P. inornata ; P. obscuripennis ; P. ustulata ). 1 st tarsomere with 7 apical teeth of approx. the same size or, with 1–2 lateral teeth larger and 2 apical teeth moderately large in the middle. 2 nd tarsomere with 7–9 apical teeth, rarely 7 and rarely 9, all with approx. the same size, or with 1–2 lateral larger teeth and intermediate teeth gradually smaller medially or of uniform size, usually 1–3 platellae ( Pintalia painensis -group, Pintalia constellaris -group) or with 3 very thin setae ( Pintalia pictipennis -group).

Male terminalia. Pygofer with or without processes between dorsal and distal margins, asymmetric in some species; ventromedian process triangular, crooked in some species (e.g., P. magnaepiprocti sp. nov.). Anal tube ( Fig. 14 ), longer than wider, in some species distally developed, bent approx. at between 45°– 90° with epiproct and paraproct small ( Pintalia painensis -group); or elongated but not developed distally, with epiproct and paraproct small ( Pintalia constellaris -group); or moderately short and wide not developed distally, with paraproct large developed distally, and in some species with epiproct large developed distally ( Pintalia pictipennis -group). Genital styles, in lateral view long and well expanded apically ( Pintalia pictipennis -group, Pintalia constellaris -group), or long with the apex slightly or moderately expanded ( Pintalia painensis -group); in ventral view, touching medially and forming the basal opening, moderately large, laterally concave, and apically triangular ( Pintalia pictipennisgroup ) or small and diamond-shaped ( Pintalia constellaris -group and Pintalia painensis -group). Aedeagus with shaft tubular, ventral ridge developed and flagellum without spines ( Pintalia pictipennis -group), or tubular usually with three conspicuous spines at the apex of the aedeagal shaft, without ventral ridge and flagellum spineless ( Pintalia constellaris- group); or tubular and wider apically near to flagellum, ventral ridge absent, usually flagellum with spines and phallotreme laterally ( Pintalia painensis -group).

Female terminalia. Anal tube not developed distally (bent) ending together with epiproct and paraproct; usually elongated, tubular, or cylindrical dorsally with lateral margins moderately developed ( P. lateralis ; Pintalia painensis - group), or slightly longer than wide and flattened ventrally ( P. obscuripennis ; P. consobrina ; P. ustulata ) and in parts Pintalia pictipennis -group, present only P. serratilis sp. nov. and P. stali sp. nov. Epiproct and paraproct flattened, and small; epiproct triangular in dorsal view, with two rounded margins bent down laterally; paraproct rounded in dorsal view, sometimes weakly elongated. Gonocoxae VIII truncated, with a rounded laterally, in caudal view usually small and ovoid. Ovipositor sabre-shaped, in lateral view curved dorsally in some specimens surpassing anal tube.

Distribution. Neotropical region. South America, Brazil ( Stål 1862; Muir 1934), Bolivia ( Fennah 1945), Chile ( Coquimbo) ( Campodonico 2021), Ecuador ( Muir 1934; Fennah 1945), Guiana Francesca ( Fennah 1945), Guyana ( Muir 1934; Caldwell 1944), Peru ( Fennah 1945), Trinidad and Tobago ( Fennah 1945), Venezuela ( Lethierry 1890). North America, United States of America ( Kramer 1983; Bartlett et al. 2014; Van Duzee 1908; Caldwell 1944; Kramer 1983; Caldwell and Martorell 1951), Cuba ( Bartlett et al. 2014), Guadeloupe ( Fabricius 1803; Stål 1862), Cayman Islands ( Fennah 1971), Martinique ( Fabricius 1803; Stål 1862), Mexico ( Caldwell 1944; Bartlett et al. 2014). Central America, Costa Rica ( Caldwell 1944; Kunz and Holzinger 2022), Dominica ( Fennah 1948), Granada ( Fennah 1948), Guatemala ( Caldwell 1944; Bartlett et al. 2014), Haiti ( Uhler 1901), Nicaragua ( Bartlett et al. 2014), Panama ( Metcalf 1938), Saint Lucia ( Fennah 1948), Saint Vincent and the Grenadines (Uhler 1895; Fennah 1948).

Remarks. Pintalia is a large genus with very varied morphology and lacks an in-depth review of the species that make up the genus. All species here treated, including P. lateralis , have the vertex divided transversely into anterior and posterior compartments by the subapical carina. However, in P. inornata and P. ustulata the apical transverse carina is medially elongated and, as a result, it exhibits a larger apical compartment than in other species. Most species present the vertex with elevated carinae in dorsal view; however, in P. consobrina Stål, 1862 the subapical carina is not foliate, and in P. fasciatipennis Stål, 1862 neither of the two carinae (apical transverse and subapical) are elevated, which is uncommon for the genus. Furthermore, P. inornata , P. consobrina , P. fasciatipennis and P. ustulata are the only studied species with developed median carina in the basal compartment of the vertex (very distinct).

In the three species-groups herein proposed, the transverse apical carina can be concave in frontal view as in Pintalia pictipennis- group, or straight as in the Pintalia painensis- group and the Pintalia constellaris- group. In dorsal view, it may be the same size or larger than the subapical carina as observed for the Pintalia pictipennisgroup or of the same size or smaller than the subapical carina as in the Pintalia painensis- group and Pintalia constellaris- group. In frontal view, there appear to be two main frontoclypeal suture patterns. Some species exhibit a semicircular frontoclypeal suture bent upwards as in P. lateralis and the Pintalia pictipennis- group; however, in other species, it is angularly bent upwards as in P. fraterna and Pintalia painensis- group.

We did not find any significant variations in the thorax (pronotum and mesonotum) among the species reported here, except for P. inornata , which dorsally exhibits almost straight and parallel carinae on mesonotum vs. narrow near the pronotum and wider distally, which is observed in most species (e.g. P. lateralis ). All species reported here have lateral spines on the posterior tibia, except for P. inornata ; P. obscuripennis and P. ustulata (very distinct). Although this appears to be a convenient character, we were not successful in associating this feature with other features relevant to species groupings. Furthermore, it was not possible to accurately assess the apical teeth of the hind tibia of the species described by Stål, 1862. On the other hand, we observed thin setae ( Pintalia pictipennisgroup ) and platellae ( Pintalia painensis- group and Pintalia constellaris- group) on the new species herein described, and this character is normally used for subgeneric-grouping in Cixiidae . However, it can be in some species absent or in a smaller number on one of the legs.

Although the tegmina can present different anastomosis and crossveins in some species, it usually exhibits stable cell patterns, with 12 apical (nodal) cells and 7 subapical (prenodal) cells ( Fig. 12–13 ). However, in some species, there may be an asymmetry of cells and veins between the right and left tegmina. For most species, the apical margin is smoothly curved between RA and icu, except in P. ustulata . Here, the Pintalia pictipennis -group exhibits punctual anastomosis in RA+RP1 veins, vs. separate by crossvein in Pintalia constellaris -group and Pintalia painensis -group). The shape of the basal margin of the pterostigma can also be useful for identification, since in the Pintalia pictipennis- group it is small and distally triangular, while in the Pintalia constellaris- group it is large and sub-rectangular weakly angled distally, and in the Pintalia painensis- group it is large and sub-rectangular, weakly, or well angled distally. Although there are conspicuous spotting patterns in some species and -groups, some patterns can be observed in species of the three groups herein proposed, such as an inclined transverse spot in the center of the tegmina (most species), two semicircular spots in pcc (most species), and another three semicircular or rounded spots between ScP+RA and A1.

It was only possible to examine the posterior wings of P. lateralis ( type), P. ustulata and P. fraterna . These species have a petiolate anastomosis in MP3 + CuA1 distally forked as Pintalia painensis- group ( Fig. 12B ) and Pintalia constellaris- group ( Fig. 12C ) (I-Type). However, Pintalia pictipennis- group exhibits complete anastomosis with distal MP3+4 + CuA1 fusion ( Fig. 12D ) (Y-type) ( Emeljanov 2002, Le Cesne 2022).

Character distribution across Pintalia Stål, 1862

The groups that we propose were based on the morphology of the new species presented.Although they were useful, we did not use the groups formed by Fennah (1945), as the species treated by him could not be fully evaluated before the construction of this work. Phylogenetic analyses will be necessary to identify whether the synapomorphies we indicate are useful for the proposal of subgenera or new genera in Pintalia .

Here, we suggest the main characters for the proposal of new species and consequent allocation in groups of Pintalia ( Table 1 ):

• Vertex with two transverse carinae (apical and subapical) and two compartments (apical and basal) present (1) or absent (0) ( AP).

• Vertex with apical transverse carina and subapical carina elevated (2), not elevated (1), or only apical transverse carinae elevated (0) ( CV).

• Apical transverse carina in frontal view concave medially (2), straight (1), medially elongated (0) ( AT).

• Median carina of the basal compartment developed (1) or undeveloped (0) ( MC).

• Frons with lateral carinae elevated (1) or no elevated (0) ( FR).

• Frontoclypeal suture semicircular bent forwards (1) or straight bent forwards (0) ( FS).

• Median carina touching the frontal ocellus (1) or evanescent near the frontal ocellus (0) ( FM).

• Tegmina with RA+RP1 with punctual anastomosis (1) or anastomosis absent (0) ( TR).

• ScP+R well forward of fork CuA (3), slightly (2), in the same level (1), behind (0) ( TV).

• Hind wing with a petiolate anastomosis in MP3+4 + CuA1 distally forked, Y-type (1) or with complete anastomosis with distal fusion between MP3+4+CuA1, I-type (0) ( WA).

• Females with anal tube long (1) or short (0) ( FA).

• Hind tibia with thin setae (1) or platellae (0) on the 2 nd, tarsomere ( PS).

• Lateral spines in hind tibia present (1) or absent (0) ( LS).

• Anal tube distally developed (1), undeveloped (0) ( AS).

• Male anal tube in dorsal view with narrowing (lesser width) just near the base (1) or not narrowing (0) ( AD). • Paraproct distally developed (1), undeveloped (0) ( PP).

• Epiproct distally developed (1), undeveloped (0) ( EP).

• Ridge on the aedeagal shaft present (1), absent (0) ( VR).

• Flagellum spine-like (1) or flagellum without spines (0) ( FS).

• Pygofer with medioventral process cone shaped laterally, wide, in caudal view (1) or without cone-shaped process laterally (0) ( PY).

• Shape and size of the genital styles, well large (1), or moderately large (0) ( GS).

• Shape and size of the basal opening of the genital styles, larger and laterally concave (1) or small in a diamond shape (0) ( OG).

• Shape of the aedeagal shaft, tubular (1) tubular and apically larger (0) ( AF).

• Pregenital segment evanescent in the middle (2), moderately evanescent (1), not evanescent (0) ( PG).

Key to the groups and new species of Pintalia Stål, 1862

1. Male with long anal tube, developed, bent distally. Flagellum of the aedeagous with spines. Hind wing Y-type.......................................................................................... Pintalia painensis -group ( 8)

–. Males with short anal tube, not developed, bent distally. Flagellum without spines.................................. 2

2(1). Apical transverse carina of vertex concave in frontal view and larger than the subapical carina in dorsal view. Paraproct long, developed distally. Genital styles well expanded apically. Aedeagal shaft with ventral ridge developed. Females with small Xsegment. Hind wing I-type....................................................... Pintalia pictipennis- group ( 3)

–. Apical transverse carina of vertex straight in frontal view and smaller than the subapical carina in dorsal view. Paraproct short, not developed distally. Genital styles moderately to well expanded apically. Aedeagal shaft without ventral ridge. Females with X-segment long or moderately long. Hind wing Y-type................................ Pintalia constellaris- group ( 7)

3(2). Males with epiproct short and flattened dorsoventrally........................................................ 4

–. Males with epiproct distinct, short, and swollen dorsally or developed distally..................................... 5

4(3). Pygofer symmetrical bilaterally. Aedeagal shaft usually with three small ventral ridges laterally; in right lateral view the bifid spine with fork distant to the base; in left lateral view with curved spine large in C-shaped on the apex of the shaft............................................................................................... P. minuta sp. nov.

–. Pygofer asymmetrical bilaterally.Aedeagal shaft with a curved ventral ridge; in right lateral the bifid spine with fork near to the base; in left lateral view with a moderately large or large spine elongated on the apex of the shaft........ P. minima sp. nov.

5(3). Males with epiproct in cone-shaped developed distally. Paraproct rectangular and flattened dorsoventrally. Ventral ridge well developed and curved occurs in basal half. Apex of the aedeagal shaft with three long and thin spines, two on the right lateral and one in left lateral.............................................................. P. magnaepiprocti sp. nov.

–. Males with epiproct short and swollen dorsally. Paraproct concave with basal half-higher, and apical half lower. Aedeagal shaft with ventral ridge distinct occurs dorsally near to the base. Apex of the aedeagal shaft with two spines.................. 6

6(5). Pygofer symmetrical bilaterally. Anal tube flattened near to the base and rounded distally, with ventral process large and rectangular distally. In dorsal view, epiproct rounded apically. Ventral ridge composed of two pairs of spines the ones above smaller. In right lateral view the shaft of the aedeagus exhibit a wide spine near to the apex, and one other small spine no movable below, medially; in left lateral view a single small spine occurs apically........................ P. stali sp. nov.

–. Pygofer weakly bilaterally asymmetrical; anal tube short and wide, well tubular, with ventral process small and triangular distally. In dorsal view, the epiproct truncate apically. Ventral ridge form two lines serrated and curved to right lateral. In right lateral view the shaft of the aedeagus exhibits two wider spines near to the apex, one bifid; in left lateral view without spine..................................................................................... P. serratilis sp. nov.

7(2). Yellowish tegmina exhibits dark spot with hyaline spots. Pygofer narrow distally with cone-shaped process sloping upwards and apically rounded. Genital styles moderately wide distally................................................. .. 7

–. Pygofer wider distally; with cone-shaped process sloping upwards and apically sharp. Genital styles well wide distally. Aedeagus exhibits the three spines on the apex of the shaft: the other two spines in opposite sides of the shaft (d and e) small, and approx. the same size. Flagellum small, without process..................................... P. fennahi sp. nov.

8(1). Anal tube distally bent in approx. 45°. Pygofer laterally with distal/dorsal inclination very large and rounded P. muiri sp. nov.

–. Anal tube distally bent in approx. 90°. Pygofer laterally with distal/dorsal inclination weakly or moderately large and rounded ................................................................................................... 9

9(8). Aedeagal shaft with a larger spine in left lateral of the apex; in right lateral the apical spine small, below in the middle of the shaft occur a small spine; flagellum with four spines, one near to the base, and three apically.... P. montalvaniensis sp. nov.

–. Aedeagal shaft with a larger spine in right lateral of the apex.................................................. 10

10(9). Pygofer laterally with process rounded and bent apically. Aedeagus with four spines on the shaft, two on each side; in right lateral the larger spine occurs on the apex of the shaft; flagellum with three spines, one medially and two apically............................................................................................... P. dorensis sp. nov.

–. Pygofer without process laterally........................................................................ 11

11(10). Flagellum with a bifid movable spine medially and a spine short not mavable apically.Aedeagal shaft with three spines; in right lateral the long spine tubular and moderately wide, below near the middle of the shaft occur a spine moderately large.............................................................................................. P. lundi sp. nov.

–. Flagellum without bifid spine, but with three common short spines, one basally, one medially and one apically. Aedeagal shaft with four spines; in right lateral, the long spine flattened, wide and curved, below occurs two small spines with approx. the same size................................................................................. P. painensis sp. nov.

12(7). 2 nd tarsomere without. Aedeagus exhibits the three robust spines on the apex of the shaft: apically wiyh three large and robust spines, slightly curved (1 st, 2 nd, 3 rd) (a)(b)(c), occurs on the apex of the shaft, towards the base; 4 th spine straight (d), is slghtly bent medially, inserted little above of the center on the aedeagal shaft towards the base; in left lateral view, 5 th spine small (e), not movable, occurs near the middle of the shaft towards to the flagellum, with a basal ridge in “u”...... P. speciosa sp. nov.

–. 2 nd tarsomere with 1-3 platellae. Aedeagus exhibits the three fine spines on the apex of the shaft: apically wiyh three large and fine spines well curved, (1 st, 2 nd, 3 rd) (a)(b)(c), (a) in S-shapped; spine (b) straight, biger the others; spine (C) in hook-shapped; 4 th spine straight (d) is straight, inserted little above of the center on the aedeagal shaft towards the base; in left lateral view, 5 th spine small (e), not movable, occurs near the apex of the shaft towards to the flagellum....... P. constellaris ( Walker, 1858) .

Remarks. This key does not include the species described by Stål (1862) and Muir (1934) as it was not possible to evaluate the aedeagus of these species, and this structure is essential for species-level recognition in the genus Pintalia . Species such as P. pictipennis and P. stali sp. nov., P. constellaris and P. speciosa sp. nov., and all species of the Pintalia painensis -group highlight the need to use genitalia for identification at the species level, until it is possible to better understand what an intraspecific or interspecific variation is in the genus. With this publication we hope to find male specimens of the species described by Stål (1862) to add them to the key. The supplementary description of the species described by Muir (1934) may be useful to clarify the morphology of the male terminalia and thus also add them to the key. However, it has not yet been possible to evaluate all holotypes and paratypes deposited at BPBM and BMNH respectively.

Figures

FIGURE 1. Head Pintalia types Stål, 1862 (scale bar A–I, 1mm.): A, P. lateralis Stål, 1862; B, P. consobrina Stål, 1862; C, P. fasciatipennis Stål, 1862; D, P. fraterna Stål, 1862; E, P. inornata Stål, 1862; F, P. obscuripennis Stål, 1862; G, P. pictipennis Stål, 1862; H, P. proxima Stål, 1862; I, P. ustulata Stål, 1862.

FIGURE 2. Pintalia lateralis Stål, 1862: Head, A, ventral view; B, dorsal view; C, transition vertex to front; D, habitus female in right lateral view.

FIGURE 3. Pintalia lateralis Stål, 1862: A, habitus female in ventral view; B, posterior leg in ventral view.

FIGURE 12. A, Pintalia lateralis Stål, 1862 tegmina (forewings): abbreviations of the cell names (red letters), longitudinal veins (black capital letters) and crossveins (in italics); Hind wing (figures: B–D); Petiolate anastomosis in MP3+4 + CuA1 distally forked (Y-type), Pintalia painensis group = Blue, Pintalia constellaris group = Yellow, complete anastomosis between MP3+4 + CuA1 (I-type), Pintalia pictipennis group (Red).

FIGURE 14. Pintalia pictipennis Stål, 1862: Head, A, ventral view; B, dorsal view; C, transition vertex to frons; D, habitus male in left lateral view.