Pintaliini Metcalf, 1938

Hoch, Júlio César Do Carmo Vaz Santos 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 : 7-9

Tribe Pintaliini Metcalf, 1938

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).

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.