Pintalia painensis, Hoch & Bartlett & Ferreira, 2025

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 : 13-17

Pintalia painensis -group

Diagnosis. This group can be distinguished mainly by characters of the male terminalia. However, some other characters are also useful for identification. Vertex moderately large, with apical transverse carina smaller than the subapical carina; apical compartment large, wider than long, moderately long medially; caudal margin triangular, slightly concave laterally with a small basal emargination; in frontal view with apical transverse carina straight. Frons approx. 1.9–2.4 times wider in the largest width than at the apically; frontoclypeal suture bent upwards medially, laterally straight. Tegmina (forewings) with pt large and sub rectangular, weakly, or well angled distally; fork of ScP+RA and RP moderately closer to bc than fork CuA1 and CuA2; rm-1 proximally, together, or distally to first MP fork; mcu-1 distally to first MP fork. Hind wing with RP bifid, MP trifid with petiolate anastomosis in MP3+4 + CuA1 distally forked (Y-type), CuA1 and CuA2 bifid. In ventral view, males with segment viii, complete, or separated medially. Hind tibia with 3–4 lateral spines, spines closest to the femur well small; 6 apical teeth as provided for the genus. 1 st tarsomere with 7 apical teeth. 2 nd tarsomere with 8 apical teeth, and with 1–3 platellae, in some species absent or in a smaller number on one of the legs (asymmetric bilaterally).

Male terminalia. Pygofer bilaterally symmetrical; in lateral view, usually without lateral process, when present, laterally flattened (thin) vs. broad in Pintalia constellaris -group; dorsal margin concave, caudal margin usually straight, in some species slightly irregular. in ventral view, ventromedian process triangular, usually as wide as long. Anal tube long, tubular dorsally and excavated ventrally, developed and flattened distally with apex truncate and irregular, usually straight laterally and slightly asymmetrical in most species; in dorsal view, occurs a narrowing near the base but gradually elongates again to the apex; epiproct and paraproct short; in dorsal view epiproct triangular, and laterally with two small bent down margin; paraproct rounded, narrow laterally and slightly larger than the epiproct distally. Genital styles moderately or weakly wide distally, excavated ventrally and with small bent margins to the ventral region; basal opening between the genital styles, in diamond-shaped usually moderately small. Aedeagus display 5–7 spines; aedeagal shaft tubular, and usually wider apically near to flagellum; aedeagal shaft with 3–4 spines; usually a long or wider spine occurs laterally on the apex of the aedeagal shaft; ventral ridge absent. Flagellum usually with 2–6 spines in right laterally.

Female terminalia. Anal tube long, in dorsal view rounded medially, with apex truncated; in lateral view with lateral margin slightly projected and straight, ventrally produced two third basal and a third apical excavated and apically produced again; epiproct and paraproct thin and short; epiproct triangular in dorsal view with lateral margin near to the base slightly developed and bent down; paraproct in dorsal view rounded. Gonocoxae VIII truncated, usually with rounded lateral carinae and irregular; in caudal view, it shows straight carinas moderately distant from the margin, closer to the genital segment, and distant medially or close to the ovipositor. Ovipositor large, in lateral view, sabre-shaped, curved upwards dorsally surpassing the anal tube.

Species here assigned to this group: P. painensis sp. nov., P. lundi sp. nov., P. montalvaniensis sp. nov., P. dorensis sp. nov., P. muiri sp. nov.

Distribution. BRA. Minas Gerais State ( Fig. 77 ).

Notes. Fennah (1945) grouped some species due to the anal tube developed distally, bent down in approx. 90° (e.g., P. bicaudata- group), bent in approx. 45° (e.g., P. albolineata- group), and to the shape of the genital styles. Here we have grouped all the new species with distally developed anal tube, with spines on the flagellum and genital styles weakly or moderately wide distally. A review of the species described by Muir 1934, and Fennah 1945 is indicated to properly allocate them to groups or new genera. Some species deposited in ISLA may have a morphology very similar to the morphology described for Pintalia painensis- group, however the flagellum does not present spines.

Morphology. Body length: 3.9–5.3 ♂ mm (n=18). 5.0– 5.8 mm ♀ (n=6).

Head. Vertex with apical compartment Square and large, in dorsal view with apical transverse carina usually almost straight and slightly irregular, totally developed ( P. painensis sp. nov., P. lundi sp. nov., P. montalvaniensis sp. nov.), or slightly evanescent laterally ( P. dorensis sp. nov., P. muiri sp. nov.); the subapical carina can be concave ( P. dorensis sp. nov., P. muiri sp. nov.) or bent medially ( P. painensis sp. nov., P. lundi sp. nov., P. montalvaniensis sp. nov.); approx. 1.9–2.4 times wider in the largest width than apically.

Thorax. Pronotum narrow, inconspicuous median carina and moderately developed but evanescent submedian carina behind the eyes. Tegmina (forewings) mostly hyaline with a dark brown spots, with four rounded spots in nodal region ( P. painensis sp. nov., P. montalvaniensis sp. nov.) or with one irregular spot ( P. dorensis sp. nov., P. muiri sp. nov.) or two irregular spots ( P. lundi sp. nov.); rm-1 ahead to first fork MP ( P. painensis sp. nov.)

together ( P. dorensis sp. nov., P. lundi sp. nov., P. muiri sp. nov.) or distally ( P. montalvaniensis sp. nov.) and mcu- 1 occurred distally to first MP fork ( P. dorensis sp. nov., P. lundi sp. nov., P. muiri sp. nov., P. montalvaniensis sp. nov.). Posterior legs, hind tibia with three lateral spines ( P. dorensis sp. nov., P. lundi sp. nov.) or four lateral spines ( P. painensis sp. nov., P. muiri sp. nov., P. montalvaniensis sp. nov.); 1 st tarsomere with two apical teeth external slightly larger, and two medially with approx. the same size; 2 nd tarsomere with 1–2 lateral apical teeth well larger ( P. painensis sp. nov., P. dorensis sp. nov., P. lundi sp. nov.) or slightly larger ( P. muiri sp. nov., P. montalvaniensis sp. nov.) and with 1–3 platellae.

Male terminalia. Pygofer bilaterally symmetric, laterally with a developed process, rounded and thin ( P. dorensis sp. nov.), very large and rounded ( P. muiri sp. nov.) or moderately rounded ( P. painensis sp. nov., P. lundi sp. nov., P. montalvaniensis sp. nov.). Anal tube in dorsal view, very narrow near to the base ( P. dorensis sp. nov., P. painensis sp. nov.) or slightly narrow ( P. lundi sp. nov., P. montalvaniensis sp. nov., P. muiri sp. nov.). Genital styles moderately wide distally ( P. dorensis sp. nov., P. montalvaniensis sp. nov.) or weakly wide distally ( P. painensis sp. nov., P. lundi sp. nov., P. muiri sp. nov.); basal opening of styles usually moderately small and diamond-shaped ( P. painensis sp. nov., P. lundi sp. nov., P. dorensis sp. nov., P. montalvaniensis sp. nov.) or weakly distinct, slightly rounded laterally and triangular elongated apically ( P. muiri sp. nov.). Aedeagus with a large spine on the right side of the apex of aedeagal shaft( P. painensis sp. nov., P. lundi sp. nov., P. muiri sp. nov., P. dorensis sp. nov.) or a large spine on the left lateral of the apex of aedeagal shaft( P. montalvaniensis sp. nov.); in left lateral view with one spine ( P. painensis sp. nov., P. lundi sp. nov., P. muiri sp. nov., P. montalvaniensis sp. nov.) or two spines ( P. dorensis sp. nov.); in right lateral view with one short spine ( P. lundi sp. nov., P. montalvaniensis sp. nov., P. dorensis sp. nov.) or two short spines ( P. painensis sp. nov.). Flagellum with two spines ( P. muiri sp. nov., P. lundi sp. nov.), one of them bifid ( P. lundi sp. nov.), with three spines ( P. painensis sp. nov., P. dorensis sp. nov.) or four spines ( P. montalvaniensis sp. nov.).

Female terminalia. Female terminalia very similar between species with little change in size of the anal tube and gonocoxae VIII.

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.

Checklist of species of Pintalia Stål, 1862 from Brazil

Table 2: See the appendix.

Amended descriptions

Figures

FIGURE 77. Distribution of Pintalia-groups in South America (A). Distribution of Pintalia painensis-group (B). Distribution of Pintalia pictipennis-group (C). Distribution of Pintalia constellaris-group (D). The extra records (not reported in the type and/or material examined) of Pintalia constellaris-group were obtained through the INaturalist platform (last access 25.x.2023), Pintalia constellaris available in: https://www.inaturalist.org/taxa/895204-Pintalia-constellaris.