Latycephala viridula (Kuoh)

Latycephala viridula (Kuoh) View in CoL

Figs 1C–F View FIGURES 1 , 17 View FIGURES 17 BC, 20BH, 21A–D, 24A–H.

Petalocephala viridula Kuoh 1984: 274-275 View in CoL , fig 4a–d.

Latycephala viridula (Kuoh) View in CoL : Jones & Deitz 2009: 45.

Diagnosis. Pale brownish green mottled with yellow spots; crown narrowly margined by red. Aedeagus with subapical pair of forked processes, proximal branch directed anteriorly and then curved dorsally, distal branch stout, tapering distally, exceeding shaft apex and meeting its counterpart at midline much beyond membranous lobe-like apex of aedeagal shaft.

Description. Pale brownish green, mottled with pale yellow; head both dorsally and ventrally anteriorly margined by red in front of eyes, ocelli reddish, eyes brownish. Face, thoracic and abdominal sternites and pleurites yellowish green, antennae and legs greenish yellow.

Crown surface punctate, longitudinally rugose (especially adjacent to median carina) with well-developed median carina, about 0.7× (in male) to 0.9 (in female) as long medially as interocular distance, 0.8–09× as long medially as median length of pronotum. Pronotal lateral margins slightly divergent posteriorly, about 1.1–1.2× as wide across posterolateral angles as between anterolateral angles, medially with a faint impressed line, surface striate-punctate on disc and rest punctate. Mesonotum as long as or slightly longer than median length of pronotum, mesoscutum punctate, mesoscutellum transversely striate punctate. Female sternite VII with slightly convex anterior margin, about 0.4× as wide proximally as long medially, narrowed slightly posteriorly, posterior margin with lateral margins slightly convex, medially concave ( Fig. 20H View FIGURES 20 ).

Male genitalia. Pygofer in lateral view triangular with straight dorsal margin, narrowed posteriorly, much more so in distal region and sharply angled, 1.6× as long as height along anterior margin, ventral process slightly curved dorsoposteriorly, prong-like, widened distally and reaching dorsal margin of pygofer. Style anterior apodeme short, about 0.2× as long as total length, with broad mid region, apophysis slightly narrowed and curved ventrally almost at right angle near apex, ventrally directed portion slightly narrowed proximally and flared near apex with rounded ventral apical margin. Aedeagus short, with well-developed dorsal apodeme, shaft directed posterodorsally, dorsal surface in lateral view with convexity at about midlength, in proximal region and with one pair of bifurcate lateral subapical processes arising on dorsal surface laterally, proximal branch of process directed anteriorly and then curved dorsally, longer, stouter distal process tapering distally and directed dorsally and then mesally meeting its partner at midline much beyond apex of shaft; shaft apex membranous lobe-like with apical gonopore indistinct.

Female genitalia. Valvula I slightly curved dorsally, and slightly narrowed, sculptured area occupying distal 0.3 length, obliquely strigate, strigae short basally, occupying almost half-width in distal half-length of sculptured area ( Figs 21 View FIGURES 21 AB). Valvula II slightly curved dorsally, of uniform width, toothed area occupying distal 0.25 length, teeth about 8, small except proximal one ( Figs 21 View FIGURES 21 CD).

Measurements. Male10.0– 10.1mm long, 3.4–3.5 mm wide across eyes and 3.8–3.9mm wide across posterolateral angles of pronotum. Female 12.4 mm long, 3.7 mm wide across eyes and 3.9 mm wide across posterolateral angles of pronotum.

Material examined. INDIA: Arunachal Pradesh: 3♂, Hunli, 15.vii.2023, Ajay Kumar ; Manipur: 1♀, Senapati: Siang, Mong Jang , 28.ix.2013, sweep net, Yeshwanth, H.M. ( UASB) .

Remarks. Kuoh (1984) described this species from Yunnan Province, China. It is a new record for the Indian subcontinent. L. viridula resembles P. chlorocephala externally but differs in being mottled with pale yellow with the crown and face margined by red. Crown shape and coloration of this species also resemble those of female P. umbrosa but the lateral margin of the crown is much straighter in the latter species than in this species.

Genus Petalocephala Stål

Petalocephala Stål 1854: 251 View in CoL . Type species: Petalocephala bohemani Stål View in CoL , subsequent designation by Atkinson 1885.

Pachyledra Schumacher, 1912: 248 View in CoL . Type: Pachyledra kamerunensis Schumacher, 1912 View in CoL . Synonymised by Linnavuori 1972:214.

Eogypona Kirkaldy 1901: 38 View in CoL , Type species. Eogypona kirbyi Kirkaldy1901: 38 View in CoL . Synonymised by China 1926: 350.

Destinoides Cai & He 2000: 57 View in CoL . Type species Destinoides fasciata Cai & He 2000 View in CoL , a junior synonym of Ledra latifrons Walker 1851 View in CoL . Syn. nov.

Diagnosis. Sexually dimorphic, males with more or less pentagonal heads and females with slightly longer often conical or parabolic heads, and anteriorly foliaceous; usually longer than pronotum, more or less flat, rarely sloping laterally; with median carina often well-developed and prominent, sometimes weak or absent. Pronotum more or less flat, rarely convex (as in P. nigrilinea ). Male pygofer with short ventral process (absent in P. uniformis ), Style apex ventrally either smoothly or abruptly curved (often at right angle). Aedeagus without preatrium or atrial ventral surface not prolonged, dorsal apodeme well-developed, bifid distally, shaft tubular, usually apically membranous, with or without lateral or apical processes.

Description. Medium to large (5.5–19.0 mm long), green, rarely brown arboreal leafhoppers often marked with dark brown on head, thorax and forewings. Sexually dimorphic, males with more or less pentagonal heads and females with slightly longer often conical or parabolic heads, and anteriorly foliaceous; usually longer than pronotum, more or less flattish, rarely sloping laterally, punctate, with median carina often well-developed and prominent, sometimes weak or absent. Face horizontal, flat, sometimes upper part concave, lower part short, nearly semicircular in outline, clypellus parallel-sided, frontoclypeus narrow parallel sided in lower part, sharply triangularly tapering upwards in front of antennal pits. Lora elongate, genae flattish, antennal pits shallow. Ocelli on crown close to posterior margin, either as far apart from each other as from adjacent eye or closer to each other than to adjacent eye (in most males) or closer to adjacent eye than to each other (in most females). Pronotum flattish, punctate or especially on posterior half rugose punctate, with or without medially longitudinally finely impressed or carinate or not; lateral margins long, subacute, either parallel-sided or divergent posteriorly, anterior margin slightly convex, posterior margin medially concave, lateral posterolateral margins adjacent to mesonotum either straight or slightly concave. Mesonotum as long as or longer than pronotum. Forewings long, coriaceous, punctate, more richly and deeply in clavus, claval veins separate, not connected by cross vein, sometimes apically forked, subapical and apical cells with accessary veins. Metatibia not foliaceous, with 4–6 prominent setae on anterodorsal (AD) row; metabasitarsomere with 35 platellae ( Figs 20C–E View FIGURES 20 ).

Male genitalia. Pygofer more or less triangular, without macrosetae or anterior marginal apodemes, dorsal margin more or less straight, without processes, ventral margin sometimes expanded, with short or long ventral process (absent in P. uniformis Distant ) that may or may not reach dorsal margin. Subgenital plates with a few short setae. Style without preapical lobe, part anterior to articulation with connective short, posterior to articulation broad proximally and narrowed distally, subapically ventrally either smoothly or abruptly curved (often at right angle), portion beyond curvature either narrowed, sinuate or not, often sculptured and with or without basal group of setae. Connective T or V-shaped with anterior median lobe. Aedeagus without preatrium or atrial ventral surface not prolonged, dorsal apodeme well-developed, bifid distally, shaft tubular, usually apically membranous, with or without lateral or apical processes, gonopore apical or subapical, in latter case on dorsal surface. Segment X with ventroposterior margin prolonged posteriorly and membranous, without sclerotized processes.

Female sternite VII about as long as or longer than sternite VI, almost rectangular with posterior margin medially concave ( Figs 20I–N View FIGURES 20 ). Valvulae I and II well-sclerotized. Valvula I with dorsal sculpturing confined to distal 0.33–0.5 length, sculpturing obliquely strigate, strigae increasing in length distally in proximal half-length. Valvula II with toothed area occupying distal 0.33–0.5 length, with or without basal tooth before toothed area, the first tooth in toothed area prominent followed by smaller teeth, each tooth with crenulate or smooth margin ( Figs 21 View FIGURES 21 GH, KL, OP; 22CD, GH, KL).

Remarks. Species of Petalocephala resemble those of Yelahanka Viraktamath, Webb & Yeshwanth , but differ in having a much longer crown (slightly to considerably longer than half interocular width). Yelahanka has its forewing deflected ventrally at level of outer claval vein which is carinate and also the claval veins are fused for some distance or connected by a cross vein and the metafemur distal macrosetae are 2+0 compared to 2+ 1 in Petalocephala .

Jones & Deitz (2009) removed the genus Eogypona from Ledrinae . China (1926) discussed in detail regarding the synonymy of Eogypona Kirkaldy with Petalocephala , which was earlier mentioned by Kirkaldy (1907) as “…apparently a synonym of Camptelasmus (= Petalocephala )”. The genus Petalocephala includes 89 valid species ( Jones & Deitz 2009), of which 23 were listed from the Indian subcontinent by Metcalf (1962). Of these, P. bainbriggei Distant , P granulosa Distant and P. tabulata Distant were transferred to the genus Yelahanka by Viraktamath et al. (2021); the remaining 20 species remain in the genus. In this study two new species records are given for the subcontinent and three new species are described. P. hornei Distant 1908 is treated as a junior synonym of P. chlorocephala ( Walker 1851) following Distant (1908).

Cai & He (2000) established the genus Destinoides with Destinoides fasciata Cai & He as the type species. Jones & Deitz (2009) in their revision of the world Ledrinae , suspected this genus to be synonymous with Petalocephala . However, Sun et al. (2014) revised Destinoides , synonymized the type species as a junior synonym of Petalocephala latifrons (Walker) and also transferred P. conspicua to this genus. Wang et al. (2024) in their molecular phylogenetic analysis included two examplars, both identified as Destinoides conspicua , ( P. conspicua 1 and P. conspicua 2). These workers did not illustrate either the habitus or male genitalia of these taxa, but in their key to Ledrine genera separated Destinoides from Petalocephala ( Wang et al. 2024: 11; couplet 33) by the “aedeagal shaft broad and laminate,” compared to the “narrow and cylindrical” aedeagal shaft of Petalocephala . The aedeagus of holotype male D. latifrons (Walker) ( Sun et al. 2014: Figs 8 View FIGURES 8 , 9 View FIGURES 9 ) and syntype male of D. conspicua Distant ( Sun et al. 2014: Figs 17 View FIGURES 17 , 18 View FIGURES 18 ) show that the aedeagus in both the species is neither broad nor lamellate. Therefore, apparently Wang et al. (2024) misidentified D. conspicua . The male genitalia of these two species of Destinoides are very similar to some species of Petalocephala and the shape of aedeagus is within the range of variation found in Petalocephala and hence, we consider Destinoides to be a junior synonym of Petalocephala . See also Discussion.

Key to species of Petalocephala View in CoL of the Indian subcontinent

[Note. Males of bicolor View in CoL , cephalotes View in CoL , confusa View in CoL , conica View in CoL , cultellifera View in CoL , glauca View in CoL , hearsayi View in CoL , insignis View in CoL , kempi View in CoL , subaquila View in CoL , and umbrosa View in CoL are not known. Similarly, females of chlorocephala View in CoL , nauniensis , nigrilinea View in CoL and remota View in CoL are not known. The female crown is variable in length within a species and sternite VII is more or less uniform in most of the species hence the couplets for females in the following key are only tentative until males of the species are discovered.]

1. Upper surface of body and forewings chocolate brown; crown, pronotum and mesonotum with bright yellow median stripe, narrowing anteriorly on crown ( Figs 8 View FIGURES 8 GK)................................................. P. perductalis (Kirby) View in CoL

- Virescent or yellowish green or greenish brown without such a stripe, margins of crown, pronotum and mesonotum may be fuscous to dark brown................................................................................. 2

2. Crown median length 1.2× or more as long as interocular distance ( Figs 1 View FIGURES 1 GI, 7AB, 10AB)........................... 3

- Crown median length less than 1.1× as long as interocular distance ( Figs 2A View FIGURES 2 , 3A View FIGURES 3 ,).................................. 5

3. Crown median length 2× or more as long as pronotal median length, pronotal lateral margins more or less parallel sided ( Fig. 7B View FIGURES 7 ) (female).............................................................................. P. kempi Pruthi View in CoL

- Crown median length 1.7× or less as long as pronotal median length, pronotal lateral margins distinctly divergent posteriorly ( Figs 1 View FIGURES 1 GI,10AB)...................................................................................... 4

4. Pronotum longer than exposed mesonotum ( Fig. 10A View FIGURES 10 ) (female).................................. P. subaquila Distant View in CoL

- Pronotum shorter than exposed mesonotum in both sexes ( Figs 1 View FIGURES 1 GI)........................ P. balehonnurensis sp. nov.

5. Females............................................................................................. 6

- Males............................................................................................. 18

6. Frontoclypeus black ( Fig. 18A View FIGURES 18 ), forewing subcostal area with dark brown longitudinal stripe extending for basal 0.66 length ( Fig. 6I View FIGURES 6 )................................................................................ P. insignis Distant View in CoL

- Frontoclypeus and forewing costal area not marked as above.................................................. 7

7. Crown about as long as interocular distance (about 0.91–1.1× as long as interocular distance)......................... 8

- Crown distinctly shorter than interocular width (less than 0.9× as long as interocular distance)....................... 13

8. Small species measuring less than 9mm long ( Fig. 9E–I View FIGURES 9 )..................................... P. sahyadrica sp. nov.

- Larger species measuring more than 10 mm long............................................................ 9

9. Crown apex conically rounded or parabolic ( Figs 3I View FIGURES 3 , 4 View FIGURES 4 AD)................................................... 10

- Crown more or less an equilateral triangle, with apex acutely angled ( Figs 2A View FIGURES 2 , 6A View FIGURES 6 )................................ 11

10. Large species measuring 15–17 mm long...................................................... P. conica Distant View in CoL

- Smaller species measuring 11 mm long....................................................... P. confusa Distant View in CoL

11. Crown and pronotum margined with dark brown ( Figs 2A–C View FIGURES 2 ); forewing apical margin dark brown ( Fig. 2A View FIGURES 2 ).................................................................................................... P. bicolor Distant View in CoL

- Crown and pronotum with margin not dark brown ( Figs 6 View FIGURES 6 AB); forewing apical margin either differently colored or not... 12

12. Pronotal lateral margins more or less parallel sided (distance between posterolateral angles less than 1.1× as long as that between anterolateral angles) ( Fig. 6A View FIGURES 6 )..................................................... P. glauca (Melichar) View in CoL

- Pronotal lateral margins distinctly diverging posteriorly (distance between posterolateral angles more than 1.2× as that of anterolateral angles) ( Fig. 6E View FIGURES 6 )............................................................. P. hearsayi Distant View in CoL

13. Large species measuring more than 17 mm long............................................ P. cultellifera (Walker) View in CoL

- Smaller species measuring less 16 mm long............................................................... 14

14. Lateral margins of crown in front of eyes almost straight for almost half length of crown then mesally curved ( Fig. 2E View FIGURES 2 )........................................................................................ P. cephalotes Distant View in CoL

- Lateral margins of crown in front of eyes mesally directed, apex subangulate or conically rounded ( Figs 10E View FIGURES 10 )........... 15

15. Head in front of eyes both dorsally and ventrally reddish brown margined ( Figs10 View FIGURES 10 EF)................. P. umbrosa Distant View in CoL

- Head in front of eyes of same color as rest of crown or face ( Fig. 11 View FIGURES 11 DE)........................................ 16

16. Sternite VII about 1.33× as wide basally as at apex, posterior margin shallowly concave ( Figs.20 View FIGURES 20 MN)... P. uniformis Distant View in CoL

- Sternite VII less than 1.2× as wide at base as at apex, posterior margin medially concavely excavated ( Figs 20 View FIGURES 20 JK)....... 17

17. Valvula II with proximal prominent tooth proximad of regular toothed area ( Figs 21 View FIGURES 21 KL)..... P. fuscomarginata Cai & Kuoh View in CoL

- Valvula II without proximal prominent tooth proximad of regular toothed area ( Figs 21 View FIGURES 21 OP)............ P. latifrons (Walker) View in CoL

18. Pygofer without process on ventral or outer margin posteriorly ( Fig. 33 View FIGURES 33 AD); aedeagal shaft with a sclerotized angulate plate on dorsal surface apically, and subapical lateral membranous lobes not exceeding plate ( Figs 33 View FIGURES 33 FGH)...... P. uniformis Distant View in CoL

- Pygofer armed with ventral, apical or dorsoposterior process ( Figs 25 View FIGURES 25 BC, 32A); aedeagal shaft without sclerotized plate on dorsal surface ( Fig. 31I View FIGURES 31 ).............................................................................. 19

19. Style apex abruptly ventrally directed, dorsal margin at point of bend almost at right angle ( Figs 25D View FIGURES 25 . 27C View FIGURES 27 ); aedeagal shaft without median sclerotized process on ventral surface apically................................................ 20

- Style apex slightly curved at apex ( Fig. 32D View FIGURES 32 ); aedeagal shaft with median sclerotized process on ventral surface apically ( Figs 32 View FIGURES 32 FH)............................................................................. P. sahyadrica sp. nov.

20. Pygofer posteriorly strongly narrowed and acutely angular distally ( Figs 26 View FIGURES 26 AB)................................... 21

- Pygofer gradually slightly narrowed distally and posterodorsal angle rounded ( Figs 25A View FIGURES 25 , 29 View FIGURES 29 AB)..................... 23

21. Aedeagal shaft in lateral view sinuate ( Figs 26I View FIGURES 26 , 27G View FIGURES 27 ); style apex without subapical tooth ( Figs 26 View FIGURES 26 DE)................. 22

- Aedeagal shaft in lateral view slightly curved, not sinuate ( Fig. 28G View FIGURES 28 ); style apex with subapical tooth ( Figs 28 View FIGURES 28 CD)............................................................................................. P. latifrons (Walker) View in CoL

22. Aedeagal shaft with one pair of short thin processes at base of dorsal expansion in lateral view ( Fig. 26H View FIGURES 26 ); pygofer ventral process lamellate and short ( Figs 26A–C View FIGURES 26 ).............................................. P. chlorocephala (Walker) View in CoL

- Aedeagal shaft in lateral view without processes at base of dorsal expansion ( Fig. 27G View FIGURES 27 ); pygofer process stout and long ( Figs 27 View FIGURES 27 AB)...................................................................... P. fuscomarginata Cai & Kuoh View in CoL

23. Aedeagal shaft with subapical pair of lateral spine-like processes exceeding shaft apex ( Fig. 29E–G View FIGURES 29 ).................. 24

- Aedeagal shaft with two pairs of subapical lateral processes, the proximal pair short, and the distal pair membranous medially convergent beyond shaft apex ( Figs 31G–I View FIGURES 31 ).................................................. P. remota (Melichar) View in CoL

24. Aedeagal shaft in proximal 0.75 length of uniform width dorsal surface without angular projection ( Fig. 29G View FIGURES 29 ); style apex with ventrally bent part sinuate ( Figs 29 View FIGURES 29 CD).................................................... P. nauniensis sp. nov.

- Aedeagal shaft broad proximally then narrowed till midlength with an angular projection on dorsal surface slightly distad of midlength ( Fig. 30D View FIGURES 30 ); style apex with bent part triangular in lateral view ( Fig. 30B View FIGURES 30 )................ P. nigrilinea (Walker) View in CoL