Proekes Theron

Figs 22–30, 33

Deltocephalus cephaleus Naudé, 1926: 53 .

Proekes cephaleus (Naudé, 1926); Theron, 1975: 190, 191, 204–205.

Type species: Deltocephalus cephaleus Naudé, 1926 .

Revised generic diagnosis and description

Diagnosis.

1. Medium leafhopper; male and female: length from apex of crown to apex of tegmina, 3.7–4.3 mm; crown length medially 0.7–0.8 mm; crown angle 82–87°.

2. Dorsum with head in male and female with brown to light brown longitudinal line from apex of coronal suture to posterior margin of head; apex of crown with narrow, longitudinal paired light brown marking; lateral margin of crown with about three dorsally extended horizontal brown arcs on face; pronotum with single, amorphous to round marking near lateral margin (Figs 22A–D, H, J, K, L, 27A–C, E, G).

Terminalia in male.

3. Valve crescentic (Fig. 23D) or narrowly rectangular (Figs 23K, 30C), length/width=0.22.

4. Pygofer deeply incised in dorsal view; pygofer lobe in lateral view acutely triangular, mediodorsal margin with straight or sublinear, edentate, acuminate process, orientation posteriad; 2–4 macrosetae, (length 40–70 µm) (Figs 23C, J, R, S, 30A, B).

5. Segment 10 large, conical or dolioform, (0.5–0.6 times as wide as width across pygofer) (Figs 23C, J, R, 28A, 30A, B), reaching apex of pygofer lobe.

6. Connective wider than long (greatest length/greatest width=0.5–0.7), stem short, wide, arms slightly shorter to much longer than stem (length arm/length stem=0.9–2.0).

7. Aedeagus with base or subbase of anterior margin of shaft 1–2 pairs of symmetrical to asymmetrical teeth; shaft compressed or tubular; dorsal apodeme commonly reduced and preatrium short, rarely about half as long as shaft and preatrium reduced.

Terminalia in female.

8. Sternite 7 trapezoid, posterior margin sublinear (Figs 22J, 24A) or shallowly concave or medially narrowly convex (Fig. 30I); greatest length/greatest width=0.47–0.57; width across apex/width across base=0.59–0.76; medial length/average lateral length=0.95–1.07.

9. Valvula 2 minutely denticulate on subapex of mediodorsal margin (Figs 24M, 25A, D, E, 29H) and submarginal lateral pore-like structures along most of forked apex (Figs 23N, O, 25B, C).

10. Valvula 3 with unique long extension; laterally with many short macrosetae apically and subapically on lateral surface (Figs 24P, 29C, D), length 23–35 µm, thin, 2–3µm, rarely thicker, 3–4 µm; 0.48–0.53 times longer than greatest length of pygofer in lateral view (Figs 22F–J, 24J).

11. Valvifer 2 sculpture pore-like and microtrichia (Figs 24E, F, 29B).

Etymology. Pro, Latin and Greek, before, forward, in front of; - eces, -ekes, Greek, suffix meaning sharp; for the acute angle of the crown and face. Gender feminine.

Description.

Male, female and nymph.

Color. Ground color ochraceous in male (Figs 22A, B, L, 27A, B, E, F), female (Figs 22C, D, J, K, 27C, D, G, H), with light brown to brown markings. Nymph in Fig. 22E, F, stramineous to yellow laterally, light brown medially, light ochraceous, narrow medial line. Nymph in Fig. 22M collected with adults, but probably not an associated species.

Head. Brown arcs of clypeus extended into lateral margin of crown, light brown to brown longitudinal line over coronal suture, distad to apex of crown variable; crown with distinct, paired, parallel light brown line or weakly developed; base of head, between eye and coronal suture with short, elongate, paired brown mark (Figs 22D, L, H, 27C, E, G) or feint to absent (Figs 22A, B, 27A).

Face. Numerous horizontal brown arcs on clypeus, extending onto margin of head (Figs 22H, I, 27D, F, H).

Pronotum. Ground color ochraceous, with paired brown marking laterally and variable amorphous markings (Figs 22A–D, J, L, H, 27A–C, E, G).

Tegmina. Ochraceous, with brown veins, apical cells distally embrowned, brown irroration in some cells (Figs 22A–D, J–L, 27A–H).

Hind wing. Greyish-brown opaque or brown translucent distally, yellowish translucent basally and costal cells, veins brown.

Morphology.

Head. Wider than pronotum (male and female head width/pronotum width=1.11–1.16; crown angle in male and female 82–87°; crown length/crown length next to eye in male and female=1.65–1.77); crown length/pronotum length=1.65–1.80.

Face. Clypellus extended beyond posterior margin of gena, margin below lorum narrow, gena with notch below eye (Fig. 22L).

Ocellus. Male and female ocellus diameter/interocular distance=0.34–0.46.

Pronotum. Lateral margin short, non-carinate, pronotum length/pronotum width=0.35–0.37.

Tegmina. Appendix absent. Male and female, length/width=2.34–2.61; four apical cells; 2–3 anteapical cells.

Hind wing. Reduced, elongate, shorter and narrower than tegmina, male and female, length/width=3.59–4.35.

Tegmina-hind wing relation. Ratio of length tegmina/length hind wing=1.13–1.22; width tegmina/width hind wing=1.41–2.11.

Chaetotaxy. Profemur row AV with 9–11 short, wide setae, row IC 8–10 long, fine setae, AV 1 single, AM 1 single. Spinulation of protibia 1+4, mesotibia 4+4, metafemur apex 2+2+1. Metatarsus 1 apical plantar setae include four (rarely 5) rounded platellae and one acute seta; metatarsus 2 apex with one acute, two rounded and one acute platellae.

Measurements.

Male (n=129). Apex of crown to apex of tegmina 3.75–4.31 mm; apex of crown to apex of abdomen 3.62–4.23 mm; crown length 0.74–0.83 mm; crown length next to eyes 0.43–0.48 mm; pronotum length 0.43–0.48 mm; head width 1.37–1.50 mm; pronotum width 1.20–1.34 mm; ocellus diameter 25–31 µm; interocular distance 65–80 µm; crown angle 83–87°; crown length/crown length next to eye=1.67–1.77; head width/pronotum width=1.10–1.16; ocellus diameter/interocular distance=0.34–0.44; crown length/pronotum length=1.65–1.80; pronotum length/ pronotum width=0.35–0.37; length from apex of crown to apex of tegmina/length from apex of crown to apex of abdomen=0.98–1.07.

Female (n=92). Apex of crown to apex of tegmina 3.68–4.20 mm; apex of crown to apex of abdomen 4.35–5.05 mm; crown length 0.75–0.85 mm; crown length next to eyes 0.44–0.50 mm; pronotum length 0.43–0.48 mm; head width 1.38–1.52 mm; pronotum width 1.20–1.34 mm; ocellus diameter 25–32 µm; interocular distance 62–76 µm; crown angle 82–87°; crown length/crown length next to eye=1.63–1.76; head width/pronotum width=1.12–1.16; ocellus diameter/interocular distance=0.36–0.47; crown length/pronotum length=1.68–1.48; pronotum length/ pronotum width=0.35–0.37; length from apex of crown to apex of tegmina/length from apex of crown to apex of abdomen=0.80–0.87.

Terminalia.

Male.

Segment 10. Large, slightly depressed, deeply incised into base of pygofer, narrowly conical in dorsal view, lateral margins straight, convex in P. diacaphalis (Fig. 30B), 1.05–1.32 times longer than wide (Figs 23C, J, R, 28A, 30A, B); greatest width at base/greatest width at apex=1.49–1.85; greatest width across pygofer/greatest width of segment 10=0.51–0.59. Ventrally with medial, membranous recess (Figs 23R, 28A, 30A, B). Apex almost reaching apex of pygofer lobe.

Pygofer. In lateral view triangular, dorsal view rectangular; basal suture right-angled (Figs 23C, 30A) or short, straight (Fig. 23R); anterior margin with short or elongate apodeme, rarely without apodeme. Pygofer lobe acutely triangular in lateral view, apex membranous, tumid, semicircular, subapically with median, ovoid lobe (Fig. 23S), base of phragma attached medioventrally, with microtrichia and short setae; dorsomedially with straight to slightly curvate, edentate process, mediad of apical, membranous lobe (Fig. 23J), protruding slightly or recessed behind apex of lobe; 2–4 short macrosetae (40–70 µm long) laterad of origin of process (Fig. 23S).

Subgenital plate. Lateral margin curvate, medial margin straight to sublinear, narrowly divergent, length/ width=1.31–1.60; macrosetae variable in position and number, 19–25 (Fig. 23D, K), in distal three quarters, 13–19 in distal half, or 7–9 subapically; length 100–175 µm. Style apex extend half-way to three quarters into subgenital plate. Dorsomedial ridge articulated with style.

Valve. Crescentic (Fig. 23D) to narrowly rectangular (Fig. 23K, 30C); length/width=0.18–0.22.

Style. Narrow, elongate, greatest length/greatest average width=5.10–5.87, length apophysis/greatest length=0.16–0.20; apophysis with fine, ventral teeth (Fig. 30K, L), preapical lobe rounded (Figs 26E, 30H) or right-angled (Fig. 23G), anterior lateral lobe produced anteriad, about as wide or wider than median width of style, anterior medial lobe short Fig. 23G in Theron (1975), Figs 26E, 30H.

Connective. Y-shaped, transverse, short, wide stem, commonly articulated to preatrium. Width stem/width arms=0.5–0.6; length arms/length stem=0.9–2.0; angle of arms 78–106°; greatest length/greatest width=0.5–0.7; length stem/width greatest=0.2–0.3; Fig. 23H in Theron (1975), Figs 26F, 28E with differentiation of sclerotized and de-sclerotized arms. Rarely fused to atrium, arms Y-shaped (Fig. 30E, J).

Aedeagus. Aedeagal shaft in lateral view, strongly to weakly compressed (medial width laterally/medial width in anterior or posterior view=0.99–2.79). Dorsal apodeme commonly reduced (Figs 23E, L, M, O, 26A, C, 28B, C), with membranous lobes (Fig. 28B, C, G, H) or 0.21–0.42 times as long as greatest length of aedeagus (Fig. 30D, E). Preatrium variable, short (Figs 23E, L, 26A, C) to very short (Figs 23M, 28B), 0.21–0.43 times as long as greatest width across base of aedeagus, or absent (Fig. 30D). Anterior margin of shaft with variable, apical and basal teeth. Anterior apical teeth short (Figs 23E, F, L–Q, 26A, B), reduced (Fig. 26C) or elongate (Figs 28B–D, F–I, 30D–F). Anterior subbasal teeth as one pair (Figs 23E, F, L, 28B, C, G), two basal, paired, symmetrical teeth (Fig. 23M–P) or asymmetrical teeth (Fig. 26A–D). Posterior margin of shaft with short, apical and median paired teeth (Figs 23E, F, L–Q, 26A–D), or median paired teeth long (Figs 28G–I, 30D–F). Gonopore from subapex of anterior margin to subapex of posterior margin, narrow anteriorly, wide with rounded base posteriorly (Figs 23F, N, P, Q, 26B, D, 28C, D, 30E, F).

Female.

Genital capsule. Ovipositor protruding beyond posterior margin (Figs 22G, 24J), length from base of pygofer to apex of V3/length of pygofer=1.47–1.53.

Sternite 7. Trapezoid; posterior margin sublinear, lateral margins straight or curvate (Fig. 24A) or with slight median projection (Fig. 30I); medial length/average lateral length=0.95–1.07; length/width=0.47–0.57; width across apex/width across base=0.59–0.67.

Valvula 1. Lanceolate (Fig. 24H), sculpture laterally imbricate apically (Figs 24K, 25F), medially strigate marginally, imbricate basally (Figs 24L, 29F); sculpture medially finely anastomosing, striate (Fig. 29E).

Valvula 2. Distal half slightly wider than base, margin edentate or minutely denticulate on subapical medial margin (Figs 24M, 25A, D, E, 29H), marginal to submarginal pore-like structures (Figs 24N, O, 25B, C), distal third forked (Fig. 24I), ratio length to join in forked apex/greatest length=0.30–0.38; sculpture laterally as in Fig. 29G, medially as in Fig. 29H.

Valvula 3. Elongate, length/width=6.01–7.62; apex and subapex with numerous (n=32–45) short, length 23–35 µm, thin (2–3 µm), rarely thicker (3–4 µm) setae (Figs 24P, 29C, D).

Valvifer 1. Longer than wide (length/width=1.42–1.92), variable dimensions between left and right valvifers, shape of margins variable (Fig. 24B, C), fused dorsally (Fig. 24G).

Valvifer 2. 2–3 times longer than wide (Figs 24D, 29A), (length/width=2.57–3.00), sculpture as in Figs 24E, F, 25H, I, 29B, with triangular microtrichia and up to eight circular, pore-like structures.

Remarks. Proekes cephaleus and three new species described below are recognized as distinct by a combination of external and internal male and female features as in the diagnosis above and species diagnoses below. Proekes is generally larger than the three other genera described here, e.g., length from apex of crown to apex of tegmina 3.72–4.26 mm and in the female the prolongation of the valvula 3 well beyond the posterior margin of the pygofer lobe. Notable is the range in shape of the aedeagal shaft, that is strongly compressed, weakly compressed or tubular, and a possible association between long and short teeth on the shaft and the degree of compression of the shaft, i.e., short teeth in the compressed shaft and long teeth in the tubular shaft. Further, the relation of the reduction or extension of the dorsal apodeme and preatrium respectively, applied here, as in Colistra . The dorsal apodeme is clearly reduced and preatrium relatively elongated in P. cephaleus (Naudé) and P. hemiplatyphalis sp. n., but somewhat developed in P. tetracaphalis sp. n. and the dorsal apodeme well developed and preatrium reduced in P. diacaphalis sp. n. The symmetry and asymmetry of 1–2 pairs of teeth at the base or subbase of the anterior margin of the aedeagal shaft is somewhat comparable to the consistent, single pair in all species of Proekoides . The single male from Clanwilliam included in P. tetracaphalis sp. n. and specimens of P. cephaleus from Verlorenvlei suggest possible additional species. The orientation of the shaft at rest in the genital capsule is considered as vertical, with the apex of the shaft at rest in the membranous recess in the segment 10. Caudal examination of whole specimens occasionally corroborated this position. During maceration the shaft is dislodged and even if pushed back, seemed to lift the anal tube, or return to a more horizontal orientation. The pygofer process in Proekes is consistently edentate and sublinear in dorsal and lateral views, in Colistra it is sublinear and toothed ventrally and in Proekoides concave and toothed dorsally. The connective and style resemble that of Colistra and Proekoides . In Proekes the connective is transversely rectangular, with ratio greatest length/greatest width=0.40–0.65; Colistra = 0.50–0.65; Proekoides = 0.62–0.76; but more equilateral in Xhoreus (0.8–1.0). The subgenital plate and valve are similar to that of Colistra and Proekoides . The long extension of valvula 3 of the female ovipositor is unique in this genus and tribe. Modelled distribution of the combined records of four species in Fig. 33B.

Key to males of Proekes species

1 Aedeagal shaft compressed (median width in lateral view/median width in dorsal or ventral view=1.73–3.28), teeth on shaft short (much shorter than width of shaft in lateral view), preatrium short (preatrium length/length across base of aedeagus=0.23– 0.45)............................................................................................... 2

1`Aedeagal shaft tubular or weakly compressed, teeth on shaft long (as long as or longer than width of shaft in lateral view), preatrium short or reduced.............................................................................. 3

2 Aedeagal shaft strongly compressed, (median width in lateral view/median width in dorsal or ventral view=2.28–3.58) (Fig. 23E, F, L–Q), preatrium short (preatrium length/length across base of aedeagus=0.16–0.34) (Fig. 23E, L, O)............................................................................................ Proekes cephaleus (Naudé)

2`Aedeagal shaft weakly compressed, (median width in lateral view/median width in dorsal or ventral view=1.54–2.02) (Fig. 26A–D), preatrium elongate (preatrium length/length across base of aedeagus=0.40–0.44) (Fig. 26A, C)............................................................................................ Proekes hemiplatyphalis sp. n.

3 Apex of shaft with two paired, tubular teeth (Fig. 28B–D, F–L); subgenital plate with many macrosetae in distal half..................................................................................... Proekes tetracaphalis sp. n.

3`Apex of shaft with one paired, depressed tooth (Fig. 30D–F), subgenital plate with few macrosetae at apex (Fig. 30C)..................................................................................... Proekes diacaphalis sp. n.