Colistra Davies

Stiller, Michael & Webb, Michael D., 2022, Leafhoppers of the Fynbos Biome of South Africa: Colistra, Proekes, Proekoides and a new genus (Insecta, Hemiptera, Cicadellidae, Deltocephalinae, Bonaspeiini), Zootaxa 5199 (1), pp. 1-79 : 5-10

Colistra Davies

Figs 1–14 , 31

Colistra Davies, 1988: 215 , 216.

Type species: Caffrolix parvulus Linnavuori, 1961 , by original designation.

Revised generic diagnosis and description

Diagnosis.

1. Small leafhopper; male and female: length from apex of crown to apex of tegmina 2.8–3.2 mm; crown length medially 0.5–0.6 mm; crown angle 92–103°.

2. Dorsum of head in male and female with brown to dark brown color pattern with apex of crown with deltate, paired, proximate marking and circular or amorphous marking on disc, sometimes small ovoid paired making near base of head ( Figs 1A–D , 2A–E , 3A–Q ), markings sometimes variable ( Fig. 3A, H, I ), rarely immaculate ( Fig. 3K ).

Terminalia in male:

3. Valve crescentic ( Figs 7C , 9C , 11C ), length/width=0.1–0.2.

4. Pygofer deeply incised in dorsal view; pygofer lobe in lateral view, acutely triangular, mediodorsal margin with straight process, with subapical, ventral tooth; 3–6 macrosetae, rarely 8 (length 63–143 µm) ( Figs 7B, F , 9B, D–F , 11B, D ).

5. Segment 10 (of anal tube) large, wide (0.6–0.7 times as wide as width across pygofer at point of greatest width of segment 10), elongate, reaching apex of pygofer lobe (dorsal view, length/width=0.9–1.1) ( Figs 7A , 9A , 11A ).

6. Connective wider than long (greatest length/greatest width=0.5–0.6), stem short, wide, arms slightly shorter to much longer than stem (length arm/length stem=0.7–2.6).

7. Aedeagus with base of anterior margin of shaft edentate; either dorsal apodeme present and preatrium reduced or dorsal apodeme absent and preatrium produced.

Terminalia in female:

8. Sternite 7 posterior margin variable, either uniformly rounded to sublinear ( Figs 8A , 10A, B , 14D, E ), or with deep or shallow notch flanked by rounded lobes ( Figs 12A–C , 14A–C ). Greatest length/greatest width=0.4–0.5, depth of notch/greatest length=0.2–0.3.

9. Valvula 3 protruding one quarter of pygofer length ( Fig. 2G ); with 1–3 rows of marginal to submarginal setae ( Figs 8M , 10F , 12K , 14H, I ).

10. Valvifer 2 sculpture pore-like and microtrichia ( Figs 8F , 10J , 12G , 14J ).

Etymology. Retrospectively, not provided by Linnavuori (1961) or Davies (1988). Latin colis, penis, masculine and two possible suffixes denoting either tools and means; e.g., -tra, -tron, -trum, or diminutive suffixes with derogatory implication, wild resemblance -aster, -tra, -trum; (e.g., Aspidistra elatior , common aspidistra, cast-iron plants, Asparagaceae , Plantae). Gender masculine, determined by the root ( Brown 1956).

Description

Male, female and nymph.

Color.

Head. Ground color ochraceous to stramineous. Apex of crown with paired triangular marking, large ( Figs 1A–C , 2C, E , 3D, F, G, N, Q ), or small ( Figs 1D , 2D , 3C, M ), sometimes absent ( Fig. 3K ). Disc with brown ( Figs 1A–C , 2C, E , 3C, D, N–Q ) to light brown ( Figs 1D, I , 2A, B , 3B ) circular or amorphous marking, sometimes with wide ( Figs 1B , 2E , 3B, 3G ) or narrow lateral extension ( Figs 1A , 3F, L, N–Q ), large ( Figs 1B , 2E , 3D, P ) or small ( Fig. 3E ) or divided medially ( Figs 1D , 2B, D , 3H, I, M ), or sometimes absent ( Fig. 3A, K ). Posterior margin of head with 1–2 small ovoid markings present (e.g., Figs 1A , 2E , 3F, O ) or absent or faded.

Face. Ground color ochraceous to stramineous, frontoclypeus with numerous dark brown lateral arcs ( Figs 1F , 2H ).

Nymph. Variable, reddish, brown or ochraceous, mottled with darker marking ( Fig. 3R–T, V ), median pale longitudinal line ( Fig. 3R–T, V ), or median reddish-brown line ( Fig. 2J, L, M ) with head marked as in adult ( Fig. 3T, U, V ) or unmarked ( Figs 2K , 3R, S ).

Tegmina. Brown marking in margin of outer costal, inner discal cell and inner claval cell. Cells appear opaque white due to reflectance of hind wing under tegmina; veins brown; costal cells yellow or translucent.

Hind wing. Elongate wing light brown to brown to grey-brown translucent, veins brown; short wing pale white opaque or light brown opaque, veins variable, light brown or without color.

Morphology.

Head. Crown angle 92–104°. Wider than pronotum.

Face. Postclypeus longer than wide, with lateral margins parallel to slightly convergent, anterior and posterior margins convex; posterior margin extended beyond lora and posterior margin of gena below lorum (=maxillary plate) narrow.

Ocellus. Ocellus diameter/interocular distance=0.4–0.6.

Pronotum. Lateral margin short, smooth.

Tegmina. Length/width=1.89–2.12; four apical cells, outer and central anteapical cells, inner anteapical cell and inner discal cell fused or without outer anteapical cell ( Figs 7I , 8N , 9O , 10O , 11M , 12P , 13Q , 14N ); appendix absent.

Hind wing. Shorter than tegmina, elongated with costal margin sublinear ( Figs 7J , 8O , 9N , 10P ) or short, about as wide as long with costal margin shallowly convex ( Figs 11N , 12Q , 13R , 14M ); opaque; about one third shorter and narrower respectively than tegmina length and width; length/width=1.1–2.4. Ratio of length and width of tegmina/length and width of hind wing of male and female=1.9–2.0.

Chaetotaxy. Profemur row AV 9–11 short, wide setae, row IC 7–9 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 rounded platellae, one acute seta (lateral) or rarely five round and one acute; metatarsus 2 apical plantar setae include one acute seta, two rounded platellae and one acute seta or rarely three round, one acute.

Measurements.

Male and female similar dimensions (n=339). Apex of crown to apex of tegmina 2.7–3.4 mm, apex of crown to apex of abdomen 2.8–3.9 mm; crown length 0.5–0.6 mm; crown length next to eye 0.3–0.4 mm; pronotum length 0.4 mm; head width 1.1–1.3 mm; pronotum width 1.0– 1.3 mm; ocellus diameter 24–31 µm; interocular distance 45–66 µm; crown angle 92–104°; crown length/crown length next to eye=1.3–1.5; head width/pronotum width=1.0–1.1; ocellus diameter/interocular distance=0.4–0.6; crown length/pronotum length=1.0–1.5; pronotum length/pronotum width=0.3–0.4.

Terminalia.

Male

Segment 10. Large, incised deeply into pygofer, almost to base of pygofer, lateral margin convergent, sublinear ( Figs 7A , 9A ) or convex ( Fig. 11A ), in dorsal view length/width=0.9–1.1, in lateral view compressed ( Figs 7B , 9B , 11B ). Greatest width of segment 10 relative to greatest width of pygofer at point of greatest width of segment 10=0.6–0.7. Posterior margin of anal tube extended beyond posterior margin of pygofer lobe, or as far as pygofer lobe.

Pygofer. In lateral view longer than greatest width; anterior margin oblique, anterior apodeme elongate, angled slightly ventrad (e.g., Fig. 7B ); basolateral suture variably curvate, commonly shallowly convex, rarely strongly convex, incised less than half-way into pygofer lobe ( Fig. 11B ) or right-angled extending about 1/2 into pygofer lobe ( Figs 7B , 9B ); region posteriad of basolateral suture glabrous. Pygofer lobe, in lateral view, acutely triangular, extending almost to base of pygofer; 4–5 macrosetae, rarely eight, laterad of medial area of origin of pygofer process, 55–76 µm in length. Sclerotized process with origin at dorsomedial margin, process straight, orientation posteroventrad, tooth of process ventral, subapical ( Figs 7B, F , 9B, D, E , 11B, D ), mediad of membranous lobes. Apex of lobe membranous or desclerotized, weakly demarcated ( Fig. 7F ) or well differentiated ( Figs 9D , 11D ) in lateral view, medially with numerous microtrichia and few short setae; folded over subapex of sclerotized process ( Fig. 9E ). Subapex medially with ovoid, membranous structure covered with microtrichia and few short setae ( Figs 7F , 9E , 11D ). Lobe with dorsal margin near origin of process with variable, recurved, tooth-like or triangular process, about as long as wide, orientation dorsal or mediolateral, or reduced, often obscured by macrosetae, viewed better dorsolaterally.

Subgenital plate. Extended beyond posterior margin of pygofer lobe and rarely beyond posterior margin of anal tube; greatest length/greatest width=0.8–1.5; medial margins contiguous or narrowly divergent distally; apex broadly rounded; lateral margin shallowly convex. Macrosetae in 2–3 rows near apex, single row medially, macrosetae absent basally, medial macrosetae about as long as median width of subgenital plate, lateral macrosetae shorter.

Valve. Crescentic ( Figs 5C , 7C , 9C ), length/width=0.1–0.2.

Style. Narrow, elongate, greatest length/greatest width=3.4–4.4, length apophysis/greatest length=0.2; apophysis acuminate, apex narrowly rounded, ventrally denticulate, curvate ( Figs 7G , 9G ) or sublinear ( Figs 11L , 13L ), orientation lateroposteriad, angle 38–70°, about half or less as wide as width across preapical lobe; preapical lobe right-angled; anterior medial lobe short in dorsal view; anterior lateral lobe subparallel to base of style ( Figs 9G , 11L , 13L ), about as wide as base or obtusely angled to base ( Fig. 7G ), narrower than base. Apex of apophysis of style inserted either less than halfway ( Figs 7C , 9C ) or three quarters ( Fig. 11C ) length of subgenital plate (halfway to three quarters in C. acapitatus sp. n.).

Connective. Transverse, stem short, wide, arms longer than stem, widely separated, sublinear. Greatest width across arms/width across apex of stem=1.5–2.1. Length arms/length stem=0.7–2.3, variable due to longer or shorter desclerotized margin medially at base of arms, e.g., Figs 7H , 11F , with wide margin; Figs 9N , 11E , 13O, P , with narrower margin. Angle of arms 75–91°. Greatest length/greatest width=0.5–0.6 ( Figs 7H , 9N , 11E, F , 13O, P ).

Aedeagus. Orientation of aedeagus in pygofer with apex of shaft dorsad, vertical, in membranous recess in segment 10. Shaft, dorsal apodeme and preatrium configured as follows:

1. Dorsal apodeme half to two thirds as long as shaft, subparallel to shaft, attached to shaft at atrium, preatrium much shorter than shaft ( Fig. 7D ).

2. Dorsal apodeme about one third as long as shaft, preatrium about as long as shaft ( Fig. 9H, I ).

3. Dorsal apodeme reduced, preatrium as long or longer than shaft ( Figs 11G–K , 13A–F ).

Shaft with teeth in distal half paired or single (possibly due to fusion of contiguous teeth) at anterior or posterior margin, medial paired teeth variable in orientation and origin (anterior, medial or posterior). Base of anterior margin of shaft edentate. Gonopore marginal, wide at apex, anterior subapical margin narrowly incised ( Fig. 13G, I, K ) or not incised ( Figs 7E , 9J ), posterior margin narrowly V-shaped to rounded incision ( Figs 7E , 9J ). Species are distinguished by relative width of apex of shaft in lateral view, compared to medial width ( Figs. 11G–K , 13A–F ), by relative length of shaft to that of the dorsal apodeme ( Figs 7D , 9I ) and by finely denticulate area at anterolateral region of apex of shaft ( Figs 7E , 9K–M ).

Female.

Measurements. As above, female similar to that of male.

Terminalia.

Sternite 7. Transverse, rectangular to isosceles trapezoid (width at posterior margin/width at anterior margin=0.6–0.8, greatest length/greatest width=0.4–0.6) with lateral margins rounded. Three shapes of posterior margins: rounded ( Figs 1G , 2I , 8A , 10A, B , 14D, E ) or with deep ( Fig. 12A–C ) (greatest depth of notch/greatest length laterally=0.61–0.77) or shallow notch ( Fig. 14A–C ) (greatest depth of notch/greatest length laterally=0.75– 0.84), posterior margins laterad of notch rounded.

Valvula 1. Parallel-sided, apex acuminate ( Figs 8I , 10G , 12H , 14F ), sculpture pattern commonly reticulate, striate towards base ( Figs 8K, L , 10K, L, M , 12L, M, N ).

Valvula 2. Serration about in distal half of dorsal margin ( Figs 8H , 10H , 12I , 14J ), narrow, short teeth with trough between successive teeth finely serrate ( Figs 8J , 10N , 12O ).

Valvula 3. 5–22 marginal to submarginal macrosetae at apex, 25–37 µm in length, short setae up to 17 µm ( Figs 8M , 10E, F , 12J, K ). One quarter of apex of valvula 3 protruding beyond pygofer.

Valvifer 1. Rhomboid or rectangular with dorsal and ventral margin somewhat rounded, anterior margin narrowly rounded, posterior margin acute; dorsally fused widely ( Fig. 8B ). Length/width=1.3–1.8 ( Figs 8C, D, E , 10C, D , 12D, E , 14K, L ).

Valvifer 2. Apices narrowly rounded ( Figs 8G , 10I , 12F , 14G ), sculpture pattern at anteroventral margin and submargin reticulate or densely set with microtrichia and few circular structures ( Figs 8F , 10J , 12G , 14H, I ), one specimen with seta. Length/width=2.6–3.0.

Remarks. The combination of external features namely, color pattern of head and tegmina and less so pronotum, and crown angle correspond well within species of Colistra . Other distinguishing features of the genus are given in the diagnosis.

Colistra forms a group with Xhoreus , Proekes and Proekoides , all having some correspondence in size, shape, color pattern and male genitalia. As some specimens appear devoid of the above markings, male genitalia have to be examined. The following similarities and differences are however apparent.

Overall length and body proportions in Colistra bears some similarities with Proekoides with respect to the length from the apex of the crown to the apex of the tegmina, i.e., 3.1–3.5 mm in Proekoides and 2.7–3.4 mm in Colistra . Proekes is much larger (length from apex of crown to apex of abdomen, all specimens, 4.0– 5.1 mm while in the female the ovipositor extends well beyond the apex of the pygofer.

Color pattern of the head, dorsally, includes crown, disc and base tabulated below:

Externally, Colistra also resembles some other Bonaspeiini , e.g., Kimbella Davies, 1988 , Bonaspeia Linnavuori, 1961 (new species in Davies, 1987 and Curvostylus Davies, 1987 . In Kimbella ( Figs 4A, B ), the length from the apex of the crown to the apex of the is tegmina 3.30–3.74 mm, the crown angle is 73–77° and color pattern of head with paired, transverse, deltate marks and small, single, light brown circular marking on the disc. However, in Kimbella the aedeagus has the dorsal apodeme long and preatrium short, and pygofer lobe at posteroventral margin with triangular, serrate process. Tentative differences between Colistra on the one hand and Bonaspeia and Curvostylus on the other, are that the latter two genera lack distinct color pattern on the head, are larger in size, have a long aedeagal shaft, shorter dorsal apodeme and preatrium and different features of the apophysis of the style. With regard to size, length from apex of crown to apex of tegmina (from values in Davies, 1987) in Bonaspeia males 3.46–4.25 mm, females 3.40–3.96 mm, Curvostylus , both sexes, 3.47–4.50 mm and Colistra , both sexes, 2.70–3.20 mm. The apophysis of the style in Bonaspeia could be considered triangular, asymmetric, generally linear, orientation posteriad, about three quarters as wide as width across subapical lobe. The style in Curvostylus has the apophysis digitate, curvate or sublinear, orientation posterolaterad, up to half as wide as width across subapical lobe. Thus, the styles of Colistra and Curvostylus correspond. The aedeagus of most species of Bonaspeia appear to have the dorsal apodeme and preatrium shorter than the shaft and often with the preatrium shorter than the dorsal apodeme, with the exception in B. karensis .The above differentiation of the three genera is tentative due to numerous examined specimens considered new for Bonaspeia and Curvostylus but also probable new genera (personal observation of first author), as well as lack of metrics. Superficial resemblance in marking and color patterns especially on the head are also found in Refrolix trangalensis Theron, 1984 ( Fig. 4D ) and some undetermined Bonaspeiini from the Albany Thicket Biome ( Figs 4C , 5F–I and 6A–H ), and in specimens from the Savanna Biome ( Fig. 6A, B ) with small markings at the apex of the crown. Specimens in Fig. 6F–H ( Fig. 6F , Theewaterskloof, female, Fig. 6G, H , Jonkershoek, male and female respectively) closely resemble the color pattern of the head of Colistra , but are larger (corresponding that of Bonaspeia ( Fig. 5A–C ) and Curvostylus ( Fig. 4E–H )). The male aedeagus of these specimens show some resemblance to the aedeagus in Fig. 13E and with a much longer, single tooth on the preatrium as in Fig. 11K and Fig. 13D .

In male genitalia the pygofer lobe process is present in three of the four genera treated here (absent in Xhoreus gen. n.), but difficult to observe, and is also present in other genera, e.g., Bonaspeia . Most reliable for recognition of genera is the denticulation at the anterior base of the aedeagal shaft, for example, in all species of Colistra , the basal anterior margin of shaft is consistently edentate. Also, the apical, subapical and medial denticulation of the shaft are variable in orientation, position and size and sometimes absent, as discussed under the remarks of each species. In Colistra and Proekes , if the dorsal apodeme is developed the preatrium is reduced and vice versa. In Proekoides , the dorsal apodeme is consistently reduced and the preatrium developed. These features are tabulated as follows:

The connective in Colistra ( Figs7H , 9N , 11E , 13O, P ), Proekoides ( Fig. 16K–O ) and Proekes ( Figs 23H , 26F , 28E , 30J ) is wider than long, with a short, wide stem and widely angled arms. In Xhoreus it least comparable to the former, about as wide as long, narrow stem, but similar angle of arms. Styles of the four genera correspond, with metrics under development.

Key to males of Colistra species

1 Aedeagus with elongate dorsal apodeme; shaft about three quarters longer than dorsal apodeme, preatrium short ( Fig. 7D, E ), apex with anterolateral margin finely denticulate ( Fig. 7D, E )............................... C. parvulus (Linnavuori)

1`Aedeagus with short dorsal apodeme, shaft about two thirds longer than dorsal apodeme ( Fig. 9H, I ), or dorsal apodeme reduced ( Figs 11G–K , 13A–F ), apex of shaft variable......................................................... 2

2 Dorsal apodeme of aedeagus short, preatrium elongate, about as long as shaft ( Fig. 9H, I ), shaft with subapex of posterior margin with single tooth and subapex of anterior margin finely denticulate ( Fig. 9K–M )............... C. semialius sp. n.

2`Dorsal apodeme of aedeagus reduced, shaft of aedeagus at apex in lateral view of variable width ( Figs 11G–K , 13A–F ).... 3

3 Apex of shaft wide, more than twice as wide as medial width of shaft ( Fig. 11G–K )................. C. bucapitatus sp. n.

3`Apex of shaft narrow, up 1.8 times as wide as medial width of shaft ( Fig. 13A–F )................... C. acapitatus sp. n.

Figures

FIGURE 1. Colistra parvulus (Linnavuori, 1961), habitus view. A, male, Ceres specimen, dorsally. B, male, Rawsonville specimen, dorsally. C, female, Rawsonville specimen, dorsally. D, female, Pearly Beach specimen, dorsally. E, female, Rawsonville specimen, laterally. F, face. G, sternite 7. H, female, Maanschijnkop specimen, dorsolaterally. I, male, allotype, dorsolaterally. J, genital capsule, glued to card. K, genitalia, glued to card. L, label, holotype. M, label, collector. N, allotype label. A–E, scale=1 mm.

FIGURE 2. Colistra species, habitus view, dorsally, laterally and ventrally. A–K, C. semialius sp. n. A, male, without marking in disc, Algeria Forestry Station specimen. B, male, small marking in disc, Jamaka specimen. C, male, large marking in disc, Algeria Forestry Station specimen. D, female, small marking in disc,Algeria Forestry Station specimen.E, female, large marking in disc, Algeria Forestry Station specimen. F, G, lateral view, male, female respectively. H, face, male. I, sternite 7. J, K, nymph, Algeria Forestry Station specimen, light brown and dark brown specimen respectively. L, M, Colistra parvulus (Linnavuori, 1961), nymph, Fisantekraal, late and early instar, respectively. A–G, J–L, scale=1 mm.

FIGURE 3. Colistra species, habitus views, dorsally. A–E, R, Colistra bucapitatus sp. n. A, male, unmarked disc, Silvermine specimen. B, male, marked disc, Silvermine specimen. C, male, reduced marking at crown, Stanford specimen. D, male, large marking at crown and disc, Stanford specimen. E, male, small marking at disc, Bettys Bay specimen. F–Q, S–V, Colistra acapitatus sp. n. F, male, lightly pigmented, Seekoevlei specimen. G, male, heavily pigmented, Seekoevlei specimen. H, male, reduced marking at crown and disc, Seekoevlei specimen. I, male, reduced marking at crown and disc, Seekoevlei specimen. J, male, Dwarsrivier. K, male, unmarked head, Uitsig specimen. L, male, disc with paired marking, Uitsig specimen. M, female, disc with paired marking, Wiedouw specimen. N, female, Sanddrift specimen. O, female, Uitsig. P, female, Seekoevlei specimen. Q, female, Stanford specimen. R, nymph, Stanford specimen. S, nymph, Seekoevlei specimen. T, nymph, mottled, Uitsig specimen. U, nymph, pale, Uitsig specimen. V, nymph, striped, Uitsig specimen. A-V, scale=1 mm.

FIGURE 4. Leafhoppers similar to Colistra. A, Kimbella sp., male, Kareedouw specimen. B, Kimbella serricauda (Linnavuori, 1961), female, Robinson’s Pass specimen. C, undetermined, male, Aberdeen specimen. D, Refrolix trangalensis Theron, 1984, male. E, Curvostylus sp., male, De Hoop specimen. F, Curvostylus sp., male, De Hoop specimen. G, Curvostylus sp., male, Driftsands specimen. H, Curvostylus choridulus (Naudé, 1926), male. A–H, scale=1 mm.

FIGURE 5. Leafhoppers similar to Colistra. A–I. Dorsal habitus views. A, Curvostylus ceresi (Theron, 1974). B, Bonaspeia attia Davies, 1987. C, B. straminea Linnavuori, 1961. D, E, unnamed leafhopper from Gansbaai, male and female respectively. F, G, unnamed leafhopper from Cradock, male and female respectively. H, I, unnamed leafhopper from Fouriesburg, female and male respectively. A–I, scale=1 mm.

FIGURE 6. Leafhoppers similar to Colistra. A–I. Dorsal habitus views. A, undetermined, apex of crown with small, paired marking, male, Happy Rest specimen. B, undetermined, apex of crown with small, paired marking, male, Venda specimen. C, Caffrolix sp., male, Keurboom River specimen. D, undetermined, male, Hottentotsholland specimen. E, undetermined, nymph, Hottentotsholland specimen. F, Curvostylus sp., female, Theewaterskloof specimen. G, undetermined, male, Stellenbosch. H, undetermined, female, Stellenbosch. A–H, scale=1 mm.

FIGURE 7. Colistra parvulus (Linnavuori, 1961), line drawings, male. A, genital capsule, dorsally. B, genital capsule, laterally. C, subgenital plate, ventrally. D, aedeagus, laterally. E, aedeagus, shaft apex, posteriorly. F, pygofer, lobe, laterally. G, style. H, connective. I, tegmina. J, hind wing.

FIGURE 8. Colistra parvulus (Linnavuori, 1961), line drawings, female. A, sternite 7. B, valvifer 1, dorsally. C, valvifer 1, laterally, rectangular, Rawsonville specimen. D, valvifer 1, laterally, rounded, Rawsonville specimen. E, valvifer 1, laterally, elongate, Rawsonville specimen. F, valvifer 2, sculpture, apex. G, valvifer 2. H, valvula 2. I, valvula 1. J, valvula 2, sculpture, apex. K, valvula 1, sculpture, subapex. L, valvula 1, sculpture, midsection. M, valvula 3, apical setae. N, tegmina. O, hind wing.

FIGURE 9. Colistra semialius sp. n., line drawings, male. A, genital capsule, dorsally.B, genital capsule, laterally. C, subgenital plate, ventrally. D, pygofer lobe, laterally. E, pygofer, lobe, dorsally. F, pygofer, lobe, caudally. G, style. H, aedeagus, anteriorly. I, aedeagus, laterally. J, aedeagus shaft apex posteriorly. K–M, aedeagus shaft, apex, laterally. K, wide region of denticulation. L, acute region of denticulation. M, narrow region of denticulation. N, connective. O, tegmina. P, hind wing.

FIGURE 10. Colistra semialius sp. n., line drawings, female. A, sternite 7, Jamaka specimen. B, sternite 7, Algeria Forestry Station specimen. C, valvifer 1, Jamaka specimen. D, valvifer 1. E, valvula 3. F, valvula 3, apical setae. G, valvula 1. H, valvula 2. I, valvifer 2. J, valvifer 2, sculpture, apex. K, valvula 1, sculpture, apex. L, valvula 1, sculpture, midsection. M, valvula 1, sculpture, base. N, valvula 2, sculpture, apex. O, tegmina. P, hind wing.

FIGURE 11. Colistra bucapitatus sp. n., line drawings, male. A, genital capsule, dorsally. B, genital capsule, laterally. C, subgenital plate. D, pygofer lobe laterally. E, connective, a–b, width across stem, c–d, width across arms, e, length stem, f, length arms. F, connective, Cedarberg specimen. G–K, aedeagus laterally. G, Papkuilsvlei specimen. H, Natures Valley specimen. I, Keurboom specimen. J, Caledon specimen. K, Remhoogte specimen. L, style, a, apophysis length, sagittal plane, a–b, midsection, b–c, basal length, d–e, angle of apophysis. M, tegmina. N, hind wing.

FIGURE 12. Colistra bucapitatus sp. n., line drawings, female. A–C, sternite 7. A, Silvermine specimen. B, Stanford specimen. C, Papkuilsvlei specimen. D, valvifer 1, Stellenbosch specimen. E, valvifer 1, Silvermine specimen. F, valvifer 2, Silvermine specimen. G, valvifer 2, apical sculpture. H, valvula 1. I, valvula 2. J, valvula 3. K, valvula 3, apical setae. L, valvula 1, sculpture, apex. M, valvula 1, sculpture, midsection. N, valvula 1, sculpture, base. O, valvula 2, sculpture, apex. P, tegmina. Q, hind wing.

FIGURE 13. Colistra acapitatus sp. n., line drawings, male. A–F, aedeagus laterally. A, Groenkol specimen. B, Uitsig specimen. C, Klondyke specimen. D, Seekoevlei specimen. E, Sanddrift specimen. F, Aggenbachskraal specimen. G, aedeagus, apex, posteriorly, Uitsig specimen. H, aedeagus, apex, laterally, Uitsig specimen. I, aedeagus, apex, anteriorly, Uitsig specimen. J, aedeagus, shaft, anteriorly, Seekoevlei specimen. K, aedeagus, shaft, posteriorly, Seekoevlei specimen. L, style. M, aedeagus anteriorly, Arniston specimen. N, aedeagus posteriorly, Arniston specimen. O, connective. P, connective. Q, tegmina. R, hind wing.

FIGURE 14. Colistra acapitatus sp. n., line drawings, female. A, sternite 7, Groenkol specimen, deep notch. B, sternite 7, Aggenbachskraal specimen. C, sternite 7, Groenkol specimen, shallow notch. D, sternite 7, Uitsig specimen, rounded posterior margin. E, sternite 7, Uitsig specimen, rounded posterior margin. F, valvula 1. G, valvifer 2. H, valvifer 2, sculpture, apex, Groenkol specimen. I, valvifer 2, sculpture, apex, Seekoevlei specimen. J, valvula 2, Seekoevlei specimen. K, valvifer 1, Seekoevlei specimen. L, valvifer 1, Seekoevlei specimen. M, hind wing. N, tegmina.

FIGURE 16. Proekoides species, line drawings, male genitalia. A–E, aedeagus, laterally. A, P. hawekwae Stiller, 1986. B, P. cedarbergensis Stiller, 1986. C, P. piketensis Stiller, 1986. D, P. koebergis Stiller, 1986. E, P. postspina sp. n. F–J, aedeagus, posteriorly. F, P. hawekwae Stiller, 1986. G, P. cedarbergensis Stiller, 1986. H, P. piketensis Stiller, 1986. I, P. koebergis Stiller, 1986. J, P. postspina sp. n. K–O, connective. K, P. hawekwae Stiller, 1986. L, P. cedarbergensis Stiller, 1986. M, P. piketensis Stiller, 1986. N, P. koebergis Stiller, 1986. O, P. postspina sp. n. P–T, style. P, P. hawekwae Stiller, 1986. Q, P. cedarbergensis Stiller, 1986. R, P. piketensis Stiller, 1986. S, P. koebergis Stiller, 1986. T, P. postspina sp. n.

FIGURE 23. Proekes cephaleus (Naudé, 1926), male, line drawings inTheron 1975 and additional drawings. A–I, Theron, 1975. A, head dorsally. B, head, laterally. C, pygofer lobe and anal tube, laterally. D, subgenital plate, in part, ventrally. E, aedeagus, laterally. F, aedeagus, anteriorly. G, style, dorsally. H, connective. I, tegmina. J–U, additional drawings. J, genital capsule, dorsally. K, subgenital plate and valve, ventrally. L–Q, aedeagus. L, laterally, Hawekwa specimen. M, laterally, Fisantekraal specimen. N, anteriorly, Fisantekraal specimen. O–Q, laterally, anteriorly, posteriorly, Verlorenvlei specimen. R, genital capsule, laterally, Fisantekraal specimen. S, pygofer lobe, laterally, Fisantekraal specimen. T, hind wing, male, Fisantekraal specimen. U, tegmina, male, Fisantekraal specimen.

FIGURE 26. Proekes hemiplatyphalis sp. n., line drawings, male. A, aedeagus, laterally, posterior paired tooth. B, aedeagus, anteriorly. C, aedeagus, laterally, edentate posterior margin. D, aedeagus, anteriorly. E, style, dorsally. F, connective.

FIGURE 28. Proekes tetracaphalis sp. n., line drawings, male. A–F, K–L, Grootwinterhoek specimens.A, pygofer, dorsally. B, aedeagus, laterally. C, aedeagus, anteriorly. D, aedeagus, posteriorly. E, connective. F, aedeagus, apex, caudally. G–J, aedeagus, Clanwilliam specimen. G, aedeagus, laterally. H, aedeagus, anteriorly. I, aedeagus, posteriorly. J, aedeagus, apex, caudally. K, aedeagus, apex, caudally. L, aedeagus, apex, caudally.

FIGURE 30. Proekes diacaphalis sp. n., line drawings, male and female. A, pygofer, laterally. B, pygofer, dorsally. C, subgenital plate. D, aedeagus, laterally. E, aedeagus, anteriorly.F, aedeagus, posteriorly. G, aedeagus, gonopore, caudally. H, style dorsally. I, sternite 7. J, connective. K, style apophysis, ventrally. L, style apophysis, laterally.

FIGURE 31. Potential natural distribution models, Colistra species. A, Colistra parvulus (Linnavuori, 1961). B, C. semialius sp. n. C, C. bucapitatus sp. n. D, C. acapitatus sp. n.

Color pattern of the head, dorsally, includes crown, disc and base tabulated below: