Corresponding author: Michael Stiller (
Academic editor: Michael Wilson
The leafhopper genus
The South African leafhopper genus
The wide range of colors and shapes of the species included in
Specimens in more than 200 records were examined, with less than half with an associated plant, but few with nymphs and few repeat collections.
More than 1000 specimens were examined. Terms and drawings follow
Distribution analysis was done in Diva GIS and MaxEnt (
Original label data with line breaks indicated by a vertical space bar (|), label breaks as ||.
Repositories and institutional acronyms for holotypes are
1 | Aedeagus, in lateral and dorsal or anterior view, with shaft with regular profile, uniformly tapered; reduced preatrium; aedeagus evenly sclerotized (Fig. |
2 |
– | Aedeagus, in lateral and dorsal or anterior view, with irregular profile, shaft widest medially, acuminate apex; rounded preatrium; desclerotized apex and medially, sclerotized laterally and basally (Fig. |
|
2 | Apophysis of style, in lateral view, widely arched dorsad, about as long as base of style (Figs |
|
– | Apophysis of style with slight ventrad curve, apophysis much shorter than base of style; male and female length from apex of crown to apex of tegmina 2.55–4.98 mm, length from apex of crown to apex of abdomen 3.27–4.53 mm, pronotum length 0.44–0.62 mm, head width 1.28–1.71 mm, pronotum width 1.23–1.69 mm | 3 |
3 | Tegmina with narrow extension at posterior margin (Figs |
4 |
– | Tegmina with posterior margin rounded or truncate (Figs |
5 |
4 | Large species (Fig. |
|
– | Smaller species (Fig. |
|
5 | Tegmina with three colors (light brown, dark brown and white cells, veins and reticulation dark brown) (Fig. |
|
– | Tegmina at most with two colors or light green to yellow-green, tegmina macropterous, submacropterous (up to 3 abdominal segments exposed) or brachypterous exposing 4–5 abdominal segments; aedeagus in lateral view curvate, dorsal apodeme basal, preatrium reduced; female sternite VII with ligula longer than wide or without ligula and deep, narrow parallel-sided notch; denticulation of valvula 2 sinuous with fine denticulation in trough (e.g., Figs |
6 |
6 | Tegmina marked with darker V-shaped band, reaching apex of abdomen; crown with 2–3 paired, amorphous to angular light brown marks (Fig. |
|
_ | Tegmina with uniform, reticulate brown to light brown marks or unmarked and yellow-green to light green, tegmina brachypterous (Fig. |
Small (2.2 mm) to moderate (6.5 mm) size.
Male and female similar size, color and shape.
Ochraceous or light green, dark brown spotted, speckled, reticulate, or unmarked.
Tegmina rarely brachypterous or macropterous, commonly submacropterous. Hind wing reduced in brachypter and submacropter, fully developed in macropter. Two species with tegmina with narrow, elongated extension at posterior margin, other species with margin rounded or truncate.
Aedeagus C-shaped, uniformly sclerotized, shaft uniformly curved, tapered or subparallel in lateral and posterior or anterior views, dorsal apodeme relatively short, transverse, preatrium reduced or weakly developed, shaft edentate or rarely submacropter of
Style slightly longer than greatest width, apophysis in dorsal view linear to sublinear, rarely curvate laterad, in lateral view curved ventrad, preapical lobe usually ventrad.
Connective Y-shaped, either longer than wide (length greatest/width greatest 1.72–2.52) in four species, or wider than long in
Valve obtusely triangular.
Pygofer lobe rounded, edentate, scattered macrosetae distally, anal tube incised half way, anterior apodemes absent.
Subgenital plate triangular, commonly with uniseriate macrosetae, rarely absent.
Four broad patterns recognized:
Ochraceous to stramineous with weak or distinct brown reticulations on tegmina and regular to irregular stipples or spotting on the head and pronotum (
Light green to yellowish green with feint, brown reticulation sometimes in posterior margin of tegmina or without markings in tegmina. (
Head and pronotum ochraceous to stramineous with head with three pairs of light brown regular to amorphous markings, tegmina with brown reticulations and dark brown, oblique band (
Head and pronotum ochraceous to stramineous, tegmina with light brown, dark brown and white color pattern (
Corresponding closely, except length from apex of crown to apex of tegmina or abdomen (370 males, 307 females) Apex of crown to apex of tegmina 1.9–4.3 mm (smallest male and female in
Most curvate:
Intermediate curvature:
Least curvate:
In posterior or dorsal view width of shaft at subapex/width of shaft medially 0.6–1.3, in lateral view width of shaft at subapex/width of shaft medially 0.7–1.0, two species (
Aedeagus in lateral view relation of relative size of dorsal apodeme to that of shaft, straight line distance from apex of point of attachment of connective, i.e., the atrium, to apex of dorsal apodeme/straight line distance from apex of point of attachment of connective to apex of shaft 0.3–0.5.
Area of the aedeagus in lateral view, 18–38 µm2 in all specimens, smallest area in
The five new species of
However, two important internal features of male terminalia define the genus.
The generic feature of the aedeagus is the strict combination of the C-shape, uniform sclerotization, short dorsal apodeme and reduced preatrium. Additionally, the aedeagus is considered edentate or rarely with single, desclerotized, subapical, anterior tooth (Figs
Style metrics vary considerably, not fully resolved and require retrospective evaluation in other members of
The ocellus diameter and interocellar distance in many specimens is poorly defined.
Morphologic differences and similarities between the brachypterous form of
|
|||
---|---|---|---|
color | mottled, spotted, speckled | light green, light brown, rarely dark brown | reticulate |
exposed abdominal segments | 5–6 | 5–6 | 3–5 |
style apophysis laterally | sublinear | sublinear | curvate ventrad |
aedeagal shaft | apex with paired process, sublinear, shaft tubular | apex edentate, shaft compressed | apex edentate, shaft tubular |
pygofer lobe | long macrosetae seriate | long macrosetae scattered | long macrosetae scattered |
dorsal apodeme of aedeagus | about half as long as shaft | about half as long as shaft | about one third as long as shaft |
male tergite X | anteroventral process less than one third as long as distal part | anteroventral process less than one third as long as distal part | anteroventral process about half as long as distal part |
subgenital plate | triangular, lateral and medial margins convergent; macrosetae long, distal, submarginal; style reaching half way | triangular, lateral and medial margins convergent; macrosetae short, medial, submarginal, style reaching half way | right-angled triangular, lateral margin sinuous or sublinear; macrosetae medial, marginal; style near base |
connective length greatest/width greatest | 1.10–1.46 | 1.06–1.62 | 1.54–2.40 ( |
sternite VII | ligula absent, sinuous, wide or narrow V-shaped notch; narrow deep notch similar to that in Fig. |
provisionally considered shallowly bilobate | ligula absent, wide notch, either deep or shallow; rarely narrow, deep notch (Fig. |
habitat | Grassland and Fynbos Biomes | Grassland Biome | Fynbos Biome |
Stramineous, ochraceous, often with reticulate pattern, light green to yellow green.
Tegmina brachypterous, submacropterous, posterior margin broadly rounded or truncate, or macropterous, with appendix and apex narrowly rounded.
Aedeagal shaft elongate, strongly curvate, preatrium reduced, sometimes with single, subapical, anterior narrow process.
Style apophysis in dorsal view sublinear, in lateral view apophysis short, apex curved ventrad. The apex of the subgenital plate is equidistant to apex of pygofer lobe.
Female sternite VII posterior margin with recessed, variable ligula or sublinear, or V-shaped with narrow, parallel-sided notch.
Brachypter rare (36 males and 29 females), with tegmina about as long as wide (length/width 1.1–1.2) (Figs
Tegmina submacropterous [see
Macropter rare (10 males and 8 females), with tegmina extended beyond apex of abdomen and hind wing well developed with large jugal lobe (Fig.
Submacropter: length/width 1.28–1.69, 4–7 macrosetae, length 111–190 µm, angle at apex of subgenital plate, by trigonometry 31–38° (Fig.
Brachypter: length/width 0.85–1.12, 2–4 macrosetae, length 71–175 µm, angle at apex of subgenital plate, by trigonometry 42–49° (Fig.
Macropter: length/width 1.69–2.01, 6–8 macrosetae, length 176–287 µm, angle at apex of subgenital plate, by trigonometry 26–31° (Fig.
More than 400 specimens were examined, with the majority submacropters (353 specimens: 204 males, 91 females, 58 nymphs, including 47 dissected males, 31 localities), brachypters were 94 specimens (54 males, 34 females and 6 nymphs, 23 localities) and macropters 19 specimens from 6 localities, with Slagboom, near Ceres in the Agterwitsenberg the only locality where all three forms were recorded, i.e., 1967, 1969 and 2004.
Color varied greatly (Figs
The aedeagal shaft in anterior or dorsal view of 70 examined specimens had 46 with a wide apex, 18 narrow and six parallel. The subapical, anterior process was short in 36 specimens, long in 21 and absent in 10, its position relative to the shaft, 38 acute, 17 subparallel and three right-angled. The distal notch was shallow in 25 specimens, deep in eight and absent in 36. Shaft curvature and shape, especially in the brachypter was variable, but not considered discrete.
Variability in the style was sometimes due to angle of dorsal or lateral observation. In general, lateral view commonly clearly depicted ventral curvature of the apophysis. Dorsal view, however, suggested greater variability as in Fig.
Variability in the connective was the degree of the membranous ingression between the arms, and somewhat the lateral margin of the stem, as in Figs
The female sternite VII was ligulate in 21 submacropters and all macropters (Figs
The female sternite VII was least typical in brachypters, and did not resemble that of the submacropter or macropter. Furthermore, examination of specimens of
Tegmina submacropterous, posterior margin rounded, stramineous, ochraceous, or light green to yellow green.
Aedeagal shaft elongate, strongly to weakly curvate, preatrium reduced.
Style apophysis in dorsal view sublinear, in lateral view apophysis about half as long as base, strongly arched dorsad, apex curved ventrad.
Subgenital apex equidistant to apex of pygofer lobe.
Female sternite VII posterior margin with wide, rounded, deep or shallow notch.
Named in Greek for the arched apophysis of the style, arch,
This was the smallest of the known species of
The valvifer 1 was uniform in all examined specimens. The shape of the posterior margin of the female sternite VII varied in depth of the notch, but consistently wide and rounded, and the sternite wider than long.
Associated plants in 20 out of 50 records, were
Tegmina macropterous, posterior margin with narrow, acuminate extension, light brown, oblique band medially.
Aedeagal shaft short, strongly curvate, preatrium reduced.
Style apophysis in dorsal view sublinear, in lateral view short, apex curved ventrad.
Subgenital apex equidistant to apex of pygofer lobe.
Female sternite VII posterior margin with short, wide ligula, recessed in deep or shallow V-shaped, wide notch.
Named in Latin, for the large specimen with the wing tail,
This species was larger than the other species of
Tegmina submacropterous, posterior margin rounded, light brown, oblique band medially.
Aedeagal shaft elongate, curvate, preatrium reduced.
Style apophysis in dorsal view curvate laterad, in lateral view short, apex with slight ventral curvature.
Subgenital apex extended beyond posterior margin of pygofer lobe.
Female sternite VII posterior margin sinuous.
Named in Latin for the brown V-shaped band across the tegmina,
The color pattern, especially of the head and tegmina, differentiated this species from the other species of
Tegmina macropterous, posterior margin with narrow, dark brown, acuminate extension. Hind wing reduced.
Aedeagal shaft elongate, widely curvate, parallel-sided in lateral view, preatrium reduced.
Style apophysis in dorsal view sublinear, in lateral view short, apex curved ventrad.
Subgenital apex equidistant to apex of pygofer lobe.
Female sternite VII posterior margin sublinear.
Named in Latin, for the small specimen with the wing tail,
Measurements for comparison of
Measurements | ||
---|---|---|
male apex of crown to apex of tegmina | 4.02–4.23 mm | 5.02–5.95 mm |
male head width | 1.10–1.15 mm | 1.67–1.81 mm |
male pronotum width | 1.05–1.09 mm | 1.63–1.82 mm |
male crown angle | 95–101° | 111–117° |
female apex of crown to apex of tegmina | 4.99–5.33 mm | 5.51–6.49 mm |
female head width | 1.29–1.33 mm | 1.76–1.92 mm |
female pronotum width | 1.23–1.28 mm | 1.74–1.93 mm |
female crown angle | 97–103° | 111–117° |
Male genitalia differed distinctly between these two species. In
Tegmina submacropterous, rounded posterior margin, with whitish, light brown and dark brown markings.
Aedeagal shaft elongate, sublinear, preatrium produced, about as long as dorsal apodeme.
Style apophysis in dorsal view sublinear, in lateral view short, apex curved ventrad.
Subgenital plate apex equidistant to apex of pygofer lobe.
Female sternite VII posterior margin with short, wide, recessed ligula.
Valvula 2 with short, square teeth.
Named in Greek, for the three colors on the tegmina,
Described from one whole and six dissected males and numerous females (six dissected) from three proximate localities in the Cedarberg Mountains. All had the same distinct color pattern on the tegmina. The aedeagus was least typical for
Distribution maps and potential natural distribution models.
Specimens of the species of
Superficially corresponding species with specimen localities.
Distribution maps and potential natural distribution models.
Colleagues, reviewers and editors are thanked for their contributions to the improvement of the manuscript. Permits issued by CapeNature (CNN44-87-16619 (2022), 0056-AAA008-00042 (2015), 0035-AAA004-00688 (2011), 001-202-00022 (2004)) facilitated acquisitions.
Additional material examined
species data