Research Article |
Corresponding author: Michael Stiller ( stillerm@arc.agric.za ) Academic editor: Michael Wilson
© 2023 Michael Stiller.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Stiller M (2023) Revision of the South African leafhopper genus Hadroca Theron (Hemiptera, Auchenorrhyncha, Cicadellidae, Bonaspeiini). Deutsche Entomologische Zeitschrift 70(1): 13-54. https://doi.org/10.3897/dez.70.90851
|
The leafhopper genus Hadroca Theron, 1974 with type species Euscelis ramosa Naudé, 1926 is re-defined here by examination of long series of specimens with varied color and wing polymorphic males and females. Five new species are added: Hadroca alacaudella sp. nov., H. alavittata sp. nov., H. bualacauda sp. nov., H. hapsistylis sp. nov. and H. treichroa sp. nov. The genus is known only from South Africa, mostly from the Fynbos Biome. All species have the aedeagus C-shaped and uniformly sclerotized, the shaft gradually tapered, commonly edentate, rarely with single, subapical, anterior process and the style with distal apophysis curved ventrad. A key to species is provided, with discussion on the differentiation from similar genera such as Bloemia Theron, 1974, Basutoia Linnavuori, 1961 and Tzitzikamaia Linnavuori, 1961 and distribution maps.
Afrotropical Region, biodiversity, Cape Floristic Region, herbivore, phytophagous, plant-feeding, shrub-association
The South African leafhopper genus Hadroca Theron, 1974 was originally known from Euscelis ramosa Naudé. This species was re-examined by
The wide range of colors and shapes of the species included in Hadroca, necessitates careful attention to features of the aedeagus and style. In lateral view the aedeagus is C-shaped, uniformly sclerotized, with short, narrow, transverse dorsal apodeme and reduced preatrium. The style has the apex or the whole apophysis curved ventrally, albeit weakly in H. alavittata sp. nov. or excessively in H. hapsistylis sp. nov.
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. |
Bloemia hieroglyphica Naudé |
2 | Apophysis of style, in lateral view, widely arched dorsad, about as long as base of style (Figs |
H. hapsistylis sp. nov. |
– | 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. |
H. bualacauda sp. nov. |
– | Smaller species (Fig. |
H. alacaudella sp. nov. |
5 | Tegmina with three colors (light brown, dark brown and white cells, veins and reticulation dark brown) (Fig. |
H. treichroa sp. nov. |
– | 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. |
H. alavittata sp. nov. |
_ | Tegmina with uniform, reticulate brown to light brown marks or unmarked and yellow-green to light green, tegmina brachypterous (Fig. |
H. ramosa (Naudé) |
Euscelis ramosa Naudé, 1926: 65–66.
Diagnosis.
Hadros, Greek, well-developed, bulky, stout, large (
Male and female, and nymph.
Four broad patterns recognized:
Face. Brown horizontal arcs on clypeus (Figs
Head. Angle at crown in both sexes 100–113°, broadly rounded to face, disc smooth, anterior margin of head shagreened, H. ramosa brachypter with fine or coarsely shagreened margin or finely rugose in H. ramosa macropter, H. alacaudella sp. nov. and H. bualacauda sp. nov.
Ocellus and interocular distance. Ocellus small, distance to eye more than three times its diameter (ocellus diameter/interocular distance 0.16–0.41), ocellus diameter 23–38 µm, interocular distance 77–159 µm.
Pronotum. Lateral margin carinate. Narrower or as wide as head.
Scutellum. Suture obtusely V-shaped to arcuate in H. ramosa macropter, sublinear in H. alacaudella sp. nov., H. bualacauda sp. nov.; absent in H. ramosa submacropter and brachypter, represented by few, fine superficial rugae.
Tegmina. Submacropter, 3–4 abdominal segments exposed, brachypter, 4–5 abdominal segments exposed. In brachypter and submacropter with posterior margin broadly rounded, sometimes truncate, appendix absent. Macropter in H. ramosa with appendix, four apical cells, three anteapical cells (Fig.
Hind wing. Reduced in brachypter and submacropter, and macropterous H. bualacauda sp. nov. and H. alacaudella sp. nov. About half as wide and long as tegmina or much smaller. In macropter of H. ramosa well developed with large, folded jugal lobe. In submacropter variable, smaller than tegmina, with narrow, folded jugal lobe (Fig.
Chaetotaxy. Protibia 1+4, mesotibia 4+4, hind knee 2+2+1. Profemur intercalary (IC) row of 9–11 long, fine setae, distal anteroventral seta (AV1) sometimes slightly longer, thicker, darker than intercalary setae, anteromedial (AM1) one seta, 7–13 anteroventral (AV) setae short, narrowly triangular (profemur in H. bualacauda sp. nov. Fig.
Anal tube. Male tergite X rectangular, wider than long, rarely square or longer than wide; lateral margins variable, parallel, weakly divergent or convergent, sometimes weakly sclerotized; in lateral view anterobasal margin produced anteriad, in dorsal view paired triangular or rounded profile, sometimes desclerotized, anterior process about as long as distal part, half as wide in lateral view; width in dorsal and lateral view similar. Length/width 0.74–1.10, width greatest/width distal 1.01–1.41. Tergite 11 length/width 2.11–2.86.
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 H. hapsistylis sp. nov. 1.8 mm, in H. bualacauda sp. nov. largest male 6.3 mm, female 7.1 mm), apex of crown to apex of abdomen 2.6–4.2 mm; crown length 0.39–0.57 mm; crown length next to eye 0.28–0.38 mm; pronotum length 0.33–0.58 mm; head width 1.01–1.60 mm; pronotum width 0.97–1.57 mm; ocellus diameter 23–38 µm; interocular distance 77–159 µm; crown angle 100–113°; crown length/crown length next to eye 1.34–1.55; head width/pronotum width 1.00–1.05; ocellus diameter/interocular distance 0.16–0.41; crown length/pronotum length 0.94–1.24; pronotum length/pronotum width 0.33–0.38; crown length/head width 0.33–0.42; crown length/pronotum width 0.34–0.43; length from apex of crown to apex of tegmina/length from apex of crown to apex of abdomen 0.73–1.09.
Male.
Aedeagus. C-shaped, uniformly sclerotized (Figs
Most curvate: H. ramosa 160–184°, H. bualacauda sp. nov. 175–189°.
Intermediate curvature: H. hapsistylis sp. nov. 139–151°, H. alacaudella sp. nov. 140–165°.
Least curvate: H. alavittata sp. nov. 127–134°, H. treichroa sp. nov. 100–130°.
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 (H. alavittata sp. nov., H. alacaudella sp. nov.) with wider subapex than medial width, i.e., width of shaft at subapex/width of shaft medially 1.1–1.5, the other width of shaft at subapex/width of shaft medially 0.5–0.8.
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 H. treichroa sp. nov. 12–18 µm2, largest area in H. alacaudella sp. nov., 38–44 µm2; perimeter in all specimens 1.21–2.35 mm,
Style. Distally with narrow, short base, preapical lobe ventral, apophysis commonly short (in H. hapsistylis sp. nov. preapical lobe absent, apophysis very long, Fig.
Connective. Y-shaped, commonly longer than wide, stem generally longer than arms, width across stem slightly less than width across arms (Figs
Subgenital plate. Generally triangular, lateral margin concave to sinuous (Figs
Valve. Right to obtusely angled (81–148°), triangular (Figs
Pygofer. Length/width 0.88–1.30; anterior apodeme absent; ventrobasal suture sublinear; pygofer lobe rounded, wide, edentate; male tergite X of anal tube incised about half way into pygofer (Figs
Connective lobe. Edentate, in lateral view with posterior margin broadly rounded, broadly merged with pygofer, scattered long 134–244 µm and short 56–88 µm macrosetae.
Female.
Sternite VII. Commonly wider than long, rectangular to trapezoid, or almost longer than wide in H. treichroa sp. nov. (Fig.
Valvula 3. Slight protrusion beyond posterior margin of pygofer. Ventral margin with 5–11, short macrosetae, length 17–34 µm.
Valvula 2. Distal third to half, forked, dorsally serrate, teeth rounded, small teeth in trough (Figs
Valvula 1. Lanceolate. Sculpture marginal, at apex imbricate, at base striate, at mid-section intermediate, rarely granulose (Figs
Valvifer 2. Elongate, length/width 2.50–3.02, sculpture ventrobasally with numerous microtrichia, absent in H. treichroa sp. nov. and 4–11 pore-like structures, with H. treichroa sp. nov. with 1–2 setae in three out of five specimens (Fig.
Valvifer 1. In dorsal view, margins free; in lateral view anterior margin broadly rounded, posterior margin narrowly rounded, apices acute (Figs
The five new species of Hadroca and especially H. ramosa display a wide range of sizes, wing shapes and forms and color patterns. Hadroca ramosa displays brachyptery, submacroptery and macroptery, with three shapes of the posterior margin of the sternite VII in females with associated males. Two species have tegmina extended well beyond the abdomen, thus considered macropters, but with reduced, narrower and shorter hind wings, albeit with jugal lobes, i.e., H. bualacauda sp. nov. and H. alacaudella sp. nov. Brachypters have 3–5 exposed abdominal segments and 1–3 in submacropters.
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
Basutoia brachyptera Linnavuori, 1961, 0.51–0.58; Bonaspeia Linnavuori, 1961, species 0.63–0.79; Curvostylus Davies, 1987, species 0.81–0.87; Flavorubivolatus Stiller, 2021, species 0.41–0.53, F. curtiverpus Stiller, 2021, 0.49–0.60, F. tensus Stiller, 2021, 0.45–0.51, F. glabrus Stiller, 2021, 0.37–0.49; select species of Geelus Stiller, 2020, 0.43–0.64; Retevolatus Stiller, 2021, species 0.31–0.54, R. flexiverpus Stiller, 2021, 0.41–0.57, R. semicurvierpus Stiller, 2021, 0.20–0.25, R. subspiniverpus Stiller, 2021, 0.38–0.47; Tzitzikamaia Linnavuori, 1961, species 0.55–0.64.
Style metrics vary considerably, not fully resolved and require retrospective evaluation in other members of Bonaspeiini. The generic feature is the ventrad curvature of the apophysis usually visible in situ in the cleared pygofer or whole specimens (Fig.
The ocellus diameter and interocellar distance in many specimens is poorly defined.
Bloemia hieroglyphica (Naudé, 1926) is similar to Hadroca in color, shape and size (Fig.
Basutoia brachyptera Linnavuori, 1961, bears some resemblance to the brachypterous form of H. ramosa, but with length from apex of crown to apex of abdomen larger, i.e., males 4.0–4.6 mm. All examined specimens in
Tzitzikamaia Linnavuori, 1961 is brachypterous (Fig.
Morphologic differences and similarities between the brachypterous form of H. ramosa (Naudé), Basutoia brachyptera Linnavuori and Tzitzikamaia species.
Tzitzikamaia spp | Basutoia brachyptera | Hadroca ramosa brachypter | |
---|---|---|---|
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 (Hadroca spp); 1.81–2.27 Hadroca ramosa brachypter |
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 |
Goniagnathus brachypterus Linnavuori, 1978, (Fig.
Johanus cypraeus (Naudé, 1926) bears some resemblance in shape and size, but has a distinct asymmetric aedeagus.
Teinopterus microphallus Stiller, 2011, is a grass-feeding leafhopper in Paralimnini, with a narrow tail-like wing extension (Fig.
Euscelis ramosus Naudé, 1926: 65–66.
Hadroca ramosa (Naudé), Theron, 1974: 161–162.
Ramosus, Latin, full of branches (
Male, female and nymph.
Dorsum. Small circular marks (Fig.
Tegmina
. Brown to dark brown reticulations (e.g., Figs
Face
. Clypeus with brown, horizontal arcs (Figs
Abdomen
. Heavily marked (Fig.
Male and female. Tegmina.
Hind wing. In brachypterous males 1.0 mm long, 0.6–0.7 mm wide and females 1.1 mm long and 0.7–0.8 mm wide. In subbrachypterous males 1.1–1.7 mm long and 0.6–0.7 mm wide, females 1.2–1.8 mm long and 0.6–0.8 mm wide. In macropterous specimens with jugal lobes well developed (Fig.
Chaetotaxy. AV 10–12, IC 9–11 (Fig.
Male.
Brachypter (n=35). Apex of crown to apex of tegmina 2.46–2.73 mm; apex of crown to apex of abdomen 3.83–4.26 mm; crown length 0.46–0.52 mm; crown length next to eye 0.36–0.42 mm; pronotum length 0.50–0.57 mm; head width 1.59–1.81 mm; pronotum width 1.52–1.75 mm; ocellus diameter 30–45 µm; interocular distance 66–117 µm; crown angle 116–124°; crown length/crown length next to eye 1.19–1.31; head width/pronotum width 1.03–1.06; ocellus diameter/interocular distance 0.29–0.60; crown length/pronotum length 0.86–0.98; pronotum length/pronotum width 0.31–0.34; crown length/head width 0.27–0.31; crown length/pronotum width 0.28–0.32; length to tegmina/length to abdomen 0.62–0.67.
Hadroca ramosa (Naudé) male and female submacropter, habitus images with specimen localities. A. Female, dorsally, Slagboom; B. Female, dorsally, Slagboom; C. Male, dorsally, Slagboom; D. Male, laterally, Ceres; E. Male, dorsally, Skurfteberg; F. Male, dorsally, Dwarsrivier; G. Female, dorsally, Dwarsrivier; H. Female, dorsally, Dwarsrivier; I. Female, laterally, Dwarsrivier; J. Female, dorsally, Cedarberg; K. Male, dorsally, Caledon; L. Male, laterally, Caledon; M. Male, dorsally, Driehoek; N. Female, Dwarsrivier, dorsally; O. Male, Dwarsrivier, dorsally. A–O. Scale bars: 1 mm.
Hadroca ramosa (Naudé) male, female and nymph submacropter and brachypter, habitus images, profemur and terminalia with specimen localities. A–H, submacropter. A. Face, male, Skurfteberg; B. Face, female, Dwarsrivier; C. Face, male, Fisantekraal; D. Profemur, female, Dwarsrivier; E. Genital capsule, Hottentotsholland; F. Sternite VII, Clanwilliam; G. Sternite VII, Dwarsrivier; H. Sternite VII, Dwarsrivier; I. Nymph, Driehoek; J. Nymph, Fisantekraal; K. Nymph, Driehoek; L–M, brachypter, male, Driftsands. L. Dorsally; M. Laterally. J–M. Scale bars: 1 mm.
Hadroca ramosa (Naudé) male and female brachypter and submacropter, habitus images, hind wing, face and terminalia with specimen localities. A–L, brachypter. A. Female, dorsally, Driftsands; B. Female, laterally, Driftsands; C. Male, dorsally, Jonkershoek; D. Male, dorsally, Muldersvlei; E. Female, dorsally, Ceres; F. Female, dorsally, Jonkershoek; G. Female, dorsally, Ceres; H. Female, laterally, Jonkershoek; I. Hind wing, male, Swellendam; J. Sternite VII, Jonkershoek; K. Face, male, Ceres; L. Face, male, Jonkershoek. M–O, holotype male. M. Dorsally, Jonkershoek; N. Laterally, Jonkershoek; O. Face, Jonkershoek. A–N. Scale bars: 1 mm.
Hadroca ramosa (Naudé) male, female macropter, habitus images and wings with specimen localities. A. Male, laterally, Clanwilliam; B. Male, dorsally, Clanwilliam; C. Female, laterally, Clanwilliam; D. Female, Clanwilliam; E. Face, female, Clanwilliam; F. Face, male, Clanwilliam; G. Sternite VII, Clanwilliam; H. Tegmina, male, Clanwilliam; I. Hind wing, male, Clanwilliam. A–D, H, I. Scale bars: 1 mm.
Submacropter (n=142). Apex of crown to apex of tegmina 2.72–3.43 mm; apex of crown to apex of abdomen 3.32–3.94 mm; crown length 0.47–0.54 mm; crown length next to eye 0.33–0.37 mm; pronotum length 0.45–0.54 mm; head width 1.28–1.56 mm; pronotum width 1.23–1.54 mm; ocellus diameter 25–43 µm; interocular distance 69–96 µm; crown angle 102–115°; crown length/crown length next to eye 1.36–1.56; head width/pronotum width 1.00–1.05; ocellus diameter/interocular distance 0.32–0.53; crown length/pronotum length 0.90–1.15; pronotum length/pronotum width 0.33–0.39; crown length/head width 0.32–0.40; crown length/pronotum width 0.32–0.42; length to tegmina/length to abdomen 0.79–0.90.
Macropter (n=10). Apex of crown to apex of tegmina 5.03–6.14 mm; apex of crown to apex of abdomen 1.10–4.72 mm; length antenna 1.76–2.58 mm; crown length 0.45–0.50 mm; crown length next to eye 0.35–0.39 mm; pronotum length 0.66–0.81 mm; head width 1.67–1.91 mm; pronotum width 1.55–1.82 mm; ocellus diameter 51–68 µm; interocular distance 108–122 µm; crown angle 119–128°; crown length/crown length next to eye 1.22–1.36; head width/pronotum width 1.05–1.08; ocellus diameter/interocular distance 0.44–0.59; crown length/pronotum length 0.57–0.74; pronotum length/pronotum width 0.42–0.45; crown length/head width 0.24–0.29; crown length/pronotum width 0.25–0.32; length to tegmina/length to abdomen 1.17–1.37.
Female. Brachypter (n=26). Apex of crown to apex of tegmina 2.44–2.65 mm; apex of crown to apex of abdomen 3.35–3.69 mm; crown length 0.46–0.54 mm; crown length next to eye 0.36–0.41 mm; pronotum length 0.50–0.55 mm; head width 1.56–1.68 mm; pronotum width 1.49–1.63 mm; ocellus diameter 29–43 µm; interocular distance 67–105 µm; crown angle 112–122°; crown length/crown length next to eye 1.21–1.37; head width/pronotum width 1.02–1.05; ocellus diameter/interocular distance 0.32–0.55; crown length/pronotum length 0.88–1.04; pronotum length/pronotum width 0.32–0.34; crown length/head width 0.28–0.43; crown length/pronotum width 0.29–0.35; length to tegmina/length to abdomen 0.69–0.76.
Submacropter (n=78). Apex of crown to apex of tegmina 2.89–3.52 mm; apex of crown to apex of abdomen 3.70–4.51 mm; crown length 0.48–0.56 mm; crown length next to eye 0.33–0.39 mm; pronotum length 0.47–0.55 mm; head width 1.35–1.60 mm; pronotum width 1.31–1.57 mm; ocellus diameter 27–46 µm; interocular distance 70–102 µm; crown angle 103–116°; crown length/crown length next to eye 1.34–1.55; head width/pronotum width 1.00–1.04; ocellus diameter/interocular distance 0.32–0.55; crown length/pronotum length 0.91–1.13; pronotum length/pronotum width 0.34–0.38; crown length/head width 0.32–0.40; crown length/pronotum width 0.32–0.41; length to tegmina/length to abdomen 0.72–0.84.
Macropter (n=8). Apex of crown to apex of tegmina 4.94–6.10 mm; apex of crown to apex of abdomen 4.04–4.92 mm; length antenna 1.64–2.18 mm; crown length 0.47–0.53 mm; crown length next to eye 0.35–0.39 mm; pronotum length 0.68–0.79 mm; head width 1.70–1.89 mm; pronotum width 1.58–1.90 mm; ocellus diameter 50–66 µm; interocular distance 107–125 µm; crown angle 118–125°; crown length/crown length next to eye 1.28–1.44; head width/pronotum width 1.05–1.08; ocellus diameter/interocular distance 0.41–0.60; crown length/pronotum length 0.62–0.76; pronotum length/pronotum width 0.42–0.45; crown length/head width 0.26–0.30; crown length/pronotum width 0.27–0.33; length to tegmina/length to abdomen 1.19–1.27.
Male.
Aedeagus. Shaft edentate or sometimes with single, narrow, anterior, subapical process at base of gonopore; commonly narrow curvate (Figs
Style. Slightly longer than greatest width, length greatest/width greatest 1.12–1.44 in submacropter, 1.62–1.92 in brachypter, 1.16–1.40 in macropter. Apophysis short (all examined specimens, length apophysis/length greatest 0.13–0.19, submacropter 0.13–0.17, brachypter 0.15–0.20, macropter 0.13–0.17); in dorsal view straight (Fig.
Connective. Longer than wide, stem and arms in submacropter and brachypter of variable length, (length stem/length arms submacropter 0.85–1.36, brachypter 0.98–1.33), in macropter stem longer than arms (length stem/length arms 1.29–1.61); length greatest/width greatest in submacropter 1.77–2.56, brachypter 1.81–2.27 (Figs
Subgenital plate. Apex narrowly rounded (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.
Valve. Broadly or narrowly triangular, length/width 0.36–0.46, angle at posterior margin by trigonometry 110–126° (Fig.
Connective lobe. In lateral view, rarely shorter or commonly longer than greatest width (length/width 1.0–1.3), posterior margin rounded. Macrosetae length in macropter 126–288 µm, submacropter 101–222 µm, brachyptera 68–173 µm (Fig.
Female.
Sternite VII. Variable, either (commonly) with narrow acute (length/width 0.6–1.2), recessed ligula on broadly rounded posterior margin (Figs
Valvula 3. Macrosetae 22–44 µm, in Driftsands specimens (n=4) 37–53 µm.
Valvula 2. Nine dissected specimens (Caledon, Cedarberg, Dwarsrivier, Fisantekraal, Hottentotsholland, Fisantekraal, Jonkershoek, Ladismith, Stellenbosch) with serrate apex (Fig.
Valvula 1. Sculpture granulose distally (Fig.
Valvifer 2. Length/width 1.6–2.1. Sculpture with circular structures and microtrichia (Fig.
Valvifer 1. Length/width 1.6–2.1. (Fig.
Type locality. Holotype male, South Africa, Western Cape province, Stellenbosch, Jonkershoek, -33.96, 18.92, 17 Dec. 1922, F.W. Pettey, CCDL01015,
Holotype
male, glued to triangle card, pinned, with abdomen missing. Original label unknown, type set label printed by J.G. Theron: “Stellenbosch | Jonkershoek | 17-12-22 | F.W. Pettey || type (red paper) || HOLOTYPE | Euscelis | ramosa |
Hadroca ramosa (Naudé) and Bloemia hieroglyphica (Naudé), male genitalia images and line drawings with specimen localities. A–G, H. ramosa. A. Aedeagus, Hottentotsholland, laterally; B. Aedeagus, Hottentotsholland, dorsally; C. Connective, Hottentotsholland, dorsally, DIC; D. Connective, Hottentotsholland, dorsally, bright field; E. Style, Hottentotsholland, dorsally; F. Style, Hottentotsholland, laterally; G. Aedeagus & connective, Hottentotsholland, laterally; H–J, B. hieroglyphica, aedeagus. H. Apex, dorsally, Bredasdorp; I. Laterally, Bredasdorp; J. Line drawing, laterally, fig. 76 copied from Theron, 1974; K. Anteriorly, Bredasdorp; L. Line drawing, posteriorly, fig. 77 copied from Theron, 1974. A–H. Scale bars: 100µm.
Hadroca ramosa (Naudé), ovipositor images with specimen localities. A. Valvula 1, medially, Jonkiespoort; B. Valvula 1, subapex, Jonkiespoort; C. Valvula 1, subbase, Jonkiespoort; D. Valvula 2, dentate, apex, Hottentotsholland; E. Valvula 2, dentate, subbase, Hottentotsholland; F. Valvula 2, edentate, subapex, Jonkiespoort; G. Valvula 2, edentate, subapex, Cedarberg; H. Valvula 2, edentate, dorsally, Cedarberg; I. Valvifer 1, sculpture, Fisantekraal; J. Valvifer 1, sculpture, Cedarberg. A–F, H, I. Scale bars: 50µm.
Hadroca ramosa (Naudé) male terminalia, line drawings with specimen localities. A. Genital capsule, dorsally, Dwarsrivier; B–D, genital capsule, laterally. B. George; C. Clanwilliam; D. Copied from Theron, 1974; E–I, subgenital plate. E. Gifberg; F. De Hoop; G. Clanwilliam; H. Copied from Theron, 1974; I. Dwarsrivier; J–O, aedeagus, laterally. J. Driftsands, annotations in material and methods; K. Stellenbosch; L. Swellendam; M. De Hoop; N. Clanwilliam; O. Copied from Theron, 1974, laterally, annotations in material and methods; P–S, aedeagus posteriorly, T, aedeagus anteriorly, U, aedeagus posteriorly, V, aedeagus anteriorly. P. Copied from Theron, 1974; Q. Clanwilliam; R. Cedarberg; S. Cedarberg; T. Stellenbosch; U. De Hoop; V. Stellenbosch.
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
Hadroca ramosa (Naudé) male and female terminalia, line drawings with specimen localities. A–D, connective; A. Stellenbosch; B. Driftsands; C. Clanwilliam; D. Copied from Theron, 1974; E–J, style. E. Dorsally, Driftsands; F. Laterally, Stellenbosch; G. Dorsally, Stellenbosch; H. Macropter, laterally, Clanwilliam; I. Ventrally, copied from Theron, 1974, annotations in material and methods; J. Apophysis, laterally, copied from Theron, 1974. K–M, Valvifer 1. K. Dwarsrivier; L. Fisantekraal; M. Dwarsrivier; N–U, sternite VII. N. Submacropter Dwarsrivier; O. Macropter Algeria Forestry Station; P. Submacropter Fisantekraal; Q. Driehoek; R. Copied from Theron, 1974; S. Swellendam; T. Driftsands; U. Driehoek.
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 Tzitzikamaia revealed some females with the sternite VII with a similar notch as in the specimens from De Hoop, Jonkershoek and Swellendam, as in Fig.
Hadroca ramosa (Naudé) male, aedeagus, apex, laterally or anteriorly with specimen localities. A–M, laterally. A. Gifberg; B. Jonkershoek; C. Slagboom; D. George; E. Hottentotsholland; F. Cedarberg; G. Dwarsrivier; H. Oudekraal; I. Dwarsrivier; J. Swartberg; K. Caledon; L. Dwarsrivier; M. Dwarsrivier; N. Anteriorly Gifberg.
Named in Greek for the arched apophysis of the style, arch, hapsis, and pillar, for the style, stylis, gender feminine.
Male, female & nymph. Crown with few, paired small circular or elongate marks (Fig.
Male.
Tegmina. Tegmina submacropterous (Figs
Hadroca hapsistylis sp. nov., male and female habitus images with specimen localities. A. Male, light brown, Calvinia; B. Male, dark brown, Calvinia; C. Male, dark head, Oudtshoorn; D. Male, light head, Oudtshoorn; E. Male, weak marking, Oudtshoorn; F. Male, light color, Oudtshoorn; G. Male, brown marking, Richmond; H. Male, light color, Ceres; I. Female, light color, Wiedouw; J. Male, brown marking, Nietgenaamd; K. Male, light color, Nietgenaamd; L. Female, dark brown, Nietgenaamd. A–L. Scale bars: 1 mm.
Hadroca hapsistylis sp. nov., male, female and nymph, habitus images and hind wings with specimen localities. A. Male, Ladismith, dorsally; B. Male, Oudtshoorn, dorsally; C. Male, laterally, Warmwaterberg; D. Female, laterally, Wiedouw; E. Male, Ceres, hind wing; F. Female, Oudtshoorn, hind wing; G. Nymph, Oudtshoorn; H. Nymph, Wiedouw; I. Nymph, Nietgenaamd; J. Nymph, Oudtshoorn, large; K. Nymph, Oudtshoorn, medium; L. Nymph, Oudtshoorn, small. A–L. Scale bars: 1 mm.
Hadroca hapsistylis sp. nov., male, female and nymph, face and terminalia images with specimen localities. A. Male, genital capsule, laterally, Richmond; B. Male, face, Richmond; C. Male, face, Warmwaterberg; D. Sternite VII, Wiedouw; E. Sternite VII, New Bethesda; F. Sternite VII, Oudtshoorn.
Hind wing. Reduced, rectangular (Fig.
Female. Tegmina. Submacropterous, not extended beyond apex of abdomen (Fig.
Hind wing. Reduced (Fig.
Chaetotaxy. AV 6–8 setae, IC 8–12 setae.
Male. (n=150). Apex of crown to apex of tegmina 1.95–2.45 mm; apex of crown to apex of abdomen 2.12–2.89 mm; crown length 0.32–0.43 mm; crown length next to eye 0.24–0.31 mm; pronotum length 0.29–0.36 mm; head width 0.90–1.08 mm; pronotum width 0.89–1.03 mm; ocellus diameter 22–29 µm; interocular distance 97–122 µm; crown angle 101–110°; crown length/crown length next to eye 1.29–1.45; head width/pronotum width 1.00–1.06; ocellus diameter/interocular distance 0.19–0.28; crown length/pronotum length 1.10–1.25; pronotum length/pronotum width 0.32–0.35; crown length/head width 0.35–0.41; crown length/pronotum width 0.36–0.43; length to tegmina/length to abdomen 0.81–0.93.
Female. (n=119). Apex of crown to apex of tegmina 2.26–2.80 mm; apex of crown to apex of abdomen 2.68–3.31 mm; crown length 0.37–0.49 mm; crown length next to eye 0.27–0.34 mm; pronotum length 0.31–0.41 mm; head width 0.99–1.22 mm; pronotum width 0.97–1.16 mm; ocellus diameter 22–29 µm; interocular distance 104–136 µm; crown angle 101–108°; crown length/crown length next to eye 1.31–1.47; head width/pronotum width 1.01–1.07; ocellus diameter/interocular distance 0.18–0.25; crown length/pronotum length 1.12–1.30; pronotum length/pronotum width 0.31–0.36; crown length/head width 0.36–0.41; crown length/pronotum width 0.37–0.43; length to tegmina/length to abdomen 0.79–0.90.
Male.
Aedeagus. Shaft edentate, widely curvate (Fig.
Style. Elongate, narrow, preapical lobe reduced, apophysis elongated, arched dorsad, apex directed posteroventrad, arch in situ vertical or partially lateral, fine denticulation ventrolaterally on base of arch. Length greatest/width greatest 2.96–3.93 (Fig.
Connective. Wider than long, stem very short (length stem/length arms 0.44–0.82), narrow, apex variable, well or poorly sclerotized; arms widely separated, length greatest/width greatest 0.66–0.90 (Fig.
Subgenital plate. Apex variable, either rounded or acute, desclerotized or uniformly sclerotized, sometimes dorsoapical margin with fine, long setae; lateral margin convex. Length to width 1.1–1.6, 4–6 macrosetae, length 84–149 µm; angle at apex of subgenital plate, by trigonometry 31–42° (Fig.
Hadroca hapsistylis sp. nov., male and female, terminalia, line drawings with specimen localities. A. Genital capsule, laterally, Ladismith; B. Subgenital plate, Ladismith; C. Subgenital plate, Theronsberg; D. Aedeagus, laterally, Brakwater; E. Aedeagus, laterally, Ladismith; F. Aedeagus, dorsally, Ladismith; G. Aedeagus, dorsally, Brakwater; H. Aedeagus, apex, dorsally, Ladismith; I. Aedeagus, apex, ventrally, Ladismith; J. Aedeagus, apex, dorsally, Brakwater; K. Aedeagus, apex, dorsally, Ladismith; L. Aedeagus, apex, laterally, Ladismith; M. Style, laterally, Brakwater; N. Style, medially, Brakwater; O. Style, dorsally, Ladismith; P. Connective, Brakwater; Q. Sternite VII, Oudtshoorn; R. Sternite VII, Nietgenaamd; S. Sternite VII, Brakwater; T. Valvifer 1, Warmwaterberg; U. Valvifer 2, Wiedouw; V. Valvifer 2, sculpture, Wiedouw.
Connective lobe. In lateral view about as long as wide (length/width 0.9–1.2), lobe slightly narrower than basal width of pygofer, posterior margin of lobe rounded. Macrosetae 141–205 µm long (Fig.
Female.
Sternite VII. Narrow, transversely rectangular, with wide deep or shallow rounded notch (Figs
Valvula 3. Macrosetae length 16–30 µm (Fig.
Valvula 2. Basally trough and crest denticulate (Fig.
Valvula 1. Sculpture granulose distally, strigate basally (Fig.
Valvifer 2. Length/width 1.6–2.1 (Fig.
Valvifer 1. Length/width 2.6–2.9 (Figs
Type locality. Holotype male, South Africa, Western Cape province Oudtshoorn 5km south, −33.649, 22.222, 5 May 2015, M. Stiller leg., DVac, Pteronia ? succulenta, Asteraceae, CCDL26403,
Holotype
male, glued to triangle card, pinned, with genitalia in microvial pinned to specimen. Original label “R.S.A. Oudts- | hoorn 5km S | −33.64904 22.22212 | 5.v.2015 | M. Stiller leg. || DVac | Pteronia | ? succulenta | Asteraceae ||
Hadroca hapsistylis sp. nov., ovipositor images. A. Valvula 1, subapically; B. Valvula 1, apically; C. Valvula 1, subbasally; D. Valvula 1, basally; E. Valvula 2, subbasally; F. Valvula 2, apically; G. Valvifer 1; H. Valvula 3, apically; I. Valvifer 2, sculpture; J. Valvifer 1, texture, medially. A–J. Scale bars: 100 µm.
Paratypes. 234♂♂, 164♀♀, 92 nymphs, total specimens 491.
This was the smallest of the known species of Hadroca, with similarly wide range of color patterns as in H. ramosa, i.e., ochraceous, stramineous, light green to yellow green. Most characteristic was the long and strongly arched apophysis of the style, and orientation medioventrad in 16 dissected specimens and ventrad in 15, possibly due to displacement during maceration. Measurements of parts of the style were complicated by the obscure distinction between the base of the apophysis and medial part of the style and the true width that was influenced by the arch of the apophysis. In other species of Hadroca the preapical lobe, whether ventral or lateroventral, marked this intersection. The subgenital plate was more variable than in other species of Hadroca. The apex in 69 examined specimens was clearly desclerotized in 12, weakly desclerotized or weakly pigmented in 12 and sclerotized in 45. The apex was acute in 44, broadly rounded in 5, and narrowly rounded in 20. No apparent correlation was recognized between color, subgenital plate, locality or associated plant. Some correlation was present in the rounded apex of the subgenital plate and the longer valve, and the acute apex of the subgenital and the shorter valve. The connective of this species was wider than long with a short, variably sclerotized stem. The aedeagal shaft was commonly smooth, but papillose in Graaf Reinet specimens.
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 Eriocephalus sp., Euryops ? oligoglossus, Felicia ? filifolia, Helichrysum tricostatum, Pentzia incana, Pentzia sp., Pteronia incana, P. paniculata, Pteronia sp., Rosenia sp., Asteraceae, Phylica cryptandroides, P. oleaefolia, Rhamnaceae and Zygophyllum sp. Zygophyllaceae. Map of potential natural distribution in Fig.
Named in Latin, for the large specimen with the wing tail, bu-, prefix meaning large, ala, wing, cauda, tail, gender feminine.
Male, female & nymph. Dorsum usually unmarked. Tegmina with some cells whitish, sometimes with brown reticulation, distally embrowned, veins light brown (Fig.
Male. Tegmina. Macropterous (Fig.
Hind wing. Reduced (Fig.
Female. Tegmina. Macropterous (Fig.
Hind wing. Reduced (Fig.
Chaetotaxy. AV 9–14, IC 8–10, single AV1 seta. (Fig.
Male. (n=41). Apex of crown to apex of tegmina 5.02–5.95 mm; apex of crown to apex of abdomen 3.87–4.63 mm; crown length 0.52–0.61 mm; crown length next to eye 0.34–0.40 mm; pronotum length 0.58–0.67 mm; head width 1.67–1.81 mm; pronotum width 1.63–1.82 mm; ocellus diameter 24–40 µm; interocular distance 175–207 µm; crown angle 111–117°; crown length/crown length next to eye 1.46–1.62; head width/pronotum width 0.99–1.04; ocellus diameter/interocular distance 0.12–0.21; crown length/pronotum length 0.83–0.97; pronotum length/pronotum width 0.35–0.38; crown length/head width 0.30–0.35; crown length/pronotum width 0.31–0.35; length to tegmina/length to abdomen 1.24–1.35.
Female. (n=63). Apex of crown to apex of tegmina 5.51–6.49 mm; apex of crown to apex of abdomen 4.57–5.23 mm; crown length 0.56–0.63 mm; crown length next to eye 0.37–0.41 mm; pronotum length 0.64–0.72 mm; head width 1.76–1.92 mm; pronotum width 1.74–1.93 mm; ocellus diameter 25–36 µm; interocular distance 188–227 µm; crown angle 111–117°; crown length/crown length next to eye 1.44–1.61; head width/pronotum width 0.98–1.02; ocellus diameter/interocular distance 0.12–0.18; crown length/pronotum length 0.82–0.94; pronotum length/pronotum width 0.36–0.38; crown length/head width 0.31–0.34; crown length/pronotum width 0.30–0.35; tegmina/length to abdomen 1.18–1.28.
Hadroca bualacauda sp. nov., male, female and nymph, habitus images, face, terminalia and profemur. A. Male, dorsally, Dwarsrivier; B. Male, dorsally, Biesiesfontein; C. Male, laterally, Gifberg; D. Nymph, dorsally, Dwarsrivier; E. Face, Biesiesfontein; F. Sternite VII, Garies; G. Profemur, Tulbagh; H. Caps, i.e., Profumur, Gifberg. A–D. Scale bars: 1 mm.
Male.
Aedeagus. Shaft edentate, narrowly curvate (Fig.
Style. Apophysis short (length apophysis/length greatest 0.1–0.2), straight, length greatest/width greatest 1.5–1.9 (Fig.
Hadroca bualacauda sp. nov., ovipositor images with specimen localities. A. Valvula 1, subbasally, Biesiesfontein; B. Valvula 1, subapically, Biesiesfontein; C. Valvula 2, apically, Biesiesfontein; D. Valvula 3, apically, Garies; E. Valvifer 1, Biesiesfontein; F. Valvula 2, immaculate, apically, Garies; G. Valvula 2, sculptured, apically, Garies; H. Valvifer 1, left, Garies; I. Valvifer 1, right, Garies; J. Valvifer 2, left, Garies; K. Valvifer 2, right, Garies. A–K. Scale bars: 100 µm.
Hadroca bualacauda sp. nov., ovipositor images with specimen localities. A. Valvula 1, narrow apex, Biesiesfontein; B. Valvula 1, wide apex, Biesiesfontein; C. Valvula 3 with valvifer 2, Biesiesfontein; D. Valvula 2 with valvifer 2, Biesiesfontein; E. Valvula 1, sculpture, medially, Biesiesfontein; F. Valvula 1, sculpture, apically, Biesiesfontein; G. Valvula 3, apically, Biesiesfontein; H. Valvifer 1, Biesiesfontein; I. Valvifer 2, Biesiesfontein; J. Valvifer 2, Biesiesfontein; K. Valvifer 2, sculpture, Biesiesfontein; L. Valvula 1, sculpture, subapically, Biesiesfontein. A–L. Scale bars: 100 µm.
Hadroca bualacauda sp. nov., male and female terminalia, line drawings with specimen localities. A. Genital capsule, dorsally, Hermon; B. Genital capsule, laterally, Hermon; C. Subgenital plate, Hermon; D. Subgenital plate, Dwarsrivier; E. Aedeagus, laterally, Piketberg; F. Aedeagus, laterally, Faure; G. Aedeagus, dorsally, Faure; H. Aedeagus, posteriorly, Dwarsrivier; I. Aedeagus, apex, posteriorly, Dwarsrivier; J. Styles and connective, anteriorly, Dwarsrivier; K. Style, dorsally, Cape Flats; L. Style, laterally, Biesiesfontein; M. Connective, Wolseley; N. Connective, Tulbagh; O. Sternite VII, Biesiesfontein; P. Sternite VII, Garies; Q. Hind wing, female, Biesiesfontein; R. Hind wing, male, Biesiesfontein; S. Tegmina, female, Biesiesfontein.
Connective. Longer than wide, stem longer or shorter than arms (length stem/length arms 1.0–1.5), length greatest/width greatest 2.0–2.7 (Fig.
Subgenital plate. Lateral margin concave, apex narrow, variably elongated, densely rugulose; length/width 1.3–1.9, 4–8 macrosetae, length 44–183 µm; angle at apex of subgenital plate, by trigonometry 29–37° (Fig.
Connective lobe. In lateral view longer than greatest width (length/width 1.1–1.4), posterior margin rounded. Macrosetae 44–178 µm long (Fig.
Female.
Sternite VII. Rectangular, with short, narrow ligula (length/width 0.2–0.6), recessed in deep or shallow, wide, V-shaped posterior margin (Figs
Valvula 3. Whole part in Fig.
Valvula 2. Whole part in Fig.
Valvula 1. Whole part in Fig.
Valvifer 2. Length/width 2.5–2.9, sculpture small triangular scales (Figs
Valvifer 1. Length/width 1.8–2.2, distal margin blunt (Figs
Type locality. Holotype male. South Africa, Northern Cape province, Biesiesfontein Farm south Springbok, −29.75, 17.93, 29 Sep.–3 Oct. 2002, 710 m, M. Stiller leg., sweeping, Pteronia divariculata, Asteraceae, CCDL18292,
Holotype
male, glued to triangle card, pinned, with genitalia in microvial pinned to specimen. Original label “South Africa | Northern Cape | Province | Biesiesfontein | Farm S Springbok | 29°45ʹS, 17°56ʹE | 29.ix.–3.x.2002 | 710 m M. Stiller | swept off | Pteronia | divariculata | Asteraceae |
Paratypes. 59♂♂, 84♀♀, 39 nymphs, 183 total specimens.
This species was larger than the other species of Hadroca, and with an acuminate extension at the posterior margin of the tegmina. Hadroca alacaudella sp. nov. also had a similar extension of the tegmina, but was much narrower and shorter. Differences are elucidated under the remarks section of H. alacaudella sp. nov. Variability in the aedeagus of H. bualacauda sp. nov. was confined to the degree of taper of the shaft in lateral view, and generally appeared to be thicker than in other species of Hadroca. The subgenital plate was more variable in the length of the apex, as in Fig.
Anacardiaceae, Searsia undulata; Asteraceae: Berkheya canescens, Dimorphotheca cuneata, Eriocephalus africanus, E. punctulatus, E. racemosus, E. umbellulatus, Euryops multifudus, Gorteria personata, Pentzia sp., Pteronia divariculata; Aizoaceae, Galenia africana; Ebenaceae, Diospyros ramulosa; Fabaceae, Wiborgia mucronata Menispermaceae, Antizoma miersiana. However, the only record with some adults and numerous nymphs was on Dimorphotheca cuneata which is widely distributed in drier parts of southern Africa (https//posa.sanbi.org.Explore, accessed 16 Jun. 2022).
Named in Latin for the brown V-shaped band across the tegmina, ala, wing, vittata, ribbon, band, gender feminine.
Male & female. Crown with paired, transverse, light brown marks apically, medially and subbasally, pronotum with some marks (Fig.
Male. Tegmina. Submacropterous (Fig.
Hind wing. Reduced (Fig.
Female. Tegmina. Submacropterous (Figs
Hind wing. Reduced (Fig.
Chaetotaxy. AV 7–9, IC 9–11.
Male. (n=39) Apex of crown to apex of tegmina 2.70–2.98 mm; apex of crown to apex of abdomen 2.82–3.24 mm; crown length 0.52–0.56 mm; crown length next to eye 0.35–0.38 mm; pronotum length 0.42–0.45 mm; head width 1.28–1.34 mm; pronotum width 1.23–1.30 mm; ocellus diameter 26–39 µm; interocular distance 98–121 µm; crown angle 99–103°; crown length/crown length next to eye 1.43–1.51; head width/pronotum width 1.02–1.05; ocellus diameter/interocular distance 0.23–0.36; crown length/pronotum length 1.19–1.29; pronotum length/pronotum width 0.33–0.35; crown length/head width 0.40–0.43; crown length/pronotum width 0.41–0.44; length to tegmina/length to abdomen 0.89–0.98.
Female. (n=32) Apex of crown to apex of tegmina 3.08–3.34 mm; apex of crown to apex of abdomen 3.53–4.08 mm; crown length 0.56–0.60 mm; crown length next to eye 0.38–0.41 mm; pronotum length 0.45–0.48 mm; head width 1.38–1.44 mm; pronotum width 1.33–1.40 mm; ocellus diameter 28–42 µm; interocular distance 110–130 µm; crown angle 99–103°; crown length/crown length next to eye 1.43–1.51; head width/pronotum width 1.02–1.05; ocellus diameter/interocular distance 0.23–0.36; crown length/pronotum length 1.21–1.29; pronotum length/pronotum width 0.33–0.35; crown length/head width 0.40–0.42; crown length/pronotum width 0.41–0.44; length to tegmina/length to abdomen 0.78–0.91.
Aedeagus. Shaft widely curvate, segment angle 127–134°, apically wider than medially, gonopore v-shaped anteropostriad (Fig.
Hadroca alavittata sp. nov., male and female, habitus images and terminalia with specimen localities. A. Female, dorsally, Swellendam; B. Male, dorsally, Graaf Reinet; C. Female, dorsally, Graaf Reinet; D. Male, dorsally, Swellendam; E. Male, dorsally, Swellendam; F. Female, laterally, Swellendam; G. Genital capsule, posteriorly, Graaf Reinet; H. Female, Swellendam; I. Sternite VII, Swellendam; J. Face, female, Swellendam. A–I. Scale bars: 1 mm.
Style. Apophysis short (length apophysis/length greatest 0.1–0.2), curved or angled posterolaterad (Fig.
Connective. Longer than wide, stem longer or shorter than arms, (length stem/length arms 0.79–1.20), length greatest/width greatest 1.33–1.70 (Fig.
Subgenital plate. Triangular, apex narrowly rounded, length/width 1.4–1.6, macrosetae absent, fine, narrow setae subapically dorsally and on ventrolateral margin; apex of subgenital plate extended beyond posterior margin of pygofer lobe, angle at apex of by trigonometry 31–36° (Figs
Hadroca alavittata sp. nov., male and female, images, tegmina, face and ovipositor with specimen localities. A. Tegmina, reflected light, female, Swellendam; B. Tegmina, transmitted light, female, Swellendam; C. Face, male, Swellendam; D. Valvifer 1, Cradock; E. Valvula 1, subapically, Cradock; F. Valvula 1, medially, Cradock; G. Valvula 1, subbasally, Cradock; H. Valvula 2, apically, Cradock; I. Valvula 2, medially, Cradock; J. Valvifer 1, sculpture, Cradock. Scale bars:1 mm (A, B), 100 µm (D–J).
Hadroca alavittata sp. nov., male and female terminalia, line drawings with specimen localities. A. Genital capsule, dorsally, Cradock; B. Genital capsule, dorsally, Swellendam; C. Genital capsule, laterally, Swellendam; D. Subgenital plate, ventrally, Graaf Reinet; E. Male tergite X, laterally, Swellendam; F. Male tergite X, ventrally, Swellendam; G. Aedeagus, apex, dorsally, Graaf Reinet; H. Aedeagus, apex, posteriorly, Graaf Reinet; I. Connective, Swellendam; J. Connective, Swellendam; K. Aedeagus, laterally, Swellendam; L. Aedeagus, dorsally, Swellendam; M. Style, dorsally, Swellendam; N. Style, laterodorsally, Swellendam; O. Hind wing, female, Swellendam; P. Hind wing, male, Swellendam; Q. Sternite VII, Graaf Reinet; R. Sternite VII, Swellendam; S. Valvifer 2, Graaf Reinet; T. Valvifer 1, Swellendam; U. Valvula 3, apex, Graaf Reinet; V. Valvula 1, Swellendam; W. Valvula 2, Graaf Reinet; X. Valvula 3, Graaf Reinet.
Connective lobe. In lateral view shorter than greatest width (length/width 0.6–0.8), posterior margin acute. Macrosetae 46–86 µm long (Fig.
Female.
Sternite VII. Transversely rectangular, posterior margin sinuous with wide, shallow notch (Figs
Valvula 3. Macrosetae length 23–41 µm (Fig.
Valvula 2. Apex serrate, fine teeth in trough, rounded teeth as in Figs
Valvula 1. Lanceolate (Fig.
Valvifer 2. Length/width 2.3–2.9 (Fig.
Valvifer 1. Length/width 2.2–2.8 (Figs
Type locality. Holotype male, South Africa, Eastern Cape province, Cradock, −32.166, 25.616, 19 Jan. 1984, J.G. Theron leg., sweeping, Asparagus sp., Asparagaceae CCDL28305,
Holotype
male, glued to triangle card, pinned, with genitalia in microvial pinned to specimen. Original label “South Africa | Cradock | 19.i.1984 | J.G. Theron || Asparagus sp., Asparagaceae ||
Paratypes. 39♂♂, 29♀♀, total specimens 68.
The color pattern, especially of the head and tegmina, differentiated this species from the other species of Hadroca, although not always well developed. The face was also distinct. The male tergite X with the ventral paired process and the glabrous subgenital plate were unique. Least typical was the style with the compressed, laterally curved apophysis in dorsal view (Fig.
Named in Latin, for the small specimen with the wing tail, ala, wing, cauda, tail, diminutive suffix –ella, gender feminine.
Male, female & nymph. Dorsum usually unmarked, ground color light brown. Tegmina smoky translucent, apex embrowned with numerous light brown veins (Fig.
Male. Tegmina. Macropterous (Figs
Hind wing. Reduced (Fig.
Female. Tegmina. Macropterous, length/width 3.9–4.3, length 4.0–4.4 mm, width 1.0–1.1 mm; posterior margin narrowly extended with numerous cross veins (as in male, Fig.
Hind wing. Reduced, elongate, length/width 3.7–4.0, length 2.5–2.8 mm, width 0.7 mm (as in male, Fig.
Chaetotaxy. Male AV 7–9 setae, female AV 8–9 setae, male and female, IC 7–10 setae.
Male. (n=8) Apex of crown to apex of tegmina 4.02–4.23 mm; apex of crown to apex of abdomen 2.80–2.94 mm; crown length 0.46–0.52 mm; crown length next to eye 0.30–0.32 mm; pronotum length 0.41–0.43 mm; head width 1.10–1.15 mm; pronotum width 1.05–1.09 mm; ocellus diameter 25–30 µm; interocular distance 116–132 µm; crown angle 95–101°; crown length/crown length next to eye 1.49–1.63; head width/pronotum width 1.04–1.06; ocellus diameter/interocular distance 0.19–0.24; crown length/pronotum length 1.09–1.23; pronotum length/pronotum width 0.38–0.40; crown length/head width 0.41–0.45; crown length/pronotum width 0.43–0.47; length to tegmina/length to abdomen 1.39–1.47.
Female. (n=20) Apex of crown to apex of tegmina 4.99–5.33 mm; apex of crown to apex of abdomen 3.71–4.01 mm; crown length 0.52–0.58 mm; crown length next to eye 0.34–0.38 mm; pronotum length 0.48–0.51 mm; head width 1.29–1.33 mm; pronotum width 1.23–1.28 mm; ocellus diameter 23–34 µm; interocular distance 140–153 µm; crown angle 97–103°; crown length/crown length next to eye 1.45–1.61; head width/pronotum width 1.03–1.06; ocellus diameter/interocular distance 0.16–0.24; crown length/pronotum length 1.06–1.17; pronotum length/pronotum width 0.38–0.40; crown length/head width 0.40–0.44; crown length/pronotum width 0.42–0.46; length to tegmina/length to abdomen 1.28–1.40.
Male.
Aedeagus. Shaft narrowly curvate, segment angle 144–157°, apically wider than medially, dorsal apodeme angled anteriad, gonopore convoluted to V-shaped, subapically between anterior and posterior margin. In lateral view, area 38–44 µm2, perimeter 1.494–2.325 mm (Fig.
Style. Apophysis short (length apophysis/length greatest 0.1–0.2), straight, length greatest/width greatest 2.4–2.7 (Fig.
Connective. Longer than wide, stem longer than arms (length stem/length arms 1.6–2.0), length greatest/width greatest 2.3–2.9 (Fig.
Subgenital plate. Lateral margin concave; length/width 1.5–1.8, 5–7 macrosetae, length 60–98 µm; angle at apex of subgenital plate, by trigonometry 29–33° (Fig.
Hadroca alacaudella sp. nov., male, female and nymph, habitus images, face, sternite VII, tegmina, hind wing with specimen locality, Klondyke farm. A. Male, dorsally; B. Female, dorsally; C. Male, laterally; D. Female, laterally; E. Nymph, dorsally; F. Face, female; G. Face, male; H. Sternite VII; I. Sternite VII; J. Tegmina, transmitted light; K. Tegmina, reflected light; L. Hind wing. A–E, I–L. Scale bars: 1 mm.
Connective lobe. In lateral view longer than greatest width (length/width 1.4–1.6), posterior margin rounded. Macrosetae, short setae length 42–62 µm, long setae length 83–134 µm, longest seta 165 µm, apices of long setae damaged by KOH (Fig.
Female.
Sternite VII. Rectangular, posterior margin straight to slightly sinuous (Figs
Valvula 3. Macrosetae length 23–41 µm (Fig.
Valvula 2. Apex serrate (Fig.
Valvula 1. Lanceolate (resembling Fig.
Valvifer 2. Length/width 2.3–2.9 (Fig.
Valvifer 1. Length/width 2.2–2.8 (Fig.
Type locality. Holotype male, South Africa, Western Cape province, Klondyke farm, near Ceres, −33.308, 19.577, 28 Jan. 2022, M. Stiller leg., sweeping, Senecio sp., Asteraceae, CCDL28915,
Holotype
male, glued to triangle card, pinned, with genitalia in microvial pinned to specimen. Original label “R.S.A. Klondyke | farm Bo Swaar- | moed, −33.308 | 19.577, 28.i. | 2022, M. Stiller || sweep | Senecio sp. | Asteraceae ||
Paratypes. 14♂♂, 47♀♀, 7 nymphs, total specimens 68.
Hadroca alacaudella sp. nov. and H. bualacauda sp. nov. had the acuminate extension of the posterior margin of the tegmina, but specimens of the former were much narrower, the crown angle more acute and shorter than the latter, and listed in Table
Hadroca alacaudella sp. nov., male and female, terminalia, line drawings with specimen locality, Klondyke farm. A. Genital capsule, dorsally; B. Genital capsule, laterally; C. Subgenital plate; D. Aedeagus, laterally; E. Aedeagus, dorsally; F. Connective; G. Style; H. Style; I. Sternite VII; J. Valvula 3; K. Valvula 2; L. Valvifer 1; M. Valvifer 1; N. Valvifer 2; O. Valvifer 2, sculpture; P. Tegmina.
Measurements for comparison of Hadroca alacaudella sp. nov. and Hadroca bualacauda sp. nov.
Measurements | Hadroca alacaudella sp. nov. | Hadroca bualacauda sp. nov. |
---|---|---|
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 H. alacaudella sp. nov. the aedeagus was widely curvate, and wider distally than medially in lateral view (Fig.
Named in Greek, for the three colors on the tegmina, treis, three; chroa, color, gender feminine.
Male & Female. Crown unmarked or with small red dots, pronotum unmarked or with light brown marks (Fig.
Male. Tegmina. Submacropterous (Fig.
Hind wing. Reduced (slightly larger than in Fig.
Female. Tegmina. Submacropterous (Fig.
Hind wing. Reduced (slightly larger than in Fig.
Chaetotaxy. AV 6–8, IC 7–9.
Male. (n=6). Apex of crown to apex of tegmina 1.98–2.22 mm; apex of crown to apex of abdomen 2.56–3.12 mm; crown length 0.38–0.43 mm; crown length next to eye 0.30–0.33 mm; pronotum length 0.35–0.38 mm; head width 1.10–1.13 mm; pronotum width 1.02–1.06 mm; ocellus diameter 14 µm; interocular distance 104–115 µm; crown angle 105–111°; crown length/crown length next to eye 1.26–1.34; head width/pronotum width 1.06–1.09; ocellus diameter/interocular distance 0.12–0.14; crown length/pronotum length 1.07–1.15; pronotum length/pronotum width 0.34–0.36; crown length/head width 0.34–0.39; crown length/pronotum width 0.37–0.41; length to tegmina/length to abdomen 0.74.
Female. (n=25). Apex of crown to apex of tegmina 2.34–2.34 mm; apex of crown to apex of abdomen 3.47–3.61 mm; crown length 0.41–0.43 mm; crown length next to eye 0.31–0.34 mm; pronotum length 0.38–0.40 mm; head width 1.17–1.21 mm; pronotum width 1.10–1.14 mm; ocellus diameter 13–21 µm; interocular distance 111–125 µm; crown angle 107–112°; crown length/crown length next to eye 1.22–1.30; head width/pronotum width 1.05–1.08; ocellus diameter/interocular distance 0.11–0.18; crown length/pronotum length 1.03–1.11; pronotum length/pronotum width 0.34–0.36; crown length/head width 0.34–0.37; crown length/pronotum width 0.36–0.39; length to tegmina/length to abdomen 0.63–0.66.
Male.
Aedeagus. Shaft sublinear, slightly curvate, segment angle 100–130°, shaft with apical and medial width similar, dorsal apodeme subbasal, preatrium produced, gonopore subapical, V-shaped (Fig.
Style. Apophysis short, length apophysis/length greatest 0.1–0.2, straight; length greatest/width greatest 1.5–1.8 (Fig.
Connective. Longer than wide, stem and arms about equidistant, length stem/length arms 0.84–1.02; length greatest/width greatest 2.40–2.88 (Fig.
Subgenital plate. Triangular, length/width 1.26–1.62, macrosetae length 57–77 µm; angle at apex of subgenital plate, by trigonometry 32–38° (Fig.
Connective lobe. In lateral view longer than greatest width (length/width 1.1–1.4), posterior margin rounded. Macrosetae length 59–110 µm (Fig.
Female.
Sternite VII. Variably rectangular, posterior margin with short, wide, shallowly recessed ligula, lateral margins rounded (Figs
Valvula 3. Macrosetae length 21–34 µm.
Valvula 2. Serrate distally, with teeth square, close-set, fine sculpture at base of tooth (Fig.
Valvula 1. Sculpture granulose apically and ovoid basally.
Valvifer 2. Length/width 2.46–2.97, sculptured area with 7–10 circular pore-like structures (Fig.
Valvifer 1. Length/width 1.83–2.05, rounding dorsally and ventrally similar, anterior and posterior margin acute (Fig.
Type locality. Holotype male, South Africa, Western Cape province, Wiedouw Farm base of Gifberg Pass, south east of Van Rhynsdorp, −31.733, 18.766, 3–10 Oct 2002, M. Stiller leg., sweeping, Passerina truncata subsp. truncata, Thymelaeaceae, CCDL18930,
Hadroca treichroa sp. nov., male and female, habitus images with specimen localities. A. Male, dorsally, Wiedouw; B. Male, dorsally, Wiedouw; C. Male, laterally, Wiedouw; D. Female, dorsally, Wiedouw; E. Face, male, Wiedouw; F. Face, female, Wiedouw; G. Female, laterally, Wiedouw; H. Genital capsule, laterally, Wiedouw; I. Sternite VII, Wiedouw; J. Valvula 2, denticulation, Wiedouw. A–D, G, scale 1 mm, J. Scale bars: 50 µm.
Hadroca treichroa sp. nov., male and female, terminalia, line drawings with specimen localities. A. Genital capsule, laterally, Wiedouw; B. Subgenital plate, Wiedouw; C. Aedeagus, laterally, Wiedouw; D. Aedeagus, posteriorly, Wiedouw; E. Aedeagus, dorsally, Wiedouw; F. Aedeagus, apex, dorsally, Wiedouw; G. Connective, Wiedouw; H. Style, dorsally, Wiedouw; I. Style, laterally, Wiedouw; J. Sternite VII, Wiedouw; K. Sternite VII, Clanwilliam; L. Valvifer 1, Wiedouw; M. Valvifer 2, Wiedouw; N. Valvifer 2, sculpture, Wiedouw.
Holotype
male, glued to triangle card, pinned, with genitalia in microvial pinned to specimen. Original label “South Africa | Western Cape | Prov. Wiedouw | Farm foot | Gifberg Pass | SE Van Rhynsdorp | 31°44ʹS, 18°46ʹE | 3-10.x.2002 120 m | M. Stiller leg. || swept off | Passerina | truncata | subsp. truncata | Thymelaeaceae ||
Paratypes. 5♂♂, 24♀♀.
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 Hadroca, with the preatrium produced more and the shaft least curvate. This configuration approached that of Tzitzikamaia, but without apical processes. At present the aedeagi of species of Tzitzikamaia have single, apical paired processes. The six dissected aedeagi of H. treichroa sp. nov. do not show signs of damage at the apex which could suggest broken processes. The ventral curvature of the style apophysis corresponded well to that of the other Hadroca species. The ligulate female sternite VII was wider than long or almost square, with a recessed ligula similar to that of H. bualacauda sp. nov., and the submacropterous specimens of H. ramosa. However, the serration of the valvula 2 was least typical, i.e. with rectangular, close-set teeth. All other species of Hadroca have small teeth in the trough between larger, rounded teeth. Distribution on map in Fig.
Distribution maps and potential natural distribution models. A. Bloemia hieroglyphica; B. H. ramosa; C. H. hapsistylis sp. nov., small yellow triangles represent all leafhopper records; D. H. bualacauda sp. nov.; E. H. alavittata, yellow squares, distribution points for H. alacaudella, single red circle, H. treichroa, three green triangles.
Asteraceae was well represented with 41 recorded plants in 55 records and 18 other plant families and 27 plant species in 33 records. Most species of Hadroca appear restricted to the Western Cape province, commonly in the Fynbos Biome (Fig.
Specimens of the species of Hadroca vary greatly in size, color and shape of the posterior margin of the tegmina. Two species, i.e., H. bualacauda sp. nov. and H. alacaudella sp. nov., have the tegmina with with a tail-like posterior process. The genus is recognized by combined features of the aedeagus and style. The male aedeagus is uniformly sclerotized, the shaft curvate, preatrium reduced, or slightly produced in H. treichroa sp. nov., and the dorsal apodeme short and transverse. The apophysis of the style is curved ventrad, commonly short relative to the style (H. ramosa (Naudé), H. bualacauda sp. nov., H. alacaudella sp. nov. and H. treichroa sp. nov.), or very long in H. hapsistylis sp. nov., and in H. alavittata sp. nov. somewhat curved laterad, with the least ventrad curvature. Species recognition is based subjectively on color (Figs
Superficially corresponding species with specimen localities. A–E, H, Basutoia brachyptera Linnavuori, 1961, dorsally. A. Koppies; B. Witsieshoek; C. Highlands; D. Carolina; E. Springfontein; F, G. Teinopterus microphallus Stiller, 2011, Eastern Cape province, dorsally; F. Male; G. Female; H. B. brachyptera, male, genital capsule, posteroventrally; I–L. Tzitzikamaia species, male; I. Belfast; J. Kirkwood; K. Sani Pass; L. Jonkiespoort; M. Goniagnathus brachypterus Linnavuori, 1978, female, DNyala. A–M. Scale bars: 1 mm.
Basutoia and Tzitzikamaia species, male and female terminalia, line drawings. A–H, M, T. adspersa Theron, 1974. A. Head, pronotum and tegmina; B. Face; C. Genital capsule, laterally; D. Subgenital plate; E. Style; F. Connective; G. Aedeagus, laterally; H. Aedeagus, posteriorly; I. T. irrorata Linnavuori, 1961, aedeagus, laterally; J. T. longipennis Linnavuori, 1961, aedeagus, laterally; K. T. silvicola Linnavuori, 1961, aedeagus, laterally; L. B. brachyptera
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
Data type: species data