Hadroca ramosa (Naude)

Figs 1 -9, 26B

Euscelis ramosus Naudé, 1926: 65-66.

Hadroca ramosa ( Naudé), Theron, 1974: 161-162.

Diagnosis.

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.

Etymology.

Ramosus, Latin, full of branches (Brown 1954), for the reticulate color pattern of the tegmina, gender feminine.

Color.

Male, female and nymph.

Dorsum. Small circular marks (Fig. 1E, G, H) or transverse narrow marks (Figs 2L, M, 3A-G), or unmarked and light green to yellow-green (Fig. 1C, D, N, O).

Tegmina. Brown to dark brown reticulations (e.g., Figs 1K, 2L, 4A, E) or yellow-green (Fig. 1N) to stramineous (Fig. 1C). Veins light brown or dark brown, with cells reticulate, rarely with extensive marking in cells or immaculate, translucent or opaque.

Face. Clypeus with brown, horizontal arcs (Figs 2A, C, 3K, L, O), or weakly marked (Fig. 2B). Rarely with facial arcs extending laterodorsally onto crown (Fig. 1K).

Abdomen. Heavily marked (Fig. 1A, E), usually with median line and small round marks (Figs 1F-H, 3E-G), or without median line (Figs 2L, M, 3A-D). Nymph color as in Fig. 2I-K.

Morphology.

Male and female. Tegmina.

Brachypter rare (36 males and 29 females), with tegmina about as long as wide (length/width 1.1-1.2) (Figs 2L, M, 3A-H), hind wing reduced (Fig. 3I), (length/width 0.6, ratio tegmina length/width to hind wing length/width 0.5-0.6).

Tegmina submacropterous [see Zahniser (2021) for terms] common (206 males and 96 females), longer than wide (length/width 1.5-1.7) (Figs 1A-O, 3M, N), hind wing reduced (length/width 0.6-0.8, ratio tegmina length/width to hind wing length/width 0.5-0.7).

Macropter rare (10 males and 8 females), with tegmina extended beyond apex of abdomen and hind wing well developed with large jugal lobe (Fig. 4A-F).

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. 4I).

Chaetotaxy. AV 10-12, IC 9-11 (Fig. 2D).

Measurements.

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.

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.

Terminalia.

Male.

Aedeagus. Shaft edentate or sometimes with single, narrow, anterior, subapical process at base of gonopore; commonly narrow curvate (Figs 5A, B, G, 7J-L, N, O) segment angle 143-185° (n=37), rarely widely curvate (Fig. 7M), segment angle 136°. In dorsal or posterior view, shaft apically narrower or wider than medially, widest subbasally. Gonopore circular or V-shaped, dorsoanteriad. Process at subapex of shaft commonly acutely angled (Fig. 9D-G) or right angled (Fig. 9H) or without process (Fig. 9A, B, D, N), sometimes deformed or damaged (Fig. 9B, D, J, I, K, L, M); process extended beyond apex of shaft (Fig. 9G, H) or shorter (Fig. 9C, E, I, J). Apex commonly notched in posterior view (Figs 7P, S, 9N), or rarely rounded (Fig. 7Q, R, T-V). Aedeagus in lateral view, area in brachypter 27-28 µm 2, in submacropter 25-35 µm 2, in macropter 24-37 µm 2, perimeter in brachypter 1.389-1.634 mm, in submacropter 1.407-2.212 mm, in macropter 1.038-2.711 mm; greatest straight line length from atrium to apex of dorsal apodeme/greatest length from atrium to apex of shaft, submacropter 0.34-0.43, brachypter 0.35-0.48, macropter 0.55-0.68.

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. 8G, I) or rarely somewhat curved laterad (Figs 5E, 8E).

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 5C, D, 8A, B, D), macropter 2.13-2.57 (Fig. 8C).

Subgenital plate. Apex narrowly rounded (Fig. 7F-H) or rounded (Fig. 7E, I), lateral margin smooth, sometimes rugulate (Fig. 7I). Macrosetae basally and distally shorter than medially, uniseriate, submarginal. Three states of length/width ratios:

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. 7E, H, I).

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. 7F).

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. 7G).

Valve. Broadly or narrowly triangular, length/width 0.36-0.46, angle at posterior margin by trigonometry 110-126° (Fig. 7E-I). Submacropter, brachypter and macropter, length/width respectively 0.36-0.45, 0.36-0.49, 0.38-0.46; angle, respectively 111-127°, 100-119°, 105-120°.

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. 7A-D). Shorter macrosetae arbitrarily 65-94 µm .

Female.

Sternite VII. Variable, either (commonly) with narrow acute (length/width 0.6-1.2), recessed ligula on broadly rounded posterior margin (Figs 2F-H, 4G, 8N-R), or (rarely) with short, parallel-sided notch in wide v-shaped posterior margin (Figs 3J, 8S), or rarely sublinear (Fig. 8T), or wide ligula on sinuous margin (Fig. 8U). Posterior margin straight or slightly recessed (lateral length/medial length 1.0-1.1), wider than long, rectangular (length/width 0.4-0.5), posterior margin with narrow recess (notch width/sternite VII greatest width 0.5-0.6).

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. 6D, E), six (Cedarberg, Driehoek and Dwarsrivier, Jonkiespoort) edentate, with or without lateral, submarginal circular structures, mediodorsally with fine denticulation (Figs 6F-H).

Valvula 1. Sculpture granulose distally (Fig. 6B), strigate medially and basally (Fig. 6A, C).

Valvifer 2. Length/width 1.6-2.1. Sculpture with circular structures and microtrichia (Fig. 6I) or circular structures (Fig. 6J).

Valvifer 1. Length/width 1.6-2.1. (Fig. 8K-M).

Material examined.

Type locality. Holotype male, South Africa, Western Cape province, Stellenbosch, Jonkershoek, -33.96, 18.92, 17 Dec. 1922, F.W. Pettey, CCDL01015, SANC.

Type specimen.

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 | Euscelis ramosa | Naudé 1926 (red paper) | Hadroca | Euscelis ramosa | ( Naudé) | Det. J.G. Theron", SANC.

Remarks.

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 1A-O, 3A-H, M, N, 4A-D) with brachypters least typical, especially with the color pattern of the crown as in Figs 2L M, 3A-H.

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. 8E and the preapical lobe visible in profile, or absent when it was oriented ventrally. This variability in profile was common in undissected, cleared specimens with styles in situ.

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 5C, D, 8A-D. For the former condition, the measurement of the stem and arm length was made to the sclerotized part of the Y-shaped frame.

The female sternite VII was ligulate in 21 submacropters and all macropters (Figs 4G, 8O), but truncate or narrowly notched in brachypters. The ligulate sternite VII was in H. bualacauda sp. nov. and H. treichroa sp. nov. Dissections of the valvula 2 of submacropters were edentate in 9 and dentate in 15 specimens and dentate in 5 brachypters and 2 macropters.

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. 8S. Additionally Tzitzikamaia resembled the brachypters of H. ramosa. The Driftsands specimens (sternite VII as in Fig. 8T) were collected by vacuum on Imperata cylindrica in 2004, but not found in 2022 at the same locality on a number of shrubs. Tzitzikamaia is well represented in SANC with 246 records with more than 800 brachypterous specimens, half of the records without host or habitat, the rest divided approximately equally between grass and forbs or grass and ten tree or shrub families, with seven records on specific grasses. Distribution models and point distribution for Tzitzikamaia show a closer association with the Grassland Biome (Fig. 30A-C). The aedeagal shaft bears paired, apical processes and the dorsal apodeme is considered longer and thicker than in Hadroca. The distal configuration of the style in dorsal view has the apophysis longer than in that of Hadroca, linear, with the preapical lobe laterad of the apophysis. In Hadroca the apophysis shorter, curved ventrad, with the preapical lobe ventrad of the apophysis.