Hadroca ramosa (Naude) Theron, 1974

Euscelis ramosa Naudé, 1926: 65-66.

Redescription of Hadroca .

Diagnosis.

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 H. ramosa with single, subapical, anterior process.

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 H. hapsistylis sp. nov. (length greatest/width greatest 0.66-0.90). In anterior view with apices of arms angled or curved dorsad. Narrow in lateral view.

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.

Etymology.

Hadros, Greek, well-developed, bulky, stout, large (Brown 1954) for the stout body and pronotum, scutellum and tegmina arched in lateral view, gender feminine.

Color.

Male and female, and nymph.

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 ( H. ramosa submacropter (Fig. 1A-M) and macropter (Fig. 4A-D), H. hapsistylis sp. nov. (Figs 10A-L, 11C, D), H. bualacauda sp. nov. (Fig. 15A-C) and H. alacaudella sp. nov. (Fig. 22A-D). Nymphs as in Figs 2I-K, 15D, 22E.

Light green to yellowish green with feint, brown reticulation sometimes in posterior margin of tegmina or without markings in tegmina. ( H. ramosa (Fig. 1N, O), H. hapsistylis sp. nov. (Figs 10K, 11A, B). Nymph similar color to adult, abdomen marked as in Fig. 2K.

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 ( H. alavittata sp. nov., Figs 19A-F, H, 20A).

Head and pronotum ochraceous to stramineous, tegmina with light brown, dark brown and white color pattern ( H. treichroa sp.n, Fig. 24A-D, G).

Face. Brown horizontal arcs on clypeus (Figs 2C, 3K, L, O, 4E, F, 22F), sometimes dark brown with thick arcs and few pale spots (Fig. 2A) or clypeus and clypellus embrowned (Fig. 12B) or half of clypeus with arcs (Figs 19J, 20C) or weakly marked or unmarked (Figs 2B, 12C, 15E, 22G, 24E, F). Arcs sometimes extended dorsally into crown.

Morphology.

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. 4H). Macropters of H. bualacauda sp. nov. (Fig. 15A-C) and H. alacaudella sp. nov. (Fig. 22A-D) with acuminate posterior extension, with apex extended well beyond posterior margin of abdomen. Tegmina length/tegmina width in brachypters 1.1-1.3; submacropter 1.5-2.0, macropters with wing extension 2.9-4.3, macropter in H. ramosa 3.2-3.5.

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. 3I) or very small scale ( H. hapsistylis sp. nov., Fig. 11E, F). Hind wing length/width in brachypters 1.4-1.7; submacropter 1.7-2.7; wing extension macropters 3.2-4.1. Macropter in H. ramosa with large, folded jugal lobe, when open, wider than tegmina (Fig. 4I).

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. 15G, H; in H. ramosa Fig. 2D).

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.

Measurements. Male and female.

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.

Terminalia.

Male.

Aedeagus. C-shaped, uniformly sclerotized (Figs 5A, B, G, 7J-V, 13D-L, 18E-H, 21K, 23D, E, 25C-E). Dorsal apodeme short, narrow in lateral view, transverse in anterior view, variable in length and orientation of connection with phragma. Shaft strongly (Figs 7J-N, 18E, F, 23D) or weakly curvate (Figs 13D, E, 21K, 25C), variable width, acuminate, subparallel, or apex slightly wider than subbase or medially (Fig. 23D, E), usually edentate, sometimes with single, desclerotized, digitate, narrow, anterior, subapical process (Fig. 7N, R, S, U). Preatrium rarely short or commonly reduced. Gonopore apical to subapical, anterior (Figs 9A-N, 18G, 23D, E, 25F), rarely anterior and posterior (Figs 13I, J, 18I, 21G, H). Orientation vertical in genital capsule (Figs 7B, C, 13A, 18B, 21C, 23B, 25A). Curvature of shaft for all specimens, represented as a segment defined by chord and height 138-178°.

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 µm 2 in all specimens, smallest area in H. treichroa sp. nov. 12-18 µm 2, largest area in H. alacaudella sp. nov., 38-44 µm 2; 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. 13M-O) in dorsal view sublinear; medial arm elongate, approximately at mid-length, with base narrow or obtusely triangular; lateral arm elongate, extended anteriad beyond medial arm (Figs 8E, G, I, 13O, 18K, 21M, 23G, 25H); apophysis orientation posteriad or lateroposteriad, commonly short or rarely slightly less than half as long as style (Fig. 13M-O), always curved ventrad or lateroventrad (Figs 8F, H, I, 13M, N, 18L, 23H, 21N, 25I). Preapical lobe rarely reduced, uniformly merged with apophysis (Figs 13M-O, 21M, N, 25H, I) or commonly ventrad to lateroventrad (Figs 5E, F, 8E-J, 18K, L, 23G, H). In anterior view with medial arm curved or angled dorsad (Fig. 18J). Style positioned basally in subgenital plate (Figs 7E-G, 13C, 18C, 21D, 23C, 25B), rarely reaching half way into subgenital plate (Figs 7F, 13B, C). Commonly length greatest/width greatest 1.20-1.80, in H. hapsistylis sp. nov. 2.95-3.97; length apophysis/length greatest 0.12-0.19; extension of lateral arm anteriad of medial arm represented as length to apex of medial arm/length greatest 0.14-0.27, this range was the smallest in macropters of H. ramosa, i.e., 0.03-0.19.

Connective. Y-shaped, commonly longer than wide, stem generally longer than arms, width across stem slightly less than width across arms (Figs 8A-D, 18M, N, 21I, J, 23F, 25G), rarely transverse (Fig. 13P), articulated to aedeagus. Variable extent of desclerotized margins medially between arms. In anterior view with arms angled or curved dorsad (Fig. 18J). Length stem/length greatest 0.41-0.57; length greatest/width greatest 1.15-2.48; wider than long in H. hapsistylis sp. nov. (Fig. 13P), length greatest/width greatest 0.66-0.90.

Subgenital plate. Generally triangular, lateral margin concave to sinuous (Figs 7G, 18C, D, 21D, 23C, 25B), rarely sublinear (Figs 7H, 21D) or convex (Fig. 13B, C), apex broadly rounded (Figs 9G, H, 13B, 21D) or narrowly rounded (Figs 9G, H, 13C, 18C, D, 23C, 25B), lateral margin sometimes rugulate (Figs 7I, 18C, D), or desclerotized (Fig. 13C). Macrosetae commonly uniseriate, submarginal, basally and apically shorter than medially, length 89-223 µm, basal width 8-11 µm; absent in H. alavittata sp.n, fine short setae present near apex and distal margin, apex dorsally with long, fine setae (Fig. 21C, D). Length/width 1.08-1.69, distal angle by trigonometry 31-43°.

Valve. Right to obtusely angled (81-148°), triangular (Figs 7E, F, 18C, D, 21D, 23C, 25B) (length/width 0.36-0.49), H. hapsistylis sp. nov. most obtuse (136-159°) and narrowest (Fig. 13B, C) (length/width 0.20-0.29).

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 7B-D, 13A, 18A, B, 21A, B, 23A, B, 25A). Pygofer length/width <1 commonly in H. ramosa brachypters and submacropters and H. alavittata sp. nov., length/width>1 in H. hapsistylis sp. nov., H. bualacauda sp. nov., and most elongate in H. alacaudella sp. nov., length/width 1.44-1.70.

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. 25J, K), posterior margin varied, either sublinear (Figs 8T, 23I), or with wide, deep (Figs 8S, 13Q-S) or wide, shallow excavation (Fig. 21Q, R), or with short, median ligula, either wider than long (Figs 8O, 18O, P, 25J, K) or longer than wide (Fig. 8N, P, Q, R), or without ligula (Figs 8S, 13Q-S, 21Q, R). Length/width 0.36-0.54, H. treichroa sp. nov. length/width 0.58-0.65, width across apex/width across base 0.52-0.71, ligula (when present) length/width 0.37-1.11.

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 6D, E, 14E, F, 16F, 20H, I), rarely uniformly rectangular teeth (Fig. 24J), rarely edentate (Fig. 6F-H).

Valvula 1. Lanceolate. Sculpture marginal, at apex imbricate, at base striate, at mid-section intermediate, rarely granulose (Figs 6A-C, 14A-C, 16A, B, 17E, F, L, 20E-G).

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. 25M), one seta in seven specimens of H. alavittata sp. nov., setae absent in other species of Hadroca . Microtrichia commonly triangular with short point, rarely small, or short lines. Sculpture was most varied in H. ramosa, i.e., microtrichia many, few, or short lines, or absent (Fig. 6I, J), most uniform in H. hapsistylis sp. nov. (Figs 13U, V, 14I).

Valvifer 1. In dorsal view, margins free; in lateral view anterior margin broadly rounded, posterior margin narrowly rounded, apices acute (Figs 8K-M, 25L) or blunt (Fig. 17H), or both states, i.e., H. hapsistylis sp. nov., blunt in Fig. 13T, acute in Fig. 14G, H. alavittata sp.n, blunt in Fig. 21T, acute in Fig. 20D. Length/width 1.59-2.30.

Remarks.

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 5A, G; 7K, N, O, R, S, U; 9C, E, F, G, I-M). The exact dimensions of the dorsal apodeme are subjective due to the nature of the transition and orientation between the union of the phragma and the dorsal apodeme. An approximation of the relation between the length of the dorsal apodeme and the length of the aedeagus, by the use of a straight line distance from the point of attachment of the connective to the apex of the dorsal apodeme divided by the straight line distance from the former point to the apex of the shaft. This relation in Hadroca species is 0.35-0.45, and in other Bonaspeiini genera as follows:

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. 12A), but sometimes obscured by other structures. The most striking style apophysis is in H. hapsistylis sp. nov. with the very long and strongly arched apophysis, and least typical is in H. alavittata sp. nov. with the apophysis curvate laterad in dorsal view and its apex with minimal ventrad curvature. The medial arm is curved or angled dorsad, when observed anteriorly.

Theron (1974) stated the connective with elongated stem was diagnostic for the genus, but now found to be longer or shorter than the arms, in approximate equal proportions. Even the width of the stem showed varied degrees of lateral sclerotization. This genus has two types of connective, i.e., longer than wide in four species and wider than long in H. hapsistylis sp. nov. and both with medial part of the arms angled dorsad.

The ocellus diameter and interocellar distance in many specimens is poorly defined. Theron (1974) noted the arched pronotum, scutellum and tegmina in lateral view in the submacropterous H. ramosa (Figs 1D, L, 3N), that is similar in H. bualacauda sp. nov. (Fig. 15C), H. alavittata sp. nov. (Fig. 19F) but less distinct in H. hapsistylis sp. nov. (Fig. 11C, D), H. alacaudella sp. nov. (Fig. 22C, D) and H. treichroa sp. nov. (Fig. 24C, G) and additional specimens of H. ramosa (Figs 1I, 3B, H, 4B, D).

Bloemia hieroglyphica ( Naudé, 1926) is similar to Hadroca in color, shape and size (Fig. 28A-F), and similar curvature of the aedeagal shaft, but not uniformly sclerotized, i.e., with lateral margins weakly sclerotized, base strongly sclerotized, apex anteriorly, posteriorly and apex and subapex medially desclerotized (Fig. 5I-L). Additionally, in Bloemia the dorsal apodeme is reduced, preatrium short, in dorsal/ventral or anterior/posterior view with the aedeagus at its widest medially, base narrower and apex narrowest. The style in Bloemia in dorsal view is curved laterad, in lateral view linear.

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 SANC holdings are brachypterous and have three color forms (Fig. 27A-E). The aedeagus is C-shaped, with a much longer dorsal apodeme, about half as long as the shaft, similar width to the shaft in lateral view, and about three times wider than the shaft in anterior or posterior view. Linnavuori (1961) illustrated a part of the aedeagus in lateral view (Fig. 29L), supplemented with own observations of terminalia in numerous specimens. Basutoia is probably grass-feeding and associated with the Grassland Biome of South Africa (Fig. 30A).

Tzitzikamaia Linnavuori, 1961 is brachypterous (Fig. 27I-L), similar in some aspects of size (length from apex of crown to apex of tegmina 2.3-2.8 mm, in Hadroca 2.8-3.3 mm, length from apex of crown to apex of abdomen 3.5-4.5 mm, in Hadroca 3.4-4.2 mm) but head width and pronotum width in Tzitzikamaia 1.5-1.8 mm and Hadroca 1.3-1.5 mm, and notably with the aedeagus with apical, paired processes. The aedeagus of four species has apical, paired processes, as in Fig. 29G-K. The female sternite VII based on own observations and of two species as in Fig. 29M, N, differs considerably, except in some specimens of Tzitzikamaia resembling the narrow incision as in Figs 3J, 8S. Differentiation between Basutoia brachyptera, Tzitzikamaia species and brachypters of H. ramosa in Table 1.

Goniagnathus brachypterus Linnavuori, 1978, (Fig. 27M) (in Goniagnathini (Zahniser and Dietrich 2013)) resembles Hadroca ramosa in color, shape and size, but has the aedeagus thick and tapered apicad with large, semicircular gonopore, the style apophysis wide and serrate and based on SANC holdings distributed in the northern parts of South Africa, and Linnavuori (1978) described the species from Zaire, now the Democratic Republic of Congo.

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. 27F, G) which resembles that of H. bualacauda sp. nov. and H. alacaudella sp. nov.