Dtsch. Entomol. Z. 70 (1) 2023, 13-54 | DO! 10.3897/dez.70.90851

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Revision of the South African leafhopper genus Hadroca Theron
(Hemiptera, Auchenorrhyncha, Cicadellidae, Bonaspetin1)

Michael Stiller!

1 Biosystematics Division, ARC-Plant Health and Protection, Private Bag X134, Queenswood 0121, South Africa

https://zoobank. org/6D8&B46D4-35BE-4D27-8E8 3-11474340563 1

Corresponding author: Michael Stiller (Stiller-M@arc.agric.za)

Academic editor: Michael Wilson @ Received 25 July 2022 Accepted 3 January 2023 @ Published 3 February 2023

Abstract

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 ala-

caudella sp. nov., H. alavitiata 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 ven-

trad. A key to species is provided, with discussion on the differentiation from similar genera such as Bloemia Theron, 1974, Basutoia

Linnavuori, 1961 and 7zitzikamaia Linnavuori, 1961 and distribution maps.

Key Words

Afrotropical Region, biodiversity, Cape Floristic Region, herbivore, phytophagous, plant-feeding, shrub-association

Introduction

The South African leafhopper genus Hadroca Theron,
1974 was originally known from Euscelis ramosa Naudeé.
This species was re-examined by Theron (1974) as a sub-
brachypterous specimen, and now with additional speci-
mens showing brachyptery and macroptery, and a range
of colors and up to three shapes of the apex of the aedeagal
shaft. Five new species are described and arranged below
by numbers of examined specimens. Until the current re-
vision, the genus has included only the type species. The
genus is included in the deltocephaline tribe Bonaspeiin1,
and in a recently phylogenomic study of Deltocephalinae
(Cao et al. 2022, supporting information) was recovered
in a close relationship to Bloemia Theron.

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 preatri-
um. The style has the apex or the whole apophysis curved

ventrally, albeit weakly in H. alavittata sp. nov. or exces-
sively 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.

Materials and methods

More than 1000 specimens were examined. Terms
and drawings follow Stiller (2020, 2021) and Zahnis-
er (2021). Hind wing lengths are measured from the
point of attachment to the apex and widths the greatest
perpendicular distance, usually at the jugal lobe. Con-
nective measurements were stem width, usually across
the apex, stem length from the apex of the stem to the
sclerotized part at the confluence of the arms, arm length
from the apex of arm to the sclerotized confluence of the
arms and arm width across the widest part of the arms.
The desclerotized margin between the arms was consid-
ered too variable, as in Figs 5C, D, 8A—D, 13P, 18M, N,

Copyright 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, distri-
bution, and reproduction in any medium, provided the original author and source are credited.

14

211, J, 23F, 25G. Due to the vertical orientation of the
aedeagal shaft in the pygofer, its anterior and posterior
views are used here, and considered equivalent to dorsal
and ventral views. Curvature of the shaft is calculated
from line drawings as an approximation of an arc most
closely following the posterior margin of the shaft to the
atrium. It is represented as an angle, using chord length
between the apex of the shaft and apex of the atrium
(Fig. 7J, dashed line ab) and height of the arc (Fig. 7J,
dashed line cd). Area and perimeter of the lateral view
of the aedeagus is based on actual greatest length of rep-
resentative specimens of species treated here, and cal-
culated with ImageJ (Schneider et al. 2012). With the
aedeagus in lateral view, the relation of relative size of
dorsal apodeme to that of shaft is calculated as a ratio.
This is a straight line distance from apex of point of at-
tachment of connective to apex of dorsal apodeme (Fig.
70, dashed line cd) divided by the straight line distance
from apex of point of attachment of the connective to
apex of the shaft (Fig. 70, dashed line ab). The style
with two anterior arms, the ventral arm of Blocker and
Triplehorn (1985) and the equivalent anterior medi-
an lobe of Zahniser (2008) and Zahniser and Dietrich
(2008), henceforth termed as medial and lateral arms.

Michael Stiller: Revision of Hadroca (Cicadellidae)

Measurements of drawings of the style were based on
the parasagittal line (along medial margin of the apoph-
ysis to the medial arm, Fig. 8I dashed line ab) with the
length of the apophysis, midsection and base and great-
est width at right angles to this parasagittal line (Fig. 81,
dashed lines cd, ef and gh respectively). Micrometers
are used for setae length and thickness, and ocellus di-
ameter and interocular distance, and millimeters for all
other measurements.

Distribution analysis was done in Diva GIS and Max-
Ent (Phillips et al. 2020) with comma separated values of
species and their geographical coordinates, 19 bioclimat-
ic worldclim (https://www.worldclim.org) variables and
10 percentile training presence. Maps in Figs 26, 30 were
compiled with DIVA GIS (methods in Scheldeman and
Van Zonneveld 2010).

Original label data with line breaks indicated by a ver-
tical space bar (|), label breaks as |].

Repositories and institutional acronyms for holotypes
are SANC (The National Collection of Insects, Pretoria,
RSA, same address as author address). Parts of long series
of paratypes will be deposited in the BMNH (The Natural
History Museum, London, UK), INHS (illinois Natural
History Survey, Campaign, Illinois, USA) and SANC.

Key to species of Hadroca and Bloemia hieroglyphica

1

Aedeagus, in lateral and dorsal or anterior view, with shaft with regular profile, uniformly tapered; reduced preatrium;
aedeagus evenly sclerotized (Fig. 5A, B, G). Apophysis of style, in lateral view, curved ventrad, distinctly (Figs 8F, H, J,
13M, N, 18L, 23H, 251) or weakly (Figs 5F, 21N). Tegmina polymorphic, submacropters as in Figs 1, 3M, 10, 19A-F,
PAPA e AS Pitatino Cee S DEG (OS ame ne ntctadsaqt trace Repcauthedanat haa ac Be een ucrmadianct iA mae Rantticctociacedtet sees. cejugenades a PAe mead tence enttagn’ hzanc el 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. 5H-L). Apophysis of style, in lateral
view, straight. Tegmina submacropterouS (Fig. 2BAF) ........cccccec ccc ee eee eceeeee sense eeeeeeeeeneeeeneeees Bloemia hieroglyphica Naudé
Apophysis of style, in lateral view, widely arched dorsad, about as long as base of style (Figs 12A, 13A, M, N); male and
female length from apex of crown to apex of tegmina 1.97-—2.56 mm, length from apex of crown to apex of abdomen
2.33-3.02 mm, pronotum length 0.29-0.37 mm, head width 0.90-1.13 mm, pronotum width 0.90-1.08 mm.............
sacadhag Ate Wanna Frees ae ind od ce ee anteget Le Pociautna dbiatanadtnt-eoke aoa qeccbort aatekh ete b boedaadtp evant eebbe mae ed ght outyte-tha dads adie 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, pro-
notum length 0.44-0.62 mm, head width 1.28-1.71 mm, pronotum width 1.23-1.69 MM........... ccc cece eee e neces eens 3
Tegmina with narrow extension at posterior margin (Figs 15A—-C, 22 A-D) ......cccccccc ccc ccc ccc nec nec eec ees eee eee eeeeeeseeseeneeneenennes 4
Tegmina with posterior margin rounded or truncate (Figs 1, 3, 4, LOA-F, H, 24A—D, G) .... 0. cece ccc ccc eee eee eeceee een een eeneenes 5
Large species (Fig. 15A, B), length from apex of crown to apex of tegmina 5.02-5.95 mm, greatest width (=pronotum
width) 1.63-1.82 mm, aedeagus with short robust shaft, strongly curvate (Fig. 18E, F); female sternite VII with wide,
V-shaped notch with medial, ligula wider than long (Fig. 180, P) ..........cccccccecc ccc cce cece eeeeseeeeeneenees H. bualacauda sp. nov.
Smaller species (Fig. 22 A-D), length from apex of crown to apex of tegmina 4.02-4.23 mm, greatest width (=head
width) 1.10-1.15 mm; aedeagus with apex wider than width medially, weakly curvate (Fig. 23D); female sternite VII with
posterior margin Sublinear CRG: -28l Ms. leh arial ehantey aticlanetcerr vy susd Mulebh of laa sesar yy Meet A pe rate adler Vale Medes H. alacaudella sp. nov.
Tegmina with three colors (light brown, dark brown and white cells, veins and reticulation dark brown) (Fig. 24A—D, G),
tegmina short, 2-4 abdominal segments exposed; aedeagus in lateral view sublinear to slightly curvate, dorsal apodeme
subbasal, preatrium present; female sternite VII with posterior margin shallowly excavated with ligula wider than long
(Figs 241, 25J, K), denticulation of valvula 2 with small rectangular teeth (Fig. 24J); length from apex of crown to apex
Of -ADGIOIGE 2S OSL EA INITIO, 5 oie c delgeulene!letodes by wed oadetrite nice lypigcbeh a oh igu a deb Mapeplams pore ae ted gdeeleteu is Hgrade a smagae debieay!s 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,

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Dtsch. Entomol. Z. 70 (1) 2023, 13-54 15

narrow parallel-sided notch; denticulation of valvula 2 sinuous with fine denticulation in trough (e.g., Figs 6D, E, 201)
rately ecemtate: (Fi oO he 6 c.1rn ects tetuhh Systm Spe UL Bi he 0 og Petal hn g pete 9 gece Moth By Beeler ne Seca ot anes alts Bat a hak 6
Tegmina marked with darker V-shaped band, reaching apex of abdomen; crown with 2-3 paired, amorphous to angular
light brown marks (Fig. 19A-F, H); subgenital plate produced beyond posterior margin of pygofer lobe, without macrose-
tae (Fig. 21C, D); male tergite X with paired, ventroposteriad denticles (Fig. 21E, F); shaft in lateral view widely curvate;
female sternite VII with posterior margin wide, shallow excavation (Figs 191, 21Q, R); length from apex of crown to apex
Of AD MOME 2 2 Sve AML ann notes coreeitenbaunheaehettdedenatsiaath rants oaeasacunasudisil nanos otatonaad dian athdnaeae aude H. alavittata sp. nov.
Tegmina with uniform, reticulate brown to light brown marks or unmarked and yellow-green to light green, tegmi-
na brachypterous (Fig. 3A-H), submacropterous (Fig. 1A-O) or macropterous (Fig. 4A-G); crown either unmarked
Fig. 1J, N, O), or with small spots (Figs 1G, H, 3M) or with narrow arcs (Fig. 3A, C, D) or many narrow lines and marks
(Figs 1A, E, K, 3E—G); subgenital plate extended posteriad as far as apex of pygofer lobe, with uniseriate, marginal mac-
rosetae (Fig. 7B—-H); male tergite X with ventroposterior margin smooth; shaft in lateral view narrowly curvate; female
sternite VII with short median ligula longer than wide or wider than long in macropter and submacropter (Fig. 8N—R), or
truncated in brachypter (Fig. 8T, U) or with narrow, parallel-sided notch in brachypter (Fig. 8S)......... H. ramosa (Naudé)

Type species Etymology. Hadros, Greek, well-developed, bulky,

Hadroca ramosa (Naudé) Theron, 1974

stout, large (Brown 1954) for the stout body and pro-
notum, scutellum and tegmina arched in lateral view,
gender feminine.

Euscelis ramosa Naudé, 1926: 65-66. Color. Male and female, and nymph.

Four broad patterns recognized:

Redescription of Hadroca. Diagnosis.

he

1. Ochraceous to stramineous with weak or distinct

Small (2.2 mm) to moderate (6.5 mm) size. brown reticulations on tegmina and regular to irreg-
. Male and female similar size, color and shape. ular stipples or spotting on the head and pronotum
. Ochraceous or light green, dark brown spotted, (H. ramosa submacropter (Fig. 1A—M) and macrop-
speckled, reticulate, or unmarked. ter (Fig. 4A—D), H. hapsistylis sp. nov. (Figs 10A-—
. Tegmina rarely brachypterous or macropterous, L, 11C, D), H. bualacauda sp. nov. (Fig. 15A—-C)
commonly submacropterous. Hind wing reduced and H. alacaudella sp. nov. (Fig. 22A—D). Nymphs
in brachypter and submacropter, fully developed in as in Figs 2I-K, 15D, 22E.
macropter. Two species with tegmina with narrow, 2. Light green to yellowish green with feint, brown
elongated extension at posterior margin, other spe- reticulation sometimes in posterior margin of teg-
cies with margin rounded or truncate. mina or without markings in tegmina. (H. ramosa
. Aedeagus C-shaped, uniformly sclerotized, shaft (Fig. IN, O), H. hapsistylis sp. nov. (Figs 10K,
uniformly curved, tapered or subparallel in lateral 11A, B). Nymph similar color to adult, abdomen
and posterior or anterior views, dorsal apodeme marked as in Fig. 2K.
relatively short, transverse, preatrium reduced 3. Head and pronotum ochraceous to stramineous
or weakly developed, shaft edentate or rarely with head with three pairs of light brown regular to
submacropter of H. ramosa with single, subapical, amorphous markings, tegmina with brown reticula-
anterior process. tions and dark brown, oblique band (H. alavittata
. Style slightly longer than greatest width, apophysis sp. nov., Figs 19A—F, H, 20A).
in dorsal view linear to sublinear, rarely curvate lat- 4. Head and pronotum ochraceous to stramineous,
erad, in lateral view curved ventrad, preapical lobe tegmina with light brown, dark brown and white
usually ventrad. color pattern (H. treichroa sp.n, Fig. 24A—D, G).
. Connective Y-shaped, either longer than wide

(length greatest/width greatest 1.72—2.52) in four Face. Brown horizontal arcs on clypeus (Figs 2C,
Species, or wider than long in H. hapsistylis sp. 3K, L, O, 4E, F, 22F), sometimes dark brown with thick
nov. (length greatest/width greatest 0.66—0.90). In —_arcs and few pale spots (Fig. 2A) or clypeus and clypel-
anterior view with apices of arms angled or curved lus embrowned (Fig. 12B) or half of clypeus with arcs

dorsad. Narrow in lateral view. (Figs 19J, 20C) or weakly marked or unmarked (Figs 2B,
. Valve obtusely triangular. 12C, 15E, 22G, 24E, F). Arcs sometimes extended dor-
. Pygofer lobe rounded, edentate, scattered macrose- __ sally into crown.

tae distally, anal tube incised half way, anterior apo- Morphology. Head. Angle at crown in both sexes

demes absent. 100—113°, broadly rounded to face, disc smooth, ante-

10. Subgenital plate triangular, commonly with rior margin of head shagreened, H. ramosa brachypter

uniseriate macrosetae, rarely absent. with fine or coarsely shagreened margin or finely rugose

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16

in H. ramosa macropter, H. alacaudella sp. nov. and
H. bualacauda sp. nov.

Ocellus and interocular distance. Ocellus small, dis-
tance to eye more than three times its diameter (ocellus
diameter/interocular distance 0.16—0.41), ocellus diame-
ter 23-38 um, interocular distance 77-159 um.

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 sub-
macropter and brachypter, represented by few, fine super-
ficial 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. buala-
cauda sp. nov. (Fig. 1SA—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 submac-
ropter, 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 sub-
macropter variable, smaller than tegmina, with narrow,
folded jugal lobe (Fig. 31) or very small scale (H. hap-
sistylis 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. ramo-
sa with large, folded jugal lobe, when open, wider than
tegmina (Fig. 41).

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 (AV,) sometimes slightly
longer, thicker, darker than intercalary setae, anterome-
dial (AM, ) 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, some-
times weakly sclerotized; in lateral view anterobasal mar-
gin produced anteriad, in dorsal view paired triangular
or rounded profile, sometimes desclerotized, anterior pro-
cess 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

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Michael Stiller: Revision of Hadroca (Cicadellidae)

crown to apex of tegmina 1.9-4.3 mm (smallest male and
female in H. hapsistylis sp. nov. 1.8 mm, in H. buala-
cauda 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 um; interocular distance 77-159 um;
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/pro-
notum 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 1n ante-
rior 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 sub-
base or medially (Fig. 23D, E), usually edentate, some-
times with single, desclerotized, digitate, narrow, anterior,
subapical process (Fig. 7N, R, S, U). Preatrium rarely short
or commonly reduced. Gonopore apical to subapical, an-
terior (Figs 9A-N, 18G, 23D, E, 25F), rarely anterior and
posterior (Figs 13], J, 181, 21G, H). Orientation vertical in
genital capsule (Figs 7B, C, 13A, 18B, 21C, 23B, 25A).
Curvature of shaft for all specimens, represented as a seg-
ment 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. tre-
ichroa 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 spe-
cies (H. alavittata sp. nov., H. alacaudella sp. nov.) with
wider subapex than medial width, 1.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, 1.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 um? in all
specimens, smallest area in H. treichroa sp. nov. 12-18
um?, largest area in H. alacaudella sp. nov., 38-44 um’;
perimeter in all specimens 1.21—2.35 mm,

Style. Distally with narrow, short base, preapical lobe
ventral, apophysis commonly short (in H. hapsistylis

Dtsch. Entomol. Z. 70 (1) 2023, 13-54

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, 130, 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, 251). 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, 1.¢., 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,
211, J, 23F, 25G), rarely transverse (Fig. 13P), articulated
to aedeagus. Variable extent of desclerotized margins me-
dially 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; wid-
er 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 mar-
gin sometimes rugulate (Figs 71, 18C, D), or desclerotized
(Fig. 13C). Macrosetae commonly uniseriate, submar-
ginal, basally and apically shorter than medially, length
89-223 um, basal width 8—11 um; 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 round-
ed, 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 com-
monly 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.

17

Pygofer lobe. Edentate, in lateral view with posterior
margin broadly rounded, broadly merged with pygofer, scat-
tered long 134—244 um and short 56-88 um macrosetae.

Female.

Sternite VIT. Commonly wider than long, rectangular
to trapezoid, or almost longer than wide in H. treichroa
sp. nov. (Fig. 25J, K), posterior margin varied, either sub-
linear (Figs 8T, 231), 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 80, 180, P,
25J, K) or longer than wide (Fig. 8N, P, Q, R), or with-
out 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 um.

Valvula 2. Distal third to half, forked, dorsally ser-
rate, 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 1m-
bricate, 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, sculp-
ture 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, 1.e., microtrichia many, few, or short
lines, or absent (Fig. 61, J), most uniform in H. hapsistylis
sp. nov. (Figs 13U, V, 141).

Valvifer 1. In dorsal view, margins free; in lateral view
anterior margin broadly rounded, posterior margin nar-
rowly rounded, apices acute (Figs 8K—M, 25L) or blunt
(Fig. 17H), or both states, 1.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 es-
pecially 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 consid-
ered 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 ter-
minalia define the genus.

The generic feature of the aedeagus is the strict com-
bination of the C-shape, uniform sclerotization, short

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dorsal apodeme and reduced preatrium. Additionally, the
aedeagus is considered edentate or rarely with single, de-
sclerotized, 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; Curvo-
stylus Davies, 1987, species 0.81—0.87; Flavorubivola-
tus Stiller, 2021, species 0.41—0.53, F. curtiverpus Still-
er, 2021, 0.49-0.60, F. tensus Stiller, 2021, 0.45-0.51,
FE. 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. subspini-
verpus Stiller, 2021, 0.38-0.47; Tzitzikamaia Linnavuorl,
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 curva-
ture 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 curva-
ture. 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, 1.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. alavit-
tata 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 11, 3B, H, 4B, D).

Bloemia hieroglyphica (Naudé, 1926) is similar to
Hadroca in color, shape and size (Fig. 23A—F), and sim-
ilar curvature of the aedeagal shaft, but not uniformly
sclerotized, 1.e., with lateral margins weakly sclerotized,

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Michael Stiller: Revision of Hadroca (Cicadellidae)

base strongly sclerotized, apex anteriorly, posteriorly and
apex and subapex medially desclerotized (Fig. 5I-L). Ad-
ditionally, in B/oemia the dorsal apodeme is reduced, pre-
atrium 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, 1.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 obser-
vations 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 Hadro-
ca 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 7zitzikamaia 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 spe-
cies 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 7zitzikamaia resembling the narrow
incision as in Figs 3J, 8S. Differentiation between Basu-
toia 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-feed-
ing 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.

Hadroca ramosa (Naudé)
Figs 1-9, 26B

Euscelis ramosus Naudé, 1926: 65-66.
Hadroca ramosa (Naudé), Theron, 1974: 161-162.

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19

Table 1. Morphologic differences and similarities between the brachypterous form of H. ramosa (Naudé), Basutoia brachyptera

Linnavuori and 7Zitzikamaia species.

Tzitzikamaia spp Hadroca ramosa brachypter
color mottled, spotted, speckled light green, light brown, rarely reticulate
exposed abdominal segments 5-6 5-6 3-5
style apophysis laterally sublinear curvate ventrad

aedeagal shaft
shaft tubular

pygofer lobe long macrosetae seriate

apex with paired process, sublinear,

apex edentate, shaft compressed apex edentate, shaft tubular

long macrosetae scattered

dorsal apodeme of aedeagus
male tergite X

about half as long as shaft

third as long as distal part
subgenital plate triangular, lateral and medial

margins convergent; macrosetae
long, distal, submarginal; style

reaching half way

long macrosetae scattered
about half as long as shaft

anteroventral process less than one

about one third as long as shaft

anteroventral process less than one | anteroventral process about half as
triangular, lateral and medial
margins convergent; macrosetae

right-angled triangular, lateral
margin sinuous or sublinear;
macrosetae medial, marginal; style
near base

short, medial, submarginal, style
reaching half way

connective length greatest/width 1.10-1.46 1.06—1.62 1.54—2.40 (Hadroca spp); 1.81-
greatest 2.27 Hadroca ramosa brachypter
sternite VII ligula absent, sinuous, wide or _| provisionally considered shallowly | ligula absent, wide notch, either
narrow V-shaped notch; narrow bilobate deep or shallow; rarely narrow,
deep notch similar to that in deep notch (Fig. 8S).
Fig. 8S.

habitat Grassland and Fynbos Biomes Grassland Biome Fynbos Biome
Diagnosis. or without median line (Figs 2L, M, 3A—D). Nymph color

1. Stramineous, ochraceous, often with reticulate pat-
tern, light green to yellow green.

2. Tegmina brachypterous, submacropterous, posteri-
or margin broadly rounded or truncate, or macrop-
terous, with appendix and apex narrowly rounded.

3. Aedeagal shaft elongate, strongly curvate, preatri-
um reduced, sometimes with single, subapical, an-
terior narrow process.

4. Style apophysis in dorsal view sublinear, in later-
al view apophysis short, apex curved ventrad. The
apex of the subgenital plate is equidistant to apex of
pygofer lobe.

5. Female sternite VII posterior margin with recessed,
variable ligula or sublinear, or V-shaped with nar-
row, parallel-sided notch.

Etymology. Ramosus, Latin, full of branches (Brown
1954), for the reticulate color pattern of the tegmina, gen-
der feminine.

Color. Male, female and nymph.

Dorsum. Small circular marks (Fig. 1E, G, H) or trans-
verse 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 stra-
mineous (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),

as in Fig. 2I-K.
Morphology. Male and female. 7Jegmina.

1. Brachypter rare (36 males and 29 females), with teg-
mina about as long as wide (length/width 1.1—1.2)
(Figs 2L, M, 3A—H), hind wing reduced (Fig. 31),
(length/width 0.6, ratio tegmina length/width to
hind wing length/width 0.5—0.6).

2. Tegmina submacropterous [see Zahniser (2021) for
terms] common (206 males and 96 females), lon-
ger 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).

3. 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—P).

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. 41).

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

Measurements. Male.

Brachypter (n=35). Apex of crown to apex of tegmi-
na 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 um; interocular distance
66-117 um; crown angle 116—124°; crown length/crown

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20 Michael Stiller: Revision of Hadroca (Cicadellidae)

Figure 1. A-O 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.

length next to eye 1.19-1.31; head width/pronotum width —length/pronotum width 0.31—0.34; crown length/head
1.03—1.06; ocellus diameter/interocular distance 0.29— width 0.27-0.31; crown length/pronotum width 0.28—
0.60; crown length/pronotum length 0.86—0.98; pronotum 0.32; length to tegmina/length to abdomen 0.62-0.67.

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Dtsch. Entomol. Z. 70 (1) 2023, 13-54 21

Figure 2. A-M 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 VU, Clanwilliam; G. Sternite VU,
Dwarsrivier; H. Sternite VII, Dwarsrivier; I. Nymph, Driehoek; J. Nymph, Fisantekraal; kK. Nymph, Driehoek; L—M, brachypter,
male, Driftsands. L. Dorsally; M. Laterally. J-M. Scale bars: 1 mm.

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22 Michael Stiller: Revision of Hadroca (Cicadellidae)

Figure 3. A-O Hadroca ramosa (Naudé) male and female brachypter and submacropter, habitus images, hind wing, face and termi-
nalia 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.

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Dtsch. Entomol. Z. 70 (1) 2023, 13-54

23

Figure 4. A-I 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, Clan-

william. A—D, H, I. Scale bars: | mm.

Submacropter (n=142). Apex of crown to apex of
tegmina 2.72—3.43 mm; apex of crown to apex of abdo-
men 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 um; in-
terocular distance 69-96 um; 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/in-
terocular 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 tegmi-
na 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 um; interocular distance 108-122 um;
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/pro-

notum 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; prono-
tum width 1.49-1.63 mm; ocellus diameter 29-43 um;
interocular distance 67-105 um; 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/in-
terocular 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 teg-
mina 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 um; interocu-
lar distance 70-102 um; crown angle 103-116°; crown
length/crown length next to eye 1.34—-1.55; head width/

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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 tegmi-
na 4.94-6.10 mm; apex of crown to apex of abdomen
4,044.92 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 um; interocular distance 107-125 um;
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/pro-
notum 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, dorso-
anteriad. 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 dam-
aged (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 tm”, in submacropter 25—
35 um’, in macropter 24—37 um? 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 great-
est/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 SE, 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 great-
est 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. 71). Macrosetae basally and distally short-

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Michael Stiller: Revision of Hadroca (Cicadellidae)

er than medially, uniseriate, submarginal. Three states of
length/width ratios:

Submacropter: length/width 1.28—1.69, 4—7 macrose-
tae, length 111-190 um, 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 um, 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 um, 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 macrop-
ter, length/width respectively 0.36—0.45, 0.36—0.49, 0.38—
0.46; angle, respectively 111—127°, 100-—119°, 105—120°.

Pygofer lobe. In \ateral view, rarely shorter or com-
monly longer than greatest width (length/width 1.0—1.3),
posterior margin rounded. Macrosetae length in macrop-
ter 126—288 um, submacropter 101—222 um, brachyptera
68-173 um (Fig. 7A—D). Shorter macrosetae arbitrarily
65-94 um.

Female.

Sternite VII, Variable, either (commonly) with nar-
row 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 sublin-
ear (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, rectangu-
lar (length/width 0.40.5), posterior margin with narrow
recess (notch width/sternite VII greatest width 0.5—0.6).

Valvula 3. Macrosetae 22-44 um, in Driftsands speci-
mens (n=4) 37—53 um.

Valvula 2. Nine dissected specimens (Caledon, Ce-
darberg, Dwarsrivier, Fisantekraal, Hottentotsholland,
Fisantekraal, Jonkershoek, Ladismith, Stellenbosch) with
serrate apex (Fig. 6D, E), six (Cedarberg, Driehoek and
Dwarsrivier, Jonkiespoort) edentate, with or without lat-
eral, 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 cir-
cular structures and microtrichia (Fig. 61) 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: “Stel-
lenbosch | Jonkershoek | 17-12-22 | F.W. Pettey || type
(red paper) || HOLOTYPE | Euscelis | ramosa | Naudé
1926 (red paper) | Hadroca | ramosa | (Naudé) | Det. J.G.
Theron”, SANC.

Dtsch. Entomol. Z. 70 (1) 2023, 13-54

25

Figure 5. A—J Hadroca ramosa (Naudé) and Bloemia hieroglyphica (Naudé), male genitalia images and line drawings with speci-
men localities. A-G, H. ramosa. A. Aedeagus, Hottentotsholland, laterally; B. Aedeagus, Hottentotsholland, dorsally; C. Connec-
tive, Hottentotsholland, dorsally, DIC; D. Connective, Hottentotsholland, dorsally, bright field; E. Style, Hottentotsholland, dorsal-
ly; F. Style, Hottentotsholland, laterally; G. Aedeagus & connective, Hottentotsholland, laterally; H-J, 5. 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: 100um.

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 Slag-

boom, near Ceres in the Agterwitsenberg the only lo-
cality where all three forms were recorded, 1.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.

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Michael Stiller: Revision of Hadroca (Cicadellidae)

=
. oe ie
:

—— oo
tr
_~
na Rf mee Bes
pa DG he

— \ Fee > > 7%
. < ry ne ee tae
ie ee ee ~
— EN AE I nage ane b

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Figure 6. A-J 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: 50um.

dez.pensoft.net

Dtsch. Entomol. Z. 70 (1) 2023, 13-54

27

Ree a= SiS So

~~.

!
I
!
i}
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4
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i

a

Figure 7. A-V 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. Gif-

berg; 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 ante-
riorly. P. Copied from Theron, 1974; Q. Clanwilliam; R. Cedarberg; S. Cedarberg; T. Stellenbosch; U. De Hoop; V. Stellenbosch.

The aedeagal shaft in anterior or dorsal view of 70 ex-
amined 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 com-

monly clearly depicted ventral curvature of the apophy-
sis. 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

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28

Michael Stiller: Revision of Hadroca (Cicadellidae)

Figure 8. A-U 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. Drichoek; R. Copied from Theron, 1974; S. Swellendam; T. Driftsands; U. Driehoek.

and arm length was made to the sclerotized part of the
Y-shaped frame.

The female sternite VII was ligulate in 21 submac-
ropters and all macropters (Figs 4G, 80), 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.

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The female sternite VII was least typical in brachypters,
and did not resemble that of the submacropter or macrop-
ter. Furthermore, examination of specimens of 7zitzika-
maia 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 col-
lected by vacuum on /mperata cylindrica in 2004, but not

Dtsch. Entomol. Z. 70 (1) 2023, 13-54

29

Figure 9. A-N Hadroca ramosa (Naudé) male, aedeagus, apex, laterally or anteriorly with specimen localities. A—M, lateral-
ly. 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.

found in 2022 at the same locality on a number of shrubs.
Tzitzikamaia is well represented in SANC with 246 re-
cords with more than 800 brachypterous specimens, half
of the records without host or habitat, the rest divided ap-
proximately equally between grass and forbs or grass and
ten tree or shrub families, with seven records on specif-
ic grasses. Distribution models and point distribution for
Tzitzikamaia show a closer association with the Grassland
Biome (Fig. 30A—C). The aedeagal shaft bears paired, api-
cal 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.

Hadroca hapsistylis sp. nov.
https://zoobank.org/8098B568-D5B0-4454-82B9-24263FBCOD4D
Figs 10-14, 26C

Diagnosis.

1. Tegmina submacropterous, posterior margin round-
ed, stramineous, ochraceous, or light green to yel-
low green.

2. Aedeagal shaft elongate, strongly to weakly cur-
vate, preatrium reduced.

3. Style apophysis in dorsal view sublinear, in lateral
view apophysis about half as long as base, strongly
arched dorsad, apex curved ventrad.

4. Subgenital apex equidistant to apex of pygofer lobe.

5. Female sternite VII posterior margin with wide,
rounded, deep or shallow notch.

Etymology. Named in Greek for the arched apophysis
of the style, arch, Hapsis, and pillar, for the style, sty/is,
gender feminine.

Color. Male, female & nymph. Crown with few, paired
small circular or elongate marks (Fig. 10C, D, E, H) or
extensively marked (Fig. 10A, B) or unmarked (Fig. 10F,
I-K) and light green to yellow-green (Fig. 11A, B) (green
color form) or ochraceous to stramineous (Fig. 1O0F, H-K)
(light color form) or dark brown (Fig. 11 A—E, G, L) (dark
color form). Tegmina with brown (Fig. 10A—C, E, G) to
dark brown (Fig. 10D, L) reticulations or light green to
yellow green (Fig. 11A, B), sometimes with distal veins
embrowned (Fig. 11B). Face with clypeus and clypel-
lus embrowned (Fig. 12B) or ochraceous (Fig. 12C).
Nymphs, late instar, Fig. 11G, H, brown, early instar,
stramineous (Fig. 11J—L), or dark brown (Fig. 111).

Morphology. Male.

Tegmina. Tegmina submacropterous (Figs 10A—F,
H, J, 11A-C), brachypterous (Fig. 10G), length/width
1.6—2.0, length 1.3—1.7 mm, width 0.8—0.9 mm; posterior
margin broadly rounded.

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30 Michael Stiller: Revision of Hadroca (Cicadellidae)

Figure 10. A-L 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: | mm.

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Dtsch. Entomol. Z. 70 (1) 2023, 13-54 Hl

Figure 11. A-L 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, Nietge-
naamd; J. Nymph, Oudtshoorn, large; K. Nymph, Oudtshoorn, medium; L. Nymph, Oudtshoorn, small. A—L. Scale bars: | mm.

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32

Michael Stiller: Revision of Hadroca (Cicadellidae)

Figure 12. A-F 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. 11E), length/
width 2.1—2.5, length 0.4—0.6 mm long, width 0.2—0.3 mm.

Female. Tegmina. Submacropterous, not extended
beyond apex of abdomen (Fig. 10I, K), brachypterous
in Fig. 10 L, length/width 1.6—-2.0, length 1.3-1.9 mm,
width 0.8—1.0 mm; posterior margin broadly rounded.

Hind wing. Reduced (Fig. 11F), elongate, length/
width 2.0—2.7, length 0.4—0.7 mm, width 0.2—0.3 mm.

Chaetotaxy. AV 6-8 setae, IC 8—12 setae.

Measurements. 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 um; intero-
cular distance 97-122 um; 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 tegmi-
na 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 um; interocular
distance 104-136 um; 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

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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.

Terminalia. Male.

Aedeagus. Shaft edentate, widely curvate
(Fig. 13D, E), apex wider or narrower than medially
(Fig. 13F, G), gonopore narrow, V-shaped (Fig. 13H, I,
M, N). Shaft of Graaf Reinet specimens minutely papil-
lose, otherwise glabrous. Curvature depicted by segment
defined by chord and height 139-151°. In lateral view,
area 13—23 um’, perimeter 1.138—1.969 mm; greatest
straight line length from atrium to apex of dorsal apo-
deme/greatest straight line length from atrium to apex of
shaft 0.35-0.42.

Style. Elongate, narrow, preapical lobe reduced,
apophysis elongated, arched dorsad, apex directed pos-
teroventrad, arch in situ vertical or partially lateral, fine
denticulation ventrolaterally on base of arch. Length
greatest/width greatest 2.96—-3.93 (Fig. 13B, C, ventrally,
in situ, M, N, laterally, O, dorsally); length apophysis/
length to apex medial arm 0.66—1.15. Arched apophysis
sometimes visible in open genital capsule (Fig. 12A).

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. 13P). In anterior
view with apices of arms angled dorsad.

Subgenital plate. Apex variable, either rounded or
acute, desclerotized or uniformly sclerotized, sometimes
dorsoapical margin with fine, long setae; lateral margin

Dtsch. Entomol. Z. 70 (1) 2023, 13-54

33

Figure 13. A-V 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 VI, Brakwater; T. Valvifer 1,
Warmwaterberg; U. Valvifer 2, Wiedouw; V. Valvifer 2, sculpture, Wiedouw.

convex. Length to width 1.1—1.6, 4-6 macrosetae, length
84-149 um; angle at apex of subgenital plate, by trigo-
nometry 31—42° (Fig. 13B, C).

Pygofer 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. Mac-
rosetae 141—205 um long (Fig. 13A).

Female.

Sternite VII, Narrow, transversely rectangular, with
wide deep or shallow rounded notch (Figs 12D-F, 13Q-
S). Posterior margin with lateral length/medial length
1.8—3.6, transversely rectangular (length/width 0.4), pos-
terior margin with wide recess (notch width/sternite VII
greatest width 0.60.7).

Valvula 3. Macrosetae length 16—30 um (Fig. 14H).

Valvula 2. Basally trough and crest denticulate
(Fig. 14E, F), apex denticulate dorsally (Fig. 14F).

Valvula 1. Sculpture granulose distally, strigate basal-
ly (Fig. 14A—D).

Valvifer 2. Length/width 1.6—2.1 (Fig. 13U). Sculpture
as in Figs 13V, 141.

Valvifer 1. Length/width 2.6—2.9 (Figs 13T, 14G), ex-
tensively sculptured (Fig. 14).

Material examined. 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,
SANC, BMNH, INHS.

Type specimen. 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 | Preronia |
?succulenta | Asteraceae || SANC Pretoria | Database No.
| CCDL26403”.

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Michael Stiller: Revision of Hadroca (Cicadellidae)

J aN 4 Anke oe e Por © a : LF,

Figure 14. A—J 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 um.

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Dtsch. Entomol. Z. 70 (1) 2023, 13-54

Paratypes. 2343.3, 16492, 92 nymphs, total speci-
mens 491.

Remarks. This was the smallest of the known species
of Hadroca, with similarly wide range of color patterns as
in H. ramosa, 1.e., ochraceous, stramineous, light green to
yellow green. Most characteristic was the long and strong-
ly arched apophysis of the style, and orientation medioven-
trad 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 distinc-
tion 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 preapi-
cal lobe, whether ventral or lateroventral, marked this in-
tersection. The subgenital plate was more variable than in
other species of Hadroca. The apex in 69 examined spec-
imens was clearly desclerotized in 12, weakly desclero-
tized 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 be-
tween 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 sclero-
tized 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 Erio-
cephalus sp., Euryops ?oligoglossus, Felicia °filifolia,
Helichrysum tricostatum, Pentzia incana, Pentzia sp.,
Pteronia incana, P. paniculata, Pteronia sp., Rosenia sp..,
Asteraceae, Phylica cryptandroides, P. oleaefolia, Rham-
naceae and Zygophyllum sp. Zygophyllaceae. Map of
potential natural distribution in Fig. 26C with large red
circles indicating point distributions.

Hadroca bualacauda sp. nov.
https://zoobank. org/2D 103BDA-95C5-4CBE-ABDE-170C A5F57842
Figs 15-18, 26D

Diagnosis.

1. Tegmina macropterous, posterior margin with nar-
row, acuminate extension, light brown, oblique
band medially.

2. Aedeagal shaft short, strongly curvate, preatrium
reduced.

3. Style apophysis in dorsal view sublinear, in lateral
view short, apex curved ventrad.

4. Subgenital apex equidistant to apex of pygofer lobe.

30

5. Female sternite VII posterior margin with short,
wide ligula, recessed in deep or shallow V-shaped,
wide notch.

Etymology. Named in Latin, for the large specimen
with the wing tail, bu-, prefix meaning large, ala, wing,
cauda, tail, gender feminine.

Color. Male, female & nymph. Dorsum usually un-
marked. Tegmina with some cells whitish, sometimes
with brown reticulation, distally embrowned, veins light
brown (Fig. 15A—C). Face unmarked (Fig. 15E). Nymph
color as in Fig. 15D with abdomen darker than head, pro-
notum and thorax.

Morphology. Male. Tegmina. Macropterous
(Fig. 15A, C), (length/width 2.9-3.2, length 4.14.8 mm,
width 1.34—1.5 mm); posterior margin narrowly extended
with numerous cross veins (Figs 15A, C, 18R).

Hind wing. Reduced (Fig. 180), elongate
(length/width 3.4-4.0, male 2.9-3.5 mm long, 0.8-
1.0 mm wide).

Female. Tegmina. Macropterous (Fig. 15B), length/
width 2.9-3.3, length 4.3—-5.1 mm, width 1.4-1.6 mm;
posterior margin narrowly extended with numerous cross
veins (Figs 15B, 18R).

Hind wing. Reduced (Fig. 18P), elongate, length/
width 3.44.0, length 3.2—3.8 mm, width 0.9-1.0 mm.

Chaetotaxy. AV 9-14, IC 8-10, single AV, seta.
(Fig. 15G, H).

Measurements. Male. (n=41). Apex of crown to apex
of tegmina 5.02—5.95 mm; apex of crown to apex of abdo-
men 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 um; interocular
distance 175—207 um; 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/interocu-
lar 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/pro-
notum width 0.31—0.35; length to tegmina/length to ab-
domen 1.24—1.35.

Female. (n=63). Apex of crown to apex of tegmi-
na 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 um; interocular
distance 188-227 um; 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.

Terminalia. Male.

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36

Michael Stiller: Revision of Hadroca (Cicadellidae)

Figure 15. A-H 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 VI, Garies; G. Profemur, Tulbagh; H. Caps, 1.e., Profumur, Gifberg. A—-D. Scale bars: 1 mm.

Aedeagus. Shaft edentate, narrowly curvate
(Fig. 18E, F), segment angle 175—189°, short, apical-
ly narrower than medially, gonopore circular, anteriad
(Fig. 18G), rarely extended posteriad, shallow in Van
Rhynsdorp specimen, deeply in Dwarsrivier specimen
(Fig. 181). Curvature depicted by segment defined by
chord and height 167—181°. In lateral view, area 28-44

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um’, perimeter 1.142—1.742 mm; greatest straight line
length from atrium to apex of dorsal apodeme/great-
est straight line length from atrium to apex of shaft
0.39-0.50.

Style. Apophysis short (length apophysis/length great-
est 0.1-0.2), straight, length greatest/width greatest 1.5—
1.9 (Fig. 18K, L).

Dtsch. Entomol. Z. 70 (1) 2023, 13-54

%

Ne le ee

%, » “ ‘> a
an ee, o
% Cp ata

Figure 16. A-K 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, Biesies-
fontein; 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 um.

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38 Michael Stiller: Revision of Hadroca (Cicadellidae)

LONG

‘
¥

Figure 17. A-L Hadroca bualacauda sp. nov., ovipositor images with specimen localities. A. Valvula 1, narrow apex, Biesiesfon-
tein; B. Valvula 1, wide apex, Biesiesfontein; C. Valvula 3 with valvifer 2, Biesiesfontein; D. Valvula 2 with valvifer 2, Biesies-
fontein; 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 um.

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Dtsch. Entomol. Z. 70 (1) 2023, 13-54

39

Figure 18. A-S 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, Dwarsri-

vier; 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 short-
er than arms (length stem/length arms 1.0—1.5), length
greatest/width greatest 2.0—2.7 (Fig. 18M, N).

Subgenital plate. Lateral margin concave, apex
narrow, variably elongated, densely rugulose; length/
width 1.3—1.9, 4-8 macrosetae, length 44-183 um; an-
gle at apex of subgenital plate, by trigonometry 29-37°
(Fig. 18C, D).

Pygofer lobe. In lateral view longer than greatest
width (length/width 1.1—1.4), posterior margin rounded.
Macrosetae 44—178 um long (Fig. 18B).

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 I15F, 180, P). Posteri-
or margin slightly recessed (lateral length/medial length
1.1-1.2), transversely rectangular (length/width 0.4—0.5),

posterior margin with wide recess (notch width/sternite
VII greatest width 0.6—0.7).

Valvula 3. Whole part in Fig. 17C. Macrosetae length
22-38 um, 1-2 irregular rows (Figs 16D, 17G).

Valvula 2. Whole part in Fig. 17D. Distally serrate
(Fig. 16C, F, G), base sclerotized, desclerotized area me-
dially between apex and base.

Valvula 1. Whole part in Fig. 17A, B. Sculpture apical-
ly granulose (Fig. 17F) and basally strigate (Figs 16A, B,
LTE, 1).

Valvifer 2. Length/width 2.5—2.9, sculpture small tri-
angular scales (Figs 16J, K, 17I-K).

Valvifer 1. Length/width 1.8—2.2, distal margin blunt
(Figs 16E, H, I, 17H).

Material examined. Type locality. Holotype male.
South Africa, Northern Cape province, Biesiesfontein
Farm south Springbok, —29.75, 17.93, 29 Sep.—3 Oct.

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40

2002, 710 m, M. Stiller leg., sweeping, Pteronia divaric-
ulata, Asteraceae, CCDL18292, SANC.

Type specimen. 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°45'S, 17°56'E |
29.1x.—3.x.2002 | 710 m M. Stiller | swept off | Preronia
| divariculata | Asteraceae | SANC — Pretoria | Database
No. | CCDL18292”.

Paratypes. 5926,
total specimens.

8422, 39 nymphs, 183

Remarks. This species was larger than the other spe-
cies 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 eluci-
dated 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. 18C,
D. The depth of the notch in the female sternite was
variable, as in Fig. 180, P. The acuminate extension of
the tegmina had more cross veins (Fig. 18S) than in H.
alacaudella sp. nov. (Fig. 23P). Point distribution and
potential natural distribution are in Fig. 26D. Twenty
two records were without an associated plant, and 17
records had the following plant associations (family,
genus, species):

Anacardiaceae, Searsia undulata, Asteraceae:
Berkheya canescens, Dimorphotheca cuneata, Erioceph-
alus africanus, E. punctulatus, E. racemosus, E. umbel-
lulatus, Euryops multifudus, Gorteria personata, Pentzia
sp., Pteronia divariculata, Aizoaceae, Galenia africana,
Ebenaceae, Diospyros ramulosa,; Fabaceae, Wiborgia
mucronata Menispermaceae, Antizoma miersiana. How-
ever, 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).

Hadroca alavittata sp. nov.
https://zoobank.org/4AC8DDF 1-ADB2-4A95-92F2-25B8C5E28F0C
Figs 19-21, 26E

Diagnosis.

1. Tegmina submacropterous, posterior margin round-
ed, light brown, oblique band medially.

2. Aedeagal shaft elongate, curvate, preatrium re-
duced.

3. Style apophysis in dorsal view curvate laterad, in
lateral view short, apex with slight ventral curvature.

4. Subgenital apex extended beyond posterior margin
of pygofer lobe.

5. Female sternite VII posterior margin sinuous.

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Michael Stiller: Revision of Hadroca (Cicadellidae)

Etymology. Named in Latin for the brown V-shaped
band across the tegmina, ala, wing, vittata, ribbon, band,
gender feminine.

Color. Male & female. Crown with paired, transverse,
light brown marks apically, medially and subbasally, pro-
notum with some marks (Fig. 19A—E, H). Face in male
(Fig. 20C) and female (Fig. 19J) with 3-4 wide, dark
brown arcs distally, face basally pale ochraceous, with
amorphous small spots in clypellus and genae. Tegmina
with brown veins and reticulation within cells, medially
with oblique V-shaped, dark brown band across both teg-
mina (Figs 19A—F, H, 20A, B).

Morphology. Male. Tegmina. Submacropterous
(Fig. 19B, D, E), about as long as abdomen, (length/width
1.5—1.7, length 1.7—2.0 mm, width 1.1—1.2 mm); posteri-
or margin broadly rounded (Fig. 19B, D, E).

Hind wing. Reduced (Fig. 21P), elongate (length/
width 3.44.0, length 2.9-3.5 mm, width 0.8—1.0 mm).

Female. Tegmina. Submacropterous (Figs 18A, C, F,
H, 20, A, B), about as long as abdomen, (length/width
1.6-1.7, length 2.0—2.2 mm, width 1.2—1.3 mm); posteri-
or margin broadly rounded (Fig. 19A, C, H).

Hind wing. Reduced (Fig. 21P), elongate (length/
width 3.44.0, length 3.2—3.8 mm, width 0.9—1.0 mm).

Chaetotaxy. AV 7-9, IC 9-11.

Measurements. Male. (n=39) Apex of crown to apex
of tegmina 2.70—2.98 mm; apex of crown to apex of abdo-
men 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 um; interocular
distance 98-121 um; 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; pro-
notum 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 um; interocular distance
110-130 um; 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.

Terminalia. Male. Aedeagus. Shaft widely curvate,
segment angle 127—134°, apically wider than medially,
gonopore v-shaped anteropostriad (Fig. 21G, H). Cur-
vature depicted by segment defined by chord and height
127-134°. In lateral view, area 31-36 um?, perimeter
1.863—2.771 mm (Fig. 21K, L); greatest straight line
length from atrium to apex of dorsal apodeme/greatest
straight line length from atrium to apex of shaft 0.31-0.46.

Dtsch. Entomol. Z. 70 (1) 2023, 13-54

Al

Figure 19. A—J 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 O.1—0.2), curved or angled posterolaterad
(Fig. 21M, N), in lateral or dorsolateral or ventrolateral
view (Fig. 21N), required to distinguish ventrad cur-
vature (as in Figs 8F, H, I 18L, 23H), length greatest/
width greatest 1.7-1.9.

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. 211, J).

Subgenital plate. Triangular, apex narrowly rounded,
length/width 1.4—1.6, macrosetae absent, fine, narrow se-
tae subapically dorsally and on ventrolateral margin; apex

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42 Michael Stiller: Revision of Hadroca (Cicadellidae)

Figure 20. A—J Hadroca alavittata sp. nov., male and female, images, tegmina, face and ovipositor with specimen localities. A. Teg-
mina, reflected light, female, Swellendam; B. Tegmina, transmitted light, female, Swellendam; C. Face, male, Swellendam; D. Valv-
ifer 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 um (D-J).

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Dtsch. Entomol. Z. 70 (1) 2023, 13-54

43

R , |
;

ae U S
CE Rg A 8 OF

Ww ee X

Figure 21. A-X 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, ventral-
ly, 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.

of subgenital plate extended beyond posterior margin of
pygofer lobe, angle at apex of by trigonometry 31—36°
(Figs 19G, 21D).

Pygofer lobe. In \ateral view shorter than greatest
width (length/width 0.6—0.8), posterior margin acute.
Macrosetae 46—86 um long (Fig. 21C).

Female.

Sternite VII. Transversely rectangular, posterior
margin sinuous with wide, shallow notch (Figs 19],

21Q, R). Posterior margin sinuous with wide shallow
or deep recess (lateral length/medial length 1.1—1.3),
length/width 0.6—0.7, notch width/sternite VII greatest
width 0.6-0.7.

Valvula 3. Macrosetae length 23-41 um (Fig. 21U, X).

Valvula 2. Apex serrate, fine teeth in trough, rounded
teeth as in Figs 20H, I, 21 W.

Valvula 1, Lanceolate (Fig. 21V), sculpture strigate
(Fig. 20E-G).

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Ad

Valvifer 2. Length/width 2.3—2.9 (Fig. 21S). Sculpture
as in other species, e.g., Figs 13V, 141.

Valvifer 1. Length/width 2.2—2.8 (Figs 20D, 21T),
sculpture with microtrichia basally (Fig. 20J).

Material examined. Type locality. Holotype male,
South Africa, Eastern Cape province, Cradock, —32.166,
25.616, 19 Jan. 1984, J.G. Theron leg., sweeping, Aspara-
gus sp., Asparagaceae CCDL28305, SANC, BMNH, INHS.

Type specimen. Holotype male, glued to triangle card,
pinned, with genitalia in microvial pinned to specimen.
Original label “South Africa | Cradock | 19.1.1984 | J.G.
Theron || Asparagus sp., Asparagaceae || SANC Pretoria |
Database No. | CCDL28305”,

Paratypes. 392.4, 292 9, total specimens 68.

Remarks. 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 ven-
tral paired process and the glabrous subgenital plate were
unique. Least typical was the style with the compressed,
laterally curved apophysis in dorsal view (Fig. 21M),
and the apex with minimal ventral curvature in lateral
view (Fig. 21N). The configuration of the aedeagus and
its uniform sclerotization conforms to that of the species
of Hadroca. It also appeared to be associated with wild
asparagus. Modelled potential natural distribution is in
Fig. 26E from eight records indicated by yellow squares.

Hadroca alacaudella sp. nov.
https://zoobank.org/7B6 16E8 1-539 1-46DF-885B-6069AF5AB8D9
Figs 22, 23, 26E

Diagnosis.

1. Tegmina macropterous, posterior margin with
narrow, dark brown, acuminate extension. Hind
wing reduced.

2. Aedeagal shaft elongate, widely curvate, paral-
lel-sided in lateral view, preatrium reduced.

3. Style apophysis in dorsal view sublinear, in lateral
view short, apex curved ventrad.

4. Subgenital apex equidistant to apex of pygofer lobe.

5. Female sternite VII posterior margin sublinear.

Etymology. Named in Latin, for the small specimen
with the wing tail, ala, wing, cauda, tail, diminutive suf-
fix —ella, gender feminine.

Color. Male, female & nymph. Dorsum usually
unmarked, ground color light brown. Tegmina smoky
translucent, apex embrowned with numerous light brown
veins (Fig. 22A—D). Nymph light brown, ochraceous nar-
row median line (Fig. 22E).

Morphology. Male. Tegmina. Macropterous
(Figs 22A—D, L, K, 23P), (length/width 3.7-4.0, length
3.2-3.4 mm, width 0.8-0.9 mm); posterior margin nar-
rowly extended with numerous cross veins (Fig. 22J, K).

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Michael Stiller: Revision of Hadroca (Cicadellidae)

Hind wing. Reduced (Fig. 21L), elongate (length/
width 3.1—3.7, length 1.7—1.9 mm, width 0.5 mm).

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. 22J, K).

Hind wing. Reduced, elongate, length/width 3.7—
4.0, length 2.5—2.8 mm, width 0.7 mm (as in male,
Fig. 22L).

Chaetotaxy. Male AV 7-9 setae, female AV 8—9 setae,
male and female, IC 7—10 setae.

Measurements. 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 um; intero-
cular distance 116-132 um; 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 tegmi-
na 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 um; interocu-
lar distance 140-153 um; 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.

Terminalia. Male.

Aedeagus. Shaft narrowly curvate, segment angle
144-157°, apically wider than medially, dorsal apodeme
angled anteriad, gonopore convoluted to V-shaped, sub-
apically between anterior and posterior margin. In lat-
eral view, area 38-44 um’, perimeter 1.494—2.325 mm
(Fig. 23D, E); greatest straight line length from atrium to
apex of dorsal apodeme/greatest straight line length from
atrium to apex of shaft 0.38-0.49.

Style. Apophysis short (length apophysis/length great-
est 0.1-0.2), straight, length greatest/width greatest 2.4—
2.7 (Fig. 23G, H).

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. 23F).

Subgenital plate. Lateral margin concave; length/
width 1.5—1.8, 5—7 macrosetae, length 60-98 um; an-
gle at apex of subgenital plate, by trigonometry 29-33°
(Fig. 23C).

Dtsch. Entomol. Z. 70 (1) 2023, 13-54

i a i

Figure 22. A—L 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.

Pygofer lobe. In lateral view longer than greatest width
(length/width 1.4—-1.6), posterior margin rounded. Mac-
rosetae, short setae length 42-62 um, long setae length
83-134 um, longest seta 165 um, apices of long setae
damaged by KOH (Fig. 23B).

Female.

Sternite VII. Rectangular, posterior margin straight to
slightly sinuous (Figs 22H, I, 231). Posterior margin lat-
erally and medially similar length (lateral length/medial
length 1.0), length/width 0.6, posterior margin wide, width
across posterior margin/sternite VII greatest width 0.6—0.7.

Valvula 3. Macrosetae length 23—41 um (Fig. 23J).

Valvula 2. Apex serrate (Fig. 23K), fine teeth in trough,
rounded teeth as in Fig. 20H, I.

Valvula 1, Lanceolate (resembling Fig. 21V), sculp-
ture strigate (as in Fig. 20E-G).

Valvifer 2. Length/width 2.3—2.9 (Fig. 23N), sculpture
in Fig. 230.

Valvifer 1. Length/width 2.2—2.8 (Fig. 23M, L).

Material examined. 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, SANC;

Type specimen. Holotype male, glued to triangle
card, pinned, with genitalia in microvial pinned to speci-
men. Original label “R.S.A. Klondyke | farm Bo Swaar- |
moed, —33.308 | 19.577, 28.1. | 2022, M. Stiller || sweep
| Senecio sp. | Asteraceae || SANC Pretoria | Database#
CCDL / 28915”.

Paratypes. 143:3, 472 2, 7 nymphs, total specimens 68.

Remarks. Hadroca alacaudella sp. nov. and H. buala-
cauda sp. nov. had the acuminate extension of the poste-
rior 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 2

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46 Michael Stiller: Revision of Hadroca (Cicadellidae)

Figure 23. A—P 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.

Table 2. 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 _ in lateral view (Fig. 18E, F). The sternite VII of the fe-
species. In H. alacaudella sp. nov. the aedeagus was male in H. alacaudella sp. nov. had the posterior margin
widely curvate, and wider distally than medially in lateral sublinear, and in H. bualacauda sp. nov. it was ligulate
view (Fig. 23D). In H. bualacauda sp. nov. the aedeagus with a wide, V-shaped notch. Distribution on the map in
was narrowly curvate and narrower distally than medially | Fig. 26E from one locality indicated by a red circle.

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Dtsch. Entomol. Z. 70 (1) 2023, 13-54

Hadroca treichroa sp. nov.
https://zoobank. org/375ED020-3347-4363-8864- 183448 1E1698
Figs 24, 25, 26E

Diagnosis.

1. Tegmina submacropterous, rounded posterior
margin, with whitish, light brown and dark brown
markings.

2. Aedeagal shaft elongate, sublinear, preatrium pro-
duced, about as long as dorsal apodeme.

3. Style apophysis in dorsal view sublinear, in lateral
view short, apex curved ventrad.

4. Subgenital plate apex equidistant to apex of pygo-
fer lobe.

5. Female sternite VII posterior margin with short,
wide, recessed ligula.

6. Valvula 2 with short, square teeth.

Etymology. Named in Greek, for the three colors on
the tegmina, treis, three; chroa, color, gender feminine.

Color. Male & Female. Crown unmarked or with small
red dots, pronotum unmarked or with light brown marks
(Fig. 24A—D). Tegmina with three colors: brown to dark
brown reticulations, interspersed with whitish opaque,
light brown and dark brown cells (Fig. 24A—D, G).

Morphology. Male. Tegmina. Submacropterous
(Fig. 24A—C), about half as long as abdomen, 3—4 abdomi-
nal segments exposed, length/width 1.5—1.7, length 1.2—1.4
mm, width 0.8—0.9 mm; posterior margin broadly rounded.

Hind wing. Reduced (slightly larger than in Fig. 11E),
elongate, length/width 0.8—-2.3, length 0.4—0.8 mm,
width 0.4 mm.

Female. Tegmina. Submacropterous (Fig. 24D, G),
length/width 1.2—1.9, length 1.4—-1.5 mm, width 0.6-1.1 mm.

Hind wing. Reduced (slightly larger than in Fig.
11F), elongate, length/width 0.8—2.3, length 0.8 mm,
width 0.4 mm.

Chaetotaxy. AV 6-8, IC 7-9.

Measurements. 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; pro-
notum width 1.02—1.06 mm; ocellus diameter 14 um; in-
terocular distance 104-115 um; 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/in-
terocular 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 tegmi-
na 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

47

1.10—1.14 mm; ocellus diameter 13—21 um; interocular
distance 111-125 um; 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/interocu-
lar 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/pro-
notum width 0.36—0.39; length to tegmina/length to ab-
domen 0.63—0.66.

Terminalia. Male.

Aedeagus. Shaft sublinear, slightly curvate, segment
angle 100—130°, shaft with apical and medial width sim-
ilar, dorsal apodeme subbasal, preatrium produced, go-
nopore subapical, V-shaped (Fig. 25F). In lateral view,
area 13-17 um’, perimeter 1.138—-1.969 mm; greatest
straight line length from atrium to apex of dorsal apo-
deme/greatest straight line length from atrium to apex of
shaft 0.42—0.51 (Fig. 25C—E).

Style. Apophysis short, length apophysis/length great-
est 0.1-0.2, straight; length greatest/width greatest 1.5—
1.8 (Fig. 25H, I).

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. 25G).

Subgenital plate. Triangular, length/width 1.26—1.62,
macrosetae length 57—77 um; angle at apex of subgenital
plate, by trigonometry 32—38° (Fig. 25B); 2-3 macrose-
tae, length 55-57 um; apex rugose.

Pygofer lobe. In lateral view longer than greatest
width (length/width 1.1—1.4), posterior margin rounded.
Macrosetae length 59-110 um (Fig. 25A), 6-12 short
macrosetae 53-80 um, 5—8 long macrosetae 99-123 um
usually uniseriate.

Female.

Sternite VII, Variably rectangular, posterior margin
with short, wide, shallowly recessed ligula, lateral mar-
gins rounded (Figs 241, 25J, K), ligula length/width 0.24—
0.48. Lateral length/medial length 1.07—1.14, greatest
length/greatest width 0.59-0.69, notch width/sternite VII
greatest width 0.34—0.45.

Valvula 3. Macrosetae length 21-34 um.

Valvula 2. Serrate distally, with teeth square, close-set,
fine sculpture at base of tooth (Fig. 24J), sometimes teeth
weakly developed.

Valvula 1. Sculpture
ovoid basally.

Valvifer 2. Length/width 2.46—2.97, sculptured area
with 7-10 circular pore-like structures (Fig. 25M, N),
sometimes with 1-2 setae.

Valvifer 1. Length/width 1.83—2.05, rounding dorsally
and ventrally similar, anterior and posterior margin acute
(Fig. 25L).

Material examined. 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., sweep-
ing, Passerina truncata subsp. truncata, Thymelaeaceae,
CCDL18930, SANC.

granulose apically and

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48 Michael Stiller: Revision of Hadroca (Cicadellidae)

Figure 24. A—J 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, denticu-
lation, Wiedouw. A-D, G, scale 1 mm, J. Scale bars: 50 um.

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Dtsch. Entomol. Z. 70 (1) 2023, 13-54

49

Figure 25. A-N 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. Ae-
deagus, dorsally, Wiedouw; F. Aedeagus, apex, dorsally, Wiedouw; G. Connective, Wiedouw; H. Style, dorsally, Wiedouw; I. Style,
laterally, Wiedouw; J. Sternite VII, Wiedouw; K. Sternite VI, Clanwilliam; L. Valvifer 1, Wiedouw; M. Valvifer 2, Wiedouw;
N. Valvifer 2, sculpture, Wiedouw.

Type specimen. Holotype male, glued to triangle
card, pinned, with genitalia in microvial pinned to speci-
men. Original label “South Africa | Western Cape | Prov.
Wiedouw | Farm foot | Gifberg Pass | SE Van Rhyns-
dorp | 31°44’S, 18°46’E | 3-10.x.2002 120 m| M. Stiller
leg. || swept off | Passerina | truncata | subsp. trunca-
ta | Thymelaeaceae || SANC Pretoria | Database No. |
CCDL18930”.

Paratypes. 53.3, 24929.

Remarks. Described from one whole and six dissect-
ed 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 aedea-
gus was least typical for Hadroca, with the preatrium pro-
duced more and the shaft least curvate. This configuration
approached that of 7zitzikamaia, but without apical pro-
cesses. At present the aedeagi of species of 7Zitzikamaia
have single, apical paired processes. The six dissected ae-
deagi of H. treichroa sp. nov. do not show signs of dam-
age at the apex which could suggest broken processes.
The ventral curvature of the style apophysis correspond-
ed well to that of the other Hadroca species. The ligulate
female sternite VI 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. 26E indicated
by three green triangles.

Conclusion

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. 26B, D, E) and H. hapsistylis sp. nov. in the
Western Cape, Eastern Cape and Northern Cape provinces
(central, south- and north-western parts of South Africa)
(Fig. 26C). Hadroca alavittata sp. nov. appears to be as-
sociated with wild asparagus based on limited records and
specimens.

Specimens of the species of Hadroca vary greatly in
size, color and shape of the posterior margin of the tegmi-
na. Two species, 1.e., H. bualacauda sp. nov. and H. ala-
caudella 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 re-

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Michael Stiller: Revision of Hadroca (Cicadellidae)

Figure 26. A—E Distribution maps and potential natural distribution models. A. Bloemia hieroglyphica; B. H. ramosa; C. H. hapsi-

Stylis 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.

duced, 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 cur-
vature. Species recognition is based subjectively on color
(Figs 1A—O, 2L, M, 3A—H, M, N, 4A—D, 1OA-L, 11 A—D,
19A-F, H, 24A—D, G) and shape (Figs 15A—C, 22A-D).
Variability in the shape of the posterior margin of the

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sternite VII is ascribed to an adaptation to different ovi-
position plants, most notable in H. ramosa and H. hapsi-
Stylis. Similar variation was noted in Colistra acapitatus
(Stiller and Webb 2022), Retevolatus flexiverpus (Stiller
2021), Geelus dundraad and G. platdraad (Stiller 2020),
Discolopeus diacaenus (Stiller 2019), Bretega quinispiv-
erpa (Stiller 2016), three species of Pravistylus (Stiller
2010a), Vilargus trunculicans (Stiller 2010b) and E/gi-
nus cultellus (Stiller 2009). The submacropter Bloemia
hieroglyphica (Naudé) is similar in shape (Fig. 28A—F)
to submacropters of Hadroca ramosa, but the aedeagus

Dtsch. Entomol. Z. 70 (1) 2023, 13-54 ey

Figure 27. A—M 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. 7einopterus microphallus Stiller, 2011,
Eastern Cape province, dorsally; F. Male; G. Female; H. 3. brachyptera, male, genital capsule, posteroventrally; I-L. 7zitzikamaia
species, male; I. Belfast; J. Kirkwood; K. Sani Pass; L. Jonkiespoort; M. Goniagnathus brachypterus Linnavuori, 1978, female,
DNyala. A—M. Scale bars: | mm.

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52

Michael Stiller: Revision of Hadroca (Cicadellidae)

Figure 28. A-F Bloemia hieroglyphica (Naudé, 1926), male and female, habitus images with specimen localities. A. Female,
Jonkiespoort; B. Male, Aberdeen; C. Male, New Bethesda; D. Male, Oudtshoorn; E. Male, Ladismith; F. Male, Jonkiespoort. A-F.
Scale bars: 1 mm.

has three levels of sclerotization and some differences in
profile. More confounding are the brachypters of H. ra-
mosa (Fig. 3A—H) which bear some resemblance to spe-
cies of 7zitzikamaia Linnavuori (Fig. 27I-L), although all
four species in 7Zitzikamaia have the aedeagal shaft with
apical, paired processes. A few indeterminable examined
specimens of the latter lacked these paired processes.

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Acknowledgements

Colleagues, reviewers and editors are thanked for their
contributions to the improvement of the manuscript. Per-
mits issued by CapeNature (CNN44-87-16619 (2022),
0056-AAA008-00042 (2015), 0035-AAA004-00688
(2011), 001-202-00022 (2004)) facilitated acquisitions.

Dtsch. Entomol. Z. 70 (1) 2023, 13-54 53

Figure 29. A—-N Basutoia and 7Zitzikamaia species, male and female terminalia, line drawings. A-H, M, 7: adspersa Theron, 1974.
A. Head, pronotum and tegmina; B. Face; C. Genital capsule, laterally; D. Subgenital plate; E. Style; F. Connective; G. Aedea-
gus, laterally; H. Aedeagus, posteriorly; I. 7) irrorata Linnavuori, 1961, aedeagus, laterally; J. 7 /ongipennis Linnavuori, 1961,
aedeagus, laterally; K. 7. si/vicola Linnavuori, 1961, aedeagus, laterally; L. B. brachyptera Linnavuori 1961, aedeagus, laterally;

M. 7: adspersa, sternite VII; N. 7: /ongipennis, sternite VII.

KRM

\
7

Figure 30. A—D Distribution maps and potential natural distribution models. A. Basutoia brachyptera Linnavuorl, 1961; B. 7Zitzika-

m]
t

4
oe

v \ ¢
“5
:

maia adspersa Theron, 1974; C. 7. irrorata Linnavuori, 1961; D. 7. si/vicola Linnavuori, 1961.

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54

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Supplementary material 1

Additional material examined

Authors: Michael Stiller

Data type: species data

Copyright notice: This dataset is made available under
the Open Database License (http://opendatacommons.
org/licenses/odbl/1.0). The Open Database License
(ODbL) is a license agreement intended to allow us-
ers to freely share, modify, and use this Dataset while
maintaining this same freedom for others, provided
that the original source and author(s) are credited.

Link: https://doi.org/10.3897/dez.70.90851.suppl1