ZooKeys 369: 25-41 (2014) A peer-rev iewed open-access journa I

doi: 10.3897/zookeys.369.6506 RESEARCH ARTICLE #ZooKey

WWwW.ZOO keys.o rg Launched to accelerate biodiversity research

On cicadas of Hyalessa maculaticollis complex
(Hemiptera, Cicadidae) of China

Xu Wang'', Masami Hayashi**, Cong Wei!

| Key Laboratory of Plant Protection Resources and Pest Management, Ministry of Education, Entomological
Museum, Northwest A&F University, Yangling, Shaanxi 712100, China 2 Department of Biology, Faculty of
Education, Saitama University, Saitama 338-8570, Japan

T Attp://zoobank.org/02020680-CDB9-41A 1-8FB8-B0662142FE35
t Attp://zoobank. org/C5SAA9237-7C33-4279-8E64-AEEDESA3DE72
§ Aitp://zoobank. org 1 BCA52BD-B926-40A7-B5B3-2A51CF211AAE

Corresponding author: Cong Wei (congwei@nwsuaf.edu.cn)

Academic editor: A. Sanborn | Received 28 October 2013 | Accepted 25 December 2013 | Published 13 January 2014
http://zoobank. org/90065 CF2-5622-4DD8-B6EF-F25E503BF75D

Citation: Wang X, Hayashi M, Wei C (2014) On cicadas of Hyalessa maculaticollis complex (Hemiptera, Cicadidae) of
China. ZooKeys 369: 25-41. doi: 10.3897/zookeys.369.6506

Abstract

The genus Hyalessa China is reviewed based on the discovery of male of the type species H. ronshana China
as well as the description of one new species (H. batangensis sp. n.). The species formerly included in the
genus Sonata Lee are removed to Hyalessa as new combinations. Intraspecific variations of H. maculaticollis
are enumerated based on materials collected from various locations from China. The identity of Sonata

and the systematic placement of Hyalessa are discussed. A key to all species of Hyalessa is provided.

Keywords

Cicadomorpha, Oncotympana, Sonata, variability, morphology, new combination

Introduction

The Oncotympana was established by Stal in 1870 as a subgenus of Pomponia Stal,
1866. Distant (1905a) raised Oncotympana to generic level and redescribed it. Later,
Ishihara (1961) established the tribe Oncotympanini to accommodate it, and Hayashi

Copyright Xu Wang et al. This is an open access article distributed under the terms of the Creative Commons Attribution International License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

26 Xu Wang et al. / ZooKeys 369: 25-41 (2014)

(1978, 1984) reviewed this genus. Chou et al. (1997) recorded four species of On-
cotympana from China, but two of them (O. stratoria and O. virescens) were just listed
in their checklist. Recently, Lee (2010) found that nine species of Oncotympana from
the continental East Asia, Japan and India aren’t congeneric with the type species O.
pallidiventris, so he established the genus Sonata to accommodate these species, i-e.,
S. fuscata (Distant), S. maculaticollis (Motschulsky), S. ella (Lei & Chou), S. expansa
(Walker), S. mahoni (Distant), S. melanoptera (Distant), S. obnubila (Distant), Sonata
stratoria (Distant), and S. virescens (Distant). More recently, Lee (2011) established
the tribe Sonatini in the subfamily Cicadinae to contain the genus Sonata, and syn-
onymised Oncotympanini with Cicadini and included the Philippine genera Oncotym-
pana Stal and Neoncotympana Lee, 2011 in the subtribe Oncotympanina of Cicadini.
However, Hayashi (2011) synonymised Sonata Lee with Hyalessa China, and trans-
ferred Oncotympana maculaticollis to the genus Hyalessa. Hayashi and Saisho (2011)
recognized Oncotympana fuscata (Distant), the type species of Sonata, as a synonym of
Hyalessa maculaticollis (Motschulsky).

In the present paper we review the genus Hyalessa based on the discovery of male of
the type species H. ronshana China and the description of one new species, H. batan-
gensis sp. n. from Southeast China. In addition, we transfer the species of Sonata to
Hyalessa, bringing the species number of Hyalyssa to 10. Furthermore, the phenotypic
variability of H. maculaticollis, the most widely distributed species among its conge-
ners, is investigated based on materials collected from different locations.

Materials and methods

This study is mainly based on specimens deposited in the following institutions ab-
breviated in the text as follows:

NWAFU Entomological Museum, Northwest A&F University, Yangling, China
BMNH. The Natural History Museum, London, UK
MNHN - Muséum National d'Histoire Naturelle, Paris, France

External morphology was observed using the Olympus SZX10 stereomicroscope,
and photographed with a Nikon Coolpix P100 digital camera. The pygofer was care-
fully extracted from the terminal abdominal segments of relaxed specimen and ob-
served and photographed using a Scientific Digital micrography system equipped with
an Auto-montage imaging system and a QIMAGING Retiga 4000R digital camera
(CCD). The extracted pygofer, if necessary, was dissected and placed in 10% KOH
boiled for 2—5 minutes, washed, and transferred to glycerin for observation, and the
aedeagus were photographed using CCD similarly. Terminology for morphological
features follows that of Moulds (2005). All measurements are in millimeter.

The type specimens of the new species are deposited in the Entomological Mu-

seum, Northwest A&F University (NWAFU), Yangling, China.

On cicadas of Hyalessa maculaticollis complex (Hemiptera, Cicadidae) of China 27

Systematics

Family Cicadidae Latreille
Subfamily Cicadinae
Tribe Cicadini Latreille, 1802

Genus Hyalessa China, 1925
http://species-id.net/wiki/Hyalessa

Pomponia (Oncotympana) Stal, 1870: 710. Type species: Pomponia (Oncotympana) pal-
lidiventris Stal

Oncotympana: Distant 1905: 60, 70. Type species: Pomponia (Oncotympana) pallidi-
ventris Stal

Sonata Lee, 2010: 20. Type species: Oncotympana fuscata Distant

Type species. Hyalessa ronshana China.

Body small to large, head including eyes wider than pronotum. Anterolateral pro-
notal collar not dentate. Medial pronotal collar about one-fourth to one-third the
length of inner area. Wings hyaline; fore wing with 8 apical cells, with fuscous spots
at bases of apical cells second, third, fifth, and seventh, a marginal series of minute
pale fuscous spots near apices of longitudinal veins to apical cells in most species; hind
wing with 6 apical cells. Male operculum wider than long, lateral margin roundly pro-
duced laterad, overlapped or nearly touching to the other one centrally. Male abdomen
slightly shorter than distance from head to cruciform elevation. Posterior margin of
male abdominal tergite II] much wider than mesonotum. Timbal concealed by timbal
cover in dorsal view; timbal cover globolised, projecting beyond corresponding lateral
margin of abdomen. Aedeagus thick and curved apically; apex with pair of sclerotized
lateral processes and pair of (membranous) saccate hooks between them. Uncal lobe
large, separated from the other one distally or connected to the other one from near
base to subapex.

Remarks. ‘This genus is closely similar to Oncotympana Stal in habitus, but differs
from the latter in the following characteristics: pronotum about or more than twice as
long as head; anterolateral pronotal collar not dentate; male operculum shorter than
wide but very large; uncal lobes bifurcated; aedeagus very thick, with apex with a pair
of sclerotized lateral processes and a pair of (membranous) saccate hooks between them.

Key to the species of Hyalessa

1 Fore wing with a faint spot merely at bases of apical cells second, third, fifth,
ABC SCveIithiy FOS ECEING byt. heteui Mts de: Me ute eect Ge Ae hatte alba seentcarale st Aces? 2
- Fore wing with a distinct large fuscous spot at bases of apical cells second,
third, fifth, and seventh and a marginal series of minute pale fuscous spots
near apices of longitudinal veins to apical cells, respectively 0.0.0... eee eee 3

28

nis

Ae

ON

Xu Wang et al. / ZooKeys 369: 25-41 (2014)

Unzcal lobes separated from the other one from middle of uncus, with poste-
rior margin rounded and outer margin weakly convex.............. H. ronshana
Uncal lobes connected to each other closely from near base to apex, with
lateral margin slightly concave basally and convex to distal margin ...............
nk ale gach ee MD oes tao ra a ech sg Peden apccik ng to H. batangensis sp. n.
Male opercula well separated from each other, or close to each other but not
OVER Birt U nln dultesh nue, kash sant wcen ad cotta Satoh oR AND Sonim ent oni? é
Male opercilasoverlappincumiediall ys .1....-.-ncrseevsoncevasonspetisneectadheolesaseaiacihedts 5
Male opercula entirely dark greenish ochraceous, and abdomen (both in
males and females) with broad olivaceous or ochraceous markings................

EA Mists nceatrs Mee tle ese ine eetausi Ben Snss bin es Meiacwstichs Sebel H. mahoni comb. n.
Male opercula entirely much infuscated to black.. H. melanoptera comb. n.
Body-smaller body leneth< 26:0 mini eo. ease ctereesaee ce caeevecsnsbddannevenstasts 6
Bodyelarger, bedyleneths 30; Ona ick seassans 22s ea saiteentis tohanitencnat boat lle dadlatie 9
Male opercula not reaching hind margin of abdominal sternite I.......0..00....
saci cia te cule bee iar leh Ou uchomweecees Cree oviduaic one eta H. stratoria comb. n.
Male opercula extending to hind margin of abdominal sternite II............... yh
Mesonotum green or ochraceous; opercula pale yellowish green «0.0.0.0...
ee AY ad ey RO ie a NS a H. virescens comb. n.
Mesonotum black; opercula restaceous:. 2025... tpsecter ent entree tre 8
Body smaller, body length about 13.0 mm; uncal lobes separated from each
OUNer ALOM Teak DASE Ol UN CUS sc tes autho aed areas H. expansa comb. n.
Body larger, body length about 25.0 mm; uncal lobes separated from each
eid overesazeyeaveoatlo a lomepager tle Cet one MMi wan nei ee H. ella comb. n.

Head narrower, about 0.73 times as wide as pronotum; uncal lobes broad,
with outer margins convexly sinuate and inner margins nearly straight.........

Fetish Ca Se attroto nn eetiea tes tte setae tes usin. Mideast) H. obnubila comb. n.
Head broader, about 0.79 times as wide as pronotum; uncal lobes long, with
inner margins parallelto- outer Margins ciisscsccssscewcsseciveseses H. maculaticollis

Hyalessa ronshana China
http://species-id.net/wiki/Hyalessa_ronshana

Figs 1-3

Material examined. Holotype: 9 (BMNH), China: Yunnan Prov., 31.VII.1922. 1¢
(NWAFU), China: Hutiaoxia, Xianggelila County, Yunnan Prov., 2.VII.2007, coll.
Cong Wei; 1033, 429 (NWAFU), China: Hutiaoxia, Xianggelila County, Yun-
nan Prov., 27.VII.2007, coll. Cong Wei; 744, 822 (NWAFU), China: Hutiaoxia,
Xianggelila County, Yunnan Prov., 6.VIII.2010, coll. Meng Zhang; 19 (NWAFU),
China: Hutiaoxia, Xianggelila County, Yunnan Prov., 6.VIII.2010, coll. Silong Xu;
13° (NWAFU), China: Hutiaoxia, Xianggelila County, Yunnan Prov., 7.VII.2010,
coll. Meng Zhang.

On cicadas of Hyalessa maculaticollis complex (Hemiptera, Cicadidae) of China 29

Figure |. Hyalessa ronshana (female, holotype), habitus, dorsal view.

Measurements of types. (18 Jd, 12 2°). Body length: male 31.3—37.8, female
32.6-35.7; fore wing length: male 45.6-50.3, female 46.6-53.3; fore wing width:
male 15.2—17.6, female 15.4—17.5; width of head including eyes: male 11.0—12.0, fe-
male 10.4—12.0; pronotum width (including pronotal collar): male 14.0-16.0, female
14.0—-16.2; mesonotum width: male 11.7—13.3, female 11.5—13.4.

Description of male. Body almost black, with short yellow-green hairs.

Head (Fig. 2A, C) about 0.77 times as wide as pronotum; eyes fuscous, ocellus red.
Postclypeus moderately swollen, black, with greenish transverse grooves on each side;
lateral margin greenish. Anteclypeus black, with yellowish green medially. Rostrum
long, extending to posterior trochanter.

Thorax (Fig. 2A, C). Pronotum generally black, with central longitudinal green-
ish yellow spot near anterior margin, smaller greenish yellow spot on disc, and central
round greenish yellow spot near posterior margin; lateral margins of pronotal collar
ampliate. Mesonotum black, with pair of greenish markings on anterior angles of cru-
ciform elevation. Metanotum and lateral part of cruciform elevation yellowish green.
Thoracic sternites greenish to black.

Legs (Fig. 2E). Black, fore femur with large ochraceous patch medially and small-
er ochraceous patch near posterior margin in lateral view. Fore tibia and mid femur
mostly black. Hind legs mostly ochraceous. Fore femur with primary spine longest and
oblique to femur, secondary spine of intermediate size and subapical spine shortest,
both angled slightly.

Wings (Fig. 2A—B). Hyaline, fore wing with distinct infuscation at bases of apical
cells second, third, fifth, and seventh; a marginal series of minute pale fuscous spots
near apices of longitudinal veins to apical cells.

Abdomen (Fig. 2A, D). Generally black dorsally, with white pollinosity between
tergite I] and II. Timbal cover black, prominently globolised. Opercula greyish green,
centrally overlapping, with rounded posterior margin extending to abdominal sternite
II. Abdominal sternites mostly black, with greenish speckle on sternite IH, VI and
VIII, sparsely covered with white pollinosity.

Genitalia (Fig. 2F, G, H). Pygofer barrel-shaped in ventral view. Uncal lobes broad

and well developed, separated from the other one from middle of uncus, with poste-

30 Xu Wang et al. / ZooKeys 369: 25-41 (2014)

Figure 2. Hyalessa ronshana (male). A habitus, dorsal view B habitus, ventral view C head and thorax,
dorsal view D head and thorax, ventral view E left foreleg, showing the spines on fore femur F male py-

gofer, ventral view G male pygofer, lateral view H aedeagus.

rior margin rounded and outer margin weakly convex. Basal lobe of uncus shorter,
ca 1/2 length of uncal lobe. Aedeagus with apical one third strongly curved ventrally,
expended subapically; sclerotized lateral processes acute, large medial (membranous)
saccate hook somewhat truncate with a pair of small lateral membranous processes
between sclerotized lateral processes.

Description of female. (Figs 1, 3). Opercula smaller than those of male, broad-
ly separated from each other. Abdominal segment IX (pygofer) greenish; ovipositor
sheath not extending beyond segment IX, posterior margin of segment VII incised at
middle. Other characteristics similar to male.

Distribution. China (Yunnan).

Remarks. Hyalessa China formerly included only the type species H. ronshana
which was established on a single female collected from Yunnan Prov., China. Re-

On cicadas of Hyalessa maculaticollis complex (Hemiptera, Cicadidae) of China 31

Figure 3. Hyalessa ronshana (female). A habitus, dorsal view B habitus, ventral view C head and thorax,
dorsal view D head and thorax, ventral view E fore and hind wings.

cently, when we investigated materials of this genus collected from different locations
from China, some specimens also from Yunnan Province were found very similar to
H. ronshana, but they can be distinguished from the holotype of H. ronshana by the
concoloured mesonotum (blackish, without paired large spots adjacent to the anterior
margin of mesonotum), the normal nodal line of fore wing (absent in the ulnar cell 3
and the medial cell), the fuscous spots at bases of apical cells second, third, fifth, and
seventh of fore wing, and a marginal series of minute pale fuscous spots near apices of
longitudinal veins to apical cells. However, the holotype of H. ronshana is an unusual
form, representing a kind of deformation on the forewing vein, i.e., the veins are some-
what asymmetric and particularly, the long nodal line presented on the ulnar cell 3 and
the medial cell. In addition, the condition of the holotype is not good in condition,
e.g., the faint markings on mesonotum are not strict, and only one very faint spot ap-
peared at the base of apical cell second of fore wing (other infuscations on the veins of
fore wing seem to be diminished due to the poor condition of the specimen). ‘This is
probably due to that the holotype was rather teneral and/or it has been deposited in the
collection for a long time. Herein, judging from the adjacency of the forementioned
materials collected from Yunnan Prov. and the holotype of H. ronshana as well as the
common characters shared by them, particularly the transverse pollinosity-like band
on base of abdominal tergite HI, coloration of veins (both upperside and underside),
maculation on fore femur, hind tibia, opercula and pygofer, we conclude that the new
materials are conspecific with the holotype of H. ronshana, and redescribe this species
based on the discovery of the male for the first time. H. ronshana is similar to H. macu-
laticollis, but can be distinguished from the latter by the generally black pronotum and
mesonotum, the rounded apex of the broad uncal lobes of male pygofer, and the shape

of apical hooks of aedeagus.

32 Xu Wang et al. / ZooKeys 369: 25-41 (2014)

Hyalessa batangensis sp. n.
http://zoobank.org/F42CAD01-CECC-408F-A4EB- 1 1B74EE23C34
http://species-id.net/wiki/Hyalessa_batangensis

Figs 4—5

Type material. Holotype: ¢ (NWAFU), China: Batang County, Sichuan Prov.,
12.VIII.2001. Paratype: 14 (NWAFU), China: Batang County, Sichuan Prov.,
12.VIII.2001.

Measurements of types. (2¢'3): Body length: 26.6—31.8; fore wing length: 37.1—
43.5; fore wing width: 12.4-15.2; width of head including eyes: 8.6-10.3; pronotum
width (including pronotal collar): 11.3—14.0; mesonotum width: 9.4—10.9.

Etymology. The species name is derived from the location of the types.

Description of male. Head (Fig. 4A, C) about 0.74 times as wide as pronotum.
Compound eye greenish brown, ocellus red. Postclypeus moderately swollen, with
black medial longitudinal fasciae and greenish yellow transverse grooves on each side.
Anteclypeus black, with yellowish green fasciae medially. Rostrum with black apex
extending to posterior trochanter.

Thorax (Fig. 4A, C). Pronotum and mesonotum almost black, pronotum with
pair of submedian markings and pair of lateral markings yellow greenish. Mesonotum
with pair of submedian markings and lateral markings respectively, or without dis-
tinct markings. Cruciform elevation black, with pair of yellow greenish markings on
anterior angles. Metanotum and lateral part of cruciform elevation yellowish green.
Thoracic sternites yellow greenish, with ochraceous patches.

Legs (Fig. 4E). Dark brown, fore femur with large yellowish ochraceous patch
medially and smaller ochraceous patch near posterior margin in lateral view. Fore tibia
and mid femur, tibia mostly dark brown. Fore femur with primary spine conical and
less angled; secondary and subapical spines erect and pointed.

Wings (Fig. 4A, B). Hyaline, fore wing with indistinct infuscation at bases of apical
cells second and third; no fuscous spots near apices of longitudinal veins to apical cells.

Abdomen (Fig. 4A, D). Black, with white pruinosity between tergite Hl and HI.
Timbal cover brownish ochraceous, circular and globose. Opercula yellow greenish,
centrally overlapping, with rounded apex extending to posterior margin of sternite II.
Abdominal sternites mostly black, with sternite VII, VHI and posterior margin of HI,
IV, V and VI yellow greenish.

Genitalia (Fig. 4F, G, H). Pygofer barrel-shaped in ventral view. Uncal lobes
connected to each other closely from near base to apex; lateral margin of uncal lobe
slightly concave basally and convex to distal margin. Aedeagus with apex curved
ventrally, expended subapically; apex with sclerotized lateral processes very broad
and rounded, pair of (membraneous) saccate hooks between sclerotized lateral
processes.

Female. Unknown.

Distribution. China (Sichuan).

On cicadas of Hyalessa maculaticollis complex (Hemiptera, Cicadidae) of China ee)

Figure 4. Hyalessa batangensis sp. n. (male, holotype). A habitus, dorsal view B habitus, ventral view
C head and thorax, dorsal view D head and thorax, ventral view E left foreleg, showing the spines on fore

femur F male pygofer, ventral view G male pygofer, lateral view H aedeagus.

Remarks. This new species can be distinguished from its congeners by the
combination of the following characters: slender body, without fuscations on veins
of fore wing, and uncal lobes connected to each other closely from near base to
apex. There are slight differences of body size, markings on mesonotum, and the
shape of aedeagus presents between the holotype and the paratype: the holotype
has a bigger body size (31.8 mm), a pair of submedian markings and a pair of lateral
markings on mesonotum, and a pair of broad, rounded sclerotized lateral processes
on aedeagus (Fig. 4); the paratype has a smaller body size (26.6 mm), without
distinct markings on mesonotum, and the sclerotized lateral processes on aedeagus
are short and acute (Fig. 5). We tentatively treat the latter as an intraspecific vari-
ation of this species, and its identity needs to be confirmed when more materials
are available.

34 Xu Wang et al. / ZooKeys 369: 25-41 (2014)

Figure 5. Hyalessa batangensis sp. n. (male, paratype). A. habitus. dorsal view B habitus, ventral view
C head and thorax, dorsal view D head and thorax, ventral view E left foreleg, showing the spines on fore

femur F male pygofer, ventral view G male pygofer, lateral view H aedeagus.

Hyalessa maculaticollis (Motschulsky)
http://species-id.net/wiki/Hyalessa_maculaticollis

Cicada maculaticollis Motschulsky, 1866: 185.
Pomponia maculaticollis Distant, 1888: 296.
Oncotympana maculaticollis: Distant 1905: 559.
Oncotympana fuscata Distant, 1905: 558.
Oncotympana coreanus Kato, 1925: 27.

Sonata maculaticollis Lee, 2010: 20.

On cicadas of Hyalessa maculaticollis complex (Hemiptera, Cicadidae) of China 35

Material examined. 1 (NWAFU), China: Mt. Emei, Sichuan Prov., 5.V.1957; 13
(NWAFU), China: Mt. Emei, Sichuan Prov., 22.VII.1974; 84 (NWAFU), China:
Mt. Emei, Sichuan Prov., 22.VII.1991; 43'4 (NWAFU), China: Mt. Emei, Sichuan
Prov., 26.VII.1991; 14 (NWAFU), China: Mt. Emei, Sichuan Prov., 23.VII.2009; 14
(NWAFU), China: Beijing, 14.VII.1961; 1d (NWAFU), China: Beijing, 18.VH.1961;
334 (NWAFU), China: Qingdao, Shandong Prov., 29.VII.2010; 334 (NWAFU),
China: Qingdao, Shandong Prov., 2.VIII.2010; 334 (NWAFU), China: Qingdao,
Shandong Prov., 4. VIII.2010; 534 (NWAFU), China: Qingdao, Shandong Prov.,
8.VIII.2010; 14 (NWAFU), China: Chunhua County, Shaanxi Prov., 6. VIII.1981;
13 (NWAFU), China: Guangdong Prov., 5.VIII.1957; 243 (NWAFU), China:
Mts. Shennongjia, Hubei Prov., 11.VHI.2004; 13° (NWAFU), China: Mts. Shen-
nongjia, Hubei Prov., 15.VHI.2004; 254 (NWAFU), China: Huoditang, Ningshan
County, Shaanxi Prov., 5.VII.2008; 24.4 (NWAFU), China: Mt. Nanwutai, Xi’an,
Shaanxi Prov., 25.VII.1951; 24 (NWAFU), China: Mt. Nanwutai, Xian, Shaanxi
Prov., 26.VIII.1957; 14 (NWAFU), China: Mt. Nanwutai, Xi’an, Shaanxi Prov.,
13.VII.1959; 14 (NWAFU), China: Mt. Tianmu, Zhejiang Prov., 15. VIII.1965; 1¢
(NWAFU), China: Mt. Tianmu, Zhejiang Prov., 26.VII.2003; 234 (NWAFU), Chi-
na: Mt. Tianmu, Zhejiang Prov., 28.VII.2003; 13 (NWAFU), China: Mt. Tianmu,
Zhejiang Prov., 29.VII.2003; 316d (NWAFU), China: Mt. Huping, Hunan Prov.,
26.VII.2013; 446.5 (NWAFU), China: Mt. Qingcheng, Sichuan Prov., 9.VIII.2013.

Main characters. Body large, head slightly shorter than base of mesonotum in dor-
sal view. Rostrum extending to the posterior trochanter. Mesonotum black with follow-
ing green markings: two large central obconical spots, three pairs of large greenish spots
around them and pair of greenish spots on each lateral margin. Abdomen black; timbal
cover ochraceous. Wings hyaline; fore wing with large fuscous spot at bases of apical
cells second, third, fifth, and seventh; a marginal series of minute pale fuscous spots near
apices of longitudinal veins to apical cells. Opercula in male broad, convex, extending
to posterior margin of second abdominal segment, and overlapping; opercula in female
smaller than those of male and broadly separated from each other. Aedeagus thick and
curve ventrally, expended subapically, with a pair of sclerotized lateral processes apically
as well as a pair of (membraneous) saccate hooks when everted.

Remarks. After examining the holotype (male) of Oncotympana fuscata Distant
preserved in the Muséum National d’Histoire Naturelle (by MH) and investigating
the intraspecific variability of H. maculaticollis, we reconfirm that H. fuscata is a junior
synonym of H. maculaticollis, as previously proposed by several authors. Among the
species of Hyalessa, H. maculaticollis has the widest range of distribution, from the
Russian Maritime Territory, Korean Peninsula, Japan to China. This species mainly
occurs in the forests, and the calling song of males is very loud with a complex trans-
position. H. maculaticollis is noted for its great intraspecific variability, including body
size, markings on thorax, timbal cover, opercula and aedeagus, which has been re-
corded by Hayashi and Saisho (2011) based on materials collected from Japan. In this
study, based on more materials collected from different locations in China, we further
investigate the intraspecific variability of this species. For details, see below.

36 Xu Wang et al. | ZooKeys 369: 25-41 (2014)

Figure 6. Hyalessa maculaticollis (male). Habitus, dorsal view, showing intraspecific variability in Chi-
nese populations. A material from Mt. Emei, Sichuan Prov. B material from Beijing C material from
Qingdao, Shandong Prov. D material from Qingdao, Shandong Prov. E material from Chunhua Coun-
ty, Shaanxi Prov. F material from Guangdong Prov. G material from Mts. Shennongjia, Hubei Prov.
H material from Huoditang, Ningshan County, Shaanxi Prov.

Intraspecific variability. Body size. (1) Medium, about 30 mm in length
(Fig. GF); (2) Large, about 36 mm in length (Fig. 6A).

Coloration of body. (1) Generally black with green or ochreous markings
(Fig. 6A, D, G, H); (2) Generally yellowish with dark ochreous markings (Fig. 6B);
(3) Variegated (Fig. 6C, E, F).

On cicadas of Hyalessa maculaticollis complex (Hemiptera, Cicadidae) of China aT

Figure 7. Hyalessa maculaticollis (male). Habitus, ventral view, showing intraspecific variability in Chi-
nese populations. A material from Mt. Emei, Sichuan Prov. B material from Beijing C material from
Qingdao, Shandong Prov. D material from Qingdao, Shandong Prov. E material from Chunhua Coun-
ty, Shaanxi Prov. F material from Guangdong Prov. G material from Mts. Shennongjia, Hubei Prov.
H material from Huoditang, Ningshan County, Shaanxi Prov.

Markings on mesonotum. (1) Mesonotum with 5 pairs of greenish spots: a pair
of very small ones near anterior margin, three large spots on disc, and a pair of very
lagre spots on lateral margins, (Fig. 6A, D); (2) Above mentioned 5 pairs of spots on

38 Xu Wang et al. / ZooKeys 369: 25-41 (2014)

Figure 8. Hyalessa maculaticollis (male). Aedeagus, lateral view, showing intraspecific phenotypic vari-
ability in Chinese populations. A material from Qingdao, Shandong Prov. B material from Mt. Tianmu,
Zhejiang Prov. C material from Huoditang, Ningshan County, Shaanxi Prov. D material from Mt.
Qingcheng, Sichuan Prov. E material from Mt. Huping, Hunan Prov. F material from Mts. Shennongjia,
Hubei Prov. G-H material from Mt. Nanwutai, Xi'an, Shaanxi Prov.

mesonotum obscure or reduced (Fig. 6C, E, G, H); (3) Mesonotum with no distinct
markings on disc (Fig. 6B, F).

Timbal cover. (1) Black (Fig. 6A, G, H); (2) Ochreous or greenish yellow, with
dark ochreous to black patch on posterior area (Fig. 6C, D, F); (3) Dark ochreous or
yellow (Fig. 6B, E).

Male opercula. (1) Black (Fig. 7A, H); (2) Light ochreous or brown (Fig. 7B, F);
(3) Variegated (Fig. 7C, D, E, G).

Shaft of aedeagus. (1) Aedeagal shaft slightly S-shaped in lateral view (Fig. 8A, B,
C, D, F, G); (2) Aedeagal shaft S-shaped in lateral view (Fig. 8E); (3) Aedeagal shaft
C-shaped in lateral view (Fig. 8H).

Apex of aedeagus. (1) Paired saccate hooks much shorter than the sclerotized
lateral processes (Fig. 8A, B, E); (2) Paired saccate hooks slightly shorter than the scle-
rotized lateral processes (Fig. 8C, D, F, G); (3) Paired saccate hooks much longer than
the sclerotized lateral processes (Fig. 8H).

On cicadas of Hyalessa maculaticollis complex (Hemiptera, Cicadidae) of China 39

Discussion

The genus Hyalessa formerly included only the type species H. ronshana which was
established on a single female collected from Yunnan Province of China. Herein, we
re-address the identity of this species based on the discovery of the male for the first
time and treat Sonata as a junior synonym of Hyalessa. This genus is retained in the
tribe Cicadini, and its tribal and sub-tribal status awaiting further phylogenetic studies.

Regarding the validity of Oncotympana fuscata, Distant (1905) stated that O.
fuscata is allied to O. maculaticollis but can be distinguished from the latter by the
coloration of body, the narrower fore wings and the broader cruciform elevation.
Kurosawa (1969) treated the populations in Korea and Far East Russia as a subspecies,
O. maculaticollis fuscata. However, O. fuscata was treated as a junior synonym of O.
maculaticollis by Chou et al. (1997). Lee (1999) applied the name O. fuscata to the
Korean population because of substantial differences in song. Recently, Lee (2008)
also treated O. fuscata and O. maculaticollis as two separated species and synonymized
O. maculaticollis fuscata with O. fuscata. More recently, Lee (2010) proposed the ge-
nus Sonata with O. fuscata as its type species. However, Hayashi and Saisho (2011)
synonymised Sonata with Hyalessa and treated H. fuscata as a junior synonym of H.
maculaticollis based on examination of related holotypes deposited in the Muséum
National d’ Histoire Naturelle.

According to Hayashi and Saisho (2011), H. maculaticollis shows a high degree of
variability in coloration. Based on more materials collected from China, we investigate
further the variability of H. maculaticollis in this study, and the results show that greater
intraspecific variations occur in this species, with respect to body size, markings on tho-
rax, coloration of timbal cover and opercula and, in particular, the morphology of ae-
deagus that has never been described in detail for this species and its allies. Remarkably,
the aedeagus of H. maculaticollis is furnished with a pair of apical sclerotized processes
as well as a pair of saccate hooks. ‘The relative lengths of the paired saccate hooks and
the sclerotized lateral processes on aedeagus may be variable due to the scalability of the
saccate hooks, which forms a continuous variation as a cline. Furthermore, the condition
of curvature at apical 1/3 of aedeagus is also variable, particularly the aedeagal shaft of
specimen collected from Mt. Nanwutai of Xi'an, Shaanxi Province (Fig. 8H) is C-shaped
in lateral view, which is unique and can be easily distinguished from others with aedea-
gal shaft S-shaped (Fig. 8A—G). We tentatively treat this specimen as a variation of H.
maculaticollis, but it may represent a new species of Hyalessa, which merits further studies
using multiple sources such as morphology, acoustics, biology and molecular data, etc.

Acknowledgement

We would like to express our sincere thanks to Dr. Stéphane Puissant, Dr. Jérdme
Sueur and Dr. Michel Boulard (Muséum National d’Histoire Naturelle, Paris, France)

40 Xu Wang et al. | ZooKeys 369: 25-41 (2014)

for offering valuable comments on the first version of this paper and providing M.
Hayashi with every support to an investigation of the material deposited in the Mu-
seum. We are indebted to Mr. M.D. Webb, who allowed C. Wei and M. Hayashi to
take pictures of identified species of the genus Oncotympanal Hyalessa deposited in The
Natural History Museum, London, UK. This work was supported by the National
Natural Science Foundation of China (Grant No. 31170360, 31093430) to CW.

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