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! Bulletin of
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THE

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| -5 JUL 1996

Entomology Series

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THE

NATURAL
HISTORY
MUSEUM

VOLUME 65 NUMBER 1 27 JUNE 1996

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World List abbreviation: Bull. nat. Hist. Mus. Lond. (Ent.)

© The Natural History Museum, 1996

Entomology Series
ISSN 0968-0454 Vol. 65, No. 1, pp. 1-103

The Natural History Museum
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London SW7 5BD Issued 27 June 1996

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Plate 1 Selenocephalinae species. 1, Drabescus ogumae (Japan, Hotoyama, BMNH); 2, D. stramineous (lectotype); 3,
D. flavicollis (holotype); 4, D. pallidus (Japan, Tsuhui, BMNH); 5, Rengatella metallica (lectotype); 6, Athysanopsis
salicis (Korea, Kwon coll.); 7, Nacula multicolor (lectotype); 8, Tenompoella difficilis (Singapore, BMNH); 9, Roxas-
ella laetisigna (Sabah, Sandakan, NMNH); 10, R. philippina (allotype); 11, R. biarcuata (lectotype); 12, R. camusi
(allotype); 13, R. losbanosa (allotype); 14, Isaca falcata (paratype, BMNH); 15, Hybrazil brani (Fiji, Viti Levu,
BPBM); 16, Mysolis flavidorsum (lectotype); 17, Parabolopona luzonensis (Philippines, Baguio, BMNH); 18, Dryado-
morpha pallida; 19, Lamia cydippe (Fiji, Viti Levu, BMNH); 20, Nirvanguina placida (holotype).

Bull. nat. Hist. Mus. Lond. (Entomol.) 65(1): 1-103 Issued 27 June 1996

A revised classification of the Asian

and Pacific selenocephaline

leafhoppers (Homoptera: LS pa he
Cicadellidae)

Y. ZHANG

Institute of Entomology, Northwestern Agricultural University, Yangling, Shaanxi
712100, P.R. China

M.D. WEBB

Department of Entomology, The Natural History Museum, Cromwell Road,
London SW7 SBD, U.K.

CONTENTS

BONITG ISIS Mees Rome ae as Seen LPC II cos canines diesen tSau te d vadasaaWaeghss spasarnh Adah odanacd Macoucao¥ss l
MARU OLS LOLA eee ates ane ne MOR ESOT Cex sao sue deusuys ss nn88es -qaaupent ««ndapeas Set-deeesscaecee oqiedeeneces 2
Classification ofvAsian and) Pacific Selenocephialitiae 7. -.<..1.....0cs.ccccccsacensacecsestnesnaeonsesnsee 2
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ISEVAGSELOMICS ice - deans aurea SOeee Cacasaie stan cancanssdentsedech eter ee ee taaeer ashen eR Cee cs toe ena. 4
ANCL ESCO HSS FECSTO NS) Ch eo ORE RE eee iain | hae es ee oa eS 2 4
NUADCCOTNONUNVES Be 30 0A ds Bos c PEE ee Maa Bec acne vaxseedassscesnevervadcdeaads sbbewuswWebscusscstethscsssanaehis 5
Sam VAS ClENOCEPMAMNAGs qiaviec -c-epsac<-scaceecortertne-veeeetatacctosetcestictactacesencebeereddbanarces 5
ING VI CW Ol CH ALAC CONS ata cantan ccd eccsusnrcneacsi<cin-Penpecemmmenteneexteeee ts sc te eeeont “cee nda eute es eee aks 5

Wy OE CHOETICLIG CHECKLIST) saree snearsccceeteacesson Steen BAUR eee epee bie eee seein hts 6

Key to tribes of Asian and Pacific Selenocephalinae .............::cccccsccsesesesceeceeseenseeneee 7
MTLCAS EEO CEP AMT teaver a: cautiect Bet os. cco Te Walch bo uayiresie vas pan dada sanddhnc doa casuanesbsthas bees 7
Minis AtraOOLODOMIMIy Yori ecpecueucdtzcrt tte. .+~- 22s saa bytie NT: Tacs gegs case aaevoaspesieueesacuendosateas )
Miirit ed) Ted HC SCUTM Tm ete cakes tate cate tae sos aa+s Seaaort este naaksestannadaagancaxgabbapac datnaseapiasarhacvsnash 22
ER CROL EL COSMO ame aen ONE SINS cs oS o cc STRAIN «25 2s anges Sega sedud snceswsplias dpuask seadeendeesdeateacsastessee 27
EN USN Ga ee snc a eI oa oR an Set Leads onan sng he sbya asp Saaasa® saa baotas ten atks spphbasns 3} ee canaas 101

Synopsis. The classification of the Asian and Pacific selenocephaline leafhoppers is
revised. The 33 genera recognised here are placed in the following three tribes:
Paraboloponini (formerly Paraboloponinae, a new senior synonym of Bhatiini, syn. n.) (28
genera), Selenocephalini (three genera) and Drabescini (two genera). Keys to these tribes
and genera are given together with a world generic check-list. Seven new genera and eight
new species are described; 31 new combinations, five generic and eight specific synonymies
are proposed. The male genitalia of most species are figured: exceptions are those species of
unknown identity or known only from females, and species previously figured or those from
the Indian subcontinent. Habitus photographs of at least one species from each genus are
provided. The following genera are transferred: Hecaloidella Osborn (from Deltocephalinae
to Selenocephalini); Sohipona Ghauri & Viraktamath, Shivipona Ghauri & Viraktamath and
Cestius Distant (from Selenocephalinae to Deltocephalinae); Gessius Distant (from
Selenocephalinae to Iassinae); Divus Distant, Mysolis Kirkaldy, Omanella Merino,
Roxasella Merino and Kutara Distant (all from Selenocephalini to Paraboloponini).

© The Natural History Museum, 1996

2

INTRODUCTION

The present work was undertaken to document the
little-known Asian and Pacific Selenocephalinae,
both as an aid to their identification and to act asa
basis for future studies. A further aim has been to
test the classification proposed by Linnavuori &
Al-Ne’amy (1983) which was based mainly on the
African fauna. However, our conclusions should be
considered preliminary, pending further work on
the large number of undescribed taxa which we
have seen but which, with a few exceptions, fall
beyond the scope of the present work.

There have been several different opinions on the
status and systematic position of the
Selenocephalinae. Workers on the European
cicadellid fauna have placed the single European
genus Selenocephalus Germar in the
Deltocephalinae, either in Selenocephalini (Ribaut,
1952) or Athysanini (Wagner, 1951 and Nast 1972).
In the most comprehensive morphological study of
the Cicadellidae Evans (1947) included 19 genera
from all parts of the World’ within
Selenocephalinae (as Selenocephalini), as a tribe of
Iassinae. This arrangement was followed in
Metcalf’s (1966) catalogue, which included a
further five genera, and was also followed in part in
Hamilton’s (1983) phylogenetic scheme. In a study
of female genitalia and external characters of
selected taxa, Hill (1969) included _ the
Selenocephalinae (as Selenocephalini) in
Deltocephalinae; seven out of the nine genera
examined, of those listed by Evans (1947), together
with three additional genera from North American,
were included in the tribe. In a revision of the
African fauna Linnavuori & Al-Ne’amy (1983)
considered the Selenocephalinae as a ‘distinctive’
leafhopper subfamily. Seven tribes were recognized
by these authors, three distributed throughout the
Old World and four (three monobasic) restricted to
Africa (Table 1). This arrangement was followed in
the recent generic catalogue of Oman et al. (1990).

In the present work we have retained the
Selenocephalinae but with some reservation as we
have found no single diagnostic character (which is
not present in some other subfamily) to define this
group. The Selenocephalinae was originally erected
without description to accommodate two
Palaearctic genera, Paramesus Fieber (now
included in Deltocephalinae) and Selenocephalus;
both possess transverse striations on the
fore-margin of the head, a feature used by most
authors (but not all — see Distant, 1908: 290), to
define the subfamily. In this context it is surprising
that Linnavuori & Al-Ne’amy (1983) included
Taneira Linnavuori and Hypacostemma Linnavuori

Y. ZHANG AND M.D. WEBB

Table 1. Distribution of Selenocephalinae tribes and
numbers of genera and species from different geo-
graphical regions.

African S.Europe Asiaand

Pacific

TRIBES Gen. Spec. Gen.Spec. Gen. Spec.
Adamini 1 43
Dwightiini 1 4
Hypacostemmini 1 5)
Taneirini 4 12

Selenocephalini 11 68 1 20 3 6

Paraboloponini 3 6 - 8) iS)

Drabescini 1 1 - - 35h

in the Selenocephalinae as both these lack
transverse striations on the fore margin of the head,
although they are present in an undescribed species
of Hypacostemma (BMNH). This character, and
the bifurcated striations (strigae) on the first
valvulae, have together been used to define the
Selenocephalinae (Linnavuori & Al-Ne’amy, 1983),
but both features are present in some other
subfamilies and neither is universal within the
group (see above and below).

CLASSIFICATION OF ASIAN AND
PACIFIC SELENOCEPHALINAE

Three of the tribes proposed by Linnavuori &
Al-Ne’amy (1983) (Selenocephalini, Bhatiini and
Drabescini) are considered applicable to the Asian
and Pacific Selenocephalinae, although none of
these is well defined and we consider Bhatiini a
junior synonym of Paraboloponini syn. nov. (see
under tribes below). The majority of Asian and
Pacific genera which we include in the subfamily
have long antennae situated from midheight of the
eyes (Fig. 7) to a point near the upper limit of the
eyes (Fig. 8) in facial view. These same genera also
have the antennal pits encroaching onto the clypeus
(except Nirvanguina) and have the tentorial
branches falcate (Fig. 21) or with a short dorsal
branch (Fig. 20). These genera are assigned to
either Drabescini or Paraboloponini. Of the
remaining Selenocephalinae, Isaca Walker, Nacula
Distant and Bhatia Distant (metcalfi) are also
included in Paraboloponini despite their slightly
lower antennae. Hecaloidella Osborn with shorter
antennae and Jambocerus gen. n. and Moluccasia
gen. n. with shorter and distinctly lower antennae,

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 3

are assigned to Selenocephalini (see under tribal
comments below).

Although it is beyond the scope of the present
work to examine in detail the African tribes we
question the status of Adamini and Ianeirini: the
former monobasic tribe is defined only on its male
genitalia and the latter on characters of the face.
The differences between these tribes and
Selenocephalini are no greater than that between
some genera of the Paraboloponini.

SUBFAMILY RELATIONSHIPS

Assuming that the Selenocephalinae was a
monophyletic group Linnavuori & Al-Ne’amy
(1983) considered that the head structure and other
characters provided evidence of its close
relationship to Penthimiinae and Acostemminae.
In fact some characters noted by these authors
apply only to certain selenocephaline tribes (in
particular to Drabescini), or are variable within
these tribes (see also ‘Review of characters’). These
characters are: fore margin of the head transversely
striated (Penthimiinae, Acostemminae and most
Selenocephalinae, except some [aneirini,
Hypacostemmini and Drabescini); antennae
situated at or above midheight of eye in facial view
(Penthimiinae, Drabescini, Dwightiini and most
Paraboloponini); epistomal suture keeled (some
Acostemminae and Drabescini); antennal ledge
strong (Penthimiini, Drabescini); forewing veins
and costal margin with whitish callosities (some
Drabescini and Acostemminae); body with hair
covering (Drabescini, Acostemminae and
Dwightiini and some Paraboloponini); fore tibia
dorsally flattened (Drabescini and some
Penthimiinae, Acostemminae and
Paraboloponini); first valvulae with strigate
sculpture (Acostemminae, most Selenocephalinae
and some Penthimiinae). To this list may be added
the following shared characters: forewing with very
narrow appendix and sometimes with reticulate
venation (Acostemminae and Hypocostemmini);
fore wing with broad appendix (Penthimiinae and
Drabescini); clypeus rugose (Drabescini and some
Acostemminae: Telopetulcus Evans and Eryapus
Evans sp.,);__—clypellus = medially ridged
(Acostemminae and Drabescini). The inclusion of
the African genus Citorus Stal in the Penthimiinae
by Linnavuori & Al-Ne’amy (1983) was given as
further evidence that Penthimiinae and
Selenocephalinae were closely related, but it is not
clear to us why this genus was excluded from the
Selenocephalinae, since it was noted as having all

the features of this group. Similarly, Jafar Kirkaldy
and Thaumatoscopus Kirkaldy have _ similar
sculpture of the first valvulae to most
Selenocephalinae (Hill, 1969) but, like most
Penthimiinae, they differ from Selenocephalinae by
their dorsal ocelli. The Acostemminae share some
characters with some Selenocephalinae (as noted
above) but, with the exception of Telopetulcus, can
be distinguished by the fusion of the male valve and
pygophore and fused bases of the subgenital plates.

The similarity of Deltocephalinae and some
Selenocephalinae is apparent both in external
appearance and in the male genitalia, while
differences that do exist are certainly no greater
than between many of the deltocephaline tribes.
The usually shagreen foremargin of the head,
Y-shaped anterior tentorial branches and
non-strigate sculpture pattern of the first valvulae
in Deltocephalinae were given as a distinction
between this subfamily and Selenocephalinae by
Linnavuori & Al-Ne’amy (1983). However, the
latter two characters are known for only a relatively
few Deltocephalinae and in one of their examples,
Phlepsius Fieber, we have found some bifurcate
striations (strigae) similar to the dorsal sculpture in
Fig. 70 and that figured by them for their
selenocephaline genus Maichewia. In addition we
have found some Selenocephalinae which lack this
feature (see below). It is possible, therefore, that the
Selenocephalinae may be either paraphyletic or
belong within the Deltocephalinae, a group which
is itself poorly defined. Further work is clearly
required on the above groups to establish their
relationships to one another.

BIOLOGY

Little is known about the biology of the
Selenocephalinae except that they are generally
arboreal and are frequently caught in light trap and
tree fogging samples. One genus (Loka Linnavuor1)
occurs on grasses in Africa and is ant attended
(Linnavuori & Al-Ne’amy, 1983: 58). The following
taxa have been recorded with host plant data.
Paraboloponini: Jamitettix guamensis Metcalf
on Morinda citrifolia (Rubiaceae); Athysanopsis
salicis Matsumura on Salix (Salicaceae); Welmaya
nigrilinea (Walker) on Shorea (Dipterocarpaceae);
Drabescoides nuchalis on Rosa rugosa (Rosaceae),
Salix and Ulmus (Ulmaceae); Favintiga camphorae
(Matsumura) on Cinnamomum — camphora
(Lauraceae); Dryadomorpha pallida Kirkaldy on
Zizyphus jujuba (Rhamnaceae); D. metrosideri on
Glochidion ramiflorum (Euphorbiaceae), Rapanea

4

sp. (Myrsinaceae), Reynoldsia sp. (Araliaceae) and
Weinmannia parviflora (Cunoniaceae); Hecaloidella
sp. indet. on Syzygium aromaticum (Myrtaceae).

Drabescini: Drabescus ogumae on Morus
(Moraceae), D. ineffectus on Quercus mongolica
(Fagaceae), D. nigrifemoratus on Lespedeza sp.
(Leguminosae).

Selenocephalini (African): Gannia Theron on
Gardenia (Rubiaceae) and Salsola esterhuyseniae
(Chenopodiaceae), Adama Dlabola on Acacia
(Leguminosae), Hypacostemma uniformis (Distant)
on Laportea alatipes (Urticaceae), H. devia Theron
on Pappea capensis (Sapindaceae) and Rhus
(Anacardiaceae),

MATERIAL EXAMINED

All available specimens from Asia and the Pacific,
with transverse striations on the fore margin of the
head and of a general Deltocephalinae-like
appearance have been examined for possible
inclusion in the Selenocephalinae. Of these, we
consider Jassus costalis Walker, with non-strigate
valvulae and very short and fine striations (almost
shagreen) on the fore margin of the head, to belong
to a new genus of Deltocephalinae: Athysanini
(new placement). Types of most of the included
taxa have also been examined. Exceptions are those
of Drabescus atratus Kato and D. fasciatus Kato,
the types of which are in the care of Kato’s family
and unavailable for study, those of D. notatus
Schumacher, Kutara nigrifasciata Kuoh and
Jamitettix katonis Matsumura, which could not be
found and D. pallidus Matsumura (EIHU) which
could not be sent because of its poor condition.

Although it was not our original intention to
describe new species we consider it desirable to
describe new species in Kutara Distant, Bhatia
Distant and Drabescus Stal for misidentified species
in the literature and in Rengatella gen. n. and Isaca
to enable the recognition of their type species (see
explanation under these genera).

For most species the male genitalia are figured,
either from the holotype or a more suitable
specimen or from a male specimen matched to a
female type. Species not figured are those species of
unknown identity or known only from the female
and those species described and figured by Webb
(1981). In addition, those species of Carvaka
Distant, Divus Distant and Megabyzus Distant
from the Indian sub-continent, currently being
revised by Dr C. Viraktamath, are omitted.

Y. ZHANG AND M.D. WEBB

DEPOSITORIES
AS Academia Sinica, Beijing, China
AM Australian Museum, Syndey, Australia

AAU Anhui Agricultural University, Anhui, China
BAU Beijing Agricultural University, Beijing, China

BMNH The Natural History Museum, London, U.K.

BPBM _ Bernice P. Bishop Museum, Honolulu, Hawaii,
US.A.

CAS California Academy of Sciences, California,
UESTAG

EIHU Entomological Institute, Hokkaido

University, Sapporo, Japan
IPE Institut fiir Pflanzenschutzforschung der
Akadamie der Landwirtschaftswissen,
Eberswalde, Germany
Institute of Zoology and Botany, Tartu,
Estonia
IZW Institute of Zoology, Warszawa, Poland

IZBT

HNHM Hungarian Natural History Museum,
Budapest, Hungary

MMB Moravian Museum, Brno, Czechloslovakia

MNHN_ Muséum National d’Histoire Naturelle, Paris,
France

MRAC_ Musée Royal de l'Afrique Centrale, Tervuren,
Belgium

NAU Northwestern
Shaanxi, China
NHMW _ Naturhistorisches Museum, Vienna, Austria

Agricultural University,

NIAS National Institute of Agro-Environmental
Scieces, Tsukuba, Japan

NMNH National Museum of Natural History,
Washington, D.C., U.S.A.

NRS Naturhistoriska Riksmuseet, Stockholm,
Sweden

QM Queensland Museum, Brisbane, Australia

RML Rijksmuseum van Natuurlijke Historie,
Leiden, Netherlands

SMTD _ Staatliches Museum fiir Tierkunde, Dresden,
Germany

SU Saitama University, Urawa, Japan

UASB University of Agricultural Sciences,
Bangalore, India

ZIRAS Zoological Institute of the Russian Academy
of Sciences, St Petersburg, Russia

ZMB Zoologisches Museum an der

Humboldt-Universitat zu Berlin, Germany

ACKNOWLEDGEMENTS. For the loan of material under
their care we thank the following: Dr S. Takagi (EIHU),
Dr C. Dietrich (formerly NMNH), Dr P. Lauterer
(MMB), Dr Cai Ping (AAU), Dr Preston (BPBM); Dr W.
Tomaszewska (IZW), Dr U. Asp6ck (NHMW), Dr Per
Lindskog (NHMYV), Dr G.B. Monteith (QM), Dr J. van
Tol (RML), Dr J. Deckert (ZMB), Dr T. Vasarhelyi
(HNHM), Dr M. Hayashi (SU), Dr A. Taeger (IPE), Dr

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 5

N. Penny (CAS), Dr M. Moulds (AM), Prof. Yang
Chikun (BAU).

We are grateful to Dr C. Viraktamath, for his helpful
communications during the course of the work, and to Dr
A. Neboiss for information on Francis Walker material in
the Museum of Victoria.

We thank the British Council for supporting the senior
author during the course of this work.

TAXONOMY

Subfamily Selenocephalinae Fieber

Selenocephalidae Fieber 1872: 10.

Selenocephalini, Evans, 1947: 217; Ribaut, 1952:
312; Evans, 1966: 244; Hill, 1969: 101; Hamilton,
1983: 21.

Selenocephalinae, Linnavuori & Al-Ne’amy, 1983:
19%

Diagnosis: Ocelli marginal; foremargin of the head
rugose or transversely striated or carinate
(exceptions: Janeira and some Hypacostemma and
Drabescini); first valvulae with dorsal strigate
sculpture pattern (exceptions: Lamia Linnavuori,
Megabyzus Distant, Sombakidia gen. n. and
Dryadomorpha Kirkaldy).

REMARKS. See above for details on validity and
classification of Selenocephalinae, its subfamily
relationships and biology.

Review of characters

The following review of characters deals primarily
with the Asian and Pacific Selenocephalinae fauna;
that of the African fauna is covered in Linnavuori
& Al-Ne’amy (1983).

Head: The head varies in length (Figs 1-4) while
there is some evidence that the nymphs of short
headed forms have longer heads (see remarks under
Paraboloponini and Drabescini). In Drabescoides
Kwon & Lee and most Kutara Distant species the
vertex is elevated posteriorly. In Dryadomorpha
Kirkaldy, Parohinka Webb and Rhutelorbus Webb
the genae of the face are weakly incurved (noted in
Fletcher & Stevens’ (1988) key to Australian
subfamilies and tribes) and visible dorsally in
Rhutelorbus, as in Scaphytopiini
(Deltocephalinae). The fore margin is transversely
striated, rugose (some Bhatia) or smooth (some
Drabescini); sometimes one or more striae are
stronger, forming carinae, with a sulcus in between.
The vertex is smooth or longitudinally striate often
with a subapical depression. Evans’s (1947) key,
couplet 33, indicated that the group had prominent

antennal ledges, although in his description of
Selenocephalinae they were described as weak. This
is an elusive character; when well developed and
carinate (Drabescini) the ledges are easily
discernable but in other Selenocephalinae they are
much less obvious, or absent altogether e.g.
Hecaloidella. In Paraboloponini (except
Nirvanguina, Fig. 15) and Drabescini the high
antennal pits encroach onto the clypeus (Figs 7-9).
The anterior tentorial branches of the head have
also been used to define some leafhopper groups
but in the Selenocephalinae these are variable (Figs
17-21) although the falcate condition (Fig. 21)
found in Drabescini and most Paraboloponini is
distinctive. The antennae vary in length and
position (see “Classification’) while the second
segment (pedicel) is often dark brown in Drabescini
and Paraboloponini. A keeled epistomal suture is
present in some Drabescus.

Legs: Despite a very extensive study of the legs
Linnavuori & Al-Ne’amy (1983) gave few
significant diagnostic features except for the
foretibia and hind femur in Drabescini (see below)
and supernumerary spines on the forefemur of
some genera (compare Figs 53 and 61). In addition
we have found that some species of Bhatia and
Kutara have the foretibia slightly flattened dorsally
which is intermediate between the distinctly
flattened foretibia in Drabescini and the more usual
rounded condition. Contrary to Hamilton (1983) in
Paraboloponini the apical spines of the hind tibia
(Fig. 59) are irregular in length (as in
Selenocephalini) and the paired apical setae of the
hind femur are similar to Selenocephalini in
position and in having prominent bases (Figs 55
and 60). In addition, the twisted hind tibia which
characterized his Scarinae (lassinae, in part,
Penthimiinae and Selenocephalinae, in part) is
variable in Selenocephalinae and also present in
Deltocephalinae. The setal formula on the fore tibia
is 1+4 (Fig. 61) but in Drabescoides and most
Kutara species it is 4+4 and on the hind femur
usually 2+2+1 (Fig. 60) but sometimes 2+1+1 or
Parle

Fore wings: The inner subapical cell of the fore
wing is open in all genera except Nirvanguina (Fig.
28) and an outer subapical cell is present in all
genera except Lamia (Fig. 27). The appendix varies
from narrow (Fig. 23) to broad (Fig. 28). An
incomplete m-cu cross vein and a cross vein
between the claval veins is present in some Kutara
species (Fig. 29).

Male genitalia. The male genitalia of
Selenocephalinae are of the basic Deltocephalinae
type but some genera lack macrosetae on the
subgenital plates or if present they are usually
arranged irregularly rather than the regular

6

arrangement found in many Deltocephalinae.
Adamini, some Paraboloponini and Hecaloidella
(Selenocephalini) are similar to a _ few
Deltocephalinae in having the aedeagus
disassociated from the connective (Figs 104, 276)
and Sombakidia (Paraboloponini) has a
racket-shaped connective (Fig. 300) as in
Paralimnini (Deltocephalinae). Several genera have
the style apical process modified from the usual
short curved tapered condition and in Omanella
Merino and Tengatka gen. n. the inner or outer
basal apophysis respectively is more elongate
compared to other genera (Figs 260, 295). A feature
of some Drabescus (Drabescini) and some
Paraboloponini is a pair of long basal aedeagal
processes (Fig. 122) and in Hecaloidella and some
Paraboloponini a posterior extension of the
connective occurs (Fig. 104). The articulation
between the connective and aedeagus is obscure in
Drabescini and Drabescoides (Fig. 216) and an
additional sclerite is present between the two in
some Paraboloponini (Fig. 114).

Female genitalia: We have examined the sculpture
of the first valvulae for most Selenocephalinae from
the study area and find it to be dorsally strigate in
most taxa to a variable extent (see Figs 63, 65),
except in Lamia where it is scale-like and in
Megabyzus, Sombakidia and Dryadomorpha, where
it is reticulate (Fig. 67). A few examples have both a
reticulate and a more distal strigate pattern
together with an intermediate pattern, similar to
Acostemminae (Figs 68-70). Most of the genera
examined have the second valvulae similar to
Bhatia (Fig. 71) but differences are found in some
genera (Figs 72-80).

GENERA REMOVED FROM THE SUBFAMILY

The following genera, appear under
Selenocephalinae or Paraboloponinae in the
generic catalogue of Oman, Knight & Nielson
(1990) but belong to other subfamilies: Cestius
Distant, belonging to Deltocephalinae, Opsiini (see
Knight, 1973); Gessius Distant, belonging to
Iassinae; Sohipona Ghauri & Viraktamath and
Shivipona Ghauri & Viraktamath belonging to
Deltocephalinae: Grypotini (see remarks under
Paraboloponini); Discocephalana Metcalf
(replacement name for Discocephalus
Kirschbaum), a genus incertae sedis.

World generic check-list of subfamily
Selenocephalinae

The taxa listed below are from Asia and the Pacific
unless otherwise indicated (see also Table 1).

Tribe Adamini Linnavuori and Al-Ne’amy
Adama Dlabola (African)

Y. ZHANG AND M.D. WEBB

subgenus Krisnella Linnavuori & Al-Ne’amy
subgenus Paracostemma Linnavuori & Al-Ne’amy
subgenus Zinjella Linnavuori & Al-Ne’amy

Tribe Drabescini Ishihara
Drabescus Stal (one African species)
Paradrabescus Kuoh syn.n.
Tylissus Stal syn.n.
subgenus Ochrescus Anufriev & Emeljanov syn.n.
subgenus Leucostigmidium Anufriev & Emeljanov
syn. n.
Rengatella gen. n.

Tribe Dwightiini Linnavuori & Al-Nea’my
Dwightia Linnavuori & Al-Nea’my (African)

Tribe Hypacostemmini Linnavuori & Al-Nea’my
Hypacostemma Linnavuori (African)

Tribe Ianeirini Linnavuori
Abimwa Linnavuori (African)
Bardera Linnavuori & Al-Nea’my (African)
Ianeira Linnavuori (African)
Kanziko Linnavuori & Al-Nea’my (African)

Tribe Paraboloponini Ishihara
Bhatiini Linnavuori & Al-Nea’my Syn.n.
Athysanopsis Matsumura
Bhatia Distant
Melichariella Matsumura syn. rev.
Koreanopsis Kwon & Lee syn. n.
Carvaka Distant
Divus Distant (new placement from Selenocephalini)
Drabescoides Kwon & Lee
Dryadomorpha Kirkaldy (S. Palaearctic and Old
World tropics)
Calotettix Osborn
Khamiria Diabola
Osbornitettix Metcalf
Paganalia Distant
Rhombopsana Metcalf
Rhombopsis Haupt
Yakunopona Ishihara
Zizyphoides Distant
Favintiga Webb
Hybrasil Kirkaldy
Isaca Walker
Jamitettix Matsumura incertae sedis
Karoseefa Webb
Kutara Distant (new placement from Ianerin1)
Lamia Linnavuori
Megabyzus Distant (new placement from Delto-
cephalinae)
Mysolis Kirkaldy (new placement from Seleno-
cephalini)
Norsia Walker
Norsiana Distant
Nakula Distant
Nirvanguina gen. n.
Oceanopona Linnavuori
Odmiella Linnavuori (African)
Omanella Merino (new placement from Seleno-
cephalini)
Parabolopona Matsumura

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS mM

Parohinka Webb

Rhutelorbus Webb

Roxasella Merino (new placement from Seleno-
cephalini)

Sombakidia gen. n.

Stenomiella Evans (African)

Tengatka gen. n.

Tenompoella gen. n.

Waigara gen. n.

Welmaya gen. n.

Tribe Selenocephalini Fieber
Angolaia Linnavuori & Al-Nea’my (African)
Distantia Signoret (African)
Foso Linnavuori & Al-Nea’my (African)
Hecaloidella Osborn (new placement from Delto-
cephalinae)
Ifeia Linnavuori & Al-Nea’my (African)
Kumba Linnavuori & Al-Nea’my (African)
Loka Linnavuori & Al-Nea’my (African)
Maichewia Linnavuori & Al-Nea’my (African)
Malendea Linnavuori & Al-Nea’my (African)
Moluccasia gen. n.
Sapoba Linnavuori & Al-Neamy (African)
Selenocephalus Germar (S.W. Palaearctic)

Levantottettix Lindberg

Tambocerus gen. n.

Key to tribes of Asian and Pacific
Selenocephalinae

1 Antennal ledge weak or absent; foretibia round or
slightly flattened dorsally; appendix of forewing
MAGLOW LO!DTOAG (FIGS 22, 28) 0. ...h0.s...:<--0ecratessnsersns D,

— Antennal ledge strong; foretibia strongly flattened
dorsally with margins sharp, sometimes expanded;
appendix of forewing broad (Figs 25—26)
eee Seat nac ica asscudearaatssccesudcedeenseseeceluanesed Drabescini

2 Antennae short (distinctly shorter than half body
length), located near middle to lower corner of eyes
(VETS S66) Ee eee eee cree e er = Selenocephalini

— Antennae long (nearly half body length or more),
located near middle to upper corner of eyes (Figs 7, 8,
NED MMR ee ters cecdesstbivcsseoutccscdvedsccesasadeeeses Paraboloponini

Tribe Selenocephalini Fieber

Diagnosis: Antennae short (distinctly shorter than
half body length), located near middle to lower
corner of eyes; clypeus strongly broadening
upward; ocelli marginal.

REMARKS. This mainly African tribe, re-described
by Linnavuori & Al-Ne’amy (1983), lacks any
unique features; its monophyly is therefore
uncertain. The three genera tentatively included
here aptly demonstrate the heterogeneous nature of
the tribe, i.e. Tambocerus has Y-shaped tentorial
branches (Fig. 17) (as in Deltocephalinae),
Moluccasia has reflexed forewing veins (Fig. 23),

flattened face and upturned vertex (Fig. 14) and
Hecaloidella (new placement from
Deltocephalinae) is without macrosetae on the
subgenital plate (Fig. 100) (present in other genera
of the tribe) and has the aedeagus disassociated
from the connective (also found in Adamini and
some Paraboloponini).

In addition to the african species removed from
Selenocephalus (Linnavuori & Al-Ne’amy, 1983)
the following species, which appear under this
genus in Metcalf’s catalogue from Asia and the
Pacific have subsequently been, or are here,
transferred to other genera. Consequently,
Selenocephalus is now confined to the south-west
Palaearctic region.

brunneus Singh-Pruthi (see Carvaka)

disparatus Melichar (see Tambocerus)

dolens Melichar (see Carvaka)

egregius Stal (see Roxasella)

guttatus Walker (Ledra) Glossocratus
(Deltocephalinae: Hecalini).

marmoreus Walker (see Moluccasia)

notulus Walker (see Moluccasia)

Fieber

taiwanus Matsumura (Selenocephalus) (to
Goniagnathus comb. n.) (Deltocephalinae:
Goniagnathini).

virescens Distant (Selenocephalus) Nephotettix
Matsumura (Deltocephalinae: Doraturini).

viridipes (Distant) (Selenocephalus) Selenomorphus
Evans (lassinae)

GENERA REMOVED FROM THE TRIBE
Divus (see Paraboloponini)

Mysolis (see Paraboloponini)

Omanella (see Paraboloponini)

Roxasella (see Paraboloponini)

Key to genera of Asian and Pacific
Selenocephalini

1 Ocelli nearly touching corresponding eye (Fig. 10);
clypeus slightly concave dorsally in profile (Fig. 10);
frontoclypeal suture indistinct (Fig. 6). Subgenital
plate without macrosetae (Fig. 100); connective with
apex well extended posteriorly (Fig. 104); aedeagus
situated dorsad of connective and linked to it by
membrane (Fig. 104) ) ooeicec.5 cceve-s ceonees Hecaloidella

— Ocelli approximately own diameter from
corresponding eye (Fig. 1); clypeus flat (Fig. 14) or
evenly curved dorsally in profile; frontoclypeal suture
distinct (Fig. 5). Subgenital plate with macrosetae;
connective without apical extension, attached to
ACHEADUS|aflapeX Geekccs..-c2tecees. eeaestscatsv sree ---osedease 2

2 Anterior margin of head upturned, clypeus flat in
profile (Fig. 14); forewing with recurved veins in
corium (Fig. 23); aedeagus with basal processes (Fig.
SA) ies Reet see Mibeat te Some stent cae aes easton acer Moluccasia

8

— Anterior margin of head not upturned; clypeus
slightly swollen; forewing without recurved veins in
corium (Fig. 22); aedeagus without process (Fig.

OS) eee eee enc FOP, cc. . SEMEN Orie eon, Tambocerus
HECALOIDELLA Osborn
Hecaloidella Osborn, 1934a: 173. Type species:
Hecaloidella nitida Osborn, by original
designation.
This genus is here’ transferred from
Deltocephalinae: Scaphytopiini to

Selenocephalinae based on the transversely striated
fore margin of the head, shape of the anterior
tentorium and the strigated second valvulae. Its
inclusion in the Selenocephalini is tentative (see
remarks under classification above). Several new
species from throughout the Pacific (BMNH) differ
from nitida in having the antennae slightly lower
rather than slightly higher than mid-height of the
eyes in facial view. Characteristics of the genus are
given in the above key.

DISTRIBUTION. Samoa (nitida) and throughout
Pacific (new undescribed species).

SPECIES INCLUDED.

nitida Osborn, 1934a: 174, figs 8a-d. Holotype 2,
SAMOA (BMNH) examined. Figs 2, 6, 10, 24,
99-104, 474.

MOLUCCASIA gen. n.

Type species: Selenocephalus marmoreus Walker.

Yellow with brown irroration on vertex and
pronotum; forewings with veins brown, costal veins
edged with dark brown, subcostal region with
alternating brown and yellow patches.

Head almost as wide as pronotum; anterior
margin acute in profile, transversely striated; ocelli
on margin, approximately equal to own diameter
from eyes. Anterior tentorial branches unknown.
Vertex roundly produced medially, midlength
about 1.5 times as long as length next to eye,
upturned apically. Face more or less straight in
profile; transclypeal suture present; laterofrontal
suture extended to lateral margin of corresponding
ocellus; antennae located at level with middle of
eyes in facial view, distinctly shorter than half body
length. Pronotum with sides short, carinated.
Forewing with apical cells short and broad; with
additional recurved veins in subcostal region;
appendix very narrow. Foretibia with, setal formula
1+4; hind femur with apical setal formula 2+2+1.

Y. ZHANG AND M.D. WEBB

Male genitalia: Pygophore lobes with tooth at
posterior margin and _ several macrosetae.
Subgenital plate elongate, triangular shaped, witha
row of stout macrosetae along basal outer margin
and some fine setae on outer margin. Style with
lateral basal apohpysis produced; apical precess
short and stout, tapered to apex, strongly curved,
subapical lobe long with several fine setae.
Connective Y-shaped, arms short, widely
separated. Aedeagus with shaft robust, curved
dorsally and tapered to broadly rounded apex, the
latter laterally compressed anteriorly and with a
flange on each side posteriorly, pair of long basal
processes from anterior margin; gonopore large,
apical on posterior surface; basal apodeme short.

Female genitalia: First valvulae with strigate
sculpture dorsally. Second valvulae similar to
Bhatia (Fig. 71).

DISTRIBUTION. Indonesia (Morotai I. Sula I. and
Misool I.).

REMARKS. Distinguishing characters of this genus
are given in the above key. The two included species
may be conspecific but both available specimens
(syntypes) of marmorea ‘Sula’ and ‘N. Gut’ are
without head and the single specimen (syntype) of
notula is a female. Another syntype recorded from
‘Morty’ (Morotai I.) has not been seen.

SPECIES INCLUDED.

marmorea (Walker, 1870: 325-326)
(Selenocephalus). LECTOTYPE oh
INDONESIA: (Sula I.) (BMNH), here

designated (examined). Comb. n. Figs 23, 83-88.

notula (Walker, 1870: 326) (Selenocephalus).
LECTOTYPE ¢ INDONESIA: Misool I.
(Wallace) (BMNH), here designated (examined).
Comb. n. Figs 13, 14, 471.

TAMBOCERUS gen. n.

Type species: Selenocephalus disparatus Melichar.

Yellow with or without brown irroration. Head
narrower than or nearly equal in width to
pronotum; anterior margin rounded to face in
profile, finely transversely striated; ocelli on margin,
approximately equal to own diameter from
corresponding eye. Anterior tentorial branches
Y-shaped. Vertex slightly produced medially,
slightly sulcate subapically; smooth with some
indistinct longitudinal striations. Face with
transclypeal suture distinct; laterofrontal suture
extended to lateral margin of corresponding
ocellus; antennae situated slightly below midheight
of eyes in facial view, moderately long, clearly

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 9

shorter than half body length. Pronotum smooth,
sides short and carinated. Fore wing with venation
as in Fig. 22, sometimes with white wax patch in
middle of costal margin. Fore tibia with dorsal
surface rounded, setal formula 5+5(6) or 1+5; hind
femur with apical setal formula 2+2+1.

Male genitalia: Pygophore lobes with distal small
sclerotized teeth. Subgenital plate elongate
triangular shaped, with a row of macrosetae,
slightly irregular in basal part of plate. Style with
apical process short, tapered to apex. Connective
with stem elongate, arms short. Aedeagus with
shaft elongate, cylindrical, curved dorsally and
tapered to apex, with lateral teeth, gonopore apical
on posterior surface; basal apodeme moderately
long.

Female genitalia: First valvulae with strigate
sculpture dorsally. Second valvulae similar to
Bhatia (Fig. 71).

DISTRIBUTION. Sri Lanka.

REMARKS. The taxonomic position of this genus is
uncertain. It is tentatively included in the
Selenocephalinae based on the transversely striated
foremargin of the head and strigated first valvulae.
Its distinctly Y-shaped tentorium (Fig. 17) are
similar to those found in some Deltocephalinae. It
can be distinguished in the male genitalia by the
distal small sclerotized teeth on the pygophore
lobes and by the lateral teeth of the aedeagal shaft.

Melichar described disparatus from several
specimens from the following localities in Sri
Lanka: Pattipola, Kandy, Negomba and Paradna,
the latter specimen(s) recorded from ‘Museum
Budapest’. In addition, he noted three damaged
examples in the Berlin Museum. We have examined
the following specimens which we consider to be
syntypes: 7 males, 1 female, ‘Pattipola, 2000m’
‘Ceylon/Biro 1902’ (NHMH, MMB (1)); | female

‘Kandy/Ceylon Horn’ (NHMH); 1 female,
‘Negomba/Ceylon Horn’ and 1 __ female,
‘Paradna/Ceylon Horn’ (IPE); 3 _ females

“Ceylon/Nicter’ (ZMB). This type series represents
five different species, four of the present genus and
one of Carvaka Distant. We have selected one of
the above males from Pattipola (NHMH) as the
lectotype. This specimen bears a type label which
may be contemporary with the date of publication
of disparatus and is probably the specimen
borrowed and figured by Distant (1908: 292). Two
paralectotypes with the same data (male and
female) (NHMH) also belong to this species.

The specimens examined and figured by Distant
(1908) as Phlepsius divaricatus Melichar (BMNH)
belong to disparatus and new species of this genus.
The specimens examined and figured of Carvaka

thoracica Distant by Pruthi (1934: 81) almost
certainly belong to this genus. The specimen
examined and figured by Datta (1973: 442 and
1988: 224) as disparatus is a new species of this
genus.

SPECIES INCLUDED.

disparatus (Melichar, 1903:169) (Selenocephalus),
LECTOTYPE 6, SRI LANKA Biro’, Pattipola,
2000m., 1902 (NHMH) here designated,
examined. Comb. n. Figs 1, 5, 17, 89-93, 472.

plumbeus (Distant, 1908:347) (Phlepsius),
HOLOTYPE 6,SRI LANKA (BHNM) here
designated, examined. Comb. n. Figs 22, 94-98,
473.

Tribe Paraboloponini Ishihara

Paraboloponidae Ishihara, 1953: 5

Paraboloponini Linnavuori, 1960: 299

Paraboloponinae Eyles & Linnavuori, 1974: 39;
Linnavuori, 1978: 457; Webb, 1981: 41.

Bhatiini Linnavuori & Al-Ne’amy, 1983: 21-22.
Syn. n.

Diagnosis: Antennae long (nearly half body length
or more), usually located towards upper corner of
eye; antennal pit encroaching onto clypeus
(exception: Nirvanguina); tentorial branches
T-shaped or falcate.

REMARKS. This group has been described by
Linnavuori (1978), Webb (1981) and Linnavuori &
Al-Ne’amy (1983), the latter as Bhatiini. The above
diagnosis is far from satisfactory because of some
exceptions and because the African genus
Acastroma Linnavuori with these characters and
the fore-margin of the head carinate is placed in the
Deltocephalinae, Athysanini. This genus was
excluded from the Selenocephalinae by Linnavuori
& Al-Ne’amy (1983: 17) because of its non-strigate
sculpture pattern of the first valvulae but as noted
above the sculpture in Selenocephalinae is more
variable than these authors thought. The more
ventral position of the antennae (and ocelli) in
Tsaca (Fig. 9), Nakula and Bhatia (metcalfi),
compared to other genera, is here considered a
secondary development resulting, at least in
Nakula, from a shortening and flexing of the head
downwards.

The earlier treatment of the Paraboloponini as
Paraboloponinae (Webb, 1981) followed
Linnavuori (1978), who included genera with
‘strongly produced triangular-shaped heads’
combined with long, highly positioned antennae.
This head shape (Figs 4, 8) may have influenced
Linnavuori & Al-Ne’amy (1983) to omit the group

10

from the Selenocephalinae whilst erecting the new
tribe Bhatiini (here considered a junior synynonym
of Paraboloponin1), for genera with shorter heads
(Fig. 3). Evidence for this comes from their
comment (1983: 21) that the ‘nymph’ figured by
Evans (1947: 218) of Kutara productus (Evans) was
mis-associated as it had a much longer head than
the adult, and that it probably belonged to a genus
of Paraboloponinae. However, this ‘nymph’ (Figs
500-501) proves to be one of two exuviae, each
associated with an adult K. productus (Fig. 499) and
therefore quite possibly of this genus, which they
placed in Bhatini. Our examination of
Paraboloponini from Asia and the Pacific indicates
that head length alone is an unreliable indicator of
relationship in this group. In addition, we cannot
agree with the exclusion of the Paraboloponini
from the Selenocephalinae based on certain leg
characters (Hamilton, 1983), beacause we can find
no significant differences in the legs between this
and other selenocephaline tribes (see ‘Review of
Characters’).

GENERA REMOVED FROM THE TRIBE.

Sohipona Ghauri & Viraktamath and Shivapona
Ghauri & Viraktamath are excluded from the
Paraboloponini and tentatively placed in the
Grypotini (Deltocephalinae). Although differing
from Grypota in having long antennae and
transverse striations on the fore margin of the head
(as in Paraboloponini) they are similar to this genus
in the shape of the clypellus and in having
non-strigate first valvulae and platellae on the hind
first tarsi.

Key to genera of Asian and Pacific
Paraboloponini

Note: Due to a paucity of external and female
characters it is not possible to distinguish the
females of several genera (couplets 13-21). Also for
this reason the genus Mysolis, known only from the
female, is omitted from the key.

1 Hind femur with 2+2+1 apical setae (Fig. 60) ....... 2
— Hind femur with 2+1+1 or 2+1 apical setae ........ 22
2 Head, thorax and margins of forewing marked with

orange (Plate i icsi7/— 101) esetese seen eee oe ene 3
—sINOtmMankedtaSia DOVE mreeeces eee eeeee eee eee 5

Ww

Vertex very short (Plate 1, Fig. 7); antennae below
midheight of eye in facial view
Nakula (pil Twises eave o.s ees <x fe

Vertex and antennae not as aDOVE ou... ...eceeeeeeeeeeee 4

|

4 Inner two apical cells of forewing small (Fig. 30);
subgenital plate with macrosetae; connective extended

Y. ZHANG AND M.D. WEBB

posteriorly; aedeagus situated dorsad of connective,
connected to midlength of latter by
Membraneseessae. wee. eee Tenompoella (p. 21)

— Inner two apical cells of forewing large; male genitalia
not as above
duels sds See ee Roxasella (in part) (p. 19)

Nn

Laterofrontal sutures on face absent (Fig. 8);
connective extended posteriorly with aedeagus fused
or attached by membrane to its midlength ............ 6

— Laterofrontal sutures on face present (Fig. 7);
connective and aedeagus not as above ..............00+ 7

6 Head and thorax yellow with brown spots (Figs
493-494): head of near uniform length, foremargin
angularly rounded with three to five carinae; vertex
longitudinally striated; aedeagus fused to midlength
of connective; ovipositor extended to apex of
forewings or beyond; valvulae very long and narrow,
second valulae with fine teeth at apex (Fig. 64, 76)

Seas cass us oas eeacectese esas ret eee Omanella (p. 18)

— Head and thorax without brown spots; head produced
medially (Fig. 4); foremargin with two carinae,
foliaceous (Fig. 12); vertex shagreen; aedeagus
connected to midlength of connective by membrane;
ovipositor not as above, valvulae moderately long and
narrow, second valvulae with very fine teeth, extended
over distal one-quarter of dorsal margin (Fig. 75)

aide cbsvcasvPnatobont devote vst sabe ceas temas Parabolopona (p. 18)

—

Laterofrontal sutures of face extended one-half
distance to corresponding ocellus; male genitalia with
valve long, subgenital plate short, connective
racket-shaped; second valvulae tapered from
midlength to apex without teeth (Fig.
Wks) Weretccteerccte e C aac Sombakidia (p. 20)

— Laterofrontal sutures of face extended to
corresponding ocellus; male and female genitalia not
ASTADOVE! Kote biecusctasanchateot ches tareere Ae iacas 7 eee ER 8

oo

Foretibia with 2+4, 4+4 or more dorsal spines; vertex
usually elevated posteriorly; forewing with a crossvein
usually present between claval veins (Fig. 29);
aedeagal shaft produced apically beyond anterior
gonopore; connective angled at midlength ............ 9

— Foretibia with 1+4 dorsal spines; vertex not elevated
posteriorly; forewings without crossvein between

Clava Veins wyrr.cery sacc' s. teu. ce coves. bzedentesats sears 10
9 Stem of connective greatly expanded laterally (Fig.
DN) AER Ae OG aes SOE Drabescoides (p. 13)
— Stem of connective not greatly expanded
laterally Sa. cessor essen ceccert reece Kutara (p. 15)

10 Distinctly marked with dark brown and yellow (Plate
1, Fig. 14); outer subapical cell of forewing extended
to or beyond apex of medial subapical cell (arrowed in
82°46) 4 Sa stuce saeses srsevce- dove seers Seve <cncpenes Isaca (p. 14)

— Markings and outer subapical cell not as above ... 11

11 Outer subapical cell of forewing extended to costal

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 11

WV AE RAIN (SIGS D7) esses Poel. Madea besteet ed Lamia (p. 16)

— Outer subapical cell of forewing not extended to costal
Wal MERI R ees cote dct acae Sa cavnaceet necsiagusedacticcsasaceccwar 12

12 Forewing with inner apical cell closed, fourth apical
cell small, appendix broad (Fig. 28). . Nirvanguina (p.
17)

EMHOLEWINP MOL AS'ADOVE! .f...cc.0hs..cecdsesoencerescdscteerere 13
13 Style with lateral apophysis extended (Fig.
2:15) cca ae Rear Tengatka (p. 20)

— Style with lateral apophysis short .............0.0 14

14 Aedeagus with a process arising basally from posterior
surface, sometimes process arising from between
aedeagus and connective (Fig. 167) or from connective
(Piya, 1 SIR) Ie Ee Se 15

=MANCUCALUS MO’ AS ADOVE ax<.2ccst-ns-sssescsseceonescoesnsersase 17

15 Aedeagus with pair of laterobasal processes ... Divus
(p. 13)

— Aedeagus without a pair of laterobasal processes . 16

16 Aedeagus with apical processes and basal apodeme
(PIS Sh) a Aeceeeece cree ees Carvaka (in part) (p. 13)

— Aedeagus without apical processes or basal apodeme
UPN) a eaca csdhe ss to-s.dceretetensacstosases Hybrasil (p. 14)

17 Aedeagus with a pair of laterobasal processes ..... 18
— Aedeagus without a pair of laterobasal processes . 19

18 Pygophore lobe with ledge along posterior margin
(Fig. 145), subgenital plate with inner apical half
constricted (Fig. 147)

SPREE Coe ars, cclencdedsssadanatess Athysanopsis (p. 11)

— Pygophore lobe with ledge, if present, extended
diagonally across lobe to its ventroposterior corner
(Fig. 105); subgenital plate with inner side nearly
Ninety Lieut areeen <P ck bess bee ae cee Bhatia (p. 11)

19 Stem of connective bifurcate (Fig. 289); aedeagal shaft
filamentous (Fig. 286)
codec co tgp htec phe SRP AEA RoR EEE A EPCRA Welmaya (p. 22)

— Connective and aedeagus not as above ................ 20

20 Subgenital plate with a row’ of _ stout
MITAGHOSCLAG see. Lavtasacsevedesuhetdtieeesis Megabyzus (p. 17)

— Subgenital plate without stout macrosetae .......... 21

21 Connective short (Fig. 235)
sce cac She st ACE CRELERECEE Ot RE Roxasella (in part) (p. 19)

— Connective moderately long
@eravaica: (tasant)! (Ds 18)? v.cssceeses-2e¥-tesctcexnedescoseonecnns

22 Hind femur knee spines 2+1

Sind femur knee spines 2am Geestseeee. covers: 23

23 Clypellus with sides parallel .......... Karoseefa (p. 15)

— Clypellus expanded apically 0.0... eeeeeeeeeee 24

24 Foremargin of head broadly rounded in dorsal and
lateral view; vertex yellow with a pair of brown spots
posteriorly (Fig. 491)
cocbire cor anne ne A RRO PARCELS EDEL REESE LOCUS OAR Waigara (p. 21)

— Foremargin of head acutely rounded in dorsal and
lateral view; vertex without posterior spots ......... 25

25 Aedeagus with a pair of basal processes; female
pregenital sternite with midlength approximately

pwiceileneth of Sides: <22-:.-..:cccn---+-s- Favintiga (p. 14)
— Aedeagus and female pregenital sternite not as
FEL YON hare aie abet ai pee Ane e Reenep eee oe 26
DGSIVMAILG ce ccst eesccees ov ses cae vaugtacs enseccee ieee eee 27
SUP CIAL esr res Met eae sckcacectsocentescnastecceatsecestnssteecas® 29
27 Aedeagal processes symmetrical ...........:::cceeeeeeeeee 28

— Aedeagal processes asymmetrical
Yh ARTI oes ctan tec tea Lae Parohinka (p. 19)

28 Lateral margin of face visible dorsally; pygophore
without an internal ledge
IRHULEIOV RUS (De lO)... cen teuutosaanatgisesgaccesesads sitions sncaas oss

— Lateral margin of face not visible dorsally; pygophore
with an internal ledge
PR ee ee ee ee Dryadomorpha (p. 13)

29 Vertex usually with pale patches; posterior margin of
pregenital sternite V-shaped or U-shaped with
posterolateral corners produced ...........:ccceeeeeee 30

— Vertex without pale patches; posterior margin of
pregenital sternite with a small protuberance each side
OINTMENT OP ox.ceeeoeeseeecescrress esac Dryadomorpha (p. 13)

30 Lateral margin of face visible dorsally
GR ee ct on RES OAS Rhutelorbus (p. 19)

— Lateral margin of face not visible
dorsaliysoss, £25,. SER... Le Parohinka (p. 19)

ATHYSANOPSIS Matsumura

Athysanopsis Matsumura, 1914: 184; Ishihara,
1954: 244-245. Type species: Athysanopsis salicis
Matsumura, by original designation.

This genus, containing two species, is similar to
Bhatia but differs in marking and in the male
genitalia as indicated in the above key. The dorsal
surface of the foretibia is slightly flattened in the
single species examined as in some species of
Bhatia.

DISTRIBUTION. Japan, China.
SPECIES INCLUDED.

salicis Matsumura, 1905: [64], plate 21, fig. 19.
LECTOTYPE <6, JAPAN: Takasago, ix,

12

remounted and labelled, here designated, from a
series of 12 syntypes (examined) (EIHU). Plate
1, Fig. 6; Figs 145-150.

katoi Metcalf, 1966: 96 (/diocerus) nom. n. for
Idiocerus quadripunctatus Kato, 1933a: 8, fig. 6;
Kato 1933b: plate 19, fig.S (Athysanopsis);
Metcalf 1967: 421 (Athysanopsis quadripunctata
Kato 1933b, n.sp. error). Holotype 2, CHINA
(Kato coll.) (not examined).

SPECIES REMOVED FROM THE GENUS.
fasciata Kato (see Drabescus ineffectus).

BHATIA Distant

Bhatia, Distant, 1908: 357. Type species: Eutettix?
olivacea Melichar, by original designation.
Melichariella Matsumura, 1914: 236—237; Ishihara,

1954: 243-245: Linnavuori, 1960: 36
(synonymized with Bhatia); 1983: 23 (stat. rev.).
Type species: Melichariella _ satsumensis

Matsumura, by original designation. Syn. rev.

Koreanopsis Kwon & Lee, 1979: 50. Type species:
Koreanopsis koreana Kwon & Lee, by monotypy.
Syn.n.

This genus, containing eight species, was originally
described from a misidentified type species (Webb,
1994); this is described below as a new species. In
addition, Linnavuori & Al-Ne’amy’s (1983) figured
specimen and Ishihara’s (1961) figure of the
supposed type species is in fact Omanella johnsoni.

Bhatia is similar to Athysanopsis (see under that
genus). Several species-groups are apparent but
because the composition of these groups is
inconsistent i.e. the same species occurs in different
groups, and until the large number of new species
seen are described, we hesitate to describe new
genera for these groups. As the type species of
Melichariella and Koreanopsis fall within these
groups we are synonymizing these genera with
Bhatia. The following differences between
species-groups apply: foremargin of the head
transversely wrinkled rather than striated, vertex
shorter medially than next to eye and antennae
slightly lower than midheight of eye (metcalfi);
foretibia slightly flattened or scored dorsally rather
than rounded (metcalfi, guamensis, lituriceps and
Javana); style apical process expanded (javana) (Fig.
135); style with an apical extension (arrowed in Fig.
109) and pygophore with an internal posterior
ledge (Fig. 105) (olivacea, satsumensis and distanti);
an additional sclerite between the connective and
aedeagus (Fig. 114) (satsumensis, distanti, metcalfi
and guamensis), spiralled internal structure of the
aedeagal processes (Fig. 114) (distanti and
koreana).

Y. ZHANG AND M.D. WEBB

It is possible that Jamitettix is a synonym of
Bhatia (Linnavuori, 1960a: 324). Both metcalfi and
guamensis were originally described in Jamitettix
and in the original description of guamensis it was
compared with the type species of Jamitettix
(Kotonis). But as the holotype of kotonis is lost the
identity of this genus and species remains
uncertain.

There is some confusion regarding the holotype
of metcalfi. A specimen labelled ‘type’ from
NMNH (the originally stated depository) has data
consistent with the originally published data of a
paratype. In this case it may not be possible to
identify the holotype as three other paratypes
(topotypic) are listed. One of these with matching
holotype data has been examined and figured
(BPBM).

DISTRIBUTION. Widespread in southern Asia and
the Pacific.

SPECIES INCLUDED.

distanti sp. n. for Bhatia olivacea (Melichar)
Distant, 1908: 357, fig. 227, misidentification.

guamensis (Metcalf, 1946: 119-120, figs 10a-e)
(Jamitettix); Linnavuori, 1960a: 325-326, figs
46a—b. Holotype d (originally stated as 92),
GUAM (BPBM), (examined). Comb. n. Figs
121-127, 489.

javana (Melichar, 1914: 132) (Kutara)
LECTOTYPE 6d, JAVA: Batavia, xii.1908 (E.
Jacobson) (MMB) here designated (examined).
Comb. n. Figs 134-138, 485.

koreana Kwon & Lee, 1979: 50, figs 1-5. Holotype
6, KOREA (KNU) (not examined). Figs
139-144, 490.

lituriceps (Walker, 1869: 323) (lassus).
LECTOTYPE 2, SULA IS. (BMNH) here
designated (examined). Comb. n. Fig. 487.

metcalfi (Linnavuori, 1960a: 326, figs 46c-e)
(Jamitettix). Holotype 36, CAROLINE IS.
(NMNH ?) (not examined, see below) Comb. n.
Figs 117-120, 486.

olivacea (Melichar, 1903: 191-192, Tab. 6, fig 1, a-c)
(Eutettix?); Distant, 1908: 357 (Bhatia).
Holotype 2, SRI LANKA (MMB) (examined).
Figs 3, 7, 37, 62, 63, 71, 105-111, 483.

satsumensis (Matsumura, 1914: 237-238, fig. 12)
(Melichariella). LECTOTYPE 6, JAPAN
(EIHU) here designated (examined). Comb. rev.
Figs 128-133, 488.

SPECIES REMOVED FROM THE GENUS.
boninensis Matsumura (see Waigara)
formosana Matsumura (see Carvaka)

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 13

Bhatia distanti sp. n.
(Figs 112-116, 484)

Bhatia olivacea (Melichar) Distant, 1908: 357, fig.
227, misidentification.

Length: 3, 6.0mm. Green to brownish yellow; a
small brown spot posteriorly on vertex between
midline and eye and a few similar spots anteriorly
on pronotum; forewings with a brown spot
posteriorly on claval veins at wing margin and a
similar spot at posterior angle of clavus.

Male genitalia similar to olivacea but apical
process of style broader, an additional sclerite
present between aedeagus and connective and the
distinctly different aedeagus (compare Figs 111 and
114).

DISTRIBUTION. Sri Lanka.

REMARKS. In addition to the characters noted
above distanti differs from olivacea in its slightly
narrower vertex (compare Figs 483 and 484).

MATERIAL EXAMINED. Holotype d, Sri Lanka:
Peradeniya, iv.1905 (BMNH).

Paratypes. Sri Lanka: | (abdomen missing)
Peradeniya, xi.1901; 1 (abdomen missing),
Kalawewa, 12—14.iv.1953 (BMNH).

CARVAKA Distant

Carvaka Distant, 1918: 40. Type species: Carvaka
picturata Distant, by original designation.

In addition to the five species of this genus from the
Indian subcontinent, currently being revised by Dr
C.A. Viraktamath, we also include formosana from
Taiwan and contempta from Australia.

The genus is very difficult to diagnose because
the male genitalia are variable and there are only
slight differences distinguishing it from Divus and
Hybrasil (see above key). Its retention as a separate
genus has required the inclusion of contempta
(transferred from Hybrasil) although the overall
appearance of this species and its Australian
distribution, compared to the Oriental distribution
of the remaining species of Carvaka, might suggest
otherwise. Also, the dark brown head of contempta
with a yellow transverse band anteriorly resembles
Hybrasil, as does the connection between the
aedeagal shaft and basal process (arrowed in Figs
158, 163, 167). This process is either articulated
with the connective (Carvaka including contempta),
articulated with the aedeagal shaft (Carvaka, in
part and Hybrasil) or absent (Carvaka, in part).

In addition to the female type of contemptus we

have examined a few additional specimens of this
species (of both sexes) from Australia (BMNH).

DISTRIBUTION. India, Sri
Australia.

Lanka, Taiwan,

SPECIES INCLUDED.

brunneus Singh-Pruthi, 1930: 37, pl. 3, figs 5, Sa
(Selenocephalus). Holotype 2 , INDIA (IM) (not
examined).
contempta (Kirkaldy, 1906: 332) (Exitianus); Ross,
1968: 22 (Hybrasil). Syntypes 2, AUSTRALIA
(BPBM) (examined). Comb. n. Figs 33, 157-160,
Si is
eurytus (Kirkaldy, 1907: 54) (Exitianus). Syntype
2, AUSTRALIA (BPBM) (examined).
Synonymised by Ross, 1968: 22.
fulvida (Evans, 1966: 244-245, fig. 32i) (Car-
vaka). Holotype 2, AUSTRALIA (AM)
(examined). Syn. n.
dolens (Melichar, 1903: 170) (Selenocephalus).
LECTOTYPE 2, SRI LANKA (Nictner); here
designated (examined) (ZMB). Comb. n.
formosana (Matsumura, 1914: 238-239)
(Melichariella). LECTOTYPE 6, TAIWAN:
30.iv.07 (EIHU) here designated (examined).
Comb.n. Figs 32, 151-156, 510.
modesta Distant, 1918: 41. Syntypes 4d 49,
INDIA (BMNH) (examined). Figs 31, 509.
ochrophara Distant, 1918: 41-42. Syntypes 2°,
INDIA (BMNH) (examined). Fig. 506.
picturata Distant, 1918: 40-41, fig. 21. Syntypes 2°
1 sex unknown, INDIA (BMNH) (examined).
Fig. 507.
thoracica Distant, 1918: 41. Syntypes 1d, 12,
INDIA (BMNH) (examined). Fig. 508.

DIVUS Distant

Divus Distant, 1908: 365. Type species: Divus
bipunctatus Melichar, by original designation.

This genus, currently known froma single species, is
being revised by Dr C. Viraktamath. It can be
distinguished from the similar Carvaka by the
paired laterobasal aedeagal processes.

DISTRIBUTION. India, Sri Lanka.
SPECIES INCLUDED.
bipunctatus (Melichar, 1903: 192,) (Eutettix).

Holotype 6, SRI LANKA (not examined). Fig.
482.

14

DRABESCOIDES Kwon & Lee stat. rev.

Drabescoides, Kwon & Lee, 1979: 53. Type species:
Selenocephalus nuchalis Jacobi, by original
designation.

Drabescus __ (Drabescoides):
Emeljanaoy, 1988: 174.

Anufriey &

This monobasic genus, which is identical to Kutara
externally, can be recognized by its very distinctive
male genitalia (see Figs 208-217).

DISTRIBUTION. Korea, China, Russia, Japan?.
SPECIES INCLUDED.

nuchalis (Jacobi, 1943: 30, fig. 9) (Selenocephalus);
Kwon & Lee, 1979: 53, figs 3-6. Lectotype 9,
CHINA (SMT), designated ? by Anufriev, 1979:
166 (examined). Figs 208-217, 504.
Kutara brunnescens Distant, Vilbaste, 1968: 118,
figs 96a, misidentification.
Drabescus striatus Anufriev, 1971: 61. Holotype
3, RUSSIA (Z-I) (not examined). Synony-
mized by Anufriev, 1979: 166.

DRYADOMORPHA Kirkaldy

Dryadomorpha Kirkaldy, 1906: 335; Linnavuori,
1960: 42; Webb 1981: 49-50. Type species:
Dryadomorpha pallida Kirkaldy, by monotypy.

(See check-list for list of synonyms and Webb, 1981,
for full citation).

Five of the eight species of Dryadomorpha occur in
the study area. The genus, revised by Webb (1981),
can be distinguished by its usually acutely produced
head (see Plate 1, Fig. 18), internal male pygophore
ledge and aedeagus with a pair of apical dorsally
directed processes. Unlike most genera of the
subfamily it has reticulate rather than strigate
sculpture on the first valvulae (see “Review of
characters’).

DISTRIBUTION. Ethiopian, southern Palaearctic
and Austro-Oriental region.

SPECIES INCLUDED.

anacryon Linnavuori, 1978b: 462, figs 4a—b.
Holotype ¢, ZAIRE (MRAC) (not examined).

bruneocephala Webb, 1981: 57-58, figs 1-5.
Holotype 36, CENTRAL AFRICAN
REPUBLIC (MNHN).

metrosideri (Osborn, 1934b: 247-248) (Calotettix);
Webb, 1981: 53-54, figs 57-60. Holotype 2°,
MARQUESAS IS (BPBM) (examined).
metrosideri vat. tincta Osborn (Calotettix).
lais Eyles & Linnavuori (Calotettix).

Y. ZHANG AND M.D. WEBB

(See Webb, 1981 for full citation of synonyms).
pacifica Webb, 1981: 55, figs 64-67. Holotype 6,
IRIAN JAYA (BPBM) (examined).
pallida Kirkaldy, 1906: 336; Webb 1981: 50-53, figs
41-56. Holotype 2, AUSTRALIA (BPBM)
(examined). Plate 1, Fig. 18; Fig. 40.
virescens Distant (Paganalia).
indicus Distant (Zizyphoides).
virens Haupt (Rhombopsis).
viridis Singh-Pruthi (Rhombopsis).
chatterjeei Singh-Pruthi (Rhombopsis).
antennalis Lindberg (Platymetopius).
yakushimensis Ishihara ( Yakunopona).
australis Evans (Platymetopius).
(See Webb, 1981 for full citation of synonyms).
quadricornis (Linnavuori, 1969: 1181, figs 40c-e)
(Zizyphoides); Linnavuori, 1978b: 462, figs 3c-e.
Holotype 6, ZAIRE (NMHN) (examined).
robustipenis Webb, 1981: 54-55, figs 61-63.
Holotype 6, SARAWAK (BMNH) (examined).
viridia Osborn, 1934b: 244-245, figs 3a—c; Webb,
1981: 54. Holotype 2, MARQUESAS IS.
(BPBM) (examined).

FAVINTIGA Webb

Favintiga Webb, 1981: 47. Type — species:
Parabolopona camphorae Matsumura, 1912, by
original designation.

This monobasic genus, revised by Webb (1981), is
distinguished by the ventral process of the
connective and narrow second valvulae (Fig. 77).

DISTRIBUTION. Japan.
SPECIES INCLUDED.

camphorae (Matsumura, Sle 288)
(Parabolopona); Webb, 1981: 48, figs 28-40.
Lectotype 2, JAPAN (EIHU) designated by
Webb, 1981: 48, (examined). Figs, 48, 77, 481.

HYBRASIL Kirkaldy

Hybrasil Kirkaldy, 1907: 41, pl. 2, fig. 18;
Linnavuori, 1960b: 37-38. Type species: Hybrasil
brani Kirkaldy, by monotypy.

The taxonomic position of this monobasic genus,
here transferred from the subfamily
Deltocephalinae, was considered to be uncertain by
Ross (1968). It can be distinguished by the unusual
process between the aedeagus and connective and
the aedeagus without a basal apodeme and with an
apical gonopore on the anterior surface (Fig. 167)
(see also remarks under Carvaka).

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 15

DISTRIBUTION. Fiji.
SPECIES INCLUDED.

brani Kirkaldy, 1907: 41, pl. 2, fig. 18; Linnavuori,
1960b: 38-39, figs 13a-c. LECTOTYPE 92, FIJI:
Levu, Rewa, iii (Muir) (BPBM), here designated
(examined). Plate 1, Fig. 15; Figs 34, 74,
161-167.

SPECIES REMOVED FROM THE GENUS.
contemptus Kirkaldy (see Carvaka).

ISACA Walker

Isaca Walker, 1857: 172. Type species: [saca bipars
by monotypy.

This genus can be distinguished by its striking dark
brown and yellow coloration (Plate 1, Fig. 14),
antennae situated slightly below midheight of the
eyes, in facial view (Fig. 9), and by its distinctive
male genitalia. We have matched the type of bipars
(without abdomen), the only previously known
species, to several specimens from Sarawak, the
type locality (BMNH). To avoid misidentifications
two other species from Sabah and Brunei, are
described below. These are externally identical to
bipars but have slightly different male genitalia.

DISTRIBUTION. Sarawak, Sabah, Brunei.
SPECIES INCLUDED.

bipars Walker, 1857: 172. LECTOTYPE (abdomen
missing), SARAWAK (Wallace) here designated
(examined). Figs 9, 46, 218-224.

falcata sp. n.

sinuata sp. n.

Isaca falcata sp. n.
(Plate 1, Fig. 14; Figs 229-233)

Length: 6, 5.1-5.8mm; 2, 5.6-6.0mm. Female
pregenital sternite with posterior margin slightly
produced each side of mid-line.

Male genitalia with pygophore lacking a short
posteroventral process. Anal tube processes falcate.
Aedeagus with shaft nearly straight in lateral view,
with a short subapical medial process posteriorly.

DISTRIBUTION. Sabah.

MATERIAL EXAMINED. Holotype <6, Sabah:
Sandakan (Baker) (NMNH).
Paratypes. Sabah: 2d , 9°, same data as holotype

(NMNH, BMHN).

REMARKS. This species can be distinguished from
other species of the genus by the male genitalia

characters noted in the above description.

Isaca sinuata sp. n.
(Figs 225-228)

Length: dg, 5.2-5.4mm. Male genitalia with
pygophore with a short posteroventral process.
Anal tube processes slightly sinuate, directed
posteriorly. Aedeagus with shaft slightly sinuate, in
lateral view, with a pair of subapical anterior
processes.

DISTRIBUTION. Brunei.

MATERIAL EXAMINED. Holotype ¢, Brunei: Ulu

Paratypes. Brunei: 23, same data as holotype
(BMNH).

REMARKS. This species can be distinguished from
other species of the genus by the male genitalia
characters noted in the above description.

JAMITETTIX Matsumura nomen dubium

Jamitettix Matsumura, 1940: 40; Linnavuori, 1960:
324. Type species: Jamitettix kotonis
Matsumura, by original designation.

As the type species of this monobasic genus has
been lost we are uncertain of its identity (see
remarks under Bhatia).

DISTRIBUTION. Taiwan
SPECIES INCLUDED.

kotonis Matsumura,
TAIWAN (lost).

1940: 40. Holotype 2,

SPECIES REMOVED FROM THE GENUS.

guamensis Metcalf (see Bhatia).
metcalfi Linnavuori (see Bhatia).

KAROSEEFA Webb

Karoseefa Webb, 1981: 70-71. Type species:
Karoseefa brevipenis Webb, by original
designation.

This genus can be distinguished from all others of
the tribe by the parallel rather than concave sides of
the clypellus.

DISTRIBUTION. Sarawak and Sabah.

16

SPECIES INCLUDED.

brevipenis Webb, 1981: 71-73, figs 149-161.
Holotype, 6 SARAWAK (BMNH) (examined).
Figs 41, 73, 476.

divergens Webb, 1981: 73, figs 162-166. Holotype
3, SARAWAK (BMNH) (examined). Fig. 475.

KUTARA Distant

Kutara Distant, 1908: 308; Linnavuori, 1978: 44.
Type species: Kutara brunnescens Distant, by
original designation.

This genus is identical to Drabescoides externally
and has similar male genitalia, with an anterior
gonopore and connective angled at midlength in
lateral view. It differs in having the stem of the
connective narrow rather than broad. Two strong
transverse carinae on the foremargin of the head
are usually present, with a sulcus between, or a
single carina, with or without finer striations. See
under tribal discussion for remarks on the nymph
(of productus).

Within the seven species of the genus some
species-groups are apparent but because the
composition of these groups is inconsistent 1.e the
same species appears in different groups, and until
the large number of new species seen are described
we hesitate to describe new genera for these groups.
The following differences between species groups
apply: crossvein present between claval veins of
forewing together with 4+4 or more setae on the
foretibia (all species except J/ucidicosta); vertex
elevated posteriorly (all species except /ucidicosta
and sinensis); pygophore with an internal posterior
ledge (brunnescens and producta); vertex with a
central brown spot and or transverse bar (all species
except grisescens); foretibia quadrate in cross
section and aedeagus with a pair of basal processes
(grisescens).

Several specimens (one of which is shown in Fig.
495), representing more than one species, are
similar to the type (without abdomen) of
lucidicosta. We have therefore been unable to figure
the male genitalia of this species which is best
regarded for the present as a species incertae sedis.

The syntype series of brunnescens is represented
by two species. A single syntype (male) (Figs
168-174) matches the genitalia figured by
Linnavuori (1978) and Anufriev (1979) for this
species and is probably the specimen that these
authors examined. Five other syntypes (all female)
match the habitus figure in the original description
of this species, with respect to the more transverse
spot on the vertex (Fig. 502), a variation noted in
the original description. To maintain stability we

Y. ZHANG AND M.D. WEBB

have retained the recently held identity for this
species (see above) and describe a new species below
for the remaining syntypes.

DISTRIBUTION. Throughout Oriental and Pacific
region.

SPECIES INCLUDED.

brunnescens Distant, 1908: 308-309 (in part);
Linnavuori, 1978: 44-45, figs 13f-h (in part);
Anufriev, 1979: 166, figs 1-6. LECTOTYPE 4,
here designated, SRI LANKA: Peradeniya
(BMNH) (examined). Figs 29, 168-174, 498.

costalis (Distant, 1908: 146). (Dabescus) [sic.
Drabescus]. Holotype 2, BORNEO (BMNH)
(examined). Comb. n. Figs 196-200, 496.

grisescens Evans, 1947: 256-257, figs 31b, c, h.
Holotype 2, SARAWAK (BMNH) (examined).
Figs 201-207, 503.

lucidicosta (Walker, 1869: 323-324) (Jlassus).
LECTOTYPE 2, NEW GUINEA (Wallace)
(BMNH) here designated (examined). Comb. n.
Fig. 495.

nigrifasciata Kuoh (in Ge & Kuoh) 1992, 300 (316),
fig. 59 a-h. Holotype ¢, CHINA (AS?) (not
examined).

producta (Evans, 1947: 256, figs 3ld, g, j)
(Drabescus). Holotype 2, MALAYA (BMNH)
(examined). Comb. n. Figs 181-188, 499.

sinensis (Walker, 1851: 871) (Bythoscopus); Metcalf,
1966: 89 (lassus). Holotype (sex unknown),
HONGKONG (BMNH) (examined). Comb. n.
Figs 189-195, 497.

transversa sp. n. for Kutara brunnescens Distant,
1908: 308-309, fig. 197 (misidentification in
part); Linnavuori, 1978: 44-45, fig. 15a
(misidentification in part).

SPECIES REMOVED FROM THE GENUS.

javana Melichar (see Bhatia)
nigrofacialis Distant (see Drabescus).

Kutara transversa sp. n.
(Figs 175-180, 502)

Kutara brunnescens Distant, 1908: 308-309, fig. 197
(misidentification, in part); Linnavuori, 1978:
44-45, fig.15a (misidentification, in part).

Length ¢, 6.0mm; ¢, 7.0—-7.3 mm. Head yellow;
vertex with a medial brown spot extended laterally
as a transverse line. Pronotum brown with
numerous yellow spots. Forewings brownish
hyaline with darker brown patches in cells and at
apex.

Head with a transverse carina around margin
above and below ocellus with a sulcus in between

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 17

and some finer striations. Fore tibia rounded
dorsally with 4+4 spines. Forewing with a crossvein
between claval veins.

Male genitalia with style expanded subapically,
with apical process digitate. Connective long,
angled at midlength. Aedeagus shaft cylindrical,
without processes, gonopore arising from an
extended anterior subapical region of shaft.

Female pregenital sternite acutely produced
posterolaterally.

DISTRIBUTION. Sri Lanka

MATERIAL EXAMINED. Holotype 6, Sri Lanka
(Thwaites) (BMNH).

Paratypes. Sri Lanka: 52% Kandy
(Paralectotypes of Kutara brunnescens Distant);
12, Udawattekelle ((NMNH).

REMARKS. This species can be distinguished from
all other species of the genus by its unusual shaped
style (Fig. 177). Its simple aedeagus is damaged
apically in the only male but is probably similar to
costalis (Fig. 200).

LAMIA Linnavuori

Lamia Linnavuori, 1960b: 39. Type species: Lamia
cydippe by original designation.

This monobasic genus can be distinguished by the
parabolically produced vertex, fore wing obliquely
truncate apically with outer subapical cell
extending to the costal margin, resulting in a
triangular-shaped fifth (outer) apical cell (Fig. 27)
and by the scale-like dorsal sculpture of the first
valvulae. The genus is known only from two female
specimens of cydippe (BPBM, BMNH). The
non-type specimen has slightly more extensive
brown markings on the pronotum (Plate 1, Fig. 19).

DISTRIBUTION. Fiji
SPECIES INCLUDED.

cydippe Linnavuori, 1960b: 39, figs. 12j, 13d.
Holotype 2, FIJI (BPBM) (examined). Plate 1,
Fig. 19; Figs 27, 79, 81.

SPECIES REMOVED FROM THE GENUS.

placida Evans (see Nirvanguina)

MEGABYZUS Distant

Megabyzus Distant, 1908: 294-295. Type species:
Megabyzus signandus Distant, by original
designation.

This monobasic genus, currently being revised by
Dr C. Viraktamath, can be distinguished by the
presence of macrosetae on the subgenital plate and
by the pair of apical aedeagal processes. The first
valvulae have reticulate rather than the more usual
strigate dorsal sculpture.

DISTRIBUTION. Sri Lanka, India.
SPECIES INCLUDED.

signandus Distant, 1908: 295, fig. 187. Holotype °,
SRI LANKA (BMNH) (examined). Figs, 44,
479.

MYSOLIS Kirkaldy

Mysolis Kirkaldy, 1904: 279. replacement name for
Norsia Walker, 1869: 326. Type species: Norsia
flavidorsum Walker, by original designation.

Norsiana Distant, 1908: 148, replacement name for
Norsia Walker, 1869: 326.

The single known specimen of this genus (without
abdomen) is distinctively marked with brown as a
transverse band above and below a yellow band on
the foremargin of the head and on lateral areas of
the pronotum and scutellum (Plate 1, Fig. 16). Its
triangularly produced vertex and colour pattern
resemble Hybrazil but it has the vertex and
pronotum shiny rather than striated and has a
transverse sulcus on the foremargin of the head,
although the latter character is variable within
some other genera. The single antenna present is
situated high on the face and extends approximately
to the apex of the clavus.

A further species (fulvescens Walker) is listed in
Metcalf’s (1966) catalogue under this genus but this
species is a member of the Fulgoroid family
Tropiduchidae and is placed under Kallitaxila
granulata Stal in the BMNH collection.

DISTRIBUTION. Misool I. (West Irian)
SPECIES INCLUDED.

flavidorsum Walker, 1869: 326; Distant, 1908: 148,
pl. 7, figs 12, 12a. LECTOTYPE (sex unknown,
abdomen missing), INDONESIA: Misool I.
(Wallace) (BMNH), here designated (examined).
Plate 1, Fig. 16; Fig. 45.

NAKULA Distant, 1918
Nakula Distant 1918: 39. Type species: Nakula
multicolor Distant, by original designation.

The single known specimen of this genus is
distinctively marked with orange (Plate 1, Fig. 7)

18

and in addition can be distinguished by its short
head (very slightly shorter medially than next to
eyes) and low position of the ocelli and antennae,
the latter situated slightly below midhight of the
eyes, in facial view.

DISTRIBUTION. Myanmar (Burma).
SPECIES INCLUDED.

multicolor Distant, 1918: 39, fig.20. LECTOTYPE
2, MYANMAR (BURMA): Mergui (W
Doherty) (BMNH), here designated (examined).
Plate 1, Fig. 7.

NIRVANGUINA gen. n.

Type species: Lamia placida Evans.

Description: small species (4.0-4.6 mm). Yellow
with orange patches on vertex and pronotum and
brown irroration on forewing.

Head slightly narrower than pronotum,
foremargin foliaceous with two transverse carinae.
Vertex smooth, parabolically produced, slightly
longer medially than basal width between eyes.
Face strongly incurved below eyes; clypellus
expanded apically, transclypeal suture present;
laterofrontal suture extended to corresponding
ocelli; antennae situated near upper corner of eye in
facial view, longer than half body length; antennal
pit not encroaching onto clypeus. Pronotum
smooth, sides moderately long, carinate. Foretibia
with dorsal surface round, setal formula 1+4. Hind
femur with apical setal formula 2+2+1. Forewings
with inner subapical cell closed; fourth (outer)
apical cell small; appendix broad.

Male pygophore lobes narrowly rounded
posteriorly with several dorsal macrosetae and an
elongate dorsal process extended from base of lobe
to near its posterior margin, dentate apically.
Subgenital plate elongate, broad basally with
lateral margin strongly incurved at midlength;
tapered over distal half to acute apex; uni-seriate
row of macrosetae ventrolaterally over basal half;
several long fine setae laterally and several
moderately long fine setae dorsoapically.
Connective, broadly Y-shaped with stem short.
Style with apical process and lateral lobe short, the
former tapered to acute apex. Aedeagus with shaft
short, strongly curved dorsally, compressed
dorsoventrally over basal two-thirds, thereafter
tubular; basal apodeme moderately long,
finger-like in lateral and posterior view.

Female genitalia with first valvulae with dorsal
sculpture alveolate at midlength (as Fig. 69, except
oblique) turning to well strigate apically (Fig. 63).

Y. ZHANG AND M.D. WEBB

Second valvulae with several small dorsal teeth
over very broad distal half (Fig. 80); dorsal
sclerotized and hyaline region present, the former
short. Third valvulae with distal region curved
ventrally (Fig. 82).

REMARKS. This genus is tentatively included in the
Paraboloponini because although the antennae are
high and long the antennal pit does not encroach
onto the clypellus (Fig. 15) as in other members of
this tribe. Other differences are the strongly
parabolically produced vertex, strongly incurved
lateral margins of the face below the eyes, and fore
wing with a closed inner subapical cell and very
broad appendix. Due to the small number of
specimens available (type and two specimens from
Papua New Guinea, CAS, BMNH) the tentorial
branches have not been examined.

DISTRIBUTION. Papua New Guinea and Australia
(N. Quensland).

SPECIES INCLUDED.

placida (Evans, 1966: 221, figs 37, BI—B2) (Lamia).
Holotype 2, AUSTRALIA (AM) (examined).
Plate 1, Fig. 20; Figs 15-16, 28, 80, 82, 308-313.

OCEANOPONA Linnavuori

Oceanopona Linnavuori, 1960: 300; Webb, 1981:
73-75. Type species: Oceanopona croceipennis
Linnavuori, by original designation.

This monobasic genus can be distinguished by the
reduced number of apical setae on the hind femur
(2+1). Its male genitalia are similar to
Dryadomorpha and Karoseefa but the internal
pygophore ledge is shorter and the apophysis of the
arms of the connective are longer.

DISTRIBUTION. Eastern Caroline Islands.
SPECIES INCLUDED.

croceipennis Linnavuori, 1960: 301; Webb, 1981: 75,
figs 167-178. Holotype 6, CAROLINE IS.
(BPBM) (examined). Figs 36, 480.

OMANELLA Merino

Omanella Merino, 1936: 361. Type species:
Omanella_ barberi Merino, by _ original
designation.

This distinctive genus, comprising two species, can
be distinguished by the brown spots on the dorsum,
long ovipositor (Figs, 64, 65, 76), aedeagus fused to
the midlength of the connective (the later extended

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 19

posteriorly), and highly modified style.

Long type series of both species from the
Philippines have been examined (BPBM, BMNH)
together with specimens of johnsoni from
throughout the range of the genus.

DISTRIBUTION. Sri Lanka, Myanmar (Burma),
S.E. Asia, Philippines.

SPECIES INCLUDED.

barberi Merino, 1936: 361-362, pl. 3, figs 5a—-d.
Holotype 6, PHILIPPINES (NMNH)
(examined). Figs 50, 264-270, 493.

johnsoni Merino, 1936: 363, pl. 3, figs 4a—d.
Holotype 36, PHILIPPINES (NMNH)
(examined). Figs 256-263, 494.
Bhatia olivacea Distant, Ishihara, 1961: 243, fig.
48, plate 4: 24, misidentification; Linnavuori &
Al-Ne’amy, 1983: 22, figs 32b-e,
misidentification.

SPECIES REMOVED FROM THE GENUS.

philippina Merino (see Roxasella).

PARABOLOPONA Matsumura

Parabolopona, Matsumura, 1912: 288; Webb, 1981:
42-43. Type species: Parabolocratus guttatus
Uhler, by original designation.

This genus, revised by Webb (1981) and comprising
five species, can be distinguished by the elongate,
foliaceous head with very high antennae and
shagreen vertex (Figs 4, 8, 12), aedeagus connected
by membrane to the midlength of the connective
(the latter extended posteriorly) and very fine
dorsal teeth of the second valvulae (Fig. 75).

DISTRIBUTION. Asia, as far south as Nepal to the
Philippines (Luzon).

SPECIES INCLUDED.

chinensis Webb, 1981: 45—46, figs 20-21. Holotype
36, CHINA (CAS) (examined).

guttata (Uhler, 1896: 291) (Parabolocratus); Webb,
1981: 43, figs 1-14. Lectotype 6, JAPAN
(NMNH) (examined). Figs 4, 8, 12, 43, 75.

ishihari Webb, 1981: 45, figs 15-19. Holotype <¢,
JAPAN (EIHU) (examined).

luzonensis Webb, 1981: 46-47, figs 22-27. Holotype
6, PHILIPPINES (NMNH) (examined). Plate
eis 17.

yangi Zhang, Chen & Shen, 1995: 11, fig. 4.
Holotype ¢, CHINA (BAU) (examined).

PAROHINKA Webb

Parohinka Webb, 1981: 57-58. Type species:
Muirella longiseta Melichar, by original
designation.

The acutely produced head (Fig. 477) and elongate
face of this genus, comprising twelve species, are
similar to Dryadomorpha and Rhutelorbus but it
differs from these and other genera by its
asymmetrical aedeagus.

DISTRIBUTION. Southeast Asia and the Pacific.
SPECIES INCLUDED.

apicalis Webb, 1981: 68, figs 126-128. Holotype 6,
SARAWAK (BPBM) (examined).

brevicephala Webb, 1981: 66, figs 111-118.
Holotype ¢, SARAWAK (BMNH) (examined).

dulita Webb, 1981: 64-65, figs 103-110. Holotype
3, SARAWAK (BMNH) (examined).

longiseta (Melichar, 1914: 135, pl. 3, figs 15, 15a)
(Muirella); Webb 1981: 68-70, figs 129-143.
Lectotype 6, JAVA (MB) (examined). Figs, 39,
477.

lotophagorum (Kirkaldy, 1907: 41)
(Dryadomorpha); Linnavuori, 1960: 42; Webb,
1981: 63-64, figs 92-97. Neotype 6, FIJI
(BPBM) (examined).

malayensis Webb, 1981: 70, figs 144-148. Holotype
3, MALAYA (BMNH) (examined).

morona Webb, 1981: 60-61, figs 79-86. Holotype d,
JAVA (BMNH) (examined).

philippina Webb, 1981: 66, figs 119-123. Holotype
3, PHILIPPINES (BPBM) (examined).

recurya Webb, 1981: 67, figs 124-125. Holotype ¢,
SARAWAK (BPBM) (examined).

sinuata Webb, 1981: 61, fig. 86. Holotype 6,
PAPUA NEW GUINEA (BPBM) (examined).

spinosa Webb, 1981: 64, figs 98-102. Holotype 6,
PAPUA NEW GUINEA (BMNH) (examined).

trispicata Webb, 1981: 61-63, figs 87-91. Holotype
3, SOLOMON IS. (BPBM) (examined).

RHUTELORBUS Webb

Rhutelorbus Webb, 1981: 56-57. Type species:
Rhutelorbus merinoi, by original designation.

This monobasic genus (Fig. 478) is similar to
Dryadomorpha and Parohinka but can be
distinguished from these and other genera by its
dorsally visible lateral margins of the face.

DISTRIBUTION. Philippines, Malaya and Sarawak.

20
SPECIES INCLUDED.

merinoi Webb, 1981: 57, figs 68-78. Holotype 6,
MALAYA (BMNH) (examined). Fig. 478.

ROXASELLA Merino

Roxasella Merino, 1936: 359. Type species:
Roxasella camusi Merino, by original
designation.

This genus, which is poorly defined at present,
contains a variable assemblage of six species. In all
species, except Josbanosa, two carinae are
discernable around the fore margin of the head,
with a variable sulcus between, in addition to finer
striations. The only male (the holotype) of the
genotype, camusi, is missing but the allotype of
camusi examined (NMNH) comes nearest to
losbanosa and biarcuatus in head length and colour
pattern. Of the remaining species, philippina (with
similar male genitalia to biarcuatus and losbanosa)
and /aetisigna both have shorter heads and bright
orange markings, similar to Nacula and
Tenompoella. R. egregius is known from only three
female specimens from Myanmar (Burma), two
misidentified by Distant (1908a, 1918) as
laetisigma. These two species are identical
externally except that in the former the pregenital
sternite is asymmetrical, which would be
considered aberrant if it were not for its presence in
all three specimens.

Attention is drawn to the male genitalia of
laetisigma where the style is reminiscent of some
species of Drabescus and to the extra sclerite
between the connective and the aedeagus which is
also found in some other genera (see “Review of
characters’).

In addition to the holotypes, series of specimens
of philippina and losbanosa from the Philippines
have been examined and a few specimens of
laetisigna from Singapore, Sarawak and Sabah
(NMNH, BMNH). No material of camusi and
biarcuatus other than the allotype and lectotype
respectively, has been seen. Two specimens of the
latter and one of /aetisigna, collected by Wallace
and deposited in F. Walker’s collection, Museum of
Victoria (pers. comm. Dr. A. Neboiss), may be
syntypes of these species but were not examined.

DISTRIBUTION. Myanmar (Burma), Philippines,
Singapore, Sarawak, Borneo, Sabah.

SPECIES INCLUDED.

biarcuata (Walker, 1857: 174) (Tartessus).
LECTOTYPE 3, SARAWAK (BMNH) here

Y. ZHANG AND M.D. WEBB

designated (examined). Comb. n. Plate 1, Fig. 11;
Figs 241-245.

camusi Merino, 1936: 360, pl. 3, fig. 1, a.
Holotype ¢, PHILIPPINES (lost). Plate 1, Fig.
12s

losbanosa Merino, 1936: 360, pl. 3, fig. 2, ad.
Holotype 36, PHILIPPINES (NMNH)
(examined). Plate 1, Fig. 13; Figs 234-240.

egregia (Stal, 1864: 66) (Selenocephalus), Holotype
2, Myanmar (BURMA) (examined). Comb. n.
Fig. 505.
laetisigna (Walker) Distant, 1908: 307 (in part),
1918: 42, fig. 22, misidentification.

laetisigna (Walker, 1857: 174) (Bythoscopus);
Metcalf, 1966: 175 (Drabescus). LECTOTYPE
3, SARAWAK (BMNH) here designated,
(examined). Comb. n. Plate 1, Fig. 9; Figs
250-255.

philippina (Merino, 1936: 362, pl. 3, figs 3a—d)
(Omanella). Holotype 6, PHILIPPINES
(NMNBH) (examined). Comb. n.. Plate 1, Fig. 10;
Figs 246-249.
Philippinensis Baker nom. nud.

SOMBAKIDIA gen. n.

Type species: Eutettix morismus Merino

Description. Small species (5.0-5.4 mm). Yellow
with brown markings as in Figs 52, 513. External
characters as in Waigara but laterofrontal sutures
short; ocelli close to eyes, approximately half own
diameter from eye; vertex smooth with very faint
longitudinal striations; pronotum obscurely
transversely wrinkled and hind femur with apical
setal formula 2+2+1.

Male genitalia: Pygophore lobes narrowly
rounded posteriorly with several macrosetae, an
oblique internal ledge from dorsal margin to ventro
posterior corner. Valve elongate. Subgenital plate
short and broad, triangular shaped with a few
irregularly arranged short fine setae dorsolaterally.
Style elongate, narrow basally, apical process short,
tapered to apex, lateral lobe long. Connective short,
club-shaped with arms fused. Aedeagus with shaft
moderately long with a single elongate ventral
process, gonopore apical; basal apodeme very
short.

Female genitalia: First valvulae with reticulate
sculpture. Second valvulae narrow, tapered from
base to apex with dorsal prominence, sclerotized
and hyaline region and teeth absent.

REMARKS. This genus is similar externally to
Tengatka but can be distinguished by its short

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 21

laterofrontal sutures and unusual male and female
genitalia as noted above.

The type series of Eutettix morismus, comprising
several specimens (male and female), was examined
(NMNH, BMNB).

DISTRIBUTION. Philippines (Mindanao and

Basilan).

SPECIES INCLUDED.

morisma (Merino, 1936: 377, pl. 4, figs Sa—d)
(Eutettix). Holotype 2, PHILIPPINES

(NMNH) (examined). Comb. n. Figs 52, 66-67,
78, 298-307, 513.

TENGATKA gen. n.

Type species: Eutettix basilana Merino

Description: Small species (4.8-5.3mm). Yellow
with brown markings as in Figs 51, 512. External
characters as in Sombakidia but with laterofrontal
sutures extended to ocelli.

Male genitalia: Pygophore lobes narrowly
rounded posteriorly with several macrosetae;
without processes. Valve very short (not or only
slightly visible in dried specimens). Subgenital plate
triangular with finger-like apex, few short fine setae
laterally. Style narrow basally, lateral apophysis
elongate, apical process short, tapered to acute
apex, few sensory pits towards lateral lobe, the
latter short. Connective Y-shaped, short and broad.
Aedeagus robust, laterally compressed, shaft short,
with subapical processes and a single ventral
process from posterior surface, gonopore apical,
basal apodeme short; socle broad in lateral aspect.

Female genitalia: First valvulae with strigate
sculpture dorsally. Second valvulae similar to
Bhatia (Fig. 71).

REMARKS. This genus is similar to Sombakidia
externally but can be distinguished from this and
other genera by its short male valve and distinctive
style as noted above.

The type series of Eutettix basilana, comprising
several specimens (male and female), was examined
(NMNH, BMNB).

DISTRIBUTION. Philippines
Basilan).

(Mindanao and

SPECIES INCLUDED.

basilana (Merino, 1936: 377, pl. 4, figs 6a—d)

(Eutettix). Holotype °, PHILIPPINES
(NMNH) (examined). Comb. n. Figs 51,
291-297, 512.

TENOMPOELLA gen. n.

Type species: Tettigonia difficilis Walker

Description: Moderately large species, 6.2—-6.5mm.
Venter pale yellow, vertex, pronotum and scutellum
green with orange markings, fore wing brownish
hyaline margined with orange and marked with
dark brown apically and as a spot in each cell and at
apex of clavus and claval veins.

External characters as in Waigara but with a fine
carina around fore margin of head above and below
ocellus in addition to finer striations and ocelli
approximately twice own diameter from eyes.
Vertex nearly smooth, with anterior margin very
slightly longer medially than next to eyes. Antennae
situated at mid-height of eye in facial view.
Pronotum nearly smooth, sides moderately long
and carinated. Hind femur with apical setal
formula 2+2+1.

Male genitalia: Pygophore lobes broadly
rounded posteriorly with several macrosetae.
Subgenital plate elongate, triangular shaped, with a
longitudinal row of stout lateral macrosetae at
midlength and a few short fine setae along lateral
and apical margin. Style broad basally, apical
process short curved and tapered to apex with fine
setae and sensary pits basally; lateral lobe
moderately long. Connective Y-shaped with stem
produced posteriorly and bifurcate apically,
connected to aedeagus by membrane at its
midlength; arms short. Aedeagus small and simple,
shaft short, tubular, curved dorsally and tapered to
apex, without processes; gonopore elongate,
situated apically on posterior surface; basal
apodeme elongate, compressed anteroposteriorly
with apical and basal lateral arms.

Female genitalia: First valvulae with strigate
sculpture dorsally. Second valvulae similar to
Bhatia (Fig. 71).

REMARKS. This genus can be distinguished from
all other genera of the tribe by the small inner
apical cell of the fore wing, posteriorly produced
connective and macrosetae on the subgenital plate.
Males and females from Sandakan (Sabah) and
Singapore (BPBM and BMNH) which match the
female type from ‘Sarawak’ have been examined.

DISTRIBUTION. Sarawak, Sabah, Singapore.
SPECIES INCLUDED.

difficilis (Walker, 1857: 169) (Tettigonia); Metcalf
1965:235, (Bothrogonia). LECTOTYPE 92°,
SARAWAK (BMNH) here designated,
(examined). Comb. n. Plate 1, Fig. 8; Figs 30,
271-277.

22
WAIGARA gen. n.

Type species: Melichariella boninensis Matsumura

Description: moderately large species (6.6-7.2mm).
Yellow with a pair of black spots posteriorly on
vertex.

Head wider than pronotum; anterior margin
angularly rounded to face in profile, transversely
striated; ocelli on margin, approximately 3 times
own diameter from corresponding eye. Anterior
tentorial branches T-shaped (similar to Fig. 19).
Vertex obliquely striated, with anterior margin
slightly longer medially than next to eyes, sulcate
subapically. Face with clypellus expanded apically;
transclypeal suture present; laterofrontal sulture
extended to lateral margin of ocellus; antennae
situated slightly above midheight of eye in facial
view, longer than half body length; antennal pit
encroaching onto clypellus. Pronotum transversely
striated, side margins short and carinated.
Foretibia with dorsal surface round, setal formula
1+4. Hind femur with apical setal formula 2+1+1.

Male pygophore lobes acutely rounded
posteriorly with several macrosetae and short fine
setae, without processes. Subgenital plate elongate,
narrowed to midlength with apical half finger-like,
with several moderately long fine setae laterally.
Style broadened at base, apical process short,
lateral lobe distinct, short; with fine setae at dorsal
subapical area. Boundary between connective and
aedeagus obscure. Connective Y-shaped, short and
broad, stem not clearly defined. Aedeagus with
shaft short, curved dorsally, laterally compressed,
with pair of short processes subapically from
posterior surface, gonopore apical, socle broad in
lateral view, basal apodeme short.

Female genitalia: First valvulae with strigate
sculpture dorsally. Second valvulae similar to
Bhatia (Fig. 71).

REMARKS. This genus is similar to Bhatia
externally but has 2+1+1 rather than 2+2+1 hind
knee spines. From this and most other genera its
male genitalia can be distinguished by the robust
aedeagus with broad socle and with both shaft and
basal apodeme short.

Several specimens from the type series of
Melichariella boninensis (EYHU) and _ other
specimens from the Bonin Islands (BMNH, SU)
have been examined.

DISTRIBUTION. Bonin Islands.
SPECIES INCLUDED.

boninensis Matsumura, 1914: 238 (Melichariella).
LECTOTYPE 6, BONIN IS: Ogasawara,
20.vii.1905 (re-mounted and labelled on own pin)

Y. ZHANG AND M.D. WEBB

(EIHU), here designated (examined). Comb. n.
Figs 35, 278-283, 491.

WELMAYA gen. n.

Type species: Bythoscopus nigrilinea Walker

Description: Moderately large species (6.6-8.3
mm). Yellowish brown with darker brown markings
(Fig. 492).

External characters as in Waigara but with head
approximately as wide as pronotum; ocelli on
margin approximately own diameter from
corresponding eyes. Vertex very slightly longer
medially than next to eye. Pronotum obscurely
transversely wrinkled, side margins moderately
long, carinated. Fore tibia with dorsal surface
somewhat flattened. Hind femur with apical setal
formula 2+2+1.

Male genitalia: Pygophore lobes broadly
rounded with several macrosetae. Subgenital plate
elongate triangular with finger-like apex, a few
moderately long fine setae along lateral margin.
Style with apical process and lateral lobe short, the
former tapered to acute apex. Connective short,
stem bifurcate with an additional sclerite between
apex of arms and aedeagus. Aedeagus with shaft
filamentous, expanded apically, gonopore obscure;
basal apodeme elongate, compressed
anteroposteriorly. -

Female genitalia with well strigate sculpture
dorsally. Second valvulae similar to Bhatia.

REMARKS. This genus can be distinguished from
all other genera of the tribe by its unusual
connective and aedeagus as noted above.

In addition to the type, a few specimens from
throughout its range have been examined (BMNH,
BPBM, NMNB).

DISTRIBUTION. Malaya, Sabah, Sarawak and

Borneo.
INCLUDED SPECIES.

nigrilinea (Walker, 1857: 174) (Bythoscopus);
Distant, 1908: 146 (Drabescus). LECTOTYPE
36, SARAWAK (Wallace) (BMNH) here
designated, (examined). Comb. n. Figs 47,
284-290, 492.

Tribe Drabescini Ishihara

Drabescidae Ishihara, 1953: 6

Drabescinae Linnavuori, 1960b: 36; Linnavuori,
1978a: 41

Drabescini Linnavuori & Al-Ne’amy, 1983:21.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 23

Diagnosis: fore margin of head smooth or
irregularly striated, sometimes with a keeled
epistomal suture; clypeus rugose; antennae
moderately long and highly placed; antennal ledge
strong; dorsal surface of fore tibia distinctly
flattened; tentorium falcate; fore wing appendix
broad.

REMARKS. Contrary to Hill (1969) we have found
that the second valvulae are generally similar to
most Paraboloponini (Fig. 71). Although the head
is usually short, in an undescribed species from
Indonesia it is very long (Fig. 544). The nymph of
one species was studied (Fig. 543) and has the head
more elongate than in the adult (Fig. 542).

Although a ‘distinctive’ group all the above
diagnostic features are present in some other
subfamilies and tribes (see ‘Subfamily
relationships’) and if these were to be considered
primitive, as suggested by Linnavuori & Al-Ne’amy
(1983), then the group would be poorly defined.
The tribe has some similarity to Paraboloponini in
the shape of the tentorium and high antennae, and
some species, with paired basal aedeagal processes,
are similar to the paraboloponine genus Bhatia. In
addition, a few species of Bhatia have the foretibia
slightly flattened dorsally which is intermediate
between the distinctly flattened condition found in
Drabescini and the rounded condition found in
other Selenocephalinae.

We include only two genera in the tribe, judging
the genera and subgenera here synonymized under
Drabescus invalid, their differences being either
small compared to some other species or, we have
found intermediates (see also remarks under
Drabescus).

Key to genera of Asian and Pacific
Drabescini

1 Vertex shorter medially than next to eyes; anterior
margin of head with a ridge between ocelli; subgenital
plate with a row of stout macrosetae (Fig. 457); style
with several stout subapical setae (Fig. 463); aedeagus
without processes, basal apodeme strongly produced
dorsolaterally (Fig. 464) oo... eee Rengatella

— Vertex shorter to longer medially than length next to
eyes; anterior margin of head without a ridge between
ocelli, sometimes with a ridge between antennae;
subgenital plate and style without stout setae,
aedeagus with or without processes, basal apodeme

not or weakly produced dorsolaterally .... Drabescus
DRABESCUS Stal
Drabescus Stal, 1870: 738. Type species:

Bythoscopus remotus Walker, by monotypy

Drabescus (Ochrescus) Anufriev & Emeljanoy,
1988: 174. Type species: Drabescus ochrifrons
Vilbaste, by original designation. Syn. n.

Drabescus (Leucostigmidium) Anufrieyv &
Emeljanov, 1988: 174. Type species: Drabescus
nigrifemoratus Matsumura, by original
designation [misidentified = vilbastei sp. nov.].
Syn. n.

Paradrabescus Kuoh, 1985: 379 (382-383). Type
species: Paradrabescus testaceus Kuoh, by
original designation. Syn. n.

Tylissus Stal, 1870: 739. Type species: Tylissus
nitens Stal, by monotypy. Syn. n.

Within this large genus (currently comprising 37
species) only natalensis, from Africa, occurs outside
the study area. There is considerable variation
within the genus but the characters give
contradictory evidence of relationship. We would
suggest therefore that new genera are not described
for the included species until the many new species
seen are described and a detailed study of all the
species is undertaken.

The following notes apply to the species
examined. The type of ineffectus lacks its abdomen
and its identity (Figs 457-464) has been inferred
from a similar specimen also from northern China
(BMNH); similar specimens from elsewhere in
China include the types of a/bostriatus (see below)
and at least one new species. The male genitalia
figures of angulatus and formosanus are from a
specimen from Pakistan (BMNH) and China
(NAU) respectively, matched to the female types.
The identity of four other species from China,
albostriatus, fuscorufous, extensus and piceus, is
uncertain as males could not be associated with the
female holotypes of the first three species, and the
holotype of piceus, the only male, is poorly
preserved and possibly poorly developed. The
original figure of this latter species should be
consulted because the male genitalia are not figured
here. The habitus photograph given of piceus (Fig.
521) is of the allotype female, which matches the
holotype male but also, with the exception of the
pregenital sternite, matches the allotype female of
testaceus. All three specimens have a slightly
shorter vertex than the holotype male of testaceus
(Fig. 533). The association of these males and
females is therefore uncertain. Only the lectotype
female of stramineus (Plate 1, Fig. 2) from India, is
known. If this is not mis-labelled we hesitate to
identify three externally similar specimens from
Sulawesi (BMNH) as this species, (with male
genitalia similar to samoana), as the available
evidence suggests that Drabescus species are not
widely distributed. There is some variation in male
genitalia between specimens identified as samoana

24

(NMNH, BPBM, BMNH). This relates to the
pygophore process (Figs 363, 365) and the width of
the aedeagal shaft in posterior view, which is
slightly broader in specimens from the Philippines.
The type series of feraminensis (BPBM) is
represented by more than one _ species.
Unexplainably, the holotype of this species now
bears different data to that recorded in the original
decription and in the BPBM type list. The correct
data label could not be traced. The aedeagus and
style preapical angle of the holotype of gressitti
(Figs 436-439) differ slightly from similar
specimens from Papua New Guinea (BPBM,
BMNBH), which may or may not be this species. The
photograph of kempi is a specimen matched to the
original description and figure only, the type being
unavailable for study. The male genitalia of the type
of nitobei are missing but from the drawings made
by Dr Viraktamath of the type genitalia this species
and e/ongatus are identical. To maintain stability we
have selected nitobei as the senior name (both
species have the same date of publication) as being
the more commonly used name for this species. The
identification of females of nitobei and
nigrifemoratus (BMNH, SU) (Figs 335 and 350) is
tentative. The identity of nitens (the type species of
Tylissus = Drabescus) is uncertain; males of two
different species match the female type and one of
these is photographed (Fig. 539). These species and
other similar new speices (BPBM) (here refered to
as the nitens group ) represent one extreme of
variation in Drabescus in having the fore margin the
head strongly rounded in profile. As the head shape
in Drabescus is very variable and as the male
genitalia of the nitens group are similar to some
other Drabescus species we feel justified in
synonymising Tylissus with Drabescus.

Two new species, evansi and vilbastei, are
described below for misidentified figured specimens
in the literature.

DISTRIBUTION. Widespread in southern Asia and
the Pacific, and one species (natalensis) in Africa.

SPECIES INCLUDED.

albostriatus Yang, 1995: 42 (44), fig. 7. Holotype °,
CHINA (BAU) (examined).

angulatus Signoret, 1880b: 210 (120), pl.7, fig.73.
LECTOTYPE INDIA (NHMW) here
designated (examined). Figs 380-384, 525.

ater (Walker) (see remotus Walker).

atratus Kato 1933c: 456, pl.14, fig.17, TAIWAN
(Kato coll.) (not examined).

breviolus Matsumura, 1912: 292-293. Syntype °,
JAPAN (EIHU) here designated (examined).
Fig. 520.

brunneus Evans, 1972: 198-199, figs 11j. Holotype

Y. ZHANG AND M.D. WEBB

36, NEW GUINEA: IRIAN JAYA (BPBM)
(examined). Figs 431-435, 519.

conspicuus Distant, 1908: 306. LECTOTYPE 6d,
SRI LANKA: Peradeniya, 11.1901 (BMNH) here
designated (examined). Figs 358-362, 530.
limbaticeps (Stal, 1858: 453) (Selenocephalus):
Melichar, 1903: 170 (Drabescus). LECTOTYPE
?, SRI LANKA (IZW), here designated
(examined). Syn. n.

elongatus Matsumura, see nitobei Matsumura

evansi sp. n. for Drabescus heroni Evans, Evans
1972: 197-198, figs lla, f (misidentification).

extensus Kuoh, 1985: 377-378 (382) figs 1-2.
Holotype 2, CHINA (AAU) (examined). Fig.
Sills

fasciata Kato (see ineffectus (Walker).

femoratiformis Kwon & Lee, see nigrifemoratus
(Matsumura).

feraminensis Evans, 1972: 198, fig. 11h. Holotype
36, PAPUA NEW GUINEA _ (BPBM)
(examined). Figs 446-449, 534.

flavicollis Linnavuori, 1960a: 329 (Figs 47a-e).
Holotype 6, CAROLINE IS. (BPBM)
(examined). Plate 1, Fig. 3; Figs 369-374.

formosanus Matsumura, CHD: 294-295.
LECTOTYPE ¢, CHINA:TAIWAN (EIHU)
here designated (examined). Figs 375-379, 540.
trichromus Yang, 1995: 41-42 (44), fig. 6.
Holotype ¢, CHINA (BAU) (examined). Syn. n.

fuscorufous Kuoh, 1985: 378 (382), figs 3-4.
Holotype 2, CHINA (AAU) (examined). Fig.
518.

gressitti Evans, 1972: 198, figs 1lc-e. Holotype d,
PAPUA NEW GUINEA (BPBM) (examined).
Figs 436-439, 535.

heroni Evans, see samoanus Osborn

ineffectus (Walker, 1858: 266) (Bythoscopus);
Distant, 1908b: 145 (Dabescus [sic]). Holotype
(sex unknown, abdomen lost), CHINA (BMNH)
(examined). Figs 72, 320-326, 541.
fasciata (Kato, 1932: 224) (Athysanopsis); Kato,
1933b; pl.26, fig.7; Lee, 1979: 363, Plate 276, figs
1-13. Holotype 2, CHINA (Kato coll.) (not
examined). Syn. n.
ochrifrons Vilbaste, 1968: 116-118, figs 95a—g
(Drabescus). Holotype 6, RUSSIA (IZBT);
synonymized with fasciata Kato by Lee, 1979:
363; Syn.n.

kaindii Evans, 1972: 199, figs 11g. Holotype d,
IRIAN JAYA (BPBM) (examined). Figs
403-407, 537.

kempi Singh-Pruthi, 1930: 36, Pl. 3, figs 4, 4a.
Holotype °, INDIA (Indian Museum) not
examined. Fig. 517.

limbaticeps (Stal), see conspicuus Distant.

modestus Linnavuori, 1960a: 329-330, figs 48a—c.
Holotype ¢, CAROLINE IS. (NMNH)

ee

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 25

(examined). Figs 314-319, 536.

nakanensis Matsumura, see e/ongatus Matsumura.

natalensis Distant, 1910: 242, pl. 23, figs 11, lla.
Syntype 2 S. AFRICA (BMNH) (examined).

nervosopunctatus Signoret, 1880: 209, pl. 7, fig. 72.
LECTOTYPE ? INDIA (Wien) here designated
(examined). Figs 413-417, 527.

nigrifemoratus (Matsumura, 1905: 63, pl. 21, fig.
17) (Selenocephalus); Matsumura 1912: 290
(Dabrescus [sic]). LECTOTYPE ¢d, JAPAN: 091
Kubo, 17.vi, remounted and labelled (EIHU)
here designated (examined). Figs 333-338, 531.
femoratiformis Kwon & Lee, 1979: 53-54, fig. 4.
Holotype 2, KOREA (depository unknown).
Syn.n.

nigrofacialis (Distant, 1918: 42-43) (Kutara).
LECTOTYPE 6, INDIA: Chikkaballapura
(TV. Campbell) (BMNH) here designated
(examined) comb. n. Figs 327-332, 524.

nitens (Stal, 1870: 739-740) (Tylissus).
LECTOTYPE ¢, PHILIPPINES (NRS) here
designated (examined). comb. n. Fig. 539.

nitobei (Matsumura, 1912: 290-291) (Drabescus).
Holoiype 6, JAPAN (EIHU) (examined). Figs
345-349, 538.
elongatus Matsumura, 1912: 292. LECTOTYPE
3, JAPAN: Ishiyama, 7.viii.1903 (EIHU) here
designated (examined). syn. n.
nakanensis Matsumura, 1912: 293-294.
Holotype 36, JAPAN (EIHU) (examined). Syn.
n.

notatus Schumacher, 1915: 99. Syntypes (sex
unknown), TAIWAN (lost ?).

ochrifrons Vilbaste (see ineffectus (Walker))

ogumae Matsumura, 1912: 291-292. Lectotype 6,
JAPAN (EIHU) (examined). Plate 1, Fig. 1; Figs
418-424.

pallidus Matsumura, 1912: 293. Syntype 2 , JAPAN
(EIHU) (not examined). Plate 1, Fig. 4; Figs
351-357.

palmerstoni Evans, see samoanus Osborn

piceatus Kuoh, 1985: 378-379 (382), figs. 5-8.
Holotype ¢, CHINA (AAU) (examined). Figs
425-430, 529.

piceus (Kuoh, 1985: 381 (383), figs 16~19)
(Paradrabescus). Holotype 6, CHINA (AAU)
(examined). Comb. n. Fig. 521.

politus (Walker, 1870: 320) (Bythoscopus); Metcalf,
1966: 76 (fassus). LECTOTYPE 2
INDONESIA: Morotai I. (Wallace), (BMNH)
here designated (examined). Comb. n. Fig. 516.

remotus (Walker, 1851: 866) (Bythoscopus);
Signoret, 1880: 208 (118), pl.7 fig. 71
(Drabescus). LECTOTYPE (sex unknown,
abdomen lost), PHILIPPINES, (BMNH) here
designated (examined). Figs 25, 385-391, 514.
ater (Walker, 1851: 871) (Bythoscopus). Holotype

3, PHILIPPINES (BMNH)
Synonymized by Merino1936a: 369.

samoanus Osborn, 1934a: 176, fig. 10. Holotype d,
SAMOA (BPBM)(examined). Figs 363-368,
542, 543.
palmerstoni Evans, 1972: 199, fig. 11i. Holotype
3, AUSTRALIA (QM) (examined). Syn.n.
heroni Evans, 1966: 221, fig. 32h; Evans 1972:
197-198, figslla, f. Holotype ¢, AUSTRALIA
(AM) (examined). Syn.n.

shillongensis Rao, 1989: 65. Holotype ¢, INDIA
(ZSI) (examined). Figs 392-396, 532.

sirunkensis Evans, 1972: 198, fig. 11k. HOLOTYPE
36, PAPUA NEW GUINEA (BPBM)
(examined). Figs 450-456, 523.

stramineus Distant, 1908: 306-307, fig. 196.
LECTOTYPE 2, INDIA Nilgiri (Hampson)
(BMNH) here designated (examined). Plate 1,
Fig. 2.

testaceus (Kuoh, 1985: 380-381 (383), figs 9-15)
(Paradrabescus). Holotype ¢, CHINA (AAU)
(examined). Comb.n. Figs 397-402, 533.

trichromus Yang (see formosanus Matsumura)

vilbastei sp. n. for D. nigrifemoratus (Matsumura)
Vilbaste,1971: 105, figs 63-68; Anufriev &
Emeljanov, 1988: 175-176, pl. 116, Fig. 2, pl. 118,
figs 9-14; Lee, 1979: 356, pl. 266, figs 7-12;
misidentification.

vitreus Schmidt, 1926: 256. LECTOTYPE 4,
INDONESIA: Baru I., 1921 (L.J4. Toxopeus)
(IZW) here designated (examined). Figs
408-412, 522.

(examined).

SPECIES REMOVED FROM THE GENUS.

costalis Distant (see Kutara).

nigrilinea Walker (see Welmaya)

laetisigna (Walker) (see Roxasella)
metallicus (Walker) (see Rengatella)
productus Evans (see Kutara)

striatus Anufriev (see Drabescoides nuchalis)

Drabescus evansi sp. n.
(Figs 440-445, 526)

Drabescus heroni Evans, Evans 1972: 197-198, figs
11a, f, misidentification.

Length: d, 7.7-8.7mm, °, 8.0-8.4mm. Yellowish
brown, foremargin of head with a yellow band
between two dark brown bands. Face with a broad
yellow band laterally, continued onto thorax;
clypellus and clypeus sometimes dark brown.
Pronotum with numerous yellow spots, anterior
margin yellow. Sides of scutellum yellow. Forewing
veins dark brown with whitish yellow spots. Female

26

with ovipositor and pregenital sternite posteriorly
and medially, dark brown.

Vertex slightly longer medially than next to eyes.
Foremargin of head with a thick rim bordered
ventrally by a transverse ridge between antennae.
Fore tibia with dorsal surface non-dilated. Hind
femur with 2+1 stout setae. Female pregenital
sternite triangularly produced medially.

Male genitalia with posteroventral margin of
pygophore with a small tooth. Connective with
stem short. Style with apical process short, digitate.
Aedeagus with shaft narrow throughout length,
cylindrical, strongly curved dorsally with long
preatrium, the latter with a pair of very long
posteriorly directed processes.

MATERIAL EXAMINED. Holotype 6, Papua New
Guinea: Morobe Distr., Herzog Mts, Vagau, 4,000’,
4-17.1.1965 (BMNH).

Paratypes. Papua New Guinea: 3d, 6°, same
data as holotype (BMNH); 1 2, Wanuma, Adelbert
Mts, 800-1000m. (BMNH); 1d, Wau, 1200m.
(AMNH); 1d, Fergusson Isl. 900m. (AMNH).
Irian Jaya: 53,135 (RML).

REMARKS. This species has similar male genitalia
to feraminensis but differs in having the aedeagus
with a pair of basal processes rather than a single
bifurcate process and the shaft less strongly curved
dorsally and in lacking basal and apical articulating
struts of the aedeagus and connective respectively.

Drabescus vilbastei sp. n.
(Figs 339-343, 528)

Drabescus nigrifemoratus (Matsumura) Vilbaste,
1971: 105, figs 63-68, misidentification.

Length: ¢, 8.3mm; 2, 9.0mm. Colour and external
characters as in evansi but lower half of face yellow
rather than with a yellow transverse band laterally.

Male genitalia similar to pallidus but apical
process of style shorter and aedeagus with shaft
and processes longer and gonopore smaller.

DISTRIBUTION. Russia, Japan, N. China.

MATERIAL EXAMINED. Holotype 6, Japan:
Honshu, Saitama Pref., Nihongi Pass, 14.viti.1984,
(M. Hayashi et al.) (SU).

Paratypes. 1d, Japan: Ibaraki Pref., Takahagi,
Mt Tsuchi-dake, 4.1x.1986, (MZ Hayashi et al.)
(BMNH, SU). China: 16, Jilin, 20.viii.1939 (AS).
Russia: 1d, Nakhodka (ZIRAS); 136, Ussuriysk
(BMNH).

REMARKS. This species has been recorded and
figured from Russia by various authors as
nigrifemoratus following Vilbaste (1971).

Y. ZHANG AND M.D. WEBB

Although similar to nitobei and nigrofemoratus
externally its male genitalia are similar to pallida
(see above). Some topotypical females examined
(ZIRAS, SU) may be this species but the posterior
margin of the pregenital sternite is deeply concave
medially in specimens from Russia while in
specimens from Japan it is slightly produced
medially (see figure 344).

RENGATELLA gen. n.

Type species: Bythoscopus metallicus Walker

Brown with a yellow band between eyes at level of
ocelli and thorax densely marked with small yellow
spots.

Head wider than pronotum; anterior margin of
face rounded more or less in profile, smooth with a
few short transverse striations between ocellus and
eye; ocelli on margin, visible from above,
approximately three times own diameter from
corresponding eye; a transverse ridge between
lateral frontal sutures immediately below ocelli.
Anterior tentorial branches falcate. Vertex with
midlength very slightly shorter than length next to
eye, shiny with faint longitudinal striations; coronal
suture indistinct. Face shagreen with gena and
dorsal area of clypeus rugose; clypellus, expanded
apically with a medial ridge; transclypeal suture
obscure; laterofrontal suture extended laterad of
ocellus; antennal ledge strong, antennae high, near
upper corner of eye, long, nearly half body length.
Pronotum and scutellum hairy, the former
transversely wrinkled with side margins moderately
long and carinated. Forewing as in Fig. 26; veins
hairy. Fore tibia with dorsal surface flattened,
dorsal lateral margins sharp; hind femur with apical
setal formula 2+2+1.

Male genitalia: Pygophore with a posterior
process. Subgenital plate elongate,
triangular-shaped, with a row of stout macrosetae
over distal half. Style with base broad, apical
process digitate with several lateral setae; lateral
lobe short. Boundary between connective and
aedeagus obscure, the former Y-shaped, short and
broad, stem not clearly defined. Aedeagus with
shaft elongate, curved dorsally and tapered to apex,
cylindrical, without processes, with or without
lateral flanges, gonopore apical on posterior
surface; basal apodeme with strongly diverging
apical arms.

Female genitalia: First valvulae with strigate
sculpture dorsally. Second valvulae similar to
Bhatia (Fig. 71).

DISTRIBUTION. Sumatra, Malaya, Borneo.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 721)

REMARKS. This genus can be distinguished
externally from Drabescus by the characters noted
in the above generic key. We have matched the
female type of metallica from Sarawak to one of
two species from this region; the remaining species
is new. A further new species is from Sumatra and
Malaya. As the differences between these species
are slight and to avoid misidentifications the two
new species are described below.

SPECIES INCLUDED.

metallica (Walker, 1857: 173) (Bythoscopus).
LECTOTYPE 6 SARAWAK_ (Wallace)
(BMNH) here designated (examined). Comb.n.
Plate 1, Fig. 5; Figs 26, 457-464.

robustipenis sp.n.

malayensis sp.n.

Rengatella robustipenis sp. n.
(Figs 465-467)

Length: ¢, 10.0mm. External characters as in
generic description. Male genitalia differ from
metallicus and malayensis in having the style apical
process shorter and the aedeagal shaft more
elongate, with the socle larger and the arms of the
basal apodeme shorter.

DISTRIBUTION. Sarawak.

MATERIAL EXAMINED.
Holotype 6, Sarawak: Gunong Mulu Nat. Park,
1000m. (BMNH).

Paratypes. Sarawak: | ¢, Foot of Mt Dullit,
junction of river Injar and Lejok (BMNH).

Rengatella malayensis sp. n.
(Figs 468-470)

Length: 6, 9.0-10.2mm; @, 10.8-12.7mm.
External characters as in generic description. Male
genitalia differ from the similar metallica in having
the style apical process and aedeagal shaft shorter
and the latter more strongly curved basally in
lateral aspect. Female pregenital sternite similar to
metallica in having the posterior margin acutely
produced each side of mid-line but in malayensis
the margin is less insinuate.

DISTRIBUTION. Malaya, Sumatra.

MATERIAL EXAMINED. Holotype 6, Malaya:
Selangor, Bukit Kutu, 3400’, 13.11.1931 (BMNH).

Paratypes. Sumatra: 1d, 4¢6 (BMNH, RNHL,
ZMA); Malaya: 1 6, Perak, 12, Selangor
(BMNH).

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Deltocephalinae (Insecta: Hemiptera). Zoological Magazine

Tokyo 63: 243-245, figs. 1-2.

1961. Homoptera of Southeast Asia collected by the Osaka

City University Biological Expedition to Southeast Asia

1957-1958. Nature & Life in Southeast Asia 1:225-257.

28

Jacobi, A. 1943. Zur Kenntnis der Insekten von Mandschukuo 12.
Beitrag. Eine Homopterenfaunula der Mandschurei.
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Arbeiten iiber Morphologische u. Taxonomische Entomologie
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Kato, M. 1932. Notes on some Homoptera from South
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216-229, pl. VIII.

Kato, M. 1933a. Notes on some Manchurian Homoptera,
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— 1933b. Homoptera. Three-colour illustrated insects of Japan
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— 1933c. Notes on Japanese Homoptera, with descriptions of
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452-471, pls 14-15; 2 figs.

Kirkaldy, G.W. 1904. Bibliographical and nomenclatorial notes on
the Hemiptera. No.3. Entomologist 37: 279-283.

— 1906. Leafhoppers and their natural enemies. (Pt. IX,
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1907. Leafhoppers supplement. (Hemiptera.) Hawaii.
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Knight, W.J. 1973. A new species of Hishimonus Ishihara (Hom.,
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Y. ZHANG AND M.D. WEBB

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Indian Archipelago by Mr. A. R. Wallace, with descriptions of

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 29

new species. Journal of the Linnean Society (Zoology) 10:
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Entomotaxonomia 17: 9-14.

30 Y. ZHANG AND M.D. WEBB

Figs 1-21 Head and thorax of Selenocephalinae. 1-4, head and thorax, dorsal view; 1, Tambocerus disparatus; 2,
Hecaloidella nitida; 3, Bhatia olivacea; 4, Parabolopona guttata. 5—9, face; 5, Tambocerus disparatus; 6, Hecaloidella
nitida; 7, Bhatia olivacea; 8, Parabolopona guttata; 9, Isaca bipars. 10-12, head and thorax, left lateral view; 10,
Hecaloidella nitida; 11, Bhatia olivacea; 12, Parabolopona guttata. 13-14, Moluccasia notula; 13, face; 14, head and
thorax, left lateral view. 15-16, Nirvanguina placida, 15, face; 16, head and pronotum, dorsal view. 17—21, left anterior
tentorial branch. 17, Tambocerus disparatus; 18, Selenocephalus griseus; 19, Hecaloidella nitida; 20, Megabyzus signan-
dus; 21, Parabolopona guttata.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS

Figs 22-30 Forewings of Selenocephalinae. 22, Tambocerus plumbeus; 23. Moluccasia marmorea; 24, Hecaloidella
nitida; 25, Drabescus remotus; 26, Rengatella metallica; 27, Lamia cydippe; 28, Nirvanguina placida; 29, Kutara brun-
nescens; 30, Tenompoella difficilis.

31

32 Y. ZHANG AND M.D. WEBB

37 38

39 40

41 42

Figs 31-42 Forewings of Paraboloponini. 31, Carvaka modesta; 32, C. formosana; 33, C. contempta; 34, Hybrasil brani;
35, Waigara boninensis; 36, Oceanopona croceipennis; 37, Bhatia olivacea; 38, Athysanopsis salicis; 39, Parohinka longi-
seta; 40, Dryadomorpha pallida; 41, Karoseefa brevipenis; 42, Rhutelorbus merinoi.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 33

Figs 43-52 Forewings of Paraboloponini. 43, Parabolopona guttata; 44, Megabyzus signandus; 45, Mysolis flavidorsum;
46, Isaca bipars; 47, Welmaya nigrilinea; 48, Favintiga camphorae; 49, Roxasella losbanosa; 50, Omanella barberi; 51,
Tengatka basilana; 52, Sombakidia morisma.

34 Y. ZHANG AND M.D. WEBB

Figs 53-61 Left legs of Selenocephalinae. 53-57, Selenocephalus griseus. 53, fore femur; 54, apex of hind tibia; 55, apex
of hind femur; 56, hind tarsomeres; 57, fore tibia. 58-61, Parabolopona guttata. 58, hind tarsomeres; 59, apex of hind
tibia; 60, apex of hind femur; 61, fore femur & tibia, anterior view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 35

Figs 62-70 First valvulae of Selenocephalinae and Acostemminae, left lateral view. 62, Bhatia olivacea; 63, detail of
same at position arrowed in Fig. 62; 64, Omanella sp.; 65, detail of same at position arrowed in Fig. 64; 66, Sombakidia
morisma ; 67, detail of same at position arrowed in Fig. 66; 68, Acostemma walkeri; 69-70, detail of same at position
arrowed in Fig. 68; (arrow in Fig. 70 indicates area of strigate or forked sculpture).

36 Y. ZHANG AND M.D. WEBB

81 82

Figs 71-82 Ovipositor valvulae of Selenocephalinae, left lateral view. 71-80, second valvulae; 71, Bhatia olivacea; 72,
Drabescus ineffectus; 73, Karoseefa brevipenis; 74, Hybrasil brani; 75, Parabolopona guttata; 76, Omanella sp.; 77, Fav-
intiga camphorae; 78, Sombakidia morisma; 79, Lamia cydippe; 80, Nirvanguina placida. 8\—82, third valvulae; 81,
Lamia cydippe; 82, Nirvanguina placida.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 37

Figs 83-88 Male genitalia of Moluccasia marmorea; 83, pygophore, left lateral view; 84:aedeagus, left lateral view; 85,
subgenital plate, dorsal view; 86, connective, ventral view; 87, style, ventral view; 88, aedeagus, posterior view.

38 Y. ZHANG AND M.D. WEBB

Figs 89-98 Male genitalia of Tambocerus species. 89-93, T. disparatus; 89, genital capsule, left lateral view; 90, apex of
style; 91, Subgenital plate, valve, connective and style, ventral view; 92-93, aedeagus, posterior view and left lateral
view respectively. 94-98, T: plumbeus; 94, aedeagus, posterior view; 95, style apex; 96, pygophore lobe, left lateral view;
97, aedeagus, left lateral view; 98, connective, ventral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 39

Figs 99-104 Male genitalia of Hecaloidella nitida; 99, pygophore, left lateral view; 100, subgenital plate, valve and
style, dorsal view; 101, style, ventral view; 102, connective, dorsal view; 103-104, aedeagus and connective, ventral and
left lateral view, respectively.

40 Y. ZHANG AND M.D. WEBB

Figs 105-111 Male genitalia of Bhatia olivacea. 105, pygophore, left lateral view; 106, Subgenital plate, valve, connec-
tive and style, dorsal view; 107-108, aedeagus, anterior and posterior view, respectively; 109, style, ventral view; 110,
connective, ventral view; 111, aedeagus, left lateral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 41

Figs 112-120 Male genitalia of Bhatia species. 112-116, B. distanti, 112, aedeagus, anterior view; | 13, connective, ven-
tral view; 114, aedeagus and apex of connective, left lateral view; 115, aedeagus, posterior view; 116, style, ventral
view. 117-120, B. metcalfi, 117, apex of adeagal shaft, posterior view; 118, aedeagus and apical sclerite of connective,
left lateral view; 119, connective, ventral view; 120, style, ventral view.

42 Y. ZHANG AND M.D. WEBB

Figs 121-127 Bhatia guamensis. 121, style; 122, aedeagus, posteroventral view; 123, connective; 124-125, aedeagus, left
lateral and posterior view respectively; 126, sclerite of connective viewed from position arrowed in Fig. 127; 127, con-
nective, left lateral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS

Figs 128-133 Male genitalia of Bhatia satsumensis. 128, genital segment, left lateral view; 129, connective, ventral
view; 130, style, ventral view; 131-133 aedeagus, left lateral, posterior and ventral view, respectively.

43

44 Y. ZHANG AND M.D. WEBB

Figs 134-138 Male genitalia of Bhatia javana. 134, connective and base of aedeagus, ventral view; 135, style, ventral

view; 136, aedeagus, posterior view; 137, pygophore, and tenth segment left lateral view; 138, aedeagus, left lateral
view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 45

ie
Ife
Ve
ti
Up
ne
'
'
'
1
t
'
!
!
i}
1
1
'
i}
"

Figs 139-144 Male genitalia of Bhatia koreana. 139, genital capsule, left lateral view; 140, style, ventral view; 141, sub-
genital plate, valve, style and connective, dorsal view; 142, aedeagus, left lateral view; 143, connective, ventral view;

144, aedeagus, posterior view.

46 Y. ZHANG AND M.D. WEBB

Figs 145-150 Male genitalia of Athysanopsis salicis. 145, genital capsule, left lateral view; 146, aedeagus, left lateral
view; 147, valve, subgenital plate, style and connective, dorsal view; 148, style, ventral view; 149, connective, ventral
view; 150, aedeagus, posterior view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 47

Ze | 152

Figs 151-156 Male genitalia of Carvaka formosana. 151, genital capsule, left lateral view; 152, aedeagus, left lateral
view; 153, style, ventral view; 154, connective and base of aedeagus, ventral view; 155, aedeagus; 156, valve, subgenital
plate, connective and style, dorsal view.

48 Y. ZHANG AND M.D. WEBB

Figs 157-160 Male genitalia of Carvaka contempta. 157-158, conective and aedeagus, posterior and left lateral view,
respectively; 159, style ventral view; 160, pyhophore and tenth segment, left lateral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 49

Figs 161-167 Male genitalia of Hybrazil brani. 161, genital capsule, left lateral view; 162, style, ventral view; 163, base
of aedeagus and apex of connective, dorsal view; 164, valve, subgenital plate, connective and style, dorsal view; 165,
connective, ventral view; 166-167, aedeagus and apex of connective, posterior and left lateral view, respectively.

50 Y. ZHANG AND M.D. WEBB

Figs 168-174 Male genitalia of Kutara brunnescens. 168, genital capsule, left lateral view; 169, aedeagus, posterior
view; 170, connective, ventral view; 171, valve, subgenital plate, connective and style, dorsal view; 172, style, ventral
view; 173, apex of same in direction arrowed in Fig. 172; 174, aedeagus, left lateral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 51

180

Figs 175-180 Male genitalia of Kutara transversa. 175, genital capsule, left lateral view. 176, valve, subgenital plate,
connective and style, dorsal view; 177, style, ventral view; 178, connective, ventral view; 179-180, aedeagus, posterior
and left lateral view, respectively.

52 Y. ZHANG AND M.D. WEBB

Figs 181-188 Male genitalia of Kutara producta. 181, pygophore, left lateral view; 182, connective, ventral view; 183,
valve, subgenital plate, connective and style, dorsal view; 184, apex of aedeagus, anterior view; 185, apex of style; 186,
left style, ventral view; 187-188, aedeagus, posterior and left lateral view, respectively.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 38)

Figs 189-195 Male genitalia of Kutara sinensis. 189, pygophore and tenth segment, left lateral view; 190, valve, sub-
genital plate, connective and style, dorsal view; 191, connective, left lateral view; 192-193, aedeagus, posterior and left
lateral view respectively, 194, style, ventral view; 195, connective, ventral view.

54 Y. ZHANG AND M.D. WEBB

Figs 196-200 Male genitalia of Kutara costalis. 196, pygophore, left lateral view; 197, aedeagus, posterior view; 198,
valve, subgenital plate, connective and style, dorsal view; 199, style, ventral view; 200, aedeagus, left lateral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 55)

oe

Figs 201-207 Male genitalia of Kutara grisescens. 201, genital capsule, left lateral view; 202, apex of aedegus, dorsopo-
sterior view; 203, connective, ventral view; 204, valve, subgenital plate, connective and style, dorsal view; 205, style,
ventral view; 206-207, aedegus, left lateral and posterior view respectively.

56 Y. ZHANG AND M.D. WEBB

209

211

Figs 208-212 Male genitalia of Drabescoides nuchalis. 208, genital capsule, left lateral view; 209-211, variation of
pygopore lobe of same; 212, style, ventral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 3//

Figs 213-217 Drabescoides nuchalis. 213-215 aedeagus, anteroventral, posteroventral and left lateral view, respectively;
216, connective, base of aedeagus, subgenital plate and style, dorsal view; 217, connective and base of aedeagus, ven-
tral view.

58 Y. ZHANG AND M.D. WEBB

Figs 218-224 Male genitalia of Isaca bipars. 218, genital capsule, left lateral view; 219, tenth segment, left lateral view;
220, aedegus, left lateral view; 221, valve, subgenital plate, connective, aedeagus & style, dorsal view; 222, style, ventral
view; 223, connective, ventral view; 224, aedeagus, posterior view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS a9

eae =
ssssv

| Figs 225-233. Male genitalia of Jsaca species. 225-228, I. sinuata. 225, pygophore and tenth segment, left lateral view;
226, tenth segment, left lateral view; 227-228, aedeagus and apex of connective, ventral and left lateral view, respec-
tively. 229-233, I. falcata. 229, pygophore and tenth segment, left lateral view; 230, tenth segment; 231, connective and
base of aedeagus; 232-233, aedeagus, ventral and left lateral view respectively.

60 Y. ZHANG AND M.D. WEBB

Figs 234-240 Male genitalia of Roxasella losbanosa. 234, genital capsule, left lateral view; 235, connective, ventral
view; 236, valve, subgenital plate connective and style, dorsal view; 237, left style, ventral view; 238-240, aedeagus, left
lateral, ventroposterior and posterior view, respectively.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 61

Figs 241-249 Male genitalia of Roxasella species. 241-245, R. biarcuata. 241, left style, ventral view; 242-243, aedea-
gus, left lateral and posterior view, respectively; 244, apex of aedeagus, posterior view; 245, connective, ventral view.

246-249, R. philippina. 246, left style, ventral view; 247-248, aedeagus left lateral and posterior view, respectively; 249,
connective, ventral view.

62 Y. ZHANG AND M.D. WEBB

Figs 250-255 Male genitalia of Roxasella laetisigna. 250, genital capsule, left lateral view; 251, valve, subgenital plate,
connective, aedeagus and style, dorsal view; 252, aedeagus, posterior view; 253, aedeagus and connective, left lateral
view; 254, connective, ventral view; 255, left style, ventral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 63

yy
4, ZZ) YY;
QD

= Foe Uy pr va = SF SE: iy

; SCS eile

ess J

—————

ae ee
=

Figs 256-263 Male genitalia of Omanella johnsoni. 256, genital capsule, left lateral view; 257, valve, subgenital plate,
connective, aedeagus and style, dorsal view; 258, apex of aedeagus, dorsoposterior view (Philippines); 259-260, left
style; 261-262, aedeagus, left lateral and dorsal view respectively; 263, apex of aedegus (Thailand), dorsoposterior
view.

H

64 Y. ZHANG AND M.D. WEBB

Figs 264-270 Male genitalia of Omanella barberi. 264, genital capsule, left lateral view; 265, left style; 266-267, aedea-
gus, left lateral and dorsal view, respectively; 268-269, aedeagus and connective, left lateral and ventral view, respec-
tively; 270, left style, ventral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 65

Figs 271-277 Male genitalia of Tenompoella difficilis. 271, genital capsule, left lateral view; 272, valve, subgenital plate,
connective, aedeagus and style; 273-274, aedeagus, dorsoposterior and ventroposterior view, respectively; 275, apex
of left style, dorsal view; 276, aedeagus and connective, left lateral view; 277, left style, ventral view.

66 Y. ZHANG AND M.D. WEBB

Figs 278-283 Male genitalia of Waigara boninensis. 278, genital capsule, left lateral view; 279, valve, subgenital plate,
connective, base of aedeagus and style, dorsal view; 280-281, aedeagus and apex of connective, posterior and left lat-
eral view, respectively; 282, left style, ventral view; 283, connective and base of aedeagus, ventral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 67

287 288
a)

Figs 284-290 Male genitalia of Welmaya nigrilinea. 284, genital capsule, left lateral view; 285, left style, ventral view;
286, aedeagus, left lateral view; 287-288, detail of aedeagal shaft at apex and base, respectively, at positions and direc-
tion arrowed in Fig. 286; 289, connective, ventral view; 290, valve, subgenital plate, connective, aedeagus and style,
dorsal view.

68 Y. ZHANG AND M.D. WEBB

Figs 291-297 Male genitalia of Tengatka basilana. 291, genital capsule, left lateral view; 292, connective, ventral view;
293, valve, subgenital plate, connective and style, dorsal view; 294, connective, left lateral view; 295, left style, ventral
view; 296-297, aedeagus, left lateral view and posterior view, respectively.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 69

Figs 298-307 Male genitalia of Sombakidia morisma. 298, genital capsule, left lateral view; 299, aedeagus, posterior
view; 300, connective, ventral view; 301, valve, subgenital plate, connective and style, dorsal view; 302, valve and sub-
genital plate, left lateral view, to show variation in shape of subgenital plate compared to fig. 298; 303, apex of aedea-
gus, viewed anteriorly in direction of arrow in Fig. 304; 304, aedeagus, left lateral view; 305, left subgenital plate,
dorsal view; 306, connective, left lateral view; 307, left style, ventral view.

70 Y. ZHANG AND M.D. WEBB

312

313

Figs 308-313 Male genitalia of Nirvanguina placida. 308, genital capsule, left lateral view; 309, left style, ventral view;
310, right pygophore process, medial view; 311, valve, subgenital plate, connective and style, dorsal view; 312, aedea-
gus, posterior view; 313, aedeagus and apex of connective, left lateral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS

Figs 314-319 Male genitalia of Drabescus modestus (holotype), 314, genitale capsule, left lateral view; 315, left style,
ventral view; 316, aedeagus and apex of connective, left lateral view; 317, connective and base of aedeagus, ventral
view; 318, aedeagus, posterior view; 319, valve, subgenital plate, connective and style, dorsal view.

71

72 Y. ZHANG AND M.D. WEBB

Figs 320-326 Male genitalia of Drabescus ineffectus. 320, genital capsule, left lateral view; 321, connective, ventral
view; 322, left style, ventral view; 323, connective and base of aedeagus, left lateral view; 324, valve, subgenital plate,
connective and style, dorsal view; 325-326, aedeagus, posterior and left lateral view, respectively.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 73

Figs 327-332 Male genitalia of Drabescus nigrofacialis. 327, genital capsule, left lateral view; 328, connective, ventral
view; 329, aedeagus, left lateral view; 330, valve, subgenital plate, connective, aedeagus and style, dorsal view; 331,
aedeagus and apex of connective, ventral view; 332, left style, ventral view.

74 Y. ZHANG AND M.D. WEBB

Figs 333-338 Male genitalia and female pregenital sternites of Drabescus nigrifemoratus (syntypes). 333, male pygo-
phore and tenth segment, left lateral view; 334, aedeagus, posterior view; 335, female pregenital sternites; 336, aedea-
gus and apex of connective, left lateral view; 337, left style, ventral view; 338, connective and base of aedeagus, ventral
view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 75

Figs 339-350 Male genitalia and female pregenital sternites of Drabescus species. 339-343, D. vilbastei; 339, connective
and base of aedeagus, ventral view; 340, aedeagus, posterior view; 341, aedeagus and apex of connective, left lateral
view; 342, left style apex; 343, left style, ventral view. 344, Drabescus spp. female pregenital sternites (see D. vilbastei
remarks); 345-349, D. nitobei, 345, aedeagus and apex of connective, left lateral view; 346, connective, ventral view;
347, aedeagus and apex of connective, ventral view; 348, male pygophore and tenth segment, left lateral view; 349, left
style, ventral view; 350, D. nitobei? (see Drabescus remarks) female pregenital sternites showing variation in pigment
between different specimens.

76 Y. ZHANG AND M.D. WEBB

Figs 351-357 Male genitalia of Drabescus pallidus. 351, pygophore and tenth segment, left lateral view; 352, apex of
style; 353, aedeagus and apex of connective, left lateral view; 354, left style, ventral view; 355, connective and base of
aedeagus, ventral view; 356, left pygophore lobe, left lateral view; 357, aedeagus, posterior view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS

Figs 358-362 Male genitalia of Drabescus conspicuus (lectotype). 358, genital capsule; 359, aedeagus and connective,
ventral view; 360, left style, ventral view; 361, aedeagus and apex of connective, left lateral view; 362, aedeagus and
apex of connective, ventral view.

Lil

78 Y. ZHANG AND M.D. WEBB

Figs 363-368 Male genitalia of Drabescus samoanus. 363, pygophore and tenth segment, left lateral view; 364, connec-
tive and base of aedeagus, ventral view; 365, pygophore lobe, left lateral view; 366, aedeagus, ventral view; 367, aedea-
gus and apex of connective, left lateral view; 368, left style, ventral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS

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i
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Figs 369-379 Male genitalia of Drabescus species. 369-374, D. flavicollis (holotype). 369, pygophore and tenth seg-
ment, left lateral view; 370, aedeagus and connective, left lateral view; 371, left style, ventral view; 372, connective, ven-
tral view; 373, aedeagus and apex of connective, ventral view; 374, pygophore lobe, left ventral view; 375-379, D.

formosanus. 375, pygophore, left lateral view; 376-377, aedeagus, posterior and left lateral view; 378, left style, ventral
view; 379, connective, ventral view.

79

80 Y. ZHANG AND M.D. WEBB

Figs 380-384 Male genitalia of Drabescus angulatus. 380, pygophore and tenth segment, left lateral view; 381, aedea-

gus, posterior view; 382, connective and base of aedeagus, ventral view; 383, aedeagus left lateral view; 384, left style,
ventral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS

Figs 385-391 Male genitalia of Drabescus remotus. 385, genital capsule, left lateral view; 386, connective and base of
aedeagus, ventral view; 387, aedeagus, posterior view; 388, valve, subgenital plate, connective and style, dorsal view;
389, connective, left lateral view; 390, aedeagus, left lateral view; 391, left style, ventral view.

81

82 Y. ZHANG AND M.D. WEBB

393

Figs 392-396 Male genitalia of Drabescus shillongensis. 392, pygophore and tenth segment, left lateral view; 393,
aedeagus, posterior view; 394, left style, ventral view; 395, connective and base of aedeagus, ventral view; 396, aedea-
gus, left lateral view. Figs 392, 394-395, Yunnan specimen, Figs 393, 396, holotype.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 83

Figs 397-402 Drabescus testaceus (holotype). 397, pygophore, left lateral view; 398, connective, ventral view; left style,
ventral view; 399, left style ventral view, 400, aedeagus, left lateral view; 401, valve and subgenital plate dorsal view;
402, aedeagus, posterior view.

84 Y. ZHANG AND M.D. WEBB

Figs 403-407 Male genitalia of Drabescus kaindii (holotype). 403, aedeagus, posterior view; 404, pygophore and tenth
segment, left lateral view; 405, aedeagus, left lateral view; 406, connective, ventral view; 407, left style, ventral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 85

Figs 408-412 Male genitalia of Drabescus vitreus (Lectotype). 408, pygophore and tenth segment, left lateral view; 409,
left style, ventral view; 410, connective, ventral view; 411-412, aedeagus, posterior and left lateral view, respectively.
Subgenital plate as in remotus, fig. 388.

86 Y. ZHANG AND M.D. WEBB

Figs 413-417 Male genitalia of Drabescus nervosopunctatus. 413, pygophore and tenth segment, left lateral view; 414,
aedeagus and apex of connective, posterior view; 415, left style, ventral view; 416, aedeagus, left lateral view; 417, con-
nective and base of aedeagus, ventral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 87

Figs 418-424 Male genitalia of Drabescus ogumae. 418, genital capsule, left lateral view, 419, valve, subgenital plate,
connective and style, dorsal view; 420, left style, ventral view; 421, connective, ventral view; 422, aedeagus, posterior
view, 423-424 aedeagus ventral and left lateral view, respectively.

88 Y. ZHANG AND M.D. WEBB

Figs 425-430 Male genitalia of Drabescus piceatus (holotype). 425-426, pygophore and tenth segment, left lateral and
ventral view respectively; 427, connective and base of aedeagus, ventral view; 428, aedeagus, ventral view; 429, aedea-
gus and apex of connective, left lateral view; 430, left style, ventral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS

Figs 431-439 Male genitalia of Drabescus species. 431-435, D. brunneus (holotype). 431, pygophore, tenth segment,
connective and aedeagus, left lateral view; 432, left style, ventral view; 433-434, aedeagus, posterior and left lateral
view, respectively; 435, connective, ventral view. 436-439, D. gressitti (holotype). 436, apex of left style, ventral view;
437-438, aedeagus, left lateral and posterior view, respectively; 439, connective, ventral view.

89

90 Y. ZHANG AND M.D. WEBB

Figs 440-445 Male genitalia of Drabescus evansi. 440, genital capsule, left lateral view; 441, aedeagus, posterior view;
442, connective, ventral view; 443, apex of left style, dorsal view; 444, aedeagus and apex of connective, left lateral
view; 445, left style, ventral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 91

Figs 446-449 Male genitalia of Drabescus feraminensis (holotype). 446-447, aedeagus and connective, ventral and left
lateral view, respectively; 448, pygophore lobe, right lateral view; 449, left style, ventral view.

92 Y. ZHANG AND M.D. WEBB

Figs 450-456 Male genitalia of Drabescus sirunkensis (holotype). 450, pygophore and tenth segment, left lateral view;
451, left style, ventral view; 452-453, connective and base of aedeagus, ventral and left lateral view; 454, aedeagus,
ventral view; 455, valve and subgenital plate dorsal view; 456, aedeagus, left lateral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 93

Figs 457-462 Male genitalia of Rengatella metallica. 457, genital capsule, left lateral view; 458, valve, subgenital plate,
connective and style, dorsal view; 459, connective, ventral view; 460-461, aedeagus, posterior view, Sarawak and Bru-
nei respectively; 462, aedeagus and apex of connective, dorsal view.

94 Y. ZHANG AND M.D. WEBB

Figs 463-470 Male genitalia of Rengatella species. 463-464, R. metallica. 463, left style, ventral view; 464, aedeagus
and connective, left lateral view. 465-467, R. robustipenis. 465, aedeagus, posterior view; 466, aedeagus, left lateral
view; 467, left style, ventral view. 468-470, R. malayensis. 468, aedeagus, left lateral view; 469, aedeagus, posterior
view; 470, left style, lateral view.

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 95

474

480

482

Figs 471-482 Selenocephalini and Paraboloponini species. 471, Moluccasia notula (lectotype); 472, Tambocerus dis-
paratus (paralectotype); 473, T. plumbeus (lectotype); 474, Hecaloidella nitida (holotype); 475, Karoseefa divergens
(holotype); 476, K. brevipenis (holotype); 477, Parohinka longiseta (Brunei, BMNH); 478, Rhutelorbus merinoi (para-
type 2); 479, Megabyzus signandus (Sri Lanka, Maha, BMNH); 480, Oceanopona croceipennis (paratype) 481, Favin-
tiga camphorae (Japan, BMNH); 482, Divus bipunctatus (Sri Lanka, Peradeniya, BMNH).

96 Y. ZHANG AND M.D. WEBB

483 485 486

488 489 490

493

Figs 483-494 Paraboloponini species. 483, Bhatia olivacea (BMNH); 484, B. distanti; 485, B. javana (lectotype); 486, B.
metcalfi (Yap Is, BMNH); 487, B. lituriceps (lectotype); 488, B. satsumensis (Taiwan); 489, B. guamensis (holotype);
490, B. koreana (Kwon coll.); 491, Waigara boninensis (Ryukyus, BMNH); 492, Welmaya nigrilinea (Sarawak,
BMNH); 493, Omanella barberi(d , Zamboanga, BMNH); 494, O. johnsoni(°, Basilan I., BMNH).

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS oT

495

504

Figs 495-505 Paraboloponini species. 495, Kutara lucidicosta? (see Kutara remarks) (Papua New Guinea, Kokoda,
BMNH); 496, K. costalis (Borneo, Sandakan, BMNH); 497, K. sinensis (China, Foochow, BMNH); 498, K. brunnes-
cens (2, Sri Lanka, Alutnuwara, BMNH); 499, K. producta (N. Borneo, Bettotan, BMNH); 500-501, exuviae associ-
ated (on pin) with holotype of previous species; 502, K. transversa (paratype); 503, K. grisescens (Holotype); 504,
Drabescoides nuchalis; 505, Roxasella egregia (Burma, Myitta, BMNH).

98 Y. ZHANG AND M.D. WEBB

509

510 511

Figs 506-513 Paraboloponini species. 506, Carvaka ochrophara (India, Kodaikanal, BMNH); 507, C. picturata (syn-
type, BMNH); 508, C. thoracica (syntype, BMNH); 509, C. modesta (syntype, BMNH); 510, C. formosana (lectotype);
511, C. contempta (Australia, New South Wales, BMNH); 512, Tengatka basilana (Basilan I., BMNH); 513, Sombak-
idia morisma (allotype NMNH).

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 99

Figs 514-525 Drabescus species. 514, D. remotus (Philippines, Baguio, BMNH); 515, D. extensus (holotype); 516, D.
politus (holotype); 517, D. kempi (India, Karnataka, UASB); 518, D. fuscorufous (holotype); 519, D. brunneus (para-
type); 520, D. breviolus (2syntype); 521, D. piceus (allotype); 522, D. vitreus (lectotype); 523, D. sirunkensis (holotype);
524, D. nigrofacialis (lectotype); 525, D. angulatus (Nepal, BMNH).

100 Y. ZHANG AND M.D. WEBB

536

Figs 526-537 Drabescus species. 526, D. evansi (paratype, BMNH); 527, D. nervosopunctatus (India, Cherrapunji,
UASB); 528, D. vilbastei (holotye); 529, D. piceatus (holotype); 530, D. conspicuus (paratype, BMNH); 531, D.
nigrifemoratus (lectotype); 532, D. shillongensis (China, BMNH); 533, D. testaceus (holotype); 534, D. feraminensis
(holotype); 535, D. gressitti (holotype); 536, D. modestus (holotype); 537, D. kaindii (holotype).

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS 101

540

542

Figs 538-544 Drabesciini species. 538. Drabescus nitobei (lectotype of D. elongatus); 539, D. nitens? (see Drabescus
remarks), Philippines, BMNH; 540, D. formosanus (lectotype); 541, D. ineffectus (China, BMNH); 542, D. samoanus
(paratype, BMNH); 543, nymph of previous species (BMNH); 544, Drabescini indet. species.

102

INDEX

Y. ZHANG AND M.D. WEBB

albostriatus 24
anacryon 14
angulatus 24
antennalis 14
Athysanopsis 11
apicalis 19

ater 25

atratus 24
australis 14

barberi 18

basilana 21

Bhatia 11

Bhatiini9

biarcuata 20

bipars 15

bipunctatus 13
boninensis 22

brani 14

brevicephala 19
breviolus 24
brevipenis 15
bruneocephala 14
brunnescens 16
brunneus (Carvaka) 13
brunneus (Drabescus) 24

Calotettix 6
camphorae 14
camusi 20
Carvaka 13
Cestius 6
chatterjeei 14
chinensis 18
contempta 13
conspicuus 24
costalis 16
croceipennis 18
cydippe 17

difficilis 21
Discocephalana 6
disparatus 9
distanti 12
divergens 15
Divus 13

dolens 13
Drabescini 22
Drabescoides 13
Drabescus 23
Dryadomorpha 13
dulita 19

egregia 20
elongatus 24
eurytus 13
evansi 25
extensus 24

falcata 15
fasciata 24
Favintiga 14
femoratiformis 24

Synonyms are in italics.

feraminensis 24
flavicollis 24
flavidorsum 17
formosana 13
formosanus 24
fulvida 13
fuscorufous 24

Gessius 6
gressitti 24
grisescens 16
guamensis 12
guttata 19
guttatus 7

Hecaloidella 8
heroni 25
Hybrasil 14

ineffectus 24
indicus 14
Isaca 14
ishihari 19

Jamitettix 15
javana 12
johnsoni 18

kaindii 24
Karoseefa 15
katoi 11
kempi 24
Khamiria 6
koreana 12
Koreanopsis \\
kotonis 15
Kutara 15

laetisigna 20
Lamia 16
Leucostigmidium 23
lais 14
limbaticeps 24
lituriceps 12
longiseta 19
losbanosa 20
lotophagorum 19
lucidicosta 16
luzonensis 19

malayensis (Parohinka) 19
malayensis (Rengatella) 26
marmorea 8

Megabyzus 17
Melichariella \1

merinoi 19

metallica 26

metcalfi 12

metrosideri 14

modesta 13

modestus 24

Moluccasia 8

morisma 20

morona 19
multicolor 17
Mysolis 17

Nakula 17
nakanensis 24
natalensis 24
nervosopunctatus 24
nigrifasciata 16
nigrifemoratus 24
nigrilinea 22
nigrofacialis 24
Nirvanguina 17
nitens 24

nitida 8

nitobei 24

Norsia 17
Norsiana 17
notatus 24

notula 8

nuchalis 13

Oceanopona 18
Ochrescus 23
ochrifrons 24
ochrophara 13
ogumae 24
olivacea 12
Omanella 18
Osbornitettix 6

pacifica 14

Paganalia 6

pallida 14

pallidus 24

palmerstoni 25
Parabolopona 18
Paraboloponini 9
Paradrabescus 23
Parohinka 19

philippina (Parohinka) 19
philippina (Roxasella) 20
Philippinensis 20

piceatus 24

piceus 24

picturata 13

placida 18

plumbeus 9

politus 25

producta 16

quadricornis 14
quadripunctatus 11

recurva 19
remotus 24
Rengatella 26
Rhombopsana 6
Rhombopsis 6
Rhutelorbus 19
robustipenis 26
Roxasella 19

CLASSIFICATION OF SELENOCEPHALINE LEAFHOPPERS

samoanus 25
salicis 11
satsumensis 12
Selenocephalus 7
shillongensis 25
Shivapona 10
signandus 17
sinensis 16
sinuata (Isaca) 15
sinuata (Parohinka) 19
sirunkensis 25
Sohipona 10
Sombakidia 20
spinosa 19
stramineus 25
striatus \3

taiwanus 7
Tambocerus 8
Tenompoella 21
Tengatka 20
testaceus 25
thoracica 13
tincta 14
transversa 16
trichromus 24
trispicata 19
Tylissus 23

vilbastei 25
virens 14
virescens (Nephotettix) 7

virescens (Dryadomorpha) 14
viridia 14

Viridipes 7

viridis 14

vitreus 25

Waigara 21
Welmaya 22

Yakunopona 6
yakushimensis 14

yangi 19

Zizyphoides 6

103

Ra

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7 ’

Bulletin of The Natural History Museum

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Earlier Entomology Bulletins are still in print. The following can be ordered from Intercept (address on inside front
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A revised classification of the Asian and Pacific selenocephaline
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Y. Zhang and M.D. Webb

ENTOMOLOGY SERIES
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