European Journal of Taxonomy 45: 1-211

http://dx.doi.org/10.5852/eit.2013.45

ISSN 2118-9773

www. europeani ournaloftaxonomv. eu

2013 • Za hni ser J.N. & Dietrich C.H.

This work is licensed under a Creative Commons Attribution 3.0 License.

Monograph

urn:lsid:zoobank.org:pub:41B10E4D-7DAB-40CA-A8FE-4ECA078E04A3

A review of the tribes of Deltocephalinae

(Hemiptera: Auchenorrhyncha: Cicadellidae)

James N. ZAHNISER 1 * & Chris H. DIETRICH 2

12 Illinois Natural History Survey, Prairie Research Institute,

University of Illinois at Urbana-Champaign,

1816 S. Oak St., Champaign, IL 61820, United States of America

* corresponding author e-mail: [email protected]

1 um:lsid:zoobank.org:author:C44D6E44-FAlC-4B29-B7BB-FAF5940CD225

2 um:lsid:zoobank.org:author:82FCB86C-54B4-456A-AE5E-D7847D271CB9

Abstract. The classification of the largest subfamily of leafhoppers, Deltocephalinae, including 38

tribes, 923 genera, and 6683 valid species, is reviewed and revised. An updated phylogeny of the

subfamily based on molecular (28S, Histone H3) and morphological data and an expanded taxon sample

(37 taxa not included in previous analyses) is presented. Based on the results of these analyses and on

the morphological examination of many representatives of the subfamily, the classification of the tribes

and subtribes of Deltocephalinae is revised. Complete morphological descriptions, illustrations, lists of

the included genera, and notes on their distribution, ecology, and important vector species are provided

for the 38 recognized tribes and 18 subtribes. A dichotomous key to the tribes is provided. All names in

the taxonomic treatments are hyperlinked to online resources for individual taxa which are supported

by a comprehensive database for Deltocephalinae compiled using the taxonomic database software

package 31. The online functionality includes an interactive key to tribes and subtribes and advanced

database searching options. Each taxon (subspecies through subfamily) has a unique taxon webpage

providing nomenclatural information, lists of included taxa, an automated description (if available),

images (if available), distributional information, bibliographic references and links to outside resources.

Some observations and trends regarding the history of taxonomic descriptions in Deltocephalinae are

reported. Four new tribes are described: Bahitini tribe nov. (25 genera), Bonsapeiini tribe nov. (21

genera), Phlepsiini tribe nov. (4 genera), and Vartini tribe nov. (7 genera). The circumscription and

morphological characterization of Scaphoideini Oman, 1943 (61 genera) is substantially revised. Eleven

new species are described: Acostemma stilleri sp. nov., Arrugada linnavuorii sp. nov., Drabescus zhangi

sp. nov., Parabolopona webbi sp. nov., Goniagnathus emeljanovi sp. nov., Hecalus hamiltoni sp. nov.,

Scaphoideus omani sp. nov., Dwightla delongi sp. nov., Abimwa knighti sp. nov., Gannia viraktamathi

sp. nov., and Doratulina dmitrievi sp. nov. Some family-group level taxonomic changes are made:

Platymetopiini Haupt, 1929, Anoterostemmini Haupt, 1929, and Allygidiina Dmitriev, 2006 are

synonymized with Athysanini Van Duzee, 1892, syn. nov.; Procepitini Dmitriev, 2002 is synonymized

with Cicadulini Van Duzee, 1892, syn. nov.; Listrophorini Boulard, 1971 is synonymized with Chiasmini

Distant, 1908, syn. nov.; Adamini Linnavuori & Al-Ne’amy, 1983, Dwightlini McKamey, 2003, and

Ianeirini Linnavuori, 1978 are synonymized with Selenocephalini Fieber, 1872 syn.nov., and all three

are now recognized as valid subtribes in their parent tribe. New placements of many genera to tribe and

subtribe are made, and these are described in individual taxon treatments.

European Journal of Taxonomy 45: 1-211 (2013)

Keywords. Leafhopper, classification, taxonomy, phylogeny, database, interactive key, cybertaxonomy.

Zahniser J.N. & Dietrich C.H. 2013. A review of the tribes of Deltocephalinae (Hemiptera: Auchenorrhyncha:

Cicadellidae). European Journal of Taxonomy 45: 1-211. http://dx.doi.org/10.5852/eit.2013.45

Introduction

Deltocephalinae contains 6683 valid species and 923 genera (Fig. 1) currently making it the largest

subfamily of Cicadellidae based on the number of described species. The subfamily is distributed

worldwide, and it contains the majority of leafhopper vectors of economically important plant diseases,

some of which cause significant damage and economic loss. Many species feed on herbaceous or

woody dicotyledonous plants, while about 1/3 of the tribes specialize on grass and sedge hosts and are

particularly diverse and abundant in grassland ecosystems.

The family-group classification of Deltocephalinae has been somewhat controversial and has seen many

changes over the past 50 years. The controversy has stemmed in part from the lack of global revisions

and phylogenetic studies that provide the scope needed to address the classification of this cosmopolitan

group of leafhoppers. Also, taxonomic concepts of many family-group taxa have often not been clearly

stated or universally agreed upon, or even if they have, their circumscriptions have often not been tested

by examination of multiple character sets or in phylogenetic analyses. These deficiencies have been

partly addressed by taxonomic revisions (e.g., Linnavuori & Al-Ne’amy 1983, Knight & Webb 1993,

Zhang & Webb 1996, Dietrich & Rakitov 2002, Dietrich & Dmitriev 2003) and recent phylogenetic

studies based on morphological (Dietrich 1999, Knight & Webb 1993, Zahniser & Dietrich 2008) and

molecular data (Dietrich et al. 2001, Zahniser & Dietrich 2010). These studies provided strong evidence

8000

7000

6000

5000 -

4000

3000

2000

1000 -

1758

38 tribes

18 subtribes

923 genera

173 subgenera

6683 species

274 subspecies

8109 valid taxa

♦«*

i I 1 I

1800 1850 1900 1950

2000

Fig. 1 . Cumulative number of valid taxa (subspecies through subfamily) described 1758 to present.

X-axis indicates the year; Y-axis indicates cumulative number of valid taxa.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

that Deltocephalinae sensu Oman et al. (1990) was not monophyletic and that several other leafhopper

subfamilies had their closest relatives within the deltocephaline lineage. Based on the accumulated data

and understanding of the relationships of Deltocephalinae and other leafhoppers, Zahniser & Dietrich

(2010) adopted a revised, broader concept of the subfamily and provided a detailed history of the

classification of the subfamily.

Despite the knowledge that has been gained in recent years, a comprehensive understanding of the

relationships of all Deltocephalinae is far from complete. More taxa and more data need to be included

in future phylogenetic studies to obtain a more resolved and supported phylogeny of the subfamily. In

the current study, the taxon sample was increased over previous phylogenetic analyses by 37 taxa. The

updated phylogenetic analyses presented here are used in combination with morphological examination

of numerous representatives of Deltocephalinae to review all of the family-group taxa (tribes and

subtribes) of Deltocephalinae and provide revised morphological descriptions, revised classifications,

complete listings of genera included in each tribe and subtribe, and information on their phylogenetic

relationships, classification status, ecology, vector species, and distribution. The revised classification

clears up some unsatisfactory elements and oversights of previous classifications. It also proposes some

changes that are supported in part by the most current phylogenetic analyses of the subfamily presented

here. While more focused effort is needed to clarify some aspects of the classification, this represents a

step forward toward a classification that reflects the phylogeny of the subfamily, as far as it is known.

Perhaps the most problematic aspects of the previous classification regard the 51 genera that were

unplaced to tribe and the subfamily’s largest tribe, Athysanini. The difficulty in the classification of

these genera appears to be due to a rapid radiation of deltocephaline leafhoppers that left very little

phylogenetic signal in morphological and molecular characters (Zahniser & Dietrich 2010) among a

morphologically diverse group of deltocephalines. They possess the typical Y-shaped connective of the

Fig. 2. Number of taxa described 1758 to present, summed over 5-year intervals. X-axis indicates the

year; Y-axis indicates the number of valid (blue) and invalid (red) taxa described for the previous 5

years, inclusive of the given year.

European Journal of Taxonomy 45: 1-211 (2013)

male genitalia but do not possess any of the characters that define other tribes, and are consequently a

polyphyletic assemblage of genera that is difficult to make sense of as a whole. Effort was given here

to separate some groups that have previously been placed in Athysanini or were unplaced to tribe,

were recovered in the phylogenetic analyses here, and have diagnosable combinations of morphological

characters which suggest that they form a monophyletic group. Several genera here retained in Athysanini

are quite distinctive morphologically and may merit separate tribal status, given that they seem at least as

“unusual” as the genera placed in monobasic tribes by previous authors (see Remarks under Athysanini

below). However, we see little value in recognizing additional monobasic tribes until more detailed

analyses can be conducted to further elucidate relationships within the group.

31 Online Interactive Key and Searchable Database

This publication also serves to formally introduce the online searchable database and interactive key to

tribes and subtribes of Deltocephalinae. These were produced using the 31 taxonomic database software

package (Dmitriev 2006a) and are available through the first author’s homepage <http://imperialis.

inhs.illinois.edu/zahniser/> and directly at the following links: Database Search <http://imperialis.inhs.

illinois.edu/zahniser/search.asp?key=Delt&lng=En> and Interactive Key to Tribes of Deltocephalinae:

<http ://imperialis. inhs. illinois. edu/zahniser/key. asp?key=Delt&lng=En&i= 1 &keyN= 1 >.

The database includes comprehensive nomenclatural data for all taxa of Deltocephalinae (Figs 1-2).

This includes information for the 38 tribes, 18 subtribes, 923 genera, 173 subgenera, 6683 valid

species, 274 subspecies, and 1458 invalid names of the subfamily. Each valid taxon has a designated

taxon webpage that includes all of the information currently available for it in the database. All of the

illustrations included in this publication are associated with online taxon pages for tribes, genera, and

species. An additional -3800 illustrations of select genera and species are also included, with emphasis

on the genera of Athysanini. Country-level distributional data taken from the literature are included for

all species. The coverage is not exhaustive of all leafhopper literature but provides a baseline level of

information on the distribution of species and higher taxa. The interactive key allows identification of

the tribes and subtribes of Deltocephalinae using 56 morphological characters which were scored for

all family-group taxa. The 31 database offers a number of other useful functions including advanced

database searching options, automated taxon descriptions, and taxon and character comparisons. In the

taxonomic treatments below, all taxonomic names are hyperlinked to their respective online taxon pages.

Material & Methods

Taxon Sampling for Phylogenetic Analysis

The taxon sample used by Zahniser & Dietrich (2010) was expanded here to include 37 additional

exemplars of Deltocephalinae. All analyses except one supplementary analysis included 152 total taxa,

5 of which were putative outgroup taxa. The supplementary analysis included one additional taxon

(. Dorycephalus baeri Kouchakewitch, 1866) for which data were made available after branch support

analyses had been performed on the previous taxon sample. Notable additions to the dataset include

representatives of some family-group taxa (some of which are considered junior synonyms of other groups

here) that were not included previously. Their previous classifications are given here if different from

the revised classification proposed below (Results): Occinirvana eborea Evans, 1941 (Occinirvanini);

Dwightla acutipennis Linnavuori & Al-Ne’amy, 1983 (Dwightlini); Magnentius clavatus Singh-Pruthi,

1930 (Magnentiini); Anoterostemma ivanhofi Lethierry, 1876 (Limotettigini: Anoterostemmina);

Proceps acicularis Mulsant & Rey, 1855 (Scaphytopiini: Procepitina); Pseudophlepsius binotatus

Signoret, 1880 (Opsiini: Eremophlepsiina); Allygidius abbreviates Lethierry, 1878 (Athysanini:

Allygidiina); Listrophora styx Linnavuori, 1979 (Eupelicini: Listrophorina). With these additions, all

currently recognized tribes and subtribes of Deltocephalinae are represented in the data set except for

Achaeticina (Opsiini) and Aglenina (Paralimnini).

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Table 1. A list of taxa included in the study with voucher numbers and GenBank accession numbers. The classification follows the revised classification

proposed here. 'Fragments I, II, IV, and V from B. insula DeLong, 1964 and fragment III from B. ortha DeLong, 1937. fragments I-III and V from

P. compacta Osborn & Ball, 1897 and fragment IV from P. weedi Van Duzee, 1892. # Species scored for morphological data, combined with 28S and

Histone H3 data from congeneric species. * Identification updated from previous study (Zahniser & Dietrich 2010). **28S data taken from previous

study (Dietrich et al. 2001; Rakitov, unpublished); vouchers are deposited at INHS.

Tribe

Subtribe

Species

28S

Histone H3

Voucher #

Locality

Aphrodinae / Aphrodini

Aphrodes bicincta

AF304579

GUI 23794

71**

USA: Maryland

Aphrodinae / Portanini

Portanus sp. (undescribed sp.)*

AF304674

—

Ecuador: Napo

Aphrodinae / Portanini

Portamis elegans #

—

n.a.

Venezuela

Aphrodinae / Xestocephalini

Xestocephctlus desertorum

AF304619

GUI 23 892

LH33**

USA: Illinois

Aphrodinae / Xestocephalini

Xestocephalus superbus*

—

n.a.

USA: Illinois

Euacanthellinae

EuacantheUa pahistris

GU123728

GUI 23 827

DEL078

Australia: NSW

Neocoelidiinae

Chinaia sp. (undescribed sp.)*

AF304676

—

lhl04* *

Ecuador: Napo

Acinopterini

A cinopterus acuminatus

JX845484

GUI 23790

DEL141

USA: Illinois

Acostemmini

Acostemma stilleri*

GUI 23696

GUI 23791

DEL014

Madagascar: Toliara Prov.

Acostemmini

Eryapus sp.

GU123727

—

DEL081

Madagascar: Toliara Prov.

Acostemmini

IkeJibeJoha cristata

JF835026

JN177306

LH177

Madagascar: Province d’Antsiranana

Acostemmini

Iturnoria insulana

JN 177307

JN 177308

DEL 132

Madagascar: Province d’Antsiranana

Arrugadini

Arrugada affinis

GU123699

GUI 23795

HEC005

Peru: Huanuco Prov.

Arrugadini

Arrugada rugosa #

—

n.a.

Bolivia: Yungas

Athysanini

Allygidius abbreviatus

JX845485

JX845526

DEL 126

France: Prov-Alpes-Cote d’Azur

Athysanini

Anoterostemma ivanhoffi

JX845487

JX845528

DEL 130

Scotland: Kirkcudbrightshire

Athysanini

Atanus sp. (undescribed sp.)*

GU123700

GUI 23796

DEL021

Peru: Huanuco Prov.

Athysanini

Athysanus argentarius

GU123701

GUI 23797

DEL044

USA: Illinois

Athysanini

Brazosa picturella

GUI 23709

GUI 23806

DEL006

Brazil: Goias

Athysanini

Caranavia separata*

GU123710

GUI 23807

DEL073

Peru: Junin Prov.

Athysanini

C err ill us sp.

GU123711

GU123808

DEL058

Peru: San Martin Prov.

Athysanini

Chimaerotettix ochrescens

JX845489

JX845530

DEL020

Ecuador: Orellana

Athysanini

Colladomis lineatus

GU123718

GU123815

DEL052

USA: California

Athysanini

Colladomis clitellarius #

—

n.a.

USA: Illinois

Athysanini

Dagama forcipata

GUI 23720

GUI 23817

DEL055

South Africa: W Cape Prov.

Athysanini

EusceJis seriphidii

GUI 23729

GU123830

DEL070

Kyrgyzstan: Dzhalal-abad

Athysanini

Eusceloidea nitida

JX845494

—

DEL060

Peru: San Martin Prov.

Athysanini

Eusama amanda

AF304590

GUI 23829

77h**

Mexico: Durango

Athysanini

Eutettix pictus

GU123730

GU123831

DEL 100

USA: Illinois

Athysanini

Idioceromimus delector

GUI 23 740

GUI 23844

DEL065

Brazil: Amazonas

Athysanini

Loralia sp.

GU123746

GUI 23851

DEL 102

Australia: South Australia

Athysanini

Napo sp. (undescribed sp.)*

GU123751

GU123856

DEL061

Peru: San Martin Prov.

European Journal of Taxonomy 45: 1-211 (2013)

Tribe

Subtribe

Species

28S

Histone H3

Voucher #

Locality

Athysanini

Nesothamnus sanguineus

GU123755

GUI 23860

DEL072

Ecuador: Orellana

Athysanini

Neohegira breviceps*

GUI 23 753

GU123858

DEL077

Peru: Huanuco Prov.

Athysanini

Neohegira sp. 075

GU123786

GUI 23891

DEL075

Peru: Huanuco Prov.

Athysanini

Orient us sp.

GU123757

GUI 23862

DEL022

Taiwan: Taipei Co.

Athysanini

Orientus ishhidae *

—

n.a.

USA: Illinois

Athysanini

Pachytettix sp. (undescribed sp.)*

GUI 23761

GUI 23 865

DEL037

Peru: Huanuco Prov.

Athysanini

Platymetopius obsoletus

GU123771

GU123875

DEL013

Kyrgyzstan: Dzhalal-abad

Athysanini

Renonus rubraviridis

JX845524

JX845552

DEL 127

Mexico: Jalisco

Athysanini

Thamnotettix confinis

GU123783

GU123888

Del**

USA: Colorado

Athysanini

Thamnotetiix sp . #

—

n.a.

USA: California

Athysanini

Twiningia pellucida *

GU123785

GUI 23890

DEL053

USA: California

Athysanini

Yungasia bidentata

GU123787

GUI 23893

DEL074

Peru: Huanuco Prov.

Bahitini

Bahita sp. (undescribed sp.)*

GUI 23702

GU123798

DEL009

Peru: Huanuco Prov.

Bahitini

Corny a sp.*

GUI 23 703

GUI 23799

DEL099

Peru: Huanuco Prov.

Bahitini

Kinrentius sp.

JX845523

JX845549

LH157

Peru: Junin Prov.

Bahitini

Menosoma sp. ca. elegans

GU123749

GU123854

DEL101

Peru: Huanuco Prov.

Bahitini

Oxycephalotettix tiputini

GU123759

—

DEL018

Ecuador: Orellana

Bahitini

Taperinha adspersa

GU123780

GU123885

DEL032

Peru: Huanuco Prov.

Bonaspeiini

Bonaspeia eriocephaia

JX845521

GUI 23804

DEL049

South Africa: W Cape Prov.

Bonaspeiini

Cents goudanus

GU123712

GUI 23 809

DEL050

South Africa: W Cape Prov.

Bonaspeiini

Renosteria waverena

GUI 23 772

GU123878

DEL048

South Africa: W Cape Prov.

Chiasmini

Chiasmus varicolor *

GU123713

GU123810

Ah4**

Australia: NSW

Chiasmini

Chiasmus varicolor #

—

n.a.

Australia: NSW

Chiasmini

Doratura stylaia

AF304589

GUI 23821

LH19**

USA: Maryland

Chiasmini

Driotura gammaroides

JX845492

JX845533

LH96

USA

Chiasmini

Gurawa minorcephala

JX845495

JX856131

CHI066

Thailand: Chiang Mai

Chiasmini

Listrophora siyx

JX845500

JX845539

DEL138

South Africa: Mpumalanga

Chiasmini

Nephotettix modulatus

GU123754

GU123859

CHI007

Madagascar: Toliara Prov.

Chiasmini

Nephotettix sp . #

—

n.a.

Sri Lanka

Chiasmini

Protochiasmus mysticus *

GU123708

GU123805

DEL035

Brazil: Goias

Cicadulini

Cicadula quadrinotata

GU123717

GU123813

DEL 106

Kyrgyzstan

Cicadulini

Cicadula melanogaster #

—

n.a.

USA: Illinois

Cicadulini

Elymana acuma

GU123726

GUI 23 826

DEL092

USA: Illinois

Cicadulini

Proceps acicularis

JX845511

JX845550

DEL110

France: Prov-Alpes-Cote d’Azur

Cicadulini

Stenometopiellus sigillatus

JX845515

—

DEL064

Kyrgyzstan: Chuy

Cochlorhinini

Ball ana insula' 1

AF304582 - AF304580

—

LH52**

USA: California

Cochlorhinini

Ball ana ortha'

AF304581

GU123801

LH17**

USA: Wyoming

Cochlorhinini

Cochlorhinus pluto

AF304586

GUI 23814

LH08**

USA: California

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Tribe

Subtribe

Species

28S

Histone H3

Voucher #

Locality

Deltocephalini

"DeltocephaAUS " (undesc. gen. & sp.)*

GUI 23722

GUI 23820

DEL098

Australia

Deltocephalini

Deltocephalus sp.

GUI 23721

GUI 23819

Del8**

USA

Deltocephalini

DeltocephaJus balli #

—

n.a.

USA: Illinois

Deltocephalini

Paramesodes sp. (undesc. sp.)*

GUI 23764

GUI 23868

DEL026

Taiwan: Taichung Co.

Deltocephalini

Polyamia compacta 1

AF304609 - AF304607

GU123876

113**

USA: Colorado

Deltocephalini

Polycimia weedi 2

AF304608

—

LH90**

USA: Illinois

Dorycephalini

Dorycephctlus baeri

JX845491

JX845532

HEC010

Kyrgyzstan: Naryn

Drabescini

Drabescina

Drabescus sp.

GUI 23724

GUI 23824

DEL028

Taiwan: Taipei Co.

Drabescini

Drabescina

Drabescus sp . #

—

n.a.

Nigeria: Ife Ife

Drabescini

Paraboloponina

Bhatia satsumensis

GU123706

GU123803

DEL030

Taiwan: Taipei Co.

Drabescini

Paraboloponina

Parabolopona guttata*

GUI 23762

GUI 23866

DEL029

Taiwan: Nantou Co.

Drakensbergenini

Drakensbergena retrospina

GU123725

GUI 23825

DEL051

South Africa: KZN Prov.

Eupelicini

Eupelicina

Eupelix cuspidata

AF304644

GUI 23 828

lhl18**

Kyrgyzstan: Chuy

Eupelicini

Paradorydiina

Paradorydium paradoxum *

AF304637

GU123877

lhl16**

Kyrgyzstan: Dzhalal-abad

Eupelicini

Paradorydiina

Chloropelix canariensis*

GU123715

GU123811

DEL071

Portugal: Madeira Isl.

Faltalini

HecuUus bracteatus

GU123737

GUI 23841

HEC011

USA: New Mexico

Faltalini

Kranierana junina

GUI 23 743

GUI 23 848

HEC004

Peru: Junin Prov.

Faltalini

Tenucephalus sp. (undesc. sp.)*

GU123781

GU123886

HEC001

Brazil: Goias

Fieberiellini

Fieberiella florii

AF304594

GU123834

LH23**

USA: Illinois

Goniagnathini

Goniagnathus guttidinervis

GU123736

GU123838

DEL002

Kyrgyzstan: Dzhalal-abad

Goniagnathini

Goniagnathus rugidosus #

—

n.a.

Kyrgyzstan: Issyk-Kul

Hecalini

Glossocratina

Glossocratus afzeJii *

GU123735

GU123837

HEC009

Madagascar: Toamasina Prov.

Hecalini

Hecalina

Attemiipyga vanduzei

AF304653

GUI 23822

De26

USA: Illinois

Hecalini

Hecalina

Hecalus viridis

AF304596

GUI 23 840

De27**

USA: Colorado

Hypacostemmini

Hypacostemma viridissima *

GU123739

GUI 23843

DEL094

South Africa: KZN Prov.

Koebeliini

Grypotina

Grypotes puncticollis #

—

n.a.

USA: Pennsylvania

Koebeliini

Grypotina

Grypotellus staurus *

AF304651

AF304652

GUI 23839

De33**

Greece: Delphi

Koebeliini

Koebeliina

Koebelia grossa

AF304599

GUI 23846

LH56**

USA: California

Limotettigini

Limotettix striola

GUI 23 745

GU123850

DEL004

Kyrgyzstan: Osh

Limotettigini

Limotettix (Scleroracus) decumanus #

—

n.a.

Kyrgyzstan: Dzhalal-abad

Luheriini

Luheria con strict a

JX845502

GU123852

DEL 105

Argentina: Salta

Macrostelini

Balclutha neglecta

GU123704

GUI 23800

De21**

USA

Macrostelini

Balclutha punctata #

—

n.a.

USA: Illinois

Macrostelini

Dal bul us gel bus

AF304587

GU123818

D7**

USA: OH (OSU lab colony)

Macrostelini

Evinus peri

GUI 23731

GUI 23832

HEC012

Kyrgyzstan

Macrostelini

Macrosteles quadrilineatus *

GU123748

GU123853

De22

USA

European Journal of Taxonomy 45: 1-211 (2013)

Tribe

Subtribe

Species

28S

Histone H3

Voucher #

Locality

Macrostelini

Macrosteles quadrilineatus #

—

n.a.

USA: Illinois

Magnentiini

Magnentiiis clavatus

JX845503

JX845541

LH199

India: Karnataka

Mukariini

Agrica arisana

GU123779

GU123884

DEL023

Taiwan: Nantou Co.

Mukariini

Mukaria maculata

GUI 23750

GU123855

DEL024

Taiwan: Changhua Co.

Mukariini

Mukaria sp . #

—

n.a.

India: Assam

Mukariini

“Mukariini 146” (undesc. gen. & sp.)

JX845505

JX845543

DEL 146

China: Shaanxi Prov.

Mukariini

Scaphotettix viridis

GUI 23774

GU123879

DEL068

Taiwan: Tainan Co.

Occinirvanini

Occinirvana eborea

JX845507

JX845545

DEL 143

Australia: W. Aust.

Opsiini

Circuliferina

Neoaliturus carbonarius

GU123752

GU123857

DEL012

Kyrgyzstan: Dzhalal-abad

Opsiini

Eremophlepsiina

Pseudophlepsius binotatus

JX845512

JX845551

DEL 125

Kyrgyzstan: Issyk-Kul

Opsiini

Opsiina

Hishimonus phycitis

GU123738

GUI 23842

DEL031

Taiwan: Taipei Co.

Opsiini

Opsima

Japananus hyalinus

JX845499

JX845538

DEL 129

USA: Illinois

Opsiini

Opsiina

Nesophrosyne maritima

JX845506

JX845544

DEL 109

USA: Elawaii

Opsiini

Opsiina

Opsins versicolor*

GUI 23 756

GUI 23861

DEL043

Israel: Negev

Opsiini

Opsiina

Opsins stactogalns #

—

n.a.

USA: Illinois

Opsiini

Opsiina

Orosins oriental is

JX845509

JX845547

Australia: New South Wales

Opsiini

“Zambia1220ps” undescribed

genus near Libengaia

JX845520

JX845560

DEL 122

Zambia: Northwest Prov.

Paralimnini

Paralimnina

Aflexia rubranura

GUI 23698

GUI 23793

Del6**

USA

Paralimnini

Paralimnina

Flexamia areolata

GU123733

GU123835

De38**

USA

Paralimnini

Paralimnina

Laevicephalus monticola

GU123744

GUI 23849

60b**

USA

Paralimnini

Paralimnina

Laevicephalns melsheimerii #

—

n.a.

USA: Illinois

Paralimnini

Paralimnina

Paralimnus angnsticeps

GU123763

GU123867

DEL001

Kyrgyzstan: Issyk-Kul

Pendarini

Bandaromimus parvicanda

GUI 23705

GUI 23 802

DEL076

Peru: Huanuco Prov.

Pendarini

Chlorotettix galbanatns #

—

n.a.

USA: Illinois

Pendarini

Chlorotettix rngicollis

GU123716

GU123812

DEL042

USA: Illinois

Pendarini

Chlorotettix unicolor

GU123714

—

138**

USA

Pendarini

Copididonus hyalinipennis

GU123719

GU123816

DEL007

Brazil: Goias

Pendarini

Dorydiella kansana*

GUI 23723

GUI 23823

DEL003

USA: Nebraska

Pendarini

Paraphlepsins nebnlosns

GU123765

GUI 23869

DEL045

USA: Illinois

Pendarini

Paraphlepsins irroratns #

—

n.a.

USA: Illinois

Pendarini

Tropicamis chiapasns

GU123784

GU123889

89a**

Mexico

Pendarini

Tropicanns flectns

JX845517

JX845557

DEL 131

Mexico: Chiapas

Penthimiini

Jafarjaveti

JX845498

JX845537

DEL 113

Togo: Sotouboua Region

Penthimiini

Penthimia americana

AF304606

GU123870

LH34

USA: Illinois

Penthimiini

Penthimidia eximia

JX845510

JX845548

DEL 148

Rep. of Congo: Iboubikro

Penthimiini

Penthimiola bella*

GU123766

GUI 23871

DEL080

Madagascar: Toamasina Prov.

Phlepsiini

Excnltanns conns

GU123732

GUI 23 833

DEL005

Mexico: Chiapas

Phlepsiini

Phlepsins intricatus

GU123768

GU123873

DEL017

Kyrgyzstan: Dzhalal-abad

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Tribe

Subtribe

Species

28S

Histone H3

Voucher #

Locality

Phlepsiini

Korana rorulenta

GUI 23742

GUI 23847

DEL095

Burkina Faso: Yako

Scaphoideini

Anoplotettix fuscovenosus

JX845486

JX845527

DEL 147

France: Provence-Alpes-Cote d’Azur

Scaphoideini

Loipothea sp. (undescribed sp.)

JX845501

JX845540

DEL 115

Zambia: Northwest Prov.

Scaphoideini

Mimotettix alboguttulatus

JX845504

JX845542

DEL 117

Zambia: Northwest Prov.

Scaphoideini

Osbornellus sp. (undescribed sp. near 0.

linnavuori DeLong)*

GU123758

GU123863

DEL033

Peru: Huanuco Prov.

Scaphoideini

Phlogotettix cyclops

GUI 23769

GU123874

DEL027

Taiwan: Ilan Prov.

Scaphoideini

Omanana sp. (undescribed sp.)

JX845508

JX845546

DEL 128

Mexico: Jalisco

Scaphoideini

Scaphoideus alboguttatus*

GUI 23773

—

DEL040

Taiwan: Nantou Co.

Scaphoideini

Scaphoideus sp. SA

GU123775

—

DEL063

South Africa: W Cape Prov.

Scaphoideini

Scaphoideus oniani

JX845513

JX845553

DEL 120

Zambia: Northwest Prov.

Scaphoideini

Scaphoidophyes sp. near pyrus Barnett &

Frey tag

JX845525

JX845554

DEL 124

Zambia: Copperbelt Prov.

Scaphytopiini

Scaphytopius frontalis

JX845514

JX845555

DEL 116

USA: Maryland

Scaphytopiini

Scaphytopius vaccinium

GU123776

GU123880

lh09b**

USA

Scaphytopiini

Scaphytopius acutus*

—

n.a.

USA: Illinois

Selenocephalini

Adamina

Adama (Krisnella) elongata

GUI 23694

GU123788

DEL083

Zambia: Copperbelt Prov.

Selenocephalini

Adamina

Adama (Paracostemma) sp.

GUI 23697

GUI 23792

DEL015

Rwanda: Nyungwe Forest

Selenocephalini

Adamina

Adama (Paracostemma) sp . #

—

n.a.

Central African Republic: Boukoko

Selenocephalini

Dwightlina

Dwightla acutipennis

JX845493

JX845534

DELHI

Cameroon: Southwest Region

Selenocephalini

Ianeirina

Abimwa knighti *

GUI 23695

GUI 23789

DEL 104

Zambia: Northwestern Prov.

Selenocephalini

laneirina

Abimwa sp . #

—

n.a.

South Africa: KZN Prov.

Selenocephalini

Selenocephalina

Citorus stipes

JX845490

JX845531

DEL 144

Zambia: Northwest Prov.

Selenocephalini

Selenocephalina

Selenocephalus deserticola*

GU123777

GU123881

DEL041

Israel: Negvev

Stegelytrini

Pachymetopius decoratus

GUI 23 760

GUI 23864

DEL025

Taiwan: Taichung Co.

Stegelytrini

Placidellus sp.

GU123770

—

Thailand

Stenometopimi

Kinonia elongata

GU123741

GUI 23845

75g**

USA

Stenometopiini

Stirellus catalinus

AF304614

GUI 23882

82k**

Mexico: Durango

Stenometopiini

Stirellus bicolor

—

n.a.

USA: Illinois

Tetartostylini

Tetartostylus parabolatus

GUI 23 782

GU123887

DEL047

South Africa: W Cape Prov.

Vartini

Stymphalus rubrolineatus

GU123778

GU123883

DEL062

South Africa: W Cape Prov.

—

“Peru059” undescribed gen.sp.

GU123767

GU123872

DEL059

Peru: San Martin Prov.

—

“Cameroon 142” undescr. gen. & sp.

JX845488

JX845529

DEL 142

Cameroon: Southwest Region

—

“Ghanall4” undescr. gen. & sp.

JX845522

JX845535

DEL114

Ghana: Western Region

—

“Thailand 112” undescr. gen. & sp.

JX845516

JX845556

DEL112

Thailand: Chaiyaphum

—

“Zambiall8” undescr. gen. & sp.

JX845518

JX845558

DEL 118

Zambia: Northwest Prov.

—

“Zambia 119” near Phlogothamnus

JX845519

JX845559

DEL119

Zambia: Northwest Prov.

—

“Zambia 121” near Houtbayana / Acacimenus

JX845496

JX845536

DEL121

Zambia: Copperbelt Prov.

European Journal of Taxonomy 45: 1-211 (2013)

Taxa included in the analyses are listed in Table 1 which follows the revised classification provided here.

GenBank accession numbers, voucher numbers, and locations are provided for all specimens included.

Morphological and Molecular Data and Phylogenetic Analyses

Morphological characters were the same as those used by Zahniser & Dietrich (2010) but with the

addition of two characters (9 and 75). The sequence of characters given here is different from that of

Zahniser & Dietrich (2010). Character names and states are listed in Appendix 1 and the data matrix is

given in Appendix 2.

Methods for PCR amplification and sequence editing follow Zahniser & Dietrich (2010). An alternative

PCR primer for the D2 region of 28S was used for some taxa (see Zahniser & Nielson 2012). 28S rDNA

sequences were aligned using the MUSCTE algorithm (Edgar 2004) implemented in MEGA5 (Tamura

et al. 2011). The MUSCLE alignment was then slightly modified by hand in BioEdit 7.0.0 (Hall 1999).

The final 28S sequence alignment is available from the first author by request. Based on the alignment,

a binary set of indel (insertion/deletion) characters was coded by hand. Alignment of the Histone H3

sequences did not require alignment algorithms.

Maximum parsimony (MP) analyses were run in PA UP* 4.0bl0 (Swofford 1998). All characters were

treated as unordered except characters 13, 14, and 113, which were treated as ordered, and character 6,

which utilized a user-defined step matrix. 10,000 search replicates were performed with nchuck=5 and

chuckscore=l, and the resulting set of trees were subsequently used as the starting tree set to search for

all most parsimonious trees. 500 bootstrap (BS) replicates were performed with 100 pseudoreplicates per

bootstrap replicate. Analyses were run to compute the decay index (DI = Bremer support) (Bremer 1988)

for each node of the strict consensus of all most parsimonious (MP) trees. A search method similar to

the original tree search was used to find MP trees with node constraints but with 1,000 initial search

replicates due to computational time constraints. Total and average Bremer support for separate data

partitions were calculated for each node.

Maximum likelihood (ML) analyses were run in GARLI 2.0 (Zwickl 2006). Separate partitions were

used for the 28S, indel, each codon position of Histone H3, and morphology. Molecular models were

explored for each molecular data partition using MrModeltest 2.2 (Nylander 2004). Results of the model

tests for all partitions suggested the GTR+I+G model except for the second codon position of Histone

H3 for which the Jukes-Cantor model was selected. These selected models were implemented in ML

analyses. The morphology and indel data sets were treated as “standard variable” data. The setting for

the number of generations without significant change to terminate a replicate (genthreshfortopoterm)

was set to 10,000. 100 search replicates were performed in the initial search. 100 bootstrap replicates

were performed with 10 pseudoreplicates per bootstrap replicate. A separate ML analysis was performed

with data for an additional taxon, Dorycephalus baeri Kouchakewitch, 1866. The analysis (50 search

replicates) was performed without the morphological data set to examine the effect of excluding it in ML

analysis; branch support was not assessed for this data set.

Bayesian analyses were run in MrBayes 3.2 (Ronquist et al. 2012). The GTR+I+G model was imple¬

mented for all molecular data partitions (28S and each codon position of Histone H3) as suggested by

Huelsenbeck & Rannala (2004) given the results of the model tests. 15 million generations were run

with 2 runs and 4 chains per run. Trees were sampled every 1000 generations.

Taxonomic Treatments

The revised classification of the tribes provided below is based on the phylogenetic analyses presented

here and on comparative morphological study of numerous genera of Deltocephalinae, representatives

of which were borrowed from major museums. Although the ultimate goal is a classification that

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

reflects phylogenetic relationships, this is difficult at present because many relationships remain poorly

resolved by available data, and representatives of many genera have not been included in phylogenetic

analyses; therefore, we chose a fairly conservative approach, in some cases retaining previously

recognized groups even if they were not always recovered as monophyletic, pending further study. In

general, changes to the higher classification (i.e., recognition of new tribes or re-definition of previously

described tribes) are proposed only when supported at least in part by the phylogenetic results.

Emphasis was also placed on the ability to diagnose any such groups with morphological characters.

Thus, we did not establish tribes or subtribes for several groups of genera (clades) that received strong

branch support in the phylogenetic results because we were unable to find diagnostic morphological

characters to facilitate their identification. We hope that our results will stimulate further research

not only to confirm their monophyly but also to determine whether such groups are diagnosable by

as yet unrecognized morphological features. Along with specimens housed at INHS, specimens were

borrowed from AMNH, OSUC, NMNH, BMNH, INSRB, RMCA, and MNHN (see acronyms below)

with special attention to genera which were placed in the large tribe Athysanini, were unplaced to

tribe in Deltocephalinae, or were perceived to be of questionable placement in their respective tribes.

Representatives of most genera of Athysanini and genera unplaced to tribe in Deltocephalinae were

examined for this study.

Tribe and subtribe descriptions were modified from the automated descriptions generated by the 31

database based on the characters that were scored for the interactive key to tribes and subtribes. In

the descriptions of size, “small” = 2 to 4 mm, “medium” = 4 to 8 mm, and “large” = more than 8 mm

The Geography and ecology sections are limited to notes on their distribution patterns in the major

zoogeographical regions, their known host plants or general host plant use patterns, ecosystem types

they are found in, and important species that are vectors of phytopathogenic diseases. Information

on vector species was taken from Nielson (1968) and Weintraub & Beanland (2006). The Selected

references were chosen with emphasis on publications that are relatively recent (mostly within the past

50 years) and provide a broad perspective on the group, describe a significant number of new taxa,

provide descriptions of the higher taxa, or are comprehensive treatments of the group over a relatively

large geographical area.

One issue regarding the validity and authorship of some family group names must be addressed. A

published abstract (Oman 1943) of RW. Oman’s Ph.D. thesis (Oman 1941) included the first validly

published instances of names for the deltocephaline family-group taxa Acinopterini, Cochlorhinini,

Dorycephalini, Mesamiini, Phlepsiini, Scaphoideini, and Scaphytopiini. Because Oman’s published

abstract did not include descriptions of these taxa or indicate which genera should belong in them,

the names were not available when proposed according to ICZN Article 13.1. Although the published

abstract referred to the taxon descriptions in the thesis, the thesis itself is not considered officially

published according the ICZN (1999: Art. 8) which Article 13.1.2 requires. Nevertheless, some of the

Oman (1943) names were taken by subsequent authors as valid (e.g., Oman 1949; Oman et al. 1990),

and according to ICZN Article 13.2.1 those taken as valid should be treated as available from their

original publication date. This article provides that family-group taxa first described after 1930 but

before 1961 which do not satisfy the provisions of Article 13.1 are available if they were used as valid

before 2000 and not explicitly rejected under Article 13. Zahniser & Dietrich (2010) overlooked Article

13.2.1 and incorrectly treated the Oman (1943) names as unavailable nomina nuda. However, according

to ICZN Article 13.2.1, the only family-group names published by Oman (1943) that are invalid (and

unavailable) are Mesamiini and Phlepsiini because these names were not treated as valid in subsequent

works published prior to 2000.

Digital photographs of the habitus and face were taken with a Nikon D lx digital SLR camera or a Canon

EOS Mark 5 DII configured with lenses by Microptics, Digital Lab XLT system. Digital photographs of

European Journal of Taxonomy 45: 1-211 (2013)

the male and female genitalia were taken with a Q Imaging Micropublisher 3.3 digital camera mounted

on an Olympus BX41 compound microscope. Multiple images were combined using the CombineZP

software program (Hadley 2010). Photographs and stacked images were modified with Adobe Photoshop

CS5. The combined photo and line drawing of the faces were produced in Adobe Illustrator CS5. All

scale bars =1.0 mm.

For the taxon illustrations, the following standard structures and views are as follows: A, dorsal habitus;

B, lateral habitus; C, face; D, lateral view of male pygofer and terminal segments; E, dorsal view of

male pygofer and terminal segments; F, ventral view of subgenital plates and valve; G, ventral view of

connective and style; H, lateral view of aedeagus; I, caudal or ventral view of aedeagus. Additional notes

and additional structures or illustrations are given in the figure captions as needed.

Acronyms for institutions used here are as follows:

AMNH

IRSNB

INHS

MNHN

BMNH

NMNH

NMNS

osuc

RMCA

American Museum of Natural History; New York, NY, USA.

Institut Royal des Sciences Naturelles de Belgique; Brussels, Belgium.

Illinois Natural History Survey; Champaign, IL, USA.

Museum National d’Histoire Naturelle; Paris, France.

The Natural History Museum; London, United Kingdom.

The National Museum of Natural History; Washington D.C., USA.

National Museum of Natural Science; Taichung, Taiwan.

The C.A. Triplehorn Insect Collection at the Ohio State University; Columbus, OH, USA.

Musee Royal de FAfrique Centrale; Tervuren, Belgium.

Results

Phylogenetic Analyses

A total of 803 parsimony-informative characters were included in the final data set. The entire aligned

28S rDNA data set included 3437 positions, and with segment II (see Dietrich et al. 2001) and the

final 22 bp excluded, the data set included 2908 characters; 500 of these were parsimony-informative,

402 were variable and parsimony-uninformative, and 2006 were constant. The indel data set included

60 characters, all of which were parsimony-informative. A total of 353 bp of the Histone H3 nuclear

protein coding gene were included; 122 characters were parsimony-informative, 8 were variable and

parsimony-uninformative, and 223 characters were constant. All 121 morphological characters were

parsimony-informative.

The parsimony analysis yielded 224 MP trees. The strict consensus is provided in Figs 3a and 3b along

with bootstrap scores and total and partitioned Bremer support values.

The ML tree with the best likelihood score was found in replicate 5 (of 100) and had a score of

-45315.2484. Best scores from 16 other replicates were within 4 likelihood points of the best scoring

tree, suggesting that the number of replicates performed here was sufficient in finding the most likely

or nearly most likely trees. The tree with the best likelihood score is shown in Fig. 5 along with ML

bootstrap scores. To allow for more space in the figure, the outgroup branches are shown separately in

Fig. 4. The best tree in the analysis including Dorycephalus and excluding morphological data is shown

in Fig. 6.

The 2 runs in the Bayesian analysis converged after 10 million generations, and the trees obtained prior

to convergence were discarded as burnin. Convergence was determined by the random distribution of

likelihood scores in a plot of the scores from the post-bumin generations. The ingroup branches of the

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Bayesian consensus tree are provided in Fig. 7, and the outgroup branching patterns and branch lengths

are provided in Fig. 4, both with Bayesian posterior probability (PP) scores given below the branches.

History of taxonomic descriptions in Deltocephalinae

The compilation of the nomenclatural database allowed us to examine of some historical patterns in

the taxonomy of Deltocephalinae, some of which are summarized in Figs 1 and 2. The first described

deltocephaline was Psammotettix striatus (Finnaeus, 1758), originally placed in the genus Cicada.

■Aprhodes

■ Euacanthella

■Xes tocephalus

Portanus

1 Chinaia

‘ Ikelibeloha

Iturnoria

■Acinopterus

-Acostemma

■Eryapus

■ Pachymetopius

■ Placidellus

■ Gonlagnathus

■Orientus

• Fieberiella

■Luheria

-Magnentius

■Penthimiola

• Penthimia

■ Penthimidia

■ Loralia

■ Ocdnirvana

■ Hypacostemma

■Cameroon 142

■Ghana 114

■Jafar

1 Cerus

■ Bonaspeia

■ Renosteria

■ Dagama

• Abimwa

■ Adama (Krisn.)

■Adama (Para.)

• Dwightla

■ Citorus

■ Selenocephalus

• Korana

■Excultanus

• Phlepsius

-Zambia 121

■Zambia 118

■Loipothea

■Omanana

■Osbornellus

■ Bhatia

■ Drabescus

■ Parabolopona

■ Scaphoidophyes

■ Phlogotettix

■Zambia 119

■Anoplotettix

■Mimotettix

■Scaphoideus om.

■Scaphoideus alb.

■ ScaphoideusSA

Outgroup

Acostemmini (part)

Acinopterini

Acostemmini (part)

Stegelytrini

Goniagnathini

Athysanini (part)

Fieberiellint

Luheriini

Magnentiini

Penthimiini (part)

Athysanini (part)

Occinirvanini

Hypacostemmini

undescribed

Penthimiini (part)

Bonaspeiini

Athysanini (part)

Selenocephalini

Phlepsiini

undescribed

Scaphoideini (part)

Drabescini

Scaphoideini (part)

Fig. 3a. Top portion of the strict consensus of224 MP trees. Numbers above branches are MP bootstrap

/ total Bremer support values. Numbers below branches are partitioned Bremer support values for 28S /

indel / Histone H3 / morphology data sets. The revised classification is given to the right.

European Journal of Taxonomy 45: 1-211 (2013)

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1.6/-0.3/1.4/1.3

- 2 . 2 /

57/3

-1.5/-0.1/-0.3/4.9

-0.5/0,7/4.2/-0.4

59/3

1.0/9.5/-5.2

99/9

51/3

7.4/0.9/2.6/-2.0

55/4

5.9/2.073.27-8.1

57/4

-76

9.2/-0.6/3.6/-6.3

-13

-0 5/-1.1/-1.3Z5 9

-0.2/0/10.6/-6.4

-13

4.9/0.6/2.5/-4.9

80/9

5.17-1.0/1.273.7

98/13

6.8/-0.4/-0.4Z7.Q

\ 100/45

23.6/0,5/12.3/8.6

100/15

■72

1.5/-1.1/0,5/1.0

8.0/-1.1/4.5/3.5

59/4

-1 8/-0 1/-8 2714.0

97/11

8.9/-0.1/-3.9Z6.1

■71

4.4/0.6/2.0/-6.0

94/7 r

2 5/-n 17D 7/3 a l 100/23

10,2/2,9/6.2/3.7

Japananus

Pseudophlepsius

Opsius

Hishimonus

Zambia 122

Nesotfiomnus

/d/oeeroznimus

Scaphytopius fro.

Scaphytopius vac.

Yungasia

Neohegira 075

Neohegira brev.

Menosozno

Taperinha

Oxycephalotettix

Kinrentius

Bahita

Caruya

Banaaromimus

Cobididonus

Chlorotettix uni.

Chlorotettix rug.

Dorydiella

Paraphlepsius

Tropicanus chi.

Tropicanus fie.

Peru059

Cerrillus

Chimaerotettix

Eutettix

Eusceloidea

Napo

Renonus

Atanus

Caranavia

Pachytettix

Brazosa

Colladonus

Eusama

Platymetopius

Twinlngia

Proceps

Elymana

Cicadula

Stenometopiellus

Arrugada

Glossocratus

Attenuipyga

Hecalus

Dalbulus

Balclutha

Macrosteles

Neoaliturus

Nesophrosyne

Orosius

Limotettix

Allygidius

Thamnotettix

Euscelis

Athysanus

Anoterostemma

Grypotes

Koebelia

Ballana

Cochlorhinus

Agrica

Mukariini 146

Mukoria

Scaphotettix

Stymphalus

Thailand 112

Tenucephalus

Hecullus

Kramerana

Tetartostylus

Laevicepnalus

Paralimnus

Aflexia

Flexamia

Deltocephalus

Polyamia

DeltocephAUS

Paramesodes

Evinus

Protochiasmus

Drakensbergena

Eubelix

Chloropelix

Paradorydium

Kinonia

Stirellus

Nephotettix

Doratura

Driotura

Chiasmus

Gurawa

Listrophora

Opsiini (part)

Athysanini (part)

| Scaphytopiini

Athysanini (part)

Bahitini

Pendarini

Athysanini (part)

Cicadulini

Arrugadini

Hecalini

Macrostelini (part)

Opsiini (part)

Limotettigini

Athysanini (part)

Koebeliini

Cochlorhinini

Mukariini

Vartini

undescribed

Faltalini

Tetartostyiini

Paralimnini

Deltocephalini

Macrostelini (part)

i Chiasmini (part)

1 Drakensbergenini

Eupelicini

Stenometopiini

Chiasmini (part)

Fig. 3b. Lower portion of the strict consensus of 224 MP trees. Numbers on branches are as indicated

for Fig. 3 a.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

There were very few and scant descriptions of new taxa from 1758 through the mid-19th century, most

of which were by the European workers Fabricius, Goeze, Panzer, Fallen, Germar, Zetterstedt, Herrich-

Schaffer, Burmeister, Curtis, Rambur and Boheman, and by the American, Thomas Say. The years 1850

to 1889 saw an increase in taxonomic descriptions, led again by European workers including Walker,

Mulsant, Rey, Stal, Fieber, Kirschbaum, Motschulsky, Signoret, Flor, Fieber, Provancher, Lethierry and

Signoret, and North and South American workers including Fitch, Uhler and Berg.

A marked increase in the description rate began in 1890 and has continued to the present day (Figs

1-2). A generation of late 19th century American workers including Van Duzee, Osborn, Ball, Gillette,

Baker and Kirlcaldy began to describe many New World Deltocephalinae. Other productive workers at

the time included Matsumura, Melichar and Distant, who produced significant works on the fauna of

India (e.g., Distant 1908, 1918). Dwight M. DeFong began publishing on Deltocephalinae in 1916 and

continued until he passed away in 1984, leaving behind a rich 68-year legacy of work on the subfamily.

He is distinguished as the most prolific author of species of Deltocephalinae, having named (usually

as first or sole author) 833 valid species. DeFong’s publication on Ballana (DeFong, 1937), in which

52 new species were described, contributed to high production during the years 1937-1941 (Fig. 2),

and his publication on the leafhoppers of Illinois (DeFong, 1948) is still useful for identification of the

North American fauna. The high productivity was also the result of the work by other pre-1950 20th

century workers including Naude, Fawson, Findberg, Haupt, Kusnezov, Sleesman, Beamer, Ribaut,

Ossiannilsson, Singh-Pruthi, Merino, Zachvatkin, Evans, Oman and Hepner. Rauno E. Finnavuori,

the second most prolific describer of Deltocephalinae, began publishing on the group in 1951, and

his high-quality revisions of regional faunas are still used today. Finnavuori’s influence, especially

beginning in 1959 with his revision of the Neotropical Deltocephalinae (Finnavuori 1959), along with

that of DeFong and other authors (Dlabola, Vilbaste, Emeljanov, Heller, Kramer, Evans, Knight, Webb,

Hamilton, Cheng, Remane, Asche, Cwikla, Blocker, Hicks, Whitcomb, Theron and Davies) made the

years 1959 through 1988 the most prolific time period in the description of Deltocephalinae. This period

was followed by a precipitous decline from 1989 to 1997 (although notable authors of this time include

Meyer-Arndt, Remane, Zanol, Blocker, Viralctamath, Knight and Webb), but a resurgence over the last

15 years has been the result of the work of numerous authors including Viralctamath, Webb, Knight,

Fletcher, Zhang, Wu Dai, Wei, Duan, Zanol, Stiller, Hamilton, Emeljanov, Dmitriev, Dietrich, Rakitov,

Zahniser, Dominguez, Godoy, Fi, Renhuai Dai and Xing.

-Aprhodes

Euacanthella

F—

Porta n us

Deltocephalinae

Chinaia

■Xestocephalus

- Aprhodes

- Euacanthella

- tt - Portanus

Deltocephalinae

Chinaia

■Xestocephalus

Fig. 4. A. Outgroup relationships inferred from ML analysis. Numbers below branches are ME bootstrap

values. B. Outgroup relationships inferred from Bayesian analysis. Numbers below branches are PP

values.

European Journal of Taxonomy 45: 1-211 (2013)

-Pachymetopius

• Placidellus

Ikelibeloha

Goniagnathinii

Fieberiellinii

Occinirvanini i

Orientus !

Penthimiini (part)i

Mocroste/es

Bolclutha

Athysanini (part)

Vartini

Stenometopiini j

_ , Chloropelix

~ , .100"- Poradorvaium

Protocwasmus '

Nephotettix

Driotura

I //■ ■ Chiasmus

fiRi_ |- Gurawa

Listrophora

Stegelytrini

Acostemmini

Acinopterini

Luheriini

Magnentiini

Penthimiini (part)

Athysanini (part)

undescnibed

Hypacostemmini

Bonaspeiini

Athysanini (part)

Selenocephalini

Phlepsiini

undescribed

Scaphoideini (part)

Drabescini

Scaphoideini (part)

Athysanini (part)

Hecalini (part)

Athysanini (part)

Pendarini

Athysanini (part)

Scaphytopiini

Athysanini (part)

Opsiini

Athysanini (part)

Bahitini

Arrugadini

Cicadulini

Limotettigini

Hecalini (part)

Macrostelini

Koebeliini

Mukariini (part)

Athysanini (part)

Cochlorhinini

Mukariini (part)

undescribed

Faltalini

Tetartostylini

Paralimnini

Deltocephalini

Macrostelini (part)

Drakensbergenini

Eupelicini

Chiasmini

Fig. 5. Ingroup relationships inferred from ML analyses of combined data. Numbers below branches

are ML bootstrap values. Shortened branches are broken with 6 // 6 marks and are shown at half of their

inferred lengths.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

- Aphrades

■ Euacanthella

■ Portanus

-Xestocephalus

-Chinaia

Pachymetopius

• Placidellus

- Acostemma

— -try

Eryat>us

- Iturnorio

■ Ikelibeloha

//-

-Adnopterus

• Goniagnathus

—Luherio

■ Orientus

-Magnentius

- penthimidia

■ Fieberiella

■ Penthimiola

■ Ocdnirvana

■ Loralia

Jafar

■ Cameroon 142

■Ghana I 14

, Hypacostemma

•Cerusn „ ■

Bonaspeia

st er

Renosteria

~Adc

Adam

Abimwa

Dagama

Dwigbtla

l~^ S Selenocephalus

-Zambia 121

- -“Zambia I 18

Scaphoidopnyes

A - 7~ Korana

£xcultanus

-Ph/ebsius

Anob/oteft/x

-- Mimotettix „

lbhoideus a lb.

Scabhoideus i

Omanana

Phlogotettix

Scqphoideus063

atia

Parabolopona

Zambia 122

- Japonanus

Pseudoptlep ^^

Orosius

Attenuipyga

Eusqma

. etopius

_rScobbytopius fro.

Pachytettix Scaphytopius vac.

- Atanus

Caranqvia

•Bandaromimus

Copididonus

~7*—r— n Chlorotettix rug,

Doryd/ela

—Parophlepsius

1ropicanusxhi.

opicanus fle.

Chlorotettix uni.

-Neohegira 075

veohegi

v " Neohegira breviceps

-Yungasia r. . r

--- rr—Nesothamnus

- Menosoma

Oxycephalotettix

■ Jopeh rjpa

- Idioceromimus

Caruva

Limotettix

Jnrentius

Bahita

■ Proceps

Oca

-fc lymana . ..

Allygidius Stenomet °l },e " us

■Thamnotettix

-Anote/ostemma

Tenucephalus

■ Kramerana

■ Tetartostylus „

- - — Paralimnus

-Laevicephalus. „

— . - 1 - Aflexia

-Flexamta _ '

-Deltocebnows

=Po(yarrlio

DeltocephAuS

Chiasmus

- Eupelix

-r - Chloropelix

Paradorydium

Nephotettix

S aldutha

Macrosteles

Fig. 6. ML tree inferred from analysis of molecular data only. This tree is included to illustrate the

difference between including or excluding the morphological data from the phylogenetic analyses

(see Fig. 5 for comparison of ML-inferred tree with morphological data). This analysis is the only one

performed here to include Dorycephalus (Dorycephalini), and its position is highlighted.

European Journal of Taxonomy 45: 1-211 (2013)

.Pachymetopius

- Plocidellus

-Acostemma

-Eryapus

Ikelibeloha

Loralia

Drabescus

Parabolopona

Balclutha

Stegetytrini

Acostemmini

„ ... ..Acinopterini

Goniagnalhimj

Fiebenellinn ... ...

.. ... . i Athysamm (part)

Magnemtimii 1 ^

| Penthimiini (part)

_ . . . . lAthysanini (part)

Occinirvanini; un( jesc ri bed

Hypacostemmmn u „ d6scribed

| Penthimiini (part)

| Bonaspeiini

Athysanini (part)

Selenocephalini

Phlepsiini

undescribed

Scaphoideini

Drabescini

Limotettigini! AmJ 9 ad ' n '

' Hecalini (part)

Macrostelini

Cicadulini

Opsiini

Athysanini (part)

Bahitini

Athysanini (part)

Pendarini

Athysanini (part)

Scaphytopiini

Athysanini (part)

Koebeliini

Athysanini (part)

Mukariini (part)

Cochlorhinini

Athysanini (part ■

| Mukariini (part)

Vartinii , ,

■ undescribed

Faltalini

Tetartostylini

Paralimnini

Deltocephalini

i Macrostelini (part)

n , , . .1 Stenometopiini

Drakensbergenim i ,, .....

■Attenuipyga i Hecalini (part)

—Chhrppehx Eupelicini

Chiasmini

Listrophora

Fig. 7. Ingroup relationships inferred from Bayesian analysis of combined data. Numbers below

branches are PP values. Shortened branches are broken with 7/‘ marks and are shown at half of their

inferred lengths.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

The most productive three years were 1959 (234 taxa), 1980 (189 taxa), and 1918 (159 taxa). The first

and third most productive years were largely the result of monographic revisions by Linnavuori (1959)

and Distant (1918), while the taxa were described by many authors in 1980. In general, the percentage of

taxa described per year that are still valid (Fig. 2) has steadily increased, with the most recent two 5-year

periods reaching 96 and 99%. This could be due to a number of factors including better taxonomic

descriptions in recent years, easier access to literature, or less time since their publication (thus, less

chance to discover invalid names). The former two factors make it more likely that fewer potentially

invalid names are currently being produced than was the case in earlier generations.

Taxonomic Treatments

A complete description of the subfamily Deltocephalinae was provided by Zahniser & Dietrich (2010);

an auto-generated description is available from the hyperlinked subfamily name below and a diagnosis

of the subfamily is given below. Changes made to the classification are discussed in the Remarks section

of each taxon treatment, along with discussion of the results of the phylogenetic analyses.

The revised concepts and new descriptions of tribes rely as much as possible on multiple independent

morphological characters for their definition and diagnosis. Based on recent revisionary work and on the

results of the recent phylogenetic analyses, it has become apparent that single morphological characters

that sometimes previously defined tribes are not necessarily reliable predictors of a close relationship

between genera. Similar character states have sometimes arisen multiple times independently in

unrelated genera and are thus homoplasious. Conversely, a unique character state may have arisen once

in the common ancestor of a lineage and subsequently be shared among some or most members of a

monophyletic lineage but secondarily lost or transformed in other closely related members. The former

case is illustrated by Scaphytopiini which was previously defined based on the wide gena visible behind

the eye in dorsal view (usually in association with a narrow and produced head). The phylogenetic

analyses here suggest that the genera previously included in Scaphytopiini (based only on this character)

and included in the analyses ( Japananus , Nesothamnus, Proceps, Scaphytopius and Stymphalus) are

not closely related to one another. Thus, this character appears to be subject to homoplasy, and other

characters need to be considered in the placement of these and other genera to tribe. An example of the

latter case is illustrated in Chiasmini. This tribe was previously defined primarily by the unique hinged

aedeagus. However Zahniser (2008a, 2011) pointed out several characters of the female ovipositor that

were also helpful in defining the tribe and suggested a revised classification that included some genera

with the ovipositor characters but without the hinged aedeagus (e.g., Leofa , Gurawa). This classification

has been supported by molecular phylogenetic analyses (see below, Zahniser 2008b) and thus, in this case,

the use of multiple characters was necessary to produce a classification that is supported by morphology

and reflects phylogenetic relationships. The tribe is no longer diagnosable by a single character (i.e.

the hinged aedeagus), but the character is still unique and occurs in most, but not all, Chiasmini. Thus,

the use of multiple characters including external characters and those of the male and female genitalia

strengthens our understanding of the phylogenetic relationships, character state variability, and generic

constituency (taxonomy) of the tribe.

Deltocephalinae Dallas, 1870

Note

Because of the size and morphological diversity of this subfamily, nearly all of the characters listed

below, which are the most common states in Deltocephalinae, occur in different states in some members

of the subfamily.

European Journal of Taxonomy 45: 1-211 (2013)

Diagnosis

Small to large, usually wedge-shaped leafhoppers. Head with ocelli on the anterior margin close to eyes;

frontoclypeus not inflated, without carinae; lateral frontal sutures extending to ocelli; antennal ledges

reduced or absent; gena with a fine erect seta laterad of lateral frontal suture; gena large, mostly concealing

proepistemum. Forewing macropterous to brachypterous; if macropterous, with apices overlapping at

rest; with 3 anteapical cells; often with 1 or more crossveins between A1 and claval suture; inner apical

cell tapered distally, not extended to wing apex. Profemur AMI seta distinct; row AV with short stout

setae extending from base to 1/2 to 2/3 length of femur; intercalary row with numerous thin setae arranged

in one row. Mesotrochanter with apical posteroventral stout seta. Metafemur macrosetal formula usually

2+2+1 with penultimate pair close-set. Metatibia usually anteriorly-posteriorly compressed, ventral side

with median ridge. Male pygofer with basolateral oblique membranous cleft. Valve produced posteriorly,

lateral margins short, articulated with pygofer laterally. Subgenital plates articulated with each other and

with valve; triangular, more or less depressed or flattened; with dorsal slot or fold articulating with style.

Connective Y-shaped or ‘linear’ (with anterior arms closely appressed), rarely T-shaped (with arms

widely divergent); without anteromedial lobe or process. Style broadly bilobed basally; apophysis not

elongate. First valvula convex to relatively straight; dorsal sculpturing pattern reaching or not reaching

dorsal margin; sculpturing pattern strigate, concatenate, reticulate, imbricate, maculose, or granulose.

Second valvula with basal fused section as long as or longer than distal paired blades; with or without

median dorsal tooth; usually with small to large, regularly or irregularly shaped dorsoapical teeth on

apical 1/3 or more; teeth sometimes restricted to apical 1/4, or absent.

Included Tribes and Subtribes:

Acinopterini Oman, 1943

Acostemmini Evans, 1972

Arrugadini Linnavuori, 1965

Athysanini Van Duzee, 1892

Bahitini Zahniser & Dietrich, 2013

Bonaspeiini Zahniser & Dietrich, 2013

Chiasmini Distant, 1908

Cicadulini Van Duzee, 1892

Cochlorhinini Oman, 1943

Deltocephalini Dallas, 1870

Dorycephalini Oman, 1943

Drabescini Ishihara, 1953

Drabescina Ishihara, 1953

Paraboloponina Ishihara, 1953

Dralcensbergenini Linnavuori, 1979

Eupelicini Sahlberg, 1871

Eupelicina Sahlberg, 1871

Paradorydiina Evans, 1936

Faltalini Za hn iser & Dietrich, 2010

Fieberiellini Wagner, 1951

Goniagnathini Wagner, 1951

Hecalini Distant, 1908

Glossocratina Dmitriev, 2002

Hecalina Distant, 1908

Hypacostemmini Linnavuori & Al-Ne’amy, 1983

Koebeliini Baker, 1897

Grypotina Haupt, 1929

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Koebeliina Baker, 1897

L imotettigini B alcer, 1915

Luheriini Linnavuori, 1959

Macrostelini Kirkaldy, 1906

Magnentiini Linnavuori, 1978

Mukariini Distant, 1908

Occinirvanini Evans, 1966

Opsiini Emeljanov, 1962

AchaeticinaEmeljanov, 1962

Circuliferina Emeljanov, 1962

Eremophlepsiina Dmitriev, 2006

Opsiina Emeljanov, 1962

Paralimnini Distant, 1908

Aglenina Dmitriev, 2004

Paralimnina Distant, 1908

Pendarini Dmitriev, 2009

Penthimiini Kirschbaum, 1868

Phlepsiini Zahniser & Dietrich, 2013

Scaphoideini Oman, 1943

Scaphytopiini Oman, 1943

Selenocephalini Fieber, 1872

Adamina Linnavuori & Al-Ne’amy, 1983

Dwightlina McKamey, 2003

Ianeirina Linnavuori, 1978

Selenocephalina Fieber, 1872

Stegelytrini Baker, 1915

Stenometopiini Baker, 1923

Tetartostylini Wagner, 1951

Vart ini Zahniser & Dietrich, 2013

Key to Tribes

1. Crown sharply produced anteriorly (Fig. 48A); head narrower than pronotum; gena not incised

laterally below eye (Fig. 48C), visible behind eye in dorsal view (Fig. 48A); frontoclypeus elongate

(Fig. 48C); forewing with numerous reflexed costal veins (Fig. 48B); connective a transverse bar

with two lateral arms directed obliquely laterad, stem absent (Fig. 48G); usually with pair of long

processes articulated with connective (Fig. 48G, J);Nearctic, Neotropical.Scaphytopiini

Without the above combination of characters; crown variable; head width variable; gena incised

laterally below eye (exceptions: some Acostemmini, Opsiini, Stegelytrini, Scaphoideini, and

Vartini); forewing with or without reflexed costal veins; connective not as above; usually without

long processes articulated with connective (exceptions: Adamina, some Scaphoideini).2

2. Connective T-shaped, bar-shaped, or platelike or, if Y-shaped, with arms forming broadly obtuse

angle (e.g., Figs 8G, 25G, 33G, 53G); style more or less linear, not broadly bilobed at base (e.g.,

Figs 8G, 25G, 33G, 53G)...3

Connective with anterior arms somewhat or not at all divergent, connective Y- or U-shaped (e.g., Figs

11G, 52G) or ‘linear’ with anterior arms closely appressed (e.g., Fig. 17G); style broadly bilobed at

base (e.g., Fig. 11G).7

3. Head much narrower than pronotum; profemur row AM with numerous preapical macrosetae;

metatibia dorsal surface with supernumerary macrosetae between rows PD and AD, especially

European Journal of Taxonomy 45: 1-211 (2013)

basally; pro- and mesotibia row PD and often row AD with numerous (> 6) macrosetae; forewing

appendix often large, extending around apex; Palearctic, Oriental.Stegelytrini

Head slightly narrower to wider than pronotum; profemur row AM with not more than 2

preapical macrosetae; metatibia dorsal surface without supernumerary macrosetae; pro- and

mesotibia with <6 macrosetae; forewing appendix reduced or absent.4

4. Anterior margin of head usually with a single distinct transverse carina between ocelli; epistomal

suture keeled (Fig. 9C) or evident from change in texture on frontoclypeus; subgenital plates

completely or partially fused to each other and/or to valve (if completely articulated then pronotum

produced dorsally, with longitudinal crest, Ikelibeloha ); Madagascar.Acostemmini (part)

Anterior margin of head shagreen, glabrous, or with several irregular striae or carinae, without single

distinct carina between ocelli; epistomal suture not keeled; subgenital plates not fused, articulated

with valve.5

5. Forewing acuminate apically (Fig. 8B); head narrower than pronotum (Fig. 8A) (exception:

Cariancha ); connective bar-shaped or plate-like, without well-defined stem (Fig. 8G); Nearctic,

Neotropical .Acinopterini

Forewing rounded apically; head as wide as or wider than pronotum; connective with well-

developed anterior arms posterior stem.6

6. Connective with median, darldy sclerotized Y- or T-shaped ridge following stem and arms of

connective (Fig. 25G); forewing central anteapical cell not strongly constricted medially; female

pregenital stemite VIII not concealed and folded beneath stemite VII, movably articulated with sternite

VII (Fig. 25 J); Palearctic, Nearctic (Fie beriellaflorii introduced in North America).Fieberiellini

Connective evenly sclerotized throughout (Fig. 33G); forewing central anteapical cell constricted

medially (Fig. 33B); female pregenital sternite VIII folded beneath and concealed by sternite

VII, stemites not articulated with each other; Neotropical.Luheriini

7. Subgenital plates fused to each other (Figs 9F, 26F); valve straplike, fused to pygofer laterally (Fig.

9D, F) OR not readily apparent (fused to plates); connective fused with aedeagus (Figs 9H, 26H)

..... 8

Subgenital plates usually free; valve present, articulated with pygofer laterally and usually

articulated with subgenital plates; connective articulated with or fused to aedeagus; rarely with

plates fused and connective fused with aedeagus, and if so then body slender or relatively small

in size, clypellus tapering apically (Deltocephalini: Miradeltaphus , Yuanamia) .9

8. Anterior margin of head with a distinct transverse carina between ocelli; forewings tectiform,

not overlapping at rest (Fig. 9A); forewing appendix reduced or absent (9B); valve strap-lilce,

fused to pygofer laterally (Fig. 9D, F); Afrotropical (especially diverse in Madagascar), Oriental

(India).Acostemmini (part)

Anterior margin of head without transverse carina; forewing, if macropterous, with apices

overlapping at rest (Fig. 26A) and appendix extending around wing apex (Fig. 25A); valve not

readily apparent, fused to subgenital plates; Palearctic, Afrotropical, Oriental, and Australian

.Goniagnathini

9. Head strongly produced, elongate or spatulate (Figs 18A, 21 A, 22A, 23A); gena acutely

incised laterally below eye (Figs 18C, 21C, 22C, 23C); pro femur intercalary row absent

or with few scattered setae; metafemur macrosetal formula 2+0 or 2+1; metatarsomere

I shorter than metatarsomeres II and III combined, expanded apically; metatibia straight

throughout length and distinctly bent distally in dorsal view .10

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Without above combination of characters; head produced or not; gena obtusely incised (e.g.,

Fig. 11C) or not incised laterally below eye; profemur intercalary row more or less well-

delimited and with several to many long fine setae; metafemur macrosetal formula usually

2+2+1; metatarsomere I longer than metatarsomeres II and III combined, usually not expanded

apically; metatibia slightly arcuate throughout its length in dorsal view.15

10. Aedeagus movably hinged between base and shaft (Fig. 14H); connective with anterior

arms closely appressed (Fig. 14G); first valvula dorsal sculpturing maculose to granulose,

submarginal; second valvula teeth obtusely triangular, serrate, uniformly shaped; Old World ....

.Chiasmini (part, Listrophora, some Chiasmus )

- Aedeagus without hinge; connective with anterior arms separate (Y-shaped); first valvula

dorsal sculpturing pattern variable; second valvula teeth variable .11

11. Ocelli on crown .12

- Ocelli on anterior margin of head.13

12. Macropterous; crown encroaching onto eyes (Fig. 22A-C); frontoclypeus with median

longitudinal carina (Fig. 22C); Palearctic .Eupelicini (part, Eupelicina)

- Brachypterous (Fig. 21A-B); crown not encroaching onto eyes; frontoclypeus and clypellus

tumid, without longitudinal carina (Fig. 21B-C); Afrotropical (South Africa, Fesotho) .

.Drakensbergenini

13. Macropterous; hindwing submarginal vein evanescent apically; valve fused to pygofer (Fig.

23D); male pygofer without basolateral cleft (Fig. 23D); style base connected or articulated

only laterally with apical part (Fig. 23J); Old World.Eupelicini (part, Paradorydiina)

- Brachypterous or macropterous, hindwing submarginal vein complete; valve articulated to pygofer;

male pygofer with basolateral cleft; style base broadly fused with apical part (e.g., Fig. 27G).14

14. Aedeagus with terminal processes (e.g.. Fig. 27H-I); first and second valvulae humpbacked

dorsally; first valvula with distinctly delimited ventroapical sculptured patch; second valvula

without teeth; Nearctic.Hecalini (part, Attenuipyga, Neoslossonia)

- Aedeagus without terminal processes (Fig. 18H-I); first and second valvulae not humpbacked

dorsally; first valvula without distinctly delimited ventroapical sculptured patch; second valvula

with distinct teeth; Palearctic.Dorycephalini

15. Anterior margin of pronotum exceeding eyes in dorsal view (Fig. 35A); integument black, shiny,

punctate (Fig. 3 5 A-B); Afrotropical, Oriental.Magnentiini

- Anterior margin of pronotum not exceeding eyes in dorsal view; integument rarely completely black

and punctate.16

16. Antennal ledge strong, forming distinct ledge, sometimes seemingly continuous with anterior

margin of head (Figs 19C, 45C); protibia dorsal side flat, margins carinate; forewing appendix

large, extending around wing apex.17

Antennal ledge absent or represented by weak carina not forming ledge; protibia rounded dorsally;

forewing appendix usually smaller, not extending around wing apex.18

17. Farge, robust leafhoppers, not squat or dorsally rounded or ventrally flattened (Fig. 19A-B);

ocelli on anterior margin of head (19C); frontoclypeus texture longitudinally striate or rugose

(Fig. 19C); Palearctic, Ethiopian, Oriental, Australian.Drabescini (part, Drabescina)

European Journal of Taxonomy 45: 1-211 (2013)

- Robust and squat, dorsally rounded, ventrally flattened leafhoppers (Fig. 45A-B); ocelli on crown

distant from eyes (Fig. 45B); frontoclypeus texture glabrous or finely shagreen; Cosmopolitan ..

.Penthimiini

18. Crown and face strongly rugose (Fig. 10A-C); body greater than 7 mm in length; Neotropical.

.Arrugadini

- Crown and face shagreen, glabrous, or striate, not rugose, or if rugose then body less than 7 mm in

length .19

19. Aedeagus bifurcate, with 2 gonopores.20

- Aedeagus with one shaft and gonopore.21

20. Face (Fig. 36B) strongly depressed, nearly horizontal, sometimes concave posteroventrally

in profile, sometimes inflated antero dors ally; anterior margin of head often with one or more

carinae or sharp ridges; antennal bases near anterodorsal (upper) corners of eyes (Fig. 36C);

stem of connective short; Afrotropical, Oriental.Mukariini (part)

- Face oblique, not strongly depressed, not concave in profile; anterior margin of head without

carinae (foliaceous in Chlidochrus)', antennal bases near middle or posteroventral (lower)

corners of eyes; stem of connective longer; Cosmopolitan.Opsiini

21. Head often somewhat produced (Fig. 36A); face flattened or frontoclypeus inflated medially or

anteriorly, lying in horizontal plane in lateral view (Fig. 36B-C); antennal bases near anterodorsal

(upper) corners of eyes (Fig. 36C); eyes large (Fig. 36C); antennae often long, longer than width

of head; profemur row AV setae absent; outer anteapical cell often confluent with or nearly

touching costal vein (Fig. 36B); Oriental, feeding on bamboo .Mukariini (part)

- Without above combination of characters; head produced or not; face flattened or not, not

distinctly inflated anteriorly, lying in horizontal plane in lateral view or not; size of eyes and

antennae variable; profemur row AV setae present or absent; outer anteapical cell usually not

touching or confluent with costal vein .22

22. Connective Y-shaped and fused to aedeagus (Fig. 16G-H); male pygofer with one or more

pairs of teeth, spines, or processes (Fig. 16D-E); male pygofer side excavated medially,

scoop-like (Fig. 16E); male segment X long, sclerotized laterally, sometimes sclerotized

ventrally and/or dorsally (Fig. 16D-E); second valvulae with a median dorsal tooth; Nearctic,

Neotropical (Cochlorhinus pluto introduced in Chile) .Cochlorhinini

- Connective shape variable, but not Y-shaped and fused to aedeagus; male pygofer with or

without processes; male pygofer side variable; male segment X variable; second valvulae with

or without dorsal tooth on shaft.23

23. Metatarsomere I with platellae on plantar surface; ocelli distant from eyes, at least 4 x own

diameter from adjacent eye (e.g.. Fig. 30C) .24

- Metatarsomere I without platellae on plantar surface; ocelli usually close to eyes (exceptions:

some Athysanini, Chi asm ini , and Faltalini) .26

24. Head not produced anteriorly (Fig. 30A); crown rounded to face (Fig. 3OB) or anterior

margin of head carinate, not foliaceous; Palearctic, Oriental, Nearctic (Grypotes puncticollis

introduced to northeastern North America) .Koebeliini, in part (Grypotina)

- Head strongly produced, foliaceous anteriorly (Figs 31A-C, 37A-C) .25

25. Ocelli on face (Fig. 31C); antennae not very long; forewing veins pustulate, appendix absent (Fig.

31A-B); Nearctic (western North America).Koebeliini (part, Koebeliina)

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

- Ocelli on crown (Fig. 36A1-A2); antennae very long; forewing veins not pustulate, appendix

present; Australian (Western Australia).Occinirvanini

26. Color often greenish or bluish with red or orange longitudinal stripes (Fig. 56A-B); crown produced,

pointed (Fig. 56A); gena not incised laterally below eye, visible behind the eye in dorsal view (Fig.

56B-C); frontoclypeus elongate (Fig. 56C); profemur intercalary row setae thick and extending

to or basad of middle of profemur; forewing macropterous, truncate apically (56B); apodemes

of male sternite II long, subrectangular, flared apically, directed posterolaterally (Fig. 56J);

connective with anterior arms closely appressed (Fig. 56G); male segment X tube-like, protruding

from pygofer, often well-sclerotized (Fig. 56D-E, K); Afrotropical, Oriental.Vartini

- Color variable; crown produced or not; gena usually incised laterally below eye, not visible behind

eye in dorsal view (exception: some Scaphoideini); frontoclypeus variable, usually not very elongate;

profemur intercalary row setae fine; if macropterous, forewing usually not truncate apically; apodemes

of male sternite II not as above; connective variable; male segment X variable.27

27. Clypellus parallel-sided ortapering apically (e.g., Fig. 17C); connective linear, triangular, orracquet-

like in shape, with anterior arms closely appressed anteriorly (e.g.. Figs 17G, 42G).28

- Clypellus variable; connective usually Y-shaped (e.g., 54G) or U-shaped (e.g., Fig. 52G), with

anterior arms separated (exception: some Athysanini, Scaphoideini).32

28. Subgenital plates fused with valve and apex of plate with sharply pointed process (Fig. 55F);

connective articulated posteriorly with a pair of long paraphyses (Fig. 55G); female sternite

VII ovate, sometimes very long; Palearctic, Afrotropical .Tetartostylini

- Subgenital plates usually articulated with valve, apices without sharply pointed processes;

paraphyses present or absent; female sternite VII not ovate, not very long.29

29. Connective fused with aedeagus (Fig. 17G) .30

- Connective articulated with aedeagus (Fig. 42G) .31

30. Body elongate, somewhat depressed; anterior margin of head transversely striate or carinate; first

valvula dorsal sculpturing pattern maculose to granulose and distinctly submarginal, with distinctly

delimited ventroapical sculptured area; second valvula without dorsal teeth; ovipositor protruding

well beyond py gofer apex; Nearctic (Mexico), Neotropical.Faltalini (part, Tenucephalus)

- Body not elongate or depressed; crown rounded to face, anterior margin shagreen; first valvula

sculpturing pattern imbricate (scale-like) and reaching dorsal margin, without distinct apicoventral

sculptured area; second valvula with irregular teeth; ovipositor not protruding far beyond

pygofer; Cosmopolitan .Deltocephalini

31. Aedeagus movably hinged between base and shaft (Fig. 14H) (exceptions: Baileyus , some

Gurawa , Paraphrodes ); ovipositor protruding beyond pygofer apex (exceptions: Gurawa ,

Baileyus , Omar anus, Paraphrodes ); first valvula dorsal sculpturing pattern maculose to

granulose, submarginal; second valvula teeth uniformly shaped, obtusely triangular, serrated;

Cosmopolitan.Chiasmini (part)

- Aedeagus entire, not movably hinged; ovipositor not protruding far beyond pygofer; first valvula

dorsal sculpturing pattern usually imbricate, reaching dorsal margin; second valvula teeth not

uniformly obtuse and triangular, serrated or not; Cosmopolitan .Paralimnini

32. Macropterous, forewing with 2 anteapical cells (Fig. 34B); usually small, slender leafhoppers

(Fig. 34A-B); Cosmopolitan.Macrostelini

- Macropterous to brachypterous, if macropterous then usually with 3 anteapical cells; if

with 2 anteapical cells, then larger, more robust leafhoppers .33

European Journal of Taxonomy 45: 1-211 (2013)

33. Small to medium sized, slender, elongate leafhoppers (Fig. 15A); usually yellow, ochraceous,

or pale green, often with paired black or brown markings or spots on head (Fig. 15A-C);

subgenital plate often with row of macrosetae distant from lateral margin, with numerous long

fine setae laterally (Fig. 15F); male pygofer incised dorsally nearly to base (Fig. 15D-E); male

segment X elongate and sclerotized dorsally (Fig. 15D-E) (not elongate in Knullana DeLong);

Palearctic, Nearctic, Neotropical.Cicadulini

- Size and color variable; subgenital plate macrosetae variable, with or without long fine setae;

male pygofer dorsal incision variable; male segment X usually not elongate and strongly

sclerotized.34

34. Macropterous to brachypterous, if macropterous then with appendix large, extending around

forewing apex; aedeagus movably hinged between base and shaft (Fig. 14H) (exceptions: Leofa ,

Picchusteles); ovipositor extending well beyond pygofer apex (exception: Paraphrodes ); first

valvula dorsal sculpturing pattern granulose to maculose, submarginal; second valvula teeth

uniformly shaped, obtusely triangular and serrated; Cosmopolitan .Chiasmini (part)

- Macropterous to brachypterous, if macropterous, with appendix not large or extending around

forewing apex; aedeagus not movably hinged; ovipositor length variable; first valvula dorsal

sculpturing variable; second valvula teeth not obtusely triangular and serrated.35

35. Crown narrow, width between eyes less or equal to median width of eye (Fig. 54A); crown

completely shagreen to base; pronotum lateral margin not carinate; male pygofer declivous,

directed posteroventrally, with few macrosetae, basolateral cleft absent or not membranous (Fig.

54D); ovipositor extending far beyond pygofer; first valvula with distinctly delimited ventroapical

sculptured area; second valvula without teeth; Cosmopolitan .Stenometopiini

- Crown not very narrow; discal portion of crown texture usually glabrous or striate (rarely shagreen

in some Athysanini [e.g., Anoterostemma], some Pendarini [Chlorotettix and Copididonus], and

Limotettigini); pronotum lateral margin usually carinate; male pygofer shape variable, basolateral

cleft present, membranous; ovipositor not protruding far beyond pygofer apex; first valvula without

distinctly delimited ventroapical sculpturing; second valvula with teeth.36

36. Dorsoventrally depressed or flattened leafhoppers (e.g., Fig. 24B); head strongly or sometimes

weakly produced; crown broad, more or less parabolically shaped (Figs 24A, 28A); crown to face

transition acutely angled; anterior margin sometimes foliaceous (Figs 24B, 28B); face lying slightly

obliquely to horizontal plane (Figs 24B, 28B); lateral margin of pronotum long, nearly as long

or longer than least width of eye (Figs 24A, 28A); first valvula dorsal sculpturing maculose to

granulose, distinctly submarginal; second valvula without dorsal teeth (exception: Glossocratus ) ...37

- Not particularly dorsoventrally depressed or flattened; head not strongly produced; crown

broad or not, rarely parabolically shaped; crown to face transition less acute; face usually

lying in a more upright plane; lateral margin of pronotum usually shorter than least width of

eye; first valvula dorsal sculpturing variable; second valvulae with teeth .38

37. Ocelli close to eyes, closer to eyes than lateral frontal sutures (Fig. 28C); male apodemes of

abdominal sternite I long, narrow, as long as apodemes of sternite II (Fig. 28J); male apodemes

of abdominal sternite II large, spatulate (Fig. 28J); first and second valvulae humpbacked

dorsally; second valvula concave ventrally; Cosmopolitan .Hecalini (part)

- Ocelli relatively distant from eyes, usually 3-4 x their diameter from ocellus or more

(exception: some Faltala , Bonamus), sometimes absent or obscure; male apodemes of

abdominal sternite I short or undeveloped; male apodemes of abdominal sternite II short

or undeveloped; first and second valvulae not strongly humpbacked dorsally; second

valvula not concave ventrally; Nearctic, Neotropical .Faltalini (part)

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

38. Body usually broad and robust (Fig. 13A-B); forewings brachypterous to submacropterous

(Fig. 13A-B) (exception: Renosteria ); hindwings when present narrowed anteriorly-posteriorly

when outstretched or otherwise reduced in size (Fig. 13J-K) (exception: Renosteria );

connective U- or sometimes Y- shaped with stem often very short or absent (Fig. 13G); male

segment X broad and sclerotized; ovipositor usually distinctly protruding beyond pygofer

apex (Fig. 13B); Afrotropical (South Africa) .Bonaspeiini

- Body shape variable; forewings usually macropterous; hindwings not narrowed anteriorly-

posteriorly when outstretched; connective variable; male segment X variable; ovipositor usually

not protruding far beyond pygofer apex.39

39. Anterior margin of head with 2 or more distinct carinae or with several distinct transverse striations

(e.g.. Fig. 50C); antennal ledge with oblique carina (e.g., Fig. 49C); medium to large-sized

leafhoppers.40

- Anterior margin of head without carinae or distinct transverse striations (rarely with single

transverse carina), texture shagreen or glabrous, crown rounded to face; antennal ledge

absent or represented by weak carina; size variable .44

40. Antenna long, at least as long as half length of body; antennal pits near upper (anterodorsal)

corners of eyes, large and encroaching onto frontoclypeus (Fig. 20C) (exception: Nirvanguina)',

profemur row AV setae reduced or absent, usually with fine setae; Afrotropical, Oriental,

Australian .Drabescini (part, Paraboloponina)

- Antenna not very long; antennal pits near lower corners of eyes (exception: Selenocephalini

[Dwightlina]), not very large or encroaching onto frontoclypeus); profemur row AV with distinct

stout setae.41

41. Profemur row AM often with 1 or more extra macrosetae basad of AMI; pygofer usually with

macrosetae reduced in number or absent (e.g., Fig. 52D) (exception: Adamina); connective stem

usually very short or absent, connective U-shaped (Fig. 52G); male segment X usually broad

and well-sclerotized dorsally; Afrotropical, Palearctic, Oriental .Selenocephalini

- Profemur row AM usually without extra macrosetae basad of AMI; connective stem present, relati¬

vely long, connective Y-shaped; male segment X variable, usually not very broad or large.42

42. Head distinctly narrower than pronotum (Fig. 46A); antennal ledges carinate (Fig. 46C);

lorum long, relatively large (Fig. 46C); clypellus long, widening apically (Fig. 46C); pronotum

lateral margin long, longer than half basal width of eye; forewing often with brown irrorate

markings; pro femur row AM sometimes with one or more setae basad of AMI; nymph

with pygofer short, with 6 rows of abdominal macrosetae .Phlepsiini (part)

- Head usually as wide as or wider than pronotum; antennal ledges variable; lorum not very

large; clypellus usually not particularly long; pronotum lateral margin short, usually less than

half basal width of eye; forewing coloration variable; profemur row AM only AMI seta;

nymph variable .43

43. Medium to large sized, robust leafhoppers (Fig. 12A-B); often predominantly brown in color (Fig.

12A-B); crown usually short, broad (Fig. 12A); often with anterior margin ofhead distinctly upturned,

raised, or with several transverse carinae (Fig. 12B-C); clypellus widening apically, parallel-

margined, or tapering apically (Fig. 12C), sometimes distinctly tumid (Fig. 12C); forewing often

with strongly pigmented reflexed costal veins (Fig. 12B); pygofer often with processes (Fig. 12D);

aedeagus often with apical processes (Fig. 12H-I); Nearctic, Neotropical.Bahitini (part)

- Without the above combination of characters; Cosmopolitan .Athysanini (part)

European Journal of Taxonomy 45: 1-211 (2013)

44. Aedeagus with single, unpaired sclerotized apophysis (= ‘dorsal connective’) projecting dorsally

from atrium, membranously connected to segment X (Fig. 32H-I); male pygofer dorsal margin

thickened, declivous, and often ending in ventrally directed curved hook or spine (Fig. 32D);

Palearctic, Oriental, Australian, Nearctic, Neotropical .Limotettigini

- Without the above combination of characters .45

45. Large, yellow to greenish species (Fig. 29A-B); forewing veins greenish (Fig. 29A-B); forewing

venation highly reticulated apically (Fig. 29B); Afrotropical (South Africa).Hypacostemmini

- Color variable; forewing venation rarely highly reticulate, but if so, reticulate portion not

restricted to forewing apex.46

46. Ocelli on face below anterior margin of head (Fig. 51C), distant from eyes; Afrotropical.

.Selenocephalini (part, Ianeirini: Ianeira)

- Ocelli on anterior margin of head, close to eyes .47

47. Head distinctly narrower than pronotum (Fig. 46A); antennal ledge carinate (Fig. 46C); lorum

long, relatively large (Fig. 46C); clypellus long, widening apically (Fig. 46C); pronotum lateral

margin longer than half basal width of eye (Fig. 46A); forewing often with brown irrorate

markings (Fig. 46A-B); pro femur row AM somet im es with one or more setae basad of AMI;

nymph with pygofer short, with 6 rows of abdominal macrosetae.Phlepsiini (part)

- Head width variable; antennal ledge variable; lorum not very large; clypellus usually not particularly

long; pronotum lateral margin usually less than half basal width of eye; forewing coloration

variable; profemur row AM without one or more macrosetae basad of AMI; nymph variable.48

48. Medium to large sized, robust leafhoppers (12A-B); often predominantly brown in color (12A-

B); crown usually short, broad (12A); often with anterior margin of head distinctly upturned,

raised (12B-C); clypellus widening apically, parallel-margined, or tapering apically (12C),

sometimes distinctly tumid (12C); forewing often with strongly pigmented reflexed costal

veins (13B); pygofer often with processes (12D); aedeagus often with apical processes (12H-

I); Nearctic, Neotropical .Bahitini (part)

- Size variable; color variable; anterior margin of head usually not upturned, raised; clypellus usually

widening apically or parallel-sided, not tumid; forewing with or without reflexed costal veins;

pygofer with or without processes; aedeagus with or without apical processes .49

49. (Note: the following characters are not present in all included genera, but some combination of

most of these characters is present in all): Head narrower than pronotum, produced anteriorly

(Fig. 47A); frontoclypeus long, narrow (Fig. 47C); antennae long, longer than width of head;

body slender (Fig. 47A); head and wings often with brown, orange, ochraceous, or ivory

markings (Fig. 47A-B); forewing with one or more darkly pigmented reflexed costal veins

(Fig. 47B); profemur row AV setae absent or reduced (not stout); metatibia row PD macrosetae

long, as long as or longer than 0.5x length of protibia; male or female pygofer with dense tufts

of long fine or regular setae (Fig. 47D); subgenital plate apex membranous or long, digitate,

somewhat membranous or weakly sclerotized; subgenital plates with long fine setae laterally

(Fig. 47F); basal processes of aedeagus or connective often present, connected or articulated to

base of aedeagus or apex of connective stem (Fig. 47J, M); Cosmopolitan.Scaphoideini

- Without some combination of above characters.50

50. Ground color yellowish or greenish OR fore wing with numerous brown, tawny, or reddish-

brown ramose or reticulate pigment lines, or connected dots forming such lines (Fig. 44A-B);

head as wide as or wider than pronotum; male pygofer often with a spine or process arising

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

from dorsal side; nymph crown-face transition carinate; face of nymph with longitudinal

carina in upper part; nymph pygofer elongate; Nearctic, Neotropical .Pendarini

- Without the above combination of characters; Cosmopolitan .Athysanini (part)

Acinopterini Oman, 1943

Fig. 8

Type genus: Acinopterus Van Duzee, 1892.

Diagnosis

Acinopterini are medium sized, yellowish, greenish, orange, or brown leafhoppers. They can be identified

by the straight commissural margin of the forewing, forewing apices not overlapping, forewing appendix

absent or reduced, connective bar-shaped or plate-like, style slender (not broadly bilobed), and female

first valvula strongly curved.

Description

Head. Head distinctly narrower than pronotum (. Acinopterus ) or subequal to or wider than pronotum

(' Cariancha ). Discal portion of crown glabrous with radial or longitudinal striae. Anterior margin of

head shagreen. Frontoclypeus not tumid; texture shagreen. Clypellus widening apically. Clypellus apex

following or slightly surpassing normal curve of gena. Lorum subequal to or wider than clypellus near

base. Antennal bases near middle or posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x

width of head. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture. Antennal

ledge wealdy developed (carinate or weakly carinate). Ocelli present; close to eyes; on anterior margin

of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; with commissural margin straight; apex acuminate; appendix absent or

reduced; with 3 anteapical cells; veins not raised; without reflexed costal veins; A1-A2 crossvein absent.

Fegs. Pro femur with AMI seta only or with one or more additional proximal setae; intercalary row

with one row of five or more fine setae; row AV with relatively long macrosetae. Protibia dorsal surface

rounded, convex. Metafemur apex macrosetae 2+2+1. Metatarsomere I not expanded apically; plantar

setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae well differentiated into several rows.

Subgenital plates free from each other; articulated with valve; without macrosetae. Style linear, median

anterior lobe not pronounced. Basal processes of the aedeagus present, fused to base of aedeagus.

Aedeagus with single shaft and gonopore. Connective anterior arms widely divergent; bar-shaped or

plate-like; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern strigate, concatenate, reticulate, or imbricate

(with overlapping scales); sculpturing reaching dorsal margin; without distinctly delimited ventroapical

sculpturing. Second valvula abruptly broadened medially or subapically or broad, gradually tapered

apically; without dorsal median tooth; teeth on apical 1/3 or more; teeth small, regularly or irregularly

shaped.

European Journal of Taxonomy 45: 1-211 (2013)

Fig. 8. Acinopterini. Acinopterus acuminatus Van Duzee, 1892. A-I. Standard views (see Material &

Methods). J. $ pygofer, ventral view.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Geography and ecology

Distribution: Nearctic, Neotropical. Acinopterus species live in meadows, grasslands, and in forest

understories and feed on a variety of herbaceous dicots. Acinopterus angulatus Lawson, 1922 is a vector

of the western strain of aster yellows and western X disease.

Remarks

Acinopterini contains 2 genera and 30 species. Acinopterus (28 spp.) is widely distributed from North

to South America and Cariancha (2 spp.) is known only from Brazil. This is an early-diverging tribe

of Deltocephalinae and it possesses plesiomorphic characters of the male genitalia. Among the other

early-diverging lineages of Deltocephalinae, the 28S rDNA sequence of Acinopterus was updated to

include sequences only from A. acuminatus Van Duzee, 1892 (previously a chimeric sequence of A.

acuminatus and A. viridis Ball, 1903 was used) and the D2 region of Lukeria was included for the

first time. However, these changes did little to resolve the relationships between Acinopterini and the

other early-diverging deltocephaline tribes, Stegelytrini, Acostemmini, Luheriini and Fieberiellini.

Future analyses should include more representatives of these tribes and representatives of Cariancha to

facilitate further understanding of their relationships.

Selected references

Beamer & Lawson (1938), Oman (1949), Linnavuori & DeLong (1977), Cwikla & Ma (1986).

Included Genera

Acinopterus Van Duzee, 1892.

Cariancha Oman, 1938.

Acostemmini Evans, 1972

Fig. 9

Type genus: Acostemma Signoret, 1860.

= Acroponinae Linnavuori & Quartau, 1975.

Diagnosis

Acostemmini are medium sized to very large yellow, greenish, reddish-brown, brown, or greyish

leafhoppers. They can be identified by the single carina on the anterior margin of the head, distinct

epistomal suture, forewing with a straight commissural margin (apices not overlapping) and without

appendix, pygofer setae reduced or absent, valve fused to pygofer or lateral margin long and articulated

with pygofer, style linear-shaped, and subgenital plates sometimes fused to each other.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae. Anterior margin of head with a single transverse carina. Frontoclypeus not tumid;

texture mostly shagreen; epistomal suture distinct or evident by change in texture from glabrous, strigate,

or irregular above suture to shagreen below. Clypellus widening apically; sometimes long and hourglass-

shaped. Clypellus apex following or slightly surpassing normal curve of gena. Lorum subequal to or

wider than clypellus near base. Antennal bases near middle or posteroventral (lower) corners of eyes.

Antennae short, less than 1.5 x width of head. Gena obtusely incised laterally; with fine erect seta beside

laterofrontal suture. Antennal ledge weakly developed (carinate or weakly carinate). Ocelli present;

close to eyes; on anterior margin of head.

European Journal of Taxonomy 45: 1-211 (2013)

Fig. 9. Acostemmini. Acostemma stilleri sp. nov. A-I. Standard views (see Material & Methods).

F. Plates shown with pygofer. G. Connective and style shown with part of fused aedeagus. H. Aedeagus

shown with fused processes, connective, and style. J. Lateral view of style, connective, and base of

aedeagus. K. Ventral view of connective, styles, and ventral processes of aedeagus. L. Detail of hairlike

extensions of egg. M. Egg. N. $ sternite VII.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous, submacropterous, or rarely subbrachypterous; appendix absent or

reduced; with 3 anteapical cells; veins not raised; veins sometimes with callosities; without reflexed

costal veins; A1-A2 crossvein present; apical venation strongly reticulate or not reticulate.

Legs. Pro femur with AMI seta only or with one or more additional proximal setae; intercalary row with

one row of five or more fine setae; row AV with short, stout setae or with relatively long macrosetae.

Protibia dorsal surface rounded and convex or flat (AD and PD margins at -90° angles but not carinate).

Metafemur apex macrosetae with 2+2+1. Metatarsomere I not expanded apically; plantar setae simple,

tapered.

Male genitalia. Valve straplike, sometimes longer laterally than medially ( Acostemella ); articulated

with or fused to pygofer; if articulated then lateral margin long, with length of articulation long;

articulated with or fused to subgenital plates. Pygofer basolateral membranous cleft absent; macrosetae

reduced or absent or well differentiated into several rows. Subgenital plates fused or articulated with

each other; articulated with or fused to valve; without macrosetae, with macrosetae irregularly arranged,

or with macrosetae uniseriate laterally. Style linear, median anterior lobe not pronounced or broadly

bilobed basally, median anterior lobe pronounced; apophysis sometimes bent at nearly 90° angle. Basal

processes of the aedeagus/connective absent or present, if present then connected to or articulated with

connective or base of aedeagus. Aedeagus with single shaft and gonopore. Connective anterior arms

widely divergent, T-shaped, or somewhat divergent, Y-shaped; sometimes with an anteromedial lobe

or process (. Acostemella , Alocoelidia , Caelidioides, Ikelibeloha , Iturnoria)', fused to or articulated with

aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex or not strongly convex; dorsal sculpturing pattern strigate, concatenate, or

reticulate; sculpturing reaching dorsal margin; without distinctly delimited ventroapical sculpturing.

Second valvula broad, gradually tapered or slender throughout, without dorsal median tooth; teeth

restricted to apical 1/4 or less; teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Afrotropical and Oriental (India, Sri Lanka, New Guinea) regions. The diversity of the

tribe lays mostly in Madagascar, where it appears to have radiated and subsequently dispersed to

mainland Africa, India, and Sri Lanka where species of Acostemma are found. Several undescribed

genera and many species have recently been discovered (Zahniser & Nielson 2012) based on samples

from an arthropod biodiversity inventory of Madagascar. Some species in the genus Eryapus resemble

bird droppings.

Remarks

Acostemmini contains 12 genera and 27 species. This subfamily is arelictual lineage of Deltocephalinae

and retains some primitive features of the male genitalia.

The morphologically enigmatic genus Ikelibeloha and four other genera were included in the tribe

by Zahniser & Nielson (2012). They noted that these genera do not share all of the male genitalic

characters that previously helped to define the tribe (e.g., valve fused to pygofer) but shared other

unique acostemmine characters that supported their placement in the tribe. Ikelibeloha and Iturnoria

were included in phylogenetic analyses using only molecular data by Zahniser & Nielson (2012) and

formed a monophyletic group with the other sampled acostemmines ( Acostemma and Eryapus). In the

MP analysis here, Ikelibeloha and Iturnoria did not group with the other acostemmines, a result which is

European Journal of Taxonomy 45: 1-211 (2013)

likely due to the inclusion of morphological data and the divergent morphology of these taxa as compared

to “typical” Acostemmini. However the ML and Bayesian analyses of combined data recovered the two

genera as sister to the other included Acostemmini, Acostemma and Eryapus , with strong branch support

(85 BS support and 1.0 PP, respectively), thus forming a monophyletic Acostemmini. The previous MP,

ML, and Bayesian analyses of only molecular data also strongly supported this relationship (Zahniser &

Nielson 2012). Despite the result of the MP analysis here, the relationships among the earliest lineages of

Deltocephalinae as those depicted in the ML and Bayesian trees here (both with moderate to high branch

support) showing Stegelytrini as the earliest diverging tribe sister to a monophyletic Acostemmini and

the remaining Deltocephalinae are the most well-supported by the available evidence.

Evans (1972b) tentatively placed Telopetulcus (not examined in this study), recorded only from New

Guinea, in Acostemmini. Its placement is still uncertain. Fig. 6A of Evans (1972b) suggests the presence

of the epistomal suture on the face which would indicate that it belongs in Acostemmini, but some other

features (e.g., Y-shaped connective articulated with aedeagus) do not occur in other genera of the tribe.

Closer examination of this genus is needed with reconsideration of its tribal placement.

Selected references

Evans (1954, 1972b), Linnavuori & Quartau (1975), Linnavuori & Al-Ne’amy (1983), Zahniser &

Nielson (2012).

Included genera

Acostemana Evans, 1954

Acostemma Signoret, 1860

Acostemmella Evans, 1954

Alocoelidia Evans, 1954

Caelidioides Signoret, 1880

Dardania Stal, 1866

Eryapus Evans, 1954

Ikelibeloha Zahniser & Nielson, 2012

Iturnoria Evans, 1954

Malicia Evans, 1954

Protonesis Spinola, 1850

Telopetulcus Evans, 1972

Arrugadini Linnavuori, 1965

Fig. 10

Type genus: Arrugada Oman, 1938.

Diagnosis

Arrugadini are large brown and yellow leafhoppers. They can be easily identified by the strongly rugose

texture of the head and face, lateral margins of the pronotum longer than the basal width of the eye,

first valvula dorsal sculpturing pattern granulose and submarginal, first valvula with distinctly delimited

ventroapical sculpturing, and second valvula gradually expanded apically and without teeth.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown rugose or longitudinally

striate. Anterior margin of head irregularly textured, rugose. Frontoclypeus tumid or not tumid; texture

rugose. Clypellus widening apically. Clypellus apex following or slightly surpassing normal curve of

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 10. Arrugadini. A-B, D-J. Arrugada linnavuorii sp. nov. C. Arrugada sp. A-I. Standard views (see

Material & Methods). J. Broad view of style.

European Journal of Taxonomy 45 : 1-211 ( 2013 )

gena. Lorum subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral

(lower) corners of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely incised laterally;

with fine erect seta beside laterofrontal suture. Antennal ledge weakly developed (carinate or wealdy

carinate). Ocelli present; close to or somewhat distant from eyes; on anterior margin of head.

Thorax. Pronotum not exceeding or exceeding (female of A. rugosa) eyes anteriorly; lateral margin

carinate; lateral margin as long as or longer than basal width of eye.

Wings. Forewing macropterous or submacropterous; appendix restricted to anal margin; with 3 anteapical

cells; veins distinctly raised or carinate; without reflexed costal veins; A1-A2 crossvein absent.

Legs. Profemur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with relatively long macrosetae. Protibia dorsal surface flat, AD and PD margins at -90° angles but not

carinate. Metafemur apex macrosetae 2+2+1 or 2+1+1. Metatarsomere I not expanded apically; plantar

setae simple, tapered.

Male genitalia. Sternal apodemes of segments I and II small, not well developed. Valve articulated

with pygofer; lateral margin short, articulating with pygofer at a point. Pygofer basolateral membranous

cleft present; macrosetae well differentiated into several rows. Subgenital plates free from each other;

articulated with valve; macrosetae uniseriate laterally. Style broadly bilobed basally, median anterior

lobe pronounced. Basal processes of the aedeagus/connective absent or reduced. Aedeagus with single

shaft and gonopore. Connective anterior arms somewhat divergent, Y-shaped; articulated with aedeagus.

Female genitalia. Pygofer macrosetae reduced or absent. First valvula not strongly convex; dorsal

sculpturing pattern granulose or maculose; sculpturing submarginal; with ventroapical sculpturing

distinctly delimited. Second valvula gradually broadened medially or subapically; without dorsal median

tooth; teeth absent.

Geography and ecology

Distribution: Neotropical. Host plants are not known. Large numbers of males of Arrugada affinis

(Osborn, 1924) were observed and collected while “mud-puddling” in Peru (Rakitov et al. 2005).

Remarks

Arrugadini contains only the type genus and 4 valid species from Bolivia and Peru. This small group was

considered a separate subfamily until it was recently included as a tribe of Deltocephalinae (Zahniser &

Dietrich 2010). Based on some characters of the female genitalia (first valvula dorsal sculpturing pattern

granulose and submarginal, first valvula with distinctly delimited ventroapical sculpturing, second

valvula without dorsal teeth), it appears to be related to the grass-feeding tribe Hecalini, a relationship

that was marginally supported in the combined parsimony analyses presented here. However, the shapes

of the first and second valvulae are not very similar to Hecalus and Attenuipyga which have humpback¬

shaped valvulae. The relationships within Hecalini and between this and other tribes seem particularly

difficult to recover given the data in this study. More data are needed to resolve the relationships between

Arrugadini and other Deltocephalinae. Preliminary phylogenetic analyses of Hecalini (Catanach,

unpublished) suggest that Arrugada is related to another South American-endemic genus, Egenus.

Egenus was removed from Hecalini by Hamilton (2000), tentatively placed in Faltalini by Zahniser &

Dietrich (2010), and is transferred to Athysanini here (see below).

Selected references

Linnavuori & DeLong (1978a), Zahniser & Dietrich (2010).

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Included genera

Arrugada Oman, 1938.

Athysanini Van Duzee, 1892

Fig. 11

Type genus: Athysanus Burmeister, 1838.

= Phrynomorphini Kirkaldy, 1907.

= Thamnotettigini Distant, 1908.

= Euscelini Van Duzee, 1917.

= Anoteroste mmini Haupt, 1929 syn. nov.

= Bobacellini Kusnezov, 1929.

= Platymetopiini Haupt, 1929 syn. nov.

= Colladonini Bliven, 1955.

= Cerrillini Linnavuori, 1975.

= Allygidiina Dmitriev, 2006 syn. nov.

Diagnosis

Because this is such a large tribe, it is difficult or impossible to provide a set of characters that will

easily diagnose it. There is substantial morphological diversity in the group, but most members have a

Y-shaped connective and lack the distinctive features of other tribes.

Description

Head. Head subequal to or wider than pronotum or distinctly narrower than pronotum. Discal portion

of crown glabrous, radially or longitudinally striate, or shagreen. Anterior margin of head shagreen,

glabrous, irregularly textured, with numerous transverse striations, foliaceous, with 2 or 3 parallel

carinae, or with numerous carinae. Frontoclypeus not tumid or tumid; texture shagreen or rugose.

Clypellus parallel-sided or widening apically; apex following or slightly surpassing normal curve of

gena. Lorum subequal to or wider than clypellus near base or distinctly narrower than clypellus near

base. Antennal bases near middle or posteroventral (lower) corners of eyes. Antennae short, less than

1.5 x width of head or long, 1.5 x width of head or longer. Gena obtusely incised laterally; with fine

erect seta beside laterofrontal suture. Antennal ledge absent or wealdy developed (carinate or weakly

carinate). Ocelli present; close to eyes or distant from eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate or not; lateral margin shorter than basal width of eye.

Wings. Forewing brachypterous to macropterous; appendix absent or reduced (in submacropterous or

brachypterous forms) or restricted to anal margin; with 3 anteapical cells; veins not raised; with or

without reflexed costal veins; A1-A2 crossvein absent or present; apical venation usually not strongly

reticulate, rarely reticulate.

Fegs. Pro femur with AMI absent or reduced, with AMI seta only, or with AMI and with one or more

additional proximal setae; intercalary row with one row of five or more fine setae; row AV with thin,

hair-like setae, without setae, with short, stout setae, or with relatively long macrosetae. Protibia dorsal

surface rounded, convex. Metafemur apex macrosetae 2+2+1 or 2+2+1 with additional proximal

macrosetae. Metatarsomere I not expanded apically or expanded apically; plantar setae usually simple,

tapered.

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Fig. 11. Athysmini. Athysanus argentarius Metcalf, 1955. A-I. Standard views (see Material & Methods).

J. Hindwing.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Male genitalia. Valve articulated with pygofer or fused to subgenital plates; lateral margin short,

articulating with pygofer at a point. Pygofer basolateral membranous cleft present; macrosetae absent

or reduced (< two rows) or macrosetae well differentiated into several rows. Subgenital plates free

from each other; articulated with or rarely fused to valve; macrosetae absent or present, scattered

and irregularly arranged, uniseriate laterally, with two lateral rows, with some irregularly arranged

macrosetae near lateral margin, or uniseriate and distant from lateral margin. Style broadly bilobed

basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent or reduced

or present, connected or articulated to connective or near base of aedeagus. Aedeagus with single shaft

and gonopore. Connective anterior arms somewhat divergent, Y- or U-shaped, or rarely anterior arms

closely appressed anteriorly; articulated with or rarely fused to aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond or protruding

far beyond pygofer apex. First valvula convex or not strongly convex; dorsal sculpturing pattern strigate,

concatenate, reticulate, or imbricate (with overlapping scales); sculpturing reaching dorsal margin or

submarginal; with indistinctly delimited ventroapical sculpturing or rarely with ventroapical sculpturing

distinctly delimited. Second valvula abruptly broadened medially or subapically or broad, gradually

tapered; with or without dorsal median tooth; dorsal teeth on apical 1/3 or more; teeth large, regularly

shaped or small, regularly or irregularly shaped.

Geography and ecology

Distribution: cosmopolitan. Athysanini can be found in nearly all terrestrial ecosystems. Athysanini

feed on a wide variety of eudicots and some species occasionally feed on grasses or sedges. This

group contains numerous vectors of crop diseases. Some of the most well-known include: Colladonus

clitellarius (Say, 1830), C. geminatus (Van Duzee, 1890) and C. montanus (Van Duzee, 1892), which

transmit Eastern and Western X-diseases of peach, aster yellows, yellow leaf roll stain, and Euscelis

incisus (Kirschbaum, 1858), a vector of numerous diseases of clover.

Remarks

This is the largest tribe of Deltocephalinae, including 228 genera and 1123 species. Habitus images are

available for most genera of the tribe through the online database. Athysanini constitute a polyphyletic

assemblage of genera which have historically been placed in the tribe mostly because they retain the most

common deltocephaline external and genitalic characters but lack the distinctive features that define other

tribes. More detailed studies of the phylogeny of Deltocephalinae including many more representatives

of Athysanini are needed to more fully understand the relationships of these genera and the evolution

of all of Deltocephalinae. Such studies may also help to determine morphological characters that define

smaller, more particular lineages, particularly those belonging to clades that received strong branch

support in our phylogenetic analyses. For now, the revised classification presented here including the

new tribes Bahitini and Phlepsiini and the revised interpretation of Scaphoideini separates some groups

of genera which were previously placed in Athysanini or were previously unplaced to tribe. Revisions to

Athysanini here include the addition of many genera that were previously unplaced in Deltocephalinae,

thus resulting in a classification of the subfamily that has no genera unplaced to tribe. This arrangement

seems preferable to an alternative option of restricting Athysanini to the interpretation of the Athysanus

genus group, as defined here, and considering all other genera once included in Athysanini to be unplaced

to tribe in Deltocephalinae. The latter option would likely create more confusion, as did the previous

classification which contained both unplaced genera and a large number of genera in Athysanini without

any clear distinction between the two.

Other changes made here include considering Allygidiina, Anoterostemmina, and Platymetopiina

synonyms of Athysanini. Emeljanov (1999) and Dmitriev (2006b) recognized up to 4 subtribes

of Athysanini (Allygidiina, Athysanina, Cicadulina, and Platymetopiina) although their generic

European Journal of Taxonomy 45 : 1-211 ( 2013 )

classifications partly conflict. Both authors included the genera treated here as Cicadulini in Athysanini

(Emeljanov in Athysanina, and Dmitriev as the subtribe Cicadulina). The phylogenetic analyses

here included representatives of 5 of the 14 genera included in Platymetopiina by Dmitriev (2006b):

Anoplotettix , Colladonus, Phlepsius , Platymetopius, and Thamnotettix. Except for a relationship

between Colladonus and Platymetopius , the analyses strongly suggest that these genera are not closely

related and that the morphological characters previously used to define the group are not informative

of phylogenetic relationships. Therefore, Platymetopiina is not recognized here. Further investigation

may allow for a revised concept of this group. The genera Eusama and Twiningia were found with

strong branch support to be related to Colladonus and Platymetopius , which could in part inform

future recognition of such a taxon. The phylogenetic analyses grouped the included representatives of

Athysanina and Allygidiina in the same area of the tree and they share some morphological features.

They are therefore recognized together in the informal Athysanus genus group, and some other genera

that were not within the geographical ranges of the previous taxonomic studies are included in the group.

The morphologically unique Neotropical genus Chimaerotettix was included in our analyses, but its

placement with respect to other Athysanini was poorly resolved.

Justification for the inclusion of other genera not covered in the previous discussion is given below.

Egenus was previously included in Faltalini by Zahniser & Dietrich (2010) who had not examined

female specimens. It is transferred to Athysanini here because female specimens became available

for study which showed that they do not possess the synapomorphies of the first and second valvulae

that in part define Faltalini. Preliminary phylogenetic work on the tribe Hecalini (Catanach, personal

communication) suggests that Egenus may be related to Arrugada.

Loralia is transferred to Athysanini from Deltocephalini. It clearly does not belong in Deltocephalini

and was found here to be closely related to Occinirvanini. However, we were unable to identify

morphological features uniting Loralia and Occinirvana, so more study of the Australian fauna will be

needed to determine whether including Loralia or other genera in Occinirvanini is justifiable.

Nesothamnus is transferred from Scaphytopiini. It is not closely related to Scaphytopius but is potentially

more closely related to some Neotropical Athysanini. In several analyses, it was resolved as sister to or near

Idioceromimus with which it shares a relatively short, broad body form and bright orange/red coloration.

Phycotettix was listed in Fieberiellini by Oman et al. (1990). It does not possess the characters that

are diagnostic for Fieberiellini and its placement there was perhaps a mistake due to the similarity

of the name to the fieberielline genus Placotettix. Phycotettix was not included in Fieberiellini in a

comprehensive revision of the tribe (Meyer-Arndt & Remane 1992a, b).

Selected references

Oman (1949), Finnavuori (1959), Finnavuori & DeFong (1978b), Cwilda & Blocker (1981), Anufriev

& Emeljanov (1988), Emeljanov (1999), Dmitriev (2006b).

Included genera

Abrus Dai & Zhang, 2002

Acacimenus Dlabola, 1979

Acunasus DeFong, 1945 placement nov. (previously unplaced in Deltocephalinae)

Aeternus Distant, 1918 placement nov. (previously unplaced in Deltocephalinae)

Aindrahamia Finnavuori, 1965

Aligia Ball, 1907

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Alladanus DeLong & Harlan, 1968

Allophleps Bergroth, 1920

Allotapes Emeljanov, 1964 placement nov. (previously placed in Platymetopiina)

Amblytelinus Lindberg, 1954

Anaemotettix Korolevslcaya, 1980 placement nov. (previously placed in Platymetopiina)

Ancudana DeLong & Martinson, 1974

Andanus Linnavuori, 1959

Angubahita DeLong, 1982

Angulanus DeLong, 1946

Antoniellus Linnavuori, 1959

Aplanatus Cheng, 1980

Aplanus Oman, 1949

Aplanusiella Nielson & Haws, 1992

Arahura Knight, 1975

Arawa Knight, 1975

Argyrilla Emeljanov, 1972

Aricanus Linnavuori, 1959

Artucephalus DeLong, 1943

Asiotoxum Emeljanov, 1964

Asthenotettix Korolevslcaya, 1980 placement nov. (previously placed in Platymetopiina)

A tanus Oman, 1938

Awasha Heller & Linnavuori, 1968

Bambusana Anufriev, 1969

Bambusananus Li & Xing, 2011

Bandar a Ball, 1931

Bardana DeLong, 1980 placement nov. (previously unplaced in Deltocephalinae)

Baroma Oman, 1938

Benibahita Linnavuori, 1959

Bergolix Linnavuori, 1959

Bicoloratnm Dai & Li, 2011

Biluscelis Dlabola, 1980

Bolotheta Kramer, 1963

Bonneyana Oman, 1949

Brachylope Emeljanov, 1962

Brachypterona Lindberg, 1954

Branchana Li, 2011

Brasilanus Linnavuori, 1959

Brazosa Oman, 1938

Cahya Linnavuori, 1959

Caladonus Oman, 1949

Caranavia L innavuori, 1959

Carelmapu Linnavuori, 1959

Cenedaens Distant, 1908

Cerrillus Oman, 1938

Cetexa Oman, 1949

Chapa re a Linnavuori, 1959

Chimaerotettix Dietrich & Ralcitov, 2002 placement nov. (previously unplaced in Deltocephalinae)

Chroocacus Emeljanov, 1962

Cocrassana Blocker & Larsen, 1991

Colladonus Ball, 1936 placement nov. (previously placed in Platymetopiina)

European Journal of Taxonomy 45: 1-211 (2013)

Comayagua Linnavuori & DeLong, 1978

Conoscelis Abdul-Nour, 1988

Consepusa Linnavuori & DeLong, 1977

Cornersana DeLong, 1967

Costamia DeLong, 1946

Cozadanus DeLong & Harlan, 1968

Crassana DeLong & Hershberger, 1947

Cyanidius Emeljanov, 1964 placement nov. (previously placed in Platymetopiina)

Cyclevertex Li & Xing, 2011

Dagama Distant, 1910

Dampfiana DeLong & Hershberger, 1948

Danbara Oman, 1949 placement nov. (previously unplaced in Deltocephalinae)

Deltorhynchus DeLong, 1943

Desertana DeLong & Martinson, 1973

Devolana DeLong, 1967

Didymotettix Yang, 1996

Divitiacus Distant, 1918

Dlabolaracus Remane, 1961

Doleranus Ball, 1936

Dolyobius Linnavuori, 1959

Egenus Oman, 1938 placement nov. (transferred from Faltalini)

Elphnesopius Nast, 1984

Ephelodes Emeljanov, 1972 placement nov. (previously placed in Platymetopiina)

Euleimonios Kirkaldy, 1906 placement nov. (previously unplaced in Deltocephalinae)

Eusama Oman, 1949 placement nov. (previously placed in Platymetopiina)

Eusceloidia Osborn, 1923 placement nov. (previously unplaced in Deltocephalinae)

Eusora Oman, 1949

Eutettix Van Duzee, 1892

Excavanus DeLong, 1946

Fistulatus Zhang, 1997 placement nov. (previously unplaced in Deltocephalinae)

Fitchana Oman, 1949

Floridonus Oman, 1949

Fridonus Oman, 1949

Garapita Oman, 1938

Gunawardenea Fletcher & Moir, 2008

Hamulotettix Dai & Zhang, 2011

Hardya Edwards, 1922 placement nov. (previously placed in Platymetopiina)

Hegira Oman, 1938

Houtbayana Linnavuori, 1961

Huancabamba Linnavuori, 1959

Idioceromimus Dietrich & Rakitov, 2002 placement nov. (previously unplaced in Deltocephalinae)

Idiodonus Ball, 1936 placement nov. (previously placed in Platymetopiina)

Inghamia Evans, 1966 placement nov. (previously placed in Platymetopiina)

Jaacunga Nielson, 1988

Jakarellus Webb, 1980 placement nov. (previously placed in Platymetopiina)

Krameraxus Maldonado-Capriles, 1968 placement nov. (previously unplaced in Deltocephalinae)

Lajolla Linnavuori, 1959

Laminacutus Abdul-Nour, 2002

Lamprotettix Ribaut, 1952 placement nov. (previously placed in Platymetopiina)

Lascumbresa Linnavuori & DeLong, 1979

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Latinocesa Kocak & Kemal, 2010 placement nov. (previously unplaced in Deltocephalinae)

Laylatina Abdul-Nour, 1988

Lineana Li & Xing, 2011

Lojanus Linnavuori, 1959 placement nov. (previously unplaced in Deltocephalinae)

Loralia Evans, 1966 placement nov. (transferred from Deltocephalini)

Malasiella Evans, 1954 placement nov. (previously placed in Platymetopiina)

Melillaia Linnavuori, 1971

Mesadorus Linnavuori, 1955

Mesamia Ball, 1907

Mexicananus DeLong, 1944

Mimodorns Linnavuori, 1959

Mocolinna McKamey, 2003

Moskgha Deeming & Webb, 1982

Nakaharanus Ishihara, 1953 placement nov. (previously placed in Platymetopiina)

Napo Linnavuori & DeLong, 1976

Neocrassana Linnavuori, 1959

Neodonus DeLong & Hershberger, 1948 placement nov. (previously placed in Platymetopiina)

Neohegira Linnavuori & DeLong, 1978

Neomacednus Xing & Li, 2011

Neomesus Linnavuori, 1959

Neoreticulum Dai, 2009

Nesophryne Kirkaldy, 1907

Nesothamnus Linnavuori, 1959 placement nov. (transferred from Scaphytopiini)

Neurotettix Matsumura, 1914

Nigridonus Oman, 1949

Norvellina Ball, 1931 placement nov. (previously placed in Platymetopiina)

Nurenus Oman, 1949 placement nov. (previously placed in Platymetopiina)

Occiplanocephalus Evans, 1941

Okaundua Linnavuori, 1969

Ollarianus Ball, 1936

Ophionotum Emeljanov, 1964 placement nov. (previously placed in Allygidiina)

Orientus DeLong, 1938

Oxytettigella Metcalf, 1952

Pachytettix Linnavuori, 1959

Paracolladonus Nielson, 1988

Paracrassana Nielson, 1988

Paraganus Linnavuori, 1955

Paralaca Lozada, 1998 placement nov. (previously unplaced in Deltocephalinae)

Paramacrosteles Dai, Li & Chen, 2006

Paranurenus Nielson, 1988

Paratanus Young, 1957

Perundanus Zanol, 1993 placement nov. (previously unplaced in Deltocephalinae)

Phlepsanus Oman, 1949

Phlepsobahita Linnavuori, 1959

Phycotettix Haupt, 1929 placement nov. (transferred from Lieberiellini)

Pingellus Evans, 1966

Platymetopius Burmeister, 1838 placement nov. (previously placed in Platymetopiina)

Poliona Emeljanov, 1972 placement nov. (previously placed in Platymetopiina)

Protensus Zhang & Dai, 2001

Pseudalaca Linnavuori, 1959 placement nov. (previously unplaced in Deltocephalinae)

European Journal of Taxonomy 45: 1-211 (2013)

Pseudaligia Kramer & DeLong, 1968

Pseudaraldus Bonfils, 1981

Pseudocephalelus Linnavuori, 1969

Pseudosubhimalus Ghauri, 1974

Pseutettix DeLong, 1967

Pygotettix Matsumura, 1940

Renonus DeLong, 1959

Retusanus DeLong, 1945

Rinconada Linnavuori & DeLong, 1977

Sanuca DeLong, 1980

Sard ins Ribaut, 1946

Saudallygus Dlabola, 1979 placement nov. (previously unplaced in Deltocephalinae)

Scaphetus Evans, 1966

Scaphomonoides Li, 2011

Scaphytoceps Dlabola, 1957 placement nov. (transferred from Scaphytopiini)

Scenergates Emeljanov, 1972 placement nov. (previously placed in Platymetopiina)

Selachina Emeljanov, 1962

Serratus Linnavuori, 1959

Sincholata DeLong, 1982 placement nov. (previously unplaced in Deltocephalinae)

So bar a Oman, 1949 placement nov. (previously unplaced in Deltocephalinae)

Spinulana DeLong, 1967

Stenometohardya Dlabola, 1981

StenomisellaE\ms, 1954

Stoneana DeLong, 1943

Streptopyx Linnavuori, 1958

Stymphalella Evans, 1954 placement nov. (previously placed in Platymetopiina)

Tambocerus Zhang & Webb, 1996

Taped a Emeljanov, 1964 placement nov. (previously placed in Platymetopiina)

Tenuisanus DeLong, 1944

Thanomahia Dlabola, 1987 placement nov. (previously unplaced in Deltocephalinae)

Tingolix Linnavuori & DeLong, 1978

Twiningia Ball, 1931 placement nov. (previously placed in Platymetopiina)

Usanus DeLong, 1947

Watanabella Vilbaste, 1969

Yungasia Linnavuori, 1959

Zabrosa Oman, 1949

Zercanus Dlabola, 1965

Zilkaria de Menezes, 1974

Athysanus- group (informal genus group)

Fig. 11

Diagnosis

The Athysanus- group can be identified by their relatively large and broad shape, frontoclypeus often

broad (never long and narrow), clypellus usually parallel-margined or sometimes slightly expanded

apically, first valvula often with imbricate sculpturing pattern, and second valvulae often with distinct

median dorsal tooth on shaft.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae, or shagreen. Anterior margin of head shagreen; crown rounded to face. Frontoclypeus

not tumid to somewhat tumid; texture shagreen or glabrous. Clypellus parallel-sided, tapering apically,

or widening apically; apex following or slightly surpassing normal curve of gena. Lorum subequal to

or wider than clypellus near base or distinctly narrower than clypellus near base. Antennal bases near

middle or posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x width of head. Gena

obtusely incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledges absent or

weakly developed (carinate or weakly carinate). Ocelli present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than half basal width of eye.

Wings. Forewings brachypterous to macropterous; appendix absent, reduced; or restricted to anal

margin; with 3 anteapical cells; veins not raised; without reflexed costal veins; A1-A2 crossvein absent

or present; apical venation not highly reticulate.

Legs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae or with relatively long macrosetae. Protibia dorsal surface rounded, convex.

Metafemur apex macrosetae 2+2+1. Metatarsomere I not expanded apically or expanded apically;

plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer or rarely fused to subgenital plates; with short point

of articulation with pygofer. Pygofer dorsoapical margin incised to near mid-length; basolateral

membranous cleft present; macrosetae well differentiated into several rows. Subgenital plates free from

each other; usually articulated with valve; without macrosetae or with macrosetae scattered, irregularly

arranged, uniseriate laterally, with two lateral rows of macrosetae, with some irregularly arranged

macrosetae near lateral margin, or macrosetae uniseriate, distant from lateral margin. Style broadly

bilobed basally; median anterior lobe pronounced. Basal processes of the aedeagus/connective absent

or reduced. Aedeagus without basal hinge; with a single shaft and gonopore. Connective anterior arms

somewhat divergent, Y- or U-shaped; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex or not strongly convex; dorsal sculpturing pattern imbricate (with overlapping

scales); sculpturing reaching dorsal margin; without distinctly delimited ventroapical sculpturing.

Second valvula broad, gradually tapered; often with dorsal median tooth; teeth on apical 1/3 or more;

teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Palearctic, Nearctic and Oriental regions. The Athysamis-group genera feed on a variety of

woody or herbaceous dicots, grasses or sedges.

Remarks

This informal genus group contains 34 genera, including genera previously included in various subtribes

established for the Palearctic fauna by Emeljanov (1999) and Dmitriev (2006b). Our phylogenetic

analyses suggest that some members of Allygidiina (e.g., Allygidius ) and Platymetopiina (e.g.,

Thamnotettix) are closely related to Athysanus and/or other genera previously placed in Athysanina

(e.g., Euscelis). Additionally, the circumscription of Platymetopiina by Dmitriev (2006b) contains

genera that are not closely related to each other based on those included in our analyses. Thus, the

previous subtribal classification is abandoned here. Also, because our analyses placed Anoterostemma in

this group, Anoterostemmini, which has previously been placed in Cicadellinae (Oman et al. 1990) or as

European Journal of Taxonomy 45: 1-211 (2013)

a subtribe of Limotettigini (Dmitriev 2002), is here synonymized with Athysan ini . The representatives

of our proposed Athysanus group were recovered as a monophyletic group in some, but not all, of

our phylogenetic analyses and this group is diagnosable based on the morphological features listed

above. More taxon sampling and more data are needed to further test the monophyly of this group.

Non-Palearctic genera conforming to the above diagnosis but not considered by Emeljanov (1999) and

Dmitriev (2006b) are also included in the genus group here.

Selected references

Emeljanov (1999), Dmitriev (2006b).

Included genera

Albicostella Ishihara, 1953

Allygidius Ribaut, 1948 placement nov. (previously placed in Allygidiina)

Allygus Fieber, 1872 placement nov. (previously placed in Allygidiina)

Anoterostemma Low, 1885 placement nov. (previously placed in Anoterostemmina)

Artianus Ribaut, 1942

Athysanus Burmeister, 1838

Bilusius Ribaut, 1942

Bobacella Kusnezov, 1929

Condylotes Emeljanov, 1959 placement nov. (transferred from Paralimnini)

Conosanus Osborn & Ball, 1902

Coulinus Beime, 1954

Doliotettix Ribaut, 1942

Ederranus Ribaut, 1942

Euscelidius Ribaut, 1942

Euscelis Brulle, 1832

Extrusanus Oman, 1949

Graphocraerus Thomson, 1869 placement nov. (previously placed in Allygidiina)

Handianus Ribaut, 1942

Hesium Ribaut, 1942

Laburrus Ribaut, 1942

Macustus Ribaut, 1942

Matsumurella Ishihara, 1953

Mimallygus Ribaut, 1948

Mimohardya Zachvatkin, 1946

Phaeida Emeljanov, 1962

Pithy otettix Ribaut, 1942

Rhytistylus Fieber, 1875

Sotanus Ribaut, 1942

Speudotettix Ribaut, 1942

Stictocoris Thomson, 1869 placement nov. (previously placed in Allygidiina)

St rep tan us Ribaut, 1942

Syringitis Emeljanov, 1966 placement nov. (previously placed in Allygidiina)

Thamnotettix Zetterstedt, 1838 placement nov. (previously placed in Platymetopiina)

Transcaucasica Kocalc, 1981

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Bahitini tribe nov.

urn:lsid:zoobank.org:act:92CF5376-39E5-4B5B-B478-B8BB9AA95917

Fig. 12

Type genus: Bahita Oman, 1938.

Diagnosis

Bahitini are medium to large sized, robust leafhoppers, often brown in color. They can be identified

by the head as wide as or wider than the pronotum, crown usually short and broad and often with the

anterior margin distinctly upturned, raised, or with several transverse carinae, the clypellus widening

apically, parallel-margined, or distinctly tumid and tapering apically, the forewing often with strongly

pigmented reflexed costal veins, the pygofer often with processes, and the aedeagus often with apical

processes.

Description

Head. Head subequal to or wider than pronotum; head usually short and broad. Discal portion of crown

glabrous with radial or longitudinal striae. Anterior margin of head with one to several transverse carinae

or striae, or shagreen; usually upturned or distinctly elevated above discal portion of crown. Frontoclypeus

not tumid or somewhat tumid; texture shagreen. Clypellus tapering apically, parallel-sided, or widening

apically; sometimes bulbous and tumid; apex following or slightly surpassing normal curve of gena.

Forum subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral

(lower) corners of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely incised laterally;

with fine erect seta beside laterofrontal suture. Antennal ledges absent or weakly developed (carinate or

weakly carinate). Ocelli present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins not

raised; veins often heavily marked with brown coloration; often with or sometimes without pigmented

reflexed costal veins; sometimes with extra reflexed costal veins; A1-A2 crossvein absent or present;

apical venation not highly reticulate.

Fegs. Pro femur with AMI seta only or rarely with an extra macroseta basad of AMI; intercalary row

with one row of five or more fine setae; row AV with short, stout setae. Protibia dorsal surface rounded,

convex. Metafemur apex macrosetae 2+2+1. Metatarsomere I not expanded apically; plantar setae

simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point.

Pygofer basolateral membranous cleft present; macrosetae well differentiated into several rows; often

with one or more spines or processes arising ventrally. Subgenital plates free from each other; articulated

with valve; macrosetae uniseriate laterally, sometimes with few other scattered macrosetae. Style

broadly bilobed basally; median anterior lobe pronounced. Aedeagus with single shaft and gonopore;

often with pair of apical processes. Connective anterior arms somewhat divergent, Y-shaped, or rarely

with anterior arms closely appressed (e.g., Menosoma); articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern strigate; sculpturing reaching dorsal margin;

without distinctly delimited ventroapical sculpturing. Second valvula broad, gradually tapered; with or

without dorsal median tooth; teeth on apical 1/3 or more; teeth small, regularly or irregularly shaped.

European Journal of Taxonomy 45: 1-211 (2013)

Fig. 12. Bahitini. A-B, D-I. Bahita infuscata (Osborn, 1923). C. Bahita mimica Linnavuori & DeLong,

1978. A-I. Standard views (see Material & Methods).

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Geography and ecology

Distribution: Neotropical andNearctic regions. This group is particularly diverse in Neotropical forested

habitats.

Remarks

Bahitini contains 25 genera and 166 species. This is an exclusively New World group as currently

understood. Although the structure of the head, particularly the presence of more than one transverse

carina or striation at the anterior margin of the crown, gives some members of this group a superficial

resemblance to the Old World tribe Selenocephalini, our phylogenetic analyses did not indicate a close

relationship between these groups. Some other New World athysanine genera are similar in having the

anterior margin of the head upturned and with reflexed costal veins (e.g., Lajolla , Bardana , Acunasus )

but these are generally more elongate, more slender and/or have a more produced head which might

suggest that they are more related to the Platymetopius/Twiningia group. More detailed analyses of the

relationships of the genera of Athysanini may indicate whether some of these should be included into this

tribe. Benibahita was included in the Bahita- group by Linnavuori & DeLong (1978b) but is excluded

here, as it appears to be more closely related to Yungasia (Athysanini; similar color pattern, with head

narrower than pronotum, more slender in shape smaller in size than Bahitini). Angubahita DeLong is

excluded based on its small size and absence of other characteristics of the group. The phylogenetic

analyses presented here resolved a monophyletic group including representatives of Bahita , Caruya ,

Kinrentius, Menosoma , Taperinha, and Oxycephalotettix. Further taxonomic and phylogenetic study of

the Neotropical deltocephaline fauna may refine the circumscription of this group.

Selected references

Linnavuori & DeLong (1978b), Zanol (1999).

Included genera

Bahita Oman, 1938 placement nov. (transferred from Athysa nin i)

Brincadorus Oman, 1938 placement nov. (transferred from Athysanini)

Caruya Linnavuori & DeLong, 1978 placement nov. (transferred from Athysanini)

Chonosina Linnavuori & DeLong, 1978 placement nov. (transferred from Athysanini)

Concepciona Linnavuori & DeLong, 1977 placement nov. (transferred from Athysanini)

Frequenamia DeLong, 1947 placement nov. (transferred from Athysanini)

Hecaloidia Osborn, 1923 placement nov. (transferred from Athysanini)

Huachia Linnavuori, 1959 placement nov. (transferred from Athysanini)

Ilagia Kramer & DeLong, 1968 placement nov. (transferred from Athysa nini )

Kinrentius Wei, Dietrich & Webb, 2010 placement nov. (previously unplaced in Deltocephalinae)

Kyra Zanol, 2011 placement nov. (transferred from Athysanini)

Megabahita Linnavuori & DeLong, 1978 placement nov. (transferred from Athysanini)

Menosoma Ball, 1931 placement nov. (transferred from Athysanini)

Oxycephalotettix Zahniser, 2005 placement nov. (transferred from Athysanini)

Parabahita Linnavuori, 1959 placement nov. (transferred from Athysanini)

Perubahita Linnavuori & DeLong, 1978 placement nov. (transferred from Athysanini)

Rineda Linnavuori & DeLong, 1978 placement nov. (transferred from Athysa nin i)

Saadevra McKamey, 2003 placement nov. (transferred from Athysanini)

Sanestebania Linnavuori & DeLong, 1978 placement nov. (transferred from Athysanini)

Serridonus Linnavuori, 1959 placement nov. (transferred from Athysanini)

Sinchonoa Linnavuori & DeLong, 1978 placement nov. (transferred from Athysanini)

Suarezilinna McKamey, 2003 placement nov. (transferred from Athysanini)

Taperinha Linnavuori, 1959 placement nov. (transferred from Athysanini)

European Journal of Taxonomy 45: 1-211 (2013)

Tingopyx Linnavuori & DeLong, 1978 placement nov. (transferred from Athysanini)

Yuraca Linnavuori & DeLong, 1978 placement nov. (transferred from Athysanini)

Bonaspeiini tribe nov.

um:lsid:zoobank.org:act:50F524A4-754D-4EA4-8466-98A20D4DFA84

Fig. 13

Type genus: Bonaspeia Linnavuori, 1961.

Diagnosis

Bonaspeiini are medium sized, broad, robust, brachypterous to submacropterous, ivory, greenish, or

brownish leafhoppers. They can be identified by the anterior margin of the head without sharp carinae

(with sharp carinae in Selenocephalini), brachypterous to submacropterous forewings (exception:

Renosteria ), hind wings when present narrowed anteriorly-posteriorly when outstretched or otherwise

reduced in size (exception: Renosteria ), connective U- or sometimes Y- shaped with stem often very

short or absent, male segment X broad and sclerotized, and ovipositor usually distinctly protruding

beyond the pygofer apex.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae. Anterior margin of head shagreen, irregularly textured, or with several to numerous

transverse striae. Frontoclypeus not tumid; texture shagreen, glabrous, or rugose. Clypellus parallel

sided, tapering apically, or widening apically; apex following or slightly surpassing normal curve of

gena. Lorum subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral

(lower) corners of eyes or near upper or anterodorsal corners of eyes. Antennae short, less than 1.5 x

width of head. Gena incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledges

weakly developed (carinate or weakly carinate). Ocelli often reduced in size; close to or distant from

eyes; on anterior margin of head or shifted slightly dorsad on crown.

Thorax. Pronotum lateral margin not carinate or carinate; lateral margin shorter than basal width of eye.

Wings. Forewing brachypterous to submacropterous, or rarely macropterous (. Renosteria ); with 3

anteapical cells; veins not raised; with or without reflexed costal veins; A1-A2 crossvein absent or

present; apical venation not highly reticulate. Hind wing, when present, narrowed anteriorly-posteriorly

when outstretched or otherwise reduced in size; with full complement of wing veins or with some veins

(e.g., M3+4 and CuA) fused.

Legs. Pro femur with AMI seta only; intercalary row with one row of 5 or more fine setae; row AV with

short, stout macrosetae or with relatively long macrosetae. Protibia dorsal surface rounded, convex.

Metafemur apex macrosetae 2+2+1. Metatarsomere I not expanded or expanded apically; plantar setae

all simple or rarely (e.g., some Gcaleka ) one or more platellae.

Male genitalia. Valve articulated with pygofer; with short point of articulation with pygofer. Pygofer

dorsoapical margin incised to near mid-length or nearly to base; basolateral membranous cleft present,

sometimes sharply angulate; macrosetae absent or reduced (< two rows) or rarely well differentiated

into several rows; often with spine-like process. Subgenital plates free from each other; articulated

with valve; macrosetae scattered, irregularly arranged, uniseriate laterally, with two or more lateral

rows of macrosetae, or with some irregularly arranged macrosetae near lateral margin. Style linear,

median anterior lobe not pronounced or broadly bilobed basally, median anterior lobe pronounced.

Basal processes of the aedeagus/connective absent or reduced or present, connected or articulated to

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 13. Bonaspeiini. A-I. Standard views (see Material & Methods), Bonaspeia attia Davies, 1987.

J-K. Hind wing. J. Cerus goudanus Theron, 1984. K. Curvostylus chloridulus (Naude, 1926).

European Journal of Taxonomy 45: 1-211 (2013)

connective or near base of aedeagus. Aedeagus without basal hinge; with a single shaft and gonopore;

often with short hooks or spines. Connective anterior arms somewhat divergent, U- or sometimes Y-

shaped; stem often very short or absent; articulated with aedeagus. Segment X broad, sometimes very

large and occupying much of dorsal part of pygofer; sclerotized dorsally and laterally.

Female genitalia. Pygofer with macrosetae reduced or absent or with numerous macrosetae. Ovipositor

not protruding or protruding far beyond pygofer apex. First valvula convex, dorsal sculpturing pattern

strigate, concatenate, or reticulate; sculpturing reaching dorsal margin; without distinctly delimited

ventroapical sculpturing. Second valvula broad, gradually tapered; with or without dorsal median tooth;

teeth on apical 1/3 or more; teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Afrotropical (South Africa). Bonaspeiini are co mm on in the fynbos biome and collected

on woody and herbaceous dicots. Gcaleka species are dorsoventrally flattened, apparently an adaptation

to feeding on their reed-like hosts in the family Restionaceae. Salsolibia and Salsocolila feed on the

ganna bush, Salsola esterhuyseniae Botsch. (Amaranthaceae), a woody-stemmed bush with succulent

leaves living in silt deposits in valleys and riverbeds. The squat, compact leafhoppers apparently mimic

the succulent leaves of their host (Theron 1979). Renosteria , Cerus and Refrolix feed on renosterbos,

Elytropappus rhinocerotis Less. (Asteraceae), a common plant in the southern part of the Western Cape

Province. Davies (1988a) reports Kimbella acusiculcita Davies, 1988 from Erica peltata Andrews

(Ericaceae) and Colistra parvulus (Linnavuori, 1961) from Erica hispudula L. and Erica muscosa

(Aiton) E.G.H.Oliv.

Remarks

Bonaspeiini contains 21 genera and 108 species. The tribe is closely related to Selenocephalini but is

considered distinct here due to the lack of distinct carinae on the anterior margin of the head which are

typical of most Selenocephalini and the presence of a number of unique characters (see Diagnosis).

Three exemplars (from Bonaspeia, Cerus , and Renosteria) were included in the phylogenetic analyses

here, and the group was resolved as paraphyletic with respect to the clade Selenocephalini + Dagama

(Athysanini). However, branch support was not high, and more detailed phylogenetic analyses are

needed to further resolve these relationships. A representative of Gcaleka was included in previous

analyses (Zahniser & Dietrich 2010) and was resolved as sister to Bonaspeia , but was excluded from

analyses here due to missing data in the 28S sequence.

Selected references

Linnavuori (1961), Theron (1974, 1984a, b), Stiller (1986), Davies (1987, 1988a, b).

Included genera

Basutoia Linnavuori, 1961 placement nov. (transferred from Athysanini)

Bloemia Theron, 1974 placement nov. (transferred from Athysanini)

Bonaspeia Linnavuori, 1961 placement nov. (previously unplaced in Deltocephalinae)

Caffrolix Linnavuori, 1961 placement nov. (transferred from Athysanini)

Capeolix Linnavuori, 1961 placement nov. (transferred from Athysanini)

Cerus Theron, 1975 placement nov. (previously unplaced in Deltocephalinae)

Colistra Davies, 1988 placement nov. (transferred from Athysanini)

Curvostylus Davies, 1987 placement nov. (transferred from Athysanini)

Gcaleka Naude, 1926 placement nov. (previously unplaced in Deltocephalinae)

Hadroca Theron, 1974 placement nov. (previously unplaced in Deltocephalinae)

Johanus Theron, 1974 placement nov. (transferred from Athysanini)

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Kaapia Theron, 1983 placement nov. (transferred from Athysanini)

Kimbella Davies, 1988 placement nov. (transferred from Athysanini)

Megaulon Theron, 1975 placement nov. (transferred from Pa.ra.1 imnin i)

Proekes Theron, 1975 placement nov. (transferred from Stenometopiini)

Proekoides Stiller, 1986 placement nov. (transferred from Stenometopiini)

Refrolix Theron, 1984 placement nov. (transferred from Athysa nin i)

Renosteria Theron, 1974 placement nov. (transferred from Athysanini)

Salsocolila McKamey, 2003 placement nov. (transferred from Athysanini)

Salsolibia Theron, 1979 placement nov. (transferred from Athysanini)

Tzitzikamaia Linnavuori, 1961 placement nov. (transferred from Athysanini)

Chiasmini Distant, 1908

Fig. 14

Type genus: Chiasmus Mulsant & Rey, 1855.

= Doratur ini Emeljanov, 1962.

= Listrophorini Boulard, 1971 syn. nov.

= Paraphrodini Linnavuori, 1979.

Diagnosis

Chiasmini are small to medium sized leafhoppers, usually white, stramineous, green, brown, grey,

or black in coloration, and sometimes iridescent. They can be identified by the tapering or parallel¬

sided clypellus, aedeagus hinged at the base (hinge usually but not always present), ovipositor usually

extending far beyond the pygofer, first valvula dorsal sculpturing pattern maculose to granulose and

usually submarginal, first valvula without distinctly delimited ventroapical sculpturing, and second

valvula teeth obliquely triangular and serrated.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae or shagreen. Anterior margin of head shagreen, glabrous, irregularly textured, or

foliaceous. Frontoclypeus not tumid; texture shagreen or glabrous. Clypellus parallel-sided or tapering

apically; apex following or slightly surpassing normal curve of gena. Lorum wider than, subequal to,

or distinctly narrower than clypellus near base. Antennal bases near middle or posteroventral (lower)

comers of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely incised or strongly incised

(nearly forming right angle) laterally; with or without ( Baileyus , Gurawa , Listrophora , some Chiasmus)

fine erect seta beside laterofrontal suture. Antennal ledge absent or weakly developed (carinate or weakly

carinate). Ocelli absent, reduced, or present; close to or distant from eyes; on anterior margin of head or

on crown {Chiasmus).

Thorax. Pronotum lateral margin carinate or not carinate; lateral margin shorter than basal width of eye.

Wings. Fore wing brachypterous to macropterous; if macropterous then appendix large and extending

around wing apex; with 3 anteapical cells or with 2 anteapical cells; veins not raised or veins distinctly

raised or carinate ( Baileyus , Gurawa , Paraphrodes ); without reflexed costal veins; A1-A2 crossvein

absent.

Legs. Pro femur with AMI seta only or rarely with an additional proximal seta ( Doraturopsis Lindberg,

Protochiasmus Zahniser); intercalary row with one row of five or more fine setae or intercalary row

reduced or absent; row AV with short, stout setae or with relatively long macrosetae. Protibia dorsal

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Fig. 14. Chiasmini. Chiasmus sp. (see note under Chiasmini on identification of species of Chiasmus).

A-I. Standard views (see Material & Methods).

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

surface rounded, convex. Metafemur apex macrosetae 2+1, 2+2, 2+1+1, or 2+2+1. Metatarsomere I not

expanded apically or expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point.

Pygofer basolateral membranous cleft present or absent, not membranous; macrosetae absent, reduced

(< two rows) or well differentiated into several rows. Subgenital plates free from each other; articulated

with valve; macrosetae absent or present, scattered, irregularly arranged, or uniseriate laterally. Style

broadly bilobed basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective

absent or reduced. Aedeagus often with or sometimes without ( Baileyus , Leofa , Gurawa, Paraphrodes)

basal hinge; with a single shaft and gonopore. Connective anterior arms somewhat divergent, Y- or

U-shaped, or anterior arms closely appressed anteriorly; articulated with aedeagus.

Female genitalia. Pygofer with macrosetae reduced or absent or with numerous macrosetae. Ovipositor

usually protruding far beyond pygofer apex (not protruding in Baileyus , Gurawa , Omaranus ,

Paraphrodes). First valvula not strongly convex; dorsal sculpturing pattern granulose or maculose;

sculpturing submarginal; without distinctly delimited ventroapical sculpturing. Second valvula abruptly

broadened medially or subapically; without dorsal median tooth; dorsal teeth present on apical 1/3 or

more; teeth obtusely triangular and serrated.

Geography and ecology

Distribution: cosmopolitan. Nearly all species are grass or sedge feeders, and they are typical members

of the grassland faunas of the world. Driotura spp. have been collected from Asteraceae. Nephotettix

virescens (Distant, 1908) and some other Nephotettix spp. are agricultural pests and transmit tungro

vims of rice in southeast Asia, and can cause severe economic loss.

Remarks

Chiasmini contains 21 genera and 317 species. The tribe appears to be most closely related to

Stenometopiini (Zahniser 2008a; Zahniser & Dietrich 2010) with which it shares some similarities of

the ovipositor and male pygofer. Phylogenetic analyses sometimes resolve it as sister to Stenometopiini,

and also suggest that the two tribes are related to Eupelicini, Drakensbergenini, and Evinus. In the

phylogenetic analyses here, the tribe was resolved as monophyletic in ML and Bayesian analyses, but

in parsimony analyses it was monophyletic with the exception of Protochiasmus , which was resolved,

sister to Evinus but with little or no branch support. The placement of Gurawa in Chiasmini is supported

by these analyses and the morphologically similar Baielyus and Paraphrodes (Zahniser 2008a, 2011)

are also included.

For the first time, molecular data for Listrophora were included in phylogenetic analyses and the genus

was found with very strong branch support to be related to Chiasmini. The morphological characters of

the male and female genitalia of Listrophora (aedeagus hinged, first valvula dorsal sculpturing pattern

granulose and submarginal, second valvula with distinct serrated oblong and triangular teeth) match

those of other Chiasmini, despite the bizarre and unique diamond-shaped process of the head, which led

Boulard (1971) to place it in a separate tribe. In our analyses it was found to be closely related to Gurawa

within Chiasmini, with which it shares the raised or carinate veins of the forewing. Listrophorina was

previously included as a subtribe of Eupelicini (Zahniser & Dietrich 2010) but is synonymized here with

Chiasmini.

Nephoris is transferred to the tribe based on the figures published by Jacobi (1912). N. chalybaea Jacobi,

1912 appears similar to Leofa thompsoni Zahniser, 2008 which was also described from central Africa.

Attempts to locate the type were unsuccessful, but for now it is considered a valid genus pending further

study.

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Selected references

Vilbaste (1965), Ross (1968), Boulard (1971),Blocker (1983), Blocker & Johnson (1988a, b, 1990a, b,

c), Emeljanov (1999), Dmitriev (2003), Zahniser & Hicks (2007), Zahniser (2008a, b, 2011), Duan et

al. (2009), Duan & Zhang (2012a, b, c).

Notes on illustration (Fig. 14)

The specimens figured are identified as Chiasmus sp. from W. Cape Province, South Africa. The species

identity of Chiasmus specimens is difficult to determine given the variability in some described species

and because of the relatively uniform male genitalia among species. C. varicolor (Kirkaldy, 1906) is

known from Australia and South Africa, C. undulatus Theron, 1982 is also known from South Africa,

and C. katonae (Melichar, 1908) is known from Tanzania. The species figured is similar to C. katonae

but differs in size, surface shape of the crown, and coloration. The genus should be revised and species

concepts reevaluated to improve the taxonomy and allow for easier identification.

Included genera

Aconura Lethierry, 1876

Aconurella Ribaut, 1948

Athysanella Baker, 1898

Baileyus Singh-Pruthi, 1930

Chiasmus Mulsant & Rey, 1855

Doratura Sahlberg, 1871

Doraturopsis Lindberg, 1935

Driotura Osborn & Ball, 1898

Exitianus Ball, 1929

Gurawa Distant, 1908

Icaia Linnavuori, 1973

Leofa Distant, 1918

Listrophora Boulard, 1971 placement nov. (previously placed in Eupelicini: Listrophorina)

Nephoris Jacobi, 1912 placement nov. (transferred from Athysanini)

Nephotettix Matsumura, 1902

Omar anus Distant, 1918

Paraphrodes Linnavuori, 1979

Picchusteles Linnavuori & DeLong, 1976

Protochiasmus Zahniser, 2010

Stenogiffardia Evans, 1977

Zahniserius Duan & Zhang, 2012

Cicadulini Van Duzee, 1892

Fig. 15

Type genus: Cicadula Zetterstedt, 1840.

= Procepitina Dmitriev, 2002 syn. nov.

Diagnosis

Cicadulini are small to medium sized, slender, stramineous, yellow, or greenish leafhoppers, sometimes

with the anterior margin of the head marked with black spots. They can be identified by the male segment

X often long and strongly sclerotized, and subgenital plates sometimes with a row of macrosetae near

the middle and with long fine setae laterally.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown shagreen. Anterior margin of

head shagreen. Frontoclypeus not tumid; texture shagreen. Clypellus widening apically; apex following

or slightly surpassing normal curve of gena. Lorum subequal to or wider than clypellus near base.

Antennal bases near middle or posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x

width of head or long, 1.5 x width of head or longer. Gena obtusely incised laterally; with fine erect

seta beside laterofrontal suture. Antennal ledge absent. Ocelli present; close to or distant from eyes; on

anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Fore wing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins not

raised; without reflexed costal veins; A1-A2 crossvein absent.

Legs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1.

Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Sternal apodemes of abdominal segments I and II usually well-developed. Valve

articulated with pygofer; lateral margin short, articulating with pygofer at a point. Pygofer basolateral

membranous cleft present or absent, not membranous; macrosetae well differentiated into several

rows. Subgenital plates free from each other; articulated with valve; macrosetae uniseriate laterally or

uniseriate, distant from lateral margin. Style broadly bilobed basally, median anterior lobe pronounced.

Basal processes of the aedeagus/connective absent or reduced. Aedeagus with single shaft and gonopore.

Connective anterior arms somewhat divergent, Y-shaped; articulated with aedeagus. Segment X long,

length usually 2x width or longer; strongly sclerotized dorsally.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern imbricate (with overlapping scales); sculpturing

reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula broad,

gradually tapered; without dorsal median tooth; dorsal teeth on apical 1/3 or more; teeth small, regularly

or irregularly shaped.

Geography and ecology

Distribution: Nearctic, Neotropical and Palearctic regions. Cicadulini feed on sedges and grasses, and

are often found in association with wetland or marsh habitats or in forest understories.

Remarks

Cicadulini contains 14 genera and 114 valid species. This group was considered a subtribe of Athysanini

by Dmitriev (2006b) and Emelj anov (1999) included cicaduline genera in his Athysanina. Two exemplars

of the tribe (Cicadula and Elymana) were included in a phylogenetic analysis of Deltocephalinae

(Zahniser & Dietrich 2010) and were supported as a monophyletic group. Molecular data for Proceps

and Stenometopiellus included here for the first time suggest with strong branch support that they belong

in Cicadulini; they are transferred to the tribe here. Dmitriev (2002) described the monotypic Procepitina

as a subtribe of Scaphytopiini, but it is considered a synonym of Cicadulini here. The characters of the

ovipositor of Proceps are consistent with those of other Cicadulini; males were not available for this

study but Emelj anov (1967) described male segment X as long and well sclerotized, consistent with

other members of the tribe. Cicadulini is considered a tribe separate from Athysanini following Oman

et al. (1990). It appears to be related to other grass-feeding tribes of Deltocephalinae and perhaps to

Limotettigini, but more data are needed to more firmly resolve its relationship to other tribes.

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Fig. 15. Cicadulini. A-C. Cicadula melanogaster (Provancher, 1872). D-I. Cicadula quadrinotata

(Fabricius, 1794). A-I. Standard views (see Material & Methods).

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Calanana was listed in Athysanini by Oman et al. (1990). However, it was clearly suggested to be

related to Cicadula , Elymana , and Paluda by Oman (1949) with which it shares the long sclerotized

segment X and the subgenital plate with a row of macrosetae running medially along the plate. It is

therefore placed in Cicadulini here.

Selected references

Hamilton (1975b), Dmitriev (2002).

Included genera

Calanana DeLong, 1945 placement nov. (transferred from Athysanini)

Cicadula Zetterstedt, 1840

Dudanus Dlabola, 1956

Elymana DeLong, 1936

Hecadula Dietrich & Rakitov, 2002

Knullana DeLong, 1941

Mocydia Edwards, 1922

Mocydiopsis Ribaut, 1939

Morinda Emeljanov, 1972

Paluda DeLong, 1937

Proceps Mulsant & Rey, 1855 placement nov. (previously in Scaphytopiini: Procepitina)

Rhopalopyx Ribaut, 1939

Stenometopiellus Haupt, 1917 placement nov. (transferred from Athysanini)

Taurotettix Haupt, 1929

Cochlorhinini Oman, 1943

Fig. 16

Type genus: Cochlorhinus Uhler, 1876.

Diagnosis

Cochlorhinini are medium sized leafhoppers, with ground color greenish, yellow, brown, black, or ivory.

They can be identified by the Y-shaped connective fused to aedeagus, male pygofer with one or more

pairs of teeth, spines, or processes, male pygofer side excavated medially and scooplike, male segment

X long and sclerotized laterally and sometimes ventrally and/or dorsally, and second valvulae with a

median dorsal tooth.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae. Anterior margin of head shagreen, glabrous, irregularly textured, or foliaceous.

Frontoclypeus not tumid; texture shagreen. Clypellus widening apically; apex following or slightly

surpassing normal curve of gena. Lorum subequal to or wider than clypellus near base. Antennal bases

near middle or posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x width of head.

Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledge absent.

Ocelli present; close to or distant from eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous or submacropterous; appendix restricted to anal margin; with 3 anteapical

cells; veins not raised; without reflexed costal veins; A1-A2 crossvein absent.

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Fig. 16. Cochlorhinini. Cochlorhinuspluto Uhler, 1876. A-I. Standard views (see Material & Methods).

H-I. Aedeagus shown with connective fused at base.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Legs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae; row AV with

short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apexmacrosetae 2+2+1. Metatibia

row PD macrosetae often long, nearly equal to or longer than half length of protibia. Metatarsomere I not

expanded apically; plantar setae with one or more platellae or without platellae.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae well differentiated into several rows;

dorsally excavated to mid-length or nearly to base; side excavated medially, scooplike; usually with

apical or subapical teeth, spines, or processes. Subgenital plates free from each other; articulated with

valve; macrosetae uniseriate laterally or irregularly arranged near lateral margin. Style broadly bilobed

basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective present (paired or

singular) or absent. Aedeagus with single shaft and gonopore; symmetrical or asymmetrical; sometimes

dorsoventrally flattened or lamellate; sometimes with 2 or 3 pairs of apical processes. Connective anterior

arms somewhat divergent, Y- or U-shaped; fused to aedeagus. Segment X relatively long, sclerotized

laterally, and sometimes sclerotized ventrally and/or dorsally.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern strigate, concatenate, or reticulate; sculpturing

reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula broad,

gradually tapered; usually with dorsal median tooth; dorsal teeth on apical 1/3 or more; teeth small,

regularly or irregularly shaped.

Geography and ecology

Distribution: Nearctic (western US), Neotropical ( Cochlorhinuspluto Uhler, 1876, introduced in Chile).

All are endemic to the western United States and are collected in grasslands, meadows, and forest edges

and understories.

Remarks

Cochlorhinini contains 12 genera and 145 species. Phylogenetic analyses recovered Ballana as sister to

Cochlorhinus with moderate to strong branch support. Based on this result and on shared morphological

characters (see Diagnosis above) and a western North American distribution, Allygianus , Allygiella ,

Ballana , Gloridonus , Limb anus, and Pasadenus are transferred to Cochlorhinini here. These genera do

not possess some characters that previously helped to distinguish a more strict interpretation of the tribe

(sensu Oman 1949): ocelli distant from eyes; metatarsomere I with some platellae. However the overall

evidence from the molecular data, morphological observations (particularly the fused connective and

aedeagus), and distribution support the placement of these genera in the tribe.

Cochlorhinini is potentially related to Koebeliini, Mukariini, Vartini or some Athysanini, but more data

are needed to resolve its relationship to other tribes.

Selected references

Beamer (1940), Oman (1949), DeLong (1964).

Included genera

Allygianus Ball, 1936 placement nov. (transferred from Athysanini)

Allygiella Oman, 1949 placement nov. (transferred from Athysa nin i)

Ballana DeLong, 1936 placement nov. (transferred from Athysanini)

Calonia Beamer, 1940

Cochlorhinus Uhler, 1876

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Drionia Ball, 1915

Eulonus Oman, 1949

Gloridonus Ball, 1936 placement nov. (transferred from Athysanini)

Huleria Ball, 1902

Limbanus Oman, 1949 placement nov. (transferred from Athysanini)

Pasadenus Ball, 1936 placement nov. (transferred from Athysanini)

Penehuleria Beamer, 1934

Deltocephalini Dallas, 1870

Fig. 17

Type genus: Deltocephalus Burmeister, 1838.

Diagnosis

Deltocephalini are small to medium sized leaflioppers and are variable in color. They can be identified

by the tapering or parallel-sided clypellus, narrow lorum, linear connective with anterior arms closely

appressed, connective fused to the aedeagus, and first valvula dorsal sculpturing imbricate.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae or shagreen. Anterior margin of head shagreen, rounded or angled to face. Frontoclypeus

not tumid; texture shagreen. Clypellus parallel-sided or tapering apically; apex following or slightly

surpassing normal curve of gena. Lorum distinctly narrower than clypellus near base. Antennal bases

near middle or posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x width of head.

Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledge absent.

Ocelli present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin not carinate; lateral margin shorter than basal width of eye.

Wings. Forewing brachypterous to macropterous; appendix absent, reduced, or present and restricted

to anal margin; with 3 anteapical cells; veins not raised; without reflexed costal veins; A1-A2 crossvein

usually absent.

Legs. Profemur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1.

Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer or rarely fused to subgenital plates (. Miradeltaphus ,

Yuanamia ); lateral margin short, articulating with pygofer at a point. Pygofer basolateral membranous

cleft present; macrosetae well differentiated into several rows. Subgenital plates free from each other;

articulated with valve or rarely fused to each other (. Miradeltaphus , Yuanamia :); macrosetae uniseriate

laterally. Style broadly bilobed basally, median anterior lobe pronounced. Basal processes of the

aedeagus/connective absent or reduced or present, connected or articulated to connective or near base

of aedeagus. Aedeagus with single shaft and gonopore. Connective anterior arms closely appressed

anteriorly, linear-shaped; fused to aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex or not strongly convex; dorsal sculpturing pattern imbricate (with overlapping

scales); sculpturing reaching dorsal margin; without distinctly delimited ventroapical sculpturing.

Second valvula broad, gradually tapered; without dorsal median tooth; dorsal teeth on apical 1/3 or

more; teeth small, regularly or irregularly shaped.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 17. Deltocephalini. Deltocephalus pulicaris (Fallen, 1806). A-I. Standard views (see Material &

Methods). G. Connective and style shown with aedeagus fused to connective. H. Aedeagus shown with

connective and style.

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Geography and ecology

Distribution: cosmopolitan. Deltocephalini feed on grasses and sedges and are diverse and abundant

in grassland ecosystems. Several species are economically important as vectors of disease agents.

Graminella nigrifrons (Forbes, 1885), Maiestas dorsalis (Motschulsky, 1859) and Endria inimica (Say,

1830) are vectors of pathogens of maize, rice, and wheat, respectively.

Remarks

Deltocephalini contains 68 genera and 587 species. Morphological and molecular phylogenetic

analyses to date suggest that it is monophyletic and closely related to Paralimnini, but further taxon

sampling in these large tribes and more data are needed. The two tribes together appear to be related to

Tetartostylini which also possesses the tapered clypellus and linear connective. Many genera and species

of Deltocephalini are morphologically quite similar to each other, and some genera, e.g., Amplicephalus

and Polyamia , are probably artificial. Revisions of many genera and identification keys, especially of the

Neotropical fauna, are needed. As presently defined, the tribe encompasses the “Deltocephalus- group”

of genera recognized by Fang et al. (1993, 1995) and Webb & Viraktamath (2009), but is considerably

narrower than the concepts of Deltocephalini employed by Oman (1949) and Linnavuori (1959).

Selected references

Oman (1949), Linnavuori (1959), Fang et al. (1993, 1995), Blocker et al. (1995), Dmitriev (2004b),

Webb & Viraktamath (2009).

Included genera

Afrosus Linnavuori, 1959

Alobaldia Emeljanov, 1972

Alodeltocephalus Evans, 1966

Amblysellus Sleesman, 1929

Amplicephalus DeLong, 1926

Arundanus DeLong, 1935

Bolarga Oman, 1938

Bolivaia Linnavuori & DeLong, 1979

Cabrellus Emeljanov, 1964

Cabrulus Oman, 1949

Crumbana Oman, 1949

Cruziella Linnavuori & DeLong, 1979

Ctenurellina McKamey, 2003

Daltonia Oman, 1949

Deltanus Oman, 1949

Deltazotus Kramer, 1971

Deltella Oman, 1949

Deltocephalus Burmeister, 1838

Destria Oman, 1949

Endria Oman, 1949

Fusanus Linnavuori, 1955

Graminella DeLong, 1936

Haldorus Oman, 1938

Heidinus Theron, 1988 placement nov. (previously unplaced in Deltocephalinae)

Horouta Knight, 1975

Kanorba Oman, 1938

Kansendria Kramer, 1971

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Limpica Cheng, 1980

Lonatura Osborn & Ball, 1898

Lore liana DeLong & Kolbe, 1975

Loreta Linnavuori, 1959

Lusitanocephalus Quartau, 1970

Maiestas Distant, 1917

Maricaona Caldwell, 1952

Matsumuratettix Metcalf, 1952

Mattogrossus Linnavuori, 1959

Mendozellus Linnavuori, 1959

Mexara Oman, 1949

Miradeltaphus Dash & Viraktamath, 1995

Neodeltocephalus Linnavuori, 1959

Nullamia DeLong, 1970

Ochromelanus Stiller, 2011

Onura Oman, 1938

Paramesodes Ishihara, 1953

Parandanus Linnavuori & DeLong, 1976

Parayuanamia Xing & Li, 2011

Peitouellus Vilbaste, 1969

Picchuia Linnavuori & DeLong, 1979

Planicephalus Linnavuori, 1954

Polyamia DeLong, 1926

Ragia Theron, 1973

Recilia Edwards, 1922

Reventazonia Linnavuori, 1959

Sanctanus Ball, 1932

Sanluisia Linnavuori, 1959

Spathifer Linnavuori, 1955

Tetramelasma Stiller, 2011

Theronns Stiller, 2009

Tideltellus Kramer, 1971

Toldoanus Linnavuori, 1954

Tumupasa Linnavuori, 1959

Unerus DeLong, 1936

Unoka Lawson, 1928

Vicosa Linnavuori & DeLong, 1978

Wakaya Linnavuori, 1960

Warlucephala Lletcher, 2006

Wyushinamia Zhang & Duan, 2006

Yuanamia Zhang & Duan, 2006

Dorycephalini Oman, 1943

Lig. 18

Type genus: Dorycephalus Kouchakewitch, 1866.

Diagnosis

Dorycephalini are medium sized, elongate, dorsoventrally flattened, greenish or ivory to brownish

leafhoppers. They can be identified by the strongly produced, spatulate, and flattened head with anterior

European Journal of Taxonomy 45: 1-211 (2013)

margin foliaceous, long pronotum, raised forewing veins, forewing with two anteapical cells, metafemur

apex macrosetae 2+0, and hind tibia distinctly bent apically. Eupelicini and some Hecalini (particularly

Attenuipyga and Neoslossonia which were previously placed in Dorycephalini) share some of these

characters, but Dorycephalini can be distinguished from Eupelicini by the crown not encroaching

onto eyes as in Eupelicina, head wider and more spatulate-shaped than Paradorydiina, and valve not

fused to the pygofer and style not gracile as in Paradorydiina. From Attenuipyga and Neoslossonia

(Hecalini), Dorycephalini can be distinguished by the forewing veins A1 and A2 separate throughout

their length, male pygofer lobate apically and not distinctly pointed posterodorsally, style apophysis

slightly expanded, rounded apically, aedeagus without processes, female first valvula without distinctly

delimited ventroapical sculpturing, second valvulae not humpbacked dorsally, and second valvulae with

distinct teeth.

Description

Head. Head subequal to or wider than pronotum; elongate; spatulate. Discal portion of crown rugose.

Anterior margin of head foliaceous. Frontoclypeus tumid; texture shagreen. Clypellus parallel-sided or

tapering apically; apex following or slightly surpassing normal curve of gena. Lorum distinctly narrower

than clypellus near base. Antennal bases near upper or anterodorsal corners of eyes. Antennae short, less

than 1.5 x width of head. Gena strongly incised laterally (nearly forming right angle); without fine erect

seta beside laterofrontal suture. Antennal ledges absent. Ocelli present; somewhat distant from eyes; on

crown.

Thorax. Pronotum lateral margin carinate; lateral margin as long as or longer than basal width of eye.

Wings. Forewing submacropterous to brachypterous; appendix absent or reduced; with 2 anteapical

cells; veins distinctly raised or carinate; without reflexed costal veins; A1-A2 crossvein absent; A1 and

A2 separate from each other throughout their length; apical venation not highly reticulate.

Fegs. Profemur with AMI absent or reduced; intercalary row reduced or absent; row AV with

relatively long macrosetae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+0.

Metatarsomere I expanded apically; plantar surface with one or more platellae.

Male genitalia. Apodemes of sternite I long and wide; those of sternite II not well-developed. Valve

articulated with pygofer; with short point of articulation with pygofer. Pygofer dorsoapical margin

incised to near mid-length; basolateral membranous cleft present; macrosetae absent or reduced (< two

rows). Subgenital plates free from each other; articulated with valve; without macrosetae. Style broadly

bilobed basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent.

Aedeagus without basal hinge; with a single shaft and gonopore. Connective anterior arms somewhat

divergent, Y- or U-shaped; articulated with aedeagus.

Female genitalia. Pygofer with macrosetae reduced or absent. Ovipositor not protruding far beyond

pygofer apex. First valvula not strongly convex; dorsal sculpturing pattern granulose or maculose;

sculpturing submarginal; without distinctly delimited ventroapical sculpturing. Second valvula broad,

gradually tapered; without dorsal median tooth; teeth on apical 1/3 or more; teeth small, regularly or

irregularly shaped.

Geography and ecology

Distribution: Palearctic. The two known species feed on grasses. Their elongate, flattened shape appears

to be an adaptation for crypsis and closely appressing against grass blades.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 18. Dorycephalini. Dorycephalus baeri Kouchakewitch, 1866. A-I. Standard views (see Material

& Methods). J. Ventral view of abdominal stemites I-III, showing large apodemes of sternite I {slap)

and apodemes of sternite II (s2ap).

European Journal of Taxonomy 45: 1-211 (2013)

Remarks

Dorycephalini as circumscribed here contains only the type genus and its 2 species. Dorycephalus was

included in molecular phylogenetic analyses here for the first time. It was added to the data set only

after branch support analyses were already completed so it is not present in the trees figured with branch

support but is included in Fig. 6. In ML analyses of the molecular data only (28S + H3) Dorycephalus

was recovered in the grass-specialist clade including Chiasmini, Eupelicini, Stenometopiini, and some

others, and was quite distant from the long branch uniting Hecalus and Attenuipyga. Based on this result

and because it does not share the characters which unite Attenuipyga , Neoslossonia , and Hecalini (see

Diagnosis of Hecalini below), Dorycephalini is retained as a monotypic tribe here, and Attenuipgyga

and Neoslossonia are placed in Hecalini.

Selected references

Oman (1985a), Dmitriev (2001).

Included genera

Dorycephalus Kouchakewitch, 1866

Drabescini Ishihara, 1953

Figs 19, 20

Type genus: Drabescus Stal, 1870.

= Paraboloponini Ishihara, 1953.

= Bhatiini Linnavuori & Al-Ne’Amy, 1983.

Diagnosis

Drabescini are medium sized to large leafhoppers, variable in color and shape. They can be identified by

the antennae situated near the upper part of the face, antennal pits large and often encroaching onto the

frontoclypeus, anterior margin of the head glabrous, irregularly textured, or with one to many carinae or

striae, long antennae, nymph often with apical process on the head and male pygofer with a pair of long

apical appendages widely separated at the base.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae, or shagreen. Anterior margin of head glabrous, irregularly textured, or with one to

many transverse carinae. Frontoclypeus not tumid; texture shagreen, glabrous, or striate (Drabescina).

Clypellus widening apically; apex following or slightly surpassing normal curve of gena. Lorum subequal

to or wider than clypellus near base. Antennal bases near upper or anterodorsal corners of eyes; antennal

pits usually large, encroaching on frontoclypeus. Antennae long, as long as width of head or longer.

Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledge wealdy

developed (carinate or weakly carinate) or strongly developed, with a definite ledge (Drabescina). Ocelli

present; close to or distant from eyes; on anterior margin of head.

Thorax. Pronotum exceeding or not exceeding eyes anteriorly; lateral margin carinate or not; lateral

margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin or large, extending around wing

apex (Drabescina); with 3 anteapical cells; veins not raised; with or without reflexed costal veins; A1-A2

crossvein absent or present.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Legs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae, or not in one

row, scattered; row AV with thin hair-like setae, without setae, or rarely with short or long macrosetae.

Protibia dorsal surface rounded, convex (Paraboloponina) or dorsal surface flat, sharply carinate along

AD and PD margins (Drabescina). Metafemur apex macrosetae 2+1, 2+1+1, or 2+2+1. Metatarsomere

I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae absent or reduced (< two rows) or

present, well differentiated into several rows. Subgenital plates free from each other; articulated with

valve; usually without macrosetae. Style broadly bilobed basally, median anterior lobe pronounced.

Basal processes of the aedeagus/connective absent or reduced or present, connected or articulated to

connective or near base of aedeagus. Aedeagus with single shaft and gonopore. Connective anterior

arms somewhat divergent, Y- or U-shaped; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex or not strongly convex; dorsal sculpturing pattern strigate; sculpturing

reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula broad,

gradually tapered; with or without dorsal median tooth; dorsal teeth on apical 1/3 or more or restricted

to apical 1/4 or less; teeth large, regularly shaped or small, regularly or irregularly shaped.

Geography and ecology

Distribution: Palearctic, Afrotropical, Oriental and Australian regions. Drabescini are often found on

woody hosts in Old World tropical or deciduous forests, and are often collected at lights.

Remarks

Drabescini contains 38 genera and 187 valid species. Dmitriev (2004a) synonymized Drabescini

and Paraboloponini based on shared characters of the nymph, and its monophyly was supported in

phylogenetic analyses. Here, we accept the synonymy by Dmitriev (2004a) but continue to recognize

separate subtribes following the circumscription of the two groups by Zhang & Webb (1996) due to the

morphological differences of the adults that distinguish them. The phylogenetic analyses here suggest

that the tribe is closely related to Scaphoideini, with which it shares long antennae and profemur row

AV with thin hair-like setae or without setae. Our analyses provided no evidence of a close relationship

between Drabescini and Selenocephalini, which some previous authors (e.g., Linnavuori & Al-Ne’amy

1983; Zhang & Webb 1996) placed together in a separate subfamily, Selenocephalinae.

Selected references

Linnavuori (1978a, b), Webb (1981), Zhang & Webb (1996), Viraktamath (1998), Dmitriev (2002,

2004a).

Included subtribes:

Drabescina Ishihara, 1953

Fig. 19

Type genus: Drabescus Stal, 1870.

Diagnosis

Drabescina are large, robust, black, grey, or fuscous leafhoppers. They can be distinguished from

Paraboloponina by the very strong antennal ledges, antennae usually somewhat shorter- 1.0 to 1.5 x

European Journal of Taxonomy 45: 1-211 (2013)

width of head, rugose or striate texture of the frontoclypeus, protibia flattened and expanded dorsally

and with dorsal margins sharply carinate, and forewing appendix large and extending to the wing apex.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae. Anterior margin of head glabrous, irregularly textured, or with numerous transverse

striations or carinae. Frontoclypeus not tumid; texture glabrous or striate. Clypellus widening apically;

apex following or slightly surpassing normal curve of gena. Lorum subequal to or wider than clypellus

near base. Antennal bases near upper or anterodorsal corners of eyes. Antennae long, as long as width of

head or longer. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture. Antennal

ledges strongly developed (with a definite ledge). Ocelli present; distant from eyes; on anterior margin

of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix large, extending to wing apex; with 3 anteapical cells; veins

not raised; without reflexed costal veins; A1-A2 crossvein absent or present; apical venation not highly

reticulate.

Legs. Profemur with AMI seta only; intercalary row setae not in one row or scattered; row AV with thin,

hair-like setae or without setae. Protibia dorsal surface flat; sharply carinate along AD and PD margins.

Metafemur apex macrosetae 2+1. Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; with short point of articulation with pygofer. Pygofer

basolateral membranous cleft present; macrosetae absent or reduced (< two rows) or well differentiated

into several rows. Subgenital plates free from each other; articulated with valve; without macrosetae.

Style broadly bilobed basally, median anterior lobe pronounced; apophysis digitate, lobate, or incrassate

with distinct strigate to reticulate texturing. Aedeagus usually with pair of fused basal processes; without

basal hinge; with a single shaft and gonopore. Connective anterior arms somewhat divergent, Y-shaped;

articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern strigate; sculpturing reaching dorsal margin;

without distinctly delimited ventroapical sculpturing. Second valvula abruptly broadened medially or

subapically or broad, gradually tapered; with dorsal median tooth; teeth on apical 1/3 or more or teeth

restricted to apical 1/4 or less; teeth large, regularly shaped or teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Palearctic, Afrotropical and Oriental regions. Species of Drabescus have been collected at

lights and sweeping woody forest vegetation.

Remarks

Drabescina contains 2 genera and 55 species.

Included genera

Drabescus Stal, 1870

Rengatella Zhang & Webb, 1996

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 19. Drabescini (Drabescina). Drabescus zhangi sp. nov. A-I. Standard views (see Material &

Methods). J. Ventral view of $ pygofer.

European Journal of Taxonomy 45: 1-211 (2013)

Paraboloponina Ishihara, 1953

Fig. 20

Type genus: Parabolopona Matsumura, 1912.

Diagnosis

Paraboloponina are medium sized to large leafhoppers, and are variable in shape and color. They can

be distinguished from Drabescina by the antennal ledges which are weak or absent, antennae usually

longer, 1.5 x width of head or longer, texture of the frontoclypeus shagreen, protibia rounded dorsally or

rarely somewhat flattened, and forewing appendix not especially large.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae, or shagreen. Anterior margin of head glabrous, irregularly textured, foliaceous, with

numerous transverse striations, with 2 or 3 parallel carinae, or with numerous carinae. Frontoclypeus

not tumid; texture shagreen or striate. Clypellus widening apically, apex following or slightly surpassing

normal curve of gena. Lorum subequal to or wider than clypellus near base. Antennal bases near upper

or anterodorsal corners of eyes. Antennal pits often very large and encroaching onto frontoclypeus.

Antennae long, 1.5 x width of head or longer. Gena not incised or obtusely incised laterally; with fine

erect seta beside laterofrontal suture. Antennal ledges absent or wealdy developed (carinate or weakly

carinate). Ocelli present; close to or distant from eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewings macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins not

raised; without or with reflexed costal veins; A1-A2 crossvein absent or present; apical venation not

highly reticulate.

Legs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae; row AV with

thin, hair-like setae or without setae, or rarely with short, stout setae. Protibia dorsal surface rounded,

convex. Mesofemur row AV sometimes with several long macrosetae. Metafemur apex macrosetae 2+1

or 2+1+1 or 2+2+1. Metatarsomere I not expanded apically, plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; with short point of articulation with pygofer. Pygofer

basolateral membranous cleft present; macrosetae absent or reduced (< two rows) or well differentiated

into several rows. Subgenital plates free from each other; articulated with valve; without macrosetae

or with macrosetae scattered, irregularly arranged or uniseriate laterally. Style broadly bilobed basally,

median anterior lobe pronounced. Basal processes of the aedeagus/connective absent or reduced or

present, connected or articulated to connective or near base of aedeagus. Aedeagus without basal hinge;

with a single shaft and gonopore. Connective anterior arms somewhat divergent, Y- or U-shaped;

articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern strigate, concatenate, or reticulate; sculpturing

reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula abruptly

broadened medially or subapically or gradually broadened medially or subapically; with or without

dorsal median tooth; teeth on apical 1/3 or more or restricted to apical 1/4 or less; teeth small, regularly

or irregularly shaped.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

■ -I

/ \ 1 /

ml a

\ •'

1 ■ A

Fig. 20. Drabescini (Paraboloponina). Parabolopona webbi sp. nov. A-I. Standard views (see Material

& Methods). G. Connective shown with fused process.

European Journal of Taxonomy 45: 1-211 (2013)

Geography and ecology

Distribution: Palearctic, Afrotropical, Oriental and Australian regions.

Remarks

Paraboloponina contains 36 genera and 132 species.

Included genera

Athysanopsis Matsumura, 1914

Bhatia Distant, 1908

Canopyana Viraktamath & Srinivasa, 2006

Carvaka Distant, 1918

Divus Distant, 1908

Drabescoides Kwon & Lee, 1979

Dryadomorpha K i rlcal dy, 1906

Eminea Seven, 1997

Favintiga Webb, 1981

Halimunella Kamitani, 2012

Hybrasil Kirlcaldy, 1907

Indokutara Viraktamath, 1998

Isaca Walker, 1857

Jamitettix Matsumura, 1940

Karoseefa Webb, 1981

Kotabala Viraktamath, 1998

Kutara Distant, 1908

Megabyzus Distant, 1908

My so l is Kirkaldy, 1904

Nakula Distant, 1918

Nirvanguina Zhang & Webb, 1996

Oceanopona Linnavuori, 1960

Odmiella Linnavuori, 1978

Odzalana Linnavuori, 1969

Omanella Merino, 1936

Parabolopona Matsumura, 1912

ParohinkaWQbb, 1981

Rkutelorbus Webb, 1981

Roxasella Merino, 1936

Roxasellana Zhang & Zhang, 1998

Sombakidia Zhang & Webb, 1996

Stenomiella Evans, 1955

Tengatka Zhang & Webb, 1996

Tenompoella Zhang & Webb, 1996

Waigara Zhang & Webb, 1996

Welmaya Zhang & Webb, 1996

Drakensbergenini Linnavuori, 1979

Fig. 21

Type genus: Drakensbergena Linnavuori, 1961.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Diagnosis

Drakensbergenini are small to large, ivory, ochraceous, to dark brown brachypterous leafhoppers,

often with a dark brown median stripe from apex of crown to abdomen. They can be identified by the

produced head, ocelli on the crown, tumid frontoclypeus, strongly tapering clypellus, and brachypterous

fore wings.

Description

Head. Head subequal to or wider than pronotum; somewhat to strongly produced. Discal portion of

crown shagreen. Anterior margin of head shagreen or foliaceous; often shagreen and rounded to face

laterally and foliaceous apically. Frontoclypeus tumid; texture shagreen. Clypellus strongly tapering

apically; apex following or slightly surpassing normal curve of gena. Lorum distinctly narrower than

Fig. 21. Drakensbergenini. A-B. Drakensbergena deorsuspina Stiller, 2009. C. Drakensbergena sp.

D-I. D. phaeogramma Stiller, 2009. A-I. Standard views (see Material & Methods).

European Journal of Taxonomy 45: 1-211 (2013)

clypellus near base. Antennal bases near upper or anterodorsal corners of eyes. Antennae short, less

than 1.5 x width of head or long, 1.5 x width of head or longer. Gena strongly incised laterally (nearly

forming right angle); with fine erect seta beside laterofrontal suture. Antennal ledge weakly developed

(carinate or wealdy carinate). Ocelli reduced or present; distant from eyes; on crown.

Thorax. Pronotum lateral margin carinate or rarely ecarinate; lateral margin shorter than basal width of

eye.

Wings. Forewing brachypterous; coriaceous; wings usually fused; veins obscure.

Legs. Profemur with AMI absent or reduced; intercalary row reduced or absent; row AV with relatively

long macrosetae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+1 or 2+1+1.

Metatarsomere I expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point.

Pygofer basolateral membranous cleft present; macrosetae absent or reduced (< two rows). Subgenital

plates free from each other; articulated with valve; without macrosetae. Style broadly bilobed basally,

median anterior lobe pronounced. Basal processes of the aedeagus/connective absent or reduced.

Aedeagus with single shaft and gonopore; apex bifid. Connective anterior arms somewhat divergent,

Y-shaped; stem often appearing to be formed by two longitudinal bars; articulated with aedeagus.

Segment X long; sclerotized dorsally and laterally.

Female genitalia. Pygofer with macrosetae reduced or absent. Ovipositor not protruding far beyond

pygofer apex. First valvula not strongly convex; dorsal sculpturing pattern concatenate, reticulate,

granulose, or maculose; sculpturing reaching dorsal margin or submarginal; without distinctly delimited

ventroapical sculpturing. Second valvula abruptly broadened medially or subapically or broad, gradually

tapered; without dorsal median tooth; dorsal teeth on apical 1/3 or more; teeth small, regularly or

irregularly shaped. Third valvula with two irregular setal rows along entire ventral margin.

Geography and ecology

Distribution: Afrotropical region (South Africa and Lesotho). Drakensbergena spp. feed on grasses in

the high elevation grassland biome of southern Africa.

Remarks

Dralcensbergenini contains the type genus and 18 species. This group was considered a separate subfamily

by Linnavuori (1979a) but placed as a tribe of Deltocephalinae by Zahniser & Dietrich (2010). Stiller

(2009) described 16 new species of the genus. Phylogenetic analyses and morphological characters

(produced head, profemur intercalary row reduced, AMI seta reduced or absent, AV setae relatively

long, metafemur apical setae 2+1+1 or 2+1) suggest that this tribe is related to Eupelicini.

Selected references

Linnavuori (1979a), Stiller (2009).

Included genera

Drakensbergena Linnavuori, 1961

Eupelicini Sahlberg, 1871

Figs 22, 23

Type genus: Eupelix Germar, 1821.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Diagnosis

Eupelicini are medium sized to large ivory, ochraceous, green, or brownish leafhoppers. They can be

identified by the strongly produced or spatulate head, gena without fine erect seta beside laterofrontal

suture, genal margins strongly emarginate (-90° angle), forewing veins raised or carinate, metafemur

apical setae 2+0, metatarsomere I expanded apically, male pygofer macrosetae absent or reduced,

subgenital plates without macrosetae, and aedeagus not hinged basally.

Description

Head. Head subequal to or wider than pronotum; somewhat to very strongly produced anteriorly. Discal

portion of crown shagreen, granulose, or punctate. Anterior margin of head foliaceous. Frontoclypeus

elongate; not tumid; texture shagreen or punctate. Clypellus parallel-sided, tapering apically, or widening

apically; apex following or slightly surpassing normal curve of gena; notched apically. Lorum subequal

to or wider than clypellus near base. Antennal bases near upper or anterodorsal corners of eyes. Gena

strongly incised laterally (nearly forming right angle); without fine erect seta beside laterofrontal suture.

Antennae short, less than 1.5 x width of head. Antennal ledge absent or wealdy developed (carinate or

weakly carinate). Antennal bases near upper or anterodorsal comers of eyes. Ocelli present; close to or

distant from eyes; on anterior margin of head or on crown.

Thorax. Pronotum usually with median longitudinal carina and sublateral keels or carinae; lateral

margin carinate or not; lateral margin as long as or longer than basal width of eye.

Wings. Fore wing macropterous or submacropterous; appendix absent or reduced (Paradorydiina) or

appendix large, extending around wing apex (Eupelicina); with 3 anteapical cells; veins distinctly raised

or carinate; without reflexed costal veins; A1-A2 crossvein absent.

Legs. Profemur with AMI absent or reduced; intercalary row reduced or absent; row AV without setae or

with relatively long macrosetae. Protibia dorsal surface flat, AD and PD margins at -90° angles but not

carinate. Metafemur apex macrosetae 2+0. Metatibia row AV macrosetae restricted to apical 1/2 or less.

Metatibia not strongly compressed laterally, more or less square shaped in cross section; distinctly bent

apically in dorsal view. Metatarsomere I expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer or fused to pygofer (Paradorydiina), if articulated then

lateral margin short, articulating with pygofer at a point. Pygofer basolateral membranous cleft present

(Eupelicina) or absent (Paradorydiina); macrosetae absent or reduced (< two rows). Subgenital plates

free from each other; articulated with or fused to valve; without macrosetae. Style broadly bilobed

basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent or

reduced. Aedeagus with single shaft and gonopore. Connective anterior arms somewhat divergent, Y- or

V-shaped; articulated with aedeagus.

Female genitalia. Pygofer with macrosetae reduced or absent. Ovipositor protruding (Paradorydiina)

or not protruding (Eupelicina) far beyond pygofer apex. First valvula not strongly convex; dorsal

sculpturing pattern reticulate, granulose, maculose, or imbricate (with overlapping scales); sculpturing

reaching dorsal margin or submarginal; with (Paradorydiina) or without (Eupelicina) distinctly delimited

ventroapical sculptured area. Second valvula broad, gradually tapered or gradually broadened medially

or subapically; with or without dorsal median tooth; dorsal teeth on apical 1/3 or more if present; teeth

small, regularly or irregularly shaped (Eupelicina) or absent (Paradorydiina).

Geography and ecology

Distribution: Palearctic, Afrotropical, Oriental and Australian regions. All members are grass feeders.

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Remarks

Eupelicini contains 7 genera and 64 species. Listrophorina is removed from the tribe here and synonymized

with Chiasmini, and Dorycephalini is considered a separate tribe following Zahniser & Dietrich (2010).

Eupelicini is supported as a monophyletic group in phylogenetic analyses and by shared morphological

characters. It is related to other grass-feeding tribes of Deltocephalinae including Drakensbergenini,

Chiasmini, and Stenometopiini.

Selected references

Evans (1966), Morrison (1973), Linnavuori (1979a, b), Viraktamath & Viraktamath (1989), Dmitriev

(2001), D’Urso (1992).

Included subtribes:

Eupelicina Sahlberg, 1871

Fig. 22

Type genus: Eupelix Germar, 1821.

Diagnosis

Eupelicina are medium sized to large, whitish leafhoppers. They can be distinguished from Paradorydiina

by the crown partly extending over the eyes anteriorly, the dorsal position of the ocelli, lateral margin

of the pronotum carinate, frontoclypeus with a median carina anteriorly, forewing appendix large and

extending around the wing apex, forewing marginal vein complete, hind wing marginal vein complete,

male valve not fused to the pygofer, style more robust, ovipositor not extending far beyond the pygofer

apex, first valvula dorsal sculpturing pattern reticulate or imbricate, second valvula with a dorsal median

tooth and with dorsal teeth on the apical 1/3.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown shagreen. Anterior margin

of head foliaceous. Frontoclypeus not tumid; texture shagreen; with median longitudinal carina toward

apex. Clypellus parallel-sided or tapering apically; apex following or slightly surpassing normal curve

of gena. Lorum subequal to or wider than clypellus near base. Antennal bases near upper or anterodorsal

corners of eyes. Antennae short, less than 1.5 x width of head. Gena strongly incised laterally (nearly

forming right angle); without fine erect seta beside laterofrontal suture. Antennal ledges absent. Ocelli

present; distant from eyes; on crown.

Thorax. Pronotum lateral margin carinate; lateral margin as long as or longer than basal width of eye.

Wings. Forewing macropterous or submacropterous; appendix large, extending around wing apex; with

3 anteapical cells; veins distinctly raised or carinate; without reflexed costal veins; A1-A2 crossvein

absent; apical venation not highly reticulate.

Legs. Profemur with AMI absent or reduced; intercalary row reduced or absent; row AV with relatively

long macrosetae. Protibia dorsal surface flat, AD and PD margins at ~90° angles but not carinate.

Metafemur apex macrosetae 2+0. Metatarsomere I expanded apically; plantar setae in adult all simple.

Male genitalia. Valve articulated with pygofer, with short point of articulation with pygofer. Pygofer

basolateral membranous cleft present; macrosetae absent or reduced (< two rows). Subgenital plates

free from each other; articulated with valve; without macrosetae. Style broadly bilobed basally; median

anterior lobe pronounced. Basal processes of the aedeagus/connective absent. Aedeagus without basal

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 22. Eupelicini (Eupelicina). Eupelix cuspidata (Fabricius, 1775). A-I. Standard views (see Material

& Methods).

European Journal of Taxonomy 45 : 1-211 ( 2013 )

hinge; with a single shaft and gonopore. Connective anterior arms somewhat divergent, Y-shaped;

articulated with aedeagus.

Female genitalia. Pygofer with macrosetae reduced or absent. Ovipositor not protruding far beyond

pygofer apex. First valvula not strongly convex; dorsal sculpturing pattern reticulate or imbricate (with

overlapping scales); sculpturing reaching dorsal margin or submarginal; without distinctly delimited

ventroapical sculpturing. Second valvula broad, gradually tapered; with dorsal median tooth; teeth on

apical 1/3 or more; teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Palearctic region.

Remarks

Eupelicina contains the type genus and its 2 included species.

Included genera

Eupelix Germar, 1821

Paradorydiina Evans, 1936

Fig. 23

Type genus: Paradorydium Kirkaldy, 1901.

= Dorydiini Fieber, 1872 (Type: Dorydium Burmeister, 1839; suppressed).

Diagnosis

Paradorydiina are medium sized to large, somewhat to very elongate, produced leafhoppers, usually

whitish, greenish, or brownish in color. They can be distinguished from Eupelicina by the crown not

extending over the eyes anteriorly, ocelli on the anterior margin of the head, lateral margin of the

pronotum not carinate, frontoclypeus without a median anterior carina, forewing appendix reduced or

absent, forewing marginal vein incomplete, hind wing marginal vein incomplete, male valve fused to the

pygofer, style gracile, ovipositor extending far beyond the pygofer apex, first valvula dorsal sculpturing

pattern maculose or granulose, second valvula without a dorsal median tooth and without dorsal teeth

on apical 1/3.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown punctate. Anterior margin of

head foliaceous. Frontoclypeus not tumid; texture punctate. Clypellus parallel-sided or tapering apically;

apex following or slightly surpassing normal curve of gena. Lorum subequal to or wider than clypellus

near base. Antennal bases near upper or anterodorsal corners of eyes. Antennae short, less than 1.5 x

width of head. Gena strongly incised laterally (nearly forming right angle); without fine erect seta beside

laterofrontal suture. Antennal ledges absent or weakly developed (carinate or weakly carinate). Ocelli

present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin not carinate; lateral margin as long as or longer than basal width of

eye.

Wings. Forewing macropterous or submacropterous; acuminate; appendix absent or reduced; veins

distinctly raised or carinate; without reflexed costal veins; A1-A2 crossvein absent; apical venation not

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 23. Eupelicini (Parodorydiina). Paradorydium quadrigonum (Naude, 1926). A-E, H—I. Standard

views (see Material & Methods). D. Pygofer shown with valve, plates, and internal genitalia. J. Ventral

view of pygofer showing valve, plates, connective and styles. K. Ventral views of sternites I (slap) and

II (s2ap).

European Journal of Taxonomy 45 : 1-211 ( 2013 )

highly reticulate; marginal vein often absent or partially absent medial to Ml+2; hind wing marginal

vein often absent.

Legs. Profemur with AMI absent or reduced; intercalary row reduced or absent; row AV with thin, hair¬

like setae, without setae, or with relatively long macrosetae. Protibia dorsal surface flat, AD and PD

margins at -90° angles but not carinate. Metafemur apex macrosetae 2+0. Metatarsomere I expanded

apically, plantar setae simple, tapered.

Male genitalia. Valve fused to pygofer. Pygofer basolateral membranous cleft absent, not membranous;

macrosetae absent or reduced (< two rows). Subgenital plates free from each other and articulated with

or fused to valve; without macrosetae. Style broadly bilobed basally, median anterior lobe pronounced;

somewhat gracile, not fully sclerotized posteriad of median anterior lobe. Basal processes of the

aedeagus/connective absent or reduced. Aedeagus without basal hinge; with a single shaft and gonopore.

Connective anterior arms somewhat divergent, Y- or U-shaped; articulated with aedeagus.

Female genitalia. Pygofer with macrosetae reduced or absent. Ovipositor protruding far beyond pygofer

apex. First valvula not strongly convex; dorsal sculpturing pattern granulose or maculose; sculpturing

submarginal; with ventroapical sculpturing distinctly delimited. Second valvula gradually broadened

medially or subapically; without dorsal median tooth; teeth absent.

Geography and ecology

Distribution: Palearctic, Afrotropical, Oriental and Australian regions.

Remarks

Paradorydiina contains 6 genera and 62 species. Paradorydium and Chloropelix were included in the

phylogenetic analyses here and were resolved together with very high branch support on a relatively

long branch sister to Eupelix.

Sectoculus is transferred to Paradorydiina here. Morrison (1973a) established the genus and indicated

that it belonged to Paradorydiini which he included in Hecalinae. It was listed in Hecalini by Oman

et al. (1990) who apparently overlooked its placement by Morrison. It is similar morphologically to

Paradorydium.

Included genera

Afralycisca Kocak, 1981

Chloropelix Lindberg, 1936

Mapochia Distant, 1910

Mapochiella Evans, 1966

Paradorydium Kirkaldy, 1901

Sectoculus Morrison, 1973 placement nov. (transferred from Hecalini)

Faltalini Zahniser & Dietrich, 2010

Fig. 24

Type genus: Faltala Oman, 1938.

Diagnosis

F altalini are small to large sized, somewhat to strongly dorsoventrally flattened, ivory, ochraceous, fuscous,

yellowish, or brown leafhoppers; sometimes sexually dimorphic; dorsum sometimes with 5-6 more or

less developed longitudinal ochraceous stripes bordered with fuscous or dark coloring; stripes may be

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

present on head, pronotum, wings, and/or abdomen. They resemble Hecalini in the following characters:

head often produced or parabolic in shape; first valvula dorsal sculpturing pattern submarginal; first

valvula relatively straight; first valvula with distinctly delimited apicoventral sculptured area; second

valvula without dorsal teeth. Faltalini can be distinguished from Hecalini and other Deltocephalinae

by the following combination of characters: dorsoventrally flattened shape, head often produced,

anterior margin of head often foliaceous or with carinae or striae, ocelli often distant from eyes (close

to eyes in Hecalini), sometimes with distinctive color pattern, first valvula dorsal sculpturing pattern

maculose to granulose, first valvula with distinctly delimited apicoventral sculpturing (usually not as

long as in Hecalini and Dorycephalini), first valvula not humpbacked dorsally (humpbacked in Hecalini

and Dorycephalini), second valvula straight to somewhat convex ventrally (concave in Hecalini and

Dorycephalini), second valvula without dorsal teeth.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae or shagreen. Anterior margin of head shagreen, glabrous, striate, or foliaceous.

Frontoclypeus not tumid; texture shagreen or glabrous. Clypellus parallel-sided or tapering apically;

apex following or slightly surpassing normal curve of gena. Lorum subequal to or wider than clypellus

near base. Antennal bases near middle or posteroventral (lower) corners of eyes or near anterodorsal

(upper) corners of eyes (some Tenucephalus , Bonamus). Antennae short, less than 1.5 x width of head or

long ( Tenucephalus , Bonamus). Gena obtusely incised laterally; with fine erect seta beside laterofrontal

suture. Antennal ledge absent. Ocelli absent or reduced or present; close to ( Tenucephalus ) or distant

from eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter ( Tenucephalus , Bonamus) or longer

than basal width of eye.

Wings. Forewings of both sexes brachypterous, quadrate, coriaceous, venation indistinct to reticulate;

or males submacropterous or macropterous (. Acrolithus , Hecullus) and females brachypterous to

submacropterous; or males and females macropterous ( Tenucephalus , Bonamus)', if macropterous,

appendix restricted to anal margin; with 3 anteapical cells or with 2 anteapical cells; veins not raised;

without reflexed costal veins; A1-A2 crossvein absent.

Fegs. Protrochanter often with stout ventroapical seta. Pro femur with AMI seta only; intercalary row

with one row of five or more fine setae; row AV with short, stout setae or with relatively long macrosetae.

Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2 or 2+2+1. Metatarsomere I

not expanded apically, plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae absent or reduced (< two rows) or

macrosetae well differentiated into several rows. Subgenital plates free from each other; articulated

with valve; macrosetae irregularly arranged, uniseriate laterally, or reduced or absent. Style broadly

bilobed basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent or

reduced or present ( Tenucephalus ). Aedeagus with single shaft and gonopore. Connective anterior arms

somewhat divergent, Y-shaped, or closely appressed anteriorly, linear shaped; articulated with or rarely

fused (some Tenucephalus) to aedeagus.

Female genitalia. Pygofer with macrosetae reduced or absent. Ovipositor protruding ( Tenucephalus ,

Bonamus) or not protruding far beyond pygofer apex. First valvula not strongly convex; dorsal

sculpturing pattern granulose or maculose; sculpturing submarginal; with ventroapical sculpturing

European Journal of Taxonomy 45 : 1-211 ( 2013 )

— ^ _ r

, — r

f w

Fig. 24. Faltalini. Clorindaia cyphora Blocker & Fang, 1992. A-I. Standard views (see Material &

Methods).

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

distinctly delimited. Second valvula broad, gradually tapered or gradually broadened medially or

subapically; without dorsal median tooth; teeth absent.

Geography and ecology

Distribution: Nearctic and Neotropical regions. Known hosts are grasses. Tenucephalus spp. have been

collected at lights. Many species appear to be predominantly or completely brachypterous, some are

sexually dimorphic for wing length (. Acrolithus , Hecullus), and some are macropterous ( Bonamus ,

Hecalocorica , Tenucephalus).

Remarks

Faltalini contains 10 genera and 26 species and is distributed in the New World from the southwestern

United States to Argentina and Chile. Tenucephalus and Bonamus form a morphologically distinct

group within the tribe, distinguished by the following characters: body long and slender; both sexes

macropterous; ovipositor extending well beyond pygofer apex. Although including these genera makes

the tribe as a whole more difficult to characterize morphologically, some characters, especially of the

female genitalia, support a relationship to other Faltalini and 28S rDNA data very strongly support the

monophyly of the tribe (Zahniser & Dietrich 2010).

Egenus Oman, 1938 is removed from the tribe here. It was tentatively included by Zahniser & Dietrich

(2010) but subsequent observations of the female revealed that the genus does not possess the characters of

the ovipositor that help to distinguish the tribe. It is transferred to Athysanini. A preliminary phylogenetic

study of Hecalini (Catanach, unpublished) suggests that Egenus is related to Arrugada Oman, 1938.

Selected references

DeLong & Thambimuttu (1973), Linnavuori & DeLong (1977a), Blocker & Fang (1992), Zahniser &

Webb (2004), Marino de Remes Fenicov & Paradell (2009), Zahniser & Dietrich (2010).

Included genera

Acrolithus Freytag & Ma, 1988

Aequecephalus DeFong & Thambimuttu, 1973

Bonamus Oman, 1938

Clorindaia Finnavuori, 1975

Faltala Oman, 1938

Hecalocorica Nielson, 1996

Hecullus Oman, 1949

Kramer ana DeFong & Thambimuttu, 1973

Tenucephalus DeFong, 1944

Virganana DeFong & Thambimuttu, 1973

Fieberiellini Wagner, 1951

Fig. 25

Type genus: Fieberiella Signoret, 1880.

= Synophropsini Ribaut, 1952.

Diagnosis

Fieberiellini are medium sized, tawny to brown leafhoppers. They can be identified by the forewing

lacking an appendix, forewings not overlapping, connective T-shaped and with a distinct stem, style

European Journal of Taxonomy 45 : 1-211 ( 2013 )

linear or not broadly bilobed basally, and female sternite VIII not concealed and folded beneath sternite

VII and movably articulated with sternite VII*.

* Sternite VIII is almost always folded beneath and concealed by sternite VII in Deltocephalinae and

is often membranous or reduced; thus, when the pygofer and ovipositor are in their resting positions,

sternite VII appears to be the one immediately basad of the genital capsule. Meyer-Arndt & Remane

(1992b) pointed out the unique configuration inFieberiellini but identified the entire structure (comprising

sternites VII and VIII) as sternite VII.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae. Anterior margin of head shagreen or with numerous transverse striations or carinae.

Frontoclypeus not tumid; texture shagreen. Clypellus widening apically; apex following or slightly

surpassing normal curve of gena. Lorum subequal to or wider than clypellus near base. Antennal bases

near middle or posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x width of head.

Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledge wealdy

developed (carinate or weakly carinate). Ocelli present, close to eyes, on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix absent or reduced; with 3 anteapical cells; veins not raised;

without reflexed costal veins; A1-A2 crossvein present or absent.

Legs. Pro femur with AMI seta only or with one or more additional proximal setae; intercalary row

with one row of five or more fine setae; row AV with relatively long macrosetae. Protibia dorsal surface

rounded, convex. Metafemur apex macrosetae 2+2+1. Metatarsomere I not expanded apically; plantar

setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at

a point. Pygofer basolateral membranous cleft present; macrosetae well differentiated into several

rows. Subgenital plates free from each other; articulated with valve; macrosetae irregularly arranged,

uniseriate laterally, with two lateral rows of macrosetae, or with some irregularly arranged macrosetae

near lateral margin. Style linear, median anterior lobe not pronounced. Basal processes of the aedeagus/

connective absent or reduced. Aedeagus with single shaft and gonopore. Connective anterior arms

widely divergent, T-shaped, with darkly pigmented ventromedial ridge contrasting with more lightly

pigmented dorsolateral areas; stem well-developed; articulated with aedeagus. Segment X sclerotized

dorsally and laterally; sometimes large; sometimes with long ventrally directed processes.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. Sternite VIII not concealed and folded beneath sternite VII, movably articulated with sternite

VII. First valvula convex; dorsal sculpturing pattern strigate; sculpturing reaching dorsal margin;

without distinctly delimited ventroapical sculpturing. Second valvula abruptly broadened medially or

subapically; without dorsal median tooth; teeth on apical 1/3 or more; teeth large, regularly shaped or

small, regularly or irregularly shaped.

Geography and ecology

Distribution: endemic to the Palearctic region and adventive in the Nearctic region (Fieberiella florii

Stal, 1864 in North America). Fieberiellini are most diverse in the southern Palearctic region, especially

the Mediterranean subregion, and have host plants in a number of different families as documented by

Meyer-Arndt & Remane (1992b).

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 25. Fieberiellini. A-C, J. Fieberiellaflorii (Stal, 1864). D-I. Placotettix taeniatifrons (Kirschbaum,

1868). A-I. Standard views (see Materials & Methods). F. Plates shown slightly obliquely and valve

shown separated from plates. J. $ sternites VI-VIII ( s6-s8 ).

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Remarks

Fieberiellini contains 10 genera and 48 valid species. Meyer-Amdt & Remane (1992a, b) produced a

detailed revision of the tribe in which they illustrated the species, documented host plants and geographic

ranges, clarified the morphological characterization and circumscription of the tribe, and proposed

phylogenetic relationships among the genera. Phylogenetic analyses suggest that Fieberiellini is a

relatively early diverging lineage of Deltocephalinae, and retains plesiomorphic characteristics of the

male genitalia. It appears to be related to the monotypic Neotropical tribe Luheriini, with which it shares

some morphological characteristics. However, the divergent 28S rDNA sequence of Lukeria constricta

and difficulty in amplifying some regions have confounded an exact and confident phylogenetic

placement of the group. Fieberiella florii is adventive in North America and is a vector of the western

strain of North American aster yellows, Western and Eastern X-disease of peach, and Western X-disease

yellow leaf roll strain. Cechenotettix quadrinotatus (Mulsant & Rey, 1855) is a vector of yellow decline

of Lavandula.

Phycotettix was listed in Fieberiellini by Oman et al. (1990). It was not treated by Meyer-Arndt &

Remane (1992a, b) and is removed from the tribe here and placed in Athysanini. It does not possess any

of the characters that define the tribe.

Selected references

Meyer-Arndt & Remane (1992a, b), Dmitriev (2003).

Included genera

Cechenotettix Ribaut, 1942

Docotettix Ribaut, 1948

Dohukia Meyer-Arndt & Remane, 1992

Erico te ttix Lindberg, 1960

Fieberiella Signoret, 1880

Habrostis Dubovsky, 1966

Heliotettix Rodrigues, 1968

Parafieberiella Dlabola, 1974

Placotettix Ribaut, 1942

Synophrops is FI au pt, 1926

Goniagnathini Wagner, 1951

Fig. 26

Type genus: Goniagnathus Fieber, 1866.

Diagnosis

Goniagnathini are medium sized to large, squat, robust leafhoppers. They can be identified by the

broad head, anterior margin of head glabrous, large forewing appendix (in macropterous individuals),

subgenital plates fused to each other, valve apparently absent or fused to subgenital plates, style with

broad basal part articulated with linear or modified apical part, and connective fused to the aedeagus.

Description

Head. Head subequal to or wider than pronotum; crown wide, parallel-margined. Discal portion of

crown glabrous with radial or longitudinal striae. Anterior margin of head glabrous or irregularly

textured. Frontoclypeus not tumid; texture glabrous, rugose, or shagreen. Clypellus widening apically;

apex following or slightly surpassing normal curve of gena. Lorum subequal to or wider than clypellus

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

near base. Antennal bases near middle or posteroventral (lower) corners of eyes. Antennae short, less

than 1.5 x width of head. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture.

Antennal ledge weakly developed (carinate or wealdy carinate). Ocelli present; close to eyes; on anterior

margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing usually macropterous, sometimes brachypterous; if macropterous, appendix large,

extending around wing apex; with 3 anteapical cells; veins not raised or raised; veins sometimes

pustulate; without reflexed costal veins; A1-A2 crossvein present.

Legs. Profemur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with relatively long macrosetae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae

2+2+1. Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve fused to subgenital plates or obscure. Pygofer basolateral membranous cleft

present; macrosetae well differentiated into several rows. Subgenital plates fused to each other; with

or without macrosetae. Style linear, median anterior lobe not pronounced or broadly bilobed basally,

median anterior lobe pronounced; broad basal part articulated with linear or modified apical part, with

cleft between the two. Basal processes of the aedeagus/connective absent or reduced. Aedeagus with

single shaft and gonopore. Connective anterior arms somewhat divergent, Y- or U-shaped, or closely

appressed anteriorly, linear shaped; sometimes highly reduced in size; fused to aedeagus.

Female genitalia. Py gofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern strigate, concatenate, or reticulate; sculpturing

reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula slender

throughout; without dorsal median tooth; teeth on apical 1/3 or more or teeth restricted to apical 1/4 or

less; teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Palearctic, Afrotropical, Oriental and Australian. Species are found in arid and grassland

regions of the Old World where they feed on herbs and shrubs (e.g., Artemisia, Tamarix).

Remarks

Goniagnathini contains 4 genera and 60 species. Goniagnathini is an early-diverging lineage and appears

to have diverged from the stem group of Deltocephalinae at nearly the same time as Acinopterini,

Fieberiellini, and Luheriini. It has a unique configuration of the style, with a broad base articulated to a

linear or modified apical part.

Selected references

Linnavuori (1978b), Dash & Viraktamath (2001), Dmitriev (2003), Fletcher & Zahniser (2008), Duan et

al. (2009), Viraktamath & Gnaneswaran (2009).

Included genera

Goniagnathus Fieber, 1866

Me gal ops ins Emelj anov, 1961

Raunothus Dlabola, 1987

Tamaricades Emeljanov, 1962

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Fig. 26. Goniagnathini. A-B, D-K. Goniagnathus emeljanovi sp. nov. C. G. rugulosus (Haupt, 1917).

A-I. Standard views (see Material & Methods). G. Connective and style shown with aedeagus fused

to connective. H. Aedeagus shown fused to connective, style also shown. I. Caudal view of aedeagus/

connective shown with median arms of styles extending ventrally. J. Ventral view of pygofer. K. §

sternite VII.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Hecalini Distant, 1908

Figs 27, 28

Type genus: Hecalus Stal, 1864.

= Reuteriellini Evans, 1947.

Diagnosis

Hecalini are medium sized to large, somewhat to strongly dorsoventrally flattened, stramineous, yellow,

green, or brown leafhoppers, sometimes with bright orange or reddish markings. They can be identified

by the produced and parabolically shaped head, dorsoventrally flattened body, lateral margin of pronotum

as long as or longer than the basal width of the eye, ocelli closer to they eyes than the laterofrontal

sutures, apodemes of male sternite I long and relatively narrow, apodemes of male stemite II broad and

well-developed, male pygofer often produced or pointed posterodorsally, segment X withdrawn into the

py gofer, ventral margins of the male pygofer often lobate, aedeagus often with one or two pairs of apical

processes, first valvula dorsal sculpturing granulose to maculose and submarginal, first valvula often

with distinctly delimited ventroapical sculpturing, second valvula usually without teeth, humpbacked

dorsally, and concave ventrally.

Description

Head. Head subequal to or wider than pronotum; often produced anteriorly and/or parabolically shaped.

Discal portion of crown glabrous with radial or longitudinal striae, shagreen or granulose. Anterior

margin of head shagreen, glabrous, irregularly textured, foliaceous, with a single transverse carina,

or with 2 or 3 parallel carinae. Frontoclypeus tumid or flat; texture shagreen. Clypellus parallel-sided,

tapering apically, or rarely widening apically; apex following or slightly surpassing normal curve of

gena. Lorum narrower than, subequal to, wider than clypellus near base. Antennal bases near middle

or posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely

incised laterally or strongly incised laterally (nearly forming right angle); with or without (. Attenuipyga ,

some Glossocratus) fine erect seta beside laterofrontal suture. Antennal ledge absent. Ocelli present;

close to eyes, closer to eyes than laterofrontal sutures; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin as long as or longer than basal width of eye.

Wings. Forewing brachypterous to macropterous; if macropterous, appendix restricted to anal margin or

rarely large, extending around wing apex; with 3 anteapical cells; veins not elevated or rarely elevated

(Attenuipyga ); without reflexed costal veins; A1-A2 crossvein absent or present; A1 and A2 sometimes

confluent (. Attenuipyga [Dorycara]).

Legs. Pro femur with AMI seta only or rarely with AMI and with one or more additional proximal setae

(some Glossocratus ) or AMI absent (. Attenuipyga :); intercalary row with one row of five or more fine

setae or setae scattered, not in one row, or setae absent or reduced; row AV with short, stout setae or

with relatively long macrosetae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae

2+2+1, 2+2+1 and with additional proximal macrosetae (some Glossocratus ), or 2+0 ( Attenuipyga ).

Metatarsomere I not expanded apically or expanded apically; plantar setae simple, tapered.

Male genitalia. Apodemes of sternite I usually long, usually as long as those of sternite II, relatively

narrow. Apodemes of stemite II large, broad (exception: Glossocratus). Valve articulated with pygofer;

lateral margin short, articulating with pygofer at a point. Pygofer basolateral membranous cleft present;

macrosetae well differentiated into several rows; often produced or pointed posterodorsally, with ‘anal

tube’ withdrawn into pygofer; ventral margins often lobate. Subgenital plates free from each other;

European Journal of Taxonomy 45 : 1-211 ( 2013 )

articulated with valve; macrosetae scattered, irregularly arranged or uniseriate laterally. Style broadly

bilobed basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent

or reduced. Aedeagus with single shaft and gonopore; often with one or two pairs of apical processes,

or rarely with preapical processes. Connective anterior arms somewhat divergent, Y- or U-shaped;

articulated with aedeagus.

Female genitalia. Pygofer with macrosetae absent, reduced, or with numerous macrosetae. Ovipositor

usually protruding or sometimes not protruding far beyond pygofer apex. First valvula not strongly

convex; dorsal sculpturing pattern granulose or maculose; sculpturing submarginal or rarely reaching

dorsal margin; usually with ventroapical sculpturing distinctly delimited. Second valvula gradually

broadened medially or subapically; humpbacked dorsally; concave ventrally; usually without dorsal

median tooth (except Glossocratus)', dorsal teeth usually absent but if present (some Glossocratus ),

teeth small, regularly or irregularly shaped, on apical 1/3 or more.

Geography and ecology

Distribution: cosmopolitan. Hecalini are grass feeders and are sometimes very brightly colored with

yellow, green (in live specimens), or orange. Some are extremely elongate and dorsoventrally flattened,

apparently an adaptation enhancing crypsis on their grass hosts.

Remarks

Hecalini contains 23 genera and 180 species. The three included exemplars ( Hecalus , Attenuipyga and

Glossocratus ) were resolved as a monophyletic group here in MP analyses, but not in ML or Bayesian

analyses. In the MP analysis, the partitioned Bremer support from the Histone H3 data was -14.4 which

apparently strongly conflicted with the signal from the 28S and morphology data sets (DI= 7.7 and 13.2,

respectively). This conflict in signal likely influenced the results of the ML and Bayesian analyses which

resolved different topologies for the hecalines included here. MP analyses of the combined data suggest

that Hecalini is closely related to Arrugadini, but more data are needed to confirm this.

Hamilton (2000) revised the concept of Hecalini and excluded several genera that have only a superficial

similarity to the tribe. He also pointed out several characters that are shared between Hecalini and

“Dory cephalic”, although of the latter he was apparently referring only to the New World Attenuipyga

and Neoslossonia, and not to Dorycephalus which does not share several of the characters he mentioned.

The characters listed by Hamilton (2000) and others noted here that are shared between Attenuipyga,

Neoslossonia , and other Hecalini and not with Dorycephalus are listed in the Diagnosis. Based on their

shared morphological characters and support from the molecular data, Attenuipyga and Neoslossonia

are transferred to Hecalini here. Dorycephalini is maintained as a separate monotypic tribe.

Morrison (1973) revised the Oriental fauna and suggested that Clavena may belong to Ledrini based on

the position of the ocelli stated in the original description of C. sulcata Melichar. He did not examine

the type, so its inclusion in Hecalini has not been confirmed. Sectoculus is removed from Hecalini and

placed in Eupelicini: Paradorydiina (see discussion under Paradorydiina).

Selected references

Linnavuori (1957, 1959, 1975), Morrison (1973), Linnavuori & DeLong (1978c), Hamilton (2000),

Dmitriev (2002).

Included subtribes:

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Glossocratina Dmitriev, 2002

Fig. 27

Type genus: Glossocratus Fieber, 1866.

Diagnosis

Glossocratina are medium sized to large brownish leafhoppers. They are distinguished from Hecalina by

the laterofrontal sutures keeled above the antennal pits, gena sometimes without a fine erect seta beside

the laterofrontal suture, apodemes of male sternite II not as developed as in Hecalina, ovipositor not

protruding far beyond the pygofer apex, and second valvula with teeth. Nymphs can be distinguished

from Hecalina by the oblique lateral carina extending from ante nn al pit to margin of head, outer margin

of the gena angulate, anal tube with a long apical process, and body covered with short setae (Dmitriev,

2002 ).

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown radially or longitudinally

striate or shagreen. Anterior margin of head foliaceous or with 2 or 3 parallel carinae. Frontoclypeus

tumid or not; texture shagreen. Laterofrontal sutures keeled above antennal pits. Clypellus parallel¬

sided, tapering apically, or widening apically; apex following or slightly surpassing normal curve of

gena. Lorum subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral

(lower) corners of eyes. Antennae short, less than 1.5 x width of head. Gena strongly incised laterally

(nearly forming right angle); with or without fine erect seta beside laterofrontal suture. Antennal ledges

absent. Ocelli present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin as long as or longer than basal width of eye.

Wings. Forewing brachypterous to macropterous; appendix restricted to anal margin or large, extending

around wing apex; with 3 anteapical cells; veins not raised; without reflexed costal veins; A1-A2

crossvein absent or present; apical venation not highly reticulate.

Legs. Pro femur with AMI seta only or with AMI and with one or more additional proximal setae;

intercalary row setae not in one row or scattered; row AV with relatively long macrosetae. Protibia dorsal

surface rounded, convex. Metafemur apex macrosetae 2+2+1 or 2+2+1 and with additional proximal

macrosetae. Metatarsomere I expanded apically; plantar setae simple, tapered.

Male genitalia. Apodemes of male sternite II not as long as those of sternite I (sternite II apodemes

less developed than in Hecalina). Valve articulated with pygofer; with short point of articulation with

pygofer. Pygofer dorsoapical margin incised to near mid-length; basolateral membranous cleft present;

macrosetae well differentiated into several rows. Subgenital plates free from each other; articulated with

valve; macrosetae uniseriate laterally. Style broadly bilobed basally, median anterior lobe pronounced.

Basal processes of the aedeagus/connective absent. Aedeagus without basal hinge; with a single shaft

and gonopore. Connective anterior arms somewhat divergent, Y- or U-shaped; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula not strongly convex; dorsal sculpturing pattern granulose or maculose; sculpturing

reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula very

broad; gradually or abruptly broadened medially or subapically; with or without dorsal median tooth;

teeth on apical 1/3 or more; teeth small, regularly or irregularly shaped.

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Fig. 27. Hecalini (Glossocratina). Glossocratus afzelii (Stal, 1854). A-I. Standard views (see Material

& Methods). G. Connective and style shown with aedeagus. J. Dorsal view of $ abdominal sternites

I-III (partial view of sternite III), slap = apodeme of sternite I, s2ap = apodeme of sternite II.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Geography and ecology

Distribution: Palearctic, Afrotropical and Oriental regions. Glossocratina are grass feeders.

Remarks

Glossocratina contains the type genus and 29 species.

Included genera

Glossocratus Fieber, 1866

Hecalina Distant, 1908

Fig. 28

Type genus: Hecalus Stal, 1864.

Diagnosis

Hecalina are medium sized to large yellow, greenish, or brownish leafhoppers, sometimes with orange

or reddish stripes. They are distinguished from Glossocratina by the unkeeled laterofrontal suture, gena

with fine erect seta beside the laterofrontal suture, ovipositor extending far beyond the pygofer apex, and

second valvula without dorsal teeth.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown radially or longitudinally

striate or shagreen. Anterior margin of head shagreen, glabrous, irregularly textured, foliaceous, with a

single transverse carina, or with 2 or 3 parallel carinae. Frontoclypeus tumid or not; texture shagreen.

Clypellus parallel-sided or tapering apically; apex following or slightly surpassing normal curve of

gena. Forum distinctly narrower than clypellus near base. Antennal bases near middle or posteroventral

(lower) corners of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely incised laterally

or strongly incised laterally (nearly forming right angle); with fine erect seta beside laterofrontal suture.

Antennal ledges absent. Ocelli present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin as long as or longer than basal width of eye.

Wings. Forewing brachypterous to macropterous; appendix restricted to anal margin; with 3 anteapical

cells; veins not raised; without reflexed costal veins; A1-A2 crossvein absent or present; apical venation

not highly reticulate.

Fegs. Profemur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1.

Metatarsomere I not expanded apically or expanded apically, plantar setae simple, tapered.

Male genitalia. Apodemes of stemite I long, as long as those of sternite II, relatively narrow.

Apodemes of sternite II large, broad. Valve articulated with pygofer; with short point of articulation

with pygofer. Pygofer basolateral membranous cleft present; macrosetae well differentiated into several

rows; anteroventral margins lobate; produced posterodorsally. Subgenital plates free from each other;

articulated with valve; macrosetae scattered, irregularly arranged, or uniseriate laterally. Style broadly

bilobed basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent.

Aedeagus without basal hinge; with a single shaft and gonopore; often with one or two pairs of apical or

subapical processes. Connective anterior arms somewhat divergent, Y- or U-shaped; stem often short;

articulated with aedeagus. Segment X partially retracted into pygofer.

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Fig. 28. Hecalini (Hecalina). A-B, D-I. Hecalus hamiltoni sp. nov. C. Hecalus viridis (Uhler, 1877).

J. Attenuipyga platyrhynchus (Osborn, 1894). Al, $. A2, A-I. Standard views (see Material &

Methods). J. Ventral views of stemites I and II. slap = apodeme of stemite I, s2ap = apodeme of sternite

II.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Female genitalia. Pygofer with macrosetae reduced or absent. Ovipositor protruding far beyond pygofer

apex. First valvula not strongly convex; humpbacked in shape; dorsal sculpturing pattern granulose or

maculose; sculpturing submarginal; with ventroapical sculpturing distinctly delimited. Second valvula

gradually broadened medially or subapically; humpbacked in shape; concave ventrally; without dorsal

median tooth; teeth absent.

Geography and ecology

Distribution: cosmopolitan. Hecalina are grass feeders.

Remarks

Hecalina contains 22 genera and 151 species.

Except for Hecalus , Attenuipyga, and Neoslossonia , all genera were previously unplaced to subtribe

and are included here in the subtribe Hecalina. Hecalus was previously the only included genus in

the subtribe (Dmitriev 2002). Attenuipyga and Neoslossonia are transferred to Hecalina from Dory-

cephalini.

Included genera

Alospangbergia Evans, 1973 placement nov. (previously unplaced in Hecalini)

Annidion Kirkaldy, 1905 placement nov. (previously unplaced in Hecalini)

Attenuipyga Oman, 1949 placement nov. (transferred from Dorycephalini)

Bordesia de Bergevin, 1929 placement nov. (previously unplaced in Hecalini)

Cephalius Fieber, 1875 placement nov. (previously unplaced in Hecalini)

Clavena Melichar, 1902 placement nov. (previously unplaced in Hecalini)

Dicyphonia Ball, 1900 placement nov. (previously unplaced in Hecalini)

Epicephalius Matsumura, 1908 placement nov. (previously unplaced in Hecalini)

Hecalocratus Evans, 1966 placement nov. (previously unplaced in Hecalini)

Hecalus Stal, 1864

Hecalusina He, Zhang & Webb, 2008 placement nov. (previously unplaced in Hecalini)

Jiutepeca Linnavuori & DeLong, 1978 placement nov. (previously unplaced in Hecalini)

Linnavuoriella Evans, 1966 placement nov. (previously unplaced in Hecalini)

Lualabanus Linnavuori, 1975 placement nov. (previously unplaced in Hecalini)

Memnonia Ball, 1900 placement nov. (previously unplaced in Hecalini)

Neohecalus Linnavuori, 1975 placement nov. (previously unplaced in Hecalini)

Neoslossonia Van Duzee, 1909 placement nov. (transferred from Dorycephalini)

Parabolocratalis Evans, 1955 placement nov. (previously unplaced in Hecalini)

Psegmatus Fieber, 1875 placement nov. (previously unplaced in Hecalini)

Reuteriella Signoret, 1879 placement nov. (previously unplaced in Hecalini)

Spangbergiella Signoret, 1879 placement nov. (previously unplaced in Hecalini)

Thomsoniella Signoret, 1880 placement nov. (previously unplaced in Hecalini)

Hypacostemmini Linnavuori & Al-Ne’amy, 1983

Fig. 29

Type genus: Hypacostemma Linnavuori, 1961.

Large, greenish lealhoppers.

European Journal of Taxonomy 45 : 1-211 ( 2013 )

Diagnosis

Hypacostemmini are large, greenish leafhoppers. They can be identified by their large size (7-10 mm),

highly reticulated apical venation of the forewing, appendix reduced in size, tectiform configuration

of the forewings, and style apex foot-like, sharply angled medially with apex projecting laterally and

darkly sclerotized.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae. Anterior margin of head shagreen. Frontoclypeus not tumid; texture shagreen.

Clypellus parallel-sided or widening apically; apex following or slightly surpassing normal curve of

gena. Lorum subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral

(lower) comers of eyes. Antennae short, less than 1.5 x width of head or long, 1.5 x width of head or

longer. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledge

weakly developed (carinate or wealdy carinate). Ocelli present; close to eyes; on anterior margin of

head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin, reduced in size; with 3 anteapical

cells; veins not raised; without reflexed costal veins; A1-A2 crossvein absent or present; apical venation

highly reticulate.

Legs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1.

Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae well differentiated into several rows.

Subgenital plates free from each other; articulated with valve; macrosetae uniseriate laterally. Style

broadly bilobed basally; median anterior lobe pronounced; apex foot-lilce, sharply angled medially

with apex projecting laterally, darkly sclerotized. Basal processes of the aedeagus/connective absent or

reduced. Aedeagus with single shaft and gonopore. Connective anterior arms somewhat divergent, Y- or

U-shaped; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex, dorsal sculpturing pattern strigate, concatenate, or reticulate; sculpturing

reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula abruptly

broadened medially or subapically; with dorsal median tooth; teeth on apical 1/3 or more; teeth small,

regularly or irregularly shaped.

Geography and ecology

Distribution: Afrotropical (South Africa). Specimens have been collected at lights and swept from trees

and bushes. Recorded host plant families include Urticaceae, Sapindaceae, and Anacardiaceae (Theron,

1987).

Remarks

Hypacostemmini contains the type genus and 5 species. Phylogenetic analyses suggest that Hypacostemma

is related to a group of South African genera including Bonaspeiini and some undescribed genera sampled

in the analyses here (Cameroon 142 and Ghanall4). This group is in turn related to Selenocephalini.

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 29. Hypacostemmini. Hypacostemma viridissima Liimavuori, 1961. A-I. Standard views (see

Material & Methods).

European Journal of Taxonomy 45: 1-211 (2013)

More detailed phylogenetic analyses of these genera, including more representatives of other genera in

the diverse Bonaspeiini, are needed.

Selected references

Linnavuori & Al-Ne’amy (1983), Theron (1987).

Included genera

Hypacostemma Linnavuori, 1961

Koebeliini Baker, 1897

Figs 30, 31

Type genus: Koebelia Baker, 1897.

= Grypotini Haupt, 1929.

Diagnosis

Koebeliini are small to medium sized, yellow, light green, or brown leafhoppers. They can be identified

by the ocelli distant from the eyes, clypellus long, narrow and extending well beyond the normal curve

of the gena, and metatarsomere I with platellae on the plantar surface.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae or shagreen. Anterior margin of head foliaceous (Koebeliina) or shagreen, with

transverse striations, or carinate (Grypotina). Frontoclypeus not tumid; texture shagreen. Clypellus

narrow; parallel-sided or tapering apically; apex far-surpassing normal curve of gena. Lorum subequal

to or wider than clypellus near base. Antennal bases near middle or posteroventral (lower) corners of

eyes or near anterodorsal (upper) comers of eyes. Antennae short, less than 1.5 x width of head. Gena

obtusely incised laterally, with fine erect seta beside laterofrontal suture. Antennal ledges absent. Ocelli

present; distant from eyes; on anterior margin of head (Grypotina) or on face (Koebeliina).

Thorax. Pronotum lateral margin carinate or not; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous or submacropterous; appendix absent, reduced, or restricted to anal

margin; with 3 anteapical cells; veins not raised or slightly raised; without reflexed costal veins; A1-A2

crossvein absent.

Legs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae or reduced

or absent; row AV with short, stout setae or with relatively long macrosetae. Protibia dorsal surface

rounded, convex or flat, AD and PD margins at -90° angles but not carinate. Metafemur apex macrosetae

2+2+1, 2+2, 2+1, 2+1+1, or 2+2+1+1. Metatarsomere I not expanded apically; plantar surface with one

or more platellae.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae absent or reduced (< two rows) or well

differentiated into several rows. Subgenital plates free from each other; articulated with valve; macrosetae

scattered, irregularly arranged. Style broadly bilobed basally, median anterior lobe pronounced. Basal

processes of the aedeagus/connective absent or reduced. Aedeagus with single shaft and gonopore; often

asymmetrical. Connective anterior arms somewhat divergent, Y-shaped; articulated with or fused to

aedeagus.

100

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Female genitalia. Pygofer with macrosetae reduced or absent or with numerous macrosetae. Ovipositor

protruding far beyond pygofer apex or not. First valvula convex or not strongly convex, dorsal sculpturing

pattern strigate, concatenate, reticulate, or with rectangular shaped cells; sculpturing reaching dorsal

margin or submarginal; without distinctly delimited ventroapical sculpturing. Second valvula broad

and gradually tapered apically, gradually broadened medially or subapically, or slender throughout;

without dorsal median tooth; teeth on apical 1/3 or more or restricted to apical 1/4 or less; teeth large

and regularly shaped or small, regularly or irregularly shaped.

Geography and ecology

Distribution: Palearctic, Oriental, and Nearctic regions. This tribe has an interesting disjunct distribution

between the two subtribes, Koebeliina (western North America) and Grypotina (Oriental and Palearctic

regions). Grypotes puncticollis (Herrich-Schaffer, 1834) is adventive in the northeastern United States

(Wheeler 1989). All host records of Koebeliini are from Pinus spp. G. puncticollis feeds on P. sylvestris

L. (Scotch pine) and has also been recorded from Swiss mountain pine, P. mugo Turra (Wheeler 1989)

and P. nigricans Host.

Remarks

Koebeliini contains 6 genera and 15 species. Phylogenetic analyses strongly support its monophyly

(Grypotellus and Koebelia included in analyses) and suggest (with little or no branch support) that the

tribe may be related to Mukariini or Cochlorhinini, with which it shares the presence of platellae on

metatarsomere I.

Selected references

Oman (1971), Dietrich & Dmitriev (2003), Dmitriev (2003).

Included subtribes:

Grypotina Haupt, 1929

Fig. 30

Type genus: Grypotes Fieber, 1866.

Diagnosis

Grypotina are small to medium sized, yellow, greenish, or tawny leafhoppers. They can be distinguished

from Koebeliina by the crown not or weakly produced; anterior margin of head shagreen, crown rounded

to face, with transverse striations, or carinate, not foliaceous; forewing with appendix restricted to anal

margin; forewing veins not pustulate; metafemur penultimate pair of macrosetae close-set, when present;

metatibia dorsal rows with distinct intercalary setae; metatibia pecten without platellae; first valvula

curved; first valvula dorsal sculpturing pattern imbricate, reaching dorsal margin; second valvula with

teeth on apical 1/3 or more.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae or shagreen. Anterior margin of head shagreen, with numerous transverse striations,

or with a single transverse carina; sometimes irregularly shaped. Frontoclypeus not tumid; texture

shagreen. Clypellus narrow; parallel-sided or tapering apically; apex extending well beyond normal

curve of gena. Forum subequal to or wider than clypellus near base. Antennal bases near middle or

posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x width of head or long, 1.5 x

101

European Journal of Taxonomy 45: 1-211 (2013)

width of head or longer. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture.

Antennal ledges absent. Ocelli present; distant from eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

•

, mwLgBSfa

. •

Fig. 30. Koebeliini (Grypotina). Grypotes puncticollis (Herrich-Schaffer, 1834). A-I. Standard views

(see Material & Methods). H-I. Aedeagus shown with partially fused connective at base.

102

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Wings. Fore wing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins not

raised; without reflexed costal veins; A1-A2 crossvein absent; apical venation not highly reticulate.

Legs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1.

Metatarsomere I not expanded apically; plantar surface with one or more platellae.

Male genitalia. Valve articulated with pygofer; with short point of articulation with pygofer. Pygofer

basolateral membranous cleft present; macrosetae well differentiated into several rows. Subgenital plates

free from each other; articulated with valve; macrosetae scattered, irregularly arranged. Style broadly

bilobed basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent.

Aedeagus without basal hinge; with a single shaft and gonopore. Connective anterior arms somewhat

divergent, Y- or U-shaped; articulated with or fused to aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula curved; dorsal sculpturing pattern concatenate, reticulate, or with rectangular shaped

cells; sculpturing reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second

valvula broad, gradually tapered; without dorsal median tooth; teeth on apical 1/3 or more; teeth large,

regularly shaped or teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Palearctic, Oriental and Nearctic (adventive) regions ( Grypotes puncticollis is introduced

in North America).

Remarks

Grypotina contains 5 genera and 10 species.

Included genera

Grypotellus Emeljanov, 1999

Grypotes Fieber, 1866

Pinopona Viraktamath & Sohi, 1998

Shivapona Ghauri & Viraktamath, 1987

Sohipona Ghauri & Viraktamath, 1987

Koebeliina Baker, 1897

Fig. 31

Type genus: Koebelia Baker, 1897.

Diagnosis

Koebeliina are medium sized to large brown leafhoppers. They are distinguished from Grypotina by

the crown strongly produced, spatulate, anterior margin of head foliaceous, forewing veins pustulate,

appendix absent, metatibia dorsal rows without or with indistinct intercalary setae, metatibia pecten with

platellae, first valvula relatively straight and not strongly curved, first valvula dorsal sculpturing pattern

reticulate to maculose and submarginal, and second valvula with teeth restricted to apical 1/4.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown shagreen. Anterior margin of

head foliaceous. Frontoclypeus not tumid; texture shagreen. Clypellus narrow; parallel-sided or tapering

apically; apex extending well beyond normal curve of gena. Lorum subequal to or wider than clypellus

103

European Journal of Taxonomy 45: 1-211 (2013)

Fig. 31. Koebeliini (Koebeliina). Koebelia grossa Ball, 1909. A-I. Standard views (see Material &

Methods).

104

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

near base. Antennal bases near middle or posteroventral (lower) corners of eyes. Antennae short, less

than 1.5 x width of head. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture.

Antennal ledges absent. Ocelli present; distant from eyes; on face.

Thorax. Pronotum lateral margin not carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous or submacropterous; appendix absent or reduced; with 3 anteapical

cells; veins not raised or slightly raised; veins pustulate; without reflexed costal veins; A1-A2 crossvein

absent; apical venation not highly reticulate.

Legs. Pro femur with AMI seta only; intercalary row reduced or absent; row AV with relatively long

macrosetae. Protibia dorsal surface flat, AD and PD margins at -90° angles but not carinate. Metafemur

apex macrosetae 2+1 or 2+2. Metatarsomere I not expanded apically; plantar surface with one or more

platellae.

Male genitalia. Valve articulated with pygofer; with short point of articulation with pygofer. Pygofer

basolateral membranous cleft present; macrosetae absent or reduced (< two rows). Subgenital plates

free from each other, articulated with valve; macrosetae scattered, irregularly arranged. Style broadly

bilobed basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent.

Aedeagus with large apical process arising from ventral margin; without basal hinge; with a single shaft

and gonopore. Connective anterior arms somewhat divergent, Y-shaped; articulated with aedeagus.

Female genitalia. Pygofer with macrosetae reduced or absent. Ovipositor protruding or not protruding

far beyond pygofer apex. First valvula not strongly convex; dorsal sculpturing pattern reticulate or

maculose; sculpturing reaching dorsal margin or submarginal; without distinctly delimited ventroapical

sculpturing. Second valvula broad, gradually tapered or slender throughout; without dorsal median

tooth; teeth restricted to apical 1/4 or less; teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Nearctic (Western North America).

Remarks

Koebeliina contains the type genus and 5 species.

Included genera

Koebelia Baker, 1897

Limotettigini Baker, 1915

Fig. 32

Type genus: Limotettix Sahlberg, 1871.

Diagnosis

Limotettigini are small to medium sized ivory, greyish, or black leafhoppers, often with dark markings.

They can be identified by the parallel-sided or tapering clypellus, pygofer with a spine-like process

arising from the dorsal margin, and aedeagus articulated with a plate-like “dorsal connective” at the

dorsal margin of the socle.

105

European Journal of Taxonomy 45: 1-211 (2013)

dorsal connective

Fig. 32. Limotettigini. A-B, D-I. Limotettix striolus (Fallen, 1906). C. L. decumanus Kontkanen, 1949.

A-I. Standard views (see Material & Methods). F. valve shown dissociated from plates.

106

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae or completely shagreen to base. Anterior margin of head shagreen. Frontoclypeus

not tumid; texture shagreen. Clypellus parallel-sided or tapering apically; apex following or slightly

surpassing normal curve of gena. Lorum distinctly narrower than clypellus near base. Antennal bases

near middle or posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x width of head.

Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledges absent.

Ocelli present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins not

raised; without reflexed costal veins; A1-A2 crossvein absent; apical venation not highly reticulate.

Legs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1.

Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; with short point of articulation with pygofer. Pygofer

basolateral membranous cleft present; macrosetae well differentiated into several rows; with spine¬

like process arising from dorsal side. Subgenital plates free from each other, articulated with valve;

macrosetae uniseriate laterally or with two lateral rows of macrosetae, sometimes with some irregularly

arranged macrosetae. Style broadly bilobed basally, median anterior lobe pronounced. Basal processes

of the aedeagus/connective absent or reduced. Aedeagus without basal hinge; with a single shaft and

gonopore; with plate-like “dorsal connective” articulated to dorsal margin of socle. Connective anterior

arms somewhat divergent, Y-shaped; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula not strongly convex; dorsal sculpturing pattern imbricate (with overlapping scales);

sculpturing reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second

valvula broad, gradually tapered; without dorsal median tooth; teeth on apical 1/3 or more; teeth small,

regularly or irregularly shaped.

Geography and ecology

Distribution: cosmopolitan. Some species are found around wetlands, bogs, and peatlands, while others

are found in grasslands or transitional habitats. Host plant families include Juncaceae, Cyperaceae,

Poaceae, Myricaceae, Ericaceae, and Asteraceae. L. (, Scleroracus ) flavopictus (Ishihara, 1953) is a

vector of Japanese aster yellows and potato witches’ broom in Japan and L. (, Scleroracus) vaccinii (Van

Duzee, 1890) is a vector of false blossom of cranberries in eastern North America.

Remarks

Limotettigini contains the type genus with 5 subgenera and 89 species, following the classification

of Hamilton (1994). In the phylogenetic analyses, Limotettix was consistently placed within the large

grass feeding clade of Deltocephalinae, but its position in the clade is uncertain. It may be related

to Cicadulini, the Athysanus- group (Athysanini), Hecalini, or Macrostelini. Dmitriev (2000, 2002)

included Anoterostemmina as a subtribe of Limotettigini but a relationship between these groups was not

supported in the phylogenetic analyses here, wherein molecular data for Anoterostemma were included

for the first time. Instead, Anoterostemma is resolved near Euscelis and other Athysanus- group genera,

and it is transferred to Athysanini here.

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Selected references

Hamilton (1994), McKamey (2001).

Included genera

Limotettix Sahlberg, 1871

Luheriini Linnavuori, 1959

Fig. 33

Type genus: Luheria Osborn, 1923.

Diagnosis

Luheriini are medium sized to large yellow, orange, and brown leafhoppers. They can be identified

by the anterior margin of the head with numerous striations or carinae, forewing appendix absent or

reduced, forewings not overlapping, central anteapical cell constricted medially, T-shaped connective,

and ‘linear’ style.

Description

Head. Head wider than pronotum. Discal portion of crown glabrous with radial or longitudinal striae.

Anterior margin of head with numerous transverse striations or carinae. Frontoclypeus not tumid;

texture shagreen. Clypellus widening apically; apex following or slightly surpassing normal curve of

gena. Lorum subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral

(lower) comers of eyes. Antennae moderately long, about as long as width of head. Gena obtusely

incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledges weakly developed

(carinate or wealdy carinate). Ocelli present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix absent or reduced; commissural margin straight; with 3

anteapical cells; central anteapical cell constricted medially; veins not raised; with or without reflexed

costal veins; A1-A2 crossvein present.

Legs. Pro femur with AMI and with one or more additional proximal setae; intercalary row with one

row of five or more fine setae; row AV with short, stout setae. Protibia dorsal surface rounded, convex.

Metafemur apex macrosetae 2+2+1. Metatarsomere I not expanded apically; plantar setae simple,

tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at

a point. Pygofer basolateral membranous cleft present; macrosetae well differentiated into several

rows. Subgenital plates free from each other; articulated with valve; macrosetae scattered, irregularly

arranged. Style linear, median anterior lobe not pronounced. Basal processes of the aedeagus/connective

absent. Aedeagus with single shaft and gonopore. Connective anterior arms widely divergent, T-shaped;

articulated with aedeagus. Segment X large; well sclerotized dorsally; with anteroventral processes.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex, dorsal sculpturing pattern strigate; sculpturing reaching dorsal margin;

without distinctly delimited ventroapical sculpturing. Second valvula relatively slender, gradually

tapered; without dorsal median tooth; teeth restricted to apical 1/4 or less; teeth small, regularly or

irregularly shaped.

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 33. Luheriini. Luheria constricta Osborn, 1923. A-I. Standard views (see Material & Methods).

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Geography and ecology

Distribution: Neotropical. The only known species, Lukeria constricta Osborn, 1923, is widely

distributed in South Am erica from northern Argentina to Bolivia and northeastern Brazil. Host plants

are unknown. Males exhibiting mud-puddling behavior have been collected on wet soil.

Remarks

The tribe contains only the type genus and species. Phylogenetic analyses and some plesiomorphic

characters of the head, wings, and male genitalia show that this is an early diverging lineage of

Deltocephalinae. It appears to be most closely related to the Old World tribe Fieberiellini. L. constricta

has an unusual 28S rDNA sequence that contains several large insertions, and some regions were

impossible to amplify using primers that worked well for most other Deltocephalinae. This divergent

sequence resulted in its long terminal branch in phylogenetic analyses.

Selected references

Linnavuori (1959), Zahniser (2005).

Included genera

Lukeria Osborn, 1923

Macrostelini Kirkaldy, 1906

Fig. 34

Type genus: Macrosteles Fieber, 1866.

= Balcluthini Baker, 1915.

= Gnathodini Baker, 1915.

= Coryphaelini Nast, 1972.

Diagnosis

Macrostelini are small to medium sized, slender, often stramineous, yellow, or greenish leafhoppers,

with or without dark markings. They can be identified by their long, slender shape, forewing with 2

anteapical cells, subgenital plates usually with membranous digitate apical lobe, and male pygofer

macrosetae sometimes plumose.

Description

Head. Head subequal to or wider than pronotum or distinctly narrower than pronotum. Discal portion

of crown glabrous with radial or longitudinal striae or shagreen. Anterior margin of head shagreen.

Frontoclypeus not tumid; texture shagreen. Clypellus parallel-sided or tapering apically; apex following

or slightly surpassing normal curve of gena. Lorum subequal to, wider than, or distinctly narrower than

clypellus near base. Antennal bases near middle or posteroventral (lower) corners of eyes. Antennae short,

less than 1.5 x width of head. Gena obtusely incised laterally; with fine erect seta beside laterofrontal

suture. Antennal ledges absent. Ocelli present; close to or distant from eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate or not; lateral margin shorter, as long as, or longer than basal

width of eye.

Wings. Forewing macropterous or submacropterous; appendix restricted to anal margin; with 2 anteapical

cells; veins not raised; without reflexed costal veins; A1-A2 crossvein absent; apical venation not highly

reticulate.

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 34. Macrostelini. Macrosteles quadrilineatus (Forbes, 1885). A-I. Standard views (see Material &

Methods). J. Ventral view of pygofer.

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European Journal of Taxonomy 45: 1-211 (2013)

Legs. Profemur with AMI seta only; intercalary row with one row of five or more fine setae; row

AV without setae, with thin hair-like setae, or with short, stout setae. Protibia dorsal surface rounded,

convex. Metafemur apex macrosetae 2+1+1 or 2+2+1. Metatarsomere I not expanded apically; plantar

setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae absent or reduced (< two rows) or well

differentiated into several rows; macrosetae sometimes plumose. Subgenital plates free from each other;

articulated with valve; macrosetae uniseriate laterally. Style broadly bilobed basally, median anterior

lobe pronounced. Basal processes of the aedeagus/connective absent or reduced. Aedeagus with single

shaft and gonopore. Connective anterior arms somewhat divergent, Y- or U-shaped; articulated with

aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex or not strongly convex; dorsal sculpturing pattern granulose, maculose,

or imbricate (with overlapping scales); sculpturing reaching dorsal margin or submarginal; without

distinctly delimited ventroapical sculpturing or with distinctly delimited apicoventral sculpturing.

Second valvula abruptly broadened medially or subapically, or broad, gradually tapered, or gradually

broadened medially or subapically; without dorsal median tooth; if present, teeth on apical 1/3 or more;

teeth absent or small, regularly or irregularly shaped.

Geography and ecology

Distribution: cosmopolitan. Some genera (e.g., Balclutha , Dalbulus ) feed mostly on grasses or sedges,

while others (e.g., Macrosteles) feed on a wide variety of hosts including herbaceous dicots, grasses,

and sedges. Macrosteles frontalis (Scott, 1875) lives on horsetail ( Equisetum sp.), a primitive vascular

plant. This tribe includes a number of economically important vectors of plant pathogens. Dalbulus

maidis (DeLong, 1923), the corn leafhopper, and D. elimatus (Ball, 1900) are especially damaging

to maize as vectors of corn stunt spiroplasma, maize bushy stunt phytoplasma, and maize rayado lino

virus in Mexico and Central and South America (Nault & Ammar 1989). Macrosteles quadrilineatus

(Forbes, 1885), the aster leafhopper, transmits American and European aster yellows to numerous

plants. Other species of Macrosteles transmit clover phyllody, clover dwarf, aster yellows, stolbur, Kok-

saghyz yellows, chrysanthemum yellows, lissers, anemone witches’ broom, eggplant dwarf, garland

chrysanthemum witches’ broom, marguerite yellows, mitsuba witches’ broom, onion yellows, tomato

yellows, white leaf phytoplasma, little cherry, and oat blue dwarf. Cicadulina mbila (Naude, 1924) and

congeneric species transmit maize streak in sub-Saharan Africa.

Remarks

Macrostelini contains 37 genera and 316 species. A morphological phylogenetic analysis (Knight &

Webb 1993) which included exemplars of all genera resolved the tribe as monophyletic. Three of the

four exemplars in phylogenetic analyses here ( Macrosteles , Dalbulus , and Balclutha) were resolved as a

monophyletic group with strong branch support. This monophyletic group is included in the large grass-

feeding clade but its relationship to other tribes in the clade is not clear. The fourth exemplar included in

analyses here is Evinus which was recently placed in the tribe (Lu et al. 2011); it was resolved near the

grass feeding tribes Chiasmini, Stenometopiini, and Eupelicini but branch support was not strong and

its exact relationship to other Deltocephalinae needs further study. Like other Macrostelini, it has two

anteapical cells in the forewing.

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Selected references

Blocker (1967), Knight (1987), Knight & Webb (1993), Webb & Vilbaste (1994), Dietrich et al. (1998),

Dmitriev (2003).

Included genera

Aderganna Knight & Webb, 1993

Afro steles Theron, 1975

Agelina Oman, 1938

Alebranus Linnavuori, 1959

Argaterma White, 1878

Atlantocella Dlabola, 1982

Balclutha Kirkaldy, 1900

Baldulus Oman, 1934

Balolina Knight & Webb, 1988

Cicadabara Knight & Webb, 1993

Cicadidella China, 1928

Cicadulina China, 1926

Cicaduloida Osborn, 1934

Cortona Oman, 1938

Coryphaelus Puton, 1886

Dalbulus DeLong, 1950

Davisonia Dorst, 1937

Elrabonia Linnavuori, 1959

Evinus Dlabola, 1977

Kadrabolina Knight & Webb, 1993

Macrosteles Fieber, 1866

Marquesitettix Metcalf, 1952

Masafuera Knight & Webb, 1993

Nesoclutha Evans, 1947

Nesolina Osborn, 1935

Nesophyla Osborn, 1934

Nesoriella Osborn, 1934

Nyhimbricus Webb, 1986

Paracicadula Osborn, 1934

Sagatus Ribaut, 1948

Sanctahelenia Dlabola, 1976

Scaphoidulina Osborn, 1934

Scaphosteles Knight & Webb, 1993

Sonronius Dorst, 1937

Stellena Theron, 1973

Teyasteles Linnavuori, 1969

Yamatotettix Matsumura, 1914

Magnentiini Linnavuori, 1978

Fig. 35

Type genus: Magnentius Singh-Pruthi, 1930.

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Diagnosis

Magnentiini are medium sized black, coarsely punctate, wedge-shaped leafhoppers. They can be

identified by their robust appearance, coarsely punctate texture of the pronotum and face, hirsute body

and face, crown very short, pronotum produced anteriorly with apex exceeding eyes in dorsal view,

ocelli distant from eyes, antennal ledge distinct, forewing appendix large, and forewing veins raised.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown punctate. Anterior margin of

head punctate. Frontoclypeus not tumid or tumid, texture punctate. Clypellus parallel-sided, tapering

apically, or widening apically; apex following or slightly surpassing normal curve of gena. Lorum

subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral (lower)

corners of eyes. Antennae short, less than 1.5 x width of head. Gena not or slightly incised laterally;

hirsute, with many fine hairs (difficult to interpret presence/absence of fine erect seta). Antennal ledges

strongly developed, with distinct ledge. Ocelli present; distant from eyes; on face.

Thorax. Pronotum exceeding eyes anteriorly; lateral margin carinate; lateral margin shorter than basal

width of eye.

Wings. Forewing macropterous or submacropterous; appendix present, extending around wing apex;

with 3 anteapical cells; veins distinctly raised; without reflexed costal veins; A1-A2 crossvein present;

apical venation not highly reticulate.

Legs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with relatively long macrosetae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae

2+0. Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer dorsoapical margin incised to near mid-length; basolateral membranous cleft present;

macrosetae well differentiated into several rows. Subgenital plates free from each other; articulated

with valve; macrosetae scattered, irregularly arranged. Style broadly bilobed basally, median anterior

lobe pronounced. Basal processes of the aedeagus/connective absent. Aedeagus with single shaft and

gonopore. Connective anterior arms somewhat divergent, Y-shaped; articulated with aedeagus.

Female genitalia. Pygofer with macrosetae reduced or absent. Ovipositor not protruding far beyond

pygofer apex. First valvula not strongly convex; dorsal sculpturing pattern strigate; sculpturing reaching

dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula abruptly broadened

medially or subapically; without dorsal median tooth; teeth on apical 1/3 or more; teeth small, regularly

or irregularly shaped.

Geography and ecology

Distribution: Afrotropical and Oriental regions.

Remarks

Magnentiini contains 2 genera and 4 species. Although it bears a strong superficial resemblance to

Nionia , it was transferred from Nioniinae to Deltocephalinae by Zahniser & Dietrich (2010) based on

the deltocephaline-like male genitalia.

Magnentius was included here for the first time in analyses with molecular data. The placement in

Deltocephalinae was supported in analyses here as it was resolved with moderate to strong branch

support (MP BS=71, DI=7) as sister to three Penthimiini in MP analyses and sister to two Penthimiini

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 35. Magnentiini. Magnentius clavatus Singh-Pruthi, 1930. A-I. Standard views (see Material &

Methods).

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European Journal of Taxonomy 45: 1-211 (2013)

in ML and Bayesian analyses with strong support (ML BS=93, PP=1.0, respectively). Some characters

that are unusual among Deltocephalinae, including its robust appearance, ocelli distant from the eyes,

strong antennal ledges, and forewing appendix extending around the wing apex, support its relationship

to Penthimiini.

Selected references

Linnavuori (1978a).

Included genera

Magnentius Singh-Pruthi, 1930

Ndua Linnavuori, 1978

Mukariini Distant, 1908

Fig. 36

Type genus: Mukaria Distant, 1908.

Diagnosis

Mukariini are small to medium sized, often dorsoventrally depressed or ventrally flattened, brown, black,

whitish, yellow, or green, leafhoppers, sometimes marked with orange or red. They can be identified

by the produced head, often with the frontoclypeus tumid distally, ventral part of face flat and lying

nearly horizontally or concave, and ocelli distant from eyes. Other distinctive characters (e.g. bifurcate

aedeagus with 2 gonopores in Mukaria and Pseudobalbillus) exist within the group but are not shared

by all taxa.

Description

Head. Head often somewhat produced; subequal to or wider than pronotum. Discal portion of crown

glabrous with radial or longitudinal striae. Anterior margin of head shagreen, striate, or with one to

several carinae. Frontoclypeus often tumid distally; texture shagreen, glabrous, or striate. Ventral part

of face lying in nearly horizontal plane, or concave. Clypellus widening apically; apex following or

slightly surpassing normal curve of gena. Lorum subequal to, wider than, or distinctly narrower than

clypellus near base. Antennal bases near upper or anterodorsal corners of eyes. Antennae short, less than

1.5 x width of head, or long. Gena obtusely incised laterally; with fine erect seta beside laterofrontal

suture. Antennal ledges strongly developed (with a definite ledge), or reduced or absent. Ocelli present;

distant from eyes; on anterior margin of head.

Thorax. Pronotum not exceeding eyes anteriorly, lateral margin carinate, lateral margin shorter than

basal width of eye.

Wings. Forewing macropterous; appendix absent, reduced, or restricted to anal margin; with 2 or 3

anteapical cells; outer anteapical cell sometimes very close to or confluent with costal vein; veins not

raised; without reflexed costal veins; A1-A2 crossvein absent; apical venation not highly reticulate.

Legs. Profemur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with thin, hair-like setae or without setae. Protibia dorsal surface rounded, convex. Metafemur apex

macrosetae 2+2+1. Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at

a point. Pygofer basolateral membranous cleft present; macrosetae absent or reduced (< two rows)

or well differentiated into several rows; often with processes arising ventrally, apically, or medially.

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 36. Mukariini. Mukaria maculata (Matsumura, 1912). A-H. Standard views (see Material &

Methods). G. Connective and styles shown with aedeagus fused to connective. H. Aedeagus shown

with connective and style.

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Subgenital plates free from each other; articulated with or rarely fused ( Agrica ) to valve; macrosetae

scattered, irregularly arranged, or uniseriate laterally. Style broadly bilobed basally, median anterior

lobe pronounced. Basal processes of the aedeagus/connective absent or reduced or present, connected

or articulated to connective or near base of aedeagus. Aedeagus with single shaft and gonopore or shaft

divided near base, with two gonopores. Connective anterior arms somewhat divergent, Y- or V-shaped;

anterior arms sometimes very widely divergent; articulated with or fused to aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex or not strongly convex; dorsal sculpturing pattern strigate; sculpturing

reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula broad,

gradually tapered; without dorsal median tooth; teeth on apical 1/3 or more; teeth large, regularly shaped

or small, regularly or irregularly shaped.

Geography and ecology

Distribution: Palearctic, Afrotropical, and Oriental regions. All known hosts are bamboos.

Remarks

Mukariini contains 12 genera and 54 species. The taxonomy of Mukariini has recently undergone

some modification. It previously only contained Mukaria , Neobassareus , and Pseudobalbillus , but

recently Chen et al. (2007) placed the Mohunia group in this tribe, Zahniser & Dietrich (2010) included

Scaphotettix and Agrica , and several new genera and species have been described (Hayashi, 1996;

Chen et al. 2007, 2008, 2009; Li et al. 2007; Dai et al. 2009). The phylogenetic analyses here included

representatives of Agrica, Mukaria , Scaphotettix , and an undescribed genus from China. Agrica and the

undescribed genus were resolved as sister to each other, as were Mukaria and Scaphotettix , but the four

were not resolved as monophyletic. However there is little or no support on the branches separating

them, and it is expected that more data will resolve the group as monophyletic in future analyses. Some

morphological characters and the restricted use of bamboos as hosts support the recognition of the group.

The tribe is potentially related to Vartini, Koebeliini, or Cochlorhinini. Future phylogenetic studies on

the group should also include representatives of the large African genus Pseudobalbillus to confirm its

placement in Mukariini, other genera of which are found in the Oriental region.

Selected references

Linnavuori (1979b), Hayashi (1996), Li & Chen (1998), Knight & Webb (2002), Chen et al. (2007,

2008,2009), Dai etal. (2009), Zahniser & Dietrich (2010), Khatri & Webb (2011), Yang & Chen (2011).

Included genera

Agrica Strand, 1942

Benglebra Mahmood & Ahmed, 1969

Flatfronta Chen & Li, 1997

Mohunia Distant, 1908

Mukaria Distant, 1908

Neobassareus Kocak, 1981

Neomohunia Chen & Li, 2007

Paramohunia Chen & Li, 2007

Pseudobalbillus Jacobi, 1912

Pseudomohunia Li, Chen & Zhang, 2007

Scaphotettix Matsumura, 1914

Tiaobeinia Chen & Li, 2008

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Occinirvanini Evans, 1966

Fig. 37

Type genus: Occinirvana Evans, 1941.

Diagnosis

Occinirvanini are medium sized, elongate, bizarrely shaped, ivory and orange colored leafhoppers, with

some fuscous coloration on the forewing apex. They can be identified by the elongate head, ocelli on the

crown, ocelli very distant from eyes, foliaceous anterior margin of the head, very long antennae, antennal

bases situated very high on the face anterad of the eyes, face strongly concave, and metatarsomere I

plantar surface with platellae.

Description

Head. Head subequal to or wider than pronotum; strongly produced, elongate. Discal portion of crown

shagreen. Anterior margin of head foliaceous. Face strongly concave. Frontoclypeus not tumid; texture

shagreen. Clypellus widening apically; apex following or slightly surpassing normal curve of gena.

Lorum subequal to or wider than clypellus near base. Antennal bases near upper part of face, anterad

of eyes. Antennae very long, 3-4 x width of head or longer; scape and pedicel large. Gena obtusely

incised laterally; hirsute, with many fine hairs (difficult to interpret presence/absence of fine erect seta).

Antennal ledges absent. Ocelli present; distant from eyes, closer to crown apex than to adjacent eye; on

crown.

Thorax. Pronotum not exceeding eyes anteriorly, lateral margin not carinate; lateral margin longer than

basal width of eye.

Wings. Fore wing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins not

raised; without reflexed costal veins; A1-A2 crossvein absent; apical venation not highly reticulate.

Legs. Pro femur short, stout; with AMI seta only; intercalary row with one row of five or more fine setae;

row AV without setae or with few short, stout setae. Protibia dorsal surface flat, AD and PD margins

at -90° angles but not carinate. Metafemur apex macrosetae 2+2+1. Metatarsomere I not expanded

apically; plantar surface with one or more platellae.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point.

Pygofer basolateral membranous cleft present; macrosetae well differentiated into several rows; without

dorsal or ventral processes; subrectangular in shape. Subgenital plates free from each other, articulated

with valve; with single row of macrosetae distant from lateral margin; apex with ~5-7 spaced macrosetae;

lateral margin with numerous short thin or thick setae. Style broadly bilobed basally, median anterior

lobe pronounced; apophysis with subapical dorsal tooth. Basal processes of the aedeagus/connective

absent. Aedeagus with single shaft and gonopore; with “dorsal connective” articulated with dorsal part

of socle. Connective anterior arms somewhat divergent, Y-shaped; stem present, 2 x length of anterior

arms; articulated with aedeagus. Segment X not strongly sclerotized.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern strigate to reticulate; sculpturing submarginal;

without distinctly delimited ventroapical sculpturing. Second valvula broad; gradually tapered; without

dorsal median tooth; teeth on apical 1/3 or more; teeth small, regularly or irregularly shaped.

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Fig. 37. Occinirvanini. Occinirvana eborea Evans, 1941. A-I. Standard views (see Material & Methods).

Al. 5 (BMNH). A2, B. S- J. Broad view of subgenital plate. K. Lateral view of apical half of style.

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Geography and ecology

Distribution: Western Australia. The only known species, Occinirvana eborea Evans, 1941, has been

collected on Casuarina.

Remarks

Occinirvanini contains only the type genus with one species. The male genitalia of Occinirvana are

illustrated and described here for the first time, and the phylogenetic analyses here are the first to include

molecular data for the tribe. The male genitalia are for the most part typical for Deltocephalinae. The

pattern of setae on the subgenital plate and the dorsal connective are somewhat unique. The characters

of the male do not obviously suggest a relationship with other deltocephaline tribes or genera, but the

molecular data very strongly supported a relationship with Loralia (Athysanini), another Australian-

endemic genus. The clade is placed in slightly different positions among the different analyses but

always basal to the large derived deltocephaline clade shown in Fig. 3b. Despite this intriguing result,

no unique morphological synapomorphies could be identified that would unite these two genera in a

common tribe. Further study of these and other unique Australian taxa (e.g., Euleimonios Kirlcaldy,

1906, Gunawardenea Fletcher & Moir, 2008 and Occiplanocephahis Evans, 1941) should be conducted

to explore their relationships to each other and to other Deltocephalinae, particularly since these

three genera have some unique features of the male genitalia in common with Loralia (but not with

Occinirvana ). For now, Occinirvanini is retained as a monotypic tribe pending further study.

Selected references

Evans (1966), Dietrich (2004).

Included genera

Occinirvana Evans, 1941

Opsiini Emeljanov, 1962

Figs 38-41

Type genus: Opsius Fieber, 1866.

Diagnosis

Opsiini are small to large, stramineous, yellow, green, or brown leafhoppers. They can be identified by

the bifurcate aedeagus with two shafts and gonopores. Some Mukariini and Ascius (Scaphytopiini) have

a similarly divided aedeagus but Opsiini lack the other characters that define those groups.

Description

Head. Head subequal to or wider than or distinctly narrower than pronotum. Discal portion of crown

glabrous with radial or longitudinal striae. Anterior margin of head shagreen, glabrous or irregularly

textured, with numerous transverse striations, or foliaceous ( Chlidochrus Emeljanov, 1962).

Frontoclypeus not tumid; texture shagreen or glabrous. Clypellus parallel-sided, tapering apically, or

widening apically; apex following or slightly surpassing normal curve of gena. Lorum subequal to or

wider than clypellus near base. Antennal bases near middle or posteroventral (lower) corners of eyes.

Antennae short, less than 1.5 x width of head. Gena obtusely incised laterally; with fine erect seta beside

laterofrontal suture. Antennal ledges absent or weakly developed (carinate or weakly carinate). Ocelli

absent or reduced or present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin not carinate or carinate; lateral margin shorter than basal width of eye.

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European Journal of Taxonomy 45: 1-211 (2013)

Wings. Forewing brachypterous to macropterous; appendix restricted to anal margin; with 3 anteapical

cells; veins not raised; with or without reflexed costal veins; A1-A2 crossvein absent or present; apical

venation not highly reticulate.

Legs. Profemur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+0, 2+1,

2+2, or 2+2+1. Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Apodemes of male sternites I and II often very well-developed. Valve articulated

with pygofer; lateral margin short, articulating with pygofer at a point. Pygofer dorsoapical margin

not strongly incised or incised to near mid-length; basolateral membranous cleft present; macrosetae

absent or reduced (< two rows) or well differentiated into several rows. Subgenital plates free from each

other; articulated with valve; without macrosetae or with macrosetae irregularly arranged or uniseriate

laterally. Style broadly bilobed basally, median anterior lobe pronounced. Basal processes of the

aedeagus/connective absent, reduced, or present and fused to base of aedeagus. Aedeagus shaft divided

near base, with two gonopores, or shaft divided toward apex and forming semicircles (Circuliferina),

with two gonopores. Connective anterior arms somewhat divergent, Y- or U-shaped; articulated with

aedeagus.

Female genitalia. Pygofer with macrosetae reduced or absent or with numerous macrosetae. Ovipositor

protruding or not protruding far beyond pygofer apex. First valvula convex or not strongly convex;

dorsal sculpturing pattern strigate, concatenate, reticulate, with rectangular shaped cells, granulose,

maculose, or imbricate (with overlapping scales); sculpturing reaching dorsal margin or submarginal;

without distinctly delimited ventroapical sculpturing. Second valvula broad and gradually tapered,

gradually broadened medially or subapically, or slender throughout; without dorsal median tooth; teeth

on apical 1/3 or more or restricted to apical 1/4 or less; teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: cosmopolitan. Opsiini feed on a wide variety of herbaceous and woody dicots. Opsius

stactogalus Fieber, 1866, a Palearctic species introduced in the New World, feeds on tamarix. This tribe

contains several species of economic importance. Neoaliturus ( Circulifer ) tenellus (Baker, 1896) is the

vector of beet curly top, tomato big bud, and 16SrV-16SrIX. N. (C.) haematoceps (Mulsant & Rey, 1855)

vectors sesame phyllody and 16SrV-16SrIX. Orosius orientals (Matsumura, 1914) transmits tomato

big bud, tobacco yellow dwarf, lucerne witches’ broom, legume little leaf, mosaic I, potato purple top

wilt, and witches’ broom of groundnuts. Orosius albicinctus Distant, 1918 transmits sesame phyllody.

Hishimonus phycitis (Distant, 1908) vectors eggplant little leaf. Hishimonoides sellatiformis Ishihara,

1965 and Hishimonus sellatus (Uhler, 1896) are vectors of mulberry dwarf and the latter leafhopper

is also a vector of Rhus yellows, jujube witches’ broom, and Cryptotaenia japonica witches’ broom.

Hishimonoides chinensis Anufriev, 1970 is a vector of jujube witches’ broom.

Remarks

Opsiini contains 36 genera and 303 species. The phylogenetic analyses here included eight representatives

of the tribe including the following that are included in analyses for the first time: Pseudophlepsius

Zachvatkin, 1924, Orosius Distant, 1918, Nesophrosyne Kirkaldy, 1907, Japananus Baft, 1931 (pre¬

viously in Scaphytopiini), and an undescribed genus near Libengaia from Zambia. The ML and Bayesian

analyses resolved it as monophyletic but with low branch support, and the parsimony analysis resolved it

as two separate monophyletic groups. The present phylogenetic analyses included more representatives

of Opsiini than previous analyses and are the first to suggest the tribe is monophyletic. Strong branch

support was recovered for some internal relationships within the tribe, but more representatives and

more branch support are needed in future analyses to test the current subtribal classification.

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Opsiini is a morphologically diverse tribe. Except for the bifurcate aedeagus with two gonopores, there

seem to be very few, if any, characters that can define the tribe entirely. It appears that the bifurcate

aedeagus has arisen at least three times in the evolutionary history of Deltocephalinae. The finding that

Japananus (previously included in Scaphytopiini based on the produced head and very broad gena)

is related to Opsiini rather than Scaphytopiini underscores the utility of the bifurcate aedeagus as a

predictor of relationships and helpful for classification at the tribal level. However, it is necessary to

corroborate any taxonomic hypothesis based on this character with other morphological characters or

molecular data. For example, Ascius DeLong, 1943 also has a bifurcate aedeagus, but it is retained in

Scaphytopiini because it shares the numerous distinct reflexed costal veins and the widely separated

anterior arms of the connective with Scaphytopius Ball, 1931, along with the produced head and broad

gena.

Several genera (. Afrascius Linnavuori, 1969; Japananus ; Kirkaldiella Osborn, 1935; Masiripius Dlabola,

1981; Navaia Linnavuori, 1960; Phlepsopsius Dlabola, 1979 and Pugla Distant, 1908) are transferred

to Opsiina from other tribes here because they share the bifurcate aedeagus and other similarities (e.g.,

brown irrorate color pattern) to some opsiines. Their placement to subtribe should be tested in future

studies. Dixianus Ball, 1918 and Lycioides Oman, 1949 are included in Circuliferina following the

suggestion of Emeljanov (1962) that Lycioides is closely related to Neoaliturus Distant, 1918, based in

part on the shape of the aedeagus.

Selected references

Z imm erman (1948), Emeljanov (1962), Ghauri (1966), Knight (1970a, b), Linnavuori (1969), Dmitriev

(2002), Dai et al. (2010), Dai et al. (2011), Bennett & O’Grady (2011).

Included subtribes:

Achaeticina Emeljanov, 1962

Fig. 38

Type genus: Achaetica Emeljanov, 1959.

Diagnosis

Achaeticina are small, squat, brachypterous leafhoppers, stramineous, greenish, or brownish in color.

They are distinguished from the other subtribes of Opsiini by the following combination of characters:

forewings brachypterous and truncate, male pygofer lobes without ventral pointed processes, valve thick

and strongly developed, valve wider than long, subgenital plates without macrosetae, aedeagal shafts

arising from base, and ovipositor protruding far beyond pygofer apex.

Description

Head. Head subequal to or wider than pronotum or distinctly narrower than pronotum. Discal portion

of crown glabrous with radial or longitudinal striae or shagreen. Anterior margin of head shagreen,

glabrous, irregularly textured, or foliaceous. Frontoclypeus not tumid; texture shagreen or glabrous.

Clypellus parallel-sided, tapering apically, or widening apically; apex following or slightly surpassing

normal curve of gena. Lorum subequal to or wider than clypellus near base. Antennal bases near middle

or posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely

incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledges absent or weakly

developed (carinate or weakly carinate). Ocelli absent or reduced in size; on anterior margin of head if

present.

Thorax. Pronotum lateral margin not carinate; lateral margin shorter than basal width of eye.

123

European Journal of Taxonomy 45: 1-211 (2013)

AA i

X* -

IfcirC *

L'v ■ ^ pDQr

Fig. 38. Opsiini (Achaeticina). Achaetica anabasidis Emeljanov, 1959. A-I. Standard views (see

Material & Methods).

124

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Wings. Forewing brachypterous or subbrachypterous; veins not raised.

Legs. Profemur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+0, 2+1,

2+2, or 2+2+1. Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; with short point of articulation with pygofer. Pygofer

dorsoapical margin not strongly incised; basolateral membranous cleft present; macrosetae absent

or reduced (< two rows). Subgenital plates free from each other, articulated with valve; without

macrosetae. Style broadly bilobed basally, median anterior lobe pronounced. Basal processes of the

aedeagus/connective absent. Aedeagus without basal hinge; shaft divided near base, with two gonopores.

Connective anterior arms somewhat divergent, Y-shaped; articulated with aedeagus.

Female genitalia. Pygofer with macrosetae reduced or absent. Ovipositor protruding far beyond pygofer

apex. First valvula convex, dorsal sculpturing pattern granulose, maculose, or imbricate ( with overlapping

scales); sculpturing submarginal; without distinctly delimited ventroapical sculpturing. Second valvula

gradually broadened medially or subapically; without dorsal median tooth; teeth restricted to apical 1/4

or less; teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Palearctic (Central Asia).

Remarks

Achaeticina contains 4 genera and 32 species.

Included genera

Achaetica Emeljanov, 1959

Chlidochrus Emeljanov, 1962

Diacra Emeljanov, 1961

Zapycna Emelj anov, 1968

Circuliferina Emeljanov, 1962

Fig. 39

Type genus: Circulifer Zachvatkin, 1935.

Diagnosis

Circuliferina are relatively small and slender ivory, greenish, or brownish leafhoppers. They are

distinguished from other subtribes of Opsiini by their relatively small, slender shape, mesal margin of

eye notched, forewing macropterous, subgenital plate with a lateral row of macrosetae, valve normal,

aedeagus bifurcate apically with branches forming semicircles, and ovipositor not protruding far beyond

pygofer apex.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae. Anterior margin of head shagreen. Frontoclypeus not tumid; texture shagreen.

Clypellus parallel-sided or tapering apically; apex following or slightly surpassing normal curve of

gena. Lorum subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral

(lower) corners of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely incised laterally;

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European Journal of Taxonomy 45: 1-211 (2013)

Fig. 39. Opsiini (Circuliferina). Neoaliturus carbonarius Mitjaev, 1971. A-I. Standard views (see

Material & Methods).

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

with fine erect seta beside laterofrontal suture. Antennal ledges absent. Ocelli present; close to eyes; on

anterior margin of head. Mesal margin of eye notched.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins

not raised; with or without reflexed costal veins; A1-A2 crossvein absent; apical venation not highly

reticulate.

Legs. Profemur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1.

Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; with short point of articulation with pygofer. Pygofer

basolateral membranous cleft present; macrosetae well differentiated into several rows. Subgenital

plates free from each other, articulated with valve; macrosetae uniseriate laterally. Style broadly bilobed

basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent. Aedeagus

without basal hinge; shaft divided toward apex, with two gonopores. Connective anterior arms somewhat

divergent, Y-shaped; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern granulose, maculose, or imbricate (with overlapping

scales); sculpturing reaching dorsal margin; without distinctly delimited ventroapical sculpturing.

Second valvula slender throughout; without dorsal median tooth; teeth on apical 1/3 or more; teeth

small, regularly or irregularly shaped.

Geography and ecology

Distribution: cosmopolitan, Neoaliturus ( Circulifer ) tenellus (Baker, 1896) is adventive in the New

World.

Remarks

Circuliferina contains 5 genera and 45 species.

Included genera

Concavifer Dlabola, 1960

Dixianus Ball, 1918 placement nov. (transferred from Athysanini)

Lycioides Oman, 1949 placement nov. (transferred from Athysanini)

Neoaliturus Distant, 1918

Pedarium Emeljanov, 1961

Eremophlepsiina Dmitriev, 2006

Fig. 40

Type genus: Eremophlepsius Zachvatkin, 1924.

Diagnosis

Eremophlepsiina are medium sized, robust, ivory or brownish leafhoppers. They are distinguished from

other subtribes of Opsiini by the macropterous forewings, pygofer lobes with ventral pointed processes,

valve thick and strongly developed, valve longer than wide and parabolically or lanceolate-parabolically

shaped, subgenital plates without macrosetae, and aedeagal shafts arising from base.

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Description

Head. Head subequal to, wider than, or distinctly narrower than pronotum. Discal portion of crown

glabrous with radial or longitudinal striae. Anterior margin of head glabrous, irregularly textured, with

numerous transverse striations. Frontoclypeus not tumid; texture shagreen. Clypellus widening apically;

apex following or slightly surpassing normal curve of gena. Lorum subequal to or wider than clypellus

near base. Antennal bases near middle or posteroventral (lower) corners of eyes. Antennae short, less

than 1.5 x width of head. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture.

Antennal ledges absent. Ocelli present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins

not raised; with or without reflexed costal veins; A1-A2 crossvein present; apical venation not highly

reticulate.

Legs. Profemur with AMI seta only; intercalary row with one row of five or more fine setae; row

AV with short, stout macrosetae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae

2+2+1. Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; with short point of articulation with pygofer; thick and

well-developed; longer than wide, parabolically or lanceolate-parabolically shaped. Pygofer dorsoapical

margin incised to near mid-length; basolateral membranous cleft present; macrosetae well differentiated

into several rows. Subgenital plates free from each other; articulated with valve; with thin hair-like setae

or with macrosetae scattered, irregularly arranged. Style broadly bilobed basally, median anterior lobe

pronounced. Basal processes of the aedeagus present, fused near base of aedeagus. Aedeagus without

basal hinge; shaft divided near base, with two gonopores. Connective anterior arms somewhat divergent,

Y-shaped; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor protruding far beyond pygofer apex.

First valvula not strongly convex; dorsal sculpturing pattern strigate, concatenate, reticulate, or with

rectangular shaped cells; sculpturing reaching dorsal margin; without distinctly delimited ventroapical

sculpturing. Second valvula broad, gradually tapered; without dorsal median tooth; teeth on apical 1/3

or more; teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Palearctic.

Remarks

Eremophlepsiina contains 2 genera and 5 species. It is closely related to Achaeticina.

Included genera

Eremophlepsius Zachvatkin, 1924

Pseudophlepsius Zachvatkin, 1924

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 40. Opsiini (Eremophlepsiina). Pseudophlepsius binotatus (Signoret, 1880). A-F, H-I. Standard

views (see Material & Methods). I. Caudoventral view. J. Connective. K. Style.

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European Journal of Taxonomy 45: 1-211 (2013)

Opsiina Emeljanov, 1962

Fig. 41

Type genus: Opsius Fkber, 1866.

Diagnosis

Opsiina are medium sized to large, robust leafhoppers, variable in color but often yellowish, greenish,

or brownish and with reticulate or irrorate brown markings; brown markings sometimes saddle-shaped.

They are distinguished from other subtribes of Opsiini by the macropterous forewings, subgenital plates

with a lateral row of macrosetae, subgenital plate apex often digitate and membranous, valve parabolically

shaped, aedeagal shafts arising from base, and ovipositor not protruding far beyond pygofer apex.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae. Anterior margin of head shagreen. Frontoclypeus not tumid; texture shagreen.

Clypellus widening apically; apex following or slightly surpassing normal curve of gena. Forum

subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral (lower)

corners of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely incised laterally; with fine

erect seta beside laterofrontal suture. Antennal ledges absent. Ocelli present; close to eyes; on anterior

margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins not

raised; without or with reflexed costal veins; A1-A2 crossvein absent or present; apical venation not

highly reticulate.

Fegs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1.

Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; with short point of articulation with pygofer. Pygofer

basolateral membranous cleft present; macrosetae well differentiated into several rows. Subgenital

plates free from each other, articulated with valve; macrosetae uniseriate laterally. Style broadly bilobed

basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent or reduced

or present, fused to base of aedeagus. Aedeagus without basal hinge; shaft divided near base, with two

gonopores. Connective anterior arms somewhat divergent, Y- or U-shaped; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern concatenate, reticulate, or with rectangular shaped

cells; sculpturing reaching dorsal margin; without or with indistinctly delimited ventroapical sculpturing.

Second valvula broad, gradually tapered or slender throughout; without dorsal median tooth; teeth on

apical 1/3 or more; teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: cosmopolitan. Opsius stactogalus Fieber, 1866 is adventive in the New World and feeds

on Tamarix.

Remarks

Opsiina is the largest of the 4 subtribes. It contains 25 genera and 218 species.

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 41. Opsiini (Opsiina). Opsius stactogalus Fieber, 1866. A-I. Standard views (see Material &

Methods).

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Included genera

Afrascius Linnavuori, 1969 placement nov. (transferred from Scaphytopiini)

Aladzoa Linnavuori, 1969

Alishania Vilbaste, 1969

Hishimonoides Ishihara, 1965

Hishimonus Ishihara, 1953

Introrsa Dai & Zhang, 2010

Japananus Ball, 1931 placement nov. (transferred from Scaphytopiini)

Kirkaldiella Osborn, 1935 placement nov. (transferred from Athysanini)

Lampridius Distant, 1918

Libengaia Linnavuori, 1969

Litura Knight, 1970

Masiripius Dlabola, 1981 placement nov. (transferred from Platymetopiina)

Naevus Knight, 1970

Navaia Linnavuori, 1960 placement nov. (previously unplaced in Deltocephalinae)

Nesophrosyne Kirkaldy, 1907

Norva Emeljanov, 1969

Opsianus Linnavuori, 1960

Opsius Fieber, 1866

Orosius Distant, 1918

Paralampridius Dai, Dietrich & Zhang, 2011

Phlepsopsius Dlabola, 1979 placement nov. (transferred from Athysanini)

Pugla Distant, 1908 placement nov. (transferred from Platymetopiina)

Satsumanus Ishihara, 1953

Xerophytacolus Stiller, 2012

Xerophytavorus Stiller, 2012

Pa.ra.1 imnin i Distant, 1908

Figs 42, 43

Type genus: Paralimnus Matsumura, 1902.

Diagnosis

Pa.ra.1 imnin i are small to medium sized leafhoppers. They can be identified by the combination of the

following characters: clypellus tapering apically or parallel-sided, lorum narrower than clypellus at

base; connective with anterior arms closely appressed, articulated with aedeagus; female first valvula

sculpturing imbricate or rarely maculose or granulose. The tribe is very similar morphologically to

the closely related Deltocephalini, from which it can be distinguished by the articulation between the

connective and aedeagus (fused in Deltocephalini), although a few species of Flexamia (Paralimnini)

have the connective fused to the aedeagus.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae, or shagreen. Anterior margin of head shagreen. Frontoclypeus not tumid or tumid;

texture shagreen. Clypellus parallel-sided or tapering apically; apex following or slightly surpassing

normal curve of gena. Forum distinctly narrower than clypellus near base. Antennal bases near middle

or posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely

incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledges absent. Ocelli present;

close to eyes; on anterior margin of head.

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Thorax. Pronotum lateral margin not carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous to brachypterous; if macropterous then appendix absent, reduced, or

present and restricted to anal margin; with 3 anteapical cells; veins not raised; without or rarely with

reflexed costal veins; crossvein r-ml often connected to R basad of its fork; A1-A2 crossvein absent;

apical venation not highly reticulate.

Legs. Profemur with AMI seta only; intercalary row with one row of five or more fine setae; row

AV usually with short, stout setae or rarely with relatively long macrosetae. Protibia dorsal surface

rounded, convex. Metafemur apex macrosetae 2+2+1. Metatarsomere I not expanded apically; plantar

setae simple, tapered or rarely with one or more platellae.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point.

Pygofer dorsoapical margin incised to near mid-length; basolateral membranous cleft present; macrosetae

well differentiated into several rows. Subgenital plates free from each other; usually articulated with

valve, or rarely fused; macrosetae uniseriate laterally, with two lateral rows of macrosetae, with some

irregularly arranged macrosetae near lateral margin, or reduced or absent. Style broadly bilobed basally,

median anterior lobe pronounced. Basal processes of the aedeagus/connective absent or reduced or

present, connected or articulated to connective or near base of aedeagus. Aedeagus with single shaft and

gonopore. Connective anterior arms closely appressed anteriorly, linear- or loop-shaped); nearly always

articulated with aedeagus, rarely fused to aedeagus (e.g., some Flexamia).

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding or rarely protruding

far beyond pygofer apex. First valvula convex or not strongly convex; dorsal sculpturing pattern

imbricate (with overlapping scales) or rarely granulose or maculose; sculpturing reaching dorsal margin

or submarginal; without or rarely with distinctly delimited ventroapical sculpturing. Second valvula

broad, gradually tapered; without dorsal median tooth; teeth on apical 1/3 or more; teeth small, regularly

or irregularly shaped or absent.

Geography and ecology

Distribution: cosmopolitan. Paralimnini feed on grasses or sedges and are often abundant in grassland

ecosystems. A case study of the North American genus Flexamia (Whitcomb & Hicks 1988; Dietrich et

al. 1997) examined the roles of host plants, geography, phenology, and phylogeny in the species level

diversification of the genus. These studies showed that related species groups feed on the same or similar

species of grass hosts and that host shifts occurred in the past, sometimes to distantly related grass

species. Geographic partitioning of the grass host ranges and phenological differences of the leafhopper

species contributed to the speciation of the group.

Psammotettix striatus (Linnaeus, 1758) and P. alienus (Dahlbom, 1850) transmit wheat dwarf virus and

mosaic virus of winter wheat.

Remarks

Paralimnini contains 139 genera and 931 species. It is closely related to Deltocephalini, with which

it shares a similar head morphology and the linear connective. Tetartostylini, which has a similar

connective articulated with the aedeagus but has the subgenital plates fused to the valve and with unique

apical spines, is also closely related to Paralimnini. Many species in understudied areas of the world

remain to be described. For example, in just three publications. Stiller (2009; 2010a, b) described 59

new species in three genera of Paral imnini from South Africa.

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European Journal of Taxonomy 45: 1-211 (2013)

Selected references

Emeljanov (1962), Knight (1974), Whitcomb & Hicks (1988), Webb & Heller (1990), Dietrich et al.

(1997), Kamitani (1999), Dmitriev (2004b), Stiller (2009, 2010a, b).

Included subtribes:

Aglenina Dmitriev, 2004

Fig. 42

Type genus: Aglena Amyot & Serville, 1843.

Diagnosis

Aglenina can be distinguished from other Paralimnini by their large size and black and yellow coloration.

Remarks

Aglena ornata Herrich-Schaffer, 1838 is the only species belonging to this subtribe. It occurs in the

Palearctic region.

Included genera

Aglena Amyot & Serville, 1843.

Paralimnina Distant, 1908

Fig. 43

Type genus: Paralimnus Matsumara, 1902.

Diagnosis

Paral imnin a can be distinguished from Aglenina by their smaller size and their coloration, which is

typically not black and yellow as in Aglena ornata.

Description

As in tribal description.

Remarks

Paralimnina contains 138 genera and 930 species.

Only the genera occurring in European Russia and adjacent territories were considered in the placement

of genera to subtribes recognized by Dmitriev (2004). All of the other genera that were left unplaced to

subtribe in Paralimnini are placed here in Paralimnina. Commellus Osborn & Ball, 1902 and Flexarida

Whitcomb & Hicks, 1993 were both listed as unplaced to tribe in Deltocephalinae by Oman et al. (1990)

and are placed here in Paral imn ina

Pasaremus is transferred to Paralimnina from Cicadulini, in which it was placed by Oman et al. (1990).

It does not possess the characteristic long sclerotized segment X or other features of Cicadulini, and

it possesses a tapered clypellus and a ‘linear’ connective articulated with the aedeagus which are

characteristic of Paralimnina.

Included genera

Acharis Emeljanov, 1966 placement nov. (previously unplaced in Paralimnini)

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 42. Paralimnini (Aglenina). Aglena ornata (Herrich-Schaffer, 1838). A-I. Standard views (see

Material & Methods). G. Connective and style shown with aedeagus.

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European Journal of Taxonomy 45: 1-211 (2013)

Fig. 43. Paralimnini (Paralimnina). A-C. Paralimnus angusticeps Zachvatkin, 1953. D-I. Psammotettix

striatus (Linnaeus, 1758). A-I. Standard views (see Material & Methods).

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Adarrus Ribaut, 1946 placement nov. (previously unplaced in Paralimnini)

Aflexia Oman, 1949 placement nov. (previously unplaced in Paralimnini)

Agudus Oman, 1938 placement nov. (previously unplaced in Paralimnini)

Alapus DeLong & Sleesman, 1929 placement nov. (previously unplaced in Paralimnini)

Altaiotettix Vilbaste, 1965

Anareia Vilbaste, 1965 placement nov. (previously unplaced in Paralimnini)

Anargella Emeljanov, 1972 placement nov. (previously unplaced in Paralimnini)

Araldus Ribaut, 1946 placement nov. (previously unplaced in Paralimnini)

Arocephalus Ribaut, 1946

Arthaldeus Ribaut, 1946

Auridius Oman, 1949 placement nov. (previously unplaced in Paralimnini)

Bhavapura Chalam & Rama Subba Rao, 2005 placement nov. (previously unplaced in Paralimnini)

Calamotettix Emeljanov, 1959 placement nov. (previously unplaced in Paralimnini)

Caloduferna Webb, 1980 placement nov. (previously unplaced in Paralimnini)

Canariotettix Lindberg, 1954 placement nov. (previously unplaced in Paralimnini)

Caphodellus Linnavuori & DeLong, 1976 placement nov. (previously unplaced in Paralimnini)

Cazenus Oman, 1949 placement nov. (previously unplaced in Paralimnini)

Cedarotettix Theron, 1975 placement nov. (previously unplaced in Paralimnini)

Changwhania Kwon, 1980 placement nov. (previously unplaced in Paralimnini)

Chelidinus Emeljanov, 1962 placement nov. (previously unplaced in Paralimnini)

Chloothea Emeljanov, 1959

Cleptochiton Emeljanov, 1959 placement nov. (previously unplaced in Paralimnini)

Coelestinus Emeljanov, 1962

Coganus Theron, 1978 placement nov. (previously unplaced in Paralimnini)

Commellus Osborn & Ball, 1902 placement nov. (previously unplaced in Deltocephalinae)

Connectivus Xing & Li, 2012 placement nov. (previously unplaced in Paralimnini)

Cosmotettix Ribaut, 1942 placement nov. (previously unplaced in Paralimnini)

Cribrus Oman, 1949 placement nov. (previously unplaced in Paralimnini)

Ctenotettix Novikov & Anufriev, 2005 placement nov. (previously unplaced in Paralimnini)

Cumbrenanus DeLong & Cwilda, 1984 placement nov. (previously unplaced in Paralimnini)

Destitutus Xing & Li, 2011 placement nov. (previously unplaced in Paralimnini)

Diplocolenus Ribaut, 1946 placement nov. (previously unplaced in Paralimnini)

Ebarrius Ribaut, 1946

Elginus Theron, 1975 placement nov. (previously unplaced in Paralimnini)

Emeljanovianus Dlabola, 1965

Enantiocephalus Haupt, 1926

Errastunus Ribaut, 1946

Erzaleus Ribaut, 1952 placement nov. (previously unplaced in Paralimnini)

Falcitettix Linnavuori, 1953 placement nov. (previously unplaced in Paralimnini)

Flexamia DeLong, 1926 placement nov. (previously unplaced in Paralimnini)

Flexarida Whitcomb & Hicks, 1993 placement nov. (previously unplaced in Deltocephalinae)

Futasiijinoidella Kwon & Lee, 1979 placement nov. (previously unplaced in Paralimnini)

Futasujinus Ishihara, 1953 placement nov. (previously unplaced in Paralimnini)

Giprus Oman, 1949 placement nov. (previously unplaced in Paralimnini)

Gobicuellus Dlabola, 1967 placement nov. (previously unplaced in Paralimnini)

Goldens Ribaut, 1946 placement nov. (previously unplaced in Paralimnini)

Hebecephalus DeLong, 1926 placement nov. (previously unplaced in Paralimnini)

Hebexa Oman, 1949 placement nov. (previously unplaced in Paralimnini)

Hengchunia Vilbaste, 1969 placement nov. (previously unplaced in Paralimnini)

Henschia Lethierry, 1892

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European Journal of Taxonomy 45: 1-211 (2013)

Hiltus Theron, 1974 placement nov. (previously unplaced in Paralimnini)

Histipagus Remane & Asche, 1980 placement nov. (previously unplaced in Paralimnini)

Ibadarrus Remane & Asche, 1980 placement nov. (previously unplaced in Paralimnini)

Jannius Theron, 1982 placement nov. (previously unplaced in Paralimnini)

Jassargus Zachvatkin, 1933

Jilinga Ghauri, 1974 placement nov. (previously unplaced in Paralimnini)

Kaszabinus Dlabola, 1965 placement nov. (previously unplaced in Paralimnini)

Kazachstanicus Dlabola, 1961 placement nov. (previously unplaced in Paralimnini)

Ladya Theron, 1982 placement nov. (previously unplaced in Paralimnini)

Laevicephalus DeLong, 1926 placement nov. (previously unplaced in Paralimnini)

Latalus DeLong & Sleesman, 1929

Lebradea Remane, 1959 placement nov. (previously unplaced in Paralimnini)

Lecacis Theron, 1982 placement nov. (previously unplaced in Paralimnini)

Lemellus Oman, 1949 placement nov. (previously unplaced in Paralimnini)

Maximianus Distant, 1918 placement nov. (transferred from Athysanini)

Mayawa Fletcher, 2000 placement nov. (previously unplaced in Paralimnini)

Mendrausus Ribaut, 1946 placement nov. (previously unplaced in Paralimnini)

Mendreus Ribaut, 1946 placement nov. (previously unplaced in Paralimnini)

Mesolimnella Dlabola, 1994 placement nov. (previously unplaced in Paralimnini)

Metagoldeus Remane & Asche, 1980 placement nov. (previously unplaced in Paralimnini)

Metalimnus Ribaut, 1948

Micrelloides Evans, 1973 placement nov. (previously unplaced in Paralimnini)

Micropedeticus Stiller, 2009 placement nov. (previously unplaced in Paralimnini)

Miraldus Tindberg, 1960 placement nov. (previously unplaced in Paralimnini)

Mocuola Emeljanov, 1964 placement nov. (previously unplaced in Paralimnini)

Mogangella Dlabola, 1957 placement nov. (previously unplaced in Paralimnini)

Mogangina Emeljanov, 1962 placement nov. (previously unplaced in Paralimnini)

Mongolojassus Zachvatkin, 1953 placement nov. (previously unplaced in Paralimnini)

Multiproductus Xing, Dai & Li, 2011 placement nov. (previously unplaced in Paralimnini)

Myittana Distant, 1908 placement nov. (previously unplaced in Paralimnini)

Nanosius Dlabola, 1974 placement nov. (previously unplaced in Paralimnini)

Naudeus Theron, 1982 placement nov. (previously unplaced in Paralimnini)

Nicolaus Lindberg, 1958 placement nov. (previously unplaced in Paralimnini)

Orocastus Oman, 1949 placement nov. (previously unplaced in Paralimnini)

Pantallus Emeljanov, 1961

Paraglena Emeljanov, 1997

Paragygrus Emeljanov, 1964 placement nov. (previously unplaced in Paralimnini)

Paralaevicephalus Ishihara, 1953 placement nov. (previously unplaced in Paralimnini)

Paralimnellus Emeljanov, 1972 placement nov. (previously unplaced in Paralimnini)

Paralimnoidella Kwon & Lee, 1979 placement nov. (previously unplaced in Paralimnini)

Paralimnus Matsumura, 1902

Paramesanus Dlabola, 1979 placement nov. (previously unplaced in Paralimnini)

Paramesus Fieber, 1866

Parapotes Emeljanov, 1975

Pasaremus Oman, 1949 placement nov. (transferred from Cicadulini)

Pazu Oman, 1949 placement nov. (previously unplaced in Paralimnini)

Peconus Oman, 1949 placement nov. (previously unplaced in Paralimnini)

Philaia Dlabola, 1952 placement nov. (previously unplaced in Paralimnini)

Phlebiastes Emeljanov, 1961 placement nov. (previously unplaced in Paralimnini).

Pinumius Ribaut, 1946 placement nov. (previously unplaced in Paralimnini)

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Platentomus Theron, 1980 placement nov. (previously unplaced in Paralimnini)

Pleargus Emeljanov, 1964 placement nov. (previously unplaced in Paralimnini)

Praganus Dlabola, 1949 placement nov. (previously unplaced in Paralimnini)

Pravistylus Theron, 1975 placement nov. (previously unplaced in Paralimnini)

Prosperellus Emeljanov, 1999 placement nov. (previously unplaced in Paralimnini)

Psammotettix Haupt, 1929

Pseudolausulus Wagner & Franz, 1961 placement nov. (previously unplaced in Paralimnini)

Pteropyx Haupt, 1927 placement nov. (previously unplaced in Paralimnini)

Quartausius Dlabola, 1974 placement nov. (previously unplaced in Paralimnini)

Quaziptus Kramer, 1965 placement nov. (previously unplaced in Paralimnini)

Restiobia Davies, 1988 placement nov. (previously unplaced in Paralimnini)

Rhoananus Dlabola, 1949

Rodezotettix Della Giustina & Wilson, 1995 placement nov. (previously unplaced in Paralimnini)

Rosemis Oman, 1949 placement nov. (previously unplaced in Paralimnini)

Samuraba Linnavuori, 1961 placement nov. (previously unplaced in Paralimnini)

Savanicus Dlabola, 1977 placement nov. (previously unplaced in Paralimnini)

Secopennis DeLong & Sleesman, 1929 placement nov. (previously unplaced in Paralimnini)

Sestrelicola Remane & Asche, 1980 placement nov. (previously unplaced in Paralimnini)

Sicistella Emeljanov, 1972 placement nov. (previously unplaced in Paralimnini)

Soracte Kirkaldy, 1907 placement nov. (previously placed in Platymetopiina)

Soractellus Evans, 1966 placement nov. (transferred from Deltocephalini)

Sorhoanus Ribaut, 1946

Spartopyge Young & Beirne, 1958 placement nov. (previously unplaced in Paralimnini)

Subhimalus Ghauri, 1971 placement nov. (previously unplaced in Paralimnini)

Takagiella Vilbaste, 1969 placement nov. (previously unplaced in Paralimnini)

Teinopterus Stiller, 2011 placement nov. (previously unplaced in Paralimnini)

Telnsus Oman, 1949 placement nov. (previously unplaced in Paralimnini)

Tiaratus Emeljanov, 1961 placement nov. (previously unplaced in Paralimnini)

Tigriculus Dlabola, 1961 placement nov. (previously unplaced in Paralimnini)

Triasargus Novikov & Anufriev, 2005 placement nov. (previously unplaced in Paralimnini)

Tytthuspiliis Stiller, 2011 placement nov. (previously unplaced in Paralimnini)

Umeqi Stiller, 2011 placement nov. (previously unplaced in Paralimnini)

Urganus Dlabola, 1965 placement nov. (previously unplaced in Paralimnini)

Vecaulis Theron, 1975 placement nov. (previously unplaced in Paralimnini)

Verdanus Oman, 1949

Vilargus Theron, 1975 placement nov. (previously unplaced in Paralimnini)

Yanocephalus Ishihara, 1953 placement nov. (previously unplaced in Paralimnini)

Pendarini Dmitriev, 2009

Fig. 44

Type genus: Pendarus Ball, 1927.

Diagnosis

Pendarini are medium sized to large ochraceous, yellowish, greenish, or brownish leafhoppers, sometimes

with brown irrorate markings or ramose pigment lines. As nymphs, they are distinguished from other

Deltocephalinae by the carinate crown-face transition, face with medial longitudinal carina in upper

part, acrometope with anterior margin tracing anterior carina of head, abdomen with 4-6 longitudinal

rows of very short macrosetae, and pygofer elongate (Dmitriev, 2009). Adults often have male pygofer

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with an acute apex and with one or more spines or teeth, and the aedeagus often with single or paired

apical or preapical processes.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae, or completely shagreen to base. Anterior margin of head shagreen or foliaceous.

Frontoclypeus not tumid; texture shagreen. Clypellus parallel-sided or widening apically; apex

following or slightly surpassing normal curve of gena. Lorum subequal to or wider than clypellus near

base. Antennal bases near middle or posteroventral (lower) corners of eyes. Antennae short, less than

1.5 x width of head. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture.

Antennal ledges absent or weakly developed (carinate or weakly carinate). Ocelli present; close to eyes;

on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than half basal width of eye.

Wings. Forewing macropterous or submacropterous; appendix present and restricted to anal margin or

rarely absent or reduced ( Dorydiella ); with 3 anteapical cells; veins not raised; without reflexed costal

veins; A1-A2 crossvein absent or present; apical venation not highly reticulate.

Legs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1.

Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae well differentiated into several rows;

sometimes elongate or extended posteriorly; often with one or more spines or processes arising dorsally

or ventrally. Subgenital plates free from each other; articulated with valve; macrosetae uniseriate

laterally. Style broadly bilobed basally; median anterior lobe pronounced. Aedeagus with single shaft

and gonopore. Connective anterior arms somewhat divergent, Y- or U-shaped; stem shorter than arms;

articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern strigate, granulose, or maculose; sculpturing

reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula broad,

gradually tapered; with or without dorsal median tooth; teeth on apical 1/3 or more; teeth small, regularly

or irregularly shaped.

Geography and ecology

Distribution: Nearctic and Neotropical regions. Species of this tribe feed on a variety of plants including

grasses, herbs, and woody hosts, including conifers. Several species apparently alternate between woody

oviposition hosts and herbaceous food plants. Some Pendarus and Paraphlepsius species specialize on

single species of perennial grasses. Chlorotettix species feed on grasses and sedges.

Remarks

Pendarini was recently described by Dmitriev (2009) based on nymphal characters and contains 8 genera

and 249 species. Five genera ( Bandaromimns, Tropicanus, Ileopeltus, Chlorotettix and Copididonus)

are added to the tribe here based on the results of our phylogenetic analyses and based on observations

of nymphs of Chlorotettix which share the elongate pygofer of other Pendarini. A clade including

representatives of Paraphlepsius Baker, 1897, Dorydiella Baker, 1897, Bandaromimus Linnavuori,

1959, Tropicanus DeLong, 1944, Chlorotettix , and Copididonus was consistently resolved in all analyses

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

!§a

V.nV.'iki

Fig. 44. Pendarini. Pendarus magnus (Osborn & Ball, 1897). A-I. Standard views (see Material &

Methods).

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European Journal of Taxonomy 45: 1-211 (2013)

and received moderate branch support in Bayesian analyses. Pendarus was not included in the analyses

but is undoubtedly related to this group. Ileopeltus is included based on the comments of Cwikla (1988b)

suggesting the genus is closely related to Chlorotettix. Further study is needed to determine if other

genera should be placed in this tribe and to discover additional morphological characters to aid in its

diagnosis. The tribe is an exclusively New World group as currently understood.

Selected references

DeLong (1945), Hamilton (1975a), Cwilda (1988a, b), Dmitriev (2009).

Included genera

Bandaromimus Linnavuori, 1959 placement nov. (transferred from Athysanini)

Chlorotettix Van Duzee, 1892 placement nov. (transferred from Athysanini)

Copididonus Linnavuori, 1954 placement nov. (transferred from Athysanini)

Dorydiella Baker, 1897

Ileopeltus Cwikla, 1988 placement nov. (previously unplaced in Deltocephalinae)

Paraphlepsius Baker, 1897

Pendarus Ball, 1927

Tropicanus DeLong, 1944 placement nov. (transferred from Athysanini)

Penthimiini Kirschbaum, 1868

Fig. 45

Type genus: Penthimia Germar, 1821.

= Thaumatoscopini Baker, 1923.

Diagnosis

Penthimiini are medium sized to large, squat, robust, often black or brown leafhoppers; often with

ventral part of face and/or entire ventral side flattened and dorsal side convex. They can be identified

by the ocelli on the crown and often distant from the eyes, strong antennal ledge, dorsally flattened and

carinate protibia, and forewing with appendix large and extending around the wing apex.

Description

Head. Head subequal to, wider than, or distinctly narrower pronotum. Discal portion of crown glabrous

with radial or longitudinal striae. Anterior margin of head with numerous transverse striations or carinae,

or foliaceous. Head and pronotum usually sloping anteriorly. Face short, broad. Frontoclypeus tumid

or not; texture striate or glabrous. Clypellus widening apically; apex following or slightly surpassing

normal curve of gena. Lorum subequal to or wider than clypellus near base. Antennal bases near upper

or anterodorsal corners of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely incised

laterally; with fine erect seta beside laterofrontal suture. Antennal ledges strongly developed (with a

distinct ledge); sometimes forming lateral part of anterior margin of head. Ocelli present; distant from

eyes; on crown.

Thorax. Pronotum lateral margin carinate; lateral margin shorter or longer than basal width of eye.

Wings. Forewing macropterous or submacropterous; sometimes coriaceous, setose, punctate, or

tuberculate; appendix large and extending around wing apex or rarely restricted to anal margin; with

3 anteapical cells; veins not raised; without reflexed costal veins; A1-A2 crossvein absent or present;

apical venation not highly reticulate.

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Legs. Pro femur with AMI seta only or with AMI and with one or more additional proximal setae;

intercalary row with one row of five or more fine setae; row AV usually with thin, hair-like setae or

without setae, more rarely with short, stout setae or with relatively long macrosetae. Protibia dorsal

surface flat, AD and PD margins sharply carinate or at ~90° (not rounded). Metafemur apex macrosetae

2+2+1. Metatarsomere I not expanded apically, plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae well differentiated into several rows.

Subgenital plates free from each other; articulated with valve; without macrosetae or macrosetae

scattered, irregularly arranged. Style linear, median anterior lobe not pronounced or broadly bilobed

basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent or reduced

or present, connected or articulated to connective or near base of aedeagus. Aedeagus with single shaft

and gonopore. Connective anterior arms somewhat divergent, Y- or U-shaped; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding or rarely protruding

(e.g., Penthimidia) far beyond pygofer apex. First valvula convex; dorsal sculpturing pattern strigate,

concatenate, or reticulate; sculpturing reaching dorsal margin; without distinctly delimited ventroapical

sculpturing. Second valvula abruptly broadened medially or subapically or broad, gradually tapered;

with or without dorsal median tooth; teeth on apical 1/3 or more; teeth large, regularly shaped or small,

regularly or irregularly shaped.

Geography and ecology

Distribution: cosmopolitan, poorly represented in North Am erica Penthimiini are collected on trees,

shrubs, and herbaceous vegetation.

Remarks

Penthimiini contains 46 genera and 203 species. It is a relatively early diverging lineage of Deltocephalinae

but has the typical derived deltocephaline characters of the male genitalia (pygofer with basolateral

membranous cleft, Y-shaped connective, and style broadly bilobed basally). The phylogenetic analyses

included 4 exemplars of the tribe as circumscribed here ( Penthimia , Penthimidia , Penthimiola , and Jafar

included in analyses; Citorus Stal, 1866 is removed from the tribe based on these analyses). Parsimony

analyses resolved the former 3 exemplars as monophyletic and sister to Magnentius Singh-Pruthi, 1930,

while ML and Bayesian analyses resolved Penthimidia and Penthimiola together and Penthimia in an

uncertain position elsewhere in the tree. It is unclear why Penthimia does not always group with other

penthimiines in phylogenetic analyses, but based on the morphological evidence supporting Penthimiini

as a taxon, we suspect that the phylogenetic placement of Penthimia in the ML and Bayesian analyses is

artifactual. Jafar was also included for the first time and grouped with one or two undescribed African

genera, somewhat distant from other Penthimiini but with no branch support. Because its phylogenetic

position is uncertain, and because it possesses several characters diagnostic for the tribe (large and

robust body, ocelli on crown, inflated frontoclypeus, strong antennal ledges, and dorsally flattened and

bicarinate protibiae), it is retained in the tribe pending further study.

Citorus , which was previously included in Penthimiini based on its body shape and large forewing

appendix, was found with strong branch support to be more closely related to Selenocephalini (Dwightla

and Selenocephalus). Based on the phylogenetic results and on some morphological characters that differ

from Penthimiini (anterior margin of the head with distinct sharp carinae, ocelli on anterior margin of

head and close to eyes, antennal ledge not very strong or ledge-like) it is transferred to Selenocephalina

here. The squat, robust body form and large appendix are similar to Penthimiini but these are apparently

convergent characters in this case.

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European Journal of Taxonomy 45: 1-211 (2013)

Fig. 45. Penthimiini. Penthimia americana Fitch, 1851. A-I. Standard views (see Material & Methods).

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Selected references

Evans (1966, 1972), Linnavuori (1977).

Included genera

Alopenthimia Evans, 1972

Amberbakia Distant, 1912

Chanohirata Hayashi & Machida, 1996

ChinaellaEvms , 1935

Ectopiocephalus Kirkaldy, 1906

Eovulturnops Evans, 1947

Eupenthimia Evans, 1972

Eusallya Evans, 1972

Foroa Linnavuori, 1977

Gressittella Evans, 1972

Haranga Distant, 1908

Irenaella Linnavuori, 1977

Jafar Kirkaldy, 1903

Kronos Distant, 1917

Ledroides Dammerman, 1910

Macutella Evans, 1972

Malichus Distant, 1918

Megalopenthimia Evans, 1954

Musosa Linnavuori, 1977

Neodartellus Evans, 1955

Neodartus Melichar, 1903

Neopenthimia Evans, 1972

Neovulturnus Evans, 1937

Nielsoniella Linnavuori, 1977

Nortoides Evans, 1972

Nubelella Evans, 1972

Nubelloides Evans, 1972

Osella Evans, 1972

Penthimia Germar, 1821

Penthimidia Haglund, 1899

Penthimiella Evans, 1972

Penthimiola Linnavuori, 1959

Pen th im i ops is Evans ,1972

Pentria Evans, 1972

Piorella Evans, 1972

Platyscopus Evans, 1941

Reticuluma Cheng & Li, 2005

Sidelloides Evans, 1972

Tambila Distant, 1908

ThaumatopoidesEvms, 1947

Thaumat os copus Kirkaldy, 1906

Tolasella Evans, 1972

Tomaloides Evans, 1972

Uzelina Melichar, 1903

Vertigella Evans, 1972

Vulturnus Kirkaldy, 1906

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European Journal of Taxonomy 45: 1-211 (2013)

Phlepsiini tribe nov.

urn:lsid:zoobank.org:act:6F2DC22C-BD71-4BFD-BEEE-8AF8BA2819B4

Fig. 46

Type genus: Phlepsius Fieber, 1866.

Diagnosis

Phlepsiini are medium sized to large, robust, ivory to brown or dark brown leafhoppers, often with brown

irrorate markings on the forewing and body. They can be identified by the head distinctly narrower

than pronotum, antennal ledges carinate, lorum long and relatively large, clypellus long and widening

apically, pronotum lateral margin longer than half basal width of eye, profemur row AM sometimes with

1 or more setae basad of AMI, and nymph with short py gofer and with 6 rows of abdominal macrosetae.

Description

Head. Head narrower than pronotum. Discal portion of crown glabrous with radial or longitudinal

striae. Anterior margin of head with one to several transverse carinae or striae, irregularly textured, or

partly glabrous. Frontoclypeus not tumid; not particularly broad; texture shagreen. Clypellus widening

apically; relatively long; apex following or slightly surpassing normal curve of gena. Lorum large;

subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral (lower)

comers of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely incised laterally; with fine

erect seta beside laterofrontal suture. Antennal ledges weakly developed (carinate or weakly carinate).

Ocelli present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin longer than half basal width of eye, somet im es

longer than entire basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins not

raised; A1-A2 crossvein absent or present; apical venation not highly reticulate; with numerous false

veins and irrorate markings.

Legs. Profemur with AMI seta only or rarely with extra macrosetae basad of AMI; intercalary row with

one row of five or more fine setae; row AV with short, stout setae or with longer setae. Protibia dorsal

surface rounded, convex; dorsal macrosetae 4+4. Metafemur apex macrosetae 2+2+1. Metatarsomere I

not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae well differentiated into several rows;

without spines or processes. Subgenital plates free from each other; articulated with valve; macrosetae

uniseriate laterally. Style broadly bilobed basally; median anterior lobe pronounced. Basal processes

of the aedeagus/connective absent or reduced, or present, connected or articulated to connective or

near base of aedeagus. Aedeagus with single shaft and gonopore. Connective anterior arms somewhat

divergent, Y-shaped; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern strigate, reticulate, or concatenate; sculpturing

reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula broad,

gradually tapered; without dorsal median tooth; teeth on apical 1/3 or more; teeth small, regularly or

irregularly shaped.

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Fig. 46. Phlepsiini. Phlepsius intricatus (Herrich-Schaffer, 1838). A-F, H-I. Standard views (see

Material & Methods). J. Connective, ventral view. K. Style, ventral view.

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Geography and ecology

Distribution: Palearctic, Afrotropical, Oriental, Nearctic, and Neotropical regions. Phlepsiini feed on

woody or herbaceous dicots. Texananus lathropi (Baker, 1925), T. latipex DeLong 1943, T. oregonus

(Ball, 1931), 72 pergradus DeLong, 1938 and T. spatulatus (VanDuzee, 1892) are vectors of the western

strain of North American aster yellows.

Remarks

Phlepsiini currently only includes 4 genera and 81 species. Phylogenetic analyses identified a well-

supported clade including Phlepsius , Excultanus , and Korana which was consistently found in a clade

along with Scaphoideini, Drabescini, and some undescribed genera. Oman (1941) recognized a tribe,

“Phlepsiini”, which is considered here to be a nomen nudum (see Materials and Methods: Taxonomic

Treatments, above). In addition to Phlepsius (which included the species of Texananus and Excultanus ),

Oman included Norvellina, Remadosus, Dorydiella, Dixianus, Paraphlepsius, and Phlepsanus. The

phylogenetic analyses here do not support this broad interpretation of the tribe. Orientus has several

characters diagnostic for this tribe but is smaller and more slender than the other genera. It was also

not placed within the Phlepsiini clade in the phylogenetic analyses, but was found on a long branch

of indeterminate relationship near the early diverging lineages of Deltocephalinae. It is retained in

Athysanini at this time pending further analyses.

Selected references

DeLong (1938, 1939), Crowder (1952), Theron (1980), McKamey (2000, 2003a).

Included genera

Excultanus Oman, 1949 placement nov. (previously unplaced in Athysanini)

Korana Distant, 1910 placement nov. (previously unplaced in Deltocephalinae)

Phlepsius Fieber, 1866 placement nov. (previously placed in Platymetopiina)

Texananus Ball, 1918 placement nov. (previously placed in Platymetopiina)

Scaphoideini Oman, 1943

Fig. 47

Type genus: Scaphoideus Uhler, 1889.

Diagnosis

Scaphoideini are medium sized (usually 4-7 mm), often slender, ivory, yellowish, ochraceous, or brown

leafhoppers, often marked with orange, brown, or black on head, pronotum, or forewings. None of the

following diagnostic characters are present in all taxa, but some combination of these characters is

present in all and a few (*) appear to be unique to this tribe: head narrower than pronotum, produced;

frontoclypeus long, narrow; antennae long; body slender; head and wings often with brown, orange,

ochraceous, or ivory markings; forewing with one or more darkly pigmented reflexed veins in vicinity

of outer anteapical cell; profemur row AV setae absent or reduced (without stout setae); metatibia

macrosetae in row PD long, as long as or longer than 0.5 x length of protibia*; male or female py gofer

with dense tufts of long fine or regular setae*; subgenital plate apex membranous or long, digitate, and

somewhat membranous or weakly sclerotized; subgenital plates with long fine setae laterally and/or

dorsally (also occurs in many other deltocephaline tribes); basal processes of the aedeagus or connective

sometimes present, connected or articulated to base of aedeagus or apex of connective stem.

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Description

Head. Head often narrower than pronotum, sometimes subequal to or wider than pronotum; anterior

margin shagreen; crown rounded to face or angulate. Discal portion of crown glabrous with radial or

longitudinal striae. Frontoclypeus not tumid; long, narrow; shagreen. Clypellus widening apically.

Lorum subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral

(lower) corners of eyes or near anterodorsal (upper) comers of eyes. Antennae long (longer than width

of head). Gena obtusely incised or not incised laterally; with fine erect seta beside laterofrontal suture.

Antennal ledge absent. Ocelli close to eyes.

Thorax. Pronotum lateral margin carinate; lateral margin shorter or longer than half basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin; veins sometimes bordered or

highlighted at apices with brown coloration; with 3 anteapical cells; often with pigmented reflexed

costal veins; usually without A1-A2 crossvein; apical venation not highly reticulate.

Legs. Profemur row AM with AMI seta only; intercalary row with one row of five or more fine setae;

row AV setae usually reduced or absent, or sometimes short, stout setae. Protibia dorsal surface rounded,

convex. Metatibia macrosetae in rows AD and PD often very long, row PD setae often equal to or longer

than 0.5 x length of protibia. Metatarsomere I long. Metafemur apical setae 2+2+1.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer with basolateral membranous cleft; often with many long thick or fine setae somet im es

arranged in dense tufts or groups. Subgenital plate with macrosetae absent, uniseriate laterally, or

uniseriate, distant from lateral margin; often with long fine hairs laterally and/or dorsally; plate

apex often long, digitate, and/or membranous or filamentous. Style broadly bilobed basally, median

anterior lobe pronounced. Basal processes of the aedeagus/connective absent or present, connected to

or articulated with connective or near base of aedeagus. Aedeagus with single shaft and gonopore.

Connective usually Y-shaped, sometimes linear or with anterior arms closely appressed; articulated

or rarely fused (Sikhamani, Thryaksha , and Univagris) with aedeagus, or aedeagus dissociated from

connective ( Scaphoideus ).

Female genitalia. Pygofer with numerous macrosetae, sometimes with dense tufts or groups of fine or

thick setae. Ovipositor not protruding far beyond pygofer apex. First valvula convex; dorsal sculpturing

pattern often concatenate, sometimes strigate or reticulate; reaching dorsal margin. First valvula

ventroapical sculpturing indistinctly delimited or absent. Second valvula abruptly broadened medially

or subapically or broad, gradually tapered; with or without dorsal median tooth; dorsoapical teeth large

or small, regularly shaped; teeth on apical 1/3 or more.

Geography and ecology

Distribution: cosmopolitan. Scaphoideini are among the most commonly encountered forest-dwelling

deltocephalines throughout the tropical and warm-temperate regions of the world. Nymphs and adults

are often collected on understory grasses or herbaceous vegetation but most species appear to require

woody hosts for oviposition. Scaphoideus titanus Ball, 1932 is a serious pest and vector of grape

flavescence doree in Europe and of chrysanthemum yellows. S. luteolus Van Duzee, 1894 is a vector of

elm phloem necrosis in the United States. Osbornellus borealis DeLong & Mohr, 1936 is a vector of

western X disease.

Remarks

Scaphoideini contains 61 genera and 631 species. Oman et al. (1990) included 12 genera, significantly

fewer than the circumscription here. Several genera included previously (Acunasus DeLong, 1945,

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European Journal of Taxonomy 45: 1-211 (2013)

Fig. 47. Scaphoideini. Scaphoideus omani sp. nov. A-F, H-I. Standard views (see Material & Methods).

J. Lateral view of connective and connective processes. K. 9 stemite VII. L. Style, ventrally.

M. Ventral view of connective and connective processes.

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Danbara Oman, 1949, Sobara Oman, 1949 and Twiningia Ball, 1931) are not included here. Twiningia

was included in the phylogenetic analyses here and was found to be closely related to Platymetopius

Burmeister, 1838 (Athysanini), and the others are excluded because they do not possess characters

considered here to be diagnostic for the tribe.

The phylogenetic analyses presented here consistently resolved a clade including Scaphoideini sensu

novo plus Drabescini. This clade was in turn resolved as sister to Phlepsiini. Scaphoideini was resolved

as paraphyletic with respect to Drabescini and three undescribed genera, but with little or no branch

support for the basal branches of the group. Genera of Scaphoideini included in phylogenetic analyses

here that were not included previously are Loipothea Linnavuori, 1969, Omanana DeLong, 1942,

Scaphoidophyes Kirlcaldy, 1906, Anoplotettix Ribaut, 1942 and Mimotettix Matsumura, 1914. Further

taxon sampling and molecular data may help to resolve the relationships within this large tribe and its

relationship to the other related tribes, but the available evidence suggests that this is a lineage distinct

from other Athysanini, the tribe in which many of the genera were previously placed. The circumscription

of the tribe here is based on the phylogenetic analyses and the morphological characters identified

here as diagnostic for most members of the group. Representatives of most genera of Athysanini were

examined or were reviewed from the literature for their potential placement in Scaphoideini. Further

adjustments to the circumscription may be needed after more detailed study of the group.

The Perugrampta group (including Perugramptella, Penthigrampta , and Stenogrampta) is placed here

pending further study. These genera possess the following characters that are diagnostic for Scaphoideini:

pro femur row AV setae absent, darkly pigmented reflexed branches of R vein, male or female pygofer

or subgenital plates with tufts of fine or thick setae, subgenital plates with long fine setae laterally, and

basal processes of the aedeagus or connective present and connected to base of aedeagus.

Selected references

Barnett (1977, 1979, 1980), Martinson (1977), Knight & Fletcher (2007), Viraktamath & Mohan (1994,

2004), Stiller (2001), Zhang & Dai (2006), Freytag (2008), Dominguez & Godoy (2010), Dai & Dietrich

( 2011 ).

Included genera

Acastroma Finnavuori, 1969 placement nov. (transferred from Athysan ini )

Afroidens Finnavuori, 1961 placement nov. (previously unplaced in Deltocephalinae)

Alemaia Heller & Finnavuori, 1968 placement nov. (previously unplaced in Deltocephalinae)

Amimenus Ishihara, 1953 placement nov. (previously unplaced in Deltocephalinae)

Anoplotettix Ribaut, 1942 placement nov. (previously placed in Platymetopiina)

Bampnrius Dlabola, 1977 placement nov. (transferred from Athysanini)

Bonus Distant, 1908 placement nov. (transferred from Athysanini)

Burakia Kocalc, 1981 placement nov. (previously placed in Platymetopiina)

Cantura Oman, 1949 placement nov. (previously unplaced in Deltocephalinae)

Caphodns Oman, 1938 placement nov. (previously unplaced in Deltocephalinae)

Capoideus Theron, 1974 placement nov. (previously unplaced in Deltocephalinae)

Coroticus Distant, 1918 placement nov. (transferred from Scaphytopiini)

Dariena Finnavuori & DeFong, 1977 placement nov. (transferred from Athysanini)

Delospilopterus Stiller, 2001 placement nov. (transferred from Athysanini)

Euscelidella Evans, 1954 placement nov. (previously unplaced in Deltocephalinae)

Ferganotettix Dubovsky, 1966 placement nov. (transferred from Athysanini)

Grammacephalus Haupt, 1929 placement nov. (transferred from Scaphytopiini)

Gunghuyana Distant, 1910 placement nov. (previously unplaced in Deltocephalinae)

Loipothea Finnavuori, 1969 placement nov. (previously unplaced in Deltocephalinae)

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Longicornis Li & Song, 2008 placement nov. (transferred from Athysanini)

Madagella Evans, 1954 placement nov. (previously placed in Platymetopiina)

Melanetettix Knight & Fletcher, 2007 placement nov. (transferred from Athysanini)

Mesotettix Matsumura, 1914 placement nov. (previously unplaced in Deltocephalinae)

Mimotettix Matsumura, 1914 placement nov. (transferred from Athysanini)

Mirzayansus Dlabola, 1979 placement nov. (transferred from Athysanini)

Monobazus Distant, 1908 placement nov. (transferred from Athysanini)

Moorada Ghauri, 1975 placement nov. (transferred from Athysanini)

Nataretus Theron, 1980 placement nov. (transferred from Athysanini)

Neolimnus Linnavuori, 1953 placement nov. (transferred from Athysanini)

Neophlepsius L i nnavuori, 1955 placement nov. (transferred from Athysanini)

Omanana DeLong, 1942 placement nov. (transferred from Athysanini)

Osbornellus Ball, 1932 placement nov. (transferred from Athysanini)

Pachodus Linnavuori, 1961 placement nov. (transferred from Athysanini)

Par allygus Melichar, 1903 placement nov. (transferred from Athysanini)

Par amelia Evans, 1954 placement nov. (previously placed in Platymetopiina)

Penthigrampta Dietrich & Rakitov, 2002 placement nov. (transferred from Penthimiini)

Perugrampta Kramer, 1965 placement nov. (transferred from Penthimiini)

Perugramptella Dietrich & Rakitov, 2002 placement nov. (transferred from Penth im i ini )

Phlogotettix Ribaut, 1942 placement nov. (previously unplaced in Deltocephalinae)

Phlogothamnus Ishihara, 1961 placement nov. (previously unplaced in Deltocephalinae)

Platyretus Melichar, 1903 placement nov. (transferred from Athysa nin i)

Premanus DeLong, 1944 placement nov. (transferred from Athysa nin i)

Prescottia Ball, 1932 placement nov. (previously unplaced in Deltocephalinae)

Scaphodhara Viraktamath & Mohan, 1994 placement nov. (previously unplaced in Deltocephalinae)

Scaphoidella Vilbaste, 1968 placement nov. (transferred from Athysanini)

Scaphoideus Uhler, 1889 placement nov. (transferred from Athysanini)

Scaphoidophyes Kirkaldy, 1906 placement nov. (previously unplaced in Deltocephalinae)

Scaphoidula Osborn, 1923 placement nov. (previously unplaced in Deltocephalinae)

Scaphomonus Viraktamath, 2009 placement nov. (transferred from Athysanini)

Sikhamani Viraktamath & Webb, 2006 placement nov. (transferred from Scaphytopiini)

Soleatus DeLong, 1971 placement nov. (previously unplaced in Deltocephalinae)

Spathanus DeLong, 1945 placement nov. (transferred from Athysanini)

Stenogrampta Dietrich & Rakitov, 2002 placement nov. (transferred from Pent him i ini )

Sudhamruta Viraktamath & Anantha Murthy, 1999 placement nov. (transferred from Scaphytopiini)

Tbilisica Dlabola, 1958 placement nov. (previously placed in Platymetopiina)

Thamnophryne Kirkaldy, 1907 placement nov. (previously unplaced in Deltocephalinae)

Thryaksha Viraktamath & Anantha Murthy, 1999 placement nov. (transferred from Scaphytopiini)

Tongdotettix Kwon, 1980 placement nov. (previously placed in Platymetopiina)

Tuakamara Webb, 1980 placement nov. (previously placed in Platymetopiina)

Univagris Viraktamath & Anantha Murthy, 1999 placement nov. (transferred from Scaphytopiini)

Wanritettix Vilbaste, 1969 placement nov. (transferred from Athysanini)

Scaphytopiini Oman, 1943

Fig. 48

Type genus: Scaphytopius Ball, 1931.

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Diagnosis

Scaphytopiini are small to medium sized usually brownish, or sometimes black, yellow, or greenish

leafhoppers, sometimes with irrorate markings and often with reflexed costal veins on the forewing.

They can be identified by the anteriorly produced and pointed head, gena not incised laterally and visible

behind eyes in dorsal view, frontoclypeus elongate, reflexed costal veins on the forewing, valve large,

and connective broad, W- or U-shaped, and without or with a short stem.

Description

Head. Head narrower than pronotum; often produced and pointed anteriorly. Discal portion of crown

glabrous with radial or longitudinal striae. Anterior margin of head angulate; texture shagreen,

subcarinate, or with transverse striae. Frontoclypeus elongate; texture shagreen. Clypellus widening

apically; apex following or slightly surpassing normal curve of gena. Lorum narrower, subequal to, or

wider than clypellus near base. Antennal bases near middle or posteroventral (lower) corners of eyes.

Antennae short, less than 1.5 x width of head. Gena broad; not incised laterally; visible behind eyes in

dorsal view; with fine erect seta beside laterofrontal suture. Antennal ledges absent. Ocelli present; close

to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins not

raised; with numerous reflexed costal veins; A1-A2 crossvein absent or present; apical venation not

highly reticulate.

Legs. Pro femur with AMI seta only; intercalary row with one row of five or more fine setae; row AV with

short, stout setae or with relatively long macrosetae. Protibia dorsal surface rounded, convex. Metafemur

apex macrosetae 2+2+1. Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve large; parabolically shaped; articulated with pygofer and subgenital plates; lateral

margin short, articulating with pygofer at a point. Pygofer dorsoapical margin not strongly incised

or incised to near mid-length; basolateral membranous cleft present; macrosetae well differentiated

into several rows; without processes. Subgenital plates free from each other, articulated with valve;

without macrosetae or with macrosetae uniseriate laterally. Style elongate; broadly bilobed basally,

median anterior lobe pronounced. Basal processes of the aedeagus/connective often present, articulated

to connective or near base of aedeagus. Aedeagus with single shaft and gonopore, or rarely (. Ascius )

divided near apex, with two gonopores. Connective anterior arms widely divergent, broadly W- or

U-shaped, or somewhat divergent, Y- or U-shaped; stem usually absent; articulated with aedeagus or

aedeagus dissociated from connective and attached only by thin membrane. Segment X sclerotized.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex, dorsal sculpturing pattern strigate, concatenate, reticulate, or imbricate

(with overlapping scales); sculpturing reaching dorsal margin; without distinctly delimited ventroapical

sculpturing. Second valvula abruptly broadened medially or subapically or broad, gradually tapered;

without dorsal median tooth; teeth on apical 1/3 or more; teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Nearctic and Neotropical regions. Host plants are herbaceous and woody dicots.

Scaphytopius ( Cloanthanus ) acutus (Say, 1830) is a vector of alfalfa witches’ broom, eastern and western

X-diseases of peach, little cherry, soybean bud proliferation, clover phyllody, and western strain of North

American aster yellows. S. ( Convelinus ) irroratus (Van Duzee, 1910) vectors the western strain of North

American aster yellows. S. (< Cloanthanus ) magdalensis (Provancher, 1889) transmits blueberry stunt.

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Fig. 48. Scaphytopiini. Scaphytopius acutus (Say, 1830). A-I. Standard views (see Material & Methods).

G. Connective and style shown with connective processes. J. Lateral view of connective, style and

connective processes.

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

S. ( Convelinus ) fuliginosus (Osborn, 1923) is a vector of machismo disease of legumes in Mexico and

South America. S. ( Convelinus ) nitridus (DeLong, 1943) is a vector of western X-disease of stonefruits

and celery in North America.

Remarks

Scaphytopiini contains 3 genera and 183 species. Based on phylogenetic analyses, the tribe as previously

delimited (Oman et al. 1990, Webb & Godoy 1993, Dmitriev 2002) was polyphyletic. Representatives

of Japananus, Nesothamnus Linnavuori, 1959, Proceps Mulsant & Rey, 1855, Stymphalus Stal,

1866 and 2 representatives of Scaphytopius were included in analyses and the genera were found in

different areas of the tree in most analyses. There was strong support for the branch including the two

Scaphytopius species, but its relationship to other Deltocephalinae was poorly resolved. In Bayesian

and ML analyses, Scaphytopius was resolved within a clade including the athysanine genus Eutettix and

3 others (PP=0.98). Japananus appears to be related to Opsiini, which is consistent with its bifurcate

aedeagus. Ascius also has a bifurcate aedeagus, but it is retained in Scaphytopiini because it shares

the numerous distinct reflexed costal veins and the widely separated anterior arms of the connective

with Scaphytopius , along with the produced head and broad gena. Nesothamnus was often resolved

near Idioceromimus Dietrich & Rakitov, 2002 in our analyses. Proceps is grouped with Cicadulini with

strong branch support. Stymphalus , which is placed here in the new tribe Vartini, usually grouped with

an undescribed genus from Thailand (DEL 112) and some Mukariini.

The characters formerly used to define the tribe, the produced head and the gena not incised laterally

and visible behind the eye in dorsal view, appear to have arisen independently multiple times in

Deltocephalinae and are therefore not reliable characters on their own at the tribal level. The tribe is

interpreted here to contain only three genera from the New World.

Selected references

Hepner (1947), Oman (1949), Linnavuori (1959), Webb & Godoy (1993).

Included genera

Ascius DeLong, 1943

Scaphytopius Ball, 1931

Tenuarus DeLong, 1944

Selenocephalini Pieber, 1872

Pigs 49-52

Type genus: Selenocephalus Germar, 1833.

= Ianeirini Linnavuori, 1978 syn. nov.

= Adamini Linnavuori & Al-Ne’amy, 1983 syn. nov.

= Dwightlini Linnavuori & Al-Ne’amy, 1983 syn. nov.

Diagnosis

Selenocephalini are medium sized to large, robust, often brownish, greenish, or yellow leafhoppers,

sometimes with dark irroration. They can be identified by the anterior margin of head usually with

2 or more carinae or striae, broad face, profemur row AM often with 1 or more extra setae basad of

AMI, profemur row AV setae often relatively long, connective stem usually short and stout, and their

distribution in the Afrotropical, Palearctic, and Oriental regions.

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Description

Head. Head wider than or narrower than pronotum. Discal portion of crown usually glabrous with radial

or longitudinal striae, or rarely shagreen. Anterior margin of head usually acute or subfoliaceous, with

2 or more distinct carinae or transverse striae, rarely without carinae or striae. Face often quite broad.

Frontoclypeus not tumid; texture shagreen. Clypellus widening apically; apex following or slightly

surpassing normal curve of gena; often with more or less distinct median keel. Lorum subequal to or

wider than clypellus near base. Antennal bases near middle or posteroventral (lower) corners of eyes

or rarely near anterodorsal (upper) corners (Dwightlina). Antennae short, less than 1.5 x width of head.

Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture, often relatively distant

from laterofrontal suture. Antennal ledge somewhat developed, carinate or weakly carinate. Ocelli

present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins not

raised; without reflexed costal veins; A1-A2 crossvein usually present; apical venation not reticulate or

rarely highly reticulate.

Legs. Profemur usually with AMI and with one or more additional proximal setae; intercalary row

with one row of five or more fine setae; row AV with short stout setae or often with relatively longer

setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1. Metatarsomere I not

expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae often absent or reduced, or sometimes

well differentiated into several rows. Subgenital plates free from each other; articulated with valve;

with macrosetae scattered, with several irregular rows near lateral margin, or without macrosetae. Style

broadly bilobed basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective

absent or present, connected or articulated to connective or near base of aedeagus. Aedeagus with single

shaft and gonopore. Connective anterior arms somewhat divergent, Y- or U-shaped; stem often short,

stout; articulated with or rarely fused ( Abimwa ) to aedeagus, or sometimes dissociated from aedeagus

(Adamina).

Female genitalia. Pygofer with numerous macrosetae. Ovipositor usually not protruding far beyond

pygofer apex. First valvula convex; dorsal sculpturing pattern usually strigate, sometimes concatenate

or reticulate; sculpturing reaching dorsal margin; without distinctly delimited ventroapical sculpturing.

Second valvula broad, tapering apically or abruptly broadened medially or subapically; often with dorsal

median tooth; teeth on apical 1/3 or more or restricted to apical 1/4 or less; teeth small, regularly or

irregularly shaped.

Geography and ecology

Distribution: Palearctic, Afrotropical and Oriental regions. Selenocephalini are usually collected on

herbaceous or woody dicots and are often collected at lights. They are particularly diverse in Afrotropical

forest ecosystems. Some Selenocephalini are ant-attended.

Remarks

Selenocephalini contains 21 genera and 196 species. It is interpreted here to contain all of the tribes

recognized in Selenocephalinae sensu Linnavuori & Al-Ne’amy (1983a) with the exception ofDrabescini

(=Paraboloponini, Bhatiini). We retain as subtribes most of the groups previously recognized as tribes of

Selenocephalinae. Representatives of each of the subtribes were included in the phylogenetic analyses

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

(Dwightla is included for the first time here) and the tribe was usually resolved as monophyletic, although

with low branch support. The Afrotropical genus Dagama was also sometimes resolved within or as

sister to this clade, and this group was resolved near Bonaspeiini. Dagama does not possess carinae on

the anterior margin of the head and is not included in the tribe at this time. Further research should aim to

determine the relationships between this and other Afrotropical athysanine genera and Selenocephalini,

Bonaspeiini, Hypacostemmini, Phlepsiini, Scaphoideini, and Drabescini.

Selected references

Linnavuori (1978a), Linnavuori & Al-Ne’amy (1983), Zhang & Webb (1996), Dmitriev (2003),

McKamey (2003b).

Included subtribes:

Adamina Linnavuori & Al-Ne’amy, 1983 status nov.

Fig. 49

Type genus: Adama Dlabola, 1980.

Diagnosis

Adamina are medium sized to large yellow, green, or black leafhoppers, sometimes with longitudinal

reddish stripes (subgenus Adama). They can be identified by the anterior margin of the head with 3

sharp carinae or with numerous irregular carinae, the aedeagus united to the connective only by a thin

membrane, with long paraphyses (or "pseudostyles") usually articulated with or fused to the connective,

and first valvulae widening apically, with irregular margins near the apex, and with reticulate sculpturing.

Description

Head. Head subequal to, wider than, or narrower than pronotum. Discal portion of crown glabrous with

radial or longitudinal striae or shagreen. Anterior margin of head with 3 parallel carinae or with numerous

irregular carinae. Frontoclypeus not tumid; texture shagreen. Clypellus widening apically. Clypellus

apex following or slightly surpassing normal curve of gena. Lorum subequal to or wider than clypellus

near base. Antennal bases near middle or posteroventral (lower) corners of eyes. Antennae short, less

than 1.5 x width of head. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture.

Antennal ledge weakly developed (carinate or wealdy carinate). Ocelli present; close to eyes; on anterior

margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins not

raised; without reflexed costal veins; A1-A2 crossvein present; apical venation highly reticulate or not

reticulate.

Legs. Pro femur with AMI and with one or more additional proximal setae; intercalary row with one

row of five or more fine setae; row AV with relatively long macrosetae. Protibia dorsal surface rounded,

convex. Metafemur apex macrosetae 2+2+1. Metatarsomere I not expanded apically; plantar setae

simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point.

Pygoferbasolateral membranous cleft present; macrosetae well differentiated into several rows. Subgenital

plates free from each other; articulated with valve; without macrosetae. Style broadly bilobed basally,

median anterior lobe pronounced. Basal processes of the aedeagus/connective present, connected or

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articulated to connective or near base of aedeagus. Aedeagus with single shaft and gonopore; dissociated

from and lying far above connective, attached to connective only by thin membrane. Connective anterior

arms somewhat divergent, U-shaped.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor protruding or not protruding far

beyond pygofer apex. First valvula convex; margins irregularly undulate or notched subapically; dorsal

sculpturing pattern strigate, concatenate, reticulate, or with rectangular shaped cells; sculpturing reaching

Fig. 49. Selenocephalini (Adamina). A-B, D-H. Adama ( Paracostemma ) amalthusia Linnavuori &

Al-Ne’amy, 1983. C. A. ( Paracostemma ) sp. A-H. Standard views (see Material & Methods). G.

Connective and style shown with connective processes or “pseudostyles”.

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dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula abruptly broadened

medially or subapically; without dorsal median tooth; teeth restricted to apical 1/4 or less; teeth small,

regularly or irregularly shaped.

Geography and ecology

Distribution: Afrotropical. Adamina are frequently collected at lights and likely feed in the forest canopy.

Remarks

Adamina contains the type genus, 4 subgenera, and 49 valid species. All members are found in the

Afrotropical region.

Included genera

Adama Dlabola, 1980

Dwightlina McKamey, 2003 status nov.

Fig. 50

Type genus: Dwightla McKamey, 2003.

Diagnosis

Dwightlina are large, robust, ochraceous, reddish-brown, or brown leafhoppers. They can be identified

by the anterior margin of head with 3 parallel carinae, face very broad, antennal pits near upper corners of

eyes, forewing and face with slight to heavy covering of fine setae, profemur with 2 or more macrosetae

basad of AMI seta, and male pygofer without macrosetae.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae. Anterior margin of head with 3 parallel carinae. Face very broad; with slight to

heavy covering of fine hairs. Frontoclypeus not tumid; texture shagreen. Clypellus widening apically;

apex following or slightly surpassing normal curve of gena. Forum subequal to or wider than clypellus

near base. Antennal bases near upper or anterodorsal corners of eyes. Antennae short, less than 1.5 x

width of head. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture. Antennal

ledge weakly developed (carinate or weakly carinate). Ocelli present; close to eyes; on anterior margin

of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewings macropterous; with slight to heavy covering of fine setae; tapering apically; apex thin

or membranous, folded over onto opposite wing; appendix large, extending around wing apex; with 3

anteapical cells; veins not raised; without reflexed costal veins; A1-A2 crossvein present.

Fegs. Fegs often with accessory setae. Profemur with AMI and with 2 or more additional proximal

setae; intercalary row with one row of five or more fine setae, sometimes with extra scattered setae; row

AV with very long macrosetae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae

2+2+1. Metatibia dorsal surface usually with extra stout setae, especially proximally. Metatarsomere I

not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point.

Pygofer basolateral membranous cleft present; without macrosetae. Subgenital plates free from each

other; articulated with valve; macrosetae irregularly arranged near lateral margin. Style broadly bilobed

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Fig. 50. Selenocephalini (Dwightlina). A-B, D-F, H-K. Dwightla delongi sp.nov. C. D. acutipennis

Linnavuori & Al-Ne’amy, 1983. A-F, H-I. Standard views (see Material & Methods). J. Ventral view

of connective. K. Ventral view of style.

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basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent. Aedeagus

with single shaft and gonopore. Connective anterior arms somewhat divergent, Y- or U-shaped; stem

short; articulated with aedeagus. Segment X large, sclerotized.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor protruding or not protruding far

beyond pygofer apex. First valvula convex; dorsal sculpturing pattern strigate; sculpturing reaching

dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula slender throughout;

without dorsal median tooth; dorsal teeth on apical 1/3 or more or restricted to apical 1/4 or less; teeth

large, regularly shaped.

Geography and ecology

Distribution: Afrotropical. Dwightlina are inhabitants of forests and have been collected at lights.

Remarks

Dwightlina contains the type genus and 5 species from the Afrotropical region. McKamey (2003b)

provided the replacement name Dwightla for the preoccupied Dwightia Linnavuori & Al-Ne’amy, 1983,

thus necessitating a change in the family group name. Phylogenetic analyses of new molecular data

provided here confirm that Dwightla is closely related to Selenocephalini, a result also found based on

morphology alone (Zahniser & Dietrich 2008).

Included genera

Dwightla McKamey, 2003

Ianeirina Linnavuori, 1978 status nov.

Fig. 51

Type genus: Ianeira Linnavuori, 1969.

Diagnosis

Ianeirina are medium sized to large, ivory to brown leafhoppers. They can be identified by their somewhat

long and slender habitus, anterior margin of the head usually with numerous carinae (shagreen and

rounded to face in Ianeira), ocelli on the face below the anterior margin of the head, male pygofer

without macrosetae, subgenital plates fused basally and to the valve (not fused in Bardera), and segment

X well-sclerotized, large to very large, sometimes occupying most of the dorsal side of the pygofer.

Description

Head. Head subequal to, wider than, or distinctly narrower than pronotum. Discal portion of crown

glabrous with radial or longitudinal striae. Anterior margin of head shagreen, with numerous transverse

striations, or with numerous carinae. Frontoclypeus narrow; not tumid; texture shagreen. Clypellus

widening apically, apex following or slightly surpassing normal curve of gena. Lorum subequal to or

wider than clypellus near base. Antennal bases near middle or posteroventral (lower) corners of eyes.

Antennae short, less than 1.5 x width of head. Gena obtusely incised laterally; with fine erect seta beside

laterofrontal suture. Antennal ledges weakly developed (carinate or weakly carinate). Ocelli present;

close to or distant from eyes; on face just below anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Fore wing macropterous; appendix restricted to anal margin; with 3 anteapical cells; veins not

raised; with or without reflexed costal veins; A1-A2 crossvein present.

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Fig. 51. Selenocephalini (Ianeirina). Abimwa knighti sp. nov. A-D, F, H-I. Standard views (see Material

& Methods). D. Pygofer shown with subgenital plates and valve. J. Ventral view of style. K. Ventral

view of connective and aedeagus fused to connective. L. Dorsal view of $ segments X, XI, and anus.

M. $ sternite VII.

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Legs. Pro femur with AMI seta only or with AMI and with one or more additional proximal setae;

intercalary row with one row of five or more fine setae; row AV with short, stout setae. Protibia dorsal

surface rounded, convex. Metafemur apex macrosetae 2+2+1. Metatarsomere I not expanded apically;

plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point.

Pygofer basolateral membranous cleft present, macrosetae absent or reduced (< two rows). Subgenital

plates free from each other and articulated with valve ( Bardera ) or fused to each other basally (apices of

plates usually separated) and to valve ( Abimwa , Ianeira ); without macrosetae or macrosetae scattered,

irregularly arranged. Style broadly bilobed basally, median anterior lobe pronounced; sometimes reduced

in size ( Abimwa , Ianeira). Basal processes of the aedeagus/connective absent or reduced or present,

connected or articulated to connective or near base of aedeagus. Aedeagus with single shaft and gonopore.

Connective anterior arms somewhat divergent, Y- or U-shaped; articulated with or fused to aedeagus.

Segment X well-sclerotized; large to very large, sometimes occupying majority of dorsal side of pygofer.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula convex; dorsal sculpturing pattern strigate; sculpturing reaching dorsal margin,

without distinctly delimited ventroapical sculpturing. Second valvula broad, gradually tapered; with

dorsal median tooth; teeth on apical 1/3 or more; teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Afrotropical.

Remarks

Ianeirina contains 4 genera and 13 valid species. Males of Kanziko Linnavuori & Al-Ne’amy, 1983

are not known. There is variability in the condition of the anterior margin of the head (rounded to face

and shagreen in Ianeira , and with numerous irregular carinae in the other genera) and the condition

of the male genital capsule (plates and valve articulated in Bardera and fused in Ianeira and Abimwa

Linnavuori, 1978) among the genera. The monophyly of the tribe needs to be tested in a more thorough

phylogenetic analysis. One representative of Ianeirini, Abimwa , was included in phylogenetic analyses

and was found to be closely related to other Selenocephalini and in some analyses to Dagama Distant,

1910 (Athysanini).

Included genera:

Abimwa Linnavuori, 1978

Bardera Linnavuori & Al-Ne’amy, 1983

Ianeira Linnavuori, 1969

Kanziko Linnavuori & Al-Ne’amy, 1983

Selenocephalina Fieber, 1872

Fig. 52

Type genus: Selenocephalus Germar, 1833.

Diagnosis

Selenocephalina are medium sized to large, robust often brownish, yellowish, ochraceous, grey, brown,

or reddish leafhoppers, sometimes with dark irroration. They can be identified by the carinae on the

anterior margin of the head, ocelli on the anterior margin of the head and close to the eyes, broad face,

male pygofer usually with few or no macrosetae, segment X broad and strongly sclerotized dorsally, and

lacking the features defining other subtribes of Selenocephalini.

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Description

Head. Head subequal to or wider than pronotum. Discal portion of crown glabrous with radial or

longitudinal striae. Anterior margin of head with 2 or 3 parallel carinae or with numerous irregular carinae

or striations. Face very broad. Frontoclypeus not tumid; texture shagreen. Clypellus widening apically;

apex following or slightly surpassing normal curve of gena; often with more or less distinct median

keel. Lorum subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral

(lower) corners of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely incised laterally,

with fine erect seta beside laterofrontal suture. Antennal ledges weakly developed (carinate or weakly

carinate). Ocelli present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin or large, extending around wing

apex; with 3 anteapical cells; veins not raised; without reflexed costal veins; A1-A2 crossvein present;

apical venation not highly reticulate.

Legs. Pro femur with AMI seta only or with AMI and with one or more additional proximal setae;

intercalary row with one row of five or more fine setae; row AV with short, stout setae or with relatively

long macrosetae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1.

Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae usually absent or reduced (< two

rows). Subgenital plates free from each other; articulated with valve; without macrosetae or macrosetae

scattered, irregularly arranged, uniseriate laterally, with two lateral rows of macrosetae, or with irregularly

arranged macrosetae near lateral margin. Style broadly bilobed basally, median anterior lobe pronounced.

Basal processes of the aedeagus/connective absent or reduced or present, connected or articulated to

connective or near base of aedeagus. Aedeagus with single shaft and gonopore. Connective anterior

arms somewhat divergent, Y- or U-shaped; articulated with aedeagus. Segment X broad; sclerotized

dorsally at least at apex; sometimes with processes.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor protruding or not protruding far

beyond pygofer apex. First valvula convex; dorsal sculpturing pattern strigate; sculpturing reaching

dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula abruptly broadened

medially or subapically or broad, gradually tapered; often with or sometimes without dorsal median

tooth; teeth on apical 1/3 or more; teeth large, regularly shaped or teeth small, regularly or irregularly

shaped.

Geography and ecology

Distribution: Palearctic, Afrotropical and Oriental. Selenocephalina are most diverse in the Afrotropical

forested ecosystems and are often collected at lights.

Remarks

Selenocephalina contains 15 genera and 129 species. Selenocephalus was included in the phylogenetic

analyses and found to be closely related to other Selenocephalini and sister to Citorus (included in

phylogenetic analyses here for the first time), which is transferred from Penthimiini and included in

Selenocephalina here. Citorus has a squat, robust body form, large forewing with the appendix extending

around the wing apex, and protibiae flattened dorsally and carinate, which account for its previous

placement in Penthimiini, but it shares other characters (ocelli on anterior margin of head and close to

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Fig. 52. Selenocephalini (Selenocephalina). A-B, D-K. Ganniaviraktamathi sp. nov. C. Selenocephalus

planus (Turton, 1802). A-I. Standard views (see Material & Methods). G. Connective and style shown

with aedeagus and processes of aedeagus. H-I. Aedeagus shown with processes of aedeagus. J. Broad

view of style. K. $ sternite VII.

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eyes; antennal ledges not very strong) with Selenocephalina. Very strong branch support was recovered

for a clade including Dwight la, Citorus and Selenocephalus.

Included genera

Angolaia Linnavuori & Al-Ne’amy, 1983

Bumka McKamey, 2003

Citorus Stal, 1866 placement nov. (transferred from Penthimiini)

Distantia Signoret, 1879

Foso Linnavuori & Al-Ne’amy, 1983

Gannia Theron, 1979 placement nov. (previously unplaced in Deltocephalinae)

Hecaloidella Osborn, 1934

Ifeia Linnavuori & Al-Ne’amy, 1983

Loka Linnavuori & Al-Ne’amy, 1983

Maichewia Linnavuori & Al-Ne’amy, 1983

Malendea Linnavuori & Al-Ne’amy, 1983

Moluccazhangia McKamey, 2003

Omanellinus Zhang, 1999

Sapoba Linnavuori & Al-Ne’amy, 1983

Selenocephalus Gq rmar, 1833

Stegelytrini Baker, 1915

Fig. 53

Type genus: Stegelytra Mulsant & Rey, 1855.

Diagnosis

Stegelytrini are medium sized to large leafhoppers, sometimes very robust, setose, and flylike

in appearance. They can be identified by the head usually much narrower than the pronotum, eyes

encroaching onto the pronotum and lying above lateral carinae of pronotum, antennae usually very long,

clypellus often with pair of stout subapical setae, legs with supernumerary setae, profemur row AM with

AMI plus two or more additional macrosetae, metatibia dorsal surface with extra setae between rows

AD and PD near base, valve often with long length of articulation with pygofer, connective with anterior

arms widely divergent, and style linear.

Description

Head. Head usually distinctly narrower than pronotum, rarely wider than pronotum; eyes lying above

lateral carinae of pronotum. Discal portion of crown glabrous with radial or longitudinal striae. Anterior

margin of head shagreen or with numerous transverse striations or carinae. Frontoclypeus not tumid,

texture shagreen. Clypellus widening apically; apex following or slightly surpassing normal curve of

gena; usually with pair of subapical stout setae. Lorum subequal to or wider than clypellus near base.

Antennal bases near middle or posteroventral (lower) corners of eyes. Antennae usually very long, 1.5 x

width of head or longer, rarely short. Gena not incised or obtusely incised laterally; with fine erect seta

beside laterofrontal suture. Antennal ledges weakly developed (carinate or wealdy carinate) or strongly

developed (with a definite ledge). Ocelli present; close to eyes; on anterior margin of head.

Thorax. Thorax often very large or inflated. Pronotum lateral margin carinate or not carinate; lateral

margin often longer than basal width of eye, rarely shorter. Mesonotum often elongate and protuberant

posteriorly.

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Wings. Forewing macropterous or submacropterous; apex sometimes truncate; appendix absent or

reduced, or present and restricted to anal margin or large and extending around wing apex; with 3

anteapical cells; veins not raised; without reflexed costal veins; A1-A2 crossvein absent or present;

apical venation highly reticulate or not; commissural margin usually thickened and elevated.

Legs. Legs often with supernumerary t hin and thick setae. Pro femur with AMI and with two or more

additional proximal setae; intercalary row with one row of five or more fine setae, or setae scattered, not

in one row; row AV with relatively long macrosetae. Protibia dorsal surface rounded, convex. Metafemur

apex macrosetae 2+2+1 or 2+2+1 and with additional proximal macrosetae and/or with extra macrosetae

clustered near apex. Metatibia dorsal surface with extra setae between rows AD and PD, especially near

base; dorsal side often distinctly wider at base than at apex. Metatarsomere I not expanded apically;

plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer or rarely fused to pygofer; lateral margin often relatively

long and with long length of articulation with pygofer, or sometimes lateral margin short, articulating

with pygofer at a point. Pygofer basolateral membranous cleft often absent, rarely present; macrosetae

absent or reduced (< two rows) or weft differentiated into several rows. Subgenital plates free from each

other; articulated with valve; without macrosetae or with macrosetae scattered, irregularly arranged, or

uniseriate, distant from lateral margin. Style linear, median anterior lobe not pronounced. Basal processes

of the aedeagus/connective absent or reduced or present, connected or articulated to connective or near

base of aedeagus. Aedeagus with single shaft and gonopore. Connective anterior arms widely divergent,

T-shaped or plate-like; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding or protruding far

beyond pygofer apex. First valvula convex; dorsal sculpturing pattern strigate or concatenate; sculpturing

reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula abruptly

broadened medially or subapically, or broad, gradually tapered; without dorsal median tooth; teeth on

apical 1/3 or more; teeth large, regularly shaped or teeth small, regularly or irregularly shaped.

Geography and ecology

Distribution: Palearctic and Oriental regions. Stegelytrini have been collected on trees. Recorded host

families include Fagaceae and Lauraceae. Aggregations of males have been observed on wet soil.

Remarks

Stegelytrini contains 30 genera and 83 species. It is the earliest diverging lineage of Deltocephalinae,

and its relictual status is reflected in the retention of primitive structures of the male genitalia such as

the narrow style base and median anterior lobe of the connective. Many members of the tribe also have

a suite of derived characters that are outwardly modified and produce a strange flylike or otherwise

“hairy” appearance. The functional significance of these modified characters is not known, although

some may be Batesian mimics of flies or bees. The tribe, which has been accorded separate subfamily

status by some authors, was recently revised in a series of papers (see Selected references). We prefer to

treat the group as a tribe of Deltocephalinae to accentuate the strong phylogenetic support for its close

relationship to other deltocephalines.

Selected references

Webb (1999), Wei et al. (2006, 2007a, b, 2008a, b, 2010).

Included genera

Aculescutellaris Zhang, Wei & Webb, 2007

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European Journal of Taxonomy 45: 1-211 (2013)

S'Jh'm. jF

fe -i

v r ¥ I

'r \I

Fig. 53. Stegelytrini. Stegelytra putoni Mulsant & Rey, 1875. A-I. Standard views (see Material &

Methods).

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Cyrta Melichar, 1902

Daochia Wei, Zhang & Webb, 2006

Doda Distant, 1908

Honguchia Wei & Webb, 2010

Kasinella Evans, 1971

Kunasia Distant, 1908

Louangana Wei & Webb, 2010

Minucella Wei, Zhang & Webb, 2008

Neophansia Wei & Webb, 2010

P achy me to pi ns M ats um u r a, 1914

Paracyrta Wei, Webb & Zhang, 2008

Paradoxivena Wei, Zhang &Webb, 2006

ParaplacideUus Zhang, Wei & Shen, 2002

Paratoba Wei, Webb & Zhang, 2007

Pataniolidia Wei, Webb & Zhang, 2007

Placidellus Evans, 1971

Platyvalvata Zhang, Wei & Webb, 2006

Pseudododa Zhang, Wei & Webb, 2007

Quiontugia Wei & Zhang, 2010

Shangonia Wei & Webb, 2010

Stegelytra Mulsant & Rey, 1855

Stenolora Zhang, Wei & Webb, 2006

Sychentia Wei & Webb, 2010

Temburocera Webb, 1999

Toba Schmidt, 1911

Trunchinus Zhang, Webb & Wei, 2007

Wadkiifia Linnavuori, 1965

Wyuchiva Zhang, Wei & Webb, 2006

Yaontogonia Wei & Webb, 2010

Stenometopiini Baker, 1923

Fig. 54

Type genus: Stenometopius Matsumura, 1914 (=Hodoedocus Jacobi, 1910).

= Stirellini Emeljanov, 1966.

Diagnosis

Stenometopiini are small to medium sized, often brightly colored or iridescent leafhoppers. They can

be identified by the narrow crown, shagreen texture of the crown, clypellus parallel-sided or tapering

apically, forewings often submacropterous to brachypterous, male pygofer sloping caudoventrally and

with few macrosetae and often with a distinct lateral tooth, female ovipositor protruding far beyond the

pygofer apex, first valvula dorsal sculpturing granulose to maculose and submarginal, first valvula with

distinctly delimited ventroapical sculpturing, and second valvula without dorsal teeth.

Description

Head. Head subequal to or wider than pronotum; sometimes strongly produced or elongate. Crown

narrow; discal portion of crown shagreen. Anterior margin of head shagreen, rarely sharply angulate

or foliaceous if head is elongate. Frontoclypeus not tumid, texture shagreen. Clypellus parallel-sided

or tapering apically, apex slightly surpassing normal curve of gena. Lorum distinctly narrower than

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clypellus near base. Antennal bases near middle or posteroventral (lower) corners of eyes. Mesial margin

of eyes notched in frontal view. Antennae short, less than 1.5 x width of head. Gena obtusely incised

laterally; with fine erect seta beside laterofrontal suture. Antennal ledges absent. Ocelli present; close to

eyes; on anterior margin of head.

Thorax. Pronotum lateral margin not carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous to brachypterous; appendix reduced, absent, or restricted to anal margin;

with 3 anteapical cells; veins not raised; usually without reflexed costal veins; A1-A2 crossvein absent;

apical venation not highly reticulate.

Legs. Profemur with AMI seta only; intercalary row with one row of five or more fine setae; row AV

with short, stout setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+0, 2+1,

2+2, or 2+2+1. Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point.

Pygofer sloping caudoventrally; basolateral membranous cleft absent, not membranous; macrosetae

absent or reduced (< two rows); often with distinct short tooth laterally. Subgenital plates often short,

rounded; free from each other; articulated with valve; without macrosetae or macrosetae uniseriate

laterally or uniseriate distant from lateral margin. Style broadly bilobed basally, median anterior lobe

pronounced and usually extending anterad of lateral lobe. Basal processes of the aedeagus/connective

absent or reduced. Aedeagus with single shaft and gonopore; often whip-like, long and narrow.

Connective anterior arms somewhat divergent, Y-, U- or narrowly V-shaped; articulated with aedeagus.

Female genitalia. Pygofer with macrosetae reduced or absent. Ovipositor protruding far beyond pygofer

apex. First valvula not strongly convex; dorsal sculpturing pattern granulose or maculose; sculpturing

submarginal; with ventroapical sculpturing distinctly delimited, long and triangular. Second valvula

gradually broadened medially or subapically; without dorsal median tooth; teeth absent.

Geography and ecology

Distribution: cosmopolitan. Stenometopiini feed on grasses or sedges and often abundant in grassland

ecosystems.

Remarks

Stenometopiini contains 8 genera and 96 species. Based on some morphological characters (Zahniser

2008a) and on phylogenetic analyses, the tribe appears to be most closely related to Chiasmini or to

some other grass-specialist groups including Eupelicini, Drakensbergenini, and Evinus Dlabola, 1977.

Most of the species are included in two genera: Doratulina , which contains only Old World species, and

Stirellus, which contains both Old World and New World species. Historically, taxonomists have been

inconsistent or unclear in their reasoning for their placement of species in these genera. The distinction

between the two genera is therefore unclear (see Zahniser & Nielson 2012 for further discussion), but

both are retained pending further study.

Proekes Theron, 1975 and Proekoides Stiller, 1986 were previously listed in the tribe by Oman et al.

(1990) but share no similarities to the tribe and are here transferred to Bonaspeiini.

Selected references

Vilbaste (1965), Emeljanov (1966, 1968), Linnavuori (1979b), Dmitriev (2004b), Zahniser (2008a),

Zahniser & Nielson (2012).

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 54. Stenometopiini. A-B, D-J. Doratulina dmitrievi sp. nov. C. Stirellus bicolor (Van Duzee,

1892), A1 c?,A2 A-I. Standard views (see Material & Methods). J. $ sternite VII.

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European Journal of Taxonomy 45: 1-211 (2013)

Included genera

Anaconura Emeljanov, 1999

Anemolua Kirkaldy, 1906

Doratulina Melichar, 1903

Giffardia Kirkaldy, 1906

Hodoedocus Jacobi, 1910

Kinonia Ball, 1933

Paivanana Distant, 1918

Stirellus Osborn & Ball, 1902

Tetartostylini Wagner, 1951

Fig. 55

Type genus: Tetartostylus Wagner, 1951.

Diagnosis

Tetartostylini are medium sized, somewhat elongate, straw colored to greenish leafhoppers. They can

be identified by the tapering or parallel-sided clypellus, subgenital plates fused to valve and with apices

modified into sharp spines, connective with anterior arms closely appressed anteriorly, connective

articulated with a pair of long processes, connective articulated with aedeagus, female sternite VII long,

ovate or subovate.

Description

Head. Head subequal to or wider than pronotum. Discal portion of crown shagreen. Anterior margin of

head shagreen. Frontoclypeus not tumid, texture shagreen. Clypellus parallel-sided or tapering apically;

apex following or slightly surpassing normal curve of gena. Lorum distinctly narrower than clypellus

near base. Antennal bases near middle or posteroventral (lower) comers of eyes. Antennae short, less

than 1.5 x width of head. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture.

Antennal ledges absent. Ocelli present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin not carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; appendix restricted to anal margin; with 2 or 3 anteapical cells; veins

not raised; without reflexed costal veins; A1-A2 crossvein absent; apical venation not highly reticulate.

Legs. Pro femur with AMI seta only; intercalary row reduced or absent; row AV with thin, hair-like

setae or without setae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1.

Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a

point. Pygofer basolateral membranous cleft present; macrosetae well differentiated into several rows.

Subgenital plates free from each other, fused to valve; produced apically into modified spines; macrosetae

uniseriate laterally. Style broadly bilobed basally; median anterior lobe pronounced. Basal processes of

the aedeagus/connective present, articulated with connective. Aedeagus with single shaft and gonopore.

Connective anterior arms closely appressed anteriorly, linear or loop-shaped; articulated with aedeagus.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor not protruding far beyond pygofer

apex. First valvula not strongly convex; dorsal sculpturing pattern imbricate (with overlapping scales);

sculpturing submarginal; without distinctly delimited ventroapical sculpturing. Second valvula broad.

172

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Fig. 55. Tetartostylini. A-B, D-J. Tetartostylus acutus Heller & Linnavuori, 1968. C. Tetartostylus

sp. A-I. Standard views (see Material & Methods). G. Connective and style shown with processes of

connective or “pseudostyles”. J. $ sternite VII.

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European Journal of Taxonomy 45: 1-211 (2013)

gradually tapered; without dorsal median tooth; teeth on apical 1/3 or more; teeth small, regularly or

irregularly shaped.

Geography and ecology

Distribution: Palearctic and Afrotropical regions. Tetartostylini feed on grasses.

Remarks

Tetartostylini contains the type genus and 11 species. It is resolved as sister to the Paralimnini +

Deltocephalini clade in all phylogenetic analyses conducted thus far, with moderate to strong branch

support. With those two tribes, it shares a tapered clypellus and a ‘linear’ connective. Further taxon

sampling in Paral imnin i could indicate that it belongs in that tribe. For now, the continued recognition

of the tribe is supported by phylogenetic analyses. Dmitriev (2003) included Hiltus Theron, 1974 in the

tribe based on nymphal characters, but because this genus does not have the subgenital plates fused to

the valve and pseudostyli articulated with connective, we follow Theron (1974) in placing that genus in

Paral imnin i

Selected references

Wagner (1951), Theron (1973), Dmitriev (2003).

Notes on illustration (Fig. 55)

The species figured in Fig. 55A-B, D-J is perhaps referable to T. acutus Heller & Linnavuori, 1968

described from Ethiopia. A series of four males and three females (INHS) were collected from:

ZAMBIA: Northwestern Prov., ~28.5 km N of Mwinilunga, 11°30’16.1” S - 24°22’44.8” E, Chitunta

plain grassland, 1384 m, 5-7 Nov. 2007, ZA-03, sweep, JN Zahniser. This series shares with T. acutus

a similar shape of the female stemite VII and similar coloration of the face. However, T. acutus was

described based only on a female specimen. The females examined here are somewhat shorter (4.8-

4.9 mm ) than that described by Heller & Linnavuori (5.5 mm). For now, the Zambian specimens are

tentatively identified as T. acutus , but confirmation by comparison to the type is needed. The specimens

were collected in a native Zambian grassland from which numerous other undescribed species of

Deltocephalinae have been identified (e.g., Doratulina dmitrievi sp. nov., below). Further exploration

of the central and east African grasslands will likely reveal a rich, nearly completely undescribed fauna

of Deltocephalinae.

Included genera

Tetartostylus Wagner, 1951

Vartini tribe nov.

um:lsid:zoobank.org:act:ElF01DC6-889F-48C3-B0D7-28C87B614FlD

Fig. 56

Type genus: Varta Distant, 1908.

Diagnosis

Vartini are medium sized to large, somewhat elongate, usually greenish or bluish leafhoppers, usually

with red or orange longitudinal stripes. They can be identified by the produced and pointed head,

gena visible behind the eye in dorsal view, elongate frontoclypeus, lorum distant from genal margin,

profemur intercalary row setae thick and extending to or beyond middle of profemur, forewings truncate

apically, apodemes of male stemite II long, subrectangular, flared apically, and pointed posterolaterally,

174

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

connective with anterior arms appressed, and male segment X tube-like and protruding from pygofer

and often well-sclerotized.

Description

Head. Head narrower than pronotum; acutely or bluntly produced. Discal portion of crown glabrous

with radial or longitudinal striae. Anterior margin of head foliaceous or acutely angled with several

transverse striae or carinae. Frontoclypeus very long, narrow; texture shagreen; sometimes with median

longitudinal carina anterodorsally. Clypellus parallel-sided or widening apically; apex following or

slightly surpassing normal curve of gena. Lorum distinctly narrower than clypellus near base; very well-

separated from genal margin. Antennal bases near middle, posteroventral (lower) corners, or anterodorsal

(upper) corners of eyes. Antennae long, 1.5 x width of head or more. Gena not incised, slightly incised,

or sinuate laterally; visible behind eye in dorsal view; with fine erect seta beside laterofrontal suture.

Antennal ledges absent. Ocelli present; close to eyes; on anterior margin of head.

Thorax. Pronotum lateral margin carinate; lateral margin shorter than basal width of eye.

Wings. Forewing macropterous; truncate apically; appendix absent, reduced, or present and restricted to

anal margin; with 3 anteapical cells; veins sometimes obscure; veins not raised; without reflexed costal

veins; A1-A2 crossvein absent; apical venation not highly reticulate.

Legs. Setal bases often darkly colored. Profemur with AMI seta only; intercalary row setae relatively

large and thick, extending to or past middle of profemur; row AV with short, stout setae. Protibia dorsal

surface rounded, convex; row AD with 1 apical macrosetae; row PD with 5 to 9 macrosetae. Metafemur

apex macrosetae 2+2+1. Metatarsomere I not expanded apically; plantar setae simple, tapered.

Male genitalia. Apodemes of sternite II long, subrectangular, flared apically, pointed posterolaterally.

Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point. Pygofer

basolateral membranous cleft present or absent; macrosetae absent or reduced or present, well

differentiated into several rows. Subgenital plates free from each other or fused or partially fused to

each other or to valve; often with many scattered macrosetae, or with one to several rows of macrosetae

near lateral margin. Style broadly bilobed basally; median anterior lobe pronounced. Basal processes of

the aedeagus/connective absent or present, if present then fused to connective or aedeagus. Aedeagus

with single shaft and gonopore. Connective anterior arms closely appressed anteriorly, or rarely slightly

separated; stem absent or present; articulated with or fused to aedeagus. Segment X tube-like; protruding

from pygofer; sometimes with ventral processes.

Female genitalia. Pygofer with numerous macrosetae. Ovipositor protruding or not protruding far

beyond pygofer apex. First valvula convex; dorsal sculpturing pattern strigate to reticulate; sculpturing

reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula broad,

gradually tapered or slightly expanded subapically; without dorsal median tooth; teeth on apical 1/4 to

1/3 or more; teeth small to medium sized, regularly or irregularly shaped.

Geography and ecology

Distribution: Afrotropical, Palearctic, and Oriental regions. Linnavuori (1978b) reported Stymphalus

Stal, 1866 “on sedges and grasses in moist locations”.

Remarks

Vartini includes 7 genera and 23 species. The tribe is newly described here to include seven genera

previously placed in Scaphytopiini. The only diagnostic characters that Vartini share with Scaphytopius

are the produced head and very broad gena visible behind the eyes in dorsal view. These two characters

175

European Journal of Taxonomy 45: 1-211 (2013)

Fig. 56. Vartini. A-C. Vartalapa robusta Viraktamath, 2004. D-K. Varta rubrofasciata Distant, 1908.

A-I. Standard views (see Material & Methods). J. Ventral view of base of $ abdomen showing long

apodemes of sternite II (sternite I obscure). K. Ventral view of $ pygofer.

176

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

tend to co-occur in many deltocephalines that are not related to one another. None of the genera previously

included in Scaphytopiini that were sampled in phylogenetic analyses here ( Scaphytopius , Nesothamnus,

Japananus , Stymphalus ) were found to be closely related to one another. Thus, the aforementioned

characters appear to be convergent and not appropriate for sole use in defining tribes of Deltocephalinae.

The Vartini are separated from Scaphytopiini here and, based on the number and uniqueness of the

characters defining the group, appear to form a monophyletic group. Three characters supporting the

group are newly reported here: profemur intercalary row setae relatively thick and extending to or

basad of midlength of femur; apodemes of male sternite II long, subrectangular, flared apically, and

pointed posterolaterally; male segment X tube-like and protruding from pygofer, often well-sclerotized.

Viraktamath (2004) revised the group and several new species have been described since.

Stymphalus was resolved near some Mukariini (. Mukaria and Scaphotettix ) and sister to an undescribed

genus from Thailand which does not share the diagnostic characters of the tribe listed above. More data

are needed to resolve its relationship to other tribes of Deltocephalinae.

Selected references

Linnavuori (1978b), Viraktamath & Anantha Murthy (1999), Viraktamath (2004).

Included genera:

Curvimonus Viraktamath & Anantha Murthy, 1999 placement nov. (transferred from Scaphytopiini)

Shivania Viraktamath, 2004 placement nov. (transferred from Scaphytopiini)

Stymphalus Stal, 1866 placement nov. (transferred from Scaphytopiini)

Varta Distant, 1908 placement nov. (transferred from Scaphytopiini)

Vartalapa Viraktamath, 2004 placement nov. (transferred from Scaphytopiini)

Vartatopa Viraktamath, 2004 placement nov. (transferred from Scaphytopiini)

Xenovarta Viraktamath, 2004 placement nov. (transferred from Scaphytopiini)

New species descriptions

A co ste m mini

Acostemma stilleri sp. nov.

um:lsid:zoobank.org:act:202D6A00-lE0B-4990-A9CF-81B700031C68

Fig. 9

Diagnosis

There are 11 valid species of Acostemma, 8 of which are known from Madagascar. The male genitalia

of the new species appear to be unique and do not match any of the 6 species for which genitalia were

illustrated by Evans (1954) or A. phylia Evans, 1959. The genitalia of A. suboecia (Jacobi) were not

illustrated, but the size of the male is reported to be 12.5 mm, much larger than the species described here.

Of the species illustrated by Evans (1954), A. stilleri may be most similar to A. sulcifrons (Signoret), but

the processes arising from the preatrium are much longer in A. stilleri and the the processes arising from

the shaft of the aedeagus are differently shaped.

Etymology

This species is named in honor of M. Stiller, the leading authority on South African Deltocephalinae.

177

European Journal of Taxonomy 45: 1-211 (2013)

Type material

Holotype

$, MADAGASCAR, Toliara Prov., Parc Nat. de Tsimanampetsotsa, Foret de Bemanateza, 20.7 km 81°

E of Efoetse, 23.0 km 131° SE ofBeheloka, 22-26 Mar. 2002, 23°59 , 32” S -43°52’50” E, coll. Fisher,

Griswold et al ., California Academy of Sciences, at light - in spiny forest thicket, elevation 90 m, code:

BLF6253. Deposited in coll. CAS.

Paratypes

3 SS and 1 $, same data as holotype; 1 $, MADAGASCAR, Toliara Prov., Mahafaly Plateau, 6.2 km

74° ENE of Itampolo, elev. 80 m, 21-25 Feb. 2002, 24°39’13” S -43°59’48” E, coll. Fisher, Griswold

et al, Calif. Acad, of Sciences, malaise trap - spiny forest thicket, code: BLF5759; 4 33 and 3

MADAGASCAR, Toliaran Prov., Parc National d’Andohahela, Foret de Manantalinjo, 33.6 km 63°

ENE of Amboasary, 7.6 km 99° E ofHazofotsy, 12-16 Jan. 2002,24°49’ 1” S -46°36’36” E, coll. Fisher,

Griswold et al ., California Academy of Sciences, at light - in spiny forest/thicket, elevation 150 m, code:

BLF4813.

One of the $ paratypes from locality BLF6253 is completely cleared and stored in glycerine. Genomic

DNA was extracted from this specimen, and DNA sequence data generated from it are included in the

analyses here. The specimen bears the voucher number “DEL 014”. 5 paratype males and 3 females

deposited at CAS, 3 paratype males and 1 female deposited at INHS.

Description

Size. $ 8.5-9.5 mm, 5 8-10 mm .

Body. Head as wide as pronotum. Crown slightly produced medially; longer medially than next to eyes;

surface texture longitudinally striate; depressed with distinct transition from vertex to frons; upturned at

apex. Anterior margin of head with single carina between ocelli. Face with epistomal suture evident just

below and mesad of ocelli; face shagreen below epistomal suture, rugose or irregular above. Clypellus

long, widening apically. Gena broad. Antennal ledge absent. Forewing wing with several extra crossveins

in anteapical and/or apical cells. A1-A2 crossvein present. Profemur row AM with AMI and 2 or more

additional proximal macrosetae; intercalary row with apical setae long, fine and with proximal setae

long, thick, extending more than half length of femur; row AV with short, stout setae. Protibia dorsal

surface rounded; macrosetae 1+5. Mesofemur row AV with numerous short, stout setae. Metafemur

apical setae 2+2+1. Metatarsomere I pecten with 6 long platellae, without tapered setae.

Male. Pygofer broad, rounded; with numerous short, thick setae posteriorly and posteroventrally; fused

to valve. Subgenital plates short; fused to each other; with several macrosetae subapically. Style in lateral

view bent apically at nearly 90° angle; apophysis curved, somewhat thick. Connective with anterior arms

separate at slightly more than 90°; median stem absent; with short posterolateral pair of arms; fused with

aedeagus. Aedeagus lying on top of and fused to connective; preatrium strongly modified into recurved

process, bifurcated near connective forming pair of slender acuminate processes; aedeagal shaft with

two asymmetrical processes, one arising from base parallel to shaft and terminating near gonopore, the

other arising near apex, curved anteriorly, extending beyond gonopore; gonopore apical. Segment X

long, narrow; 2.25 x longer than wide.

Female. Sternite VII length -1.75 x width; pointed and V-shaped apically; lateral margins convex

anteriorly, concave posteriorly. First and second valvulae illustrated in Zahniser & Dietrich (2008: figs 5b,

7d) as “Acostemma sp.”. First valvulae relatively slender; dorsal sculpturing pattern strigate to reticulate

or imbricate. Second valvulae slender; with -6 knob-like dorsal teeth restricted to less than apical 1/4

of valvulae. The eggs of this species are very unusual in having a tuft of lamellate seta-like extensions

178

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

at one end. This has not been observed in any other Deltocephalinae, but a similar modification was

reported by Wei et al. (2007b) in the stegelytrine Patanolidia magnified Wei & Webb. The setae appear

to be more filamentous in that species, and occupy a greater portion of the egg than in A. stilleri. Eggs

of numerous species of Deltocephalinae have been observed and are usually unremarkable in structure.

Remarks

At least 20 other species of Acostemma were collected in the Terrestrial Arthropod Inventory of

Madagascar project conducted by the California Academy of Sciences (CAS) and Tsimbazaza Botanical

and Zoological Park (PBZT), and none were assignable to described species. The available illustrations

of the genitalia of Acostemma spp. are not very detailed, and a revision of the genus would add much

needed clarity to the taxonomy of the currently known species.

Arrugadini

Arrugada linnavuorii sp. nov.

um:lsid:zoobank.org:act:24E57486-2093-4068-8054-57D22C2F59BD

Fig. 10 A-B, D-J

Diagnosis

A. linnavuorii is similar to A. rugosa Osborn, 1924 and A. affinis (Osborn, 1924) in the shape of the

pygofer but can be distinguished from them by the medially unproduced head (produced in A. rugosa

and A. affinis) and the shape of the aedeagus which, is longer, more bowed, and apex not spinose in the

new species.

Etymology

This species is named in honor of Dr. Rauno E. Linnavuori, author of 5 valid tribes, 2 subtribes, 127

genera and 671 species of Deltocephalinae.

Type material

Holotype

c?, PERU, Pasco, Villa Rica, 100 m, 23 Nov. 1971, R.T. & J.C. Schuh, coll. AMNH.

Paratypes

5 paratype sama data as holotype. Two specimens have an additional label reading: Collected on

mud road in forested area just east of town. Two other specimens have pink labels, one with #2885 and

the other with #2990. 2 paratypes are at AMNH and the remaining paratypes are at INHS.

Description

Size. S 7.5-8.0 mm.

Body. Head wider than pronotum. Crown dark brown to black in color, sometimes yellowish in part;

not produced medially; median length equal to length next to eye; texture strongly striate to rugose;

depressed; sharply angled to face. Ocelli on anterior margin of head; 2-3x their diameter from adjacent

eyes. Face strongly rugose. Clypellus widening apically. Antennal ledges distinct. Pronotum yellow

anteriorly, dark brown to black on posterior 2/3; carinate laterally; lateral carina longer than half basal

width of eye. Mesonotum yellow with pair of dark spots medially. Forewing brown; claval veins yellow;

sometimes with a few extra crossveins; usually with 1 to several crossveins between Al and claval

suture. Protrochanter with stout apical seta. Pro femur row AM with AMI only; intercalary row with long

fine setae; row AV with ~4 to 7 relatively long thick setae. Protibia flattened dorsally; dorsal margins

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European Journal of Taxonomy 45: 1-211 (2013)

angulate; macrosetae 5+1. Mesotrochanter with stout apical PV seta. Mesofemur row AV with ~4 to 6

relatively long thick setae. Metafemur apical setae 2+1+1. Metatarsomere I plantar setae irregularly

arranged.

Male. Pygofer incised dorsally to midlength; with several rows of macrosetae on posterior half; hind

margin notched posterodorsally; lobate ventrally; on either side of basolateral cleft, pygofer lobes

overlap, with posterior lobe articulating with dorsal extension of subgenital plate. Subgenital plates

constricted medially; macrosetae uniseriate distant from lateral margin. Connective Y-shaped; stem as

long as or longer than anterior arms; articulate with aedeagus. Style broadly bilobed basally; apophysis

short, digitate. Aedeagus with base of shaft tall, turning dorsad after some length; tapering apically;

gonopore subapical on ventral side. Segment X strongly sclerotized dorsally; elongate, narrow; basally

with protuberant dorsal lobe.

Remarks

One specimen is especially lightly pigmented but has identical male genitalia. Also collected at the same

time were three specimens of A. rugosa. These individuals, including the new species, were apparently

mud-puddling, which has been reported before in A. affinis by Rakitov et al. (2005).

Drabescini

Drabescus zhangi sp. nov.

um:lsid:zoobank.org:act:899B96Bl-7890-4256-8AF9-599E63134DB2

Fig. 19

Diagnosis

Externally, the new species is very similar in coloration to D. natalensis Distant, 1910, the only other

known African species of Drabescus. It can be distinguished from that species by the following

characters: crown not as strongly produced, coloration of crown not as mottled, style apophysis spatulate

(digitate in natalensis ), aedeagus with pair of long processes arising from ventral margin ( natalensis

with pair of short apical processes), and segment X with pair of long filamentous setae arising from

ventral margin (absent in natalensis). The holotype was compared with specimens of D. natalensis

which were collected at the same locality.

Etymology

This species is named in honor of Dr. Yalin Zhang, who has contributed extensively to knowledge of the

Oriental leafhopper fauna.

Type material

Holotype

c?, CENTRAL AFRICAN REPUBLIC, Maboke, 6 May 1969, Michel Boulard, coll. AMNH.

Description

Size. $ 9.2 mm

Body. Head wider than pronotum. Crown very slightly produced medially; surface shiny with longitudinal

and radial striations; dark brown medially at posterior margin, with transverse ivory band reaching

posterior margin laterally, with brown band anteriorly; with pair of dark brown spots near eyes; sharply

angled to face. Anterior margin of head with numerous irregular carinae; with distinct transverse ivory

band leading onto eyes. Frontoclypeus sharply keeled below dorsal margin; keel confluent laterally with

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ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

distinct oblique antennal ledges; texture longitudinally striate; dark brown. Clypellus constricted near

base; expanding apically; dark brown. Lorum dark brown; wider than clypellus at base. Gena ivory

below eye, dark brown above. Antennal sockets near upper corners of eyes. Antennae long, 1.5 x width

of head or longer. Pronotum and mesonotum ivory with 5 broad longitudinal brown or black stripes;

median 3 stripes tapering anteriorly and posteriorly. Pronotum carinate laterally. Forewings smoky;

veins brownish with very small whitish callosities, each bearing a very small seta; without extra

crossveins; with crossvein between A1 and claval suture; with A1-A2 crossvein. Pro femur row AM

without distinct setae; intercalary row with numerous scattered fine setae, not in a distinct row; row

AV with only scattered fine setae. Protibia dorsal surface flat; dorsal margins carinate. Mesofemur with

scattered longer ventral setae. Mesotibia dorsal surface flattened, margins carinate. Metafemur apical

setae 2+1. Metatibia row PD setae uniform in length.

Male. Pygofer incised dorsally past midlength; laterally with several macrosetae subapically; each side

with spine arising from ventral margin, slightly sinuate, directed posteriorly, surpassing pygofer sides.

Subgenital plates long, triangular; lateral margins straight to slightly concave; without macrosetae; with

several short fine hairs laterally. Connective Y-shaped; stem about as long as anterior arms; articulate

with aedeagus. Style broadly bilobed basally; preapical lobe squarish; apophysis incrassate, spatulate,

with stippled texture. Aedeagus U-shaped; with pair of slender process arising from shaft near ventral

margin, following shaft, slightly surpassing apex of shaft; gonopore apical. Segment X broad; sclerotized

laterally and dorsally on posterior half; with pair of very long filamentous setae arising from ventral side,

extending beyond pygofer.

Parabolopona webbi sp. nov.

um:lsid:zoobank.org:act:08E017C2-9224-4514-9FlF-CA86CCBFAA6A

Fig. 20

Diagnosis

P. webbi is similar to P. luzonensis Webb, 1981 with which it shares the preapically and sharply bent

style apophysis. It can be distinguished from that species by shorter, wider stem of the connective, the

aedeagus with a single basal process (P. luzonensis with a pair of basal processes), and apex of aedeagus

with a pair of processes (P. luzonensis without processes). It is also similar to the recently described P.

mutabilis Ohara & Kogure, 2012, with which it shares the unpaired ventral process of the aedeagus.

Ohara & Kogure (2012) report significant variation in the male genitalia of P. mutabilis , but P. webbi

appears to fall outside of the range of variability described and can be distinguished by the pygofer more

deeply excavated caudodorsally, the laterally directed apical processes of the aedeagus, the aedeagal

shaft narrower in lateral view, the ventral process of the aedeagus more widely separated from shaft, and

style apophysis longer and more sharply bent.

Etymology

The new species is named in honor of Michael D. Webb of the Natural History Museum, Fondon, a

leading world authority on leafhoppers.

Type material

Holotype

c?, TAIWAN, Nantou Co., 1550m, 24°5 , 5 ,, N- 121°9’5”E, 13 Jun. 2004, J.N. Zahniser, 08-1, sweeping,

coll. NMNS.

Description

Size. S 7.0 mm.

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European Journal of Taxonomy 45: 1-211 (2013)

Color. General color yellowish to greenish with two light orange longitudinal stripes from anterior

margin of head to mesonotum.

Body. Head about as wide as pronotum; produced; about 2 x longer medially than next to eye. Crown

texture shagreen with slight longitudinal or radial striations; transversely furrowed medially. Ocelli on

anterior margin of head, slightly dorsad; 3-4 x their own diameter from adjacent eye. Anterior margin of

head sharply angled; with two sharp carinae and numerous smaller irregular carinae. Frontoclypeus flat;

shagreen; depressed near anterior margin; relatively long. Clypellus expanding apically. Lorum wider

than clypellus at base. Antennal sockets near upper comers of eyes; antennal pits very large, encroaching

onto frontoclypeus; antennae very long, about 0.5 length of body. Pronotum carinate laterally; lateral

margin nearly half as long as eye width at base. Forewing yellowish-greenish; smoky and more hyaline

apically; apical veins darkly pigmented at apex; commissural margin with 3 small dark spots, 1 each

where anal veins meet margin, 1 at apex; with small dark spot at Cu-Ml and at base of M3+4. Legs

lightly colored and with numerous dark pigment spots, especially at bases of setae. Profemur row AM

with AMI seta only; intercalary row with 11-13 fine setae; row AV reduced to very small fine hairs.

Protibia rounded dorsally; macrosetae 1+4. Mesotrochanter with stout apical PV seta. Mesofemur

rows AV and PV each with 3 distinct long macrosetae. Mesotibia dorsal macrosetae 4+4. Metafemur

apical setae 2+2+1. Metatibia row PD setae relatively long, longer setae nearly twice length of row AD

macrosetae; row PD setae alternating shorter and longer. Metatarsomere I pecten setae with 3 platellae

between tapered AV and PV setae.

Male. Pygofer sclerotized dorsally nearly to mid-length; sides somewhat produced posteroventrally; with

numerous setae posteroventrally; inner wall with blade-like process arising from posterodorsal margin,

directed ventrally. Subgenital plates broad, rhomboidal in shape; apices digitate; without macrosetae.

Connective Y-shaped; anterior arms long, slender; stem fused to or modified into single lanceolate

process, process with longitudinal striate or rough sculpturing apically. Style broadly bilobed basally;

preapical lobe sharply angulate; apophysis sharply pointed medially; apex sharp, directed laterally; with

some stippled sculpturing preapically. Aedeagus with socle large; base broad; with unpaired medial

lanceolate process arising ventrally near base; shaft nearly straight, slightly upturned apically; apex with

pair of short apical processes directed dorsally then bent laterally; gonopore subapical on ventral side.

Segment X sclerotized laterally and somewhat dorsally; longer than wide.

Remarks

The holotype specimen is slightly teneral, but the genitalia appear to be fully formed and sclerotized

and indicate that it is a new species. P. webbi geographically occurs between the two most closely

related species, P. luzonensis and P. mutabilis. P. luzonensis occurs on the Philippine island of Luzon

lying approximately 400 km south of Taiwan. P. mutabilis is known from the Ryukus (Tokunoshima,

Kunigami and Nago) which lie -500-600 1cm NE of Taiwan. Small islands occur between the larger

islands of Luzon, Taiwan, and the Ryukyus, and thus dispersal via island hopping and subsequent

isolation may have promoted speciation in this group.

Goniagnathini

Goniagnathus emeljanovi sp. nov.

urn:lsid:zoobank.org:act:7A4C507B-38Al-483F-BA4A-D722D26ECBA4

Fig. 26 A-B, D-K

Diagnosis

G. emeljanovi can be distinguished from other species of the genus by the unique shapes of the style,

connective, and aedeagus. G. punctifer (Walker, 1858) is also recorded from Madagascar, and the new

182

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

species can be distinguished from it by the shorter pygofer, absence of a pygofer process, and absence

of paired apical or medial processes of the aedeagus.

Etymology

This species is named in honor of Dr. A.F. Emeljanov, who has made extensive contributions to the

knowledge of the Deltocephalinae.

Type matrial

Holotype

<$, MADAGASCAR, Diego-Suarez Prov., Parc National Montagne d’Ambre, 1125 m., 29 Jan.-ll Feb.

2001, 12 0 31’13” S -49°10 , 45” E, malaise trap, R. Harin ‘Hala coll., MA-01-01D-03, coll. CAS.

Paratype

1 5, same data as holotype, but with additional label with collection code: CASLOT 007672, coll. CAS.

Description

Size. S 5.2 mm, $ 5.5 mm.

Color. Generally brownish, mottled with darker brown and ivory markings.

Body. Head slightly wider than pronotum. Crown broad; slightly longer medially than next to eye;

glabrous with longitudinal striations; convex; rounded to face. Ocelli on anterior margin of head; 3 x

their own diameter from adjacent eye. Frontoclypeus tawny, speckled with dark brown and red dots;

slightly longer than wide. Clypellus widening apically. Loram wider than clypellus at base. Gena with

fine erect seta distant from laterofrontal suture. Antennal sockets near lower comers of eyes. Antennal

ledges weakly carinate. Antennae short. Pronotum carinate laterally; lateral margin slightly less than

0.5 x width of eye. Forewing with extensive brown irrorate markings; veins whitish, interrupted by brown

markings; with crossvein between A1 and claval suture; with A1-A2 crossvein. Protrochanter with stout

apical seta. Profemur row AV setae relatively long. Protibia dorsal surface rounded; macrosetae 3+4.

Mesotrochanter with stout apical PV and AV setae. Mesofemur row AV with ~7-8 relatively long setae.

Mesotibia dorsal setae 4+4. Metatibia pecten setae nearly equal in size. Metatarsomere I pecten with 5

platellae; PV seta platellate.

Male. Pygofer short; rounded; incised dorsally to midlength; with ~8 long macrosetae near posterior

margin; with numerous shorter thick setae posteroventrally; pygofer lobes nearly touching ventrally.

Subgenital plates fused; with numerous macrosetae subapically. Connective short; Y-shaped; fused to

aedeagus. Style very broad basally; median anterior arm long, extended dorsally to meet connective;

preapical lobe knob-like; apophysis with setae basally; apex expanded, with two points; apex textured

with several rows of small bumps. Aedeagus very broad at base in ventral view; excavated ventromedially;

bifurcate at apex; dorsal margin dentate apically.

Female. Sternite VII undulately excavated. Pygofer with medium-sized macrosetae ventrally and

subapically. First valvulae slender; strongly convex; dorsal sculpturing pattern strigate basally,

concatenate apically, reaching dorsal margin. Second valvulae slender; with short irregular dorsal teeth

and serrations on apical 1/3. Third valvulae with ventroapical row of ~6 distinct setae.

183

European Journal of Taxonomy 45: 1-211 (2013)

Hecalini

Hecalus hamiltoni sp. nov.

urn:lsid:zoobank.org:act:lE2332E7-7260-41B4-81B5-70077EBA59C5

Fig. 28 A-B, D-I

Diagnosis

Males of H. hamiltoni can be distinguished from other species of the genus by the aedeagus with very

distinctive long lamellate process arising ventromedially. All other species have shorter processes arising

apically or subapically.

Etymology

This species is named in honor of Dr. K.G.A. Hamilton, who has contributed greatly to the taxonomy of

North American Deltocephalinae and whose recent revision of the New World Hecalini facilitated the

discovery of this species.

Type material

Holotype

c?, USA, CA, Riverside Co., Rt. 37 1 mi E jet 79, Aguanga 6, 75 m, N 33°27’18” - W 116°51’34”, 24

Apr. 1998, RARakitov & CH Dietrich sweeping, 98-035, coll. INHS.

Paratypes

1 s, USA, CA, San Diego County Rd„ SI mile 15, 1479 m, 32°49 , 33”N- 116°29’45” W, 23 Apr. 1998,

C.H. Dietrich, vacuum, 98-031; 1 2 § § and 1 nymph, USA, CA, San Diego Co., Kitchen Crk. near

Cibbets Flat Cmpgd., 32°46 , 54” N- 116°26’39” W, 23 Apr. 1998, C.H. Dietrich, vacuum, 98-029. Two

additional nymphs from the last locality are stored in ethanol at INHS. All specimens deposited at INHS.

Description

Size. $ 5.7-5.8 mm, § 7.5-8.9 mm.

Color. Generally yellow to orangish-yellow.

Body. Head produced; narrower than pronotum. Crown slightly shorter than least width between eyes;

texture completely shagreen, basal half more shiny. Anterior margin of head foliaceous; slightly upturned;

in male, anterior margin colored white with fuscous above and below. Face of male mostly fuscous,

lighter anteriorly and near sutures; face of female yellow. Legs and ventral sclerites of male fuscous to

dark brown, of female yellow. Pronotum lateral margin longer than width of eye; carinate. Forewing of

male reaching mid-length of pygofer, slightly darkened apically; forewing of female reaching apex of

final pregenital tergite, not darkened apically; outer anteapical cell long, expanded apically; with several

extra straight costal veins arising from outer anteapical cell. Hindwings of male smoky, of female not

strongly colored.

Male. Pygofer pointed dorsoapically; broadly rounded ventrally; narrowly incised dorsally, incision not

reaching midlength of pygofer; with numerous macrosetae and some smaller setae. Subgenital plates

long; uniseriate laterally and with some smaller scattered setae and short fine setae near apex. Connective

Y-shaped; stem very short. Style very broad basally; preapical lobe distinct, rounded; apophysis slender,

pointed apically. Aedeagus laterally compressed; shaft robust, serrate ventrally; gonopore apical; with

pair of long lamellate processes arising ventromedially, extending dorsally and recurved; processes

serrate ventrally. Segment X basal half sclerotized dorsally and laterally; apical half membranous. With

small pair of sclerites in tissue connecting aedeagus and segment X.

184

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Female. Sternite VII broadly rounded; slightly undulate. Ovipositor protruding from pygofer.

Remarks

The aedeagal processes became somewhat more coiled after soaking in KOH.

Scaphoideini

Scaphoideus omani sp. nov.

urn:lsid:zoobank.org:act:3CB10731-F785-4BB6-B17D-E5D75DA4E39D

Fig. 47

Diagnosis

S. omani can be distinguished from other species of the genus by the color pattern and the shape of

the aedeagus. At least one other undescribed species with a very similar color pattern is known from

sub-Saharan Africa, but differs from S. omani in details of the color pattern, especially on the crown,

mesonotum, and base of the forewing, and in the male genitalia.

Etymology

This species is named in memory of Prof. Paul Wilson Oman, whose seminal work on the New World

leafhopper fauna made the present work possible.

Type material

Holotype

<$, ZAMBIA, Northwestern Prov., Zambezi Source NatT. Mon., S 11°22.172’ -E 024°18.622’, 1474 m,

6 Nov. 2007, Hg-vapor light, J.N. Zahniser, ZA-06, coll. INHS.

Paratypes

3 (?(?, same data as holotype, coll. INHS; 2 $$, 1 §, ZAMBIA, Northwestern Prov., ~15 km N of

Mwinilunga, FwakeraNational Forest, 11°34’38.2” S - 024°23’40.1” E, 1445 m, 5 Nov. 2007, Hg-vapor

light, J.N. Zahniser, ZA-05, coll. INHS; 1 specimen, sex unknown, RHODESIA, Victoria Falls NatT.

Park, 3-6 Apr. 1968, Paul Spangler, coll. NMNH.

Description

Size. Male 4.8-5.0 mm, female 5.3 mm

Color. Strikingly colored with contrasting bright yellow or ivory with dark brown areas. Crown bright

yellow/cream colored, with irregular brown markings near anterior corners of eyes. Face brown with

slight yellow markings. Pronotum and mesonotum brown laterally; bright yellow/cream colored medially,

expanded toward apex of mesoscutellum. Forewing brown on basal 1/3; brown patch acuminate apically

toward base of middle anteapical cell; hyaline or whitish on apical 2/3 except reflexed costal veins and

apical veins tinged with brown and brown spot in 3rd apical cell. Fegs and sternites mostly yellow

except mesosternum brown; legs with brown pigment at bases of large setae.

Body. Head narrower than pronotum; produced somewhat anteriorly; median length about 1.5 x length

next to eye. Crown depressed. Anterior margin of head mostly shagreen, glabrous at tip; angled to face.

Ocelli fairly large; situated laterally on anterior margin close to eyes. Frontoclypeus very long, narrow;

shagreen. Antennal pits large, encroaching onto frontoclypeus. Antennal sockets situated above middle

of eye. Antennae long, 1.5 x width of head or longer. Eyes notched along median margin. Clypellus

expanded apically. Forum wider than clypellus at base. Forewing veins R1 and R2+3 reflexed, strongly

pigmented; with an extra reflexed costal vein; outer anteapical cell acuminate basally and apically; inner

185

European Journal of Taxonomy 45: 1-211 (2013)

anteapical cell open; inner apical cell short, broad, triangular; with crossvein between A1 and claval

suture; A1 strongly curved distally, meeting commissural margin nearly at right angle. Profemur row

AV setae absent or reduced; intercalary row with ~10 long fine setae; row AM with AMI only, situated

near ventral margin. Protibia dorsal setae 1+4. Mesofemur row AV with numerous short, stout setae.

Mesotibia dorsal setae 4+4. Metafemur apical setae 2+2+1. Metatibia row PD setae very long; longest

setae nearly equal to or longer than protibia. Metatarsomere I long; apex with 5 platellae; apical row

inner seta platellate.

Male. Pygofer somewhat long, narrow; dorsal incision not reaching midlength; with numerous

macrosetae medially on posterior half; posterodorsal apex with vertical row of ~8 very long macrosetae;

internally with sclerotized bars reaching from base of segment X to anteroventral corner of aedeagus.

Valve distinctly pointed medially. Subgenital plates long, triangular; with ~3 macrosetae laterally; with

several long thin lateral and apical setae. Connective anterior arms touching apically; stem short; dorsal

margin of stem produced in lateral view; fused to pair of processes. Processes of connective fused to

each other along basal 2/3; fused to connective; in lateral view extending posterodorsally and curved

posteroventrally near apex; processes flared toward apex in ventral view. Style preapical lobe distinct,

with several long setae; apophysis long, gracile, with minute texturing. Aedeagus subquadrate in shape;

posterior and dorsal margins indented; with flange from dorsoapical corner extending anteroventrally,

spade-shaped in posterior view; gonopore on posterior surface near dorsal margin; disassociated from

connective and connective processes. Segment X long; lightly sclerotized dorsally at base.

Female. Sternite VII median length longer than width; produced medially; notched apically; with lateral

lobes. Sternite VIII very darkly sclerotized medially. Pygofer with numerous ventral macrosetae; apex

of each pygofer side with two bunches of 7-8 very long macrosetae; dorsoapical rim with group of 6

long macrosetae. First valvifers closely associate or lightly fused anteriorly; with distinct sclerotized

margins. First valvula convex; dorsal sculpturing pattern strigate to reticulate, reaching dorsal margin.

Second valvula with small irregular dorsal teeth reaching nearly to midlength; shaft with blunt dorsal

tooth.

Remarks

This species is placed in Scaphoideus based on the following characters: forewing outer anteapical cell

small, acuminate apically; forewing vein A1 strongly curved apically, meeting commissural margin at

nearly right angle; connective with long pair of processes (in this case, processes are fused to each other

basally and fused to connective); aedeagus disassociated from connective and connective processes.

Selenocephalini: Dwightlina

Dwightla delongi sp. nov.

um:lsid:zoobank.org:act:D2B3ECC6-5D8E-43A2-8DlA-543DllCABC40

Fig. 50 A-B, D-F, H-K

Diagnosis

D. delongi can be distinguished from other species of Dwightla by the long narrowly pointed pygofer

apex (similar in D. Hercules but lobe is wider in that species) and the aedeagus with a pair of long apical

falcate processes directed ventrally.

Etymology

This species is named after Dwight M. DeLong, author of 86 valid genera and 833 species of

Deltocephalinae.

186

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Type material

Holotype

<$, CAMEROON, Southwest, nr. Korup National Park, at bridge, 05°03’02.1” N - 08°56’41.0” E, 15-30

Mar. 2009, J.R. Cryan & G.J. Svenson, #8, coll. INHS.

Description

Size. $9.1 mm

Color. General color reddish-brown.

Body. Head slightly wider than pronotum. Crown broad; slightly longer medially than next to eye; about

4 x wider than width of eye; texture glabrous with longitudinal striae; depressed preapically; with 8 more

or less well defined black maculae. Anterior margin of head with 3 distinct carinae. Ocelli on anterior

margin of head; about 2 x their own diameter from adjacent eye. Face light brown; broad. Frontoclypeus

shagreen; distinctly broadening dorsally. Clypellus widening apically; somewhat ovoid. Lorum large.

Gena wide; with long fine erect seta laterad and distant from frontoclypeal suture. Antennal sockets

large; near upper corners of eyes. Antennal ledges distinctly carinate. Antennae somewhat long, about as

long as width of head. Pronotum carinate laterally; lateral margin about as long as width of eye; with ~10

more or less well defined black maculae in a fluctuate transverse line near anterior margin. Forewing

with numerous short fine hairs on veins and wing surface; apices folded over, forming pointed apex;

medial part of wing apex texture delicate; costal margin whitish, with small black spots near apex; with

1 or 2 crossveins between A1 and claval suture; with A1-A2 crossvein. Pro femur row AM with AMI

and 2 or 3 more proximal macrosetae; intercalary row with fine setae apically, apparently with larger

setae basally, basal setae continuous with row AV setae, generally in one row but with a few scattered

setae; row AV setae long; dorsally with 2 apical and 4-5 more proximal macrosetae. Protibia rounded

dorsally; macrosetae 1+7 (or so). Mesofemur row AV with ~10 short setae. Mesotibia dorsal setae ~5+7.

Metafemur apical setae 2+2+1. Metatibia row PV macrosetae thick with 1-2 slightly smaller lighter

setae between; dorsal surface with supernumerary short stout setae between rows AD and PD; pecten

setae nearly equal in length. Metatarsomere I pecten setae with 6 platellae; PV seta platellate.

Male. Pygofer incised dorsally nearly 2/3 its length; without macrosetae; produced caudoventrally;

sharply pointed at apex. Subgenital plate triangular; with scattered macrosetae laterally. Connective

broadly Y-shaped; stem very short. Style slender; preapical lobe evident in lateral view; apophysis

digitate, turned slightly medially at apex, ventral side with rough texture and subapical small tooth.

Aedeagus with pair of long falcate apical processes directed ventrally (holotype with one of these

processes broken at base and directed anteriorly). Segment X long; sclerotized laterally and dorsally,

and ventrally near apex.

Selenocephalini: Ianeirina

Abimwa knighti sp. nov.

urn: 1 sid: zoobank, org: act: 54F1AF 2D -EB 8D-4E4C-8C5B-FCF 0E A93 EB1B

Fig. 51

Diagnosis

A. knighti can be distinguished from other species in the genus by the relatively long and constricted

subgenital plate/valve complex; very short aedeagus with one pair of dorsal knob-like processes at apex,

and size and shape of the basal processes of the aedeagus.

187

European Journal of Taxonomy 45: 1-211 (2013)

Etymology

The species is named in honor of Dr. W.J. Knight, who has made significant contributions to the

knowledge of Old World Deltocephalinae.

Type material

Holotype

<$, ZAMBIA, Northwestern Prov., ~15 km N of Mwinilunga, 11°34’28.2” S - 024°23 , 40.1” E, 1445 m,

5 Nov. 2007, Hg-vapor light, ZA-05, J.N. Zahniser, coll. INHS.

Paratypes

1 S, 2 same data as holotype, coll. INHS. The $ paratype is completely cleared and DNA was

extracted and stored at INHS. Its voucher number is ‘DEL 104’. It was included in previous phylogenetic

analyses of Deltocephalinae (Zahniser & Dietrich 2010) listed as “Abimwa n.sp.”, and the same data are

included in analyses here.

Description

Size. $ and $ 6.3 mm.

Color. General color brown with extensive whitish coloration and small dots.

Body. Head slightly narrower than pronotum. Eye wider than 1/2 width of crown. Crown depressed;

texture smooth; sharply angled to face. Anterior margin of head with numerous transverse carinae.

Ocelli on face below carinae; about 1.5 x their own width from adjacent eye. Face somewhat darker in

color. Frontoclypeus shagreen. Laterofrontal sutures extending beyond ocelli to anterior margin of head.

Clypellus expanding apically. Antennal sockets near lower comers of eyes. Antennal ledges carinate.

Antennae about as long as width of head. Pronotum carinate laterally; lateral margin less than half width

of eye. Forewing whitish hyaline with numerous brown pigment lines and dots; veins brown. Pro-,

meso-, and metafemora with 2 broad dark brown longitudinal stripes or blotches, basal stripe or blotch

broader than subapical one. Pro femur row AM with AMI and 1 extra macroseta near base of intercalary

row; intercalary row with ~5-6 long fine setae; row AV setae stout. Protibia dorsal macrosetae 3(or 4)+6.

Mesofemur row AV with numerous short stout setae. Mesotibia dorsal setae ~6+6. Metafemur apical

setae 2+2+1. Metatarsomere I apical setae with 4 platellae; PV seta platellate.

Male. Pygofer reduced and modified; sclerotized dorsally only as transverse band; with short rounded-

rectangular lateral lobes without setae; lobes articulating posteriorly with anteroventral lobes of segment

X. Subgenital plates fused to valve; plates separated apically; with short fine hairs laterally; with several

uniseriate or scattered macrosetae near apex. Valve/plate complex very broad basally, constricted

medially. Connective Y-shaped; stem nearly as long as anterior arms; fused to aedeagus. Style small;

broadly bilobed basally; lateral anterior arm stalked, knob-like; preapical lobe absent; apophysis short,

rounded, blunt. Aedeagus fused to connective; with pair of long widely separated lateral processes

arising from base, directed caudodorsally; shaft very short; shaft with pair of short, knob-like processes

on dorsal side; gonopore large, apical. Segment X very large, occupying majority of dorsal side of genital

capsule; produced anterolaterally into pair of dentate lobes; with pair of medial darkly sclerotized plates

arising from inner margin; medial plates with digitate less sclerotized lobes anteriorly; lobes of medial

plates articulating with separate small, broad, chevron-shaped sclerotized plate.

Female. Sternite VII broadly excavated posteriorly. Pygofer with numerous setae ventroapically. First

valvifers fused anterodorsally. First valvula convex; dorsal sculpturing pattern strigate, reaching dorsal

188

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

margin. Second valvula with small knob-like dorsal teeth on apical 1/3 to 1/4; teeth not serrate; with

blunt median dorsal tooth. Third valvula with -8 distinct close-set setae ventroapically.

Remarks

Previous descriptions of species in this genus did not mention the inner plate-like processes of the male

segment X and the short median plate described here, so it is unknown if they are unique to this species.

Selenocephalini: Selenocephalina

Gannia viraktamathi sp. nov.

urn:lsid:zoobank.org:act:76324C0E-0625-483B-ACC6-AABCF336CE49

Fig. 52 A-B, D-K

Diagnosis

G. viraktamathi can be distinguished from other species of the genus by the unique shapes of the

aedeagus, ventral processes of the aedeagus, the pygofer, and male segment X.

Etymology

This species is named in honor of Dr. C.A. Viraktamath, a leading expert on Oriental Cicadellidae.

Type material

Holotype

S, ZAMBIA, ~12km SSE of Kitwe, Grey stone Farm, 12°55’03.8” S-028°15’52.6” E, 1247 m, 11 Nov.

2007, Hg-vapor light, ZA-09, J.N. Zahniser, coll. INHS.

Paratypes

1 <$, 1 5, same data as holotype, coll. INHS.

Description

Size. S 6.5-7.0 mm, $ 8.2 in in

Color. General color rusty brown to ochraceous with extensive dark brown mottled markings.

Body. Head wider than pronotum. Crown broad, more than 2 x width of eye; not produced; barely longer

medially than next to eye; texture glabrous with slight longitudinal or radial striations; sharply angled

to face. Anterior margin of head with 3 distinct parallel carinae between eyes; with a fourth less distinct

carina below distinct carinae. Ocelli large; close to eyes; less than 1 diameter width from adjacent eye.

Face broad. Frontoclypeus light tannish in color, with dark brown or black beneath anterior carinae;

widening dorsally; depressed below anterior margin of head; texture shagreen. Clypellus widening

apically. Forum large. Gena wide; with fine erect seta distant from laterofrontal suture. Antennal sockets

near middle of eyes. Antennal ledges not carinate; apparent as small bulges. Antennae shorter than width

of head. Forewing veins with short fine hairs. Forewing with crossvein between A1 and claval suture;

with A1-A2 crossvein. Pro femur ta nni sh in color, with dark brown or black ventrally near apex and

along intercalary row and spots of coloration dorsally near apex on anterior side; row AM with AMI

and with a smaller seta more proximally near base of intercalary row; intercalary row with -8-10 fine

setae; row AV with -6-9 relatively long thick setae. Protibia dorsal macrosetae 4+4. Mesofemur row

AV with -8 short stout setae; row PV with distinct somewhat longer setae; anterior side with extra setae

near apex. Mesotibia dorsal setae 4(or 5)+5. Metafemur apical setae 2+2+1. Metatarsomere I apex with

6 platellae; PV seta platellate.

189

European Journal of Taxonomy 45: 1-211 (2013)

Male. Pygofer not strongly incised dorsally; short; rounded; ventroapically with small lobate extension;

without macrosetae; with numerous short setae ventrally. Valve large; triangular. Subgenital plates

somewhat long; triangular; lateral margin convex; with numerous macrosetae scattered near lateral

margin. Connective short; U-shaped; without stem; with posterior angles sharp; articulating with

ventral aedeagal processes at meeting point of two processes. Style elongate; narrow; preapical lobe

squarish; apophysis digitate, slightly bent, with rough stippled texture medially. Aedeagus with socle

expanded; anteroventral corners extended ventrally and articulating with ventral processes; base of

aedeagus squarish with posteroventral corner sharp, with dorsal margin notched; shaft curved; gonopore

subapical on ventral side. Ventral processes of aedeagus extending from anteroventral lobes of socle of

aedeagus, meeting ventrally, then separating and continuing as long skinny sinuous processes to near

top of aedeagus. Segment X very large; distinctly protruding from pygofer; sclerotized for long length

laterally and as broad transverse band dorsally; ventroapically with several short, sharp teeth.

Female. Sternite VII slightly excavated, posterior margin broadly parenthesis-shaped. First valvifers

fused anterodorsally. First valvula convex; dorsal sculpturing strigate, reaching dorsal margin. Second

valvula broad, tapering apically; with small irregular teeth on apical 1/3; with blunt dorsal median tooth.

Third valvulae pigmented ventrally; with numerous short setae ventrally in one row and with other

scattered setae.

Stenometopiini

Doratulina dmitrievi sp. nov.

urn:lsid:zoobank.org:act:D44FB522-2ED8-4B79-97B7-FCA0EB60359E

Fig. 54A-B, D-J

Diagnosis

Other than the four species of the distinctive genus Hodoedocus Jacobi, 1910, there are three described

species of Stenometopiini from Africa: Doratulina sobrina (Melichar, 1912) from Chad, Gabon, the

Republic of the Congo, and Central African Republic; Stirellus flavovirescens (Stal, 1855) from South

Africa; and S. Sagittarius (Naude, 1926) from South Africa. The new species does not conform to any

of their descriptions.

Etymology

This species is named for Dr. D.A. Dmitriev in recognition of his contributions to the understanding

of the morphology of the nymphs of Deltocephalinae and for his contributions to the 31 database and

interactive key to Deltocephalinae.

Type material

Holotype

S, ZAMBIA, Northwestern Prov., ~28.5 km N of Mwinilunga, 11°30’ 16.1” S - 024°22 , 44.8” E,

Chitunta plain grassland, 1384 m, 4-7 Nov. 2007, ZA-03, sweep, J.N. Zahniser, coll. INHS.

Paratypes

9 SS, 6 $ 5, same data as holotype. All specimens deposited at INHS.

Description

Size. $ 2.9-3.2 mm, § 3.2-3.5 mm.

Color. General color light fuscous to dark brown or black. Males usually more darkly colored than

females.

190

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Body. Crown shagreen to base; coronal suture lightly colored; with pair of small ovoid dark markings

near base. Frontoclypeus fuscous with dark brown arcs from lateral frontal suture reaching only part

way across. Clypellus parallel-sided; extending slightly beyond normal curve of gena. Gena with large

dark brown spot below antennal bases; with small dark brown spot below eye. Pronotum, forewings,

and abdomen fuscous to dark brown or black dorsally; laterally with bright yellow or greenish stripe.

Forewings brachypterous, exposing 4 pregenital tergites; venation reticulated. Profemur row AM with

AMI and with an additional more proximal macroseta; intercalary row with a few fine setae; row AV

with many close-set, short stout setae. Protibia dorsal macrosetae 1+3. Mesotibia dorsal macrosetae 3+3.

Metafemur apex macrosetae 2+1 or 2+1+1; with several smaller scattered setae.

Male. Pregenital sternite lightly sclerotized on apical half. Pygofer dorsally incised to base; long, sloping

caudoventrally; apex subquadrate; with few macrosetae dorsally; with group of short setae ventrally; with

short tooth on apical half near ventral margin; with median lightly colored stripe separating dorsal and

ventral parts; apex darkly colored. Segment X long; sclerotized dorsally and laterally. Valve long, triangular.

Subgenital plates rounded laterally; macrosetae uniseriate laterally. Connective Y-shaped; anterior arms

with small lateral lobes; stem nearly as long as anterior arms; stem divided for most of length. Style

narrow; median anterior arm somewhat produced anteriorly; apophysis digitate. Aedeagus with narrowly

triangular base; flagellum long, parallel to base, about 2 x longer than longest length of base.

Female. Sternite VII slightly notched medially. Pygofer with few apical macrosetae.

Acknowledgements

We sincerely thank Dmitri Dmitriev (INHS) for his assistance with the 31 database and for helpful

discussion regarding the taxonomy of Deltocephalinae. We thank Stuart McKamey (NMNH) for

providing initial lists of taxonomic names that were used as the basis for the nomenclatural data entered

into the database. We are grateful to the following curators for assisting with museum visits and loans

that were instrumental in this project: Mick Webb (BMNH), Adeline Soulier-Perkins (MNHN), Jerome

Constant (IRSNB), Eliane de Coninck (RMCA), Stuart McKamey (NMNH), Randall Schuh (AMNH),

and Fuciana Musetti (OSUC). For providing specimens used in the phylogenetic analyses, we are

grateful to Mike Wilson, Roman Rakitov, Michael Stiller, Mike Irwin, Terry Erwin, Adam Wallner,

Vinton Thompson, Daniela Takiya, Murray Fletcher, Mick Webb, Charles Bartlett, Andrew Deans,

Norman Johnson, Fuciana Musetti, Mike Sharkey, Norman Penny, and Gordon Bennett. We thank

Allison Keever, Minerva Dorantes, Elizabeth Gonzalez Maza, and Stephanie Tham for their assistance in

photographing leafhopper specimens, editing images, and entering data into the database. We gratefully

acknowledge that this work was supported by NSF grant #DEB 08-41612 ARRA.

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Manuscript received: 30 August 2012

Manuscript accepted: 15 March 2013

Published on: 29 May 2013

Topic editor: Koen Martens

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d'Histoire Naturelle, Paris, France; National Botanic Garden

of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History

Museum, London, United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium;

Natural History Museum of Denmark, Copenhagen, De nm ark

Discla im er: The EJT consortium cannot guarantee that the internal links in this publication will remain

operable in the future.

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Appendix 1. List of morphological characters and states.

1. Head width: 0) as wide as or wider than pronotum; 1) narrower than pronotum.

2. Crown length: 0) parallel margined or somewhat produced; 1) strongly produced.

3. Crown width: 0) less than or equal to 2 x width of eye; 1) more than 2 x width of eye.

4. Crown surface shape: 0) convex; 1) flat or concave.

5. Crown texture at base: 0) shagreen; 1) punctate; 2) irregularly rugose; 3) glabrous and/or with

longitudinal or radial striae.

6. Texture of anterior margin of head: 0) shagreen; 1) striate or irregularly textured; 2) with numerous

carinae; 3) with 2 or 3 parallel carinae; 4) with a single transverse carina.

7. Position of ocelli: 0) on crown, posterad of anterior margin; 1) on anterior margin; 2) on face,

posteroventrad of anterior margin.

8. Proximity of ocelli to eyes: 0) close; 1) distant.

9. Lateral corner of eye: 0) even with or just above lateral carina of pronotum; 1) lying well above

lateral carina of pronotum.

10. Mesial margin of eye: 0) entire; 1) notched.

11. Lateral incision of gena: 0) not incised; 1) somewhat incised; 2) strongly incised, ca 90° angle.

12. Position of antennal pit: 0) near middle or posteroventral (lower) corner of eye; 1) near anterodorsal

(upper) corner of eye.

13. Antennal ledge: 0) absent; 1) wealdy carinate to carinate; 2) strongly developed, with ledge.

14. Ante nn al length: 0) short; 1) long; 2) very long.

15. Frontoclypeus length: 0) longer than wide; 1) shorter than or subequal to width.

16. Frontoclypeus texture: 0) shagreen; 1) rugose; 2) punctate; 3) striate; 4) glabrous.

17. Frontoclypeus in profile, anterodorsal part: 0) not inflated; 1) inflated.

18. Frontoclypeus in profile, posteroventral part: 0) not inflated; 1) inflated.

19. Sharply carinate ridge on frontoclypeus below antennal pit: 0) present; 1) absent.

20. Median longitudinal carina on frontoclypeus: 0) absent; 1) present.

21. Long fine setae laterally on frontoclypeus: 0) present; 1) absent.

22. Single fine erect seta on gena near lateral frontal suture: 0) present; 1) absent.

23. Proximity of fine erect seta to lateral frontal suture: 0) close; 1) distant.

24. Lateral frontal suture extent: 0) reaching ocellus; 1) absent or reaching only part way to ocellus.

25. Lateral frontal suture length above antennal pit: 0) distinctly shorter than clypeo-genal suture; 1)

subequal to or longer than clypeo-genal suture.

26. Lateral frontal suture direction: 0) mesad of ocelli; 1) toward middle of ocelli; 2) laterad of ocelli;

3) strongly divergent, ca. 90° angle.

27. Ratio of frontoclypeal-loral suture to clypellar-loral suture: 0) 1/3 or less; 1) more than 1/3 to 1; 2)

more than 1.

28. Lorum apex: 0) widely separated from genal margin; 1) near genal margin.

29. Lorum width: 0) distinctly narrower than clypellus at base; 1) subequal to or wider than clypellus at

base.

30. Clypellar suture completeness: 0) complete; 1) faint or obsolete medially.

31. Clypellar suture: 0) straight; 1) arcuate.

32. Clypellus shape: 0) tapered apically or parallel-sided; 1) expanded apically ovoid.

33. Clypellus apex: 0) not far surpassing normal curve of gena; 1) far surpassing normal curve of gena;

2) not attaining normal curve of gena.

34. Clypellus in lateral aspect: 0) not inflated; 1) inflated.

35. Apical margin of clypellus: 0) notched or concave; 1) straight or convex.

36. Pronotum lateral carina: 0) absent; 1) present.

37. Pronotum transverse striae: 0) absent; 1) present.

38. Forewing veins: 0) not raised, elevated, or carinate; 1) raised, elevated, or carinate.

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39. Number of anteapical cells: 0) 3; 1) 2.

40. R1 position: 0) basad of Rs; 1) distad of Rs.

41. Reflexed costal veins: 0) absent; 1) present.

42. Crossvein m-cu2: 0) absent (inner anteapical cell open); 1) present (inner anteapical cell closed).

43. A1 crossvein between A1 and claval suture: 0) absent; 1) present.

44. A veins: 0) gradually curved distally, meeting claval suture obliquely; 1) strongly curved distally,

meeting claval suture at ca 90° angle.

45. A1-A2 crossvein: 0) absent; 1) present.

46. Appendix: 0) absent or reduced; 1) restricted to anal margin; 2) large, extending around wing apex.

47. Hind wing submarginal vein: 0) complete; 1) obscured apically.

48. Protrochanter: 0) attaining posterior margin of mesosternum; 1) not attaining posterior margin of

mesosternum (mesosternum enlarged).

49. Protrochanter stout ventroapical seta: 0) absent; 1) present.

50. Profemur row AV: 0) reduced or absent; 1) with numerous (> 3) distinct setae; 2) with a few widely

spaced distinct setae.

51. Profemur row AV setae, length: 0) short; 1) long.

52. Profemur row AV, apex: 0) without setae in apical 1/4; 1) with several thin seta in apical 1/4.

53. Profemur intercalary row: 0) not in rows or scattered; 1) with five or more fine setae in one row; 2)

reduced or absent.

54. Profemur extra row of setae above intercalary row: 0) absent; 1) present.

55. Pro femur row AM: 0) with only AMI present; 1) with AMI and one or more additional proximal

setae; 2) absent.

56. Profemur dorsoapical setae: 0) with 2 apical setae; 1) with 1 seta or absent.

57. Profemur row PV basal and median setae: 0) reduced or absent; 1) present; 2) with one distinct

median seta.

58. Profemur row PV apical seta: 0) reduced or absent; 1) present.

59. Protibia dorsal margins: 0) rounded; 1) sharply carinate; 2) angulate (not carinate).

60. Protibia row AD: 0) without macrosetae; 1) with 1 macroseta; 2) with 2 or more macrosetae.

61. Protibia row PV macrosetae: 0) numerous; 1) 1 to 4; 2) absent.

62. Mesotrochanter stout apical PV seta: 0) absent; 1) present.

63. Mesotrochanter stout apical AV seta: 0) absent; 1) present.

64. Mesotrochanter extra setae: 0) absent; 1) present, with numerous extra thick or fine setae.

65. Mesofemur row AV setae: 0) reduced or absent; 1) with numerous distinct setae.

66. Mesofemur row AV setae, length: 0) short; 1) long.

67. Mesofemur row AV apical seta: 0) reduced or absent; 1) present.

68. Metafemur penultimate set of macrosetae: 0) none; 1) one; 2) two; 3) three.

69. Metafemur penultimate pair, proximity: 0) widely separated; 1) closely set.

70. Metafemur antepenultimate set of macrosetae: 0) none; 1) one; 2) two.

71. Metatibia shape: 0) arched throughout its length; 1) straight and distinctly bent apically in dorsal

view.

72. Metatibia cross-section shape: 0) square; 1) rectangular.

73. Metatibia dorsal surface, distinct proximal setae: 0) absent; 1) present.

74. Metatibia row PD setae: 0) alternating long and short; 1) macrosetae interspersed with 3-4 smaller

setae.

75. Metatibia row PD setae, length: 0) short to moderate; 1) long, nearly as long as or longer than half

length of protibia.

76. Metatibia row AD: 0) with macrosetae only; 1) with macrosetae and smaller intercalary setae.

77. Metatibia row AV macrosetae: 0) 6 or fewer; 1) 7 or more.

78. Metatibia row AV macrosetae, extent: 0) extending nearly to base; 1) restricted to apical 2/3 or less.

79. Metatibia row AV: 0) with macrosetae only; 1) with macrosetae and smaller intercalary setae.

205

European Journal of Taxonomy 45: 1-211 (2013)

80. Platellae at apex of metatibia: 0) absent; 1) present.

81. Metatarsomere I length: 0) shorter than tarsomeres II & III combined; 1) equal to or longer than

tarsomeres II & III combined.

82. Metatarsomere I apex: 0) straight; 1) expanded apically.

83. Metatarsomere I dorsoapical pair of setae: 0) absent or reduced; 1) present.

84. Metatarsomere I plantar setae: 0) all simple, tapered; 1) with one or more platellae.

85. Metatarsomere I plantar setae arrangement: 0) in two rows; 1) scattered.

86. Metatarsomere IPV apical seta: 0) platellate; 1) tapered.

87. Metatarsomere I apical platellae: 0) 3 or less; 1) 4 or more.

88. Metacoxa macrosetae: 0) with several macrosetae along midline; 1) absent or reduced.

89. Female pygofer macrosetae: 0) reduced (< 10) or absent; 1) present.

90. Ovipositor: 0) not protruding far beyond pygofer; 1) apex protruding far beyond pygofer apex.

91. First valvifers: 0) not fused; 1) fused anteriorly.

92. First valvula shape: 0) convex; 1) not strongly convex to straight; 2) concave.

93. First valvula dorsal sculpturing pattern: 0) strigate, concatenate, or reticulate; 1) granulose, maculose,

or imbricate (with overlapping scales); 2) with rectangular scales; 3) conchoid.

94. Position of first valvula dorsal sculpturing: 0) reaching dorsal margin; 1) distinctly below dorsal

margin.

95. Basal elongation of first valvula dorsal sculpturing pattern: 0) absent; 1) present.

96. First valvula distinctly delimited ventroapical subtriangular sculptured area: 0) absent; 1) present.

97. Bases of first valvulae: 0) not extended anteriorly; 1) extended anteriorly.

98. Second valvula shape: 0) broad, gradually tapered; 1) abruptly broadened medially; 2) slender

throughout; 3) gradually expanded medially or subapically.

99. Second valvula dorsal tooth on shaft: 0) absent; 1) present.

100. Second valvula teeth, shape: 0) large and prominent; 1) not large, regularly or irregularly shaped;

2) absent; 3) obtusely triangular, scalene.

101. Second valvula teeth, extent: 0) present on apical 1/3 or more; 1) restricted to apical 1/4.

102. Male pygofer macrosetae: 0) absent or reduced (< two rows); 1) present, well differentiated into

several rows.

103. Pygofer basolateral membranous cleft: 0) absent; 1) present.

104. Valve shape: 0) not triangular; 1) triangular, produced posteromedially.

105. Lateral margin of valve, length: 0) short, articulating with pygofer at a point; 1) long, with longer

length of articulation with pygofer.

106. Valve condition: 0) free; 1) fused to pygofer; 2) fused to subgenital plates; 3) partially fused to

pygofer.

107. Subgenital plate shape: 0) subrectangular; 1) triangular; 2) expanded medially and tapered apically;

3) plates fused; 4) rounded, with lateral lobe; 5) elongate, lobate, rising laterally.

108. Subgenital plate macrosetae: 0) absent; 1) scattered or irregularly arranged; 2) uniseriate laterally;

3) with two lateral rows and/or with other lateral setae; 4) one or two rows in middle of plate; 5)

with one or two rows near median margin.

109. Dorsolateral fold on subgenital plate, articulating with style: 0) absent; 1) present.

110. Style shape: 0) linear, median anterior lobe not pronounced; 1) broadly bilobed basally, median

anterior lobe pronounced.

111. Style preapical lobe: 0) absent or undeveloped; 1) present.

112. Style apophysis: 0) not elongate; 1) elongate.

113. Lateral anterior arms of connective: 0) widely divergent (connective T- or bar-shaped, or plate¬

like); 1) somewhat divergent (connective Y- or U-shaped); 2) closely appressed anteriorly (=

“linear”).

114. Connective anteromedial or ventral process: 0) absent; 1) present.

115. Processes of the connective or base of aedeagus: 0) absent; 1) present.

206

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

116. Connective articulation with aedeagus: 0) articulated; 1) fused.

117. Base of aedeagus: 0) not hinged; 1) hinged.

118. Number of gonopores: 0) one; 1) two.

119. Position of gonopore: 0) ventral; 1) apical; 2) dorsal; 3) dorsolateral.

120. Male segment X long lateroventral processes: 0) absent; 1) present.

121. Male segment X: 0) elongate, 2-3 longer than wide, sclerotized dorsally; 1) not strongly sclerotized

dorsally; 2) sclerotized dorsally, not particularly elongate.

207

European Journal of Taxonomy 45: 1-211 (2013)

Appendix 2. Morphological data matrix,

A.Krisnella

Abimwa

Acinopterus

Acostemma

Adama

Aflexia

Agrica

Allygidius

Anoplotettix

Antoerostemma

Aphr

Arrugada

Atanus

Athysanus

Attenuipyga

Bahita

Balclutha

Ballana

Bandaromimus

Bhatia

Bonaspeia

Brazosa

Cameroonl42

Caranavia

Caruya

Cerrillus

Cerus

Chiasmus

Chimaerotett

Chinaia

Chlo_uni

Chlor_rug

Chloropel

Cicadula

Citorus

Cochlorhinus

Colladonus

Copididonus

Dagama

Dalbulus

1111111111111111111111

1111111111222222222233333333334444444444555555555566666666667777777777888888888899999999990000000000111111111122

1234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901

101132100010200000100010021110110001100?0??0?1001100101001020101101211010001100010100010001000100211111100101100101000101

100132201010100000100010121110110011100000101100010010100102????10121101000110001010001?101000100011001?04121100101100102

1000311000101000001000100111101100111001001000001110101001020111111211010011100010100010100000100101011000001000001000101

00113410001010000010001002111011000110010100100011101010110101111012110100011100101000111010001002011100P1310010101100100

0001031000101000001000100211101100111000001011001110101001020111101211010001100010100011111000100101111100101110101000001

00013010?1100000001000001100001000101??????????001001000000211001012110100011100101001P1100010000001011100121110????00?02

00013310011010001010001001010001011110000010P10000P01000000101011012110101011100101001P1100000000001011P04131110100000102

000030100110100000101000011110110011100P01101100010010000102P100101211010001100010100010100010000011011100131110100000102

10003010011011000010100002111011001110001010010000P01000PP01P100101211010001100010100010100000100011011100101110100000101

00100001P0100010011000100111001000101PPPPPPPPPP011000000110111001012100100011000101000110001100000010010004011P0100000100

0011310100102001000001?100101000000111010a0000100211000010021101111211010001100010111110001100000001010010510011000000101

Oal1211000101001001000010P1110110001010000000100011010001122010111lbll000001110010100?110??111010302?01100121110100000100

00003010P1100000001000000111101100111001001001000100100000012100101211010001110010100100PPPPPPPPPPPPP11100121110100000101

0010311000100010001000100111001000111001000001000110100011020101101211010001100011100010100110000011011100111110100000100

011004PP00210000001001P10P110010001111PPPP0000P011102001110201011010P01001011110011100P1000111000001001100101110100000101

0001321001100000001000000211100001111001101111000100100001010101101211010001110010100100100000100011011100131110101000101

100000100010000000100000100110110010001P0000010000P01001000120000P0211010001110010100011100111010302P11100121110100000101

0000311000100000001000000211101000111001001001000100100000011100101211010001100010100010101000100011011100121110100100101

0000301001100100001000000211101100111001001001000100100000011100101211010011110010100010100100000001001100121110200100101

00113110001111000000000002111011001110010010010000P010000001P100111211010001100010100100101000100011011P00101110101000002

00013110P0101000001000100211101000111000011010000100100P01021101101211010001100010100010101000100001001100131110100000002

101030100000000000100000011110110001100100P0110001001000010111011012110100011100101000P0PPPPPPPPPPPPP11100121110100000101

100130101010100000010010021011000111100101101100010000000101P100101211010001100010100010100000100011011100101110100000101

0000301001100000001000000111101100111001001001000100100000012100101211010001110010100100100000000000011100121110101000101

000131100010000000100PP01211101101111001101011000100101001011100101211010001110010100101PPPPPPPPPPPPP01100121110100000102

10112311001010010010000100111011001110010001010000P0100011012101111211010001110010100100100000100000011100101110100000102

00003010 PI10100000100010021110110011100a0al0?0000100100001011101101211010001100010100010101000100001011100111110100000102

00010101002010000010001101110010001000010010020001102000010211011011P0000000110100000011010111000103000100101110200010201

00010011P011010000101001PP1011P1001010010010110000P010010001P0001111P00100011000101P100110P000P00012P11100101110100000101

10000021P11122000000101011101011201000PPPPPPP00100P01000010110010P1201010001100010100101010001000001100P10500011010000102

001000100010000000101000021110100011100100100100010010000102010010121101000110001011001010000001010101100012111010000010?

000000100110000000100000121110000011100100100100010010000001P1001012110100011100101001P0100000000001011100121110101000001

00111410P0210002001101P10P000010001001010000001001102021112111011010P00001000100010000P1010111010002P00PP110P100100000100

00000011P0100100001000000211100100111001000001000010100001011100101211010001110010100011100010000101010100141100100000100

001133100010101300100011PP1110010011100000101200111010101121P101111211010001100010100010101000100011001100111110101000102

0001311100100000001000000111001100111001010001000100100011021101101211010001110110110010101000100011011100121110100100100

0000301000100000001000000211101100111001000001000100100001011100101211010001110010100010100000000100011100121110100000201

000030100110000000100000011110010011100100a0al000100100000020100101211010011110010100110101000100011011100121100100000102

0001301000101100000000100211101100111000101011000100101011011100101211010011100010100010101000100011011100131110100000112

00003010P110010000100000010100000011101P001001000100100000011P001012110100P11100101000PP100010100001011100121110100000101

208

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

Deltocephalus

DeltocephAUS

Doratura

Dorydiella

Drabescus

Drakensbergen

Driotura

Dwightla

Elymana

Eryapus

Euacanthella

Eupelix

Eusaama

Euscelis

Eusceloidea

Eutettix

Evinus

Excultanus

Fieberiella

Flexamia

Gcaleka

Ghanal14

Glossocratus

Goniagnathus

Grypotes

Gurawa

Hecalus

Hecullus

Hishimonus

Hypacostemma

Idioceromimus

Ikelibeloha

Iturnoria

Jaf ar

Japananus

Kinonia

Kinrentius

Koebelia

Korana

Kramerana

Laevicephalus

Limotettix

Listrophora

Loipothea

1111111111111111111111

1111111111222222222233333333334444444444555555555566666666667777777777888888888899999999990000000000111111111122

1234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901

00003010P110000000100000010101P0001010010110010001001000000111001012

10010010P11000000010000011001010001010010110110001101000010211011012

00013110?0101000001000001201101000101????????20001101000010201011012

00113110001000000010000001111011001110010010000001001000110201001012

00113211001121030010000001111001001110010011120100P00000011011011111

01110011001100000110000001010010011111?????????001102021000210011001

00003000P1100010001010001210001000101PPPPPPPP20001101000110211011112

0010331000111010001010110P1110010011100000111200111010101102P1011112

000100100110010000100000010101010011100100101100010010000001P1001012

00113410P010200000100010021110110011100101PPP0001P1PPPP01122P1P11112

01113101P0211100001001P100000010000010PPPPPPPPP01210100P110111011111

0111000100210000001101P10P111000000101010000020011102021112211011010

00013010001000000010000001111011001110010010110001001000000111001012

00003010011010100010000002110010001110010010010001001000010211001012

000031110110001001101000111010100011100100100200010010000101P1001012

00003010001001000010000001111001001100010010010001001000000111001012

0100000101200000001000010P011000000100110010010000P01000010121000P02

10013010011010000010001002111011001110010010110001001010010211101012

00013210001010000010001003111011001110010010010011101010010111111012

00013010P110000000100000P10000000010100P1PPPPP00010010000001P0011012

0011321000101P000010001002111011001110000010100001101000010211011012

00003010001001001010c000021110010011101?00?0010000?010000101?1001012

011004100020000000100000011110000011100P0110100011001000110201011012

00003110001010100010001002111011000110000110120011101000010211111112

00000011P01000000010000102111000101110010010010010001000010110000P12

Oilldlll?0201000001011?0010010102001110100100000010020210021P0001001

00100110001000000010000000010010001010010000a00001001000010101001012

0110011100100000001000010?1110100011a00001101100110010000?0111011012

00003010001000000010000000111011001110010010110001001000000101001112

00103010001011000010001002111011011110010010P1000100100000011PPP1012

00100011001000000010001011P111P10001100000PPP10P00P01000010101000P12

101131211000220e0000101001?011?10001000100?0120100?010000021?1010?01

1000310010002P00001010P0121010110001100101P020001101100PPP01P1011112

101131000111201310100010002110110011100101101200111010001111P1P11113

110101101000000000101010021010010011000P00102100010010000001P1011012

00010010?11001000010000012010010001010?????????00100100001011101100f

101134101000000g00101000120000100111100000?0?201011???101122?0011112

01110021001000000010000001111000101000010000000011102000112200001112

10000010001010000010001001111011000110010010110001101010100201001012

OOllOal100200000001000011P11101000011PPPPPPPPPP011101000010211011012

00013010P110000000100000120100000010100101000100010010000P0111001012

00000010011000000010000002010010001110010010010001001000010111001012

OilPd41110102000001011P00100100100010101000000P0010020210021PPPP1000

00003010011000000010100001111001001110011110110000P01000110201011112

11010001110010100101100110000001011100121110200100101

110100P1100010100110100011010302P11100121110200100101

11010001110010100011010111000103000100111110200010201

11010001110010100011100010000001011100121100100000102

P001000011001010011010000000011P001100101100100000102

P0100101011001100011000110000001001100101110100000102

11010001100010100010010121000103011100101110200010101

11011001100010100010111000100200101100111010101000100

11010011110010100010100010000001011100121100100000100

110111P11000101000100011000001000100P13P0010100100100

P10000P1100001100111PPPPPPPPPPPPP10110550011010000100

P0100101011101000010000111000011001100101110100000101

11010001110010100010100010000011001100121110100000101

11010001100010100010101110000001011100121110100000202

11010001100010100100PPPPPPPPPPPPP11100121100100000102

110100011000101000P0PPPPPPPPPPPPP11100121110100000102

1101000111001010001P000111010302P01110401110100000101

11010001100010100110101000000001011100121110100000102

11010001100010100010101000100101011000120000000000111

11010001110010100110100010001011011100121110200100102

11110001100000110011?????????????01000131110101000102

11010011100010100100100000100111011100131110101000101

110100011000011000P0100110000301011100121110100000102

11010001100010100010101000100201011PP2300100P00100001

11000001110010110011100020000001011100111110100100301

P0100001100110100010010111000103001100101110200000101

110100011100001001P0010111010302P11100121110100000001

110100011100101001P0000111010002P11100121110101000101

11010011100010100010100000000001011100131110100001102

1101000111001010001P100000100011011100121110100000102

1101100110001010110110P000000101001100131010100000101

P1010001100010100010000100P00101000110100010010000102

1101000110001010001P10P000P00000000P123010P0011000101

11010001100010100100100000P000P1001100121110101000102

11010001100010100010100000100011111100101110200001101

P00100P11100101000P0110111010302P0010014111010001010?

12011001100010100laOPPPPPPPPPPPPPlllOOlOlllOlOOOOOlOl

00000000111110110011010101000301111100111110100000301

11011001100010100010100000000001011100121110100000101

110100011100111001P0000111010002P01100111110100000101

11010001110010100101100010000001011100121110200000101

11010001110010100010100110000001011100131110100000101

P000010001010000001P010111000103001100101110200010201

11010011100010100010100000100101011P02121110100000112

209

European Journal of Taxonomy 45: 1-211 (2013)

liiiiiiiiiiiiiiiiiiin

1111111111222222222233333333334444444444555555555566666666667777777777888888888899999999990000000000111111111122

1234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901

Loralia

Luheria

Macrosteles

Magnentius

Menosoma

Mimotettix

Mukaria

Napo

Neoaliturus

Neoheg075

Neohegira_br

Neomohunia

Nephotettix

Nesophrosyne

Nesothamnus

Occinirvana

Omanana

Opsius

Orientus

Orosius

Osbornellus

Oxycephalotet

Pachymeto

Pachytettix

Parabolopona

Paralimnus

Paramesodes

Paraphlepsius

Penthimia

Penthimidia

Penthimiola

Peru059

Phlepsius

Phlogotettix

Placidellus

Platymetopius

Polyamia

Portanus

Paradorydium

Proceps

Protochiasmus

Pseudophlep

Renonus

Renosteria

00013010?0101?0000100010011110110011100?00?0?00?0110100001020

0001321001101000001000100211101100111000000010001100101001020

00000010?0100000001000010?1100000010101?0010010000?0100000012

001?11210010200201100??1??1010110001110100101200?11010001101?

0000301001100000001000001111101101111001101011000100100000011

000030100110110000101000111110010011100100100100010010000001?

000132010011200310100001??11001101111001000001010010100000012

00003010001000000010000011111001001110010010010000P0100000011

00003010011000000010000001111011001110010010al000100100000011

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0001301001101000001000001111101100111000001001000100100001011

000132100111000010001000011100100011100100100100001010000101?

000131100010001000100000010100000101100a000002000100100001021

00013010?110010000101000021010110010100?1010010000?010000001?

100130101000010000100000111111?1001110010010?1000100100000211

01103401?011020000100011??1111?11010100?1010010000?010000021?

000030100010100000101000121110110011100?1?100100010010000001?

001030000010001000100000011110110011101?001001000100000000011

1001401010101?0000100000021110110011100000111100010010100101?

00013010?110000000101000121110110010100100102100010010000001?

00003010?1100100001000000111100100110001101001000100100000011

000134100010000e00100000011111?0011110011010110?0100100001011

100132101010210000100010011111?100010001000002001110111011010

0000311000101000001000000111101100111001001001000100100001011

0000031000111100001000010?1110010011100100?1010000?0100000011

00013010?1100000001000000100000000101001011001000100100000011

00003010?1100?0000100000?20001?0001110000000010?010010000002?

0000001000100000001000000211100100111001001001000100100000011

101021010011201310100001??111011011110000110?2011110101011100

111131010010200410100??0?21111?20111100101102201011?0??01110?

000031110011200010100001??111011001110010110110000?010000021?

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1001311001101000001000100111101100111001001011001100100001020

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100134101000020400100010121110010010100100101201?1111110110?2

000130100010000000100000011110110011100?0110?1000100100001021

00003010?110000000100000110001?000101001011011000100100000021

10003011?1102200000011?000111000200100010100010000?0100000011

01101110?0210002001101?10?01101000000101000000101010202111221001

010100100010010000101000011111?10011100100100100110010000001?100

00010001?1101000001000101110100000010??????????00110100000012101

101131101010000000101010121110110011100100?0?1001100100011020101

00013010?01001000010100011111011001010011010210000?010000001?001

0001301000100?00001000100211101000111000001001001100100001011101

1001012110100?110001010001?10100010000100?????????????100???

1?1101211010001100010100010101000100001011000111010000000112

000100211010001110010100111100110000101011100121110100000101

1011111?001000al10001101010000100000101001100101110100000001

100101211010011100010100100100000100011011100121110201000101

100101211010011100010100100?????????????11100121110100000101

1000?0211010011110010100100100100000001001100111110101101101

1001002110100011100101001?0?????????????11100101110100000002

100101211010001110010100010101010000201011100121110100001101

100101211010001110010100100?????????????11000101110101000102

1011012110100011100101000?1101000100013011000101110101000102

000101211010001100010100111100000000101000100101110100100102

1011012110100011100101000?1110111000303000100121110100010201

1000?0211010011100010100011100000000201011100121110100001101

1001012110100011000101001?0100000000101011100121110101000101

00010121100000110001011001110?001?0?0?1011100141110100000101

100101211010001100010100010100000100011011100101110101000101

100101211010001100010110010100000100000011100121110101001101

100101211010001100010100010101000100011011100101110101000101

100100211010011100010100010100010000202?11100121110100001101

1001012110100?1100010100010100000100000011100101110101000101

1011010110100011100101001?0100000000000011?04121110100000112

11011100000100110100000100103011100101001000000102

10001110010100010100000000101011100121110100000100

10011110010100100101100100101011100101110101000101

10011110010100111101110000001011100121110200000100

10001??????0?????????????11100121111200100100

110010100110100000001101011000121110100000101

100010100011100000000100011100111110100000101

10001010001011010000000101?000131110100000102

100010100100100000100101011100121110100000101

1100101001?0?????????????11100121110100000201

100010100010100000000001011100121110100000101

100010100100100000100011011100121110101000101

1000101001?01100001001010????????????????????

110010100011100000000000001100121110100000101

110010100100100010110001011100121110200100101

11000101001011100001001000011?3540010110000100

1011001000011010111010302?00??1401??0100000100

11000101000101001101000010????????????????????

11100111000?1?????????????01100101110100000102

1100010100010100000100011011100101110101001101

1100010100100100010000001011100101110100000101

1100010100010100000?00001011100131110101000101

? 11101???0

1001012110

1011112110

1001002110

? 001012110100?1

100101211010001

1?1111211010001

001111211010101

101101211010001

100101211010001

100101211010011

?? 11112120110?1

100101211010001

1011012110100?1

10010121101001

1110?010010

10121101000

1012100100 ?

10121101000

210

ZAHNISER J.N. & DIETRICH C.H., A review of the tribes of Deltocephalinae (Cicadellidae)

liiiiiiiiiiiiiiiiiiin

1111111111222222222233333333334444444444555555555566666666667777777777888888888899999999990000000000111111111122

1234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012345678901

Scaph040

000130

Scaph063

000130

Scaphl20

110130

Scaphoidophy

110130

Scaphotettix

000131

Scaphytopius

110131

Scpyl

100131

Selenocephal

001131

Stenometopi

000130

Stirellus

000000

Stymphalulus

010131

Taperinha

000130

Tenucephalus

000031

Tetartostylus

000000

Thailandll2

000134

Thamnotettix

000030

Trop 89a

000030

Tropicanl31

000030

Twiningia

000130

Xestocephalus

101000

Yungasia

100030

Zambiall8

000130

Zambiall90sb

000030

Zambia121

000000

Zambial220ps

000130

10011101000010000001111001001100011011010000P010000001110010121

10011101001010000000111001001100011011?10000?010000001?10010121

101110010000101000011110010011100?1011?10000?010000001?10010121

100110010000101000011110110011101?10100100010010000001?10010121

10?1110?00101000010?1101?00011100?00???10000?01000000120000?021

101000000000101000101110010011100?1?10?100010010000001?10010121

100000000000100000001110010011100P11111100010010000001010010121

1000101010001000110P1110110011100000101100111010001101010110121

100110000000100000021110110011100100100100010010000002110011121

10P110000000100000110100100000100000100100010010000101110110121

1010111100001100010P1010010001100000000100010010000P21110010121

10011000000010000001111011001110011110110000P010000001210010121

10011001000010000100010010001110010010010000P010000001100010121

10P01000000010000011110000001000000010010000P020000001100010121

11001010140110P001PP1000100111110100P0P100001011P00001P10101121

10001000000010000002110010001110010010010001001000??02110110121

100110000000100000111110110011100100P0P100010010000001110010021

10011000000010100001111011001110010010P100010010000001P10010021

100010000000100000011110110011100P11101100010010000001110110121

1101101100000001P1001111P0001110010110001000P01000200120000P01?

100010010000000000011110110011100101101100010010000001110010121

100110010000101000011110110011100111101100010010000001P10010121

100110010000101000021110110011100110100100010010000001P10010121

100110000000101000011110110011100011101100010010000101P00010121

100010000000101000101010110011100100101100010010000001P10011121

1010011100010100110101000100011011100121110101000101

10100111000101001101010000000110????????????????????

lOlOOlllOOOlOlOOal010000010001101110012 1111201000101

1010011100010100010100000001001011100121110100000101

10100P11100101001P110110000000101110012111010000000?

1010001100010100la0?????????????11100121110101000102

1010011100010100laO101000000100011100121110101000101

1011001100010100010?????????????01100131110100000101

1010001110010100011100110000101000100441110100000100

1010001110010100011010111010302P00100441110100000101

10110011000101001?01000000000000?110011?1?010?1001??

1010011110010100100100000100011011100131110100000101

10100111100101001011111P1010202P11100121110100000001

1010001110010100110100111000001011P0012 1110201000101

1010001100010100100100000000011011100111110100000102

1010001100010100010100000000011011100121100100000102

10100111100101001P0100011000103011100121110100000101

1010001100010100laO100000000103011100121110100000101

10100011000101000?0?????????????01100121110101000102

1010001101010100101100230000401000010550011010000101

10100111100101001P0101000000001011100121110100000102

1010011100010100011100000100103011100101110100000100

1010011100010100100100000001011011100121110101000102

1010001100010100010100000000001011100121110100000102

1010001100010100010100000000011011100121110101001101

(0,l)=a; (1f 2)=b; (l,4)=c;

(0,2)=d; (0,3)=e;

(0,1,2) f;

(0,4)=g