M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 40 20 10 B 80 60 40 20 10 M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 40 20 10 B 80 60 40 20 10 M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 40 20 10 B 80 60 40 20 10

Cover illustration (from top to bottom): Zygina flammigera (Geoffr.),

The Leafhoppers

Herbert Nickel
Errastunus ocellaris (Fall.), Conomelus anceps (Germ.),
by courtesy of Ekkehard Wachmann. Background by courtesy of Thomas Fechtler.

and Planthoppers
of Germany

The Leafhoppers and Planthoppers of Germany

Auchenorrhyncha, the planthoppers and leafhoppers, occur in high
density and species richness in almost all terrestrial ecosystems,
forming an important component of food webs. They may transmit
plant pathogens such as viruses and mycoplasmas, and a few species
Patterns and strategies
are among the worlds’ worst pests of cereals and other crops. Due to in a highly diverse group of
the lack of a comprehensive key, the central European fauna has long
been neglected by entomologists, and as a consequence, the knowledge phytophagous insects
of their life history was rather small. Host relations, in particular, were
insufficiently known. This work summarises an extensive data collec-
tion combined with a widely scattered literature on a large geographical
scale. For the first time, a systematic overview of phenology, food Herbert Nickel
plants, habitat requirements, geographical and altitudinal distribution
and economic importance of all Auchenorrhyncha species known from
Germany is provided, based on up to date taxonomy. The guild compo-
sition of all plant groups on family and species level is analysed and
discussed in detail. Further analyses deal with life strategies such as
host specificity, dispersal and life cycles. The book is a highly welcome
synthesis and reference for all students of biodiversity as well as for
ecologists, entomologists, botanists and phytopathologists.

Co-published by

Pensoft Publishers Goecke & Evers Pensoft Publishers, Sofia–Moscow

ISBN 954-642-169-3 ISBN 3-931374-09-2 Goecke & Evers, Keltern

M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 40 20 10 B 80 60 40 20 10 M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 40 20 10 B 80 60 40 20 10 M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 40 20 10 B 80 60 40 20 10
 Contents i

THE LEAFHOPPERS AND PLANTHOPPERS OF GERMANY

(HEMIPTERA, AUCHENORRHYNCHA):
Patterns and strategies in a highly diverse group of phytophagous insects
Herbert Nickel
ii Contents

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 Contents iii

The Leafhoppers and

Planthoppers of Germany
(Hemiptera, Auchenorrhyncha):
Patterns and strategies in a highly diverse
group of phytophagous insects

Herbert Nickel

Co-published
by
Pensoft Publishers, Sofia-Moscow
Goecke & Evers, Keltern
iv Contents

THE LEAFHOPPERS AND PLANTHOPPERS OF GERMANY

(HEMIPTERA, AUCHENORRHYNCHA):
Patterns and strategies in a highly diverse group of phytophagous insects

© Herbert Nickel

Pensoft Series Faunistica No 28

ISSN 1312-0174

Co-published
by
Pensoft Publishers, Sofia-Moscow (ISBN 954-642-169-3)
Goecke & Evers, Keltern (ISBN 3-931374-09-2)

Pensoft Publishers, Acad. G. Bonchev Str., Bl.6, 1113 Sofia, Bulgaria

Fax: +359-2-870-45-08, e-mail: pensoft@mbox.infotel.bg, www.pensoft.net

Vertrieb in Deutschland, den anderen Mitgliedsstaaten der EU und der Schweiz

Goecke & Evers, Keltern (in Antiquariat Goecke & Evers)

Inh. Erich Bauer

Sportplatzweg 5, D-75210 Keltern, Germany
Fax: 07236-7325; International Fax (++49) (0)7236-7325;
E-mail: books@insecta.de

First published 2003

© PENSOFT Publishers

All rights reserved. No part of this publication may be reproduced, stored in a

retrieval system or transmitted in any form by any means, electronic, mechanical,
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copyright owner.

Printed in Bulgaria, May 2003

 Contents v

Dedicated to
Günter and Eva Groh
(† Feb 2nd 1996, and Feb 5th 1996)
vi Contents

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 Contents vii

Contents

1 Introduction ........................................................................................................................... 1
2 Study area .............................................................................................................................. 3
2.1 General remarks ............................................................................................................. 3
2.2 The north German plain ............................................................................................... 3
2.3 The region of the Mittelgebirge ..................................................................................... 5
2.4 The Alps, their foothills and foreland ......................................................................... 7
3 Material and methods ......................................................................................................... 9
3.1 Literature data ................................................................................................................ 9
3.2 Museum collections ....................................................................................................... 9
3.3 Own data ....................................................................................................................... 12
3.4 Sampling deficiencies .................................................................................................. 12
3.5 Sampling methods ....................................................................................................... 13
3.5.1 General remarks ................................................................................................. 13
3.5.2 Quantitative methods ........................................................................................ 14
3.5.3 Semi-quantitative methods .............................................................................. 15
3.5.4 Qualitative methods .......................................................................................... 16
3.6 Literature on identification ........................................................................................ 16
4 Review of species ............................................................................................................... 20
4.1 Fulgoromorpha Evans, 1946 – Planthoppers .......................................................... 22
4.1.1 Cixiidae Spinola, 1839 ....................................................................................... 22
4.1.2 Delphacidae Leach, 1815 .................................................................................. 29
4.1.2.1 Asiracinae Motschulsky, 1863 .............................................................. 29
4.1.2.2 Kelisiinae W. Wagner, 1963 .................................................................. 30
4.1.2.3 Stenocraninae W. Wagner, 1963 ........................................................... 36
4.1.2.4 Delphacinae Leach, 1815 ...................................................................... 38
4.1.3 Achilidae Stål, 1866 ............................................................................................ 64
4.1.4 Dictyopharidae Spinola, 1839 .......................................................................... 65
viii Contents

4.1.5 Tettigometridae Germar, 1821 .......................................................................... 65

4.1.6 Issidae Spinola, 1839 .......................................................................................... 69
4.1.6.1 Caliscelinae Amyot & Serville, 1843 .................................................. 69
4.1.6.2 Issinae Spinola, 1839 ............................................................................. 69
4.2 Cicadomorpha Evans, 1946 – Leafhoppers (s.l.) ..................................................... 70
4.2.1 Cicadidae Leach, 1815 (s.l.) – Cicadas ............................................................ 70
4.2.2 Cercopidae Leach, 1815 – Froghoppers .......................................................... 72
4.2.2.1 Cercopinae Leach, 1815 ........................................................................ 72
4.2.2.2 Aphrophorinae Amyot & Serville, 1843 ............................................ 74
4.2.3 Membracidae Rafinesque, 1815 – Treehoppers ............................................. 78
4.2.4 Cicadellidae Latreille, 1825 - Leafhoppers (s.str.) ......................................... 80
4.2.4.1 Ulopinae Le Peletier & Serville, 1825 ................................................. 80
4.2.4.2 Megophthalminae Kirkaldy, 1906 ....................................................... 81
4.2.4.3 Ledrinae Kirschbaum, 1868 ................................................................. 82
4.2.4.4 Macropsinae Evans, 1935 ..................................................................... 82
4.2.4.5 Agalliinae Kirkaldy, 1901 ..................................................................... 91
4.2.4.6 Idiocerinae Baker, 1915 ......................................................................... 94
4.2.4.7 Iassinae Amyot & Serville, 1843 ........................................................ 101
4.2.4.8 Penthimiinae Kirschbaum, 1868 ....................................................... 103
4.2.4.9 Dorycephalinae Oman, 1943 ............................................................. 103
4.2.4.10 Aphrodinae Haupt, 1927 .................................................................. 103
4.2.4.11 Cicadellinae Latreille, 1825 .............................................................. 109
4.2.4.12 Typhlocybinae Kirschbaum, 1868 .................................................... 111
4.2.4.13 Deltocephalinae Fieber, 1869 ........................................................... 163
4.3 Dubious records ......................................................................................................... 228
4.4 Short-term changes .................................................................................................... 230
4.4.1 Declines ............................................................................................................. 230
4.4.2 Range expansions ............................................................................................. 233
4.4.3 Introductions ..................................................................................................... 234
4.4.4 Occasional influxes .......................................................................................... 235
4.5 General remarks on the German Auchenorrhyncha fauna ................................. 235
5 Utilization of plant resources ........................................................................................ 237
5.1 Plant taxa ..................................................................................................................... 237
5.1.1 General remarks and problems ..................................................................... 237
5.1.2 Plant groups and their associated Auchenorrhyncha guilds .................... 242
 Contents ix

5.1.2.1 Pteridophyta ........................................................................................ 242

5.1.2.2 Gymnospermae ................................................................................... 244
5.1.2.3 Ranunculaceae ..................................................................................... 246
5.1.2.4 Ulmaceae .............................................................................................. 246
5.1.2.5 Urticaceae ............................................................................................. 248
5.1.2.6 Fagaceae ................................................................................................ 250
5.1.2.7 Betulaceae (incl. Corylaceae) ............................................................. 253
5.1.2.8 Tiliaceae ................................................................................................ 257
5.1.2.9 Salicaceae .............................................................................................. 258
5.1.2.10 Ericaceae ............................................................................................. 262
5.1.2.11 Rosaceae .............................................................................................. 264
5.1.2.12 Fabaceae .............................................................................................. 273
5.1.2.13 Aceraceae ............................................................................................ 275
5.1.2.14 Apiaceae .............................................................................................. 276
5.1.2.15 Lamiaceae ........................................................................................... 277
5.1.2.16 Asteraceae ........................................................................................... 279
5.1.2.17 Juncaceae ............................................................................................ 284
5.1.2.18 Cyperaceae ......................................................................................... 284
5.1.2.19 Poaceae ............................................................................................... 294
5.1.2.20 Further plant groups ......................................................................... 322
5.1.3 Differences between plant taxa ...................................................................... 329
5.1.3.1 Major plant clades ............................................................................... 329
5.1.3.2 Plant families ....................................................................................... 331
5.2 General conclusions on food plant choice ............................................................. 338
5.3 Stratification and plant architecture ....................................................................... 340
5.4 Utilized substrates ..................................................................................................... 344
5.4.1 Phloem ............................................................................................................... 345
5.4.2 Xylem ................................................................................................................. 345
5.4.3 Mesophyll .......................................................................................................... 348
5.4.4 Fungi .................................................................................................................. 349
5.5 Pests on cultivated plants ......................................................................................... 350
6 Life strategies .................................................................................................................... 352
6.1 Host specificity and its possible causes .................................................................. 352
6.1.1 Diet width in Auchenorrhyncha .................................................................... 352
6.1.2 Plant apparency ................................................................................................ 355
x Contents

6.1.3 Plant diversity .................................................................................................. 357

6.1.4 Plant defence ..................................................................................................... 358
6.1.5 Nitrogen ............................................................................................................. 361
6.1.6 Interspecific competition ................................................................................ 361
6.1.7 Resource predictability ................................................................................... 365
6.1.8 Neural constraints ............................................................................................ 365
6.1.9 Regional monophagy ...................................................................................... 365
6.1.10 Endophytic fungi ........................................................................................... 366
6.2 Dispersal ...................................................................................................................... 366
6.2.1 Wing length ....................................................................................................... 366
6.2.2 Migration ........................................................................................................... 369
6.2.3 Colonization ...................................................................................................... 372
6.3 Life cycle ...................................................................................................................... 374
6.3.1 Voltinism ............................................................................................................ 374
6.3.2 Dormancy .......................................................................................................... 377
7 Summary ............................................................................................................................ 381
8 References .......................................................................................................................... 385
Appendix ............................................................................................................................... 430
Index of Auchenorrhyncha genera and species .......................................................... 430
Index and vernacular names of plants ......................................................................... 442
Acknowledgements ......................................................................................................... 459
1 Introduction

And God said, Let the waters under the heaven be

gathered together unto one place, and let the dry land
appear: and it was so.
And the earth brought forth grass,
and herb yielding seed after his kind, and the tree
yielding fruit, whose seed was in itself, after his kind.
And God made the beast of the earth after his kind, and
cattle after their kind, and every thing that creepeth upon
the earth after his kind: and God saw that it was good.
Genesis 1: 9, 12, 25

Auchenorrhyncha, the planthoppers and leafhoppers, feed on plant sap and occur in
high density and species richness in most terrestrial ecosystems. Their functional signifi-
cance is poorly known, but due to their high abundances in the herb layer as well as in
tree canopies, their biomass in relation to other arthropod groups is high, and thus, they
are likely to form an important component of terrestrial food webs (Schiemenz 1969; An-
drzejewska 1979a; Waloff 1980; Curry 1994; Achtziger 1995; Körner et al. 2001; Nickel et
al. 2002a). They serve as hosts of diverse groups of insect parasitoids such as Pipuncul-
idae (Diptera), Dryinidae (Hymenoptera) and Strepsiptera, and they are preyed upon by
ants, spiders and birds (Waloff 1975; Andrzejewska 1979b; Sander 1985; Waloff & Jervis
1987; Guglielmino & Olmi 1997; Moreby & Stoate 2001). Further, a number of species has
been found to transmit plant pathogens such as viruses and mycoplasmas, and a few are
among the worlds’ worst pests of cereals (Maramorosch & Harris 1979; Nault & Rodrigu-
ez 1985). In most parts of western and central Europe, however, damages to cultivated
plants are limited, and the role of Auchenorrhyncha in ecosystem functioning is rather
discrete, because phloem and xylem feeders do not consume living plant biomass. How-
ever, their importance may have been underestimated, because they must ingest large
amounts of an unbalanced substrate to receive adequate quantities of nutrients. Honey-
dew of Auchenorrhyncha is rather thinly distributed, but may be a significant and pre-
dictable carbon source for ants and microconsumers such as bacteria, fungi and protozoa.
Finally, by feeding on specific host plants, herbivores may alter competitive relationships
between plants, which may potentially result in changes of the vegetation (Crawley 1997).
The two final points are in need of further experimental study.
2 Introduction

Auchenorrhyncha identification often requires dissection and examination of the gen-

italia. There is, however, no modern, comprehensive identification book covering the whole
of the central European fauna, although Holzinger et al. (2003) have started to work on
such a compilation. Students have to use various keys of adjacent countries (Haupt 1935;
Ribaut 1936, 1952; Dlabola 1954; Ossiannilsson 1978, 1981, 1983), monographs and origi-
nal descriptions, which may diverge in species interpretations and differ in accuracy. Hence,
the interest in this group has been rather limited for a long time. Most sampling methods
are rather unspecific and do not allow an assignment to certain host species and micro-
habitats. This can only be done by labourious suction sampling of single plants or by
direct observation and collecting of single specimens.
Hence, the knowledge of distribution, life histories and insect – plant relationships in
Auchenorrhyncha is rather incomplete. The available literature includes a number of
ecological and faunistic contributions, although most species statements are brief, un-
precise, unsystematic, of only regional validity or even contradictory. On the one hand,
many papers have been written by ecologists who had only little time to become ac-
quainted with the complicated taxonomy and who often changed the subject afterwards,
on the other hand, most experienced taxonomists deal with the fauna of more southern
countries. Therefore, a critical review and summary of information on a current taxo-
nomic state with special reference to food plant relationships does not yet exist. The
same is true for an analysis of a complete Auchenorrhyncha fauna and the utilized
flora.
This work is intended to be the foundation of such a summary, based on all the avail-
able data on life cycle, food plants, habitats, life strategies, as well as geographical and
altitudinal distribution of all species found in Germany. Many sections of this work will
be purely descriptive, and discussions on a number of points will be far from exhaus-
tive. Some aspects will be analysed elsewhere in more detail. Special emphasis is laid
on correct taxonomy, which is regarded here as a major basis for research on all aspects
of biological diversity.
During the process of writing this book, morphological and biomolecular evidence
has grown that the Auchenorrhyncha may not be a monophyletic taxon (Bourgoin 1986,
1993; Campbell et al. 1995; Sorensen et al. 1995). However, very recently, Yoshizawa &
Saigusa (2001) insisted on the classical opinion of a sister group relationship between
the Fulgoromorpha and Cicadomorpha proposed by Hennig (1969) and Kristensen
(1975), although based only on the reduction of a sclerite in the fore wing base, which
was interpreted as a synapomorphy. Hence, the term Auchenorrhyncha may become
obsolete in the future, but is certainly practical from an ecological and historical point
of view. Further ecological studies of plant-sucking insects should treat Fulgoromorpha
and Cicadomorpha separately and include other groups, notably Psylloidea, Aphidoidea
and most Heteroptera.
 Study area 3

2 Study area

2.1 General remarks

The study area is the Federal Republic of Germany (Fig. 1). Species of neighbouring
countries were not taken into consideration, even if they have been found in adjacent
regions, e.g. the Vosges, the central Alps and the Bohemian Forest. All information on
life history is taken exclusively from Germany, if not otherwise stated. Thus, certain
ecological traits of species may be different in other parts of the range, particularly life
cycle, food plant utilization and altitudinal distribution.
This introduction to the study area includes only a brief overview of the most impor-
tant geographical units and their position, geology, morphology, climate, soils, vegeta-
tion and human impact. Much of this information is taken from Liedtke & Marcinek
(1994) and Ellenberg (1996).

2.2 The north German plain

The north German plain generally comprises the glacially affected lowlands to the
north of the hilly regions which occupy much of the central part of Germany. It also
includes the basins of Westphalia, the lower Rhine, Cologne and Leipzig. The major
part is below 100 m a.s.l., although very small portions near the southern edge are above
200 m a.s.l. The area is covered by vast Pleistocene and Holocene deposits. The soils are
usually sandy and poor except in the more loamy marshes near the coast and the river
valleys (including the periglacial drainage systems called Urstromtäler), as well as the
loess belt stretching along the southern fringe, which is among the most productive
farming grounds all over Europe.
The climate is suboceanic, but with more atlantic traits in the western parts, the win-
ters usually being rather mild (with mean January temperatures above freezing point)
and – compared with other regions – somewhat cooler summers (mean July tempera-
tures around 17 oC, although slightly lower near the coast). Continental features in-
crease towards the eastern half. The winters are colder (mean January temperatures
slightly below freezing point), the summers are warmer. In the valleys of the Saale,
Spree and along the middle courses of the Elbe and Oder, mean July temperatures ex-
ceed 18 oC and are thus comparable with those in southwest German viticultural re-
gions. Likewise, the annual rainfall decreases from west to east, with usually more than
4 Study area

Fig. 1: Map of Germany

 The region of the Mittelgebirge 5

700 mm to the west of a line from Schwerin to Braunschweig (roughly the 11th meridi-
an), but below 600 mm to the east. Locally they even drop below 500 mm, notably in the
rainshadow area to the east of the Harz Mountains and in the Oder valley.
Much of the potentially natural vegetation is dominated by beech and common oak,
the former preferring well-drained and loamy terminal moraines of eastern Holstein and
Mecklenburg. Proportions of hornbeam, hazel and silver birch are variable, depending
on geographical and local conditions. These species are joined or replaced by durmast
oak and scots pine on rather dry sites, by downy birch and black alder on moist or wet
sites. Elm, maple, ash and small-leaved lime are rather uncommon and restricted to more
nutritious localities, mainly along the southern loess fringe and in the valley bottoms. The
latter are dominated by willows and poplars on frequently flooded sites.
Other habitats are not covered by forests, notably bogs, which are generally om-
brotrophic in western parts, but rather minerotrophic in the east. Peculiar to central
Europe are the coastal habitats, notably dunes formed by strong wind and constant
translocation of the substrate, as well as the saltmarshes, which also occur locally in-
land in the vicinity of geologic faults and salt works.
The formerly vast forests have been strongly reduced by human activities, their orig-
inal tree species composition having been altered. In particular spruce and pine have
substituted beech and oak in many places. Forests have been turned into arable land,
meadows and pastures, formerly also into heathland, which had developed after heavy
woodland grazing. Nowadays, most of these heaths have been reforested with pine.
The bogs have been destroyed almost completely; the few remnant sites are heavily
disturbed by former (and even present) drainage and peat digging, as well as by nutri-
ent immissions.

2.3 The region of the Mittelgebirge

The area between the foreland of the Alps and the north German plain is treated here
as a unit, for reasons of simplicity and due to the lack of zoogeographical borders. The
northern limit is formed by the drop-off towards the northern plains, which is locally
obscured by loess deposits of the Börde belt. The Danube valley and the slopes of the
Swabian Jura down to the Lake Constance basin (Hegau) are the southern limit. The
morphological diversity of the region is conspicuous. Major parts consist of various
Paleozoic and Mesozoic rocks, except the Pleistocene and Holocene sediments of the
larger river valleys (notably the upper Rhine plain) and the volcanoes of the Eifel, Wes-
terwald, Knüll, Vogelsberg and Rhön, most of which are of Tertiary origin. The altitude
ranges from 50 to 250 m a.s.l. in valley bottoms up to over 800 m a.s.l. on the limestone
uplands of the Swabian Jura, the Paleozoic plates of the Eifel, Rothaargebirge, Harz,
Thuringian Forest, Fichtelgebirge, Oberpfälzer Wald, Erzgebirge and the volcanic Rhön.
The Black and Bavarian Forest locally extend even up to almost 1500 m a.s.l., thus reach-
ing the subalpine tree limit. Nutrient contents and pH of soils are generally high in
regions of Jurassic and Triassic limestone, certain volcanic effusive rocks and alluvial
deposits, but rather low on sand stone and most Paleozoic rocks, particularly in higher
altitudes receiving more precipitation.
6 Study area

Climatic conditions range from the rather submediterranean upper Rhine plain to
permanently cool or cold uplands of the Mittelgebirge. Mean July temperatures are
around 19 oC along the upper and middle Rhine, in the valleys of Neckar, Main, Nahe
and Moselle, as well as in the Lake Constance basin, but only 14 oC or less in higher
altitudes above 800 m a.s.l., locally even below 10 oC along the ridges of the Erzgebir-
ge and the Bavarian Forest. Mean January values are highest in the upper and middle
Rhine valley and its major tributaries and usually exceed 1 oC. On the ridges of the
eastern Mittelgebirge (Thuringian Forest, Fichtelgebirge, Erzgebirge, Bavarian Forest)
and the Harz Mountains they are –4 oC or even lower. Continentality (pronounced
differences between summer and winter temperatures, low precipitation with sum-
mer peaks) is highest in the rainshadow region to the east of the Harz Mountains
(Saale-Unstrut region) and parts of northeastern Bavaria (the Grabfeld in the vicinity
of Bad Königshofen, the basin of Mittelfranken around Nuremberg and the lower
Naab valley to the north of Regensburg). Annual rainfall is below 500 mm in the Saale-
Unstrut region and parts of the northern upper Rhine plain. Rainshadow regions to
the east of the Hautes Fagnes (Belgium) and the Eifel Mountains, the basins of Hes-
sen, Thuringia and Mittelfranken, as well as the middle Main and the Naab valley are
also fairly dry, receiving 600 mm or less. On the other hand values are highest in the
mountains, locally reaching 1500 mm or even 2000 mm, notably in the Black and Ba-
varian Forest and the Harz Mountains.
Except in very small areas of bogs (mainly on rain-exposed mountain tops and in
valley bottoms), on rocky outcrops and in inland salt marshes, the potentially natural
vegetation consists of woodland. Beech plays a major role in many areas, in lower alti-
tudes with varying proportions of common and durmast oak, hornbeam, hazel, ash,
lime, maple and elm. With increasing altitude, most of these species step back, except
sycamore, wych elm and large-leaved lime. Beech, spruce and also fir prevail in the
montane belt, joined by rowan, downy birch and locally whitebeam on more open sites.
The overall predominance of beech is only reduced in regions with less than 600 mm
annual rainfall, as well as in cool, wet or temporarily flooded valley bottoms, on loamy
or clayey, temporarily wet soils and on shady slopes with moving scree or seeping wa-
ter. On these sites beech is often replaced by oaks, hornbeam, lime and ash, locally with
field maple and service tree, on shallow soils also with whitebeam, downy oak and
Montpellier maple. In permanently wet sites black alder, downy birch, willows and
poplars are found, replaced in the large valley bottoms, which are subject to annual
flooding, by ash, smooth-leaved and fluttering elm and common oak.
Man has reduced the woodland and strongly changed the primeval vegetation.
In the lowlands large areas are intensively managed by agriculture, while grazing is
prevailing in less productive soils of higher altitudes. Formerly, grazing by sheep
and goat was also widespread in lower altitudes, particularly in dry situations on
hilltops, plateaus and south-facing hillsides, thus developing and maintaining dry
grassland, which provided habitat for numerous Mediterranean and steppe spe-
cies. Nowadays this type of grassland has often been abandoned due to its low pro-
ductivity. In some places, however, it is sustained for reasons of conservation and
landscape management.
 The Alps, their foothills and foreland 7

2.4 The Alps, their foothills and foreland

These regions represent a single unit in terms of physical geography, but not from a
biogeographical point of view. The northern border is the valley of the Danube, which
flows along the southern slopes of the Swabian and Franconian Jura and the Bavarian
Forest. Most of the southern border is formed by the rugged limestone crests of the
northern Alps. The altitudes range from 300 to 400 m a.s.l. in the Danube valley, the
Lake Constance basin and along the lower courses of the Inn and Isar, to 500 to 900 m
a.s.l. in the moraine regions and the alpine valley bottoms, and to almost 3000 m a.s.l.
on the highest mountains.
The foreland of the Alps is essentially a depression filled with debris of diverse ori-
gin washed down from the mountains. At its southern fringe these Tertiary deposits are
overlain by moraines of at least 4 glaciations. Much of the surface is rather flat or roll-
ing, except the hilly regions predominated by ground or terminal moraines. The Alps
themselves are of orogenic origin and comprise the Low and the High Alps (Voralpen
and Hochalpen). The former essentially consist of thick layers of molasse, the latter mainly
of Jurassic limestone.
Most soils are loamy and rich in nutrients; the pH is usually high, except in bogs and
on moraine deposits originating from the interior of the Alps. Below 700 m a.s.l. the
climate is rather uniform, despite the large size of the area. Mean July temperatures are
between 17 and 18 oC (although slightly higher near Lake Constance and slightly lower
along the edge of the Alps). The January means decrease from around freezing point
near Lake Constance towards the east and northeast, reaching –2 to –3 oC in the Donau-
moos and Dungau and at the edge of the Alps. The annual precipitation is largely deter-
mined by the southward increase of elevation, ranging from 700 mm in the Danube
valley to almost 1500 mm near the Alps. In the higher parts of the Alps conditions change
drastically. Mean July temperatures may be less than 10 oC, whereas January means
drop below –5 oC. Depending on the exposure, peaks may receive 2000 mm or even
more annual rainfall.
Due to good drainage the potentially natural vegetation of the plains and moraine
regions is dominated by beech. However, in the glacial basins there are large lakes as
well as vast bogs, which are almost completely devoid of woodland. Furthermore, the
river floodplains, which are subject to considerable fluctuations of the water table, hold
forests of common oak, ash, wych elm and sycamore or open stands of grey alder, pop-
lar and willow. Locally on coarse riverine gravel open pine forests with spring heath
prevail (see below); the early successional stages in these sites are almost bare of vege-
tation and offer a unique habitat for heliophilous species. These types of vegetation also
predominate in the valleys of the Alps, although sun-exposed slopes on porous lime-
stone may be covered with monospecific stands of scots pine. Most deciduous tree spe-
cies vanish between 700 and 1000 m a.s.l., except beech and sycamore, which may reach
the upper montane belt. Higher up spruce and also fir form dense forests, in drier sites
and near the forest limit accompanied by larch and stone pine. The subalpine belt is
covered by a mosaic of dense stands of mountain pine, replaced by green alder and
willows in wet sites. Tall herbs and also grasses prevail where woody plants cannot
grow due to long-lasting snow cover, seeping water, disturbance by avalanches, graz-
8 Study area

ing animals and other unfavourable factors. This belt approximately extends from 1700
to 2000 m a.s.l., but conditions may be strongly modified by the local climate, exposure
and land use. The alpine belt is dominated by dwarf shrubs and grasses. It extends at
least up to 2500 m a.s.l. and is topped only by the nival belt, where the cover of vegeta-
tion becomes more and more scattered and finally disappears.
Man has cut large parts of the lowland forests and turned them into arable land,
particularly along the Danube, on the Tertiary hills and on the larger gravel plains. In
the Alps and the moraine region meadows and pastures mainly grazed by cattle pre-
vail, except on steep slopes which are still covered by forest. The original tree species
composition has been altered by planting spruce and other conifers. The minerotrophic
fens (notably along the Danube and the lower Isar) have almost completely been drained
and turned into pastures and fields, but the ombrotrophic bogs along the edge of the
Alps are still in good condition, compared with those of north Germany. After centuries
of cattle grazing, a dramatic expansion of pastures also occurred in the upper montane
and subalpine belt, thus lowering the actual tree line for several 100 meters in many
places.
 The Alps, their foothills and foreland 9

3 Material and methods

3.1 Literature data

The most important literature on distribution and ecology is listed in Table 1, al-
though there are numerous papers on further aspects of Auchenorrhyncha biology which
contain valuable data. These papers will be cited in the relevant chapters of species and
ecology. However, analysis of literature data was done with great care, since much ma-
terial, particularly from older papers, is in need of revision. Even recent papers may
include misidentifications if specialists have not been consulted. In a few extreme cases
more than 50% of the species in published or submitted lists were found to be misiden-
tified. Due to the fact that the most important progress in Auchenorrhyncha taxonomy
was made after focusing on genitalic morphology, which was applied in Germany for
the first time only by Wagner (1935 and thereafter), most publications before that were
excluded from analysis, unless the material had been revised.
In general, it appears that northern and eastern parts of Germany are better studied
than southern and western parts. Moreover, the coverage is very uneven, with regions
sampled for almost 150 years by a number of collectors, e.g. the vicinity of Mainz, as
well as conspicuous gaps almost without any data, e.g. the Saarland (see chapter 3.4).

3.2 Museum collections

Compared to some other insect groups, e.g. Lepidoptera and Coleoptera, Auchenor-
rhyncha are not well represented in most museum collections. Moreover, collectors have
focused on large and conspicuous species, like Cicadoidea, Cercopidae and Mem-
bracidae, whereas the highly diverse groups of Delphacidae, Typhlocybinae and Delto-
cephalinae have largely been neglected. Another problem is the bad condition of many
specimens, which have been mounted on micro-pins. Particularly small specimens are
much vulnerable to vibrations and have often lost abdomen, wings or other body parts.
Among the important public collections, most material has been checked during the
recent decade, or by experienced taxonomists in earlier years (see Table 2). A part of the
collection of W. Wagner (formerly Hamburg), which is now in the Museum of the North
Carolina State University (NCSU), was not revised. However, most data on distribution
and ecology have been published, and a few specimens or even types of most species
from northern Germany are kept in the Zoologisches Institut und Museum Hamburg
10 Material and methods

Table 1. References of important data on ecology and distribution of Auchenorrhyncha in Germany

Reference Study area Period Focus

Achtziger (1991, 1995) Franconia 1989-94 Hedges, forest edges

Bieman (1987a) Europe Review of Ribautodelphax
Bittner & Remane (1977) Rhön 1973-75 Bog fauna
Bornholdt (1991) Hessen, Franken 1986-88 Effects of management
Emmrich (1966) Vicinity of Greifswald 1963 Faunistics, ecology
Fischer (1972) Bavarian Swabia since 1813 Faunistic review
Fröhlich (1996a, 1997) Central Europe 1991-94 Salt marshes
Frommer (1996) Urban areas of Cologne 1994 Faunistics, ecology
Funke & Witsack (1998) Vicinity of Halle 1996-97 Former mining areas
Haupt (1925) Bavarian Alps 1924 Faunistics, taxonomy
Heller (1987a) Vicinity of Ludwigsburg 1979-86 Faunistics, ecology
Hildebrandt (1990a, 1995b) Weser estuary 1985-88 Effects of inundation
Kirschbaum (1868) Vicinity of Mainz mostly 1860s Taxonomy, faunistics
Kuntze (1937) Mecklenburg mostly 1936 Faunistics, ecology
Marchand (1953) Weser-Ems region 1951 Grassland habitats
Müller (1978) Leutratal near Jena 1971-74 Hillside gradient
Nickel (1994) Southern Niedersachsen 1990-94 Faunistics, ecology
Nickel (1997) Niedersachsen 1990-96 Faunistics, ecology
Nickel (1999a, 1999b) Bavaria 1993-98 Faunistics, ecology
Nickel & Achtziger (1999) Central Europe 1990-1999 Meadow fauna
Nickel & Remane (1996) Bavaria mainly since 1950 Species list, faunistics
Nickel & Voith (in prep.) Bavarian and Allgäu Alps 1987-2001 Altitudinal distribution
Nicolaus (1957) Eastern Thuringia mainly since 1925 Faunistics, ecology
Niedringhaus (1991) East Frisian Islands 1977-88 Island colonization
Niedringhaus (1997) Emsland 1989-94 Faunistics, ecology
Niedringhaus & Olthoff (1993) North-western Germany 1982-92 Faunistics, ecology
Nikusch (1976) Vogelsberg 1972-74 Faunistics, ecology
Post-Plangg & Hoffmann (1982) Eifel 1974-75 Dry hillsides
Remane (1958) Weser-Ems region 1954-55 Land use effects, ecology
Remane (1961a) Western Palearctic Review of Mocydiopsis
Remane (1987) Mainzer Sand 1955-85 Ecology, faunistics
Remane (1994) Central Europe Review of Zygina
Remane & Fröhlich (1994b) Western Palearctic 1956-93 Faunistics, ecology
Schaefer (1973) Bottsand near Kiel 1971-72 Coastal habitats
Schiemenz (1969) Eastern Germany 1963-66 Dry grassland
Schiemenz (1971a, 1975, 1976) Eastern Germany 1969-74 Bog fauna
Schiemenz (1987, 1988, 1990) Eastern Germany Faunal review
Schiemenz et al. (1996) Eastern Germany Faunal review
Schwörbel (1957) Spitzberg near Tübingen 1952-55 Faunistics, ecology
Trümbach (1959) Vicinity of Erlangen 1954-57 Faunistics, ecology
Wagner (1935 ff.) North-western Germany mostly after 1920 Faunistics, taxonomy
Wagner (1939a) Mainzer Becken mostly 1930s Faunistics, taxonomy
Wagner (1950) Central Europe Review of Macropsis
Table 2. Overview of the collections included in this study (study areas specified for larger collections only refer to Germany)

Collection Study area Revision

Naturwissenschaftliches Museum der Stadt Aschaffenburg Unterfranken Wagner (1951a)

Naturkundemuseum Bamberg Oberfranken Nickel (in prep.)
Zoologisches Museum Berlin (ZMB) Mainly vicinity of Berlin Schiemenz (1987, 1988, 1990), Schiemenz et al.
(1996), Nickel (unpublished)
Zoologisches Forschungsinstitut und Museum Alexander König (ZFMK) Middle Rhine Nickel (unpublished)
Staatliches Museum für Tierkunde Dresden (MTD) Mainly eastern Germany Schiemenz (1987, 1988, 1990), Schiemenz et al.
(1996), Emmrich (pers. comm.)
Deutsches Entomologisches Institut, Eberswalde (DEI) Mainly Brandenburg Schiemenz (1987, 1988, 1990),
Schiemenz et al. (1996)
Naturkundemuseum Erfurt (NME) Thuringia Nickel & Sander (1996)
F. Heller (now partim SMNS) Baden-Württemberg Nickel (unpublished)
Museum für Naturkunde Gera Eastern Thuringia Nickel & Sander (1996)
Museum der Natur Gotha (NMPG) Thuringia Nickel & Sander (1996)
Zoologisches Institut der Universität Halle (ZIH) Mainly eastern Germany Schiemenz (1987, 1988, 1990),
Schiemenz et al. (1996)
Zoologisches Institut und Museum Hamburg (ZIMH) Mainly north Germany Nickel (unpublished)
H. Haupt (merged in H.J. Müller’s, now MTD) Germany Schiemenz (1987, 1988, 1990), Schiemenz et al.
(1996), Wagner (1939a), Nickel (unpublished)
Staatliches Museum für Naturkunde Karlsruhe (SMNK) Mainly Baden Voigt (pers. comm.)
C.L. Kirschbaum (now Museum Wiesbaden, MWNH) Mainly Mainz basin Wagner (1939a)
H.J. Müller (now MTD) Mainly former GDR Schiemenz (1987, 1988, 1990), Schiemenz et al.
(1996), Nickel (unpublished)
Zoologische Staatssammlung München (ZSM) Southern Bavaria Schönitzer & Oesterling (1998a, 1998b)
R. Struve (now Westfälisches Landesmuseum für Naturkunde, Münster) Borkum Niedringhaus (1989)
Staatliches Museum für Naturkunde Stuttgart (SMNS) Mainly Swabia Nickel (unpublished)
W. Wagner (now partim ZIMH) Germany Nickel (unpublished)
Museum collections
11
12 Material and methods

(ZIMH). Furthermore, Wagner used to exchange material freely with numerous col-
leagues, e.g. F. Heller, H.-J. Müller and M. Nicolaus, which is now found in their collec-
tions. Due to Wagner’s excellent work, a revision is neccesary only in a few species
subject to very recent taxonomic changes.
A more serious problem is the almost entire loss of G.A.W. Herrich-Schäffer’s collec-
tion during World War II. Although many species have been properly described, doubts
remain about the identity of others, which cannot be clarified any more, e.g. Chloriona
unicolor (H.-S.), Zyginidia scutellaris (H.-S.), Rhopalopyx preyssleri (H.-S.) and Handianus
procerus (H.-S.). In these cases, the current interpretation follows the most relevant revi-
sions (Fieber 1866; Ribaut 1936; Vilbaste 1962; Emelyanov 1964a).

3.3 Own data

Gathering of material and data started in 1990, focusing on food plant relationships,
habitat requirements and distribution, with emphasis on the southern and eastern half
of Niedersachsen, Franconia and northern and central parts of Thuringia, furthermore
on the Bavarian and Allgäu Alps, Palatinate and southern Baden. Smaller collections
have been taken in the following regions: eastern Holstein, East Friesia, Lüneburger
Heide, western Mecklenburg, Drömling, Oderbruch, northern Hessen, Oberlausitz, Nahe
valley, Swabian Jura and southeastern Bavarian Forest. Altogether, more than 500 sites
have been sampled, the number of specimens taken exeeds 250.000.

3.4 Sampling deficiencies

Sampling deficiencies can either exist on a specific level (meaning that certain spe-
cies or groups are under-recorded by conventional methods due to peculiar life habits)
or be caused by the geographical distribution of taxonomists and their sampling areas.
In the first case, there is often a lack of knowledge of the life history in general, while the
latter results only in gaps of faunistic recording.
Most collectors use the sweep net. Hence, epigeic species are generally under-repre-
sented in many samples, particularly certain species of Delphacidae (e.g. among the gen-
era Paraliburnia, Megamelodes, Delphacodes, Criomorphus), Aphrodinae, Errhomenus, Endria
and Cosmotettix. This gap can be filled by the use of pitfall traps and by direct search on
the base of herbaceous plants. For this purpose, the grassy vegetation should be pressed
down to the ground (e.g. with the sweep net stick or the forearm) as flat as possible.
Insects appearing on the stalks and leaves can easily be taken with the aspirator.
With some experience, it is even possible to assign them to plant species. Dwellers of
thorny and spiny woody plants form another group, which is often overlooked, notably
those on Rosaceae, e.g. Rosa, Prunus spinosa, Rubus and Crataegus. Sweep-netting them
as well as beating is difficult and can cause holes in the net; thus, only few collectors
sampled on these plants.
Species and developmental stages living in the mineral soil and the litter evade most
conventional sampling methods. The knowledge of the nymphal ecology of many spe-
 General remarks 13

cies of Cicadidae, Cercopinae and Cixiidae is actually based only on a handful of casual
records and observations. For instance, there is only one single record of a resident
population of Trigonocranus emmeae Fieb., which is supposed to be the only central Eu-
ropean Auchenorrhyncha species spending its whole life cycle in the ground, although
single macropterous specimens may be caught on dispersal flights (section 4.1.1). Fur-
thermore, cixiid nymphs cannot be identified down to species level.
Some species spend the nymphal stages in the herb or shrub layer and ascend to the
tree canopy immediately after emergence. In certain cases, notably in Lamprotettix niti-
dulus (F.) and Colobotettix morbillosus (Mel.), perhaps also Colladonus torneellus (Zett.),
there are only few records of adults, since they prefer higher layers of the vegetation,
and their nymphs are yet undescribed. On the other hand most unidentified nymphs
can easily be reared to adulthood in the laboratory, provided their host has been identi-
fied. Usually a few twigs or stems put into a vase or cellophane bag are sufficient.
It has been stated that known distribution patterns of many invertebrate taxa rather
show the distribution of taxonomists and their favoured collecting sites. This is also
true for the majority of Auchenorrhyncha species. Thus, the surroundings of Bremen,
Dresden, Göttingen, Halle, Hamburg, Jena, Kiel, Marburg, München, Oldenburg, Stutt-
gart, Wiesbaden and Würzburg are by far the best-studied regions. Conversely, there is
an almost complete lack of data in some of the more remote areas, e.g. Schleswig, Uck-
ermark, middle and southern parts of Brandenburg, lower Rhine, Münsterland, Moselle
basin, Saarland, north Baden, eastern Bavaria and certain parts of Württemberg.

3.5 Sampling methods

3.5.1 General remarks

Auchenorrhyncha occur in almost all types of terrestrial habitats in humid to semi-
arid climates. Their abundance in the herb and canopy layer often exceeds 1000 ind./
m2, and even the soil and litter layer are utilized by some taxa or developmental stag-
es. Thus, due to high densities, non-quantitative sampling can be done easily. Howev-
er, it must be taken into consideration that the small-scale distribution of host special-
ists is often clumped. Gathering of quantitative data requires more effort and is main-
ly applied to the herb layer, but not to the shrub and canopy layer. Frequent blurring
of sampling results is caused by individuals which migrate or passively drift, or have
simply fallen down from trees. Such specimens do not belong to the autochthonous
fauna, but may affect results considerably, particularly species number and diversity
indices. In general, records of single and macropterous individuals as well as of
monophages away from their host plants should be treated with care, especially dur-
ing windy wheather, on hot summer days with strong thermals and on mountain ridges.
In the following sections, the commonly-used sampling methods will be enumerated
and discussed.
Much of the author’s material was gathered by non-quantitative sweep-netting with
special reference to potential food plants. Additional material originated from suction
catches, pitfall and Malaise traps as well as ground photo-eclectors.
14 Material and methods

3.5.2 Quantitative methods

At present, suction sampling is the only practicable way of gaining data on the abun-
dance of herb layer Auchenorrhyncha (Haas 1980; Törmälä 1982). For this purpose, a
conventional, petrol-driven leaf-blower can be converted by fixing a flexible or stiff
tube at the suction opening. The tube is supplied with a fine-mesh collection bag, fixed
around the inlet either by a cuff or by rubber bands (Stewart & Wright 1995). In addi-
tion, the apparatus may be mounted on a backpack frame, but can also be carried with
one hand. Finally, the sample is put into a plastic bag or a glass jar, either in alcohol or a
cold bag. Depending on the vegetation, there can be a large amount of litter and soil,
which requires a considerable effort for sorting out the insects. For preliminary studies
and, if only adults and older nymphs shall be sampled, it may be sufficient to put the
whole catch into an open box. Auchenorrhyncha climbing and hopping up the sides can
be collected with an aspirator, thus reducing the sorting effort, although some mac-
ropterous individuals may escape. After long experience they may even be identified
and released on the spot. However, a number of species, notably those belonging to
Macrosteles, Psammotettix and Jassargus, cannot be identified in the field even by experi-
enced taxonomists, as well as most smaller instars of nymphs. Hence, such individuals
should always be checked in the laboratory. Unknown nymphs can be reared until emer-
gence.
For experiments requiring statistical analysis, this method has shown to be very effi-
cient in most habitats with a complete cover of grasses and herbs, particularly in vari-
ous types of grassland and in open forests. In heterogenous habitats it yields much
better results than sweep-netting, which almost entirely neglects epigeic species and is
much affected by surface qualities and vegetation structure.
Ground photo-eclectors can be used for estimating densities of species spending the
nymphal stage in the soil and litter and move up to higher layers after molting into the
adult stage, namely among Cixiidae, Cicadidae and Cercopinae. Distribution may be
clumped around patches of oviposition, however. Hence, larger numbers of samples
may be needed for obtaining statistically significant results. This method is only practi-
cable within more extensive programs sampling all groups of arthropods emerging from
the soil. Although cixiid planthoppers may be common in some forest habitats, they
have rarely been treated on a specific level. On the other hand a tent-shaped photo
eclector put up only for a few hours for sampling the herb layer Auchenorrhyncha was
shown to be of little efficiency (Törmälä 1982).
The soil and litter fauna can be investigated by taking cores plus vegetation and
succeeding extraction. The recommended diameter of the corer should be at least around
20 cm. Extraction according to Kempson et al. (1963) requires a special apparatus, but is
very efficient, although estimates for the herb layer are usually too low due to escaping
individuals.
Canopy sampling for quantitative results is critical. Chemical knockdown, which is
now commonly applied in tropical forests, has occasionally been done in temperate
latitudes (e.g. Southwood et al. 1982; Floren & Schmidl 1999). Auchenorrhyncha have
not been treated on a specific level, but there may be potential for gaining accurate
estimates for canopy abundances. However, it is difficult to relate these numbers to
 Semi-quantitative methods 15

area size (Basset et al. 1997, see this paper also for further methods and discussion of
canopy sampling).

3.5.3 Semi-quantitative methods

The term „semi-quantitative“ is here referred to methods, which can be standard-
ized, but do not yield estimates, which can be related to the ground area size. Compara-
bility of the results is more or less limited.
In sites with similar height and structure of the vegetation and even surface, for in-
stance in most anthropogenic grasslands, standardized sweep-netting provides good re-
sults, which can be compared within the same study and on similar plots (Witsack 1975).
The sweep net mouth should be D-shaped or square in order to cover much of the soil
surface. Numbers of individuals and species caught are often high, although epigeic spe-
cies are usually much under-represented. However, comparison can be critical in tall and
inhomogeneous vegetation, with changing time of day and weather conditions and bet-
ween different collectors. Furthermore, comparisons between different species of the same
plot are critical because of differing vertical distribution (Andrzejewska 1965; Novotný
1992; Peter 1981). In fenland habitats with tussock sedges and on forest floors with only
scattered vegetation this method is not appropriate. For special questions, it may be used
along homogeneous fringes of forests and hedges (Achtziger 1995).
Sweep-netting was done for a large proportion of the existing studies on central Eu-
ropean grassland Auchenorrhyncha (e.g. Emmrich 1966; Marchand 1953; Niedringhaus
1991; Nikusch 1976; Remane 1958; Schiemenz 1969; Achtziger & Nickel 1997; Nickel &
Achtziger 1999).
Pitfall traps also yield high numbers of individuals, but fewer species. This may be
caused by the clumped distribution of many species, particularly those living monopha-
gously on tussock grasses and sedges. The traps usually catch all the epigeic species not
sampled by sweep-netting (Agalliinae, Anoscopus spp., Streptanus spp.). Thus, using
pitfall traps for Auchenorrhyncha is only efficient as a supplement of sweep-netting.
However, both methods combined sample almost the complete range of herb-layer spe-
cies, although a direct search may be necessary in large tussocks. An important short-
coming is the fact, that relation to area size is not given, and comparability may be
critical even within the same study.
Individual and species numbers caught in tree eclectors may show high variability,
depending on tree species, site and exposure. Nymphs and adults of obligate canopy
species, which have fallen down and climb up afterwards, are most numerous. Also,
obligate vertical migrants move up the tree trunks after emergence and are caught. All
these species are capable of flight, so the proportion of sampled individuals in relation
to the total number is unknown. Also, it is difficult to relate catch numbers and area
size. Altogether, tree eclectors offer information, which may be valuable, although diffi-
cult to interpret, particularly in closed forests. Some work has been done by Büchs (1988)
and Nickel (unpublished data). Other catches have been counted on supra-specific lev-
el only (e.g. Ellenberg et al. 1986).
At present, the Malaise trap is one of the most efficient methods for sampling species
living in taller and woody vegetation, in particular those migrating either vertically
16 Material and methods

(from one layer to another) or horizontally (from one plant to another), notably among
Cixiidae, Empoasca, Zyginidia, Zygina, Arboridia, Allygus, Allygidius and Thamnotettix.
Numbers of individuals and species are often very high and may exceed 3000 speci-
mens and 50 species per trap and season. However, the results must be interpreted with
care. Again, there is no relation to the area size, and nymphs are represented only in
small numbers, so reproduction on the plot is not proved, even if sampled quantities
are large. In autumn, there may be mass catches of Typhlocybine leafhoppers (see above)
migrating to their winter quarters. Despite the trap’s high efficiency and widespread
application, Auchenorrhyncha catches have been identified only a few times on specific
level (Fröhlich & Nickel, unpublished data; Nickel & Voith, unpublished data).
Sticky traps are frequently used for sampling insects in vineyards (e.g. Herrmann et
al. 1999; Louis & Schirra 1997), although more quantitative parameters have to be gained
by other methods. Density per leaf, for instance, is determined by direct counting, but
selection of a representative leaf sample may be critical due to differing nutritive qual-
ities in relation to sun-exposure and individual plant properties.
On shrubs and along forest margins with deep-hanging branches, a standardized
beating was carried out. A few good flyers and momentarily sucking individuals may
escape, but, depending on the study question, samples within the same study and on
similar sites may be comparable (Achtziger 1995).

3.5.4 Qualitative methods

If only species lists and food plant records are needed, non-quantitative sweep-net-
ting and direct search on the plants are very efficient. Specimens are taken selectively
with an aspirator, thus avoiding large catches of other arthropod groups and common
species, which have to be sorted out in the laboratory. When sampling large and heter-
ogeneous plots with quantitative or semi-quantitative methods, search on single plant
species may offer useful supplementary data on communities and regional host prefer-
ences. Therefore, two to four samples per season can produce almost the complete range
of species, provided the student has a good knowledge of food plants. Moreover, resi-
dence of most species can be proved by laying emphasis on nymphal records. In forests,
however, recording of arboricolous species is critical, except along glades and rides
with deep-hanging branches. Thus, some arboricolous species are frequently under-
recorded because they spend most of their life in tree canopies.

3.6 Literature on identification

There is no single concise identification key for central European Auchenorrhyncha

yet. Earliest major works, mostly dating from the second half of the 19th century, nota-
bly by Herrich-Schäffer (1829 ff.), Flor (1861), Kirschbaum (1868), J. Sahlberg (1871),
Fieber (1875, 1876, 1877, 1878, 1879), Edwards (1896) and Melichar (1896) form an in-
dispensable foundation for taxonomic research, but can hardly be used for identifica-
tion today. However, Fieber, and particularly, Edwards had a surprisingly well-deve-
loped knowledge of genital morphology, despite insufficient optical gear at that time.
 Literature on identification 17

This knowledge has long been ignored, notably in the series „Danmarks Fauna“ (Jens-
en-Haarup 1920), „Fauna van Nederland“ (Blöte 1927, 1943) and „Tierwelt Mitteleuro-
pas“ (Haupt (1935).
Later Ribaut (1936, 1952) presented an excellent monograph of the French fauna of
Cicadellidae, which was published in the series of „Faune de France“, setting a land-
mark for future work worldwide. Most species are described in detail, together with
excellent drawings of the genital apparatus. It encompasses most of the German leaf-
hopper fauna and shows deficiencies only in groups, the taxonomy of which is critical
still today (notably Macropsinae, Zygina). Della Giustina (1989) completed both vol-
umes with new information on taxonomy, nomenclature and faunistics, including nu-
merous additional species.
Ribaut’s style was adopted by Ossiannilsson (1946, 1947), who wrote a two-volume
book on the Swedish Auchenorrhyncha fauna for the series “Svensk Insektfauna”,
with numerous precise original drawings of genital structures. For a long time, this
work was the main identification tool for large parts of northern Europe including
northern parts of Germany. Later Ossiannilsson (1978, 1981, 1983) presented a stan-
dard work on the Auchenorrhyncha fauna of all Fennoscandia and Denmark, which
was published in the series „Fauna Entomologica Scandinavica“. It contains numer-
ous accurate drawings of all distinguishing features, including the complete male
genitalia plus song apparatus, as well as detailed descriptions of approximately 430
species, thus covering about two third of the German fauna and almost all species of
the north German plain.
For former Czechoslovakia Dlabola (1954) published an Auchenorrhyncha volume
in the series of „Fauna ČSR“. It includes some species not treated in other books, but in
some cases it lacks accuracy and detail.
The British Auchenorrhyncha were treated by Le Quesne (1960, 1965, 1969) and Le
Quesne & Payne (1981) in the series of „Handbooks for the Identification of British
Insects“. The whole work generally reaches a high standard. The German fauna was
covered by only 60%, but features of the male song apparatus were demonstrated for a
number of difficult taxa, after Ossiannilsson (1949, 1951) had done fundamental re-
search on Auchenorrhyncha bioacoustics, demonstrating that apodemes of male ab-
dominal tergites and sternites can be used for species discrimination.
Ribaut’s style was also adopted by Emelyanov (1964a), who presented a brief and
accurate key of the fauna of European Russia in the series of „Opredelitel nasekomych
evropeiskoj chasti SSSR“, offering an important supplement to the remaining works by
covering most of the eastern species.
Linnavuori (1969a, 1969b) compiled an excellent overview of the Finnish Auchenor-
rhyncha by collecting numerous figures from the original literature and encompassing
two volumes. Written in Finnish and published in the series “Suomen Eläimet” (Ani-
malia Fennica), this work received little attention, despite its high standard. Similarly,
the Fulgoromorpha, Cicadidae, Cercopidae and Membracidae of Estonia were compre-
hensively described by Vilbaste (1971). However, treating only a small portion of the
central European fauna and written in Estonian, it is of limited value for many users.
The series of „Fauna Ukraini“ includes a volume on the Ukrainian Fulgoromorpha,
written by Logvinenko (1975), which allows the identification of numerous central Eu-
18 Material and methods

ropean species, particularly of delphacids, thus forming an important supplement to

the remaining works. In some cases, however, species interpretations should be checked
and compared with those of western European authors.
Remane & Wachmann (1993) offered an introduction provided with numerous excel-
lent colour photographs, with a broad and concise overview of biology and diversity of
the central European Auchenorrhyncha fauna. Although it is not intended to be an iden-
tification guide, identification of conspicuous species can easily be done. Furthermore,
it can be used for visual confirmation of dichotomous keys.
Work on the compilation of a new key to the central European Auchenorrhyncha is in
progress and has just resulted in the completion of the first volume covering the Fulgor-
omorpha and Cicadomorpha excl. Cicadellidae (Holzinger et al. 2003, in print).
Useful supplementary information can also be taken from Mitjaev (1971) on the
Auchenorrhyncha of Kazakhstan and from Anufriev & Emelyanov (1988) for those of
the Far East of the former Soviet Union.
Apart from these, there are numerous taxonomic papers and monographs on certain
groups or geographical areas, which are indispensable for identification of some spe-
cies. Above all, W. Wagner has to be mentioned, who did an enormous amount of work
on the Auchenorrhyncha fauna of large parts of central Europe as well as the Mediter-
ranean region. By describing and revising taxa and clarifying uncertain names, he paved
the way for modern research on this group (Wagner 1937a, 1939a, 1939b, 1941a, 1944,
1948a, 1949a, 1949b, 1950, 1952, 1953, 1955, 1963b, 1964 and others).
More recent descriptions and revisions, which have not or only partially been inte-
grated into the above mentioned major works, have been presented by Asche (1980,
1982a), den Bieman (1987a), Booij (1981), Dworakowska (1971, 1972a, 1976, 1993), Eme-
lyanov (1964b), Gillham (1991), Hoch & Remane (1983, 1985), Lang (1945), Lauterer
(1958), Lindberg (1948), Linnavuori (1957), Nast (1981), Remane (1961a, 1965, 1994),
Remane & Holzinger (1995), Remane & Jung (1995), Ribaut (1934), Tishechkin (1998,
2002), Vilbaste (1972), Wilson & Claridge (1999) and Nickel (1999a). Further important
help for identification of certain taxa is offered by Asche (1994), della Giustina (1983),
Heller (1987a, 1987b), Holzinger (1999a), Nast (1966) and Remane & Fröhlich (1991).
For Cicadula and Jassargus it has been demonstrated that females within diverse gen-
era, which were by then considered indistinguishable, can be identified after dissection
of the oviposition apparatus (Remane & Fründ 1986; Schulz 1976). Thus, further re-
search on female genitalic morphology of difficult taxa may have the potential to solve
urgent taxonomic problems (see Bourgoin 1993).
Usually identification of nymphs is possible at least in later instars and down to ge-
neric level. Study of nymphs can be easily done by sampling separate food plants, thus
collecting both adults and nymphs together, and by rearing unknown nymphs in the
laboratory, if possible on their field host plant. However, literature on nymphal mor-
phology is scanty. Wagner (1950) was the first to realize that for Macropsis, which was
one of the most difficult Auchenorrhyncha groups by then, the nymphs are often easier
to identify than adults. It was Vilbaste (1968a, 1975, 1982), who achieved great progress
in nymphal morphology, who stressed its importance for systematics and who com-
piled a key for northern European taxa. A little later, Kathirithamby (1971, 1973) stud-
ied and described the nymphs of 22 selected British species of Cicadellidae as was done
 Literature on identification 19

by Walter (1975, 1978) for a number of central European Euscelinae. Wilson (1978) pro-
duced a key and descriptions for the genera of the British arboricolous Typhlocybinae.
Stewart (1986a, 1986b) almost completed this taxon with the British species of Eupteryx.
More nymphs have been described by Dmitriev (1999a, 2000, 2001, 2002), Guglielmino
& Virla (1997), Heller (1987b), Linnavuori (1951) and some others. Continuation and
synthesis of this research is urgently needed for further research on Auchenorrhyncha
systematics and ecology.
20 Review of species

4 Review of species

The earliest list of the German Auchenorrhyncha species was compiled by Hüeber
(1904), largely based on Puton’s (1899) catalogue of the Palearctic Hemiptera, who in
turn drew much information from Fieber (1875, 1876, 1877, 1878, 1879). However, this
happened before major development of genital morphology. Moreover, courses of po-
litical borders differed much from those of today, and the fauna of adjacent areas was
included. Later Nast (1972, 1979, 1982) presented an update by listing the Palearctic
Auchenorrhyncha species, together with data for single countries. This work was main-
ly based on the catalogue of Metcalf compiled between 1932 and 1968.
Afterwards, Nast (1987) compiled an overview of the European fauna in tabular form,
including separate columns for the former FRG and GDR, partially based on species
lists published by Schiemenz (1970), Vilbaste (1974), Gravestein (1976), Nast (1976a),
Dlabola (1977), Drosopoulos (1980), Drosopoulos et al. (1983), Le Quesne & Payne (1981),
Ossiannilsson (1983), Courcy Williams (1989), D’Urso (1995), Holzinger (1996a) and
(Holzinger et al. 1997). After the reunification of Germany, Remane & Fröhlich (1994a)
produced a critical species list, with clear discrimination between new and verified
records on the one hand and old and uncertain reports on the other. The basis for the
present analysis is a modified version of the former list, compiled after revising more
collections and sampling new material, and supplemented with data on food plants,
diet breadth, life cycle and stratum preferences (Nickel & Remane 2002).
The following section is divided into brief monographs summarizing data on life
history and distribution of each species. Generally, information is taken only from the
study area, sometimes supplemented by data from adjacent countries. For reasons of
shortness, more detailed studies of life history (e.g. bioacoustics, embryonic and nymphal
development, population ecology), are referred to by citation only.
Emphasis shall be laid upon the fact that in many cases conspecificity of central
European populations with those currently bearing same names in the south European
and Asiatic literature is not proven. Revisions and further taxonomic studies are needed
for clarifying the identity and distinction of many taxa, notably among Cixius, Reptalus,
Kelisia, Issus, Tettigometridae, Cicadetta, Macropsis, Idiocerinae, Anoscopus, Empoasca,
Fagocyba, Edwardsiana, Eupteryx, Zyginidia, Zygina, Neoaliturus/Circulifer, Macrosteles,
Platymetopius, Hardya, Ophiola, Metalimnus and Psammotettix, just to mention the most
critical groups.
It should also be stressed that in a number of taxa, differences between individuals or
populations commonly treated as different species were considered here as intraspecif-
 Review of species 21

ic variation. This view is particularly held for some typhlocybid leafhoppers belonging
to the genera Kybos, Fagocyba, Edwardsiana and Arboridia.
Only those synonyms are given, which are not mentioned by Nast (1972) and Ossiannils-
son (1978, 1981, 1983). In critical cases the species name is followed by taxonomic remarks,
including notes on recent changes, critical aspects, as well as diverging interpretations. The
next line contains abbreviated information on the life cycle, as was proposed by Schiemenz
(1987), including periods of adult occurrence, overwintering stage and annual generation
number (see below). These data are usually from warmer lowland areas of Germany or
from warmer years. Thus, it should be taken into account that phenology is usually delayed
in cooler years and in more northern and higher parts, resulting in differences of earliest
occurrence of several weeks. Furthermore, there may be geographical differences in gener-
ation numbers. In some cases, adults and nymphs of species usually overwintering in the
egg stage may survive in warmer areas or during mild winters until the next spring. Cases
of remarkable intraspecific variation are specifically mentioned.
Further, a habitat description is given with special regard to food plants, moisture
and light conditions. In addition, pH preferences are specified, if data are significant.
No data on acidity are given, if preferences are unspecific or unknown. Again, these
requirements may be subject to geographical shift. For instance, in the Alps Mocydia
crocea (H.-S.) is common up to at least 1000 m a.s.l., whereas in the central parts of Ger-
many it is confined to much lower altitudes and rarely occurs above 400 m a.s.l. The
encounter frequency in host plant stands is roughly divided into three classes (high,
intermediate, low). However, this parameter does not refer to individual numbers and
is much biased by sampling methods and conditions.
Distributional borders are described only roughly due to shortage of faunistic data.
Normally, larger cities are mentioned (at distances of no more than 30 km from the
locality) as well as more important geographical units. Maximum altitudinal records
are only stated, if residence is proved or likely. If mountain data from Germany are
insufficient, literature from alpine areas of Austria, Switzerland, Italy and France are
quoted. It has to be stressed here that mountain slopes and particularly ridges often
attract migrating and drifted Auchenorrhyncha, including many species originating from
much lower altitudes and different habitats. Some unusually high altitude data in the
literature may refer to such records. Another important aspect is the downward shift of
vegetation zones towards the north, which amounts to almost 500 m from the Alps to
the Harz Mountains (Ellenberg 1996).
After each species chapter references are listed. References for identification are only
given in critical cases as well as for species which are not or not sufficiently treated in
the main identification works (della Giustina 1989; Holzinger et al. 2003; Ossiannilsson
1978, 1981, 1983; Ribaut 1936, 1952). The following abbreviations were used:

I – XII Months of occurrence of adults

B Beginning (1st to 10th day of month)
M Mid (11th to 20th day)
E End (21st to 31st day)
egg Overwintering in the egg stage
nymph Overwintering in the nymphal stage
22 Review of species

adult Overwintering in the adult stage

gen. Generation
m a.s.l. Meters above sea level
MNHN Muséum National d’Histoire Naturelle, Paris
MTD Museum für Tierkunde Dresden
SMNS Staatliches Museum für Naturkunde Stuttgart
ZFMK Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn
ZIMH Zoologisches Institut und Museum Hamburg
ZMB Zoologisches Museum Berlin
ZMH Zoological Museum Helsinki
ZSM Zoologische Staatssammlung Munich
HN Collection of Herbert Nickel, Göttingen

4.1 Fulgoromorpha Evans, 1946 – Planthoppers

4.1.1 Cixiidae Spinola, 1839

Nymphal ecology of the Cixiidae is poorly known; most species live in the soil and litter layer
feeding on roots and perhaps fungi (Wilson et al. 1994). Adults are usually found above ground
on various dicotyledonous, often woody plants and are usually considered to be polyphagous.
However, the host range of nymphs may in fact be narrower, simply because roots of other plant
species are out of reach at least under trees. Numbers of sampled adults are generally low,
except in Malaise traps and around patches of emergence shortly after it. In some groups there
is an evolutionary trend towards retaining cryptic life habits during adult stage. Much of the
taxonomy of species treated here was clarified by Wagner (1939a), who studied material mainly
from central and southern parts of Europe, and China (1942), who focused on western European
material. However, the Cixiidae still comprise some of the most critical taxa among European
planthoppers (see below). Very recently, Holzinger et al. (2003, in press.) compiled an up to date
key to central European species.

Cixius nervosus (Linnaeus, 1758)

Two morphs occur in central Europe: the nominate morph and the var. longispinus W. Wagner,
1955. The latter’s aedeagal theca has a much longer lateral spine. Apparently, in the study area
they differ neither in ecology nor distribution. They are of equal abundance and often occur
together; intermediate specimens have been found. However, it is uncertain which morph cor-
responds to the Linnean type material.
M V – E IX; nymph, 1 gen.
Adults are found on various deciduous shrubs and trees, often Salix, also Alnus, Betula, Cory-
lus, Tilia and others; in wet to damp habitats of various types (along forest roads and margins,
hedges, in open forests, on solitary trees, even among subalpine alder scrub and alpine dwarf
shrubs), often near water. In Germany the most common and widespread species among
cixiid planthoppers, frequently up to at least 1850 m a.s.l. in the Bavarian Alps and also occur-
ring at highest altitudes of the Mittelgebirge, as well as on the Frisian Islands.
Achtziger (1991), Fischer (1972), Niedringhaus (1991, 1997), Remane (1987), Schiemenz (1987), Schwoerbel
(1957), Voigt (1978), Wagner (1935, 1939a, 1951a, 1955), HN and others
 Cixiidae Spinola, 1839 23

Cixius cunicularius (Linnaeus, 1767)

B V – E IX; nymph, 1 gen.
Usually along sunny margins of forests and hedges; adults are found on various decid-
uous shrubs and trees (Corylus, Betula, Alnus, Ulmus and others). The species is wide-
spread throughout Germany (although numbers of collected specimens are usually low),
up to at least 1200 m a.s.l. in the Mittelgebirge and the Bavarian Alps.
Fischer (1972), Schiemenz (1987), Schönitzer & Oesterling (1998a), Wagner (1935, 1939a), HN and others

Cixius simplex (Herrich-Schäffer, 1835)

Mainly E VI – M X, according to Schiemenz (1987) also M V; nymph, 1 gen.
Habitat requirements are poorly known; the species is found in dry to moist habitats,
adults mainly along woody margins (published food plants are Alnus, Betula, Tilia, Pru-
nus spinosa, Pinus), occasionally also among herbaceous vegetation of wet meadows
and fens. Widespread in the lowlands of Germany, but seldom recorded. Northernmost
localities are near Bremen, Lauenburg, Rostock, Greifswald and on the island of Rügen,
up to at least 600 m a.s.l. in the Mittelgebirge; not found in Denmark and Scandinavia.
Achtziger (1991), Fischer (1972), Nikusch (1976), Post-Plangg & Hoffmann (1982), Remane (1987), Schie-
menz (1987), Schönitzer & Oesterling (1998a), Schwoerbel (1957), SMNS, Wagner (1939a, 1940a, 1951a),
Weis & Schönitzer (2001), HN; see also Nast (1987), Ossiannilsson (1978)

Cixius wagneri China, 1942

After Wagner (1939a) revised Cixius species of central and southern Europe, China (1942) point-
ed out that Wagner’s interpretation of C. intermedius Fieber, 1870 referred to a new species and
created a name for it.
E V – E IX; nymph, 1 gen.
In semi-open stands of deciduous woody plants in damp to dry habitats, often in vineyard
situations, but also in floodplains. Food plants of adults are probably various deciduous trees,
shrubs and tall-growing herbs. In Germany restricted to the western parts, recorded from the
Rhine valley between the Kaiserstuhl and Bacharach, the valley of the Nahe (Schloßböckel-
heim) and the northern Eifel Mountains (near Mechernich), between 80 and 400 m a.s.l.
Remane & Fröhlich (1994b), Rombach (1999a), HN

Cixius distinguendus Kirschbaum, 1868

Most records E VI – E IX, according to Schiemenz (1987) also E V; nymph, 1 gen.
In closed forests and along their margins; most records of adults in stands of Fagus,
Quercus, Picea and Pinus. Widespread in Germany, but rather localized, mostly collected
in low numbers only. The highest known localities are at 650 m a.s.l. in the Bavarian
foothills of the Alps.
Fischer (1972), Mölleken & Topp (1997), Niedringhaus & Olthoff (1993), Rombach (1999b), Schiemenz (1987),
Wagner (1939a, 1940a), Nickel (1999b), HN
24 Review of species

Cixius sticticus Rey, 1891

According to Remane & Fröhlich (1994b) the relationship between this species and C. carniolicus
Wagner, 1939, described from Austria, should be investigated. The description of the latter is mainly
based upon slight differences in wing venation. Furthermore, there is a strong similarity to C. alpes-
tris W.Wg. (see below). In fact, all three taxa may be conspecific (see also Holzinger et al. 2003).
M VI – E VIII; nymph, 1 gen.
In warm and rather dry sites, mainly on dry grassland with scattered shrubs and along
sunny forest margins, occasionally also in open oak and pine forests. Adults are usually
swept from various shrubs (Rosa and others). In Germany rather localized and at the
northeastern edge of the range (but see taxonomic remark above), at present known
from southern Niedersachsen (Bodenwerder, Bad Sachsa), the Ahr valley (Altenahr),
the Rhön Mountains (near Hünfeld), Thuringia (near Arnstadt, Jena and Gera), along
the middle Rhine (Bacharach) and the Nahe valley (e.g. Schloßböckelheim, Bad Mün-
ster), up to 500 m a.s.l.
Reimer (1992), Remane & Fröhlich (1994b), Schiemenz (1987), Nickel (1997), HN

Cixius alpestris W. Wagner, 1939

See taxonomic remarks above.
So far E VI – B IX; nymph, 1 gen.
Reported from high mountains of Poland, Slovakia, Austria, northern Italy and France.
This species has recently been recorded in the Berchtesgaden and Chiemgau Alps
(Königsbach- and Gotzentalalm, St. Bartolomä, Geigelstein) and the Fricken above
Garmisch-Partenkirchen, between 700 and 1700 m a.s.l. Furthermore, ‡‡ which cannot
be indentified with certainty have also been found in other sites (e.g. Rotspitz above
Hindelang). Thus, it is probably more widespread in the Bavarian and Allgäu Alps. It
has been reported to favour “besonnte Schutthänge und Heidewälder” in Austria; spec-
imens from the Engadine (Switzerland) have been taken among a group of Pinus mugo
in subalpine scree.
Nickel & Voith (unpublished data), HN; see also Günthart (1987a), Wagner & Franz (1961)

Cixius heydenii Kirschbaum, 1868

B VI – B IX; nymph, 1 gen.
Largely confined to the Alps and the Carpathians; in Germany known only from the
Bavarian and Allgäu Alps and the Bavarian Forest. Adults are usually swept in the
shrub and herb layers of damp to moist habitats, mainly in open stands of trees and
shrubs of the upper montane and subalpine belt, as well as in alpine dwarf shrub heaths
between 1200 and 2000 m a.s.l. (in North Tyrol, Austria, up to 2200 m a.s.l.), although
single individuals have been collected down to 600 m a.s.l.
Biedermann (pers. comm.), Fischer (1972), Remane & Fröhlich (1994b), HN; see also Christandl-Peskoller
& Janetschek (1976), Leising (1977)
 Cixiidae Spinola, 1839 25

Cixius beieri W. Wagner, 1939

Cixius haupti Dlabola, 1949
C. beieri Wagner, 1939 has been described from the South Tyrolese Alps (Italy). Specimens from
Mittenwald in the Bavarian Alps were named C. montanus Haupt, 1924, which turned out to be
a primary homonym, later replaced by C. haupti Dlabola, 1949. According to Remane & Fröhlich
(1994a) C. haupti Dlab. and C. beieri W.Wg. are likely to be conspecific. The only character sepa-
rating them is the length of a lateral spine on the aedeagal theca, which, as in C. nervosus (L.), is
variable. Specimens with intermediate length of this spine from the Vogelsberg were described
as C. haupti hassicus Nikusch, 1976, but are also found in other regions of the Mittelgebirge. Hence,
all three morphs are treated here as a single species.
B VI – E VIII; nymph, 1 gen.
In Germany the more common and widespread form with short aedeagal spine (haupti
morph) occurs in the Bavarian Alps, the Rhön Mountains, Black Forest, Vogelsberg,
Meißner, Solling, Harz Mountains, Thuringian Forest and the Erzgebirge, between 400
and 1200 m a.s.l. German specimens with an elongate aedeagal spine (beieri morph) have
been found only in the Harz Mountains and Black Forest. This is mainly a species of open,
montane coniferous forests also found along the edges of bogs, meadows and clearings.
In central Europe it is confined to the Alps as well as rather cool parts of the Mittelgebirge.
Adults are often swept from spruce but also from pine and deciduous shrubs.
Bittner & Remane (1977), Haas (1975), Nikusch (1976), Reimer (1992), Remane & Wachmann (1993), Schie-
menz (1975, 1987), HN

Cixius cambricus China, 1935

C. borussicus Wagner, 1939 was described from East Prussia and C. austriacus Wagner, 1939 from
South Tyrol (Italy). Later, however, Wagner considered them both as conspecific with C. cambri-
cus China (see Christandl-Peskoller & Janetschek 1976; Nast 1987; Ossiannilsson 1978). On the
other hand slight differences in the morphology of the aedeagus and the anal tube as well as
diverging habitat preferences may indicate the existence of distinct species.
M V – M IX; nymph, 1 gen.
Described from the Scottish Highlands, but found in Germany mainly in xerothermic
sites with scattered shrubs in the Saale-Unstrut region (between Halberstadt, Sonders-
hausen, Jena, Naumburg and Eisleben), in the Elbe valley near Meißen and in the Kaiser-
stuhl. All these localities are below 400 m a.s.l. On the other hand there are several sites in
the subalpine belt of the Bavarian and Allgäu Alps between 1800 and 2200 m a.s.l. Re-
ported from alpine locations between 1750 and 2430 m a.s.l. in North Tyrol (Austria) and
from xerothermic hillsides on Quercus, Acer, Carpinus and Rhododendron in Ukraine.
MTD, Schiemenz (1987), ZIMH, HN; see also Christandl-Peskoller & Janetschek (1976), Logvinenko (1975)

Cixius stigmaticus (Germar, 1818)

M V – M VII, according to Schiemenz (1987) until B IX; nymph, 1 gen.
In the interior and along the margins of deciduous forests in moderately moist to moderate-
ly dry sites. Adults were swept from Betula, Alnus, Salix, Quercus, Rubus and other woody
plants. Widespread in the lowlands of Germany, although scattered and usually found in
26 Review of species

low numbers only; highest localities are at 550 m a.s.l. in the Mittelgebirge. It is likely that the
species has declined in abundance since most records date back before 1970.
Fischer (1972), Post-Plangg & Hoffmann (1982), Schiemenz (1987), Wagner (1935, 1939a), HN

Cixius dubius W. Wagner, 1939

B V – E VI, occasionally until E VII; nymph, 1 gen.
Adults usually in the shrub and canopy layer of dense to open forests of beech, oak and
pine in damp to moderately dry sites (on Corylus, Quercus and others). There are only
scattered records from the middle parts of Germany, notably between the Danube and the
northern edge of the Mittelgebirge, notably Thuringia (locus typicus: Kleinfurra near Nord-
hausen), southern Niedersachsen, middle Hessen and Unterfranken, but also from near
Bitterfeld, Quedlinburg, Cologne, Bonn and Tübingen, up to at least 500 m a.s.l. The spe-
cies is not reported from the Netherlands, France and Switzerland, but from Belgium, the
Czech Republic, Austria and northern Italy. The Polish records are uncertain.
Reimer (1992), Remane & Fröhlich (1994b), Schiemenz (1987), Schwoerbel (1957), Wagner (1939a, 1951a),
Nickel (1994), HN

Cixius similis Kirschbaum, 1868

M V – B VIII; nymph, 1 gen.
In raised and intermediate bogs, occasionally also on peaty and acidic substrates of
other sites (e.g. moist heathlands, open forests and clearings). Adults are usually swept
from low-growing woody plants, preferentially Betula pubescens, Pinus, Vaccinium myr-
tillus and V. uliginosum. Occurs rather localized in Germany, mainly in bog areas of the
north German plain, in rather cool parts of the Mittelgebirge (notably Harz Mountains,
Thuringian Forest, Erzgebirge, Rhön Mountains, Spessart), as well as in the foothills of
the Alps, up to at least 1000 m a.s.l.
Bittner & Remane (1977), Fischer (1972), Krause et al. (1992), Niedringhaus & Olthoff (1993), Reimer (1992),
Schiemenz (1987), Strübing (1955), Wagner (1939a, 1951a), Nickel (2002), HN

Tachycixius pilosus (Olivier, 1791)

E IV – M VII; nymph, 1 gen.
Adults on various deciduous woody plants (Prunus spinosa, Quercus, Betula, Populus
and others) in damp to dry sites, in grassland with scattered shrubs, as well as in open
forests. Widespread and rather common at lower elevations in Germany (up to at least
500 m a.s.l.), but apparently absent from large parts of the Mittelgebirge and the Alps.
Fischer (1972), Niedringhaus (1991, 1997), Remane (1987), Rombach (1999b), Schiemenz (1987), Schönitzer
& Oesterling (1998a), Schwoerbel (1957), Wagner (1935, 1939a), HN

Trigonocranus emmeae Fieber, 1876

E V – E VII; nymph, 1 gen.
 Cixiidae Spinola, 1839 27

Life habits poorly known. According to very few existing data, nymphs and brachypter-
ous adults (which are unpigmented, with a reduced number of ommatidia) live in the
top soil and leaf litter in at least moderately warm sites with incomplete vegetation
cover, feeding on roots of shrubs. However, most field records comprise macropterous
and pigmented individuals on dispersal flight caught in the sweep net or in Malaise
traps. The species is probably more widespread in Germany but has been recorded only
northward to the edge of the Mittelgebirge, with only three records of autochthonous
populations, all from sun-exposed embankments or hillsides with scattered shrubs on
damp to moderately dry soils: Warburg, 270 m a.s.l., 24.V.1985, 1 †, 4 ‡‡ (brach.),
3 nymphs, and 11.VII.1986, 1 † (brach.), Wesseln, Steinberg (near Hildesheim), 150 m
a.s.l., 1997, altogether 2 ††, 3 ‡‡ (macr.), and Herne, 80 m a.s.l., 02.VII.2001, 2 †† (macr.).
All records except the first, which is from a soil sample (2-5 cm depth, extraction after
Kempson), are from pitfall traps. Also single macropterous ‡‡ near Selters (Vogelsberg),
Bacharach (middle Rhine), Merseburg, Leipzig (east German coal mining region) and
Garmisch-Partenkirchen (Bavarian Alps, 900 m a.s.l.). Otherwise the species is known
only from a few specimens (mostly macropterous) from the southern half of the Euro-
pean deciduous forest zone (northeastern Spain, France, England, northern Italy, Swit-
zerland, Austria, Croatia, Slovakia, Bulgaria, southern Russia).
Emmrich (2001), Funke & Witsack (1998), Coll. R. Köhler, Remane & Fröhlich (1994b), Coll. M. Sayer, Ni-
ckel & Voith, unpublished data), Nickel (2001), HN; also Emeljanov et al. (2002), Gnezdilov (2000)

Myndus musivus (Germar, 1825)

B VI – M VIII; nymph, 1 gen.
Adults are found on temporarily flooded sand and gravel banks along unchanneled
rivers on willows, notably Salix viminalis and S. triandra. In Germany, this species occurs
at the western and northern edge of the range, notably in the valleys of the Elbe and
Rhine and their larger tributaries (near Dresden, Meißen, Leipzig, Merseburg, Halle,
Stendal, Aschaffenburg, Wiesbaden and Bonn), as well as near Rinteln an der Weser. All
localities are below 150 m a.s.l. It is likely to have suffered a dramatic decline due to
canalization of rivers, since most records are from before 1950.
Haupt (1924, 1935), Kirschbaum (1868), Schiemenz (1987), Wagner (1951a, and unpublished data), HN

Pentastiridius beieri (W. Wagner, 1970)

So far only B VI – E VI, according to Nast (1977) in Poland and Ukraine E V – E VII;
nymph, 1 gen.
On willows on almost bare gravel banks of unchanneled mountain rivers. Has also
been taken from Myricaria germanica in Ukraine and from Alnus in the Engadine (Swit-
zerland). This species is only known from the Alps, Beskids and Carpathians. In Ger-
many there are only two records from the upper course of the Isar near Vorderriß, 800 m
a.s.l.: 23.VI.1997, 1 †, 1 ‡, as well as 04.VI.1998, 11 ††, 7 ‡‡, on tumbled shrubs of Salix
purpurea and S. eleagnos. Very recently populations in eastern France which have been
suspected to transmit stolbur phytoplasma in sugar beet fields have been provisionally
28 Review of species

assigned to this species. But regarding its specific habitat requirements, it is likely that
another species is involved.
Nickel (1999a); see also Gatineau et al. (2001), Günthart (1987a), Nast (1977), Remane & Fröhlich (1994b)

Pentastiridius leporinus (Linnaeus, 1761)

E V – B VIII, occasionally until B IX; nymph, 1 gen.
In moderately wet to wet sites which are at most rarely flooded, mainly in fens, along
rivers, as well as in ruderal and brackish habitats. Adults usually on Phragmites austra-
lis, according to published data also on Scirpus, Carex and Eriophorum. In Germany this
species occurs only locally, with strongholds in the foothills of the Alps and in the upper
Rhine plain; further scattered records are from the Wetterau, the upper Main valley, the
island of Amrum, Holstein, the rain shadow area to the east of the Harz Mountains and
the Oberlausitz, up to at least 750 m a.s.l. Some of the older literature records from
southern Germany may refer to P. beieri (W.Wg.).
Fischer (1972), Fröhlich (1996a), Schiemenz (1987), Wagner (1935, 1939a, 1970), Wagner & Wagner (1938),
Walter (pers. comm.), HN

Reptalus panzeri (P. Löw, 1883)

M VI – B VIII; nymph, 1 gen.
In xerothermic sites with scattered shrubs, usually on sunny hillsides or on plateaus.
Adults are often found on Rosa and Prunus spinosa, but also on other woody plants
(Clematis, Salix, Crataegus, Pinus and others). In Germany restricted to rather warm re-
gions between the Danube and a line running from Koblenz to Frankfurt am Main,
Fulda, Gotha, Magdeburg, Leipzig and Dresden, notably in viticultural areas (Saale-
Unstrut region, Elbe valley around Meißen, Unterfranken, upper and middle Rhine
valley, valleys of the Moselle, Nahe and Neckar, and southern Baden). The highest lo-
calities are at 500 m a.s.l. only.
Maixner (pers. comm.), Reimer (1992), Schiemenz (1987), Schönitzer & Oesterling (1998a), Schwoerbel (1957),
SMNS, Wagner (1939a, 1951a), HN

Reptalus quinquecostatus (Dufour, 1833)

Specimens published under this name show considerable morphological variation and may
belong to several distinct species; see, for instance, Dlabola (1954), Logvinenko (1975), Musil
(1956), Mityaev (1971), Holzinger et al. (2003).
So far only B VII – B VIII; nymph, 1 gen.
Reported mainly from southern Europe and southwestern parts of Asia. In Germany there
are only 6 records from the upper Rhine plain and Franconia: Speyer, Bruchgraben, 100 m
a.s.l., July 1989, in Malaise trap catches; Staffelbach, 240 m a.s.l., 11.VII.1949, 1 †; Coburg,
Krebsgrund, 300 m a.s.l., 15.VII.1950, 1 †, 1 ‡; Kitzingen, Klosterforst, 09.VII.1994, 1 ‡, on
Salix cinerea; Gerolzhofen, Eichelmannsee, 08.VIII.1994, 1 ‡, on Salix cinerea; Erlangen,
Exerzierplatz, 280 m a.s.l., 23.VII.2001, 1 †, 1 ‡. Apparently the species prefers moist sites
 Asiracinae Motschulsky, 1863 29

with scattered shrubs and tall herbs on sandy or loamy substrates in warm situations. In
Ukraine and Slovakia it is considered to be mesophilous and was collected on tall herbs
and low-growing shrubs, notably Melilotus, Carduus, Cirsium, Caragana, Cerasus and Spi-
raea. Has also been recorded in salt marshes in Hungary.
Remane (1995), Nickel (1999b), HN; see also Logvinenko (1975)

Hyalesthes obsoletus Signoret, 1865

E V – M VIII; nymph, 1 gen.
This species is restricted in central Europe to rather warm sites mainly in viticultural
areas. It lives in sun-exposed vineyard situations on abandoned land, along walls and
sunny embankments, also in old gardens and in disturbed patches in dry grassland,
usually on stony or crumby soils. In Germany it is only known from the valleys of the
Rhine (between Lörrach and Koblenz), Neckar, Main, Nahe and Moselle, as well as the
Saale-Unstrut region near Naumburg and Halle, up to 350 m a.s.l. From the Moselle
valley Convolvulus arvensis and, less frequently, Ranunculus bulbosus, Urtica dioica, Sene-
cio erucifolius and Artemisia vulgaris have been reported as the main food plants of nymphs
and adults. Adults were also recorded on Solanum nigrum, Galeopsis angustifolia, Daucus
carota, Chrysanthemum vulgare and Calystegia sepium. Woody plants such as Vitis and
Salix were only found to hold single specimens. The species is known as a potential
vector of grapevine yellows (also in southwestern Germany) and is reported from south-
ern and eastern Europe as a vector of Stolbur disease of tomatoes and potatoes.
Hoch & Remane (1985), Maixner (1994), Remane (1987), Schiemenz (1987), Schwoerbel (1957), Sergel (1984a,
1986), Wagner (1939a), Weber (pers. comm.), Weber & Maixner (1998), HN; see also Brčák (1979), Sforza et
al. (1999) (see this paper also for nymphal morphology)

4.1.2 Delphacidae Leach, 1815

4.1.2.1 Asiracinae Motschulsky, 1863

Asiraca clavicornis (Fabricius, 1794)

Recently Dolling (1996) suggested that Asiraca clavicornis (F.) might be a junior synonym of Ci-
mex aequinoctialis Scopoli, 1763 and Cicada quadristriata Gmelin, 1790, but this was dismissed by
Wilson & Asche (1998). On the one hand it is not quite clear to which species these names refer,
on the other hand synonymisation would cause far-reaching nomenclatural changes concern-
ing several supraspecific taxa.
Adults I – XII, mainly B VIII – M VI; adult, 1 gen.
This species is found in sunny, moderately dry to dry, usually disturbed sites. It is usu-
ally taken in low numbers only, but is widespread in the warmer regions in the south
and east of Germany (mainly in the valleys of Rhine, Main and Danube, the rain shad-
ow area to the east of the Harz Mountains and Thuringian Basin), where it is found in
abandoned vineyards, ruderal sites, along waysides, etc., preferentially near patches of
both bare ground and tall herbs or shrubs. The food plants are poorly known; adults are
usually found in the herb layer as well as on shrubs. Hence, the species is probably
30 Review of species

polyphagous. The northern edge of its range runs through the Mittelgebirge, approxi-
mately following a line from Cologne to Schlüchtern, Eschwege, Sondershausen, Halle
and Berlin. The highest localities are at 400 m a.s.l. Older records, some of which are not
verified, are from northwestern Poland, Mecklenburg and Lüneburg. However, the spe-
cies is strikingly coloured and easy to identify, thus, it is likely to have declined in cen-
tral Europe, as has been presumed for England.
Frommer (1996), Krause & Emmrich (1996), Nikusch (1976), Post-Plangg & Hoffmann (1982), Schiemenz
(1987), Schwoerbel (1957), Wagner (1939a, 1941a, 1951a), HN; see also Kirby (1992), Nast (1976a)

4.1.2.2 Kelisiinae W. Wagner, 1963

Kelisia guttula (Germar, 1818)

Most published records of this taxon are in need of revision due to possible confusion with K.
sima Rib. and K. sabulicola W.Wg. (see Remane & Jung 1995); this is particularly true for all coast-
al sites of north Germany as well as for most other countries outside central Europe.
B VI – E X; egg, 1 gen.
On Carex flacca, perhaps also on other low-growing sedges, in temporarily dry to mod-
erately wet sites usually on nutrient-poor soils, mainly calcareous grassland, montane
meadows and fens. Widespread but local in Germany with strongholds in the foothills
of the Alps (up to at least 1200 m a.s.l.) and the limestone areas of the Mittelgebirge
region; there are also a few confirmed localities in the north German plain.
Fischer (1972), Müller (1978), Remane (1962, and pers. comm.), Remane & Jung (1995), Rombach (1999b),
Schiemenz (1987), Nickel & Achtziger (1999), HN; see also Bieman & Booij (1984)

Kelisia sima Ribaut, 1934

The identity of this species, which was originally described as Kelisia pascuorum var. sima Ribaut,
1934, was clarified by Remane & Jung (1995). Without any further comments it was treated as a
distinct species by Wagner & Franz (1961). Moreover, recent field studies showed that the habi-
tats and host plants are different from those of K. guttula (Germ.), although both species may
occur syntopically.
B VII – B X; egg, 1 gen.
In wet to temporarily wet, sunny to moderately shady sites on basic to acidic substrates,
mainly in spring mires and fens, as well as on fairly overgrown gravel banks of alpine
rivers, sometimes also in temporarily wet forest glades and clearings. Sedges of the Carex
flava group are the host plants. Most German locations are in the Alps and their foothills
(valleys of the Iller, Lech and Isar, as well as the Chiemgau and Berchtesgaden Alps); there
are also scattered records from the Mittelgebirge (southern Black Forest, 1100 m a.s.l., Rhön
Mountains, Main valley, middle Hessen, Thuringian Forest, Erzgebirge) and on the island
of Borkum. The distribution is poorly known because many older records of K. guttula
(Germ.) are in need of revision. Outside Germany the species is known from the French
Pyrenees, Trentino (Italy), Styria (Austria) and the Baltic Sea island of Gotland.
Remane & Jung (1995) (see this paper also for identification), Walter (1998), ZIMH, Nickel & Achtziger
(1999), Nickel & Sander (1996), HN; see also Wagner & Franz (1961)
 Kelisiinae W. Wagner, 1963 31

Kelisia irregulata Haupt, 1935

E VII – E X; egg, 1 gen.
On Carex flacca in wet to temporarily dry, sunny to slightly shady, usually basic and nutri-
ent-poor sites, mainly calcareous grassland, montane meadows, calcareous small-sedge fens,
occasionally also in open forests (coppice-with-standards, waysides, clearings, etc.). Wide-
spread in southern and middle parts of Germany and fairly common in areas of Jurassic
and Triassic limestone, notably in the Alps and their foothills, on the Swabian and Franco-
nian Jura (locus typicus: Blaubeuren near Ulm), in Mainfranken and Thuringia; usually found
in intermediate frequency on the host. At present, localities at the edge of the range include
Bad Münstereifel, Osnabrück, Warburg, Göttingen, Nordhausen and Jena. Occurs up to at
least 1200 m a.s.l. Otherwise reported from Slovakia, Austria and France only.
Nikusch (1976), Reimer (1992), Rombach (1999a), Schiemenz (1987), Schwoerbel (1957), Trümbach (1959),
Nickel & Achtziger (1999), Nickel (1994), HN; see also Jansky & Okali (1993), Remane & della Giustina
(1991), Remane & Guglielmino (2002), Wagner & Franz (1961)

Kelisia haupti W. Wagner, 1939

This delphacid was first collected by Haupt (1935) near Freyburg an der Unstrut (locus typicus)
and Karlstadt am Main, but erroneously identified as K. melanops Fieber, 1878. Later Wagner
(1939a) revised this material and described it as a new species. Older literature records have to
be considered doubtful due to the rather complicated situation within the genus in general and
due to the recent splitting of the Kelisia haupti group by Remane & Jung (1995).
B VII – E X; egg, 1 gen.
Monophagous on Carex humilis on sparsely vegetated dry grassland, as well as in open
pine and oak forests, usually on xerothermic hillsides and plateaus on limestone or
gypsum, occasionally also on more acidic substrates (recorded up to at most 500 m a.s.l.).
In Germany only scattered, although locally dominant in suitable sites. Known from
warmer regions of Thuringia (Kyffhäuser, Thuringian Basin and surrounding Triassic
limestone plates, notably in the basins of Saale, Gera, Werra and Wipper), of Saxony-
Anhalt (northeastern edge of the Harz Mountains, Unstrut valley) and the Main-Saale-
Tauber area (between Tauberbischofsheim, Würzburg, Karlstadt, Bad Kissingen and
Bad Neustadt), also found in southern Niedersachsen (southern edge of the Harz Moun-
tains near Bad Sachsa).
Müller (1978), Remane & Jung (1995), Schiemenz (1969, 1987), SMNS, Nickel (1994), HN

Kelisia hagemini Remane & Jung, 1995

This species and the following have only recently been split from K. haupti W.Wg. after morpho-
metric differences of the male genitalia (Remane & Jung 1995). In Germany all three species
apparently occur parapatrically.
M VIII – E VIII, in northern Italy E IX; egg(?), 1 gen.
Also on Carex humilis and until very recently known only from high mountains of south-
ern Europe (Pyrenees, southern Alps and Mt. Olymp/Greece) as well as Carinthia (Aus-
tria). In Germany it was recorded from the Swabian Jura and the Lake Constance Basin:
32 Review of species

Fridingen, 800 m a.s.l., 21.VIII.1998, 2 ††, 4 ‡‡, and Sipplingen, 450 m a.s.l., 20.VIII.1998,
1 †, in both sites on xerothermic calcareous cliff-tops with scattered trees.
Nickel (1999b), HN; see also Remane & Jung (1995), Holzinger (1999b)

Kelisia halpina Remane & Jung, 1995

See taxonomic remark above.
In Austria and the Czech Republic B VIII – E IX (Remane & Jung 1995); egg(?), 1 gen.
This species is only known from a few localities in Niederösterreich, Carinthia, North
Tyrol (Austria), Südtirol (Italy), Graubünden (Switzerland) and Moravia (Czech Re-
public). It lives in xerothermic, although temporarily seepy, usually basic sites, on Carex
humilis. There is a recent record from the Bavarian Alps: Vorderriß, 850-1000 m a.s.l.,
22.VIII.1998, 12 ††, 12 ‡‡, on steep, south-facing calcareous hillsides with scattered
stands of pine.
Nickel (1999b), HN; see also Remane & Jung (1995), Holzinger (1999b)

Kelisia vittipennis (J. Sahlberg, 1868)

M VII – B XI, occasionally until V; egg, 1 gen.
Mainly in raised and intermediate bogs where it is often among the dominant species
during mid and late summer; less commonly found in peaty meadows and calcareous
sedge fens. Host plants are cottongrasses (Eriophorum vaginatum, E. angustifolium, E. latifo-
lium). Records published from dry calcareous grassland in Britain are likely to refer to
other species, perhaps K. irregulata Hpt. Rather widespread in Germany but local, mainly
in the remnant bog areas of the north German plain, as well as in the Mittelgebirge, the
Alps and their foothills. Occurs up to at least 1900 m a.s.l. in North Tyrol (Austria) and
Switzerland, although highest known localities in Germany are only at 1000 m a.s.l.
Nikusch (1976), Remane (1958), Schiemenz (1987), Wagner (1935, 1939a), Nickel & Achtziger (1999), Nickel
(2002), HN; see also Günthart (1987a), Leising (1977)

Kelisia punctulum (Kirschbaum, 1868)

M VII – M XI; egg, 1 gen.
In tall-sedge swamps in moderately wet to temporarily flooded, moderately eutrophic,
sunny to shady sites (fens, fen woods, also wet meadows and along ditches), mainly
(perhaps exclusively) on Carex acutiformis. Widespread in southern and middle parts of
Germany and locally dominant in suitable sites. There are also scattered records from
the north German plain (in Holstein extending northward at least to Itzehoe, Bad Ol-
desloe and Bad Schwartau), from Ireland, England and eastern Poland, but not from
Denmark and Scandinavia. Highest localities are at 550 m a.s.l. only; single individuals
collected at higher altitudes are probably mostly vagrants.
Heller (1987a), Marchand (1953), Nikusch (1976), Reimer (1992), Remane (1958), Schiemenz (1987), Wagner
(1935), Nickel & Achtziger (1999), Nickel & Remane (1996), HN; see also Le Quesne (1960), Nast (1976a,
1987)
 Kelisiinae W. Wagner, 1963 33

Kelisia guttulifera (Kirschbaum, 1868)

According to Wagner (1939a) K. fallax Ribaut, 1934, described from southern France, is conspe-
cific with K. guttulifera (Kbm.), although Wagner did not see Ribaut’s type material. However, in
central Europe there are two ecologically distinct forms (see below), which may turn out to
show also morphological differences after a more thorough study, as was recently demonstrated
for Kelisia guttula (Germ.) and K. sima Rib. by Remane & Jung (1995) (see above).
M VII – M X; at least ‡‡ frequently until IV/V; ad.(?), 1 gen.
Usually in beech and oak forests (often along roads and ditches) in moist or seepy sites
on Carex sylvatica and C. remota, occasionally in alder fen woods on C. elongata and in
open wetlands (spring mires, wet meadows, sand pits, etc.) on Carex vulpina, C. distans,
C. paniculata and perhaps other species. In Germany at least scattered and presumably
often overlooked; mainly in middle and southern parts, although apparently absent or
rare to the south of the Danube and in the northwest German plain. The northern edge
of the range runs from Wales through England, the Netherlands, northern Germany
(extending at least to Höxter, Braunschweig, Uelzen, Rendsburg, Rostock and Neustre-
litz) and Poland to Lithuania. Highest localities in the Mittelgebirge are at 500 m a.s.l.
The locus typicus is near Wiesbaden.
Schiemenz (1987), Trümbach (1959), Wagner (1935, 1939a), ZIMH, Nickel (1994), HN

Kelisia praecox Haupt, 1935

M VIII – B VI; adult, 1 gen.
In temporarily wet sites on Carex brizoides, usually in open forests, clearings and forest
meadows, also on other sedge species in straw meadows of the foothills of the Alps.
Widespread in the eastern half of Germany up to 700 m a.s.l., but rather scattered, al-
though locally abundant and probably under-recorded. Localities at the western edge
of the range are near Biberach, Tübingen, Aschaffenburg, Gießen, Halle and Neubran-
denburg. Not reported from Fennoscandia, Denmark, the Netherlands, Belgium, Swit-
zerland and Italy, but from two isolated localities in the French Rhone Alps.
Fischer (1972), Remane & Fröhlich (1994b), Schiemenz (1987), Schwoerbel (1957), SMNS, Trümbach (1959),
Wagner (1951a), HN; see also della Giustina & Remane (1999), Nast (1987)

Kelisia pallidula (Boheman, 1847)

M VII – E X; egg, 1 gen.
Monophagous on Carex panicea in temporarily wet to peaty, basic to acidic, usually sun-
ny sites, preferentially spring mires, small-sedge fens, peaty meadows and intermedi-
ate bogs. Widespread in Germany, though sporadic, fairly common in the foothills of
the Alps, but in rather small and scattered populations in most regions of the Mittelge-
birge and the north German plain. The highest localities are at 1100 m a.s.l.
Niedringhaus & Olthoff (1993), Nikusch (1976a), Reimer (1992), Schiemenz (1987), SMNS, Wagner (1935,
1939a), Nickel & Achtziger (1999), Nickel & Remane (1996), HN
34 Review of species

Kelisia nervosa Vilbaste, 1972

Described from Lithuania, but strongly resembling K. confusa Lnv. (see below), which occurs
mainly in southeastern Europe.Vilbaste (1972) did not discuss the relationship towards the lat-
ter species and was apparently not aware of it. Thus, the relationship between K. nervosa Vilb.
and K. confusa Lnv. is in need of revision; they are probably conspecific.
In Lithuania M VIII – E VIII (Vilbaste 1974); egg(?), 1 gen.
This taxon is altogether known only from 6 localities in Lithuania, eastern Poland and east-
ern Germany. The two German records are both from the margins of intermediate bogs in
the vicinity of Lübben and comprised fairly large populations: Halbe, end of VIII.1995, and
Krausnick, 29.VIII.2001. The latter population lived in tussocks of Carex elata.
Remane (1995), HN; see also Nast (1976a), Vilbaste (1972)

Kelisia confusa Linnavuori, 1957

See taxonomic remarks above.
This appears to be an essentially southeast European species living in swampy and
moderately eutrophic sites among yet undetermined tall sedges (Carex sp.). Recently it
was recorded for the first time in Styria (Austria) and in the south of the upper Rhine
plain: Rust, 160 m a.s.l., 03.VIII.1997, 1 † (brach.), and Endingen, 200 m a.s.l., 18.VIII.1998,
1 † (macr.), all in tall-sedge swamps, perhaps on Carex acutiformis.
HN; see also Holzinger et al. (1997), Nast (1987)

Kelisia minima Ribaut, 1934

So far M VII – B IX, in France according to Ribaut (1934) until XI; egg(?), 1 gen.
Apparently this is a very rare species, which is only known from altogether 8 localities:
three older records from northern and southern France, „sur Cyperus longus“, another
one from Switzerland (Valais, Dugny, in a spring mire on a hillside, 600 m a.s.l.) and
4 from Germany: island of Hiddensee, Vitte, 03.IX.1965, 1 †, in brackish grassland; Wet-
terau, Münzenberg, July and August 1966 and 1967; eastern Niedersachsen, Schreyahn
near Lüchow, 21.VII.1994, 5 ††, 1 ‡, and Thuringian Basin near Erfurt, Stotternheim,
13.VII.1997, 1 †, 3 nymphs, the latter two in inland saltmarshes in tussocks of Carex
distans. The Hiddensee record was already published as K. guttulifera (Kbm.).
Emmrich (pers. comm.), Remane & Fröhlich (1994b), Nickel (1997), HN; see also Schiemenz (1987)

Kelisia ribauti W. Wagner, 1938

Belongs to a species group which is subject to diverging interpretations of species distinction
and nomenclature among various authors (Asche & Remane 1982). In particular there are pop-
ulations living in rather montane or boreal habitats, whereas others are widespread in the Med-
iterranean basin. However, morphological distinction is difficult.
B VII – E X; egg, 1 gen.
In peaty meadows, intermediate bogs and spring mires, also along the edge of raised
bogs (in the lagg zone, along ditches, in peat diggings, etc.). Probably lives on sedges
 Kelisiinae W. Wagner, 1963 35

(notably Carex nigra), although often found in patches with Eriophorum angustifolium or
E. latifolium. Is also reported from Carex elongata. Status and distribution in Germany
are very similar as in K. vittipennis (J. Shlb.), with which it often co-occurs, although
usually in lower abundance. Found up to 1900 m a.s.l. in North Tyrol (Austria) and the
Engadine (Switzerland), up to at least 1200 m a.s.l. in the Bavarian and Allgäu Alps and
the southern Black Forest.
Marchand (1953), Nikusch (1976a), Reimer (1992), Schiemenz (1987), Strübing (1955), Wagner (1939a, 1951a),
Wagner & Franz (1961), Nickel & Achtziger (1999), Nickel (1997, 2002), HN; see also Günthart (1987a),
Leising (1977)

Kelisia sabulicola W. Wagner, 1952

Most older records of this species were published as K. guttula (Germ.) until Wagner (1938a,
1939a, 1952) pointed out that there are morphologically and ecologically different populations.
Some authors (e.g. Nast 1987) consider K. sabulicola W.Wg. a junior synonym of K. pannonica
Matsumura, 1910, which was described only after 2 ‡‡ from Hungary and Germany. This book
follows the interpretation of Ossiannilsson (1978) and Remane & Fröhlich (1994a), who suggest
to retain Wagner’s unambiguous name.
Adults I – XII, mainly E VI – V; adult, 1 gen.
Monophagous on Carex arenaria on sand dunes near the coasts of the North and Baltic
Sea, as well as inland in open sandy grassland of the north German plain. Most records
are from the islands (Heligoland, East and North Frisian Islands, Hiddensee, Rügen);
locally also found inland, southward at least to Oldenburg, Dömitz, Magdeburg, Des-
sau, Potsdam and Dresden; all sites are below 150 m a.s.l. Has been reported from the
region of the Mittelgebirge, but these specimens should be revised. The locus typicus is
Horst an der Seeve near Hamburg.
Niedringhaus (1991, 1994), Remane (1958), Schaefer (1973), Schiemenz (1987), Wagner (1952a), HN

Kelisia monoceros Ribaut, 1934

M VI – E X; egg, 1 gen.
Largely confined in central Europe to the Alps and their foothills, preferentially living
in various kinds of low-productivity grassland, often with Carex sempervirens, notably
on sunny slopes, in open pine forests and on riverine gravel banks. Much more scat-
tered to the north of it where usually found on Carex vulpina ssp. otrubae and C. muricata,
in inland salt marshes and disturbed, dry or temporarily dry sites on loamy or clayey
substrates, mainly in mining areas and pastures. Most German records are from south-
ern Bavaria (up to at least 1600 m a.s.l., but frequently found above 2000 m a.s.l. in the
southern Alps), in the lowlands extending northward to Mainz, Gießen, Halle, Leipzig,
Dresden and Görlitz. Apart from a single specimen from the botanical garden in Ha-
nover, there are no records from the north German plain, although the species is report-
ed from Scandinavia.
Fischer (1972), Marchand (1953), Remane (1987), Remane & Fröhlich (1994b), Schiemenz (1987), Nickel
(1999b), HN; see also Ossiannilsson (1978)
36 Review of species

Anakelisia fasciata (Kirschbaum, 1868)

Mainly B VII – X, ‡‡ occasionally until V; mainly egg, 1 gen., according to Witsack
(1985) overwintering takes place mainly in thermic quiescence (also a few ‡‡); see also
Lauterer (1984).
In tall-sedge swamps in wet to temporarily flooded, moderately eutrophic, sunny to
shady sites (fens, lake shores and fen woods; absent from mown meadows), on Carex
riparia, perhaps also on other species of tall sedges (C. acutiformis and C. acuta). Wide-
spread in Germany, although locally absent or overlooked (for instance, not recorded
from Schleswig-Holstein, Brandenburg, Northrhine-Westphalia, Bavaria to the south of
the Franconian Jura). Appears to be restricted to lowlands and river valleys; highest
localities are at 350 m a.s.l. only. The locus typicus is near Frankfurt am Main.
Heller (1987a), Hildebrandt et al. (1998), Remane & Fröhlich (1994b), Schiemenz (1987), Witsack (1985),
Nickel & Achtziger (1999), Nickel & Remane (1996), Nickel (1994), HN; see also Lauterer (1984)

Anakelisia perspicillata (Boheman, 1845)

E VII – E X; egg, 1 gen.
Rather eurytopic (although more xerophilous towards the north) in sunny and olig-
otrophic sites with short vegetation, ranging from dry to temporarily wet and from
basic to acidic conditions, mainly pastures, submontane meadows, dry forest margins
and calcareous small-sedge swamps. The main host plant on basic substrates is Carex
flacca, and C. pilulifera has been recorded in more acidic sites. Widespread in Germany
with strongholds in the Alps and their foothills (frequently up to at least 1050 m a.s.l. in
Bavaria, but extending to 1700 m a.s.l. in eastern Austria) and the limestone regions in
middle parts, but rather scarce in the north German plain.
Nikusch (1976), Reimer (1992), Schiemenz (1987), Wagner (1935), Nickel & Achtziger (1999), Nickel & Re-
mane (1996), Nickel (1994), HN; see also Wagner & Franz (1961)

4.1.2.3 Stenocraninae W. Wagner, 1963

Stenocranus longipennis (Curtis, 1837)

E VII – E X; mainly egg(?), 1 gen. (see also Strübing 1963).
This is a western European species occurring in rather scattered populations in large
tussocks of Carex paniculata in wet fenlands of western and northern parts of Germany.
From there it is known only from 5 localities: Friedrichsruh near Hamburg, Stolzenau
an der Weser, Denkershausen near Northeim, as well as Erlenbach and Fischbach (220 m
a.s.l.), both near Pirmasens. Otherwise it is only reported from Ireland, England, the
Netherlands, France and Spain.
Marchand (1952), Strübing (1963), Wagner (1947a), ZIMH, Nickel (1994), HN; see also Bieman (1993), Nast
(1987), Remane & Fröhlich (1994b)
 Stenocraninae W. Wagner, 1963 37

Stenocranus major (Kirschbaum, 1868)

Adults I – XII, mainly E VIII – M VI; adult, 1 gen. (see also Strübing 1963).
In moderately wet to temporarily flooded, sunny to slightly shaded, rather eutrophic
sites, often near standing or running water, also along ditches, in wet ruderal sites (sand
pits, mining areas, etc.) and in fen woods, rarely on mown meadows. Phalaris arundina-
cea is the host plant. Widespread and common in most parts of Germany; in autumn the
species is locally extremely abundant and may also be found in stands of Calamagrostis
epigejos (but apparently without breeding). Found in Schleswig-Holstein northward only
to Pinneberg, Kiel and Lübeck, although there are few records from the Baltic Sea is-
lands of Zealand (Denmark), Bornholm and Öland, and from Skane (all Sweden). High-
est localities are at 800 m a.s.l. in the Mittelgebirge and Bavarian Alps.
Fischer (1972), Niedringhaus (1997), Remane (1987), Schiemenz (1987), Schwoerbel (1957), Wagner (1939a,
1947b), Nickel & Achtziger (1999), Nickel (1997), HN

Stenocranus minutus (Fabricius, 1787)

Adults I – XII, mainly E VIII – E VI; adults overwinter in primary parapause before
oviposition, 1 gen. (Müller 1957, 1958; Witsack 1981).
On Dactylis glomerata and probably D. polygama in sunny to slightly shaded, moderately
dry to moist and moderately eutrophic sites, mainly pastures, meadows, field margins,
waysides and open forests; usually absent or only as a temporary stray in most conven-
tionally managed meadows. Widespread and common in Germany, particularly in mid-
dle and southern parts, although largely confined to locations below 600 m a.s.l.; single
specimens were recorded up to 1000 m a.s.l.
Achtziger & Nickel (1997), Fischer (1972), Müller (1978), Post-Plangg & Hoffmann (1982), Niedringhaus
(1991), Reimer (1992), Remane (1987), Schiemenz (1987), Schöpke (1996), Schwoerbel (1957), Wagner (1935,
1939a, 1951a), Nickel & Achtziger (1999), HN; see also May (1978)

Stenocranus fuscovittatus (Stål, 1858)

B VIII – B VII; adult, 1 gen.
In moderately wet to temporarily flooded, oligotrophic to moderately eutrophic sites
(mainly fens, spring mires, peaty meadows, etc.); host plants are various species of sedges
(Carex elata, C. paniculata, C. panicea, presumably also C. acutiformis, C. acuta and others).
The species is widespread and fairly common in the middle and southern parts of Ger-
many, but there are only scattered localities in the north German plain where it lives at
the edge of the range. It frequently occurs up to 600 m a.s.l. in the foothills of the Alps
(possible vagrants found up to 950 m a.s.l.), although only below 350 m a.s.l. in south-
ern Niedersachsen. Northernmost localities are near Stolzenau an der Weser, Mölln,
Rostock and Neustrelitz. Also reported from England and northern Poland; records
from the Netherlands, however, have turned out to be erroneous.
Heller (1987a), Kuntze (1937), Marchand (1953), Nikusch (1976), Schiemenz (1977, 1987), Wagner (1939a,
1951a), Nickel & Achtziger (1999), Nickel (1997), HN; see also Bieman (1993), Le Quesne (1960), Nast (1976a)
38 Review of species

4.1.2.4 Delphacinae Leach, 1815

Jassidaeus lugubris (Signoret, 1865)

Adults I – XII, mainly E VIII – E V; adult, 1 gen.
In sparsely vegetated dry grassland in sunny, basic to acidic sites, preferentially near
the base of fine-leaved species of fescue (Festuca ovina group), perhaps also on Stipa
capillata. Rather localized in Germany and on the northern edge of the range, mainly
confined to the warmer regions of southern and eastern parts, but perhaps overlooked.
Northernmost records are from near Bad Neuenahr, Warburg, Wolfenbüttel, Sonder-
shausen, Halberstadt, Parchim and Waren an der Müritz; the highest localities are at
550 m a.s.l.
Kuntze (1937), Nikusch (1976), Reimer (1992), Remane (pers. comm.), Schiemenz (1987), SMNS, Wagner
(1939a), Walter (pers. comm.), Nickel & Remane (1996), Nickel (1997), HN

Megamelus notula (Germar, 1830)

B VI – E X, according to Schiemenz (1987) single ‡‡ until V; mainly egg, 1(?) gen.
Eurytopic in stands of tall sedges of various species in moderately wet to temporarily
flooded sites (from oligotrophic intermediate bogs to eutrophic meadow depressions,
even ditches within fields). Host plants are various species of Carex (C. acutiformis, C.
acuta, C. rostrata, C. nigra, C. disticha and others). Widespread in Germany and often
dominant in suitable sites, extending up to at least 1350 m a.s.l. in the Mittelgebirge and
the Bavarian Alps, up to 1500 m a.s.l. in North Tyrol (Austria).
Heller (1987a), Niedringhaus (1991), Nikusch (1976), Remane (1958), Schiemenz (1971a, 1975, 1987), Wag-
ner (1935, 1939a), Nickel & Achtziger (1999), HN and others

Conomelus anceps (Germar, 1821)

E VI – E X; egg, 1 gen. (see also Witsack 1985).
In moist to temporarily flooded, sunny to moderately shady sites (notably meadows
and pastures, fens, intermediate bogs, forest clearings, ruderal sites). Host plants are
various species of rushes (Juncus effusus, J. inflexus, J. articulatus and others). Widespread
in Germany and common in many places, found up to at least 1200 m a.s.l. in the Mittel-
gebirge and the Bavarian Alps. Maximum densities within tussocks of Juncus effusus in
England were over 8000 nymphs (1st and 2nd instar) or 1000 adults per sqm.
Bittner & Remane (1977), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Schiemenz (1971a, 1987),
Schwoerbel (1957), Wagner (1935), Witsack (1985), Nickel & Achtziger (1999), HN and others; see also
Rothschild (1964, 1966)

Conomelus lorifer Ribaut, 1948

This species occurs from Ukraine, the Balkans and Italy, to Poland, Germany and France. Two
endemic subspecies, C. l. calabricus Dlabola, 1979 and C. l. lorifer Ribaut, 1948, inhabit southern
and middle Italy. C. l. dehneli Nast, 1966 occurs in the remaining parts of the range. These sub-
 Delphacinae Leach, 1815 39

species show clinal morphological intergradation into each other (Remane & Asche 1979; Hoch
& Asche 1983; Hoch & Remane 1983).
B VI – M X; egg, 1 gen.
On Juncus effusus and perhaps other species of rushes, usually in seepy or temporarily
wet sites. Appears to favour disturbed habitats and sandy substrates such as sand
and gravel pits, mining areas and forest tracks, but has also been found in spring
mires and floodplain depressions. In Germany at the northern edge of the range and
apparently rather scarce; only found near Gardelegen, Nauen, Nordhausen, Halle,
Bad Salzungen, Bad Kissingen and Speyer, but probably under-recorded. All German
localities are situated below 300 m a.s.l., but the species was found up to 1400 m a.s.l.
in northern Italy.
Funke & Witsack (1998), Remane & Fröhlich (1994b), Sander (pers. comm.), Wagner (unpublished data), HN

Delphacinus mesomelas (Boheman, 1850)

M V – M IX; nymph, 1(?) gen.
In sunny, low-vegetated, dry to wet sites of low productivity. In the north German plain
on heaths and other dry and sandy habitats, to the south of it mainly on moderately dry
to wet, even peaty meadows usually at submontane altitudes, as well as on various
types of dry grassland. Host plants are fine-leaved species of fescue (mainly Festuca
ovina group and F. rubra) and perhaps additional grasses. Widespread in northern and
middle parts of Germany, but rather local; not recorded to the south of the Danube, but
occurring up to at least 1100 m a.s.l. in the southern Black Forest.
Bornholdt (1996), Kuntze (1937), Niedringhaus (1991), Reimer (1992), Rombach (1999b), Schaefer (1973),
Schiemenz (1969, 1971, 1987), Wagner (1935, 1951a), Nickel & Achtziger (1999), HN

Stiromella obliqua (W. Wagner, 1948)

So far B V – M VI, in Austria according to Moosbrugger (1946) until B VII; nymph, 1 gen.
The range of this species extends from Yakutia through Mongolia and Kazakhstan to
Europe, where it is only known from a few records from Estonia, Latvia, Ukraine, Aus-
tria (Niederösterreich and Vorarlberg, reported from Schoenus ferrugineus, up to at least
1000 m a.s.l.) and a single locality in Unterfranken near Schweinfurt (Grettstadt, 210 m
a.s.l., 15.V.1993, 17.VI.1994 and 06.V.2001, a large population in a temporarily dry cal-
careous sedge fen, perhaps on Schoenus nigricans).
Nickel & Remane (1996), HN; see also Anufriev & Emelyanov (1988), Moosbrugger (1946, and unpublished
data), Nast (1987), Vilbaste (1974), Wagner & Franz (1961)

Ditropis pteridis (Spinola, 1839)

M V – B VIII; nymph, 1 gen.
Monophagous on Pteridium aquilinum in moderately moist to moderately dry, usually
acidic and moderately shady sites, mainly in open forests and along their margins, oc-
casionally in clearings and pastures. Widespread in Germany and in intermediate to
40 Review of species

high frequency on the host, but not recorded to the south of the Danube, where it is
probably overlooked. Highest localities are at 1100 m a.s.l. in the southern Black Forest.
Reimer (1992), Schiemenz (1987), Wagner (1935, 1939a, 1951a), HN; see also Morcos (1953)

Eurysa lineata (Perris, 1857)

B V – E VII; nymph, 1 gen.
On various substrates in the grass layer of open, usually thermophilous deciduous and
coniferous forests (mainly oak and pine) and along their margins, as well as around hedg-
es, solitary trees and shrubs on rather dry grassland. Host plants are various grasses (Poa
nemoralis, Deschampsia flexuosa, Melica uniflora and others). In Germany mainly confined
to lower altitudes of middle and southern parts; the northernmost localities are near Bonn,
Warburg, Hildesheim and Malchin, the highest ones at 500 m a.s.l. To the north of these
there are only single records from southern Sweden, England and Ireland.
Achtziger (1991), Post-Plangg & Hoffmann (1982), Rombach (1999b), Schiemenz (1987), Wagner (1939a),
Nickel (1994), HN; see also Le Quesne (1960), Ossiannilsson (1978)

Eurysella brunnea (Melichar, 1896)

M V – E VII, occasionally until B IX; nymph, 1 gen.
Like E. lineata (Perr.) in open, grassy forests and near woody margins in dry grassland, vine-
yards, etc., but usually in higher-vegetated sites and rarely syntopic. The host plants are grass-
es, often Brachypodium pinnatum in basic sites, but the species is also found on siliceous sub-
strates, where Poa nemoralis is likely to be the host. Lives mainly near the base of its hosts and
is probably much under-recorded, but is usually collected in pitfall traps and with suction
samplers. In Germany living at the northern edge of the range and known mainly from rather
warm regions in the valleys of the Rhine and Main, and along their tributaries (Ahr, Nahe,
Lahn, Fränkische Saale), as well as in the rain shadow area to the east of the Harz Mountains;
highest localities are at 550 m a.s.l.; the northernmost records (which appear to be more or less
isolated) are from the vicinity of Bonn, Biedenkopf, Warburg, Göttingen, Quedlinburg and Jena.
Biedermann (pers. comm.), Kupka (1941), Reimer (1992), Remane & Fröhlich (1994b), Schiemenz (1987),
Wagner (1951a), HN; see also Asche (1994), Stewart (1997)

Eurysula lurida (Fieber, 1866)

B V – B IX; nymph, 2 gen.
In moderately dry to wet sites on small-reed, mainly in forest clearings, sand and coal
pits, along forest roads, etc., on Calamagrostis epigejos, locally also on C. canescens in fen
woods, fens and intermediate bogs. Single macropterous individuals, however, are fre-
quently found in other habitats. The species is widespread in Germany, but is found
only at lower altitudes (to at least 600 m a.s.l.). It is often overlooked due to its secretive
habits, although easily seen after direct search on the base of the host.
Fischer (1972), Niedringhaus & Olthoff (1993), Schiemenz (1987), Strübing (1955), Walter (pers. comm.),
Nickel (1999b), HN
 Delphacinae Leach, 1815 41

Eurybregma nigrolineata Scott, 1875

E IV – M VII; nymph, 1 gen.
In sunny, moderately dry to damp, occasionally moderately moist, usually disturbed
sites, which can be quickly colonized by macropterous individuals (along waysides,
railway lines and ditches, in abandoned fields and vineyards, low-input meadows,
etc., also inland salt marshes). Host plants are tall grasses (mainly Elymus repens, at
least locally also Holcus lanatus and Dactylis glomerata). In central Europe this species
is suspected to expand its range north- and westward since the 1950s and 60s, al-
though it is known from England since the 19th century (including brachypterous
specimens). From eastern Germany (Saale-Unstrut region) it is recorded since 1942, in
the west only since 1960. At present it is widespread and locally common in most
parts of the country, although not yet recorded to the south of the Danube. Localities
on the Baltic Sea island of Rügen, near Schnackenburg, Elmshorn and Bonn may be
near the northern and western edge of the range, since there are no records from the
Netherlands and Denmark. Largely confined to lowlands below 500 m a.s.l.; mac-
ropterous and also brachypterous individuals from higher altitudes and the lower
Ems area are perhaps vagrants. In France it is widespread at least in middle and north-
ern parts, in southern Sweden, Belgium, Switzerland and northern Italy it was recent-
ly found for the first time.
Achtziger (1991), Bornholdt & Remane (1993), Niedringhaus (1997), Reimer (1992), Remane (pers. comm.),
Remane & Fröhlich (1994b), Rombach (1999a), Schiemenz (1987), SMNS, Nickel & Achtziger (1999), Nickel
(1994), HN; see also Baugnée (1995), della Giustina & Remane (1999), Gillerfors (2002), Günthart (2000), Le
Quesne (1960), Nast (1972, 1987), Webb (pers. comm.)

Stiroma affinis Fieber, 1866

M V – E VIII; nymph, 1 gen.
In deciduous and coniferous forests, usually in moderately dry to moist sites on various
grasses (Brachypodium pinnatum, Calamagrostis arundinacea, Poa nemoralis, Deschampsia
flexuosa, Festuca altissima and others). Widespread in the lowlands of Germany and fair-
ly common, locally extending up to 1000 m a.s.l. in the Mittelgebirge and the foothills of
the Alps, up to 1500 m a.s.l. in the central Alps of Switzerland.
Fischer (1972), Niedringhaus (1997), Reimer (1992), Schiemenz (1987), Schwoerbel (1957), Wagner (1935,
1939a, 1951a), HN; see also Günthart (1987a)

Stiroma bicarinata (Herrich-Schäffer, 1835)

M V – E VIII; nymph, 1 gen.
Among rather tall grasses in moderately moist to wet sites, usually in low-input mead-
ows, pastures and abandoned fields (in the lowlands only in cool valley bottoms or in
the vicinity of shrubs and trees), in deciduous and coniferous forests (in more moist and
shady sites than St. affinis Fieb., but locally syntopic) and in subalpine pastures and tall
herb stands. Often in tussocks of Deschampsia cespitosa, but also on other grasses (e.g.
Holcus mollis, Glyceria maxima, Dactylis glomerata, Elymus repens, Arrhenatherum elatius).
42 Review of species

Widespread in Germany and fairly common, also in the higher parts of the Mittelgebirge
and the Alps, where frequently recorded up to at least 1800 m a.s.l.
Bittner & Remane (1977), Fischer (1972), Reimer (1992), Schiemenz (1987), Wagner (1935, 1939a), Nickel &
Achtziger (1999), HN and others

Metropis latifrons (Kirschbaum, 1868)

M V – B VII; nymph, 1 gen.
On fine-leaved species of fescue (Festuca ovina group) in xerothermic, basic as well as
acidic, sunny to moderately shady sites on hillsides and plateaus. In Germany at the
northeastern edge of the range and only known from a few localities in the valleys and
basins of the Rhine, Nahe, Main and Fränkische Saale, mostly on Triassic limestone,
aeolian sand and porphyry; the highest localities are at 550 m a.s.l. Furthermore, there
is a confirmed record from inland dunes of the lower Oder valley (Bielinek, Poland).
Asche et al. (1983), Remane (1987), Remane & Fröhlich (1994b), Wagner (1939a, 1951a), Nickel et al. (2002b),
HN; see also Haupt (1931)

Metropis inermis W. Wagner, 1939

So far B V – E VI, nymph, 1 gen.
This is apparently an eastern steppe species with isolated populations in eastern parts
of Germany, from where it is known only from 6 sites in eastern Bavaria (Naab valley
near Kallmünz and Krachenhausen, Laaber valley near Eilsbrunn and Undorf, Altmühl
valley near Meihern, all between 350 and 450 m a.s.l.) and eastern Thuringia: Steudnitz
near Jena. The latter record comprised a single macropterous † only, probably a stray
individual. The Bavarian records were from dry and sun-exposed hillsides and pla-
teaus of Jurassic limestone on a green subspecies of Festuca ovina. The Thuringian spec-
imen was collected in a dry grassland strongly influenced by phosphate immissions.
Achtziger (pers. comm.), Remane (1961b), Schiemenz (1987), Nickel et al. (2002b)

Achorotile albosignata (Dahlbom, 1850)

In Scandinavia VI – VIII (Ossiannilsson 1978), nymph(?), 1(?) gen.
The range of this species extends from the Altai through western Siberia to central and
western Europe, where it is known only from rather isolated localities. From Germany
there are only two records: a single † from the vicinity of Erlangen, which has been
taken in high summer in the mid 1950s „ … in einem trockenen Kiefernwaldstück auf
Waldgräsern …“. There is also a published record from Berlin dating from 1901: “In der
Umgebung von Berlin ist dieses Insekt häufig auf Festuca- und Agrostis-Arten, besonders
auf sterilen, sandigen Hängen.” Was also collected on species of Festuca in open pine
forests in the mountains of eastern and central Spain and southern France. Due to the
widespread occurrence of sensory pits in the nymphal stage in delphacids, some older
reports of this species from central Europe are likely to be based on misidentifications.
Matsumura (1906), Nast (1976a), Trümbach (1959) (Remane vid.); see also Remane & Fröhlich (1994b)
 Delphacinae Leach, 1815 43

Euconomelus lepidus (Boheman, 1847)

E VI – M X; egg, 1 gen.
Usually in sunny to moderately shady, temporarily or permanently wet, basic to acidic
sites with fresh to brackish water (meadow depressions, spring mires, inland salt marsh-
es, sand and coal pits). The main host plant is Eleocharis palustris s.l.; various reports
from Juncus must be considered as unreliable regarding host identification. Widespread
in Germany, but often occurring in small pockets only, although locally abundant; high-
est localities are at c. 700 m a.s.l. in Upper Bavaria, but found at 1000 m a.s.l. in the
Tyrolian Alps (Austria).
Niedringhaus (1991), Schiemenz (1987), Wagner (1935, 1939a), Nickel & Achtziger (1999), Nickel & Re-
mane (1996), HN

Delphax crassicornis (Panzer, 1796)

B VII – B IX; egg, 1 gen.
Monophagous on Phragmites australis, usually near the base of the stems. Apparently
prefers reeds which are not mown and flooded at most during winter; locally also in
open fen and floodplain forests and salt marshes. In Germany widespread though spo-
radic; most records are from the northeastern half (mainly to the east of a line from Sylt
to Hamburg, Braunschweig, Gotha and Würzburg), but also found in the Weser-Ems
area, the upper Rhine plain and the Allgäu (c. 800 m a.s.l.).
Fischer (1972), Remane (pers. comm.), Schiemenz (1987), Strübing (1960), HN

Delphax pulchellus (Curtis, 1833)

E VI – M IX; egg, 1 gen.
Monophagous on Phragmites australis, like the preceding species mainly near the ground
and in sites which are flooded not at all or only in winter. In exceptional cases, both
species are found together. The strongholds are in saltmarshes inland and along the
coast, as well as in low-input meadows and pastures with only scattered reed; appar-
ently absent from forests. Widespread in Germany but only localized, mainly near the
coast and in the north German plain, but also recorded from middle parts (although
restricted to valley bottoms) and the foothills of the Alps, where extending up to at least
700 m a.s.l.
Niedringhaus (1991), Schaefer (1973), Schiemenz (1987), Wagner (1939a), Nickel & Achtziger (1999), Nickel
& Remane (1996), Nickel (1997), HN

Euides basilinea (Germar, 1821)

Delphax speciosa Boheman, 1845
B V – E VIII; nymph, 1 - 2 gen., depending on spring weather conditions (Strübing 1960).
Monophagous on Phragmites australis. This is the most widespread among central Euro-
pean reed-dwelling Auchenorrhyncha species (both in geographical and ecological
44 Review of species

terms); along shores of lakes and ponds, in fen woods, fens and intermediate bogs, near
fresh and brackish water, along the coast as well as inland. Is often overlooked due to
secretive habits; thus, there are only few published data. However, it is widespread in
Germany, extending to at least 800 m a.s.l. in the Bavarian Alps.
Frommer (1996), Heller (1987a), Jöst (1966), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1987),
Strübing (1960), Voigt (1978), Wagner (1935), Nickel & Remane (1996), HN

Chloriona unicolor (Herrich-Schäffer, 1836)

Is often cited to be originally described from western Germany (e.g. Nast 1987). In fact
Herrich-Schäffer (1836) only states: „Ein Weib von Wien. Mann und Weib von Zürich von
Hrn. Bremy.“ The type material is lost anyway, so the identity cannot be clarified any-
more. However, Fieber (1866), who was the first revisor, published illustrations of the
male genital segment of several Chloriona species. His interpretation was also adopted by
Asche (1982a).
So far M VI – E VIII; nymph, 2(?) gen.
Monophagous on Phragmites australis. In Germany only known from three localities in
inland salt marshes and calcareous spring mires in Hessen (Wetterau near Münzen-
berg, 150 m a.s.l., several records, between 1966 and 1993) and Thuringia (Heilingen
near Orlamünde, 30.VI.1965, and Röttelmisch near Kahla, 16.VII.1968, both at c. 200 m
a.s.l.). Reported from 1400 m a.s.l. in the Engadine (Switzerland).
Asche (1982a), Fröhlich (1996a), Schiemenz (1987); see also Günthart (1987a), Nast (1984)

Chloriona sicula Matsumura, 1910

Chloriona flaveola Lindberg, 1948 sec. Asche (1982a)
So far E V – M VIII; nymph, 2 gen.
Monophagous on Phragmites australis. In Germany at the northeastern edge of the range
and only known from 5 sites along the upper course of the Rhine between Waldshut
and Speyer, but probably under-recorded. The habitats in these localities included var-
ious freshwater sites (along streams, in sand pits, a quarry and a garden pond), between
100 and 400 m a.s.l.
Remane & Fröhlich (1994b), HN; see also Nast (1984)

Chloriona dorsata Edwards, 1898

So far E V – M VII; nymph, 1(?) gen.
Monophagous on Phragmites australis in various habitats (along rivers, oxbow lakes,
ponds, fens, ditches, inland salt marshes and others). The first records for Germany
have been published only recently, but the species is rather widespread at least in mid-
dle parts. Furthermore, it is recorded from the foothills of the Alps (Murnau, 620 m
a.s.l.), the upper Rhine plain (Speyer), the lower part of the Elbe valley (Magdeburg,
Bleckede) and eastern Holstein (Kiel). Apparently, the German localities are at the south-
 Delphacinae Leach, 1815 45

ern and eastern edge of the range; however, the taxonomic relationship with taxa de-
scribed from central Asia should be checked.
Remane & Fröhlich (1994b), Nickel (1994, 1999b), HN; see also Nast (1976a, 1987)

Chloriona glaucescens Fieber, 1866

M V – M IX; nymph, 2 gen.
Monophagous on Phragmites australis, but largely confined to saline sites, rarely in fresh-
water sites with strongly fluctuating water tables and temporary salt accumulation in
the upper soil layer. In Germany mainly near the coasts of the North and Baltic Sea, also
in inland saltmarshes, sand pits and coal mines, mainly in Saxony-Anhalt and Thurin-
gia, locally in southern Niedersachsen, Hessen and Saxony (Werra valley, Wetterau,
near Bautzen and Boxberg); all localities are below 150 m a.s.l.
Fröhlich (1997), Funke & Witsack (1998), Schaefer (1973), Schiemenz (1987), Strübing (1960), Wagner (1935,
1939a), HN

Chloriona smaragdula (Stål, 1853)

E V – B VIII, according to Schiemenz (1987) until B IX; nymph, 1 - 2 gen.
Monophagous on Phragmites australis, almost exclusively inland. Eurytopic in freshwa-
ter sites (even along ditches within fields), also at the edge of inland salt marshes, but
largely absent from coastal salt marshes near the North and Baltic Sea. Single speci-
mens from straw meadows are presumably vagrants. Apart from that, this species is
widespread in low-lying regions of Germany; it is usually the most common among
reed-dwelling Auchenorrhyncha species; in the foothills of the Alps it frequently occurs
up to 800 m a.s.l.
Fischer (1972), Heller (1987a), Niedringhaus (1997), Reimer (1992), Schiemenz (1987), Strübing (1960), Wagner
(1935), HN

Chloriona stenoptera (Flor, 1861)

So far B VI – B VII, in Poland until E VII; nymph, 1(?) gen.
Monophagous on Phragmites australis; recorded only a few times in central Europe: Bia-
lowieza (eastern Poland), 09.VI.1949, 1 †, and 21.VII.1949, 4 ††, 15 ‡‡, in an open, peaty
forest of spruce, pine and birch; Baden-Württemberg: Horrheim near Stuttgart, 200 m
a.s.l., 07.VI.1983, 1 †, 3 ‡‡, in a “non-flooded stand of reed“; Schopfloch near Esslingen
(Swabian Jura), 750 m a.s.l., 07.VII.1975, 2 ††. There are three more recent records near
the Austrian and Bavarian edge of the Alps: Pinswang am Lech (North Tyrol, Austria),
830 m a.s.l., 30.VI.1997, 1 †, in reeds along a small brooklet in a meadow; Isar floodplain
near Vorderriß, 800 m a.s.l., 04.VI.1998, 3 ††, in reeds growing among willow scrub;
Gröbenzell near Munich, 500 m a.s.l., June 1999. Otherwise this species is only known
from Estonia, Latvia and Kazakhstan.
Heller (1987a), SMNS, Weis & Schönitzer (2001), Nickel (1999a); see also Mityaev (1971), Nast (1958)
46 Review of species

Chloriona vasconica Ribaut, 1934

So far mainly E V – M VII, according to Schiemenz (1987) until B IX; nymph, 1 - 2 gen.
Monophagous on Phragmites australis, usually along rivers and streams, lakes, ponds,
oxbow lakes, in wet ruderal sites and inland salt marshes. Widespread in the lowlands
of Germany, but only scattered, recorded up to 600 m a.s.l. only.
Heller (1987a), Schiemenz (1987), Wagner (1935), Nickel & Remane (1996), Nickel (1994), HN

Unkanodes excisa (Melichar, 1898)

E V – B IX; nymph, 2 gen.
In Europe largely confined to the coasts of the Baltic Sea; otherwise reported only from
Yakutia, the Kurile Islands and Alaska. Lives on Leymus arenarius, usually on primary
and white dunes. Found in Germany only locally along the Baltic Sea and in one inland
site near Königs Wusterhausen. The latter record may refer to vagrant individuals or
originate from a secondary plantation.
Schaefer (1973), Schiemenz (1987), Wagner (1935), HN; see also Anufriev & Emelyanov (1988), Wilson (1988)

Megadelphax haglundi (J. Sahlberg, 1871)

In Sweden B VI – E VII, in Austria E VIII (Ossiannilsson 1978; Nast 1986); nymph(?),
1(?) gen.
There is only a single German record from the Swabian Jura: Warmberg near Ober-
stetten, c. 800 m a.s.l., 23.VI.1968, 1 † on a xerothermic, stony, calcareous upland with
dwarf blackthorn, juniper and groups of various trees. In Europe otherwise recorded
from southern Sweden, Bohemia, Moravia (both Czech Republic), Niederösterreich
(Austria) and the French Massif Central; also reported from Mongolia. The host plants
are unknown, but are likely to include species of Poaceae.
Heller (1996); see also Anufriev & Emelyanov (1988), Nast (1987), Remane & Fröhlich (1994b)

Megadelphax sordidula (Stål, 1853)

B V – E IX; nymph, 2 gen.
On Arrhenatherum elatius in moderately dry to damp, occasionally also moist meadows,
although in most agricultural areas nowadays largely confined to narrow strips along way-
sides and ditches. The species is widespread in middle and southern parts of Germany, but
has only rarely been recorded in the north German plain. To the north of the Danube it
occurs only at lower altitudes (usually below 500 m a.s.l.), in the Bavarian Alps it is found
up to at least 800 m a.s.l. Is not known from Denmark and Britain, but common in southern
and middle parts of Sweden and Finland. The species is reported from there to invade
wheat and oat fields and to be a potential vector of Phleum green stripe virus (PGSV).
Achtziger (1991), Fischer (1972), Günthart (1988), Marchand (1953), Müller (1978), Reimer (1992), Schie-
menz (1987), Schwoerbel (1957), Wagner (1935, 1951a), Nickel & Achtziger (1999), HN; see also Ossiannils-
son (1978), Raatikainen (1960, 1970a)
 Delphacinae Leach, 1815 47

Laodelphax striatella (Fallén, 1826)

E IV – B X; nymph, 2 gen.
This species lives in unstable populations in dry to moist, usually anthropogenically
affected sites (mainly fallow fields, fertilized meadows, waysides) on various grasses,
including cereal crops, perhaps also on Cyperaceae and Juncaceae. The bulk of individ-
uals is macropterous and highly vagile. In Germany continuously reproducing popula-
tions with a higher proportion of brachypters are largely confined to rather warm low-
lands in the southern half (mainly to the south of the Main, perhaps also in Thuringia
and Saxony). Macropterous individuals may occur in larger numbers, but with much
varying reproductive success, more or less in the whole country, with single individuals
also in the north German plain and the higher parts of the Mittelgebirge and the Alps,
were they may be found up to 1800 m a.s.l. The species is reported to be noxious to
cereal crops (including rice) by transmission of various diseases, notably Maize rough
dwarf virus (MRDV) and Barley yellow striate mosaic virus (BYSMV) in warmer coun-
tries. In most parts of central Europe, however, the species is probably too scarce to
have an economic impact.
Fischer (1972), Günthart (1988), Marchand (1953), Niedringhaus & Olthoff (1993), Remane (1987), Schie-
menz (1971a, 1975), Schwoerbel (1957), Wagner (1935), Nickel & Achtziger (1999), Nickel (1997), HN; see
also Brčák (1979), Conti (1988)

Ditropsis flavipes (Signoret, 1865)

B V – B X; nymph, 2 gen.
Rather widespread and locally common in the Bavarian Alps, the Allgäu and the Swa-
bian and Franconian Jura; in sunny, moderately or temporarily dry pastures and mead-
ows, scattered and less abundant in base-rich and increasingly dry grassland habitats
of lower altitudes towards the north. The host plant is Bromus erectus. The northernmost
localities, some of which appear to be rather isolated, are on a line from Mechernich
(northern Eifel Mountains) to Lorch (Rheingau), Bad Wildungen, Bad Frankenhausen
and Eberswalde. Found up to 1000 m a.s.l. on the Swabian Jura and in the Bavarian
Alps, up to 1400 m a.s.l. in Vorarlberg (Austria).
Bornholdt (1991), Fischer (1972), Remane & Fröhlich (1994b), Rombach (1999a), Schiemenz (1987), Wagner
(1939a), Nickel & Achtziger (1999), HN; see also Moosbrugger (1946)

Paraliburnia adela (Flor, 1861)

M V – B IX; nymph, (1-?) 2 gen.
Monophagous on Phalaris arundinacea, usually in temporarily flooded, rather eutrophic,
sunny to shady sites, notably along rivers, streams and ditches, in willow-poplar flood-
plain woods and in low-input meadows. Lives near the base of the host plant and is
rarely caught with the sweep-net. In the older literature Glyceria has been reported as
host plant, which is caused by confusion of ‡‡ with those of Struebingianella lugubrina
(Boh.). There are only few published records; however, P. adela (Fl.) is widespread in
48 Review of species

Germany, fairly common in most parts and can be easily found when searching directly
on the ground. Extends up to at least 800 m a.s.l. in the foothills of the Alps.
Fröhlich (1996a), Kirschbaum (1868), Remane (1962), Schiemenz (1987), Nickel & Achtziger (1999), Nickel
& Remane (1996), HN; see also Lauterer (1980)

Paraliburnia clypealis (J. Sahlberg, 1871)

E V – B IX; nymph, (1-?) 2 gen.
This is an essentially north European species, which is known in central Europe only
from scattered records in the Netherlands, the Czech Republic and the northern half of
Germany. There are more or less isolated localities in the north German plain, the Harz
Mountains (Clausthal-Zellerfeld, 550 m a.s.l.), as well as middle Hessen (Schweinsberg
and Großenmoor, up to 300 m a.s.l.). Lives in wet, usually acidic and peaty, sunny to
shady sites (mainly fen woods, intermediate bogs and fens) in tall stands of Calamagros-
tis canescens; but there are also host reports of Rhynchospora and Eriophorum from the
Netherlands. Is perhaps under-recorded due to secretive habits and clumped occur-
rence.
Niedringhaus & Olthoff (1993), Remane (1962), Remane & Fröhlich (1994b), Schiemenz (1987), Nickel &
Sander (1996), HN; see also Bieman & Booy (1984), Dlabola (1970)

Hyledelphax elegantula (Boheman, 1847)

E IV – E IX; nymph, 2 gen., perhaps 1 at higher altitudes.
In the grass layer of deciduous and coniferous forests in moderately dry to moist sites,
also under alleys and even solitary trees and shrubs. Rarely and in low numbers in
treeless heaths and bogs, but presumably only as vagrant. Lives on various grasses,
notably Calamagrostis arundinacea, Brachypodium pinnatum, Holcus mollis, Molinia caer-
ulea, Deschampsia flexuosa, in the lowlands of southern and middle parts of Germany,
but at higher altitudes and in the north German plain, it is largely confined to the latter
species. Widespread in Germany, found up to at least 1300 m a.s.l. in the Mittelgebirge
and the Bavarian Alps, up to the subalpine belt in Austria and Switzerland.
Bittner & Remane (1977), Fischer (1972), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1987),
Wagner (1935, 1939a), HN and others; see also Günthart (1987a), Wagner & Franz (1961)

Megamelodes lequesnei W. Wagner, 1963

Was misinterpreted by Scott (1870) and since then erroneously published as M. quadrimaculatus
(Sign.) – see below. Much later Wagner (1963), who had revised Signoret’s types, discovered this
error and created a nomen novum (in a footnote!). Hence, older records of M. quadrimaculatus
(Sign.) are in need of revision.
In Britain VIII – III (Le Quesne 1960); adult(?), 1 gen.
This is a species with insufficiently known biology and distribution, which was record-
ed from Germany only from a single site in Holstein: Vollstedter See near Rendsburg,
November 1970, 1 † in a pitfall trap in a bog. In Britain it is reported to live mainly in
 Delphacinae Leach, 1815 49

base-rich fens and has been suspected to be associated with Juncus subnodulosus; in France
it is widespread and lives in fens with dominating tall sedges.
Remane (pers. comm.), Remane & Fröhlich (1994b); see also Kirby (1992)

Megamelodes quadrimaculatus (Signoret, 1865)

Liburnia fieberi Scott, 1870
Adults I – XII, mainly VIII – VI; adult, 1 gen.
In wet to temporarily wet, mesotrophic to moderately eutrophic, neutral to basic sites,
mainly spring mires, fens and wet meadows, but also inland salt marshes and over-
grown sand pits. The host relationships are not well known. Adults are often found in
the vicinity of rushes (Juncus inflexus and J. articulatus) and sedges (Carex spp.), and the
species has also been published from grasses (notably Poa spp.) and dicotyledons. How-
ever, the few definite host records (of large numbers of adults and nymphs) are from
mixed stands of non-flowering and thus inconspicuous Festuca pratensis and F. rubra. In
Germany this species is found only sporadically, at the edge of the range, and often in
very small pockets only, at least up to 500 m a.s.l. Northernmost localities are near Bonn,
Herne, Hanover, Hamburg, Neumünster, Rostock and Frankfurt an der Oder. It is not
found in northern Europe and Poland; from Denmark there is only a single record from
the 19th century. It is probably often overlooked due to its specific habitat requirements,
the late appearance and epigeic habits.
Köhler (pers. comm.), Maczey (pers. comm.), Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b),
Schiemenz (1987), Wagner (1935, and unpublished data), Nickel & Achtziger (1999), Nickel (1997, 1999b),
HN; see also Bieman & Booy (1984), Nast (1987), Ossiannilsson (1978)

Calligypona reyi (Fieber, 1866)

Mainly E VI – M IX, at least in favourable years M V; nymph(?), 1 - 2 gen.
Usually in permanently flooded sites near fresh or moderately saline, usually standing
water, rarely also in temporarily dry sites or near running water; mostly along shores of
lakes and ponds, occasionally in sand and gravel pits, saltmarshes and garden ponds.
Host plants are Schoenoplectus lacustris and Sch. tabernaemontani. Until recently there
were only a few published records from Germany (mainly near the coasts of the North
and Baltic Sea, as well as near Waren an der Müritz, Leipzig, Gera and Frankfurt am
Main). However, the species is widespread all over the country and extends at least up
to 730 m a.s.l. in the Allgäu and Upper Bavaria. May be fairly abundant in larger stands
of host plants.
Harz (1988), Remane & Fröhlich (1994b), Schiemenz (1987), Wagner (1939a), Nickel & Remane (1996), Nickel
(1997), HN

Mirabella albifrons (Fieber, 1879)

B IV – E IX; nymph, 2 gen.
50 Review of species

On Calamagrostis epigejos in sunny to slightly shady, moderately dry to moderately moist

sites, mainly in forest clearings, ruderal sites, sand and gravel pits, mining areas and on
railway embankments; rarely in rather moist sites on C. canescens. Widespread in south-
ern and eastern parts of Germany and common in favourable sites, but at the edge of
the range. Northern- and westernmost localities are on a line from Marburg to Göttin-
gen, Braunschweig, Uelzen and Greifswald. However, the species is also reported from
the Netherlands and the Paris basin. The highest localities are at c. 650 m a.s.l. in Upper
Bavaria.
Fischer (1972), Remane & Fröhlich (1994b), Schiemenz (1987), Walter (pers. comm.), Nickel (1994), HN; for
identification see Dlabola (1954)

Delphacodes capnodes (Scott, 1870)

Adults I – XII, mainly E VIII – E VI; adult, 1 gen.
In tall-sedge swamps, reeds (also in moderately saline sites) and intermediate bogs, the
latter usually with open stands of birch or pine. It lives close to the ground, often deep
in hummocks of Sphagnum and is rarely collected; most specimens are macropterous
individuals on dispersal flight. Host plants are sedges and cottongrasses. In fens it is
found on tall sedges (Carex acutiformis, C. acuta and C. riparia are reported from the
Czech Republic and Slovakia); in peaty sites of Germany and the Netherlands it is ap-
parently associated with Eriophorum angustifolium. Populations in fenland habitats con-
sist mainly or even exclusively of ‡‡, the biology of which is poorly known. Pseudoga-
mous, triploid ‡‡ from a Greek population have been found to respond to acoustic
signals of Dutch †† of D. capnodes and were also inseminated, but without passing the
male genome to their offspring. In Germany this species appears to be rare and rather
localized although it is widespread; the highest localities are at 600 m a.s.l. only.
Fröhlich (1996a), Funke & Witsack (1998), Niedringhaus & Olthoff (1993), Remane (1958), Schiemenz (1987),
Strübing (1955), Wagner (1938b, 1940a), Nickel (1999b), HN; see also den Bieman & de Vrijer (1987), Laut-
erer (1980)

Delphacodes venosus (Germar, 1830)

Adults I – XII, mainly B IX – E VII; adult, 1 gen.
In tall grass stands in moderately shady to sunny, peaty to moist, sometimes also damp
or moderately dry, usually cool sites, mainly low-input meadows and pastures, fens,
intermediate bogs and fen woods. The host plants include Deschampsia cespitosa, Festuca
arundinacea, Agrostis spp., Molinia caerulea and other grasses, and probably sedges (Car-
ex spp.). Has repeatedly been considered as tyrphophilous or even tyrphobiotic. How-
ever, in central Europe most populations are clearly found in moderately eutrophic,
moist to temporarily wet sites. The species is widespread in Germany and fairly com-
mon, but often overlooked. Most specimens are caught in pitfall traps. Frequently found
up to at least 1000 m a.s.l. in the Mittelgebirge and the Bavarian Alps.
Fischer (1972), Schiemenz (1987), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999),
HN; see also Lauterer (1980), Ossiannilsson (1978)
 Delphacinae Leach, 1815 51

Gravesteiniella boldi (Scott, 1870)

B V – E X; nymph, 2 gen.
On Ammophila arenaria on white and grey dunes near the coast, occasionally also on
open inland dunes of the north German plain. In Germany locally common on the East
and North Frisian Islands, Heligoland and along the Baltic Sea coast, also found inland
near Hamburg and Dömitz. Is also reported from eastern parts of Austria, where A.
arenaria is absent.
Niedringhaus (1991, 1994), Schaefer (1973), Schiemenz (1987), Wagner (1935), HN; see also Holzinger &
Remane (1994), Ossiannilsson (1978)

Muellerianella brevipennis (Boheman, 1847)

E VI – E X; egg, 1 - 2 gen. (see Drosopoulos 1977, Witsack 1971).
On Deschampsia cespitosa in sunny to shady, moderately moist to wet, usually tempo-
rarily waterlogged sites, mainly meadows, pastures, abandoned grassland, fens, clear-
ings, open forests, also forest roads. In Germany widespread and in intermediate to
high frequency on the host, also at higher altitudes of the Mittelgebirge, found up to at
least 1500 m a.s.l. in the Allgäu Alps.
Fischer (1972), Nikusch (1976), Reimer (1992), Schiemenz (1987), Wagner (1939a), Nickel & Achtziger (1999),
HN

Muellerianella fairmairei (Perris, 1857)

Was formerly considered to be conspecific with the following species, until Booij (1981) demonstrat-
ed that they are clearly distinct. Thus, most older records are in need of revision. The complicated
biology was studied by Drosopoulos (1977), Booij (1981, 1982) and Booij & Guldemond (1984).
E VI – E X; egg, 2 gen., probably 1 at higher altitudes.
In wet to damp, usually cool and moderately eutrophic sites, mainly meadows, pas-
tures and forest glades. Food plants are Holcus lanatus and H. mollis. At least the former
is also the oviposition plant under long-day conditions. Under short-day conditions,
overwintering eggs are laid exclusively on Juncus effusus, which is left, however, imme-
diately after hatching. The species reaches the eastern edge of its range in Germany
where it is restricted to oceanic climates of the western and northwestern parts. In the
Mittelgebirge reproducing populations were found up to 1000 m a.s.l. Easternmost lo-
calities are near Lörrach, Todtnau (Black Forest), Emmendingen, Bad Nauheim, Laas-
phe, Brilon, Hannoversch Münden, Lüneburg, Boizenburg and Rendsburg, as well as
in Jutland (Denmark) and southern Sweden. At present, the species is expanding its
range and has apparently colonized most parts of eastern Germany. Recently it has
been recorded in numerous sites where it was absent some years ago, e.g. near Gardele-
gen, Northeim, Göttingen, Brandenburg, Schwedt (Oder) and Pirna. Populations near
the edge of the range may include up to 100% of triploid, pseudogamous ‡‡.
Booij (1982), Booij & Guldemond (1984), Drosopoulos (pers. comm.), Fröhlich (1996a), K. Heller (pers.
comm.), Niedringhaus & Olthoff (1993), Rothenbücher (pers. comm.), Walter (pers. comm.), Nickel & Acht-
ziger (1999), Nickel (1997), HN
52 Review of species

Muellerianella extrusa (Scott, 1871)

Was formerly considered to be conspecific with the preceding species (e.g. by Ossiannilsson
1978) or confused with it (see above). Older records of M. fairmairei (Perr.), mainly those from
the central and eastern parts of central Europe, usually refer to M. extrusa (Scott).
E VI – E X; egg, according to Schiemenz (1987) occasionally adult, 1 gen.
On Molinia caerulea in peaty to moist, also temporarily moist sites, mainly straw mead-
ows, margins and degenerated stages of bogs and open forests. Recorded from most
parts of Germany, rather wide and densely spread in the Alps and their foothills and
the Mittelgebirge, but more scattered in the lowlands, where the species has probably
much decreased due to drainage, fertilizing and ploughing of former habitats. Occurs
up to at least 1000 m a.s.l. in the Mittelgebirge, to at least 1200 m a.s.l. in the Allgäu Alps.
Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1987), Nickel & Achtziger (1999), Nickel & Re-
mane (1996), HN; for identification see Booij (1981), Ossianilsson (1983)

Chlorionidea flava P. Löw, 1885

B V – B IX; nymph, 1 - 2 gen.
On Sesleria albicans, usually in open pine forests in dry to temporarily dry and basic sites
on sun-exposed calcareous and gypsum hillsides; in the foothills of the Alps also on
rather dry riverine gravel banks, at higher altitudes locally in open pastures. In Germa-
ny at the northwestern edge of the range and known from Thuringia (basins of Saale,
Unstrut, Ilm and Werra), Mainfranken, the Franconian and Swabian Jura and the Alps
and their foothills, also found near Eschwege and Göttingen. Highest localities are at c.
1000 m a.s.l. in the Swabian Jura and at c. 1300 m a.s.l. in the Bavarian and Allgäu Alps,
but the species is reported from the alpine belt in Austria.
Haupt (1925), Remane & Fröhlich (1994b), Schiemenz (1987), Nickel (1994), HN; see also Wagner & Franz
(1961)

Litemixia pulchripennis Asche, 1980

B VIII – B X; egg, 1 gen.
This striking species was described only 20 years ago from western France. It lives deep
in tussocks of Molinia caerulea in open pine forests on temporarily moist, acidic and
sandy substrates. More recently it was found in three sites in the northern upper Rhine
plain (near Speyer, Germersheim and Neustadt an der Weinstraße, all at c. 100 m a.s.l.).
Remane & Fröhlich (1994b), HN; see also Asche (1980), della Giustina & Remane (1992), Remane & della
Giustina (1991)

Muirodelphax aubei (Perris, 1857)

E IV – B X; nymph, 2 gen.
In grass stands in rather dry, sunny, often disturbed sites usually with some bare ground,
for instance on open sand of inland and tertiary coastal dunes, in military training ar-
eas, dry abandoned fields, in mining areas, etc., also on strongly grazed calcareous hill-
 Delphacinae Leach, 1815 53

sides and plateaus. The host plant is perhaps Poa pratensis. In Germany mainly found
near the coasts of the North and Baltic Sea and in the sand regions of the north German
plain, but also on limestone, gypsum and other substrates in more southern regions,
where it is rather scattered and more confined to warm and dry localities, notably in the
rain shadow area to the east of the Harz Mountains, Mainfranken, Kaiserstuhl, as well
as inland dunes in the valleys of Rhine, Main, Regnitz and Danube. Most sites are be-
low 400 m a.s.l., one locality in the foothills of the Alps is at c. 800 m a.s.l.
Fischer (1972), Niedringhaus (1991), Nikusch (1976), Remane (1987), Schiemenz (1987), Schwoerbel (1957),
Trümbach (1959), Wagner (1935, 1939a, 1951a), HN

Acanthodelphax denticauda (Boheman, 1845)

M IV – B X; nymph, 2 gen.
On Deschampsia cespitosa in moist to wet meadows, pastures and abandoned grassland,
locally also in forests. Widespread in Germany and in intermediate frequency on the
host, although often underrepresented in sweep-net samples. Found up to at least 850 m
a.s.l. in the Mittelgebirge and the foothills of the Alps.
Bittner & Remane (1977), Fischer (1972), Niedringhaus & Olthoff (1993), Schiemenz (1987), Schwoerbel
(1957), Wagner (1935, 1939a), Wagner & Franz (1961), Nickel & Achtziger (1999), Nickel (1997), HN

Acanthodelphax spinosa (Fieber, 1866)

M IV – M X; nymph, 2 gen., probably 1 at higher altitudes.
On Festuca rubra and, less frequently, F. ovina in sunny, damp to moist, occasionally also
peaty to dry sites (mainly oligotrophic or wet meadows, fens, dry grassland, ruderal
sites, waysides, etc.). The species is largely absent from intensively managed meadows,
which may be invaded by single macropterous individuals on dispersal flight, but with-
out long-termed reproductive success. It is widespread in Germany and common in
many places, although more scattered in northern parts, reaching the edge of the range
in Holstein (Heide and Lübeck). Occurs up to at least 1500 m a.s.l. in the Mittelgebirge
and the Bavarian Alps, up to c. 2000 m a.s.l. in North Tyrol and Styria (both Austria).
Bittner & Remane (1977), Fischer (1972), Marchand (1953), Nikusch (1976), Reimer (1992), Schiemenz (1987),
Wagner (1935), Nickel & Achtziger (1999), HN; see also Leising (1977), Wagner & Franz (1961)

Nothodelphax albocarinata (Stål, 1858)

Was not distinguished from N. distincta (Fl.) until Kontkanen (1952) clarified the identity of both
species. The majority of the older central European records, for instance Haupt (1935) and Wag-
ner (1948b), refer to N. distincta (Fl.).
B V – B IX; nymph, 2 gen.
This is a rare and tyrphobiotic species, occurring in central Europe only in very isolated
sites near the south-western edge of the range. Found along hollows in raised and inter-
mediate bogs; the host plant is probably Carex limosa. In Germany there are only 12
confirmed localities between Halle, Eilenburg, Bautzen, Weißwasser and Luckau, as
54 Review of species

well as in Upper Bavaria (Sachsenkam and Seeshaupt), up to at most 700 m a.s.l. To the
west of these, there is only a single specimen from the French Jura and a more recent
record from near Apeldoorn (Netherlands).
Remane & Fröhlich (1994b), Schiemenz (1976, 1987), Walter (pers. comm.), HN; see also Bieman & Roze-
boom (1993), Remane & della Giustina (1993)

Nothodelphax distincta (Flor, 1861)

M IV – B IX, at higher altitudes mainly V – VII; nymph, 1 - 2 gen.
Tyrphobiotic in open raised bogs on Eriophorum vaginatum. In Germany occurring rather
scattered in the remnant bog areas of the north German plain, the highest parts of the
Mittelgebirge (mainly Solling, Harz Mountains, Thuringian Forest, Erzgebirge, Rhön Moun-
tains, Black Forest) and the foothills of the Alps. The highest localities are at 900 m a.s.l.
Bittner & Remane (1977), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1958), Schiemenz (1971a,
1975, 1976, 1987), Wagner (1948b), Nickel & Remane (1996), Nickel (1997, 2002), HN

Dicranotropis hamata (Boheman, 1847)

M IV – E X; nymph, 2 gen., according to Schiemenz (1987) only 1 at higher altitudes.
Eurytopic on various grasses in moist to moderately dry, sunny to moderately shady,
often disturbed sites (mainly waysides, ruderal sites, forest clearings, abandoned fields,
occasionally also fertilized meadows and pastures). Host plants are Elymus repens, Dacty-
lis glomerata, Holcus lanatus, H. mollis, Festuca arundinacea and other grasses. Widespread
and common in Germany, extending to at least c. 1100 m a.s.l. in the Mittelgebirge and the
Alps. Is reported from Scandinavia and former Czechoslovakia as a potential vector of
cereal virus diseases, but is unlikely to cause major damage at least in central Europe.
Achtziger (1991), Fischer (1972), Niedringhaus (1991), Remane (1987), Schiemenz (1987), Wagner (1935),
Nickel & Achtziger (1999), HN and others; see also Brčák (1979), Lindsten (1979)

Dicranotropis divergens Kirschbaum, 1868

M V – B X; nymph, 1 gen.
In oligotrophic to moderately eutrophic meadows and pastures of higher altitudes in
moderately dry to moist, occasionally also wet sites. Lives on Festuca rubra, perhaps
also on additional grasses. In Germany particularly widespread and common in the
Bavarian and Allgäu Alps and their foothills up to at least 1800 m a.s.l. (but found up to
2200 m a.s.l. in Austria and Switzerland). Also occurring in most of the higher Mittelge-
birge: Bavarian Forest, Oberpfälzer Wald, Fichtelgebirge, Frankenwald, Rhön Moun-
tains, Thuringian Forest, Erzgebirge, as well as Black Forest, Swabian Jura and Harz
Mountains, usually above c. 600 m a.s.l., locally down to 400 m a.s.l. Repeated records
of single brachypters in lowland areas are probably caused by passive drift or short-
term reproduction of vagrants.
Achtziger (1991), Bittner & Remane (1977), Kupka (1941), Reimer (1992), Schiemenz (1987), Wagner (1958),
Nickel & Achtziger (1999), HN; see also Günthart (1987a), Leising (1977)
 Delphacinae Leach, 1815 55

Dicranotropis montana (Horváth, 1897)

Dicranotropis gratiosa Dlabola, 1997 (see Holzinger 1999a)
In Austria B VI – E VIII (Dlabola 1997; Moosbrugger 1946); nymph, 1 gen.
In Europe only known from a few records from the higher parts of the Carpathians (Ro-
mania) and Alps (Austria, northern Italy, Bavaria). In Germany the only locality is in the
Allgäu: Rotspitze above Hindelang, 1800 m a.s.l., 04.VIII.1995, 7 ††, 6 ‡‡ on Deschampsia
cespitosa in a subalpine pasture close to a brooklet. Is probably more widespread in the
German Alps and has been found near Scharnitz (North Tyrol, close to the Bavarian bor-
der). Was recorded in the Dolomites (Italy) from grasses in an open coniferous forest.
Nickel (1999b); see also Asche (1982b), Remane & Fröhlich (1994b); for identification see Holzinger (1999a)

Florodelphax leptosoma (Flor, 1861)

E IV – M X; nymph, 2 gen., probably 1 at higher altitudes.
On rushes (Juncus articulatus, J. effusus, J. subnodulosus, J. conglomeratus and others) in
moderately wet to temporarily flooded, sunny to slightly shady sites, mainly low-input
meadows, spring mires and intermediate bogs, locally also mining areas and sand pits.
Wide and densely spread in Germany, but often only in small populations, extending
up to at least 1100 m a.s.l. in the Mittelgebirge and the Alps, but found at 1500 m a.s.l. in
North Tyrol (Austria).
Fischer (1972), Nikusch (1976), Niedringhaus (1991), Reimer (1992), Schiemenz (1987), Wagner (1935, 1939a),
Nickel & Achtziger (1999), HN and others

Florodelphax paryphasma (Flor, 1861)

M V – E IX; nymph, 2 gen.
On Carex disticha in moderately eutrophic and temporarily waterlogged or flooded sites
(mainly wet meadows, fens, spring mires, floodplain depressions, shores of ponds and
lakes, etc.). In Germany the species has often been overlooked and was until recently
only known from very few localities mainly in the northern plain. However, according
to more recent data, it is also widespread in the middle parts where it often occurs in
high frequency and individual numbers on the host. At present, southernmost localities
are known from the vicinity of Speyer, Ludwigsburg, Ansbach, Ilmenau and Dresden,
only up to 450 m a.s.l. Has been recorded in central France, the Pyrenees and the Sierra
de Cuenca of central Spain, but not yet in Austria, Italy and Switzerland.
Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Schiemenz (1987), SMNS, Wagner (1952a),
Walter (pers. comm.), Nickel & Achtziger (1999), HN; see also della Giustina & Remane (2001)

Kosswigianella exigua (Boheman, 1847)

M III – B X; nymph, 2 gen.
In sunny, oligotrophic and low-vegetated, often grazed grasslands in moderately dry to
dry, occasionally also damp or very dry sites on various substrates (both sandy to solid,
56 Review of species

basic to acidic). Host plants are fine-leaved species of fescue (Festuca ovina group), al-
though additional grasses (Corynephorus canescens, Deschampsia flexuosa) have been re-
ported. Widespread in Germany and locally abundant, extending up to at least 1100 m
a.s.l. in the Mittelgebirge and the Alps.
Fischer (1972), Niedringhaus (1991), Remane (1987), Rombach (1999b), Schiemenz (1987), Schwoerbel (1957),
Wagner (1935, 1939a, 1951a), HN and others

Scottianella dalei (Scott, 1870)

Delphacodes gravesteini Dlabola, 1971 sec. Anufriev (1980)
In England according to Waloff & Solomon (1973) IV – VIII; nymph, 2 gen.
Occurs from Morocco through western Europe northward to England, known from
Germany only from two records of single, brachypterous †† in the extreme west: Wah-
ner Heide near Cologne, c. 50 m a.s.l., 08.VIII.1982, and Dudenhofen near Speyer, c.
110 m a.s.l., 25.IV.1989. Was found among grasses in moist sites in France („around ponds,
lakes, marshes, etc.“), on Agrostis capillaris in England.
Bieman & Booy (1984), Remane (1995); see also della Giustina (pers. comm.), Waloff & Solomon (1973)

Struebingianella lugubrina (Boheman, 1847)

E IV – M IX; nymph, 2 gen.
On species of sweet-grass (Glyceria maxima, G. fluitans group) in wet, usually temporari-
ly flooded, rather eutrophic, sunny to slightly shady sites, mainly near standing and
running water (along rivers, streams, ditches, in floodplain depressions), in wet mead-
ows, alder fens, as well as on temporarily dry bottoms of ponds. Widespread and fairly
common in the lowlands of Germany, in intermediate to high frequency on both host
species; rather scattered at higher altitudes and extending up to at least 900 m a.s.l.
Fischer (1972), Niedringhaus (1997), Reimer (1992), Schiemenz (1987), Wagner (1935, 1939a, 1951a), Nickel
& Achtziger (1999), Nickel (1997), HN

Xanthodelphax flaveola (Flor, 1861)

B VI – B IX; nymph, 1(?) gen.
In sunny to slightly shady grass stands in moderately acidic, moderately dry to moist
sites, mainly low-productivity pastures and meadows, usually on loamy or sandy sub-
strates. The host plant is Poa pratensis. In Germany rather localized and uncommon,
mainly in the eastern parts of the north German plain and the Mittelgebirge (Harz Moun-
tains, Eichsfeld, Eifel Mountains, Rhön Mountains, Thuringian Forest, Erzgebirge, Ober-
pfälzer Wald, Donnersberg, Swabian Jura, Black Forest), as well as in the foothills of the
Alps, extending up to at least 1200 m a.s.l. Has apparently declined due to intensifica-
tion of grassland management. It is not reported from Switzerland; from France it is
only known from a few localities in the southern half.
 Delphacinae Leach, 1815 57

Fischer (1972), Fröhlich (1996a), Nikusch (1976), Schiemenz (1987), Walter (1998), Wagner (1939a), HN; see
also della Giustina & Remane (1999)

Xanthodelphax straminea (Stål, 1858)

B V – M IX; nymph, 2 gen.
In sunny to moderately shady, wet to moderately dry, acidic to neutral, oligotrophic to
moderately eutrophic sites, mainly in spring mires, intermediate bogs and wet mead-
ows, usually on sandy or peaty soils, locally also in moderately saline sites; on bent-
grass (Agrostis canina, A. stolonifera, A. capillaris.). Widespread in Germany, although
scattered; more common in the sand and bog areas in the northeastern half, but rather
localized in the southwest, where recorded from Upper Bavaria, Oberpfalz, Mainfran-
ken and the upper Rhine between Lörrach and Wiesbaden. Highest localities are at
850 m a.s.l. in the Thuringian Forest and the foothills of the Alps.
Bittner & Remane (1977), Günthart (1988), Schiemenz (1971a, 1987), Wagner (1939a), Nickel & Achtziger
(1999), Nickel (1999b), HN

Xanthodelphax xantha Vilbaste, 1965

So far E VI – B VIII; nymph, 1(?) gen.
Described from the Altai and Poland, in Germany found only in two localities: Gauting
near Munich, c. 580 m a.s.l., 1959 and 1961, in a clearing within a spruce forest, and
Serrahn near Neustrelitz, c. 70 m a.s.l., 1973 and 1974, along the edge of an intermediate
bog. Possible host plants in both sites are species of Calamagrostis (C. canescens or C.
epigejos) and Molinia caerulea. Otherwise known in Europe only to occur in Latvia, Be-
lorussia and north-eastern Poland.
Remane & Fröhlich (1994b), Schiemenz (1976); see also Nast (1966), Vilbaste (1965)

Paradelphacodes paludosa (Flor, 1861)

E IV – B IX; nymph, 1 - 2 gen.
In wet, usually peaty, acidic as well as basic sites, mainly intermediate bogs, spring
mires and fens. Host plants are sedges, probably Carex rostrata on acidic, C. panicea on
more basic substrates. Widespread all over Germany, but scattered, with strongholds in
the bog areas of the north German plain and along the edge of the Alps, rather uncom-
mon elsewhere. The highest sites are at 1200 m a.s.l. in the Allgäu Alps.
Fröhlich (1996a), Marchand (1953), Remane (1958), Schiemenz (1987), Wagner (1946), Nickel & Achtziger
(1999), Nickel & Remane (1996), HN

Oncodelphax pullula (Boheman, 1852)

M V – M VIII; nymph, 1 gen.
In sedge swamps of wet to peaty, acidic as well as basic sites, mainly intermediate bogs,
calcareous small-sedge fens and birch swamps. The species lives on sedges, mainly –
58 Review of species

perhaps exclusively – on Carex nigra. It is widespread in Germany, but only very scat-
tered, highest localities are at 800 m a.s.l. in the Bavarian Alps.
Fischer (1972), Hoffmann (1980), Niedringhaus & Olthoff (1993), Remane (1962), Remane & Fröhlich (1994b),
Walter (1998), Nickel & Achtziger (1999), HN; see also Lauterer (1978)

Criomorphus albomarginatus Curtis, 1833

B V – E VII; nymph, 1 gen.
In wet to damp, sunny to shady sites, mainly low-input meadows, abandoned grass-
land, open forests, fens, waysides, ditches, forest margins, etc., avoiding high-intensity grass-
land. Host plants are various species of grasses (Festuca rubra, Deschampsia cespitosa, D.
flexuosa, Calamagrostis arundinacea and others), a few records are also from Juncaceae (Luzula
spp.). The species is widespread in the lowlands of Germany and locally common; extend-
ing to at least 1100 m a.s.l. in the Mittelgebirge, up to 1500 m a.s.l. in the Bavarian Alps.
Bornholdt (1996), Haas (1975), Niedringhaus (1991), Nikusch (1976), Remane (pers. comm.), Schiemenz
(1987), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN and others

Criomorphus borealis (J. Sahlberg, 1871)

B V – E VII; nymph, 1 gen.
This is a Siberian species, its range extending through northern Germany westward to
the Netherlands and through the Erzgebirge, the Thuringian and Bohemian Forest to
the Harz Mountains and eastern Bavaria. In the north German plain it lives only locally
on Calamagrostis canescens in intermediate bogs, fens, and fen woods (found near Pin-
neberg, Hamburg, Sassnitz/Rügen, Neustrelitz, Berlin and Niesky). In the Harz Moun-
tains, Thuringian Forest and Erzgebirge, however, it is common in many places and
often dominant in stands of Calamagrostis villosa in spruce forests and in clearings main-
ly of the submontane belt (between 350 and 1000 m a.s.l.).
Niedringhaus & Olthoff (1993), Schiemenz (1976, 1987), Strübing (1960), Remane & Fröhlich (1994b), HN;
see also Bieman & Booy (1984), Novotný (1995)

Criomorphus moestus (Boheman, 1847)

So far E V – B VII; nymph, 1 gen.
Largely confined to peatlands of northern Europe. From Germany there are several
published records most of which are based on misinterpretations. The only confirmed
records are from two intermediate bogs near Berlin, both at 50 m a.s.l.: Teufelsmoor
near Müggelheim, a population among Calamagrostis canescens in 1951 and 52, and
Maggelhansluch near Zernsdorf, 30.V.2002, 2 ††, 2 ‡‡). C. canescens is also reported as
host plant from Sweden, but the second German site was a mixed stand of Carex, Phrag-
mites and Eriophorum vaginatum without Calamagrostis. Moreover, most recently studied
stands of C. canescens in northern Germany were infested by C. borealis (J. Shlb.) only.
Strübing (1955), Coll. Maczey (vid. HN); see also Ossiannilsson (1944), Vilbaste (1960), Walter & Emmrich
(1995), Nickel & Sander (1996)
 Delphacinae Leach, 1815 59

Criomorphus williamsi China, 1939

So far E V – B VII, in England according to Hassan (1939) B V – M VII; nymph, 1 gen.
Described from England, elsewhere in Europe only recorded from Moravia (Czech Re-
public), Hungary, middle Russia and northern Germany, where known from only three
localities (all below 50 m a.s.l., all specimens brachypterous): Bremen, Stadtwald, 1982,
1 †, in an open oak-birch forest; Hamburg, Vier- und Marschlande, 01.VII.1982 and
20.VI.1983, 1 †, respectively, in abandoned fields; Amt Neuhaus near Lüneburg: Zeetze,
30.V.2000 and 07.VI.2002, a large population, in lush grass stands along a ditch, perhaps
on Poa palustris or Poa trivialis. The only record from Moravia (1 brachypterous †) is
from an open forest. The species is reported to live on grasses usually in moist sites in
England, in meadows with Rumex in middle Russia, in floodplain meadows in Kazakh-
stan. Is probably more widespread at least in lowland central Europe.
Hildebrandt (1986), Tharsen (1987), ZIMH, HN; see also Anufriev & Kirillova (1998), Emelyanov (pers.
comm.), Hassan (1939), Lauterer (1983), Le Quesne (1960), Nast (1987), Orosz (1983)

Pseudodelphacodes flaviceps (Fieber, 1866)

Specimens of Calligypona tricolorata Dlabola, 1961a and Delphacodes fenestratus Emelyanov, 1972
from central Asia have been kindly sent by A.F. Emelyanov (St. Petersburg) and exchanged for
Ps. flaviceps (Fieb.) from the Bavarian Alps. All turned out to be at least congeneric. Nomencla-
tural changes will be published elsewhere.
So far B VI – E VIII, according to Fischer (1972) also M IV; nymph, 2 gen.
On sparsely vegetated, often almost bare sand and gravel banks of unchanneled rivers
in the Alps and their foothills. In Germany very rare and only found along the Lech
(near Augsburg, 1936, i.e. before drainage) and the upper Isar: 1961 and 1994 single
macropterous †† near Ascholding and Wallgau, from 1996 to 1998 several records (in
pure stands of Calamagrostis pseudophragmites and including brachypterous individu-
als) in altogether 4 sites between Krün and Fall as well as along the Rißbach near Vorder-
riß. All localities are between 500 and 900 m a.s.l. Otherwise, in Europe only known
from old records from the Rhine floodplains in Vorarlberg (Austria) and Switzerland
(probably Rhine valley, too). Another record from the vicinity of Trieste (now Italy, but
erroneously cited from former Yugoslavia) is in need of verification.
Fischer (1972), Fröhlich (1996b), Remane (pers. comm.), Nickel (1999a); see also Melichar (1899), Moos-
brugger (1946), Nast (1987)

Toya propinqua (Fieber, 1866)

So far VIII – B X; nymph, 2(?) gen.
This is a pantropical species, in Europe largely confined to southern parts. Its occurrence
to the north of the Alps is perhaps only erratical. It lives on Cynodon dactylon and perhaps
other grasses usually in disturbed sites. Macropterous individuals on dispersal flight can
be drifted over long distances. Found in Germany only in the northern upper Rhine plain
(Mainz, Darmstadt and Speyer) and in the Lech floodplains near Landsberg (the latter
probably stray individuals). A published record from the surroundings of Berlin is based
60 Review of species

on misidentified specimens of Paraliburnia clypealis (J. Shlb.). In urban areas of Mainz the
species was found in 1993 through 1995, but not in 1996 and 1997, with brachypterous
individuals at least in 1994. Was recently reported from the French and Swiss side of the
southern upper Rhine plain near Colmar and in urban areas of Basel.
Fischer (1972), Remane (1987), Strübing (1955, and pers. comm.), Thüs (pers. comm), Wonn (1956); see also
della Giustina & Balasse (1999); Drosopoulos (1982), Mühlethaler (2001), Nast (1975), Wagner (1954)

Javesella discolor (Boheman, 1847)

In the lowlands mainly B V – M VII, at higher altitudes VI – IX; nymph, 1 gen.
Lives in the undergrowth of deciduous and coniferous forests, in shady forest glades, at
the edges of bogs (invading into the interior along ditches) and in subalpine pastures and
scrub. Host plants are mainly grasses (Deschampsia cespitosa, D. flexuosa, Calamagrostis ca-
nescens and others), probably also various species of Cyperaceae and Juncaceae (Carex,
Scirpus, Juncus, Luzula). Widespread in Germany and particularly common in the sub-
montane and montane belt (in the Bavarian Alps and the Allgäu frequently extending up
to at least 1800 m a.s.l.); less common in the lowlands, locally absent from warm regions.
Fischer (1972), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1971a, 1975, 1987), Schwoerbel
(1957), Wagner (1935, 1939a), HN; see also Novotný (1995), de Vrijer (1986)

Javesella simillima (Linnavuori, 1948)

According to Remane & Fröhlich (1994a) the status of German specimens require further inves-
tigation, particularly their relationship with J. discolor (Boh.). However, J. simillima (Lnv.) is much
smaller and paler, and the aedeagus is triangular. German specimens seen by the author were
morphologically and ecologically clearly distinct from the preceding species. Furthermore, both
taxa are reported as distinct species from large parts of arctic Europe, Asia and North America
(Anufriev & Emelyanov 1988; Ossiannilsson 1978; Wilson 1988, 1992). Thus, there is no reason
to doubt their specific separation.
E V – E VII; nymph, 1 gen.
According to existing data this taxon is largely confined to arctic regions of Asia and North
America and occurs very locally in bogs of northern Europe, the Polish Tatra and the moun-
tains of northern Bohemia and Moravia (Czech Republic). Furthermore, there are 6 locali-
ties in intermediate and raised bogs in the Erzgebirge, the Thuringian Forest, the Dresdener
Heide and the Niederlausitz, between 180 and 960 m a.s.l. Most specimens in these sites
were found on Eriophorum angustifolium, but Carex rostrata is reported as a further host.
Emmrich (1975), Schiemenz (1971a, 1975), Nickel (2002), HN; see also Anufriev & Emelyanov (1988), Lau-
terer & Novotný (1991), Ossiannilsson (1978), Wilson (1988, 1992)

Javesella pellucida (Fabricius, 1794)

B IV – M X; nymph, 2 gen.
This is a eurytopic species living in various wet to dry, sunny to moderately shady sites.
Food plants are predominantly grasses (Festuca, Elymus, Dactylis, Poa, Deschampsia, Agros-
tis, Lolium, Calamagrostis, Phleum and others), less commonly Cyperaceae, Juncaceae
 Delphacinae Leach, 1815 61

and perhaps also Equisetaceae. Populations with brachypterous individuals are usual-
ly restricted to low-input meadows, fens, abandoned fields and slightly saline sites. In
contrast, macropterous individuals fly into almost all kinds of non-forested habitats,
particularly in high summer, however, with much varying reproductive success. High-
est abundances are usually found in moderately moist to damp, moderately eutrophic
meadows and pastures (including salt marshes). J. pellucida (F.) is among the very few
species, which are able to survive at least temporarily in intensively managed grass-
land, as well as in other extreme sites (e.g. lawns, cereal fields, etc.). In Germany it is
widespread up to at least 1000 m a.s.l.; in most parts of cultivated lowlands of central
Europe it is probably the most abundant Auchenorrhyncha species. In many European
countries it is known as a vector of plant viruses, notably Oats sterile dwarf (OSDV),
Maize rough dwarf (MRDV) and European wheat striate mosaic (EWSMV).
Fischer (1972), Haas (1975), Marchand (1953), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Re-
mane (1987), Remane & Wachmann (1993), Schiemenz (1971a, 1975, 1987), Wagner (1935), Nickel & Acht-
ziger (1999), HN and others; see also Brčák (1979), Nuorteva (1965), Raatikainen (1967, 1970b)

Javesella dubia (Kirschbaum, 1868)

E IV – M X; nymph, 2 gen.
On grasses, usually Agrostis stolonifera and A. capillaris, probably also on other species,
in damp to moderately wet, sunny to shady sites (mainly meadows and pastures, forest
roads, ruderal sites, abandoned fields, moist forests and low-input lawns of gardens
and parks), under natural conditions presumably in salt marshes, along shores of lakes
and rivers and in fen woods. Widespread and common in most parts of Germany, in the
Mittelgebirge and the Bavarian Alps up to at least 900 m a.s.l., probably only temporary
populations at higher altitudes; in the Swiss Alps nymphs were found at 1200 m a.s.l.
Has been reported as a potential vector of several plant viruses, but is probably only of
minor economic importance.
Fischer (1972), Kuntze (1937), Reimer (1992), Remane (1987), Schiemenz (1987), Wagner (1939a), Nickel &
Achtziger (1999), HN and others; see also Brčák (1979), Günthart (1987a)

Javesella obscurella (Boheman, 1847)

M IV – M IX; nymph, 2 gen.
In temporarily wet, sunny to moderately shady sites on loamy and clayey substrates, local-
ly also on slightly saline soils. Primary habitats are scarcely vegetated mud banks along lake
and pond shores and in floodplain depressions, secondary ones are pastures and meadows,
waysides and mining areas. Host plants are mainly grasses (adults mostly on Alopecurus
geniculatus and Glyceria fluitans), perhaps also species of Cyperaceae and Juncaceae. Wide-
spread and common in Germany, also in higher parts of the Mittelgebirge and the Alps,
brachypterous individuals usually below 1200 m a.s.l., macropters frequently found up to
2000 m a.s.l., where breeding seems unlikely, however. Is reported as a vector of Oats sterile
dwarf virus (OSDV) from Finland, but probably without economic importance.
Fischer (1972), Heller (1987a), Leising (1977), Niedringhaus (1991, 1997), Schiemenz (1987), Schwoerbel
(1957), Wagner (1935), Nickel & Achtziger (1999), HN; see also Brčák (1979), de Vrijer (1986), Lindsten (1979)
62 Review of species

Javesella salina (Haupt, 1924)

E V – E VIII; nymph, 1(?) gen.
In saline sites along the coast and inland, rarely in fens and peaty meadows. The host
plants are unknown, perhaps Juncus gerardii, Puccinellia distans or Briza media. Found in
Germany only very locally, mainly along the coast of the Baltic Sea (westward at least to
Heiligenhafen) and in the rain shadow area to the east of the Harz Mountains (between
Magdeburg, Halberstadt, Bad Frankenhausen and Halle). Also recorded in rather iso-
lated sites near Hamburg, Bad Salzungen, Bad Nauheim and Ansbach (400 m a.s.l.).
Fröhlich (1996a), Kuntze (1937), Schiemenz (1987), Wagner (1935, 1940a), ZIMH, Nickel (1999b), HN; see
also Lauterer (1980)

Javesella forcipata (Boheman, 1847)

E V – E VIII, according to Schiemenz (1987) B V – B IX; nymph, 1 gen.
In cool and moderately eutrophic, temporarily wet or flooded, shady to moderately
shady sites, often with lush grassy vegetation (mountain and fen forests, forest mead-
ows, tall herb stands, spring mires, etc.). Host plants are various grasses (Deschampsia
cespitosa, Holcus mollis, probably also Poa trivialis, Calamagrostis canescens, Agrostis
stolonifera; recorded on Poa annua in the Netherlands). The species is widespread in
Germany, with strongholds in the submontane and montane belt, more scattered in the
lowlands, found to at least 1000 m a.s.l. in the Mittelgebirge, frequently up to 1500 m
a.s.l. in the Bavarian and Allgäu Alps.
Nikusch (1976), Reimer (1992), Schiemenz (1975, 1987), Wagner (1935), Nickel & Achtziger (1999), Nickel &
Remane (1996), Nickel (1997), HN; see also de Vrijer (1986)

Javesella stali (Metcalf, 1943)

So far B V – E VIII; nymph, 2 gen.
This is a pioneer species in stands of horsetail in early successional stages on sandy to
loamy substrates, mainly sand pits and river banks, in southwestern Germany also found
along waysides, in fallow fields and ruderal sites; the host plant is Equisetum arvense. In
Germany only rarely recorded (though probably overlooked) and at the northern and
western edge of the range ; at present there are 16 known localities near Berlin, in the
Spreewald, along the middle course of the Main and the upper Isar (750 m a.s.l.) and
along the southern and middle Rhine (between Freiburg and Bonn). It is also reported
from near Stettin (northwestern Poland) and the eastern half of France (Cahors/Lot,
Savoie, Paris).
Fechtler (pers. comm.), Maczey (pers. comm.), Schiemenz (1987), Nickel & Remane (1996), Nickel (1999a),
HN; also della Giustina (pers. comm.), della Giustina & Remane (1991), Remane & della Giustina (1993),
Wagner (1941a)

Ribautodelphax albostriata (Fieber, 1866)

B IV – B X; nymph, 2 gen.
 Delphacinae Leach, 1815 63

In grass stands usually in sunny, oligotrophic to moderately eutrophic, moderately dry to

damp, occasionally also moist sites (e.g. various types of dry grassland, meadows, ruderal
sites, waysides, forest clearings); single macropterous individuals on dispersal flight, how-
ever, may be found in most terrestrial habitats. Lives monophagously on Poa pratensis (in-
cluding ssp. angustifolia). Widespread in Germany and common in many places, although
apparently absent from the northwest German plain; border localities are near Aachen,
Cologne, Hildesheim, Hamburg and Lübeck. Highest records of brachypterous individuals
are from 950 m a.s.l. on the Swabian Jura, from 1200 m a.s.l. in the Allgäu Alps.
Fischer (1972), Frommer (pers. comm.), Günthart (1988), Reimer (1992), Remane (1987), Schiemenz (1987),
Schwoerbel (1957), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN and others

Ribautodelphax angulosa (Ribaut, 1953)

So far B V – E VIII; nymph, 2 gen.
In sunny, moderately dry, occasionally also moderately moist, usually acidic and olig-
otrophic sites (mainly dry grassland on sandy or siliceous substrates as well as low-
productivity pastures and meadows). The host plant is Anthoxanthum odoratum. De-
spite wide host distribution this species is apparently rare and very localized, but has
often been misidentified. At present there are only 10 verified records from Germany,
most of which have been gathered in or near the northern upper Rhine plain: near Frank-
furt am Main, Wiesbaden, Ebernburg an der Nahe, Speyer and Bruchsal, as well as
Donnersberg near Rockenhausen. Other localities are widely scattered: Oberstdorf/
Allgäu, c. 800 m a.s.l., Höhbeck near Lüchow, Bornhagen near Bad Heiligenstadt, Schkeu-
ditz near Halle, Sprey near Bautzen and Saupsdorf (Elbsandsteingebirge). Records pub-
lished until 1987 may refer to other species and are in need of revision.
Bieman (1987a, and pers. comm.), Wagner (1939a), Walter (pers. comm.), Nickel (1997), HN

Ribautodelphax collina (Boheman, 1847)

E IV – B X; nymph, 2 gen.
Monophagous on Agrostis capillaris in damp to dry, sunny, oligotrophic and usually acidic
sites (dry grassland on sand and other siliceous substrates, abandoned grassland, occa-
sionally low-productivity meadows). The species is widespread and locally common in
the north German plain, but in middle and southern parts of Germany, localities are
rather scattered and largely confined to areas of sand, sandstone and other acidic sub-
strates (mainly valleys of Rhine and Main, Mittelfranken, Thuringia); highest localities
are at 600 m a.s.l. Some of the older published records may refer to R. vinealis Bieman.
Bieman (1987a, 1987b), Remane (1987), Schiemenz (1987), Walter (1999), Nickel & Achtziger (1999), HN

Ribautodelphax imitans (Ribaut, 1953)

M V – E VIII; nymph, 2 gen.
Monophagous on Festuca arundinacea, in temporarily moist to temporarily wet, rather
eutrophic, usually sunny sites, notably meadows, pastures and abandoned grassland.
64 Review of species

The range includes southern and western parts of Europe; in Germany the species is
apparently restricted to the southwest (found in the Kaiserstuhl, Lake Constance Basin,
upper valley of the Neckar near Tübingen, Main and Regnitz valley near Schweinfurt
and Forchheim, as well as middle Hessen near Selters and Marburg), up to at least
400 m a.s.l.
Bieman (1987a, 1987b), Remane & Fröhlich (1994b), Schwoerbel (1957), Nickel & Achtziger (1999), Nickel
& Remane (1996), HN

Ribautodelphax pungens (Ribaut, 1953)

Older records, particularly some from northeastern Germany, are in need of revision and may
refer to other species of the genus.
E III – B X; nymph, 2 gen.
Monophagous on Brachypodium pinnatum in damp to dry, sunny to moderately shady
and usually basic sites; particularly common and often among the dominant species in
calcareous grassland and open pine forests, but some populations are reported to com-
prise varying proportions of triploid, pseudogamous ‡‡. Widespread in middle and
southern parts of Germany, less common and at the edge of the range in the north.
Northernmost localities are on a line from Aachen to Bonn, Bielefeld, Hildesheim,
Parchim, Rostock and Rügen; extending up to 1200 m a.s.l. in the Allgäu Alps.
Bieman (1987a, 1987b, 1988), Reimer (1992), Remane (1987), Rombach (1999b), Schiemenz (1987), Nickel
(1994), HN

Ribautodelphax vinealis Bieman, 1987

In the Netherlands M V – M IX; nymph, 2 gen. (Bieman 1987a).
Monophagous on Agrostis vinealis in sandy dry grassland. Was until recently only re-
ported from the Netherlands. From Germany there are three known localities not high-
er than 150 m a.s.l.: Stolzenau an der Weser, 1951; Quedlinburg, Harslebener Berge,
31.VII.1957; Pevestorf near Schnackenburg an der Elbe, 21. and 29.V.1959. A revision of
older records up to 1987 may reveal more localities.
Bieman (1987a, 1987b, and pers. comm.), Marchand (1953), Remane & Fröhlich (1994b), Nickel (1997)

4.1.3 Achilidae Stål, 1866

Cixidia confinis (Zetterstedt, 1828)

In Sweden according to Ossiannilsson (1978) M V – B VIII; nymph, 1 gen.
Nymphs as well as adults in fissures and under the bark of dead trunks of pine, feeding
on mycelium of fungi. From central Europe there are only two records, both from the
Bavarian Alps in the vicinity of Bad Reichenhall: Jochberg near Weißbach an der Alpen-
straße, 800 m a.s.l., 22.VII.1974, 1 †, and Winkl near Berchtesgaden, 700 m a.s.l.,
22.VIII.1996, 1 †, on a tree trunk in an open pine forest on a southwest-facing slope.
Otherwise known only from Sweden, Finland, Estonia and middle Russia.
 Dictyopharidae Spinola, 1839 65

SMNS (leg. Dynort), Nickel (1999b); see also Anufriev & Kirillova (1998), Linnavuori (1951) (see this paper
also for identification of nymphs), Ossiannilsson (1978)

Cixidia pilatoi D’Urso & Guglielmino, 1995

Populations of this species living in south-central Europe were published as Cixidia marginicollis
(Spinola, 1839) by most authors. The true C. marginicollis (Spin.), however, is so far only known
from Sicily (D’Urso & Guglielmino, 1995), and differences between German specimens and the
newly described C. pilatoi D’Urso & Gugl. from Italy appear to be insignificant.
So far B VI – M VI, in Austria and the Czech Republic until M IX (Holzinger 1995;
Lauterer 1978; Wagner & Franz 1961); nymph, 1 gen.
Known from Germany only from the Kaiserstuhl (Badberg, c. 350 m a.s.l., 10.VI.1952,
2 ‡‡, and 20.VI.1967, 1 †, 1 ‡). Probably lives – like most other species of Cixidia – on
and under the bark of dead trees feeding on hyphae of fungi. Is considered to be associ-
ated with deciduous trees, notably Quercus.
Coll. H.J. Müller, Remane (pers. comm.), Wagner & Franz (1961); see also Holzinger (1995a, 1996), Lauterer
(1978)

4.1.4 Dictyopharidae Spinola, 1839

Dictyophara europaea (Linnaeus, 1767)

Mainly M VII – M X, according to Schiemenz (1987) also E VI; egg, 1 gen.
In sunny, xerothermic sites on various substrates, preferentially in disturbed patches
with some bare ground (oviposition is done into the soil), locally even in dry ruderal
sites. Probably polyphagous, most individuals found on dicotyledonous herbs, but the
species is also reported from dwarf shrubs and even trees. In Germany it lives at the
northern edge of the range, with strongholds in warmer regions in southern and east-
ern parts (rain shadow area to the east of the Harz Mountains, upper Rhine plain, val-
leys of Neckar, Main, Nahe, Danube). Northernmost localities (which appear to be more
or less isolated) are on a line from Bonn to Gießen, Gotha, Sondershausen, Stendal,
Nauen and Stralsund, the highest ones are only at 400 m a.s.l. in the Lake Constance
basin.
Haupt (1924), Müller (1942), Nikusch (1976), Remane (1987), Schiemenz (1987), Schönitzer and Oesterling
(1998a), Schwoerbel (1957), Sergel (1984b), SMNS, Voigt (1978), Wagner (1939a, 1951a), HN

4.1.5 Tettigometridae Germar, 1821

Among central European Auchenorrhyncha the taxon Tettigometra Latr. is probably the most
critical group concerning identity and distinction of species. This is caused by only poor differ-
ences in genitalic morphology and a considerable intraspecific variation of head shape and
body colouration. Life history is another enigma: presumably all – at least central European –
species are associated with ants; the character of these relationships, however, is largely un-
known. Müller (1942) has observed oviposition on the surface of stones, which may, to some
extent, explain the extreme reduction of the female genitalia and the preference of habitats with
bare patches. It has been suggested that the nymphs live in ants’ nests feeding on plant roots
66 Review of species

(Emelyanov 1987), but also nymphal development near the plant base has been observed (see
Müller 1956). Rearing over more generations has failed so far. Hence, species interpretations in
this book may not be identical with those of the original describers, particularly since the where-
abouts of some type material are not known. According to Emelyanov (1980), all species known
from Germany belong to the genus Tettigometra Latr., and Brachyceps Kirk. as well as Mitricepha-
lus Sign. are only subgenera.
The situation among black taxa in central Europe is particularly problematic. Apparently, there
are at least two species in Germany. For this book, they were preliminarily distinguished by the
following characters: larger specimens (5 - 5.5 mm), which are rather broad-built, usually shiny
black, but also blackish- to orange-brownish above (particularly on the vertex), are interpreted
as T. fusca Fieb. The front is usually concave, the vertex showing two lateral bulbosities. In
contrast, smaller specimens (usually 3.3 – 4.3 mm), most of which are blackish, but occasionally
with dark or even pale brown fore wings (particularly after degreasing with benzine) and with
a short, but much variable head, are treated as T. atra Hag. This opinion is based on the study of
museum material from many parts of Europe determined by various authors, but is perhaps not
in accordance with the interpretation of Fieber (1865). It should be stressed that brown-winged
specimens have been labelled and also published as T. atra laetifica Metcalf, 1932, but also as T.
laeta H.-S. It should also be noted that the identity of T. depressa Fieber, 1865 and T. atrata Fieber,
1872 is considered here as dubious.

Tettigometra atra Hagenbach, 1825

T. atra laetifica Metcalf, 1932 (see taxonomic remarks above!)
B VIII – M VI; adult, 1 gen.
In xerothermic, sunny to moderately shady sites on basic to acidic substrates (mainly
dry grassland and margins of pine and oak forests, occasionally also managed mead-
ows). Is reported to live in ant nests (Lasius, Tetramorium). Most adults are swept in the
low vegetation, a few also from woody plants. In Germany at the northern edge of the
range, rather scattered and with strongholds in warmer parts (Saale-Unstrut region,
Rhine-Main region, Mainfranken, Neckar valley, Franconian Jura, slopes of the upper
Rhine valley). Northernmost localities, some of which are not verified, are near Ba-
charach (middle Rhine), Schlüchtern, Jena, Quedlinburg, Neustrelitz and Frankfurt an
der Oder, up to at least 600 m a.s.l. in the foothills of the Alps.
Nikusch (1976), Schiemenz (1987), Schönitzer & Oesterling (1998a), SMNS, Schwoerbel (1957), Wagner
(1939a, 1951a), HN

Tettigometra fusca Fieber, 1865

See taxonomic remarks above! Due to a misunderstanding between the authors, this species
was published for Thuringia as T. atrata Fieb. by Nickel & Sander (1996).
B VIII – M VI; adult, 1 gen.
In spatially diverse dry grassland with open stands of trees or shrubs, usually on basic
substrates on xerothermic plateaus or on upper slopes. Was found in nests of Formica
pratensis Retzius several times. In Germany only very locally and mainly in limestone
areas. Most records (including recent ones) are from the Triassic limestone regions of
eastern Thuringia (near Jena and Arnstadt), Franconia (between Würzburg, Karlstadt
and Bad Kissingen, also near Coburg), the valleys of Tauber and Kocher (Schweinberg
 Tettigometridae Germar, 1821 67

near Buchen, Crispenhofen near Künzelsau). Also published from near Erlangen,
München and Starnberg. The highest sites are situated at 600 m a.s.l. Apparently, many
populations have vanished during recent decades.
Biedermann (pers. comm.), Heller (pers. comm.), Schiemenz (1987), Schönitzer & Oesterling (1998a), SMNS,
Trümbach (1959), Wagner (1951a), ZIMH, ZMB, Nickel & Sander (1996), Nickel & Remane (1996)

Tettigometra griseola Fieber, 1865

So far B VIII – M VI; adult, 1gen.
Usually in xerothermic sites with at least scattered shrubs or trees, mainly on limestone,
loess and schist; another record is from a seepy, temporarily wet slope. In Germany
known only from 5 localities in the Kaiserstuhl area (Schelingen, Vogtsburg, Istein),
along the middle Rhine (Lorch) and in the foothills of the Alps (Fischen am Ammersee),
up to at most 600 m a.s.l.
Remane (pers. comm.), Wagner & Franz (1961), HN

Tettigometra impressopunctata Dufour, 1846

Populations living in Germany may belong to at least two different species. Moreover, there are
older reports of T. depressa Fieber, 1865, T. atrata Fieber, 1872, and others, the identity of which is
unclear and which are perhaps conspecific with T. impressopunctata Duf.
Mainly B VIII – B VI, occasionally also VII; adult, 1 gen.
Along sunny woodland margins in dry to temporarily dry, basic to acidic, usually rath-
er warm sites, mainly in grazed dry grasslands, along spatially diverse forest margins
and on gravel banks of alpine rivers with scattered shrubs. There are also older records
from fenland habitats in the foothills of the Alps. Adults are found among herbaceous
vegetation as well as on shrubs and trees (Quercus, Taxus, Myricaria and others). In Ger-
many at the northern edge of the range, presumably declined in recent decades. Most
localities are in the foothills of the Alps and in warmer regions of southern and eastern
parts (valleys of Rhine, Main and Neckar, as well as in Thuringia), northernmost sites,
which appear to be more or less isolated, are near Siegen, Göttingen, Bad Frankenhau-
sen and Magdeburg. The highest known sites are at 900 m a.s.l. in the Bavarian Alps, at
1200 m a.s.l. in Switzerland.
Schiemenz (1987), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Wagner (1939a, 1951a), Nickel (1994),
HN; see also Günthart (1987a)

Tettigometra laeta Herrich-Schäffer, 1835

Has been confused with brown-winged specimens of T. atra Hag. (= T. atra var. laetifica Metc.;
see above).
This is apparently a Mediterranean species, which is reported to live in ants’ nests, but no
further details are known on the life history. The only record from Germany is the original
description: “Bei Regensburg selten”, but the type material is unfortunately lost.
Herrich-Schäffer (1835a); see also Kirschbaum (1868), Metcalf (1932), Wagner (1939a)
68 Review of species

Tettigometra leucophaea (Preyssler, 1792)

Tettigometra obliqua (Panzer, 1799)
Mainly B VIII – M VI, occasionally also VII; adult, 1 gen.
This species is published from various, usually spatially diverse xerothermic sites, of-
ten with scattered trees or shrubs, bare sand or rocks. In older papers, notably from the
beginning of the 20th century, mass occurrences have been reported with subsequent
damage to rye, wheat and oats, but the species has declined dramatically in Germany,
as in most parts of central Europe. Formerly it was widespread in middle and southern
parts and has occurred northward at least to Frankfurt am Main, Lauenburg, Lud-
wigslust, Waren an der Müritz and Angermünde, extending to at least 500 m a.s.l. in the
foothills of the Alps. The current status is uncertain; the latest records are from the
1960s and 1970s (Saale valley near Jena, lower Oder valley near Seelow and Bad Freien-
walde). Many of the older localities have been checked since without positive records.
Thus, the species is probably extinct in Germany.
Fischer (1972), Kirschbaum (1868), Kuntze (1937), Lindberg (1948), Remane (1987), Schönitzer & Oester-
ling (1998a), Schiemenz (1987), Schwoerbel (1957), SMNS, Torka (1905), Wagner (1935, 1939a, 1951a, and
unpublished data)

Tettigometra macrocephala (Fieber, 1865)

E VII – E VI; adult, 1 gen.
Along xerothermic and spatially diverse woodland margins in basic as well as acidic
sites, mostly in vine growing regions. In Germany known only from a few isolated lo-
calities at the northern edge of the range and apparently declining, altogether known
from 17 sites, mainly in Triassic limestone areas of eastern Thuringia, Franconia and
northern Baden (near Jena, Bad Kissingen, Hammelburg, Karlstadt, Tauberbischofsheim),
but in the Rhine-Nahe region (Schloßböckelheim, Bad Münster, Wöllstein, Lorch) also
on porphyry, Keuper and other substrates; also found along the upper courses of the
Main, Neckar and Danube (Bamberg, Tübingen, Ulm), between 150 and 500 m a.s.l.
Biedermann (pers. comm.), Dynort (pers. comm.), Lindberg (1948), Schiemenz (1987), Schwoerbel (1957),
SMNS, Wagner (1939a, 1951a), HN

Tettigometra virescens (Panzer, 1799)

Most vivid green individuals of Tettigometra smaller than 5 mm in central Europe have been
labelled as T. virescens (Panz.). However, they have also been confused with T. laeta H.-S. and
perhaps other species. Furthermore, the identity of numerous varieties mostly described from
southern Europe (see Metcalf 1932) is not clear. In contrast, the morphological variation of spec-
imens originating from Germany and adjacent areas is surprisingly low.
E VIII – M VI; adult, 1 gen.
Along woody margins in xerothermic sites on basic to acidic substrates (mainly Triassic
limestone, gypsum, schist). Most adults are swept from low vegetation, in Poland also
from Pinus and Juniperus. Rare in Germany and at the northern edge of the range; only
known from a few localities, usually on upper slope and plateau situations: Saale valley
 Caliscelinae Amyot & Serville, 1843 69

(near Jena), Thuringian Basin (Gotha), Main and Tauber region (near Frankfurt am Main,
Tauberbischofsheim, Karlstadt, Hammelburg, Bad Kissingen, Bad Königshofen) and the
valley of the middle Rhine (near Bacharach), between 150 and 350 m a.s.l.
Biedermann (pers. comm.), Haupt (1924), Müller (1978), Reimer (1992), SMNS, Wagner (1939a, and unpub-
lished data), Nickel (1999b), HN; see also Smreczynski (1954)

4.1.6 Issidae Spinola, 1839

4.1.6.1 Caliscelinae Amyot & Serville, 1843

Ommatidiotus dissimilis (Fallén, 1806)

Older German reports of O. concinnus Horváth, 1905, a species otherwise published only from
Hungary (see Nast 1987), are dubious and probably refer to O. dissimilis (Fall.).
B VII – M X; egg, 1 gen.
In raised and intermediate bogs on Eriophorum vaginatum. In Germany only scattered,
but locally abundant, mainly in the north German plain and along the edge of the Alps,
also in the Black Forest; apparently absent from most regions of the Mittelgebirge. The
highest localities are at 900 m a.s.l.
Fischer (1972), Remane (1958), Schiemenz (1976, 1987), Wagner (1935), HN

4.1.6.2 Issinae Spinola, 1839

Issus coleoptratus (Fabricius, 1781)

Mainly E VI – M X, in warmer regions from E V onwards; nymph, 1 gen.
On woody plants in damp to dry sites, in closed deciduous and mixed forests as well as
in semi-open tree stands (e.g. along forest margins, in parks, cemeteries, etc., also on
solitary trees on grassland). Adults on various deciduous trees (mainly Quercus and
Acer, also Corylus, Betula, Ulmus, Tilia, Sorbus, Clematis and others), hibernating nymphs
found on Hedera, Ligustrum, Juniperus and Taxus, as well as in the litter layer. In Germa-
ny the species occurs at the northern limit of the range, which runs approximately along
the northern edge of the Mittelgebirge (northernmost records near Cologne, Northeim,
Quedlinburg, Leipzig and Kamenz); densely spread to the south of this line, although
sweep net catches usually contain only low numbers. Frequently found up to 1000 m
a.s.l. in the Bavarian Alps.
Achtziger (1991), Büchs (1988), Fischer (1972), Frommer (1996), Post-Plangg & Hoffmann (1982), Schönitzer
& Oesterling (1998a), Schwoerbel (1957), Wagner (1939a, 1951a), Nickel (1997), HN

Issus muscaeformis (Schrank, 1781)

So far E V – E VIII; nymph, 1 gen.
Apparently not ecologically distinct from I. coleoptratus (F.), although extending further
northward in Europe. Found on Quercus and Corylus along sunny forest margins and on
70 Review of species

solitary trees and shrubs. Is reported from almost the whole of the European deciduous
forest zone, but recorded from Germany from 9 localities only: upper and lower course
of the Elbe (Pirna, Harburg, and Winsen an der Luhe), Bergisches Land (Hückeswagen),
Fulda valley (Kassel), Saale-Unstrut region (Freyburg), Bavarian Forest (Hochobern-
dorf and Ulrichsberg, both near Deggendorf) and Berchtesgaden Alps (St. Bartolomä),
extending to at least 700 m a.s.l.
Emmrich (pers. comm.), Remane (pers. comm.), Coll. Sander, SMNS, Wagner (1935, and unpublished data),
HN; see also Holzinger (1996b), Ossiannilsson (1978), Wagner & Franz (1961)

Agalmatium bilobum (Fieber, 1877)

This is a Mediterranean species, known from Germany only from a single record on the
East Frisian Islands: Wangerooge, 17.VI.1960, in white dunes „eine gute Population“,
which was no more found in the 1980s. Hence, this was apparently a temporary occur-
rence.
Harz (1965, 1988), Niedringhaus (1991), Remane & Fröhlich (1994a)

4.2 Cicadomorpha Evans, 1946 – Leafhoppers (s.l.)

4.2.1 Cicadidae Leach, 1815 (s.l.) – Cicadas

Cicada orni Linnaeus, 1758

VI – B IX; nymph, development takes about 4 years (Schedl 2000).
This species is widespread and common in Mediterranean regions. Obviously, there are
occasional influxes into viticultural regions of southwest Germany. Two more recent
records are based on observations only, but the species can be easily identified: Stutt-
gart, 30.VII. -12.VIII.1986, 1 † singing; Öhringen, 05.VII.1997, 1 ‡, 3 weeks later song
heard. Furthermore, there is a record from the Pliocene of Willershausen (southern Nie-
dersachsen). Nymphs and adults are reported to be polyphagous, adult feeding has
been recorded on Pinus, Pistacia, Olea, Fraxinus and others.
Dynort (pers. comm.), Heller (1987c), Wagner (1967); see also Boulard & Mondon (1995), Schedl (2000)

Tibicina haematodes (Scopoli, 1763)

M IV – B VIII; nymph, development takes several years, three according to Schedl (2000).
In dry grassland and abandoned vineyards with scattered shrubs and trees, usually in
xerothermic upper slope situations on limestone and Triassic sandstone. According to
older reports, the nymphs feed on roots of Vitis vinifera and Prunus spinosa, oviposition
takes place on the latter. Adults are usually found in the tree and shrub layer and are
reported from Pinus sylvestris, Quercus pubescens, Fraxinus excelsior, Acer spp., Prunus
spinosa and others. Single individuals may perform long dispersal flights and often land
 Cicadidae Leach, 1815 (s.l.) – Cicadas 71

in exposed sites such as hilltops, solitary trees and even vineyard workers. In Germany
the species is restricted to isolated localities in viticultural regions in southwestern parts
between 100 and 350 m a.s.l., but many records date back before 1940. The strongholds
are in Rheinhessen (between Bad Kreuznach, Alzey and Nierstein – also recently), the
Main valley near Volkach and between Karlstadt and Ochsenfurt (all recently), as well
as in the valleys of Neckar and Enz and in the adjacent Stromberg and Kraichgau (be-
tween Ludwigsburg, Pforzheim, Sinsheim and Lauffen, also near Tübingen – although
recently reported only from near Pforzheim). Furthermore, there is a number of older
records, some of which are not verified or which are based on single specimens only,
notably from the Ruwer valley near Trier, the Palatinate Haardt near Grünstadt, and the
Rheingau near Lorch. Is also reported from the Pliocene of southern Niedersachsen
(Willershausen).
Back (1976), Eitschberger (1972), Hess (pers. comm.), Niehuis (1995), Niehuis & Simon (1994), Schedl (2000),
Schwoerbel (1957), SMNS, Stadler (1922), Vogel (1935, 1937), Wagner (1967), HN; see also Boulard & Mon-
don (1995)

Cicadetta montana (Scopoli, 1772)

According to Gogala & Trilar (1998, 1999) this is a complex of several species, which are distinct
mainly bioacoustically. Central European populations probably belong to at least two species.
The song of the more widespread form is long-lasting and continuous, with only short and
irregular interruptions. Another form is apparently rare and known from Germany only from a
few localities at the northern edge of the Thuringian Forest (Tieftal near Arnstadt) and the Tau-
ber valley near Tauberbischofsheim and Bad Mergentheim. Its song consists of two different and
alternating echemes, the first of which lasts several seconds, the second is very short. However,
it should be noted, that in many localities the type of song has not been recorded. Thus, the
short-singing form is perhaps more widespread. For French populations with a short song, Bou-
lard & Mondon (1995) use the name C. petryi (Schuhmacher, 1924), which has been created for
specimens from the Kyffhäuser in Thuringia, based on a slightly different colouration. Howev-
er, it is uncertain whether this name is correct. All Kyffhäuser populations recorded during
recent years clearly performed the long-lasting and continuous type of song, and morphologi-
cal characters for distinction between the two groups have not yet been found (Gogala & Trilar,
pers. comm.). See also Schedl (2000).
M V – M VII; nymph, development takes several years.
Among open stands of shrubs or trees in moderately dry to dry, sun-exposed sites, usu-
ally on hillsides or plateaus, mainly in dry pastures and meadows and along sunny
forest margins. Widespread but scattered in the southern and middle parts of Germany,
with strongholds in viticultural regions in the southwest and in the rain shadow area to
the east of the Harz Mountains, usually below 500 m a.s.l., but frequently at 750 m a.s.l.,
along the upper course of the Isar and on the Swabian Jura, locally even up to 1000 m
a.s.l. in the Black Forest. The northernmost localities are near Ahrweiler, Beckum/West-
phalia, Ibbenbüren, Holzminden, Göttingen, Nordhausen, Quedlinburg, Naumburg an
der Saale, Finsterwalde and Fürstenwalde. The species is absent from Denmark, but
occurs very locally in southern Scandinavia.
Andres (pers. comm.), Ant (1991), Back (1976), Bernhardt (1991), Bußmann & Feldmann (1995), Eitschber-
ger (1972), Emmrich (1984), Felzmann (pers. comm.), Hoffmann (1991), Kless (1971), Niehuis & Weitzel
72 Review of species

(1996), Reimer (1992), Schedl (2000), Schiemenz (1988), Schönitzer & Oesterling (1998a), Schwoerbel (1957),
Stein & Bogon (1990), Vogel (1935), Wolf (1976), Nickel (1994), HN and others; see also Ossiannilsson (1981)

4.2.2 Cercopidae Leach, 1815 (s.l.) – Froghoppers

4.2.2.1 Cercopinae Leach, 1815

Cercopis arcuata Fieber, 1844

In eastern Austria B V – B VII (Holzinger 1996b); nymph, 1 gen.
In Germany known only from 4 records from the Alps and their foothills, all probably
dating before 1950: Schellenberg, 03.VI.1938, and Almbachklamm, 05.VI.1938, both in
the vicinity of Berchtesgaden (ssp. haupti Priesner, 1923), also „Bayerisches Allgäu“ and
Geiselgasteig to the south of Munich, c. 550 m a.s.l. In warmer regions of the Czech
Republic, Austria and France this species lives in pine or oak forests; most adults were
found on dicotyledonous herbs.
Wagner & Franz (1961), ZIMH, ZSM; see also Holzinger (1996b), Lauterer (pers. comm.)

Cercopis sanguinolenta (Scopoli, 1763)

B V – E VII; nymph, 1 gen.
In sunny, moderately dry to dry sites on basic to acidic substrates in warm situations,
notably in dry grassland and abandoned vineyards, preferably in disturbed patches
with tall, but scattered vegetation. Food plants are various grasses and herbs; most feed-
ing adults are reported from Arrhenatherum elatius, also on Salvia pratensis, Brachypodi-
um pinnatum, Falcaria vulgaris, Cardaria draba, Galium verum, Thalictrum minus and oth-
ers. Germany is at the northern edge of the species’ range, with only isolated popula-
tions, mainly in the rain shadow area to the east of the Harz Mountains (between Sonder-
shausen, Halle, Leipzig, Eisenberg, Jena and Kölleda, as well as up along the Saale to
Rudolstadt) and in the Rhine-Nahe region (Bacharach, Schloßböckelheim, Bad Mün-
ster). A few isolated localities are situated in the Elbe valley near Meißen and the foot-
hills of the Alps near Starnberg, c. 600 m a.s.l. In the northeastern Alps of Styria (Aus-
tria) the species is less xerophilous and frequently occurs in the upper montane belt,
locally up to 1600 m a.s.l.
Biedermann (1998), Emmrich (1983), Jöst (1966), Reimer (1992), Remane & Wachmann (1993), Schiemenz
(1988), Schönitzer & Oesterling (1998a), HN; see also Wagner & Franz (1961)

Cercopis vulnerata Rossi, 1807

In the lowlands B V – M VII, at higher altitudes VI – E VIII; nymph, 1 gen.
In moderately dry to moderately wet, sunny to moderately shady sites, mainly aban-
doned grassland, forest glades, open forests, along waysides, streams and ditches, ab-
sent from mown meadows. Adults mainly on various tall herbs and grasses (Urtica,
Aegopodium, Lupinus, Chrysanthemum, Filipendula, Arrhenatherum and others); the only
 Cercopinae Leach, 1815 73

published records of nymphal feeding (from Switzerland) are from Arrhenatherum ela-
tius and Dactylis glomerata. Widespread in Germany and common in southern and cen-
tral parts, particularly in the lowlands, although frequently found in higher parts of the
Mittelgebirge, up to at least 1600 m a.s.l. in the Bavarian Alps. Rather scattered in north-
ern Germany and at the edge of the range, extending at least to the East Frisian Islands,
Bremerhaven, Itzehoe, Ratzeburg, Rostock and Strasburg.
Fischer (1972), Kehlmaier (2000), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Wagner (1935,
1939a), HN and others; see also Mauri (1982)

Haematoloma dorsatum (Ahrens, 1812)

E IV – E VII, occasionally until E VIII; nymph, 1 gen.
Adults on pine (Pinus sylvestris, additional species, notably Pinus nigra, in more south-
ern and western parts of Europe) in open forests in damp to dry sites, usually on
limestone or sandy soils. Nymphs live in the litter layer and feed on fine roots and
basal stems of grasses (preferentially Deschampsia flexuosa on acidic substrates, prob-
ably Brachypodium pinnatum and others on basic ones). Numbers of swept adults may
be large in grassy forest floor vegetation after emergence in May, but rather small in
the lower part of the tree layer from June onwards. Here they feed on pine needles
and mate; oviposition takes place in the grass layer. In the Netherlands and the Med-
iterranean region there may be local damage in pine plantations. In Germany at present
the species occurs only in the western half (mainly below 400 m a.s.l., but to at least
700 m a.s.l. on the Swabian Jura) and apparently is extending its range: collected for
the first time in 1935 near Bonn, from 1936 onwards in the Rhine-Nahe region, from
where it was published as new to Germany, recorded in 1952 near Karlsruhe, in 1955
in the Kaiserstuhl, since 1956 along the northeastern edge of the Black Forest near
Calw, later in large parts of Swabia (eastward at least to Schwäbisch Gmünd and Bad
Urach), since 1969 in middle Hessen and in the Weser-Ems area, since the mid 1970s
in the middle Main region between Karlstadt, Würzburg, Kitzingen and Schweinfurt,
as well as along the Fränkische Saale, at least since 1984 in the foothills of the Alps of
Upper Swabia north of Ravensburg, found in 1987 on the East Frisian island of Bor-
kum, later also on Norderney, since 1989 in the Senne (Westphalia), in 1993 along the
upper Main near Staffelstein, in 1996 near Lake Constance (Sipplingen), in 1997 in the
Lüneburger Heide to the south of Fallingbostel, 2001 in the Grabfeld of southern Thu-
ringia, and 2002 in the Altmark (Salzwedel, Klötze). Apparently, pine forest areas in
central Thuringia, the Wendland of Niedersachsen and in Mecklenburg, which have
been investigated during recent years, were not yet colonized.
Achtziger (pers. comm.), Bernhardt & Arnold (1991), Bußmann & Feldmann (1995), Fritzlar (pers. comm.),
Gauss (1987), Haeseler & Niedringhaus (1988), Heller (1960), Hess (pers. comm.), Jöst (1966), Kupka (1941),
Niedringhaus & Olthoff (1993), Nikusch (1976), Remane (1987), Remane & Wachmann (1993), Schulze
(1992), SMNK, SMNS, Voigt (pers. comm.), Wagner (1939a), Witsack (pers. comm.), ZFMK, Nickel & Re-
mane (1996), HN; see also Holzinger (pers. comm.), Moraal (1996)
74 Review of species

4.2.2.2 Aphrophorinae Amyot & Serville, 1843

Lepyronia coleoptrata (Linnaeus, 1758)

E V – M X; egg, 1 gen.
Usually in sunny, dry to moist, occasionally also temporarily moist or even wet sites, mainly
ruderal habitats, waysides, dunes, dry grassland (often in disturbed patches) and low-in-
put meadows; also reported from saline sites near the coast of the Baltic Sea. Adults often on
grasses; in the foothills of the Alps, nymphs were found on Equisetum, Pedicularis, Eupatori-
um, Solidago, Artemisia, Carex, Phragmites, Menyanthes and various umbelliferous herbs, in
Sweden on Vaccinium, Polygonum, Anthriscus, Chamaenerion, Potentilla, Hypericum, Ranuncu-
lus, Plantago, Galium, as well as shrubs and trees (Salix, Populus, Betula, Corylus). Widespread
in Germany, locally common in southern and eastern parts, but otherwise rather scattered
or rare, extending to at least 1250 m a.s.l. on sun-exposed hillsides in the Mittelgebirge and
along the edge of the Alps. Apparently absent from northwestern parts and extending at
least to Bonn, Marburg, Eschwege, Nordhausen, Magdeburg, Lüchow and Bad Oldesloe.
Fischer (1972), Kuntze (1937), Heller (1987a), Nickel (1979), Nikusch (1976), Post-Plangg & Hoffmann (1982),
Reimer (1992), Remane (pers. comm.), Schiemenz (1988), Schönitzer & Oesterling (1998a), Schwoerbel (1957),
Wagner (1939a, 1951a), Nickel & Achtziger (1999), Nickel (1997), HN; see also Ossiannilsson (1981)

Neophilaenus albipennis (Fabricius, 1798)

Mainly B VI – B IX, occasionally E V – B X; egg, 1 gen.
Monophagous in stands of Brachypodium pinnatum in moderately dry to damp, also
temporarily moist, usually slightly shady and basic sites, notably in dry grassland and
open stands of oak and pine. Widespread and locally common in limestone areas in
middle and southern parts of Germany, but with distribution border approximately
along the northern edge of the Mittelgebirge. Northernmost localities are near Bonn,
Höxter, Wolfenbüttel, Halberstadt, Halle, Gera and Görlitz. In the Bavarian and Allgäu
Alps it frequently occurs up to at least 1200 m a.s.l., single individiuals were even re-
corded at 1600 m a.s.l. This is one of the very few Auchenorrhyncha species of which
metapopulation dynamics have been studied in detail.
Biedermann (1997, 2000), Deutschmann (pers. comm.), Rabeler (1962), Reimer (1992), Rombach (1999a),
Schiemenz (1988), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Nickel (1994), HN

Neophilaenus campestris (Fallén, 1805)

E V – B X; egg, 1 gen.
In ruderal or grazed, usually moderately dry and sunny sites on various substrates
(sandy to clayey, basic to acidic), mainly grassy ruderal sites, mining areas, waysides
and military training areas, also grazed or trampled patches in dry grassland and in-
land dunes. Host plants are various grasses (Agrostis capillaris, probably Elymus repens,
Arrhenatherum elatius, Poa compressa and others), some adults ascend onto Pinus and
other woody plants during hot summer days. Widespread in the lowlands of Germany
(below 600 m a.s.l.), but usually at low densities, with strongholds in warmer regions.
 Aphrophorinae Amyot & Serville, 1843 75

Neither recorded to the northwest of a line running from Düsseldorf to Osnabrück,

Hamburg and Lübeck, nor to the south of the Danube.
Reimer (1992), Remane (1987), Schiemenz (1988), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), Nickel
(1997), HN; see also Badmin (1997)

Neophilaenus exclamationis (Thunberg, 1784)

According to Wagner (1955) three subspecies occur in Germany: ssp. exclamationis (Thunberg, 1784)
in the north German plain, ssp. monticola W. Wagner, 1955 in the middle parts (up to at least
1000 m a.s.l.) southward to the edge of the Alps and ssp. alpicola W. Wagner, 1955 in the Bavarian
and Allgäu Alps above 600 m a.s.l. The exact borders between their ranges are unclear, however.
E V – B X; egg, 1 gen.
In open forests and along their margins (mostly under pine or oak), usually in oligotrophic,
damp to moderately dry, basic as well as acidic sites, also in grassland of the subalpine
and alpine belt of the Alps. In most lowland regions the main host plant is clearly Festuca
ovina, but the species may also live on Deschampsia flexuosa and additional grasses. It is
widespread in Germany, but uncommon, frequently recorded in the Bavarian and Allgäu
Alps at 2000 m a.s.l., in Austria and Switzerland up to at least 2500 m a.s.l.
Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998a), Wagner (1935, 1939a, 1955), HN; see
also Günthart (1987a), Leising (1977), Whittaker (1965)

Neophilaenus infumatus (Haupt, 1917)

E V – B X; egg, 1 gen.
In sun-exposed, usually low-growing grass stands with incomplete cover of vegetation in
dry to moderately dry sites. Host plants are usually (perhaps exclusively) greyish-green
subspecies of the Festuca ovina group. In Germany this species is restricted to steppe sites
on gypsum and porphyry of Thuringia and Saxony-Anhalt. These localities are at the
western edge of the range, although the species is locally dominant. Recorded between
Halberstadt, Bernburg, Halle, Jena, Erfurt, Gotha and Sondershausen, up to 350 m a.s.l.
Single specimens (perhaps vagrants) were also found in Brandenburg and Saxony. It is
reported up to 1600 m a.s.l. in dry valleys of the central Alps of Austria and Switzerland.
Schiemenz (1969, 1988), HN; see also Günthart (1987a, 1997), Wagner & Franz (1961)

Neophilaenus lineatus (Linnaeus, 1758)

According to Remane & Wachmann (1993) this may be a complex of several species. In Germa-
ny, besides ssp. lineatus (Linnaeus, 1758), there are populations, which have been described as
Philaenus lineatus pallida Haupt, 1917 (on dunes and in other types of dry sandy grassland of
northwestern Germany) and Philaenus aterrimus J. Sahlberg, 1871 (in raised bogs of the Mittelge-
birge and the Alps). Ossiannilsson (1981) and most other authors consider them both as ecolog-
ical forms of N. lineatus (L.).
M VI – B XI; egg, 1 gen.
Eurytopic usually in tall stands of Poaceae and Cyperaceae in sunny to shaded, wet to
moderately dry sites, mainly in bogs of various types, low-input meadows and pas-
76 Review of species

tures, ruderal sites, open forests, salt marshes along the coast and inland, and coastal
dunes. Widespread and common all over Germany, frequently found up to 1600 m a.s.l.,
occasionally at 1800 m a.s.l. in the Bavarian Alps. Host plants are species of Poaceae
(Calamagrostis spp., Deschampsia spp., Festuca spp., Agrostis spp., Holcus spp., Ammophi-
la arenaria and others), Cyperaceae (Carex spp., Trichophorum cespitosum and others),
probably also Juncaceae (Juncus spp.) and other families.
Fischer (1972), Nickel (1979), Niedringhaus (1991), Reimer (1992), Remane & Wachmann (1993), Schiemenz
(1988), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN and others; see also Halkka et al.
(1977), Novotný (1995), Whittaker (1965a, 1965b, 1971)

Neophilaenus minor (Kirschbaum, 1868)

E V – E X; egg, 1 gen.
In sunny, oligotrophic, moderately dry to dry sites on various substrates (acidic to ba-
sic, preferably sand, but also on solid rock), with sparse cover of vegetation. Host plants
are mainly Corynephorus canescens, Festuca ovina, Koeleria glauca, and probably addition-
al species of fine-leaved grasses. Widespread in the sand areas of the north German
plain (northward at least to Pinneberg and Eutin) and often among the dominant spe-
cies in heaths and Corynephorus swards, but rather scattered to the south, where largely
restricted to inland dune areas and dry hillsides; highest localities are at 500 m a.s.l. in
the foreland of the Alps, at 600 m a.s.l. in the Mittelgebirge. Not reported from Denmark,
Norway and Sweden, but from southeastern Finland, Latvia and Lithuania.
Frommer (pers. comm.), Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987,
and pers. comm.), Schiemenz (1988), Schönitzer & Oesterling (1998a), Wagner (1935, 1951a), HN; see also
Ossiannilsson (1981)

Aphrophora alni (Fallén, 1805)

B VI – E X; egg, 1 gen.
Eurytopic among various stands of woody plants in wet to dry sites, usually along
shores of lakes, ponds, rivers, streams and hedges, as well as in parks, forests (mainly
along edges and roads) and xerothermic sites with scattered shrubs. Adults are usually
swept from deciduous shrubs and trees (Salix, Alnus, Populus, Betula, Prunus spinosa,
Corylus, Rosa, Cytisus and many others), nymphs in Scandinavia were found close to the
ground on various herbaceous plants (Polygonum, Filipendula, Trifolium, Angelica, Gali-
um, Ranunculus, Hieracium, Viola and others), also on adventitious shoots of Salix, Betula
and Alnus. Widespread and common in most parts of Germany, frequently occurring
up to at least 1500 m a.s.l. in the Mittelgebirge and the Alps.
Achtziger (1991), Fischer (1972), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz (1988),
Wagner (1935, 1939a), HN and others; see also Halkka et al. (1977), Ossiannilsson (1981)

Aphrophora major Uhler, 1896

Aphrophora alpina Melichar, 1900 sec. Anufriev & Emelyanov (1988), Komatsu (1997b)
M VII – E IX; egg, 1 gen.
 Aphrophorinae Amyot & Serville, 1843 77

In moist to wet sites with scattered trees or shrubs and along forest margins, mainly in
fens and intermediate bogs, along the edge of raised bogs and in cool meadow valleys.
Adults on Betula and Salix, but also in stands of tall sedges and reeds. The nymphal host
plants are not properly known; Phragmites and Pedicularis have been recorded in Bavar-
ia, Salix and Artemisia in Japan. Thus, it is likely that adults after emergence ascend
from the herbaceous layer to shrubs and trees, as in A. alni (Fall.). The species is rather
localized in Germany, it is usually collected in low numbers only. Most localities are in
the Bavarian and Swabian foothills of the Alps (between 500 and 1000 m a.s.l.) and in
the northwest German plain (near Lingen, Meppen, Vechta, Cloppenburg, Hanover,
Bremen, Pinneberg). Furthermore, there are a few localities along the upper and middle
Rhine (Germersheim, Düsseldorf, Cologne), as well as an unverified record from near
Berlin. In Poland it is only known from northeastern parts, in France only from the
north; it is not reported from Fennoscandia, Denmark and Belgium.
Frommer (1996), Haupt (1924), Jöst (1966), Mölleken & Topp (1997), Nickel (1979), Niedringhaus & Olthoff
(1993), Schiemenz (1988), Schönitzer & Oesterling (1998a), SMNS, ZMB, Nickel (1999b), HN; see also della
Giustina & Remane (2001), Komatsu (1997b), Nast (1976a, 1987)

Aphrophora corticea Germar, 1821

M VI – M X; egg, 1 gen.
In forests of Pinus sylvestris and along their margins, in temporarily moist to dry, basic
as well as acidic sites. Nymphal development takes place among low vegetation on
pine saplings, on Vaccinium myrtillus, Calluna vulgaris and other woody plants, proba-
bly also on herbs. After emergence, adults migrate up into the canopy layer, but ovipo-
sition presumably takes place near the ground. Widespread in Germany, but locally
absent, with strongholds in the pine forest areas of Jurassic and Triassic limestone, sand-
stone and aeolian sand, as well as in the north German plain. Occurs up to at least 1400
m a.s.l. in the Bavarian Alps.
Fischer (1972), Schiemenz (1988), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Wagner (1935, 1939a),
HN and others

Aphrophora pectoralis Matsumura, 1903

Aphrophora salicis forneri Haupt, 1919; Aphrophora costalis Matsumura, 1903 sec. Komatsu (1997b)
Recently some material of the extensive Matsumura collection was revised by Komatsu (1997a,
1997b, 1997c). A. costalis Mats., which was in use in the European literature since Vilbaste (1968b),
was declared a junior synonym of A. pectoralis Mats.
B VI – E IX; egg, 1 gen.
Along shores of running and standing water, in meadow valleys, forest glades and other
cool sites; adults and nymphs live on various species of willows (mainly Salix caprea, S.
purpurea, S. cinerea, S. aurita, locally S. viminalis, S. alba, S. myrsinifolia and others). Wide-
spread in Germany, but usually in low to intermediate frequency, with strongholds in cool
situations in the Mittelgebirge and along the edge of the Alps, where extending up to at
least 1200 m a.s.l. Is not recorded from Mecklenburg-Vorpommern, the northern half of
Poland and Denmark, but from large parts of Fennoscandia and the Baltic countries.
78 Review of species

Frommer (1996), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998a),
Wagner (1935, 1939a), HN

Aphrophora salicina (Goeze, 1778)

Aphrophora grisea Haupt, 1919; Aphrophora salicis unicolor Haupt, 1919
Many published reports, including recent ones, are in need of revision due to former taxonomic uncer-
tainties, which were only resolved by Wagner (1948c) in a paper, which has received little attention.
B VI – B X; egg, 1 gen.
Adults and nymphs on Salix alba and S. purpurea, less frequently on S. viminalis, S. ci-
nerea, S. fragilis, a handful of records also from S. aurita, S. caprea, S. triandra and others.
A few nymphs (probably fallen individuals) have been found in the herb layer. Wide-
spread in Germany and usually in intermediate frequency, but more confined to low-
lands and the larger river valleys than A. pectoralis Mats. Usually found below 600 m
a.s.l., locally up to 850 m a.s.l.
Fischer (1972), Heller (1987a), Lauterer & Malenovsky (1995), Niedringhaus (1991), Reimer (1992), Schönitzer
& Oesterling (1998a), Schwoerbel (1957), Wagner (1935, 1939a), HN

Philaenus spumarius (Linnaeus, 1758)

E V – B XI; egg, 1 gen. (see also Witsack 1973)
Extremely eurytopic and polyphagous. Adults and spittle masses usually on dicotyle-
donous herbs (but also on grasses, ferns, horsetails, dwarf shrubs, and even adventi-
tious shoots of shrubs and trees) in peaty to dry, sunny to moderately shady sites such
as meadows, pastures, fens, open forests, shores of running and standing water, way-
sides, ruderal sites, dry grassland (where usually confined to margins and disturbed
patches), primary dunes, salt marshes, subalpine stands of herbs, etc. Often abundant
in low-input meadows and abandoned fields with dominating dicotyledonous herbs,
where spittle masses may be extremely abundant on Silene flos-cuculi, Cirsium arvense,
Urtica dioica, Ranunculus repens, Filipendula ulmaria and others. Also found in low num-
bers on plant taxa largely avoided by other Auchenorrhyncha, e.g. Rubiaceae, Boragi-
naceae, Primulaceae, Brassicaceae and Orchidaceae, but usually rare or absent in inten-
sively managed grassland, where reproductive success is uncertain. Widespread and
common in Germany, highest localities at 1800 m a.s.l. in the Bavarian and Allgäu Alps.
Achtziger (1991), Fischer (1972), Nickel (1979), Niedringhaus (1991), Reimer (1992), Remane & Wachmann
(1993), Schiemenz (1988), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN and others; see also Halkka
(1978), Halkka et al. (1967, 1978), Stewart & Lees 1996, Wiegert (1964), Whittaker (1968, 1973)

4.2.3 Membracidae Rafinesque, 1815 – Treehoppers

Gargara genistae (Fabricius, 1775)

E VI – M X; egg, 1 gen.
In sunny to moderately shady, damp to dry sites on acidic to basic substrates. Adults
and nymphs usually stay in small aggregations on woody species of Fabaceae and are
 Membracidae Rafinesque, 1815 – Treehoppers 79

attended by ants. The northern edge of the range runs through Jutland, but in Germany
the species is widespread, though scattered, with strongholds both in areas of sand and
sandstone (where usually occurring in intermediate to high frequency on Cytisus sco-
parius in heaths, on roadside embankments and along forest margins) and on sun-ex-
posed, acidic to basic sites in warm regions of southern and eastern Germany (where
found in low to intermediate frequency, mainly on Ononis, Genista and Colutea arbore-
scens, rarely on herbaceous species such as Coronilla, perhaps also Onobrychis). Food
plant records from other countries include Cytisus nigricans, Trifolium campestre, Coronil-
la sp., Melilotus sp., Medicago sp., Glycyrrhiza sp. and Caragana arborescens. Highest local-
ities in Swabia and the Bavarian foothills of the Alps are at 650 m a.s.l. only.
Frommer (pers. comm.), Niedringhaus (1997), Post-Plangg & Hoffmann (1982), Schiemenz (1988), Schönitzer
& Oesterling (1998a), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN and others; see also Müller
(1956), Ossiannilsson (1978), Schedl (1998)

Centrotus cornutus (Linnaeus, 1758)

B V – M X, mainly M V – B VIII; nymph (usually 3rd and 5th instar), semivoltine (Müller
1984a).
In moderately moist to dry sites with tall herbs or shrubs, usually along woodland mar-
gins and among herb stands in dry grassland, along hedges, forest roads and margins,
along shores as well as in the interior of open forests with luxuriant undergrowth.
Nymphs usually stay near the base of the host (mainly Vincetoxicum, Cirsium, Carduus
and Urtica), adults ascend to the upper herb and shrub layer for feeding and oviposi-
tion and are often swept from low-growing woody plants (Populus, Quercus, Rubus, Pru-
nus and others). Widespread in Germany, but often collected in low numbers only. Found
up to at least 1200 m a.s.l. in the Bavarian Alps, up to 1500 m a.s.l. in Switzerland.
Bußmann & Feldmann (1995), Fischer (1972), Heller (1987a), Müller (1984b), Reimer (1992), Schiemenz
(1988), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN and others;
see also Günthart (1987a)

Stictocephala bisonia Kopp & Yonke, 1977

Several nomenclatural changes have been proposed for this species in recent years. In most
older papers the name Ceresa bubalus (Fabricius, 1794) was used. Caldwell (1949) placed it into
the genus Stictocephala Stål, 1869. Kopp & Yonke (1977) studied Fabricius‘ type and found that it
is not conspecific with the species introduced into the Old World, and created the new name
bisonia. More recently De Andrade (1997), after a revision of Walker’s material, suggested that
the species has already been described as Ceresa alta Walker, 1851. However, according to Diet-
rich et al. (1999) this synonymy is doubtful, and the identity of Walker’s type, which is a female,
is uncertain.
M VII – E X; egg, 1 gen.
Along woody margins and in tall herb stands, usually near rivers, streams and ditches;
oviposition is reported to take place on low-growing, woody species of Rosaceae (ap-
ple, pear, plum, cherry, peach, blackthorn and others, thus locally becoming noxious),
willows (e.g. Salix purpurea, S. aurita) and others, nymphs are usually found on herba-
ceous plants (Solidago gigantea, Medicago sativa, also invading cultivations). Additional
80 Review of species

food plants reported in the literature include Rosa, Cornus, Crataegus, Vitis, Populus, Ul-
mus, Coronilla, Melilotus, Chrysanthemum and Artemisia. The species was introduced from
North America into Europe (presumably with twigs of fruit trees) where it was report-
ed for the first time in 1912 from former Hungary. Later it spread over most parts of
southern Europe (extending northward to Belgium in oceanic climates) and has reached
parts of North Africa, Kirghizia and Kazakhstan. In Germany recorded for the first time
in 1966 along the southern upper Rhine (Isteiner Klotz) and in 1972 in numerous sites
between Lörrach and Kehl, although already found in 1952 in Alsace near Strasbourg
(France). Meanwhile it occurs in the valleys of the Black Forest, down the Rhine valley
at least to Speyer (since 1996) and Darmstadt (since 2000), up the Rhine to Radolfzell
near Lake Constance, 400 m a.s.l. (1998) and up the Neckar valley to Nürtingen (since
1995). Has been reported to cause damage on grapevine and fruit trees by oviposition
into young twigs in Italy.
Dynort (pers. comm.), Gauss (1987), Hoffmann (1990), Hoffrichter & Tröger (1973), Müller (1956), Reder
(pers. comm.), Remane (1972), Schuler (1952), SMNS, Voigt (pers. comm.), HN; see also Arzone et al. (1987),
Emelyanov (1993), Günthart (1980), Nast (1987), Schedl (1995), Vidano (1963a, 1963b)

4.2.4 Cicadellidae Latreille, 1825 - Leafhoppers (s.str.)

4.2.4.1 Ulopinae Le Peletier & Serville, 1825

Ulopa carneae W. Wagner, 1955

In Austria adults and nymphs recorded IV – VIII (Wagner & Franz 1961); probably
semivoltine, like the following species (see below).
This species is known only from a few records in the eastern parts of the Alps between
700 and 2200 m a.s.l. on Erica carnea in open pine and spruce forests and in subalpine
scrub. So far there are only two German records from the Bavarian Alps: Winkl near
Berchtesgaden, 700 m a.s.l., 22.VIII.1996, 2 nymphs; Schwarzeck near Ramsau, 1000 m
a.s.l., 24.VIII.1996, 1 †, 2 ‡‡, 2 nymphs.
Nickel (1999b); see also Leising (1977), Wagner & Franz (1961)

Ulopa reticulata (Fabricius, 1794)

Adults and nymphs I – XII; nymph and adult, semivoltine (Melber 1989).
In peaty to moderately dry, sunny to moderately shady sites on acidic substrates, main-
ly heaths, bogs and forest margins, occasionally also in open pine and oak forests; the
host plant is Calluna vulgaris. Widespread in Germany and particularly common in the
north German plain and along the edge of the Alps, more scattered in middle parts,
where locally absent. Frequently found at 1300 m a.s.l. in the Mittelgebirge and the Ba-
varian and Allgäu Alps, up to 1600 m a.s.l. in Styria (Austria).
Fischer (1972), Melber (1989), Niedringhaus (1991), Reimer (1992), Schiemenz (1971a, 1975, 1988), Schönitzer
& Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN; see also Wagner & Franz (1961)
 Megophthalminae Kirkaldy, 1906 81

Utecha trivia (Germar, 1821)

According to Orosz (1977) specimens which probably originate from the same locus typicus, but
named Ulopa lugens Germar, 1821, belong to a distinct species. The majority of European taxon-
omists, however, consider it only as a macropterous form of U. trivia (Germ.). The latter view is
adopted here.
Adults I – XII, mainly E VII – V; adult (apparently only ‡‡), 1 gen.
In sunny and oligotrophic, moderately dry to dry and low-vegetated (preferentially
grazed) sites usually on basic substrates. The host relationships are poorly known; the
few published reports include Hippocrepis comosa, Echium vulgare, in England Plantago
lanceolata. In Germany rather scattered and mainly in southern and eastern parts; north-
ernmost localities are on a line from Bad Münstereifel to Mainz, Aschaffenburg, Bad
Kissingen, Gotha, Sondershausen, Wittenberg and Dresden, frequently occurring at
700 m a.s.l. on the Swabian Jura, but recorded up to 1300 m a.s.l. in the Chiemgau Alps.
Reimer (1992), Rombach (1999a), Schiemenz (1988), Schönitzer & Oesterling (1998b), SMNS, Wagner (1939a,
1951a), HN; see also Morris (1972), Szwedo (2001)

4.2.4.2 Megophthalminae Kirkaldy, 1906

Megophthalmus scabripennis Edwards, 1915

So far only E VI; egg, 1 gen.
This species‘ range includes the Mediterranean region and parts of western Europe,
where it is normally reported from sunny margins and open, xerothermic forests. From
Germany there are only two records from sun-exposed rocky slopes in the valleys of the
southern upper Rhine and the Nahe: Isteiner Klotz near Lörrach, c. 300 m a.s.l. and Bad
Münster, c. 200 m a.s.l., 21.VI.1965, 2 ††, 1 ‡.
Remane & Fröhlich (1994b), Coll. Kobel-Lamparski (Remane det.); see also Guglielmino (1993), Holzinger
(1995b)

Megophthalmus scanicus (Fallén, 1806)

E VI – E X; egg, 1 gen.
In sunny, wet to moderately dry sites, mainly low-input meadows and pastures, heaths,
fens, abandoned fields and waysides. Single macropterous individuals may be swept
among tall herbs, shrubs and trees. Host plants are various species of Fabaceae (Trifoli-
um dubium, Medicago lupulina, Lotus pedunculatus and others). Widespread and fairly
common in most parts of Germany, frequently found in the Allgäu at 1200 m a.s.l., in
the Bavarian Alps occasionally at 1500 m a.s.l.
Achtziger (1991), Fischer (1972), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Schiemenz (1988),
Schönitzer & Oesterling (1998b), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN and others; see also
Wagner & Franz (1961)
82 Review of species

4.2.4.3 Ledrinae Kirschbaum, 1868

Ledra aurita (Linnaeus, 1758)

M VI – B X, rarely E III; nymph (hibernating twice), semivoltine (Schiemenz 1988).
In moist to moderately dry sites, in closed forests as well as in semi-open tree stands.
Lives on the bark of various deciduous trees, preferentially Quercus, but frequently also
Betula, Populus, occasionally Tilia, Fagus, Malus and Acer, also reported from Alnus and
Corylus. Widespread in the lowlands of Germany, but collected in low numbers only,
usually in light traps and tree eclectors, rarely in the sweep-net. Highest localities in the
Mittelgebirge and the foreland of the Alps at 550 m a.s.l. only.
Achtziger (1991), Büchs (1988), Fischer (1972), Frommer (1996), Niedringhaus (1997), Schiemenz (1988),
Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN

4.2.4.4 Macropsinae Evans, 1935

Oncopsis alni (Schrank, 1801)

M V – E VIII, at higher altitudes until IX; egg, 1 gen.
Monophagous on alders (Alnus glutinosa and A. incana) near standing or running water,
in fen woods and on seepy slopes. Widespread in Germany, common and in high fre-
quency on both alder species (also on planted A. incana in lowlands), extending up to at
least 1000 m a.s.l. in the Mittelgebirge and the Bavarian Alps, found up to the subalpine
belt in the northeastern Alps of Austria. Was recently found to be a potential vector of
the alder yellows phytoplasma.
Fischer (1972), Günthart (1988), Maixner & Reinert (1999), Niedringhaus (1991, 1997), Reimer (1992), Schie-
menz (1988), Schönitzer & Oesterling (1998b), Wagner (1949a, 1951a), HN; see also Wagner & Franz (1961)

Oncopsis appendiculata W. Wagner, 1944

E V – B VIII; egg, 1 gen.
On Betula pendula, rarely on B. pubescens, usually on solitary trees or along forest mar-
gins and alleys. Often occurs together with other species of Oncopsis, but apparently
confined to damp or dry sites of lower altitudes. Widespread in Germany, but uncom-
mon, locally absent and only in low to intermediate frequency. The highest localities are
in the foreland of the Alps at c. 600 m a.s.l.
Freese (pers. comm.), Frommer (pers. comm.), Schiemenz (1988), SMNS, Wagner (1949a), Nickel & Remane
(1996), HN

Oncopsis tristis (Zetterstedt, 1840)

B VI – B VIII, occasionally until B IX; egg, 1 gen.
Monophagous on birches (Betula pendula and B. pubescens), but highly eurytopic, in raised
bogs as well as dry grassland with scattered trees, on solitary trees as well as in open
 Macropsinae Evans, 1935 83

forests, often also in urban areas. Widespread and common in Germany, in high fre-
quency on the hosts, although in smaller numbers than O. flavicollis (L.). Highest local-
ities are at 1200 m a.s.l. in the Alps and the Black Forest; in North Tyrol (Austria) breed-
ing is reported from 1900 m a.s.l.
Fischer (1972), Niedringhaus (1991, 1997), Reimer (1992), Remane & Reimer (1989), Schiemenz (1988), Wagner
(1951a), HN; see also Leising (1977)

Oncopsis avellanae Edwards, 1920

So far E V – E VI; egg, 1 gen.
On Corylus avellana (in Britain also on Alnus glutinosa) in damp to dry woodland sites,
usually along forest roads and margins and on solitary shrubs in meadows and pas-
tures. Widespread at least in the western half of Germany where occurring in low to
intermediate frequency on the host. Recorded from the Kaiserstuhl, Allgäu, Rhön Moun-
tains, Mainfranken, middle Hessen, southern Niedersachsen and adjacent parts of Thu-
ringia, the Lüneburger Heide, the island of Borkum, eastern Holstein and Angeln, up to
800 m a.s.l. According to current knowledge these localities are at the eastern edge of
the range, but the species is likely to be under-recorded or misidentified and may be
more widespread in Europe. Otherwise it is only known from France, the Netherlands
and Great Britain.
Fischer (1972), Remane & Fröhlich (1994b), HN; see also Claridge et al. (1977), della Giustina & Remane
(2001), Nast (1987)

Oncopsis carpini (J. Sahlberg, 1871)

M V – E VIII; egg, 1 gen.
On Carpinus betulus in various sites, usually along forest margins and on solitary trees,
but also in the interior of forests (where difficult to sample with the sweep-net), in parks
and even in hedges in urban areas. Widespread in Germany and usually in high fre-
quency on the host, although only scarcely recorded from the northwest German plain;
highest localities are at 800 m a.s.l. in the Mittelgebirge and the Alps.
Achtziger (1991), Fischer (1972), Frommer (pers. comm.), Günthart (1988), Post-Plangg & Hoffmann (1982),
Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1949a), HN

Oncopsis flavicollis (Linnaeus, 1761) group

This taxon is morphologically extremely variable. In Wales, according to bioacoustic and mor-
phometric studies of Claridge & Reynolds (1973) and Claridge & Nixon (1986), it is likely to
consist of three distinct species. This may also be true for central European populations.
M V – VII, occasionally until B IX; egg, 1 gen.
Eurytopic on birches (Betula pendula and B. pubescens) in much variable sites, in forests
as well as on solitary trees, in bogs, parks, gardens, in dry grassland and others. Breed-
ing has been documented at 1900 m a.s.l. near the tree line of the Tyrolian Alps (Aus-
tria). Widespread in Germany, extremely abundant and in very high frequency on the
84 Review of species

hosts. Frequently recorded at 1200 m a.s.l. in the Black Forest and the Allgäu, but prob-
ably extending higher up.
Fischer (1972), Niedringhaus (1991, 1997), Reimer (1992), Remane & Reimer (1989), Schiemenz (1988), Wagner
(1935, 1939a, 1951a), HN and others; see also Leising (1977)

Oncopsis subangulata (J. Sahlberg, 1871)

E V – E VIII; egg, 1 gen.
On birches, preferentially on Betula pendula, in Wales also reported from B. pubescens.
Widespread in Germany, but less common than the preceding species, only in low to
intermediate frequency and apparently confined to lower altitudes (found only up to
700 m a.s.l.), with strongholds in river floodplains.
Fischer (1972), Freese (pers. comm.), Günthart (1988), Niedringhaus (1997), Schiemenz (1988), Wagner
(1949a), HN; see also Claridge & Reynolds (1972)

Pediopsis tiliae (Germar, 1831)

B VI – B VIII, occasionally M IX; egg, 1 gen.
On lime (Tilia cordata, T. platyphyllos, perhaps also ornamental species), mainly in forests,
along alleys and on solitary trees (also in urban areas). Widespread in the lowlands of
Germany and in intermediate to high frequency, but usually low abundance; more scat-
tered in north Germany and the foreland of the Alps. Highest localities at 600 m a.s.l. only.
Fischer (1972), Günthart (1988), Reimer (1992), Schönitzer & Oesterling (1998b), Remane & Wachmann
(1993), Schwoerbel (1957), Wagner (1935, 1939a), HN

Most members of the genus Macropsis are very difficult to identify, due to only slight differences in
genitalic structures and considerable intraspecific variation of colouration. Hence, distribution pat-
terns shown in this book are often incomplete. Wagner (1941a, 1950, 1953, 1964) contributed funda-
mentally to the knowledge of taxonomy and morphology of many species. In particular he pointed
out that most of them show a rather strict association with their host plants. Nevertheless, there may
remain a number of undescribed or misidentified species in central Europe. Later, Nast (1981), Tishech-
kin (1999) and Nickel (1999a) added further information. Just before this book was printed, Tishech-
kin (2002) presented a detailed revision of east and central European species, including a key with
all relevant characters. In many cases, a reliable identification is only possible after these papers.

Macropsis albae W. Wagner, 1950

Hamilton (1980) suggested that this name was a junior synonym of M. ocellata Provancher, 1872,
which is suspected to be introduced from Europe to North America. However, Provancher’s type
is lost; the author did not even give information on sex and type locality and thus, its identity is
highly uncertain. Later, Hamilton (1983b) designated a neotype from Canada, choosing an indi-
vidual of the introduced species dwelling on Salix alba. Although Nast (1987), Schiemenz (1988),
and others followed him, this opinion is rejected here, due to the extremely critical taxonomic
situation among green species of Macropsis, and the unambiguous description of M. albae W.Wg.
E V – E VIII; egg, 1 gen.
On Salix alba, probably also on its hybrids, usually near running or standing water.
Widespread particularly in middle and southern parts of Germany and usually in high
 Macropsinae Evans, 1935 85

frequency on the host; less common in the north German plain, where reaching the edge
of the range. Northernmost localities are on the island of Langeoog, near Hamburg,
Kiel and Neubrandenburg, recorded up to at least 750 m a.s.l. near the edge of the Alps.
The locus typicus is Campow near Ratzeburg.
Lauterer & Malenovsky (1995), Mölleken & Topp (1997), Niedringhaus (1991), Reimer (1992), Remane (pers.
comm.), Schiemenz (1988), Wagner (1950, 1951a), HN

Macropsis gravesteini W. Wagner, 1953

So far M VI – M VIII; egg, 1 gen.
Like the preceding species on Salix alba, but rather uncommon and in low frequency.
The distribution in Germany and Europe is imperfectly known, but the species has prob-
ably often been overlooked or misidentified. German records are from near Lübeck,
Dannenberg, Düsseldorf, Halle, Marburg and Bad Nauheim, mostly in larger river val-
leys below 200 m a.s.l.
Mölleken & Topp (1997), Remane (pers. comm.), Remane & Fröhlich (1994b), HN

Macropsis prasina (Boheman, 1852)

Recently Tishechkin (1994) clarified the status of two further pale green Macropsis species, both
described from central Asia and living on Salix viminalis. At least one of them – M. microcera
Vilbaste, 1982 – is also found in European Russia. Hence, much of the central European speci-
mens – including most of those cited in this book – are in need of revision. However, a number
of German specimens which have been revised by Tishechkin (pers. comm.) belonged to M.
prasina (Boh.).
B VI – B IX; egg, 1 gen.
On hairy-leaved willows in damp to wet sites, in the study area Salix cinerea, S. vimina-
lis, S. aurita and S. caprea. Apparently widespread and common in the lowlands of Ger-
many, but imperfectly documented; in many places it occurs in intermediate to high
frequency on the hosts. Frequently recorded to at least 600 m a.s.l. in the Mittelgebirge
and the Alps, occasionally up to 1200 m a.s.l.
Mölleken & Topp (1997), Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992), Wagner (1950), Nickel
& Remane (1996), HN; also Tishechkin (pers. comm.)

Macropsis viridinervis W. Wagner, 1950

B VI – M VIII; egg, 1 gen.
On Salix triandra, presumably also S. pentandra, in river floodplains, along streams and
lake shores. Often overlooked or misidentified, but occurring at least scattered in the
lowlands of southern and middle parts of Germany, with strongholds in the larger river
valleys, usually in intermediate frequency on the host, although reaching the edge of
the range. Northernmost records from near Rinteln an der Weser, Hamburg (locus typi-
cus), Dannenberg, Halle and Dresden. Also recorded to the south of Stettin (northwest-
ern Poland) and up to 700 m a.s.l. in the foothills of the Alps.
Nicolaus (1957), Wagner (1950, and unpublished data), HN; see also Holzinger (1999a)
86 Review of species

Macropsis notata (Prohaska, 1923)

Macropsis vestita Ribaut, 1952 sec. Tishechkin (2002)
B VI – E VIII; egg, 1 gen.
In intermediate frequency on Salix triandra, less commonly on S. fragilis, usually in river
floodplains. Imperfectly documented from Germany, but probably widespread in the
lowlands – at least between the Danube and the coast. Northernmost localities are near
Höxter, on the island of Amrum, near Plön, Heiligenhafen, Bad Frankenhausen and
Meißen, only up to at most 400 m a.s.l. Not recorded from Denmark and Scandinavia,
in Poland apparently confined to southern parts.
Nicolaus (1957), Remane (pers. comm.), Schwoerbel (1957), SMNS, Wagner (1950, 1951a), HN; see also
Nast (1976a), Ossiannilsson (1981)

Macropsis marginata (Herrich-Schäffer, 1836)

M V – E VIII; egg, 1 gen.
Monophagous on Salix purpurea, eurytopic near running or standing water. Distribution in
Germany imperfectly documented, but common and in high frequency on the host, at least in
middle and southern parts. The northern edge of the range apparently runs through northern
Germany; northernmost localities are near Cologne, Rinteln an der Weser, Hamburg and Greif-
swald, up to 1300 m a.s.l. in the Bavarian Alps. The locus typicus is near Regensburg.
Fischer (1972), Frommer (pers. comm.), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b),
Schwoerbel (1957), Wagner (1950), HN

Macropsis infuscata (J. Sahlberg, 1871)

B VI – E VIII; egg, 1 gen.
At lower altitudes (including the Mittelgebirge and the foothills of the Alps, up to at least
900 m a.s.l.) this species lives on Salix caprea in forest glades and clearings, along forest
margins and roads, in hedges and ruderal sites; in the Alps and their foothills (recorded
between 700 and 1200 m a.s.l.) it is also found on S. myrsinifolia along running water
and on seepy slopes. It is widespread and common in Germany and occurs in interme-
diate to high frequency on S. caprea, less frequently on S. myrsinifolia.
Fischer (1972), Frommer (pers. comm.), Niedringhaus (1997), Reimer (1992), Schiemenz (1988), Schönitzer
& Oesterling (1998b), Wagner (1950, 1951a), HN

Macropsis cerea (Germar, 1837)

Central European populations currently interpreted as M. cerea (Germ.) are likely to comprise
several distinct species.
B VI – B IX; egg, 1 gen.
On various species of willows, in the lowlands mainly Salix viminalis, S. triandra, S.
fragilis, S. cinerea and S. caprea, in the Mittelgebirge and Alps mainly S. aurita, S. purpurea
and S. myrsinifolia. Widespread in Germany, occurring in intermediate to high frequen-
 Macropsinae Evans, 1935 87

cy, but rather low individual numbers on the hosts. Highest localities are at 1200 m a.s.l.
in the Mittelgebirge and the Allgäu Alps.
Fischer (1972), Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992), Schiemenz (1988), Schönitzer &
Oesterling (1998b), Schwoerbel (1957), Wagner (1950, 1951a), HN

Macropsis remanei Nickel, 1999

E VI – M IX; egg, 1 gen.
Monophagous on Salix eleagnos along streams and rivers in mountains and their foreland.
This species was described only recently from the Bavarian Alps. At present, there are 10
German localities, near Berchtesgaden (locus typicus: Ramsau), along the upper Isar be-
tween Mittenwald and Bad Tölz, and along the Loisach near Garmisch-Partenkirchen, at
altitudes between 650 and 1400 m a.s.l. Furthermore, the species has been found in vari-
ous localities in northern Italy (Friuli Venezia Giulia: Tagliamento) and Slowenia. Proba-
bly it is more widespread in the Alps and perhaps other European mountains.
Nickel (1999a), HN; see also Löcker (2003), Tishechkin (2002)

Macropsis najas Nast, 1981

M VI – E VIII; egg, 1 gen.
Until recently this species was only known from the original description from Poland,
but later it was found in large parts of central Europe. Meanwhile there are more than
25 records mainly from middle parts of Germany (approximately between a line run-
ning from Cologne to Göttingen, Stendal and Görlitz in the north and another one from
Speyer to Würzburg and Bayreuth in the south, also near Lüneburg). It lives monopha-
gously on Salix alba along rivers and streams, recorded only below 400 m a.s.l. Other-
wise it is also known from northern Italy, eastern France and Austria.
Frommer (pers. comm.), Remane (1995), Nickel (1997), HN; see also Holzinger et al. (1996), Nast (1981)

Macropsis haupti W. Wagner, 1941

This species has been mistaken for M. cerea (Germ.) for a long time, until Wagner (1941a), after
studying Germar’s types, recognized its distinctiveness and created a new name. However,
Ossiannilsson (1948) studied the type of Pediopsis planicollis Thomson, 1870, originally described
from Sweden, and erroneously declared them to be conspecific, thus rendering M. haupti W.Wg.
a junior synonym. Subsequently, Wagner (1950), in his exhaustive monograph on central Euro-
pean Macropsis species, accepted this opinion. After studying this paper, Ossiannilsson (1954)
realized his error and declared M. planicollis (Thoms.) a junior synonym of M. cerea (Germ.).
B VI – M IX; egg, 1 gen.
Monophagous on Salix purpurea, mainly along rivers and streams. Often syntopic with
M. marginata (H.-S.), although usually in lower numbers and frequency, recorded only
up to 1000 m a.s.l. Widespread in middle and southern parts of Germany, but only scat-
tered, reaching the edge of the range in the north German plain. Northernmost locali-
ties are near Rinteln an der Weser, Dannenberg and Frankfurt an der Oder; also report-
ed from northwestern parts of Poland (to the south of Stettin).
88 Review of species

Fischer (1972), Reimer (1992), Schönitzer & Oesterling (1998b), Wagner (1950, 1951a), HN

Macropsis impura (Boheman, 1847)

B VI – M VIII; egg, 1 gen.
On Salix repens, occasionally on low-growing individuals of Salix aurita. Strongholds in Germa-
ny on coastal dunes near the North and Baltic Sea, where locally common; very sporadic else-
where, usually in peaty wetlands of the northern plains, the Mittelgebirge and along the edge of
the Alps, where found to at least 800 m a.s.l. (but up to 1050 m a.s.l. in Styria, Austria); not
recorded from southwestern parts (Baden-Württemberg, Rheinland-Pfalz, Nordrhein-Westfalen).
Niedringhaus (1991), Remane & Reimer (1989), Schiemenz (1988), Schönitzer & Oesterling (1998b), Trüm-
bach (1959), Wagner (1950), HN

Macropsis fuscinervis (Boheman, 1845)

M VI- M VIII, according to Wagner (1951a) from M V onwards; egg, 1 gen.
Monophagous on Populus tremula, usually along forest margins and roads, in clearings and
ruderal sites. Widespread in Germany, but only in intermediate frequency and low individual
numbers on the host, recorded up to 700 m a.s.l. in the Mittelgebirge and the Bavarian Alps.
Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b),
Schwoerbel (1957), Wagner (1935, 1941a, 1941c), HN

Macropsis graminea (Fabricius, 1798)

B VI – M VIII; egg, 1 gen.
Monophagous on Populus nigra, on the nominate form as well as on var. italica, occasionally
also on hybrids, where reproductive success may be uncertain; mainly in river floodplains,
parks and urban settlements, usually in intermediate to high frequency, but low individual
numbers. In Germany apparently confined to southern and middle parts (to at least 600 m
a.s.l.), localities near Cologne, Göttingen, Hamburg, Lübeck and Berlin are probably near the
northern edge of the range, although there are a few older records from Norway and Sweden.
Frommer (pers. comm.), Günthart (1988), Nikusch (1976), Reimer (1992), Schiemenz (1988), Wagner (1941a,
1941c, 1951a), HN; see also Ossiannilsson (1981)

Macropsis vicina (Horváth, 1897)

So far M VI – B VIII; egg, 1 gen.
In Europe this species occurs mainly in southern parts, but extends locally northward to
Champagne (France), Germany and eastern Poland. The host plant is Populus alba. From
Germany it is only known from 4 records in the Kaiserstuhl of southwestern Baden, the
Kyffhäuser of Thuringia and the vicinity of Berlin: Achkarren, c. 200 m a.s.l., 18.VI.1971,
3 ††, 12 ‡‡ and nymphs; Oberbergen, c. 400 m a.s.l., 15.VI.2002, 1 †, 1 ‡; Günserode, 160 m
a.s.l., 08.VII.1995, 2 ††; Berlin, Tegel, c. 40 m a.s.l., 10.VIII. 2002.
Remane & Fröhlich (1994b), Stewart (pers. comm.), HN; see also Nast (1976a)
 Macropsinae Evans, 1935 89

Macropsis elaeagni Emelyanov, 1964

In the Czechia Republic, Slovakia and Austria B VI – B IX (Holzinger & Remane 1994;
Lauterer 1984); egg, 1 gen.
Originally this species was confined to Kazakhstan and southern Russia and lives on
Elaeagnus. In recent decades it has expanded its range and appeared in the Balkan re-
gion and southeastern parts of central Europe, where it has colonized secondary plan-
tations along waysides and in public parks. Dispersal is suspected to take place mainly
through cuttings. In Germany it was found only in two localities in the northern upper
Rhine plain: Weiterstadt, 120 m a.s.l., 03.VIII.1996, 29 ††, 15 ‡‡, and Germersheim,
17.VIII.1998, 3 ‡‡, both along waysides on E. angustifolia, c. 120 m a.s.l., but it is proba-
bly more widespread at least in southern parts.
HN; see also Emelyanov (1964b), Lauterer (1984)

Macropsis glandacea (Fieber, 1868)

Pediopsis mendax Fieber, 1868; Pediopsis ulmi Scott, 1873
B VI – B IX; egg, 1 gen.
On elm, usually Ulmus minor and U. x hollandica, at least single specimens found on U.
laevis, usually in river floodplains, parks, along alleys and hedges. In Germany at the
northern edge of the range, largely restricted to the valleys of larger rivers and their
basins (so far found along the Rhine, Main, Neckar, Lahn, Danube, Altmühl, Leine,
Unstrut, Saale, Elbe and Neiße). The northernmost localities are near Cologne, Göttin-
gen, Hamburg, Plön, Stendal and Berlin, the highest ones at 400 m a.s.l.
Frommer (pers. comm.), Nikusch (1976), Remane (1987), Remane & Wachmann (1993), Schiemenz (1988),
SMNS, Wagner (1939a, 1941a, 1941c), Nickel (1994, 1999b), HN

Macropsis brabantica W. Wagner, 1964

The specific identity of this taxon has yet to be demonstrated. According to Wagner (1964) it
lives monophagously on Rubus caesius and could not be reared on R. idaeus in the laboratory.
However, German populations of Macropsis on R. caesius, most of which clearly belong to M.
fuscula (Zett), often include some individuals showing intermediate colouration of M. fuscula
(Zett.) and M. brabantica W.Wg.
So far only E VIII; probably egg, 1 gen.
In Germany only reported from two sites in Brandenburg to the south of Berlin: Klein
Köris and Groß Köris, 22. and 24.VIII.1995, „an einer niedrigwüchsigen Rubus-Art (Ru-
bus caesius?) ( ... ) sowohl im Schatten eines Mischwaldes als auch in einem Erlenbruch-
wald“. The species was described from several localities in the Netherlands and has
since been reported only from Germany and middle Russia (see below).
Remane (1995); see also Tishechkin (2002), Wagner (1964)

Macropsis fuscula (Zetterstedt, 1828)

B VI – M IX, occasionally until B X; egg, 1 gen.
90 Review of species

On Rubus idaeus, R. fruticosus and R. caesius in various types of forests, in clearings,

along hedges, in tall herb stands and gardens. Widespread in Germany, frequently oc-
curring to at least 1300 m a.s.l. in the Mittelgebirge and the Alps, but single specimens
were found up to 1600 m a.s.l. Has been identified as a potential vector of Rubus stunt
disease (RSD) in various parts of Europe.
Fischer (1972), Frommer (1996), Niedringhaus (1991, 1997), Reimer (1992), Schiemenz (1988), Schönitzer &
Oesterling (1998b), Schwoerbel (1957), Wagner (1939a, 1964), HN; see also Brčák (1979)

Macropsis scotti Edwards, 1920

So far E VII – M VIII, in Britain VII – X (Le Quesne 1965); egg, 1 gen.
This species occurs from Morocco through western Europe to England and Ireland.
From Germany it is recorded only in two localities in the extreme west: Eifel Moun-
tains, Niederzissen, c. 300 m a.s.l., mid August 1974, 1 ‡, along a hedge, and Stut-
tgart, c. 250 m a.s.l., 21.VII.1937, 1 specimen. Is reported to live monophagously on
Rubus fruticosus.
Post-Plangg & Hoffmann (1982) (Remane det.), Tishechkin (2002, and pers. comm.); see also Wagner (1964)

Macropsis megerlei (Fieber, 1868)

B VII – M VIII; egg, 1 gen.
Until recently this species was known to occur in central Europe from a few records
only. It lives on shrubs of roses in sunny, xerothermic sites (mainly on limestone, gyp-
sum, schist, etc.). Host plants are mainly Rosa rubiginosa, less frequently R. spinosissima
and R. canina; occasionally also ornamental R. glauca and R. rugosa. In Germany it is
widespread at least in the middle parts and difficult to collect, but probably fairly com-
mon on favourable sites. There are records from Mainfranken, northern Baden, middle
Hessen, Thuringia, Palatinate, as well as along the northern and southern edge of the
Harz Mountains, up to 400 m a.s.l. To the north of Germany it is known only from two
single specimens from southern Norway.
Kupka (1925), Wagner (1939a), Schiemenz (1988), SMNS, Nickel (1997), HN; see also Lauterer (2000), Os-
siannilsson (1981)

Macropsis scutellata (Boheman, 1845)

B VI – M IX; egg, 1 gen.
Lives monophagously on Urtica dioica in damp to moderately wet, sunny to moderately
shady sites, usually in river floodplains, along shores of lakes and ponds and in ruderal
sites with tall herbaceous vegetation. Widespread in Germany, although seldom record-
ed due to secretive life habits, and apparently largely confined to lowland areas (up to
at most c. 600 m a.s.l.); however, there is a single specimen from the Allgäu Alps at
1200 m a.s.l.
Fischer (1972), Frommer (pers. comm.), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Schönitzer
& Oesterling (1998b), Wagner (1935, 1939a, 1952, 1964), HN
 Agalliinae Kirkaldy, 1901 91

Macropsis mulsanti (Fieber, 1868)

In Switzerland and Austria so far M VII – B IX; egg, 1 gen.
Monophagous on Hippophae rhamnoides, reported to live on sunny and rocky slopes in Swit-
zerland. In Europe this species is mainly confined to southern parts; to the north of the Alps
it has only been found in a few localities near Lake Constance and along the upper course of
the Rhine: Bludenz and Röthis (both Vorarlberg, Austria), Strasbourg (Alsace, France) and
southern Baden: Heitersheim, 06.VIII.1997, 3 ‡‡ on a roadside embankment.
HN; see also Moosbrugger (1946), Remane & Fröhlich (1994b), Tishechkin (1993)

Hephathus nanus (Herrich-Schäffer, 1835)

Palearctic species of this genus were recently reviewed by Tishechkin (1999), proving that the
Asian H. achilleae Mityaev, 1967 also occurs in the eastern half of Europe, notably in large parts
of European Russia, southern Moravia (Czech Republic), Hungary and probably Finland. Thus,
it may be more widespread in central Europe as well. The author also pointed out that the
drawings of Ossiannilsson (1981) do not refer to H. nanus (H.-S.), but to H. achilleae Mity.
So far E VI – E VIII; egg, 1 gen.
In sunny, oligotrophic and xerothermic sites with short vegetation, usually in heavily
grazed pastures on basic substrates, perhaps associated with Cirsium acaule. In Germa-
ny only locally and at the northern edge of its range, mainly in warmer regions of east-
ern and southern parts (Saale-Unstrut region, Thuringian Basin, Mainfranken, valleys
of the Neckar, Nahe and upper Rhine). Northernmost localities are near Mainz, Nort-
heim, Magdeburg and Berlin; found up to at least 500 m a.s.l. near Arnstadt and Munich,
but there is also an unconfirmed record from at least 1000 m a.s.l. from the Bavarian
Alps. The locus typicus is near Regensburg.
Haupt (1925), Remane (1987), Schönitzer & Oesterling (1998b), Schiemenz (1969, 1988), SMNS, Wagner
(1939a), ZIMH, Nickel (1994), HN; for identification see Tishechkin (1999)

Macropsidius sahlbergi (Flor, 1861)

M VI – B VIII; egg, 1 gen. (Schiemenz 1988).
This species occurs mainly in steppe regions of central Asia and southeastern Europe,
with rather isolated populations in xerothermic grassland sites of central Europe, from
where it is reported to live on Artemisia campestris. In Germany it is confined to north-
eastern parts and known only from 5 localities near Pasewalk, Angermünde, Eberswal-
de, Stendal and Meißen, all below 150 m a.s.l. The last record dates from 1965; thus the
current status is uncertain.
Schiemenz (1969, 1988), Wagner (1941a)

4.2.4.5 Agalliinae Kirkaldy, 1901

Agallia brachyptera (Boheman, 1847)

B VII – E X; eggs overwinter in secondary parapause terminated by cold, 1 gen. (Witsack 1985)
92 Review of species

In sunny to shady, damp to wet, occasionally moderately dry or peaty sites, notably
low-input meadows and pastures, abandoned fields and forests. Presumably this spe-
cies lives polyphagously on various species of Fabaceae and Asteraceae (Taraxacum,
also reported from Trifolium, Onobrychis, Achillea, as well as Rumex). Widespread in Ger-
many and fairly common, found up to at least 900 m a.s.l. in the Mittelgebirge and the
Alps; frequently recorded at 2000 m a.s.l. in Trentino (northern Italy).
Niedringhaus (1991), Nikusch (1976), Reimer (1992), Remane (1958), Schiemenz (1988), Wagner (1935),
Nickel & Achtziger (1999), HN; see also Ossiannilsson (1981)

Agallia consobrina Curtis, 1833

M VII – M V, from M VI onwards along the upper Rhine; adult (only ‡‡?), 1 gen.
In moderately shady, moderately dry to moist sites on acidic to basic substrates, usually
in rather warm situations, mainly open deciduous forests with luxuriant growth of herbs
(coppices, coppice-with-standards, occasionally also floodplain forests and alder fen
woods), dry grassland and abandoned vineyards with scattered shrubs and trees. Lives
on various species of Lamiaceae (Glechoma, Teucrium, probably also Lamium, Stachys
and others), perhaps also grasses and Urtica dioica. In Germany widespread between
the Danube and the northern edge of the Mittelgebirge, and particularly common in
viticultural regions, rather scattered in northern Germany and not recorded to the south
of the Danube. Most localities are below 500 m a.s.l., although found up to at least 900 m
a.s.l. in the southern Black Forest.
Kuntze (1937), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schiemenz (1988), Schwo-
erbel (1957), Trümbach (1959), Wagner (1935, 1939a, 1951a), HN

Anaceratagallia austriaca W. Wagner, 1955

So far B III – M VI, in the Valais (Switzerland) according to Dlabola (1971) E IX; adult,
1(?) gen.
According to existing data, this species is a rare endemic of the Alps and south German
limestone regions. In Germany it was only found in the Franconian Jura, Kipfenberg, c.
350 m a.s.l., 06. and 08.III.1961, „eine individuenreiche Population ( ... ) auf einem Step-
penheide-Südhang ( ... ) unter Thymus-Polstern“, as well as in the middle Main valley,
Gambach, 300 m a.s.l., 04.VI.1993, 3 ‡‡, and 19.VI.1994, 2 ‡‡, along the upper edge of a
sun-exposed, steep and rocky hillside. Otherwise only reported from a few xerothermic
sites in Styria (Austria), the Vintschgau (Italy) and the Valais (Switzerland), between
800 and 2300 m a.s.l.
Remane (1961b), HN; see also Wagner (1955), Dlabola (1971), Günthart (1987a)

Anaceratagallia frisia (W. Wagner, 1939)

This taxon appears to be closely related to A. laevis Ribaut, 1935, which is widespread from
central Asia to western Europe, but it is geographically isolated and confined to Poland (inland
dunes along the lower Oder), northern and middle parts of Germany, the Netherlands (West
Friesian Islands), and perhaps Great Britain (see Remane 1995).
 Agalliinae Kirkaldy, 1901 93

E VI – E V; adult (only ‡‡), 1(?) gen.

Very scattered, but locally abundant in various xerothermic sites with incomplete cover
of vegetation, in dune areas with dominating Corynephorus and Festuca on the East Fri-
sian Islands (Norderney, Baltrum, Langeoog, Spiekeroog, Wangerooge; locus typicus:
Borkum), as well as in steppe-like dry grassland in the Kyffhäuser and along the lower
Oder, one single record is from a mining area to the south of Berlin. Highest localities
are at 250 m a.s.l. The host plants are not known, but may include various species of
Fabaceae or Thymus.
Bröring & Niedringhaus (1989a, 1989b), Niedringhaus & Olthoff (1993), Remane (1995), Schiemenz (1969,
1988), Wagner (1939b), HN

Anaceratagallia ribauti (Ossiannilsson, 1938)

I – XII, mainly VIII – VI; adult (at least a few ††), oocytes overwinter in photoperiodi-
cally controlled oligopause, ‡‡ in thermic quiescence, 1 gen. (Witsack 1985, and pers.
comm.).
In sunny, dry to damp, occasionally also moist sites on acidic to basic substrates, mainly
disturbed, grazed or trampled patches in dry grassland, in low-and medium-input
meadows, abandoned fields, mining areas, along waysides and on inland dunes. Often
syntopic with A. venosa (Geoffr.), but more dominant in acidic and sandy sites and con-
fined to the lowlands. Most populations apparently live on Plantago lanceolata and P. major,
but various dicotyledonous herbs have been reported as additional hosts from labora-
tory rearings (particularly Medicago, Trifolium and Onobrychis, as well as some species of
Lamiaceae and Scrophulariaceae). Widespread in the lowlands of Germany, with strong-
holds in warmer regions and the north German plain, where locally common and dom-
inant in favourable sites, but rare or absent in most parts of the Mittelgebirge, the Alps
and their foreland. The highest localities are only at 500 m a.s.l., but at 1500 m a.s.l. in
the Engadine (Switzerland).
Achtziger (1991), Fischer (1972), Niedringhaus & Olthoff (1993), Post-Plangg & Hoffmann (1982), Reimer
(1992), Remane (1987), Schwoerbel (1957), Wagner (1939a, 1951a), Witsack (1985), Nickel & Achtziger (1999),
HN; see also Günthart (1987a)

Anaceratagallia venosa (Geoffroy, 1785)

Mainly M VII – M X, in southwestern Germany from E V onwards; according to Witsack
(1985) eggs overwinter in secondary parapause terminated by cold and followed by
thermic quiescence, 1 gen.
In similar habitats as A. ribauti (Oss.) and occasionally syntopic, but generally prefer-
ring dryer, more basic and less vegetated sites (including quarries and old farm roads)
and extending up to higher altitudes in the Mittelgebirge and Alps. Lives on low-grow-
ing dicotyledonous plants (apparently mainly Lotus corniculatus, but also reported from
Hippocrepis comosa and Thymus). Widespread in Germany and locally common, usually
among the dominant species in sparsely vegetated calcareous grasslands of southern
and eastern parts of Germany; less common in other habitats, but found to at least
94 Review of species

1000 m a.s.l. in the Erzgebirge, the Bavarian and Black Forest, frequently up to 2000 m
a.s.l. in the Bavarian and Allgäu Alps. In North Tyrol (Austria) breeding has been re-
corded in the alpine belt at 2450 m a.s.l.
Fischer (1972), Müller (1978), Niedringhaus (1991), Reimer (1992), Remane (1987), Rombach (1999b), Wag-
ner (1937b, 1939a, 1951a), Witsack (1985), HN; see also Leising (1977)

Dryodurgades antoniae (Melichar, 1907)

B VIII – B XI; egg, 1 gen.
This is a western European species known only from Portugal, Spain, France, the Neth-
erlands and western Germany. Records from Bulgaria are in need of revision. In Germa-
ny there are 8 known sites. Localities at the edge of the range include Cologne, Braun-
schweig, Marburg and Landau/Pfalz. Lives on Cytisus scoparius in clearings and aban-
doned fields, along margins, embankments, etc.; highest records are from c. 350 m a.s.l.
only.
Frommer (pers. comm.), Jöst (1962), Remane & Fröhlich (1994b), Nickel (1994), HN

Dryodurgades reticulatus (Herrich-Schäffer, 1834)

B VIII – E V; adult, 1 gen.
Along forest margins and in the vicinity of shrubs in xerothermic limestone sites. Vicia
tenuifolia is the host plant, although adults were swept from Pinus sylvestris, where hi-
bernation may take place. In Germany confined to the southern parts, only known from
a few records from near Regensburg (locus typicus), Stuttgart and the Triassic limestone
belt of Mainfranken and adjacent areas of Baden and Thuringia (near Tauberbischof-
sheim, Würzburg, Karlstadt, Bad Neustadt and Meiningen). Highest localities in these
regions are at c. 550 m a.s.l., but at 1300 m a.s.l. in the Engadine (Switzerland).
Remane & Fröhlich (1994b), Wagner (1963), HN; see also della Giustina (1989), Günthart (1987a), Wagner
(1963)

4.2.4.6 Idiocerinae Baker, 1915

Rhytidodus decimusquartus (Schrank, 1776)

B VII – X, occasionally ‡‡ until E IV; mainly egg, 1 gen., published statements of 2 gen.
(Müller 1956) refer to North America.
On Populus nigra nigra and P. n. italica, probably also on hybrids, in river floodplains,
urban settlements and open cultivated areas. Single individuals are swept from other
woody species (e.g. Salix) and often fly into houses during autumn and winter nights.
Widespread in the lowlands of Germany and usually in intermediate frequency, but
low individual numbers; highest localities are at 550 m a.s.l. near the edge of the Alps.
Feldtmann (1938), Fischer (1972), Müller (1956), Niedringhaus & Olthoff (1993), Reimer (1992), Remane
(1987), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1939a, 1951a), HN;
see also Tromellini et al. (1988)
 Idiocerinae Baker, 1915 95

Idiocerus lituratus (Fallén, 1806)

B VII – B X; egg, 1 gen.
Usually in rather cool sites (notably fens, intermediate bogs, meadow valleys, forest glades,
coastal dunes), mainly on hairy-leaved species of willows (Salix cinerea, S. aurita, S. ca-
prea), near the coast preferentially on Salix repens. Single vagrants may be swept from S.
viminalis, S. purpurea, S. alba, S. fragilis and other woody plant species. Widespread in
Germany, occurring in intermediate to high frequency on the host plants, frequently found
up to at least 900 m a.s.l. in the Mittelgebirge and the foothills of the Alps.
Fischer (1972), Niedringhaus (1991, 1997), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b),
Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN

Idiocerus vicinus Melichar, 1898

B VIII – B VI; adult (only ‡‡?), 1 gen.
Along rivers and streams and in gravel pits, occasionally on roadside embankments,
mainly on Salix purpurea, at least occasionally also on S. eleagnos. In Germany found
only along the edge of the Alps and in the upper Rhine plain between Freiburg and
Speyer, between 100 and 1150 m a.s.l., usually in low to intermediate frequency, but
locally in large numbers.
Nickel (1999a), HN

Idiocerus similis Kirschbaum, 1868

M VII – B X; egg, 1 gen.
Monophagous on Salix purpurea along rivers and streams, in sand and gravel pits, occa-
sionally also on roadside embankments. Widespread in middle and southern parts of
Germany, found in low to intermediate frequency and rather low numbers on the host.
The northern edge of the range runs roughly along the northern edge of the Mittelgebir-
ge. Northernmost localities are near Cologne, Göttingen, Hamburg, Lüchow, Nauen and
Potsdam, frequently found up to at least 900 m a.s.l. in the Bavarian Alps.
Frommer (pers. comm.), Günthart (1988), Reimer (1992), Schiemenz (1988), Wagner (1935, 1939a), Nickel &
Remane (1996), Nickel (1994), HN

Idiocerus stigmaticalis Lewis, 1834

M VI – E IX, occasionally until B XI; egg, 1 gen.
On narrow-leaved species of willows near running and standing water, preferentially
on Salix alba, but found in lower numbers on many others, notably S. fragilis, S. triandra,
S. eleagnos, also S. viminalis, S. purpurea, S. caprea, S. cinerea and others; single individu-
als on dispersal flight may be swept from other deciduous trees such as Populus, Betula,
Alnus and Acer. Widespread in Germany and common in most regions, in high frequen-
cy at least on S. alba; the highest records are at 1000 m a.s.l. in the Bavarian Alps on
S. eleagnos.
96 Review of species

Fischer (1972), Lauterer & Malenovsky (1995), Mölleken & Topp (1997), Niedringhaus (1991, 1997), Reimer
(1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Trümbach (1959), Wagner
(1935, 1939a), HN

Idiocerus herrichii (Kirschbaum, 1868)

B VII – E V; adult, 1 gen.
On Salix alba and S. fragilis near running or standing water. Widespread in the lowlands
of Germany, but rather scattered and usually in low frequency and individual numbers,
with strongholds in the north German plain and in larger river valleys (recorded along
the Rhine, Neckar, Main, Lahn, Danube, Isar, Saale and their main tributaries, Elbe,
Oder). The highest localities in these regions are at 600 m a.s.l., but at 1200 m a.s.l. in the
Engadine (Switzerland).
Achtziger (1991), Fischer (1972), Mölleken & Topp (1997), Niedringhaus & Olthoff (1993), Reimer (1992),
Schiemenz (1988), Schönitzer & Oesterling (1998b), SMNS, Wagner (1939a, 1951a), HN; see also Günthart
(1987a)

Metidiocerus elegans (Flor, 1861)

Mainly E VII – X, occasionally III – M VI; egg (also adult?), 1 gen.
On grey-leaved willows (Salix caprea, S. cinerea, S. aurita) in cool and moist situations,
notably meadow valleys, river floodplains, fens and forest glades. Widespread in Ger-
many, but rather scattered, in low frequency and individual numbers, found to at least
1100 m a.s.l. in the Mittelgebirge and the Bavarian Alps, at 1730 m a.s.l. in Switzerland.
Fischer (1972), Niedringhaus & Olthoff (1993), Reimer (1992), Remane & Reimer (1989), Schiemenz (1988),
Wagner (1935), Nickel (1999b), HN; see also Günthart (1987b)

Metidiocerus rutilans (Kirschbaum, 1868)

Adults I – XII, mainly M VII – B VI; adult, 1 gen.
Usually on narrow-leaved species of willows (Salix viminalis, S. purpurea, S. triandra, S.
alba, less frequently S. eleagnos, S. fragilis), occasionally on S. caprea; recorded on Picea
abies in autumn and winter. Widespread in middle and southern parts of Germany and
in intermediate to high frequency on the 4 first-mentioned willow species, but often in
low individual numbers. The edge of the range runs through northern Germany; the
northernmost localities are near Lingen (Ems), Goslar, Dannenberg, Stendal and Neu-
brandenburg. Found up to at least 900 m a.s.l. in the Mittelgebirge, up to 1000 m a.s.l. in
the Bavarian Alps.
Fischer (1972), Mölleken & Topp (1997), Niedringhaus (1997), Schiemenz (1988), Schwoerbel (1957), Wag-
ner (1951a), Nickel (1994), HN

Metidiocerus impressifrons (Kirschbaum, 1868)

Adults I – XII, mainly M VII – M VI; Ad, 1 gen.
 Idiocerinae Baker, 1915 97

On narrow-leaved species of willows, preferentially Salix viminalis, occasionally also S.

purpurea and S. triandra, usually in river floodplains near water. In Germany recorded
only from scattered sites and living at the northern edge of the range, although presum-
ably under-recorded or misidentified. Most localities are along the Rhine, Main, Danube,
Elbe, Saale, Oder and Neiße, single records also from the valleys of the Leine, Ahr, Lahn,
Dill, Lech (near Füssen, 800 m a.s.l.) and Isar (near Mittenwald, 900 m a.s.l.); extending
northward at least to Düsseldorf, Northeim, Lauenburg, Magdeburg, Berlin and An-
germünde. At least single individuals have been found at 1200 m a.s.l. in the Engadine
(Switzerland).
Fischer (1972), Mölleken & Topp (1997), Remane (pers. comm.), Remane & Wachmann (1993), Schiemenz
(1988), Wagner (1939a), Nickel (1994, 1999b), ZIMH, HN; see also Günthart (1987a)

Stenidiocerus poecilus (Herrich-Schäffer, 1835)

M VII – M VI; adult, 1 gen.
On Populus nigra nigra and P. n. italica, usually in river floodplains, although often away
from water. Hibernating individuals were taken from coniferous trees in Piemonte (Ita-
ly). In Germany rather localized and mainly along the larger rivers (Elbe between Dres-
den and Boizenburg, Saale near Jena and Rudolstadt, Main near Aschaffenburg, Lahn
near Marburg, Rhine between Waldshut and Cologne, Nahe near Bad Münster); a few
records also from the surroundings of Berlin and Leipzig, the Oberlausitz, as well as
near Böblingen and Balingen (up to 600 m a.s.l.). The locus typicus is near Regensburg.
Frommer (pers. comm.), Günthart (1988), Remane (pers. comm.), Schiemenz (1988), SMNS, Wagner (1939a,
1951a), HN; see also Giordano et al. (2001)

Tremulicerus tremulae (Estlund, 1796)

M VI – B XI; egg(?), 1 gen.
Monophagous on Populus tremula, mainly in forest glades and along margins, but also
in closed stands. Widespread in middle and northern parts of Germany and in low to
intermediate frequency on the host, but not recorded to the south of the Main, where
perhaps overlooked. Reported from the French side of the upper Rhine plain and most
parts of Austria (although some of these records are in need of revision). The highest
localities are at 700 m a.s.l. in the Mittelgebirge.
Niedringhaus (1997), Reimer (1992), Schiemenz (1988), Schwoerbel (1957), Wagner (1935, 1939a, 1951a),
HN; see also Ribaut (1952), Wagner & Franz (1961)

Tremulicerus vitreus (Fabricius, 1803)

B VI – B X; egg, 1 gen.
On Populus nigra nigra, P. n. italica and their hybrids, in river floodplains and open cul-
tivated land (also in urban settlements). Widespread in lowland areas of middle and
southern parts of Germany and in intermediate to high frequency on the hosts; less
common in northern Germany, where apparently at the edge of the range. Northern-
98 Review of species

most localities are on the island of Norderney, near Lübeck, Halle and Berlin, found
only up to 550 m a.s.l. in the Mittelgebirge and the foreland of the Alps. To the north of it,
there is only one record from the 19th century from Denmark; another record is from
near Stettin (northwestern Poland).
Feldtmann (1938), Günthart (1988), Lauterer & Malenovsky (1995), Schiemenz (1988), Schwoerbel (1957),
Wagner (1935, 1939a, 1941a), ZMB, Nickel & Remane (1996), Nickel (1997), HN; see also Ossiannilsson
(1981)

Tremulicerus distinguendus (Kirschbaum, 1868)

E VI – E X; egg, 1 gen.
On Populus alba in river floodplains, parks and open cultivated land, vagrants also on
other poplar species. Widespread in the lowlands of Germany, occurring northward to
the East Frisian Islands, Holstein and Mecklenburg, usually in intermediate frequency
and rather low individual numbers on the host, to the north of it only known from a few
records from Denmark and southern Sweden. Highest localities are at 500 m a.s.l. in the
Mittelgebirge and the foreland of the Alps.
Frommer (pers. comm.), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Wagner (1935), HN; see
also Gillerfors (2002), Ossiannilsson (1981)

Tremulicerus fulgidus (Fabricius, 1775)

Idiocerus mesopyrrhus Kirschbaum, 1868
Idiocerus fuchsii Kirschbaum, 1868 (see Nickel & Remane 2002)
Adults I – XII, mainly E VII – B VII; adult (also ††), 1 gen.
On Populus nigra italica and P. n. nigra along rivers and streams and in open cultivated
land, occasionally also on poplar hybrids (but breeding has not been recorded on these);
in winter on Picea, reported from France also on Calluna, Hedera, Pinus and Cupressus.
Widespread in middle and southern parts of Germany and usually in intermediate fre-
quency on the host, but confined to lowland areas (up to at most 600 m a.s.l.) and appar-
ently absent from the north, the range extending at least to a line running from Cologne
to Marburg, Göttingen, Stendal and Leipzig.
Frommer (1996), Remane & Fröhlich (1994b), Schiemenz (1988), Nickel (1994), HN; see also Ribaut (1952)

Viridicerus ustulatus (Mulsant et Rey, 1855)

Adults I – XII, mainly M VII – M VI; adult, 1 gen.
On Populus alba in river floodplains, parks and along waysides. Widespread in middle
and southern parts of Germany, occurring in high frequency, although rather low indi-
vidual numbers, but more scattered to the north of it, where presumably only for some
decades. Northernmost records – all dating from the 1990s and many from urban settle-
ments – are from the island of Norderney, near Göttingen, Dannenberg, Berlin and Frank-
furt an der Oder; found only up to 500 m a.s.l. in the foreland of the Alps.
 Idiocerinae Baker, 1915 99

Fischer (1972), Frommer (pers. comm.), Remane & Fröhlich (1994b), Remane & Wachmann (1993), Wagner
(1939a, 1951a), Weis (pers. comm.), Nickel (1994), HN

Populicerus albicans (Kirschbaum, 1868)

M VI – B X; egg, 1 gen.
Monophagous on Populus alba in river floodplains, parks and along waysides, also in
urban settlements. Widespread in the lowlands of Germany (up to at least 600 m a.s.l.)
and usually in high frequency and individual numbers on the host, even on much iso-
lated tree individuals.
Fischer (1972), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b),
Wagner (1935, 1939a, 1951a), HN

Populicerus confusus (Flor, 1861)

M VI – B X; egg, 1 gen.
In river floodplains, along streams, in forest glades and among subalpine scrub, mainly
on grey-leaved species of willows (Salix cinerea, S. aurita, S. caprea, S. viminalis, in the
Alps and their foothills also S. myrsinifolia and S. appendiculata), occasionally found in
rather low numbers on additional species (S. alba, S. fragilis, S. triandra). Widespread in
Germany and fairly common, occurring up to at least 1200 m a.s.l. in the Mittelgebirge,
up to 1500 m a.s.l. in the Bavarian Alps.
Fischer (1972), Mölleken & Topp (1997), Niedringhaus (1991, 1997), Reimer (1992), Remane (1987), Remane
& Reimer (1989), Schiemenz (1988), Wagner (1935), 1939a, 1951a), HN and others

Populicerus nitidissimus (Herrich-Schäffer, 1835)

B VI- M X; egg, 1 gen.
On Populus nigra italica, P. n. nigra and their hybrids, in cultivated land, urban settle-
ments and river floodplains (also in closed forests). Widespread in the lowlands of Ger-
many, in high frequency and individual numbers on the hosts, although less common in
the north German plain, extending only up to 600 m a.s.l. in the Mittelgebirge and the
foreland of the Alps.
Büchs (1988), Günthart (1988), Mölleken & Topp (1997), Niedringhaus (pers. comm.), Reimer (1992), Re-
mane (1987), Schiemenz (1988), ZIMH, HN

Populicerus laminatus (Flor, 1861)

M VI – M X; egg, 1 gen.
Monophagous on Populus tremula, mainly in forests and along their margins. Wide-
spread in Germany, apparently only in low to intermediate frequency, but locally high
individual numbers. A single tree stem eclector catch in northern Thuringia revealed
33.000 individuals within one season. The highest localities are at 700 m a.s.l. in the
Mittelgebirge and the foothills of the Alps.
100 Review of species

Frommer (1996), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a,
1951a), HN

Populicerus populi (Linnaeus, 1761)

M VI – M X; egg, 1 gen.
Like the preceding species monophagous on Populus tremula, but more eurytopic, in
higher frequency and usually high individual numbers, often also found on solitary
trees and shrubs. Widespread in Germany, extending up to at least 1000 m a.s.l. in the
Mittelgebirge and the Bavarian Alps, found at 1480 m a.s.l. in Switzerland.
Achtziger (1991), Fischer (1972), Niedringhaus (1991, 1997), Reimer (1992), Remane & Reimer (1989), Schi-
emenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1951a), HN; see also
Günthart (1987b)

Acericerus heydenii (Kirschbaum, 1868)

E VII – E VI; adult, 1 gen.
Usually on Acer pseudoplatanus in forests, river floodplains and urban settlements, in
lower numbers also on A. platanoides, A. campestre and A. monspessulanum. Often flies
into houses during autumn and winter nights. Widespread in middle and southern
parts of Germany and – at least on A. pseudoplatanus – in intermediate frequency, but
low individual numbers. The northernmost localities are near Cologne, Göttingen, Kiel,
Stendal, Berlin and Fürstenwalde. Found up to at least 1000 m a.s.l. in the Bavarian
Forest, up to at least 1250 m a.s.l. in the Bavarian and Allgäu Alps.
Fischer (1972), Frommer (pers. comm.), Reimer (1992), Remane (pers. comm.), Remane & Wachmann (1993),
Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1939a), Weis & Schönitzer (2001), Nickel (1994),
HN

Acericerus ribauti Nickel & Remane, 2002

Idiocerus rotundifrons Kirschbaum, 1868 sensu Ribaut (1952) (see Nickel & Remane 2002)
Adults I – XII, mainly B VIII – V; adult, 1 gen.
Preferentially on Acer campestre, in lower numbers also on A. pseudoplatanus and A. pla-
tanoides, usually along sun-exposed forest margins, in parks and urban settlements, fly-
ing into houses during autumn and winter nights. Widespread in middle and southern
parts of Germany and in intermediate to high frequency on A. campestre, but at the edge
of the range; northernmost localities are near Cologne, Göttingen, Halle and Berlin,
mainly below 600 m a.s.l., but recorded at 800 m a.s.l. on the Swabian Jura.
Fischer (1972), Frommer (1996), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1988), Schönitzer
& Oesterling (1998b), Weis & Schönitzer (2001), Nickel (1994), HN

Acericerus vittifrons (Kirschbaum, 1868)

Idiocerus rotundifrons Kirschbaum, 1868 (see Nickel & Remane 2002)
Adults I – XII, mainly B VIII – V; adult, 1 gen.
 Iassinae Amyot & Serville, 1843 101

Life history and distribution in Germany are much as in A. ribauti Nick. & Rem.; the
species occurs on Acer campestre and A. pseudoplatanus, presumably also on A. platanoides,
but is apparently more confined to lower altitudes, so far found only below 600 m a.s.l.
The northernmost localities are near Cologne, Göttingen, Stendal and Potsdam, anoth-
er site near Neubrandenburg may be rather isolated, although there is a very recent
record from southern Sweden (Skåne).
Büchs (1988), Frommer (1996), Kuntze (1937), Reimer (1992), Schiemenz (1988), Schwoerbel (1957), Wagner
(1951a), Weis & Schönitzer (2001), Nickel (1994), HN; see also Gillerfors (2002)

Balcanocerus larvatus (Herrich-Schäffer, 1835)

E VI – M X; egg, 1 gen.
On Prunus spinosa in sun-exposed, damp to dry sites, mainly pastures and meadows
with scattered shrubs, embankments, forest margins and hedges. In Germany wide-
spread between the Danube and the northern edge of the Mittelgebirge, extending to at
least 750 m a.s.l. and occurring in high frequency at least in warmer regions, although
seldom recorded. Northernmost localities are near Bonn, Brilon, Göttingen, Wolfenbüt-
tel, Halle, Leipzig and Dresden; there is also an old record from near Neubrandenburg.
Not reported from Denmark and Scandinavia.
Achtziger (1991, 1994), Heller (1987a), Kuntze (1937), Post-Plangg & Hoffmann (1982), Remane & Fröhlich
(1994b), Rombach (1999a), Schiemenz (1988), Schwoerbel (1957), Trümbach (1959), Wagner (1939a, 1951a),
Walter (1998), Nickel (1994), HN; see also Ossiannilsson (1981)

Balcanocerus pruni (Ribaut, 1952)

M VIII – B VI; adult, 1 gen.
Like the preceding species on Prunus spinosa, but more thermophilous, in Germany large-
ly confined to viticultural regions of southwestern parts. Recorded from the Ahr valley
near Altenahr, the middle Rhine near Cologne and Bacharach, the Nahe valley near
Schloßböckelheim and Mainfranken (Karlstadt and Hammelburg), between 50 and 350 m
a.s.l. Otherwise only reported from France, Italy, the former Yugoslawia, Bulgaria, Greece
and southern Russia (Dagestan).
Frommer (pers. comm.), Remane & Fröhlich (1994b), Nickel & Remane (1996), HN; see also Dlabola (1961a),
Drosopoulos et al. (1986), Nast (1987)

4.2.4.7 Iassinae Amyot & Serville, 1843

Batracomorphus allionii (Turton, 1802)

E VI – B IX; egg, 1 gen.
In forest glades, clearings, heaths, etc., usually in oligotrophic, more or less acidic, damp
to moderately dry sites. The host plant is Cytisus scoparius, occasionally also Genista
tinctoria and perhaps additional species of woody Fabaceae. The distribution in Germa-
ny is imperfectly known; strongholds are apparently in sand and sandstone regions in
102 Review of species

western parts (northern upper Rhine plain, Pfälzer Wald, middle Hessen), although
there are widely scattered records from most other regions, only up to 450 m a.s.l.
Fischer (1972), Frommer (1996), Jöst (1966), Reimer (1992), Remane (pers. comm.), Schiemenz (1988), Wag-
ner (1935, 1939a), Nickel (1999b), HN

Batracomorphus irroratus Lewis, 1834

E VI – M IX; egg, 1 gen.
In xerothermic grassland and along sunny margins of pine and oak forests, usually on
limestone and other basic substrates. The main host plant is Helianthemum nummulari-
um, although other species of Helianthemum have rarely been swept. In Germany occur-
ring rather localized and only in middle parts, mainly in limestone regions (Saale-Un-
strut region, southern Niedersachsen, Thuringian Basin, Eifel Mountains, Rhine-Main
region, Mainfranken, Tauber valley, as well as the Franconian and Swabian Jura, usual-
ly found below 500 m a.s.l., but up to at least 800 m a.s.l. on the Swabian Jura. The
northern edge of the range roughly follows a line from Bonn to Marburg, Göttingen,
Wernigerode, Eisleben and Leipzig.
Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Remane (pers. comm.), Rombach (1999a),
Schiemenz (1988), SMNS, Wagner (1939a, 1951a), Nickel (1994), HN

Iassus lanio (Linnaeus, 1761)

E VI – B XI; egg, 1 gen.
Eurytopic on oaks (Quercus robur, Qu. petraea), in the interior of forests as well as along
their margins and on solitary trees, also in urban settlements. Widespread in the low-
lands of Germany, in high frequency on both oak species, also on the East Frisian Is-
lands, extending to at least 600 m a.s.l. in the foothills of the Alps.
Achtziger (1991), Fischer (1972), Gotsmy & Schopf (1992), Niedringhaus (1991), Reimer (1992), Remane
(1987), Schiemenz (1988), Wagner (1935, 1939a), HN and others

Iassus scutellaris (Fieber, 1868)

E VI – B X; egg, 1 gen.
Monophagous on elms, usually on Ulmus minor and U. x hollandica, perhaps also on U.
laevis, but not recorded from U. glabra. Usually along alleys and hedges and on solitary
trees, mainly in river floodplains, parks and along waysides, presumably also in closed
forests, where difficult to sample with the sweep-net. Widespread in lowland middle
parts of Germany (to at least 500 m a.s.l.), in high frequency but low individual num-
bers at least on U. minor and U. x hollandica. Furthermore, there are single records from
the area to the south of the Danube and the north German plain. Northernmost locali-
ties are near Cologne, Bielefeld, Hanover, Hamburg, Parchim and Rostock. Apparently
absent from Denmark and Scandinavia.
Fischer (1972), Feldtmann (1937), Frommer (1996), Remane (1987), SMNS, Wagner (1935), ZIMH, Nickel
(1994), HN; see also Ossiannilsson (1981)
 Penthimiinae Kirschbaum, 1868 103

4.2.4.8 Penthimiinae Kirschbaum, 1868

Penthimia nigra (Goeze, 1778)

B V – M VIII; nymph, 1 gen.
On shrubs and low-growing trees in dry to damp sites in warm situations (notably
abandoned vineyards, dry grassland, ruderal sites, forest margins, coppice-with-stan-
dards, etc.). Food plants are various deciduous shrubs and trees (adults found on Quer-
cus, Populus and others). Occurs in Germany only locally and at the northern edge of the
range (usually below 400 m a.s.l.), with strongholds in viticultural regions, notably along
the upper and middle Rhine and its tributaries (Ahr, Moselle, Nahe, Main), the Elbe
near Dresden and Meißen, as well as in the Saale-Unstrut region between Naumburg,
Halle and Leipzig; further localities, some of which appear to be rather isolated, are
near Augsburg, Bonn, Berlin and in the Oberlausitz.
Fischer (1972), Maixner (pers. comm.), Post-Plangg & Hoffmann (1982), Remane (1987, and pers. comm.),
Remane & Wachmann (1993), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1939a, 1951a), HN

4.2.4.9 Dorycephalinae Oman, 1943

Eupelix cuspidata (Fabricius, 1775)

I – XII, probably semivoltine, hibernation as nymph and adult.
In sunny to moderately shady, dry to damp, occasionally also moist or even temporari-
ly wet sites (various types of dry grassland, dunes, low-productivity meadows and
pastures, ruderal sites, waysides, sunny forest margins, moderately saline sites, etc.),
mainly on Festuca ovina, perhaps also on F. rubra and additional grasses. Widespread in
Germany and fairly common in favourable sites, recorded up to 1300 m a.s.l. in the
Black Forest, up to 1500 m a.s.l. in the Engadine (Switzerland).
Bornholdt (1996), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Reimer (1992), Remane (1987),
Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1939a, 1951a), Nickel & Achtziger (1999), HN;
see also Günthart (1987a)

4.2.4.10 Aphrodinae Haupt, 1927

For a long time, there have been diverging opinions on systematics of the genus Aphrodes Curt.
(s.str.). Until recently, most authors distinguished only two species in central Europe (Emmrich
1980; Nast 1976b). The interpretation followed here is adopted from Tishechkin (1998), who
essentially clarified biosystematic, although not nomenclatural problems. Accordingly, the Aph-
rodes bicincta group includes 4 distinct species with differences in bioacoustics, morphology and
life history (see below). Hence, most published records from central Europe are in need of revi-
sion. Furthermore, there is a number of older names, which might have priority for some taxa,
but this would require investigation of a vast material. The grammatical gender of Aphrodes is
female (see Metcalf 1963).

Aphrodes aestuarina (Edwards, 1908)

Perhaps only a subspecies or ecological form of A. makarovi Zachv.
104 Review of species

In England VII – IX; egg(?), 1 gen. (Kirby 1992).

At present this taxon is only known from coastal regions of the North and Baltic Sea
(England, Poland, Niedersachsen). Recorded in Germany only from the East Frisian
Islands and the vicinity of Cuxhaven, where it is probably restricted to coastal salt marsh-
es and moderately saline meadows and pastures. The host plants are unknown.
Niedringhaus (1991, and pers. comm.), Wagner (1937c); see also Kirby (1992); for identification see Tishech-
kin (1998)

Aphrodes bicincta (Schrank, 1776) sensu Tishechkin (1998)

Aphrodes makarovi Zachv. sensu Ossiannilsson (1981)
B VI – X, single specimens until I; egg, 1(?) gen.
Due to very recent taxonomic changes this species’ distributional status in central Eu-
rope is rather unclear. Apparently mainly in sun-exposed, damp to dry, basic to acidic
sites, notably calcareous hillsides, inland dunes, heaths, low-productivity meadows,
dry waysides and abandoned fields. Probably widespread in Germany, but with strong-
holds in the lowlands, most verified records are from below 500 m a.s.l. Host plants are
species of Fabaceae and perhaps other plant families. Was identified as a vector of Clo-
ver phyllody (CP), Clover dwarf (CD) and perhaps other plant diseases in various parts
of Europe, but the species identity of these populations is uncertain (see above). The
latter is also true for populations found in inland salt marshes and on dry hillsides at
higher altitudes of the Alps and the Mittelgebirge.
Emmrich (1980), HN; see also Brčák (1979); for identification see Tishechkin (1998)

Aphrodes diminuta Ribaut, 1952

Aphrodes centrorossica Zachvatkin, 1953; A. bicincta (Schrank, 1776) sensu Ossiannilsson (1981)
E VI – E X; egg, 1 gen.
In oligotrophic, wet to temporarily dry sites, mainly in fens dominated by Carex, in low-
input meadows and pastures. Is reported to live on species of Fabaceae in Russia. The
distribution in Germany is imperfectly known, but the species is probably widespread,
although uncommon, with strongholds in the submontane and montane belt in middle
and southern parts, but also in the north German plain. It is frequently found at 1200 m
a.s.l. in the Mittelgebirge, up to at least 1500 m a.s.l. in the Bavarian Alps.
Emmrich (1980), Remane (pers. comm.), Remane & Wachmann (1993), HN; for identification see Tishech-
kin (1998)

Aphrodes makarovi Zachvatkin, 1948

Mainly E VI – E X, single individuals occasionally until winter; egg, 1 gen.
Eurytopic in moderately wet to damp, sunny to shady sites, often in disturbed or
eutrophic patches, notably fertilized meadows and pastures, tall herb and grass stands,
abandoned fields, ruderal sites, shores of running and standing water, ditches, forest
roads, alder fen woods, etc., also slightly saline sites inland and near the coast. Host
 Aphrodinae Haupt, 1927 105

plants are Taraxacum, Cirsium, Urtica dioica, Rumex, and probably additional dicotyle-
donous herbs. Widespread and common in the lowlands of Germany, but usually below
c. 800 m a.s.l., although imperfectly documented.
Remane (pers. comm.), HN; for identification see Tishechkin (1998)

Planaphrodes bifasciata (Linnaeus, 1758)

B VI – E IX; egg, 1 gen.
In sunny to shady, moderately wet to damp sites, mainly meadows and pastures of the
submontane belt dominated by Nardus or Trisetum and dicotyledonous herbs, also in
lowland areas in moist deciduous forests with a luxuriant herb layer (e.g. floodplain
and fen woods), among stands of tall herbs as well as along margins of bogs. No host
plants are known from central Europe, but in field choice experiments conducted in
Britain, adults and nymphs have been found in numbers on several grasses, notably on
Helictotrichon pubescens, Brachypodium pinnatum, Briza media, Bromus erectus, Festuca ovi-
na, F. rubra, Trisetum flavescens, as well as on the sedge Carex flacca. The distribution in
Germany is imperfectly documented due to secretive habits. Apparently, most localities
are in the Alps and the Mittelgebirge (mainly between 400 and 1200 m a.s.l., up to 1500 m
a.s.l. in Switzerland), e.g. Solling, Harz and Rhön Mountains, Frankenwald, Thuringian
Forest, Erzgebirge and Vogelsberg, but there are also records from valley situations and
the north German plain.
Bornholdt (1996), Fischer (1972), Nikusch (1976), Reimer (1992), Remane & Wachmann (1993), Schiemenz
(1988), Schönitzer & Oesterling (1998b), Wagner (1937c), Nickel & Achtziger (1999), HN; see also Cook
(1996), Günthart (1987a)

Planaphrodes nigrita (Kirschbaum, 1868)

M VI – E X; egg, 1 gen.
Like the preceding species, mainly in mountainous areas, often syntopic and difficult to
distinguish ecologically. Less frequently found, in the lowlands apparently preferring
forests in damp sites, but rather wet habitats at higher altitudes (e.g. spring mires, bog-
gy spruce forests, occasionally grassland). The host plants are unknown, a few individ-
uals were taken from Calamagrostis canescens and Juncus articulatus, but the species oc-
curs in many sites where these plants are absent. Widespread in Germany, but imper-
fectly documented, from lowland areas up to at least the upper montane belt, found up
to 1500 m a.s.l. in the Bavarian Alps, at 2250 m a.s.l. in North Tyrol (Austria).
Emmrich (1969), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer (1992), Remane & Wachmann
(1993), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1937c), Nickel & Achtziger (1999),
Nickel & Remane (1996), Nickel (1997), HN; see also Christandl-Peskoller & Janetschek (1976)

Planaphrodes trifasciata (Geoffroy, 1785) sensu Ribaut (1952)

The name given in the original description is not valid, since it is a primary homonym of Cicada
trifasciata De Geer, 1773, which is in turn a junior synonym of P. bifasciata (Linnaeus, 1758) (see
Metcalf 1963). Therefore, Hamilton (1975) has, apparently without studying type material, pro-
106 Review of species

posed the name P. laeva (Rey, 1891), the description of which is based only on characters of
colouration. Thus, Rey’s material is in need of revision in order to clarify the nomenclatural
situation.
E VI – E X; egg, 1 gen.
In sunny, occasionally also slightly shady, often grazed or sparsely vegetated sites on
acidic as well as basic substrates, mainly bogs, heaths and xerothermic grassland, rarely
in open pine forests. The host plants are not known; but the species may be associated
with dwarf shrubs (mainly Calluna and Thymus). Widespread in Germany, with strong-
holds in bog and heath areas of the north German plain, the Mittelgebirge and the Alps,
as well as in warmer regions of southern and middle parts. Highest localities are at
1000 m a.s.l. in the Thuringian Forest and at 1300 m a.s.l. in the Chiemgau Alps.
Achtziger (1991), Fischer (1972), Remane (1987), Schiemenz (1969, 1975, 1988), Schwoerbel (1957), Wagner
(1935, 1939a, 1951a), HN

Anoscopus albifrons (Linnaeus, 1758)

M VI – E X, single individuals until winter; egg, 1 gen.
In sunny to moderately shady, moderately dry to moist sites, mainly heaths, open for-
ests, low-productivity meadows and pastures. Host plants are various grasses (Des-
champsia flexuosa, Holcus mollis, Brachypodium pinnatum and others). Widespread in Ger-
many and common in favourable habitats, also in higher parts of the Mittelgebirge, up to
at least 1500 m a.s.l. in the Bavarian Alps.
Achtziger (1991), Fischer (1972), Haas (1975), Niedringhaus (1991, 1997), Reimer (1992), Schiemenz (1988),
Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN

Anoscopus limicola (Edwards, 1908)

M VII – E IX; egg, 1 gen.
This species is apparently an endemic of western European coasts, living among grass-
es in salt marshes. The host plant is probably Puccinellia maritima. In Germany it is known
only from the islands of Sylt and Amrum, all East Frisian Islands, where locally domi-
nant, as well as from a few mainland sites (near Varel, Bremerhaven and Cuxhaven).
Further verified records exist only from Ireland, England, southern Sweden, the Neth-
erlands and the Atlantic and Mediterranean coasts of France.
Hildebrandt (1995), Niedringhaus (1991), Niedringhaus & Olthoff (1993), Wagner (1937c), ZIMH; see also
della Giustina & Remane (2001), Gravestein (1965), Kirby (1992), Nast (1987), Ossiannilsson (1981)

Anoscopus albiger (Germar, 1821)

E VI – M XI; egg, 1 gen.
Usually in moderately saline sites inland and near the coast, also in ruderal habitats
(preferentially coal mining areas) and peaty sedge swamps. Host plants are not known,
but are likely to include grasses. Rather scattered in Germany, reaching the northern
edge of the range; mainly along the coast of the North Sea and in inland salt marsh and
 Aphrodinae Haupt, 1927 107

coal mining areas of eastern parts, notably Saxony-Anhalt, Thuringia and Saxony, sin-
gle records are from near Oldenburg, Wolfenbüttel, Göttingen, Cologne, Gießen, Frank-
furt am Main, as well as the French part of the upper Rhine plain, usually below 250 m
a.s.l. To the north of Germany it is known only from the Baltic Sea islands of Bornholm
and Gotland.
Fröhlich (1996a), Frommer (1996), Funke & Witsack (1998), Niedringhaus & Olthoff (1993), Schiemenz (1988),
Wagner (1937c, 1939a), Nickel (1997), HN; see also Gillerfors (2002), Ribaut (1952)

Anoscopus alpinus (W. Wagner, 1955)

Described from northeastern parts of the Alps, but is perhaps conspecific with A. assimilis (Si-
gnoret, 1879), which is reported mainly from southwestern parts of Europe. The description of
A. duffieldi (Le Quesne, 1964), known only from a single coastal site in England (see also Remane
& Fröhlich 1994b) is based merely upon slight differences of the aedeagus tip, body colouration
and vertex length. However, in most Anoscopus species, these characters are subject to consider-
able intraspecific variation. Hence, morphological and zoogeographic evidence suggests that at
least the two latter taxa are conspecific. In France A. assimilis (Sign.) is reported from coastal
lowlands of western and northwestern parts (Ribaut 1952; della Giustina 1989).
M VII – E IX; egg, 1 gen.
In heaths and bogs dominated by dwarf shrubs, extending from the submontane to the
alpine belt. In Germany largely confined to the Bavarian and Allgäu Alps (between
1100 and 2100 m a.s.l., but up to 2970 m a.s.l. in North Tyrol, Austria) and to the higher
Mittelgebirge, notably the Erzgebirge, Thuringian Forest and southern Black Forest (be-
tween 650 and 1450 m a.s.l.) The host plants are presumably grasses associated with
Vaccinium myrtillus, Calluna vulgaris and other dwarf shrubs.
Remane & Fröhlich (1994b), Schiemenz (1971a, 1975, 1988), Walter (pers. comm.), Nickel & Voith (unpub-
lished data), Nickel (2002); see also Christandl-Peskoller & Janetschek (1976), Wagner (1955)

Anoscopus flavostriatus (Donovan, 1799)

B VII – B XI, according to Schiemenz (1988) single individuals until IV; egg, 1 gen.
In grass stands of various types, usually in rather cool, wet to damp sites (mainly low-
input meadows and pastures, forest glades, fens, abandoned fields and open forests).
The host plants are grasses (Deschampsia flexuosa, Dactylis glomerata, Elymus repens and
others). The species is widespread in Germany and fairly common in most regions (up
to at least 1000 m a.s.l.), more scattered or locally absent only in warm and dry areas
(notably the rain shadow area to the east of the Harz Mountains, Mainfranken, upper
Rhine plain).
Achtziger (1991), Fischer (1972), Niedringhaus (1991), Reimer (1992), Remane (1958), Schiemenz (1988), Schönitzer
& Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN

Anoscopus histrionicus (Fabricius, 1794)

Despite distinctive colouration of ††, this species has been misidentified or misinterpreted. Thus,
a number of records compiled by Schiemenz (1988) are in need of revision and refer to other
species, including most, if not all Thuringian records.
108 Review of species

M VI – M IX; egg, 1 gen.

Is reported to live in moist to moderately dry, usually sunny and oligotrophic sites, e.g.
heaths, low-input meadows and open forests. Host plants are probably grasses. In Ger-
many mainly in northern and eastern parts and perhaps at the edge of the range. The
westernmost localities are on the East Frisian Islands from Borkum to Wangerooge, near
Hamburg, Hanover, Dessau, Leipzig, Fürth, Regensburg and Munich, up to 500 m a.s.l.
The majority of records dates back at least 40 years, more recent ones are only known
from the East Frisian Islands. In France the species is apparently confined to the Alps
and the Pyrenees, but it is reported to occur in most European countries including the
Netherlands, Belgium and the British Islands.
Herrich-Schäffer (1834), Niedringhaus (1991), Schönitzer & Oesterling (1998b), Wagner (1935); see also
della Giustina (pers. comm.), Holzinger (pers. comm.), MNHN (Bourgoin pers. comm.), Nast (1987)

Anoscopus serratulae (Fabricius, 1775)

B VII – M X; egg, 1 gen.
In grass stands in moderately dry to moist, occasionally also wet, usually sunny sites,
mainly more or less strongly fertilized meadows and pastures, ruderal sites, lawns in
parks and gardens, also moderately saline sites. Host plants are grasses (Elymus repens,
Dactylis glomerata, Holcus spp.). The species is widespread in the lowlands of Germany
and fairly common, the highest localities are at c. 700 m a.s.l. only, but it is recorded
from 900 m a.s.l. in North Tyrol (Austria).
Achtziger (1991), Niedringhaus (1991), Nikusch (1976), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwo-
erbel (1957), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN; see also Günthart & Thaler (1981)

Stroggylocephalus agrestis (Fallén, 1806)

M VII – E X, at least occasionally until VI; egg(?), 1 gen.
In sedge stands in various wet sites (mainly fens, intermediate bogs, wet meadows, fen
woods, also slightly saline sites); host plants are usually tall species of Carex (C. acutifor-
mis, C. acuta, C. nigra and others), perhaps also Bolboschoenus maritimus. Widespread in
the lowlands of Germany and fairly common in favourable sites; the highest localities
are at c. 800 m a.s.l.
Fischer (1972), Hoffmann (1980), Kuntze (1937), Reimer (1992), Schiemenz (1976, 1988), Schönitzer & Oester-
ling (1998b), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN

Stroggylocephalus livens (Zetterstedt, 1840)

E VII – E VI; adult (and nymph?), 1 (perhaps only half) gen.
This is a tyrphophilous species usually living in intermediate bogs and fens, but it is
also reported from raised bogs (where probably restricted to margins) and birch fens.
Presumably it lives on Carex, perhaps on Eriophorum. Uncommon in Germany and mainly
in remnant bog areas of the north German plain, small populations are also found in the
Bavarian and Swabian foothills of the Alps and the Mittelgebirge (Harz Mountains, Thu-
 Cicadellinae Latreille, 1825 109

ringian Forest, Erzgebirge, Rothaargebirge, Rhön Mountains, Fichtelgebirge), as well as

in the upper Rhine plain, up to at least 900 m a.s.l.
Achtziger (pers. comm.), Bittner & Remane (1977), Hoffmann (1980), Kuntze (1937), Schönitzer & Oester-
ling (1998b), Remane (1958), Schiemenz (1971a, 1973, 1975, 1976, 1988), Wagner (1935), Walter (pers. comm.),
Nickel (2002), HN

4.2.4.11 Cicadellinae Latreille, 1825

Evacanthus acuminatus (Fabricius, 1794)

B VI – B IX; egg, 1 gen.
In moderately shady to shady, moist to moderately dry, more or less eutrophic sites,
usually with a luxuriant herb layer, mainly in deciduous and mixed forests of various
types and along woody margins (in dry grassland, abandoned vineyards, etc.); absent
from mown meadows. Host plants are various dicotyledonous herbs; adults were taken
from Glechoma, Lamium, Stachys, Valeriana, Verbascum and others. Widespread in Ger-
many, particularly in middle and southern parts, although usually collected in low in-
dividual numbers only, more scattered in the north German plain. Frequently found up
to 1200 m a.s.l. in the Bavarian and Allgäu Alps, up to at least 1500 m a.s.l. in Carinthia
(Austria) and the Engadine (Switzerland).
Büchs (1988), Fischer (1972), Niedringhaus (1991), Post-Plangg & Hoffmann (1982), Reimer (1992), Schie-
menz (1988), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN and others; see also Günthart (1987a),
Prohaska (1923)

Evacanthus interruptus (Linnaeus, 1758)

M VI – M X; egg, 1 gen.
In sunny to shady, wet to damp, usually rather eutrophic sites, mainly in low-input
meadows, fens, along various herbaceous and woody margins (shores of running or
standing water, forest roads and edges, ditches, etc.), in forests and subalpine tall herb
stands. Adults on Arctium, Epilobium angustifolium, Cirsium, Eupatorium, Lamium, Peta-
sites, Senecio, Urtica and many others, occasionally also Rubus idaeus. Widespread and
common in Germany, particularly in rather cool situations (but apparently absent from
the East Frisian Islands). Large populations have been found up to at least 1850 m a.s.l.
in the Bavarian and Allgäu Alps; the highest localities in North Tyrol (Austria) and the
Engadine (Switzerland) are at 2200 m a.s.l.
Achtziger (1991), Fischer (1972), Niedringhaus (1991), Reimer (1992), Remane & Wachmann (1993), Schie-
menz (1988), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN and others; see also Günthart (1987a)

Errhomenus brachypterus Fieber, 1866

Has been subject to taxonomic confusion, which was finally cleared by Lauterer (1983). Accord-
ingly, there is a considerable intraspecific variability of colouration, head shape and body sculp-
turation, resulting in the description of Errhomenellus flavopunctatus Melichar, 1914, which is,
however, a younger synonym.
110 Review of species

I – XII, probably semivoltine, hibernation as nymph and adult.

In the litter layer of deciduous and coniferous forests (mainly under Fagus, Quercus,
Picea, Abies and Pinus) in wet to damp, basic as well as acidic sites, also found up to at
least 1850 m a.s.l. in subalpine scrub in the Bavarian and Allgäu Alps (reported to live
under Alnus alnobetula, Pinus mugo and Rhododendron, up to 2000 m a.s.l. in Austria).
Usually recorded in pitfall traps and sieved litter, rarely in sweep-net samples and un-
der stones. The host plants are unknown. Apparently, there is not a single plant species
or even a plant family common to all sites. Moreover, there is often a complete lack of
any herbaceous undergrowth. Thus, the species may live polyphagously on fine roots
of trees and shrubs, which are within easy reach of its long rostrum. The distribution in
Germany is imperfectly documented, but it seems to be widespread and locally even
common in southern and middle parts (mainly above 300 m a.s.l.). According to present
data, the border of the range runs along des northern edge of the Mittelgebirge (north-
ernmost localities are on a line from Aachen to Dortmund, Minden, Goslar, Quedlin-
burg, Leipzig and Dresden); a singe record from near Hamburg dates from 1895. Large
numbers have been sampled in acidic and boggy spruce forests in higher parts of the
Harz Mountains, Thuringian Forest and Erzgebirge, also in limestone beech forests of
southern Niedersachsen and northern Hessen and in mixed forests on Cretaceous chalk
and dolomite in the Veldensteiner Forst near Nuremberg. More or less scattered records
are known from other regions (Solling, Eifel Mountains, Franconian Jura, Breisgau,
Swabian and Bavarian Allgäu, also in the Vosges (eastern France) and the southern
Dutch province of Limburg.
Achtziger (1991), Coll. Bauchhenss, Drees (2000), Haupt (1935), Schiemenz (1971a, 1975, 1988), Coll. Sprick,
SMNS, Wagner (1938c), Nickel (1994), HN; see also Cobben & Rozeboom (1978), Hölzel (1965), Wagner &
Franz (1961)

Cicadella lasiocarpae Ossiannilsson, 1981

So far E VI – B IX; egg, 1 gen.
Was described not long ago from Sweden, but has been found in large parts of the
Palearctic since (from eastern Siberia and Korea to Ireland, Wales and the northern half
of Germany). At present there are about 20 known localities in Germany from near
Lüchow, Haldensleben, Cologne, Dresden, Bautzen, Marburg, and Fulda, up to 300 m
a.s.l., usually in wet to peaty, mesotrophic sites (peaty meadows and pastures, fens and
intermediate bogs), often with dominating sedges, particularly Carex nigra. In most cas-
es it occurs at high densities, syntopically with C. viridis (L.).
Frommer (pers. comm.), Remane (pers. comm.), Remane & Fröhlich (1991), Nickel & Achtziger (1999),
Nickel (1997), HN; see also Dmitriev (1998), Le Quesne (1987), Ossiannilsson (1981), Gȩbicki & Szwedo
(1998), Tishechkin (2000), Walter (pers. comm.)

Cicadella viridis (Linnaeus, 1758)

Mainly B VII – M X; 1 gen.; in warmer regions E V – B XI, 2 gen., according to Witsack
(1985), hibernation takes place in the embryonic stage, in secondary parapause termi-
nated by cold.
 Typhlocybinae Kirschbaum, 1868 111

In various sunny to moderately shaded, temporarily flooded to moist, occasionally also

rather dry sites. Polyphagous on rushes, sedges and grasses, probably also on dicotyle-
dons, although most commonly on Juncus. Oviposition into saplings and low branches of
trees has been reported to cause damage in forest and fruit-tree plantations. Highest abun-
dances usually occur in fens, intermediate bogs, wet meadows and pastures, but the spe-
cies is also found along bog margins, in open forests, clearings, ruderal sites, mining ar-
eas, along forest roads and ditches, occasionally even in dry grassland (including nymphs).
It is widespread in Germany and common in most regions, in the Bavarian Alps frequent-
ly found up to at least 1200 m a.s.l., but also in the subalpine belt in Styria (Austria).
Fischer (1972), Müller (1956, 1972), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Schmutterer (1953),
Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN and others; see also Wagner & Franz (1961)

Graphocephala fennahi Young, 1977

M VII – M X, in southwestern Germany B VII – XI; egg, 1 gen.
This is a Nearctic species which was introduced probably with its host plants and which
occurs in Europe at least since the 1930s, initially in England, since the 1970s also in
Switzerland, France, the Netherlands and Germany (at that time generally reported as
G. coccinea Forster, 1771). Discovered in Germany in 1978 in Mönchengladbach, after-
wards found in Oldenburg in 1982, in Düsseldorf, Frankfurt am Main and Stuttgart in
1983, in Bremen and Hamburg in 1984, in Berlin in 1988. At present, it is widespread in
much of the western half below 550 m a.s.l., and particularly common in the basins of
the Rhine and Ems. In recent years there is a number of records from the eastern half,
indicating a further eastward range expansion. Hamburg, Dannenberg, Hanover,
Gardelegen, Berlin, Göttingen, Freiberg, Dresden, Bayreuth, Schweinfurt, Munich, Stut-
tgart and Freiburg are localities near the spreading front, although it is also reported
from Vienna (Austria). The species lives in parks, cemeteries and gardens (in England
also in open forests) on ornamental species of Rhododendron, which is necessary for
nymphal development, although some individuals may be swept from other woody
plants such as Acer, Platanus, Tilia and others. In England and France it is often men-
tioned in connection with Pycnostysanus azaleae (Peck.), a phytopathogenic fungus causing
bud blast on Rhododendron. However, it is uncertain whether pathogen transmission is
conducted or facilitated by the leafhopper, since detailed studies are lacking.
Achtziger (pers. comm.), Bußmann (1992), Feldmann & Bußmann (1993), Hoffmann (1990), Körner (pers.
comm.), Niedringhaus & Olthoff (1986, 1993), Olthoff (1986), Remane (pers. comm.), Remane & Wach-
mann (1993), Sergel (1987), SMNS, Walter (pers. comm.), Weis (pers. comm.), Nickel & Remane (1996),
Nickel (1994), HN; see also Arzone et al. (1987), della Giustina (1989), Kaiser-Mauer (1977), Morcos (1953)

4.2.4.12 Typhlocybinae Kirschbaum, 1868

Alebra neglecta W. Wagner, 1940

E VI – B IX; egg, 1 gen.
Oligophagous on Carpinus betulus, Prunus padus and Crataegus, occasionally also on Pru-
nus avium; most individuals are swept in open stands of trees and shrubs and along forest
112 Review of species

margins. Widespread in Germany, but not common and imperfectly documented. The
edge of the range runs through north Germany; northernmost localities are near Bremen,
Hamburg, Plön and Neubrandenburg, extending to at least 700 m a.s.l. in the Allgäu.
Frommer (1996), Günthart (1988), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1990), SMNS,
Wagner (1940b), Nickel & Remane (1996), HN

Alebra coryli Le Quesne, 1977

M VI – B IX; egg, 1 gen.
At present only known from the western half of Europe (British Isles, Netherlands,
France, Switzerland, Poland, Czech Republic and Italy) and Canada (where presum-
ably introduced), but probably more widespread and formerly misidentified. Lives on
Corylus avellana in forests, along their margins, as well as on solitary shrubs, also on C.
maxima and C. colurna in urban settlements. Widespread in Germany, apparently com-
mon and in rather high frequency in most better studied areas, e.g. large parts of Bavar-
ia, middle Hessen and southern Niedersachsen; also recorded along the upper and mid-
dle Rhine, the Wendland of Niedersachsen, the Oberlausitz and the Oderbruch, up to at
least 700 m a.s.l., found at 860 m a.s.l. in Switzerland.
Frommer (pers. comm.), Remane & Fröhlich (1994b), Schönitzer & Oesterling (1998b), Nickel (1994), HN;
see also della Giustina (1989), Dworakowska (1993), Gillham (1991) (see this paper also for identification),
D’Urso (1995), Günthart (1987b), Lauterer & Novotný (1991)

Alebra wahlbergi (Boheman, 1845)

E V – M IX; egg, probably 1 gen. (see also Demichelis & Bosco 1995).
Polyphagous and eurytopic on various deciduous trees, the majority of individuals found
on Acer campestre, Tilia cordata, Ulmus minor and Carpinus betulus, in lower numbers also
on Acer platanoides, A. pseudoplatanus, Ulmus glabra, Sorbus intermedia, S. aria and Betula
pendula; in Britain also on Aesculus hippocastanum, Prunus avium, Alnus glutinosa and
Castanea sativa. Widespread in Germany and usually among the commonest arboricolous
leafhopper species of lower altitudes, found up to at least 800 m a.s.l. in the Mittelgebir-
ge and the foothills of the Bavarian Alps, at 1300 m a.s.l. in the Valais (Switzerland).
Achtziger (1991), Frommer (1996), Niedringhaus (1997), Olthoff (1986), Remane (1987), Schiemenz (1990),
Wagner (1935, 1939a), HN, and others; see also Cerutti (1939a), Claridge & Wilson (1981)

Alebra albostriella (Fallén, 1826)

This taxon and the following have long been treated as conspecific, but are considered to be
distinct species since Gillham (1991). Accordingly, there are differences in colouration and in the
morphology of sternal apodemes of ††, as well as in host preferences. The latter, however ap-
pear to be variable in different parts of Europe: A. albostriella (Fall.) lives monophagously on
Quercus robur in Wales, but on various Fagaceae species in Piemont (see below). The taxonomic
situation (also within remaining Alebra species) is further complicated by the occurrence of var-
ious colour morphs.
E VI – M X; egg, 1 gen., in northern Italy according to Demichelis & Bosco (1995) occa-
sionally 2.
 Typhlocybinae Kirschbaum, 1868 113

Usually on Quercus robur, at least adults occasionally on Qu. petraea, Alnus, Betula, and
additional deciduous species, in closed forests as well as in open stands and on solitary
trees. From Italy Quercus petraea, Qu. cerris and Castanea sativa have been reported as
additional host plants. Apparently widespread in the lowlands of Germany, although
published records partially refer to the following taxon. Highest localities in the Mittel-
gebirge and the foothills of the Alps are at 750 m a.s.l.
Achtziger (1991), Frommer (1996), Niedringhaus (1997), Remane (1987), Schiemenz (1990), Wagner (1935,
1939a), HN, and others; see also Demichelis & Bosco (1995); for identification see Gillham (1991)

Alebra viridis (Rey, 1894)

M VII – M X; egg, 1 gen., 2 according to Demichelis & Bosco (1995) in northern Italy.
Usually on Quercus petraea in forests and along their margins, at least single individuals
also on Qu. robur; in Wales breeding has also been recorded on Qu. cerris, in Italy addi-
tionally on Castanea sativa. Apparently widespread in the lowlands of Germany (ex-
tending northward at least to Cologne, Lüneburg and Berlin), but less common than the
preceding taxon and – particularly in the eastern half – only poorly documented. The
highest localities are at 550 m a.s.l. in the Mittelgebirge, at 1300 m a.s.l. in the Valais
(Switzerland).
Frommer (1996), Nickel & Remane (1996), Nickel (1994), HN; see also Cerutti (1939a), Demichelis & Bosco
(1995), Gillham (1991); for identification see Gillham (1991)

Erythria aureola (Fallén, 1806)

M V – E X; egg, 2 gen., perhaps 1 at higher altitudes, perhaps 3 in warmer regions.
On dwarf shrubs in sunny, sparsely vegetated, moderately dry to dry sites; in the north
German plain mainly on heathland on Calluna vulgaris, in middle and southern parts on
various substrates and usually on Thymus. Widespread in Germany, scattered and lo-
cally absent in the Mittelgebirge, but often among dominant species in favourable plac-
es; highest localities are at 1000 m a.s.l. in the Mittelgebirge, at 1800 m a.s.l. in the Bavar-
ian and Allgäu Alps. Is reported to live in on Globularia cordifolia and Teucrium mon-
tanum in the alpine belt of Switzerland (up to 2280 m a.s.l.) and eastern parts of Austria.
Fischer (1972), Reimer (1992), Remane (1987), Schiemenz (1969, 1990), Trümbach (1959), Wagner (1935,
1939a, 1951a), HN; see also Günthart (1987a), Wagner & Franz (1961)

Erythria manderstjernii (Kirschbaum, 1868)

Adults I – XII, mainly B VIII – B VI; adult, 1 gen.
On dicotyledonous herbs and dwarf shrubs in sunny to shady, usually damp to moder-
ately wet sites of the Mittelgebirge and the Alps; mainly in open coniferous and mixed
forests, subalpine scrub and alpine grassland, also near streamlets. Adults are usually
found on various dicotyledons, e.g. Ranunculus (at least R. montanus, R. repens), Thymus,
Senecio, Eupatorium, Vaccinium, in southwestern parts of the Alps also Chrysanthemum,
Crepis, Leontodon, Trifolium, Prunella, Plantago and Chaerophyllum. Widespread and com-
114 Review of species

mon in the Bavarian and Allgäu Alps, between 800 and 2000 m a.s.l., recorded at 2400 m
a.s.l. in North Tyrol (Austria) and the Valais (Switzerland). Also in higher parts of the
Mittelgebirge (usually above c. 600 m a.s.l., but extending down to c. 450 m a.s.l. in cool
and shady situations), known from the Black Forest (syntype from near Freudenstadt),
the Swabian Jura, Rhön Mountains, Fichtelgebirge, Vogelsberg, Thuringian Forest, Erzge-
birge, Eifel Mountains, Rothaargebirge and Harz Mountains.
Fischer (1972), Kirschbaum (1868), Nikusch (1976), Rabeler (1952), Reimer (1992), Remane (pers. comm.),
Schiemenz (1990), HN; see also Cerutti (1939a), Christandl-Peskoller & Janetschek (1976), Dobler (1985),
Leising (1977), Vidano (1959a)

Liguropia juniperi (Lethierry, 1876)

So far B VIII – IX; adult, 1(?) gen.
This species is known to live on scaly species of Cupressaceae (Cupressus sempervirens,
Juniperus phoenicea, Tetraclinis articulata) in the Mediterranean region. In Germany it was
found only along the upper and middle Rhine and in the Nahe valley (urban areas of
Mainz, Frankfurt, Darmstadt and Cologne, as well as near Schloßböckelheim), below
200 m a.s.l. In Mainz the species was reported from 1993 through 1998 in parks and
gardens on ornamental Chamaecyparis. Specimens in Darmstadt were swept from Ch.
lawsoniana. Recently it was also published from the French side of the southern upper
Rhine plain near Colmar. This is probably one of the rare cases of an invading host
specialist exclusively attacking a plant which was introduced from other countries than
the insect.
Frommer (1996), Thüs (pers. comm.), Fröhlich & Nickel (unpublished data), HN; see also della Giustina &
Balasse (1999)

Emelyanoviana contraria (Ribaut, 1936)

So far M VIII – B IX; egg(?), 1(?) gen.
Only known from the Pyrenees (France, Andorra), the Alps (North Tyrol, Piemonte,
Hautes-Alpes) and mountainous regions of England and Scotland. Most records are
from sun-exposed, rocky, rather dry calcareous hillsides from the upper montane to the
alpine belt (found between 1500 and 2500 m a.s.l. in the Alps). Helianthemum sp. is re-
ported as host plant in Britain. Recently the species has been recorded in Malaise trap
samples from the Allgäu Alps: Ponten above Hindelang, c. 1850 m a.s.l., mid August
until mid September 1998, altogether 6 ††, 4 ‡‡.
Nickel & Voith (unpublished data); see also della Giustina & Meusnier (1982), Kirby (1992), Leising (1977),
Remane (1961b), Remane & Fröhlich (1994b), Ribaut (1952), Woodroffe (1971)

Emelyanoviana mollicula (Boheman, 1845)

M V – E X; egg (occasionally adult), 2 gen., perhaps 1 at higher altitudes.
In sunny, dry to damp, occasionally also moist sites, mainly dry grassland, low-input
meadows and pastures, and ruderal sites, occasionally also gardens. Host plants are
 Typhlocybinae Kirschbaum, 1868 115

various herbs, mainly Salvia pratensis, Teucrium, Origanum, Thymus, and other species of
Lamiaceae, as well as Verbascum spp.; is also reported from Heracleum, Mentha, Satureja,
Cannabis, Parietaria, Artemisia, Fragaria, and additional dicotyledons. Widespread in
Germany and common particularly in warmer regions, but apparently absent from most
parts of the northwest German plain, frequently found up to at least 1350 m a.s.l. in the
Mittelgebirge and the Bavarian Alps. Is also reported from the alpine belt in Switzerland
(up to 2200 m a.s.l.), but breeding is uncertain in these sites.
Achtziger (1991), Fischer (1972), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Rombach
(1999b), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), Nickel
(1997), HN; see also Günthart (1987a), Vidano (1965)

Dikraneura variata Hardy, 1850

Adults I – XII; but hibernation probably mainly in the egg stage, 2 gen.
Among grasses, usually in acidic, damp to moderately dry, shady to moderately shady
sites, locally and in lower abundances also on basic substrates; mainly in open forests
and along woody margins, occasionally in parks and gardens. Often occurs in high
abundance on Deschampsia flexuosa, preferentially under pine, beech, oak or spruce, in
lower numbers on fescue (Festuca ovina, F. rubra, F. heterophylla). Widespread and com-
mon in northern and middle parts of Germany (including the Bavarian Forest, Swabian
and Franconian Jura, and southern Black Forest, where found up to at least 1200 m
a.s.l.), but apparently absent to the south of the Danube.
Achtziger (1991), Reimer (1992), Remane (1987), Schiemenz (1990), Trümbach (1959), Wagner (1935, 1939a),
HN, and others

Micantulina stigmatipennis (Mulsant et Rey, 1855)

M V – B X; egg, 2 gen.
On mullein (Verbascum lychnitis, perhaps also additional species) in sunny, moderately
dry to dry, often disturbed sites on basic as well as acidic substrates. In Germany the
species is rather scattered in lowland areas, at the western edge of the range; western-
most localities are near Frankfurt an der Oder, Potsdam, Dessau, Sondershausen, Jena
and Meißen, three localities in the Franconian Jura near Eichstätt and Regensburg be-
tween 400 and 450 m a.s.l. appear to be rather isolated.
Schiemenz (1990), Schumacher (1923), ZIMH, HN; see also Vidano (1965)

Micantulina micantula (Zetterstedt, 1840)

In Italy B VII – E V; adult, probably 1 gen. (Vidano 1965).
In Germany only known from two localities in Bavaria: Mertingen near Donauwörth, c.
400 m a.s.l., 05.V.1935, and Altmühl valley near Arnsberg, 450 m a.s.l., 27.IX.1996, 4 ††,
4 ‡‡, on a xerothermic hillside on Jurassic limestone with scattered shrubs and trees on
Thalictrum minus. From Italy the species is reported from various species of Thalictrum
in meadows, near shrubs and in forests, up to 1500 m a.s.l. Otherwise known from Scan-
116 Review of species

dinavia, northern Italy, former Yugoslavia, Greece, and large parts of boreal Asia; records
from other countries may be dubious.
Fischer (1972), HN; see also Drosopoulos et al. (1986), Nast (1972, 1987), Vidano (1965)

Wagneriala incisa (Then, 1897)

So far M VII – E VIII; 1(?) gen.; in northern Italy according to Vidano (1965) E V – B XI;
egg, at least 2 gen.
In Germany only known from three records from Westphalia and Bavaria: Weser valley
near Höxter, c. 200 m a.s.l., 1955 or 56, 1 specimen in an open beech forest on a steep,
south-facing slope of Triassic limestone; Mainfranken: Sulzthal, 250 m a.s.l., 11.VII.1994,
1 ‡, and Berchtesgaden Alps: Winkl near Berchtesgaden, 700 m a.s.l., 22.VIII.1996, 2 ‡‡,
both in open pine forests on xerothermic limestone plateaus and slopes. From northern
Italy the species is reported to live in the undergrowth of chestnut and oak coppices
with sedges, notably Carex caryophyllea, C. muricata and C. montana. In the German sites,
the latter species is likely to be the host plant. Otherwise it is only known from Roma-
nia, former Yugoslavia, Austria, Italy, and the Baltic Sea islands of Gotland and Wollin.
Rabeler (1962), Nickel (1999b); see also Holzinger (1999b), Nast (1987), Ossiannilsson (1981), Vidano (1965),
Wagner (1955)

Wagneriala minima (J. Sahlberg, 1871)

Mainly B VII – E VIII, occasionally also M VI and M X, nymphs so far B VII; egg(?), 1(?) gen.
In sun-exposed, xerothermic pine forests on limestone, occasionally also on basic blown
sand; mainly on Carex humilis, perhaps also on C. alba and additional sedge species. In
Germany only known from the basin of the Saale (near Freyburg an der Unstrut, Bad
Frankenhausen, Jena, Eisenberg and Arnstadt), the valley of the Altmühl between Eich-
stätt and Kelheim, the upper Isar between Vorderriß and Wolfratshausen, and the vicin-
ity of Mainz, up to at least 1000 m a.s.l., but found at 1500 m a.s.l. in South Tyrol (Italy).
Fischer (1972), Remane (1987, and pers. comm.), Schiemenz (1990), HN; see also Günthart (1987a)

Wagneriala sinuata (Then, 1897)

E VI – B X; egg(?), 1(?) gen.
On Carex flacca usually in moderately shady, rather dry and basic sites, mainly open
pine forests and thermophilous margins of forests and shrubs on calcareous hillsides. In
Germany at the northern edge of the range, known at present only from 12 localities in
the northern Eifel Mountains, the Thuringian Eichsfeld, the Vogelsberg, Mainfranken
and the Franconian Jura, between 200 and 500 m a.s.l.
Remane & Fröhlich (1994b), Rombach (1999b, and pers. comm.), Wagner (1955), Nickel (1999b), HN

Forcipata citrinella (Zetterstedt, 1828)

E V – E X; egg, 2 gen., probably 1 at higher altitudes.
 Typhlocybinae Kirschbaum, 1868 117

In sedge stands in oligotrophic, usually peaty to temporarily moist (locally temporarily

dry), basic to acidic sites, mainly straw meadows, fens and intermediate bogs; also in raised
bogs, where confined to marginal slopes, ditches and peat diggings, rarely and in low num-
bers also in dry (often temporarily dry) grassland. Host plants are low-growing species of
sedges, perhaps C. flacca, C. nigra and others. Widespread in Germany and locally common,
particularly to the south of the Danube and in the Mittelgebirge, but rather scattered or even
locally absent at lower altitudes where there was probably a dramatic decline due to inten-
sification of agricultural management in recent decades. Found up to at least 1700 m a.s.l. in
the Bavarian and Allgäu Alps, up to 1870 m a.s.l. in the French Alps.
Fischer (1972), Marchand (1953), Reimer (1992), Schiemenz (1971a, 1976, 1990), Schwoerbel (1957), Wagner
(1935, 1939a), Nickel & Achtziger (1999), HN; see also della Giustina (1989)

Forcipata forcipata (Flor, 1861)

This species has often been confused and misinterpreted. Ribaut (1936) and Wagner (1939a)
treated it as Dicraneura citrinella (Zett.).
B VI – E X, according to Schiemenz (1990) also M V; egg, 2 gen., probably 1 at higher altitudes.
In open deciduous and coniferous forests and along their margins (mainly under pine and
spruce, but also beech, oak and alder), in moist to dry sites from the lowlands up to the
upper montane belt, at higher altitudes (above c. 1000 m a.s.l.) more in sunny and also wet
to peaty sites (tall- and small-sedge swamps, alpine grassland). Host plants are various
sedges (Carex flacca, C. humilis, probably also C. sylvatica, C. brizoides and others), woodrush-
es (Luzula spp.) and perhaps grasses. Widespread in Germany and common particularly in
the limestone regions of middle and southern parts, found in high abundances up to 2000 m
a.s.l. in the Bavarian and Allgäu Alps, in the French Alps even at 2500 m a.s.l.
Fischer (1972), Niedringhaus (1997), Reimer (1992), Schiemenz (1990), Schwoerbel (1957), Wagner (1935,
1939a), HN; see also della Giustina (1989)

Notus flavipennis (Zetterstedt, 1828)

M V – E X; egg, 2 gen., probably 1 at higher altitudes.
In moderately wet to temporarily flooded, usually sunny sites, mainly in tall-sedge
swamps, meadows, along ditches and margins of bogs, in wet ruderal sites, etc., usual-
ly on various species of tall sedges (notably Carex acutiformis, C. acuta, C. nigra, C. rostra-
ta, C. vesicaria, C. paniculata, C. disticha), also found in slightly saline sites on Bolboschoe-
nus maritimus. Widespread and common in Germany and one of the dominant species
of many wetland sites, extending up to at least 1600 m a.s.l. in the Alps.
Bittner & Remane (1977), Fischer (1972), Niedringhaus (1991), Reimer (1992), Schaefer (1973), Schiemenz
(1988), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others

Kybos butleri (Edwards, 1908)

B VI – B X; egg, 2 gen.
On Salix triandra, less frequently on S. cinerea and S. aurita, along shores of rivers, streams,
lakes and ponds, also on S. repens on dunes along the coasts of the North and Baltic Sea.
118 Review of species

In Germany known only from scattered records at lower altitudes, only to the north and
west of a line running from Mainz to Würzburg, Arnstadt, Bad Frankenhausen and
Kiel, where locally fairly common to at least c. 400 m a.s.l. However, the species is re-
ported from all adjacent countries and is probably much under-recorded.
Frommer (pers. comm.), Niedringhaus (1991, 1997), Remane (pers. comm.), Wagner (1935, 1939a, 1955),
Nickel & Remane (1996), HN; see also Dworakowska (1976), Nast (1987)

Kybos rufescens Melichar, 1896

B VI – B X; egg, 2 gen.
On Salix purpurea along rivers and streams. Widespread in Germany, in intermediate to
high frequency on the host, although less common in the north German plain. Frequent-
ly found up to at least 1000 m a.s.l. along alpine rivers in Bavaria, up to 1500 m a.s.l. in
the Engadine (Switzerland). The locus typicus is near Munich.
Fischer (1972), Frommer (pers. comm.), Nikusch (1976), Reimer (1992), Remane (1987), Schwoerbel (1957),
Voigt (1978), Wagner (1951a, 1955), HN; see also Günthart (1987a)

Kybos limpidus (W. Wagner, 1955)

B VI – E IX; egg, 2 gen.
In floodplains of rivers and streams and along lake shores, mainly on Salix viminalis,
less frequently on S. triandra, presumably also on S. pentandra. Widespread in the low-
lands of Germany, in intermediate to high frequency and often large numbers on the
main host, recorded up to 600 m a.s.l. in the foreland of the Alps. The locus typicus is
Niedermarschacht near Hamburg.
Dworakowska (1976), Emmrich (1975), Reimer (1992), Wagner (1955), Nickel & Remane (1996), HN

Kybos abstrusus (Linnavuori, 1949)

M VI – M X; egg, 2 gen.
On Populus nigra italica and P. n. nigra, preferentially in river floodplains, also in urban settle-
ments (parks, sports grounds, etc.). Rather scattered in Germany, at the northwestern edge of
the range (although perhaps under-recorded), most localities are known from larger river
valleys: upper Rhine (from Speyer to Mainz, also on the Swiss and French side), Main (from
Frankfurt to Haßfurt), Isar (Munich), Lahn (Marburg), Weser (Hannoversch Münden), Leine
(Göttingen), Elbe (near Lüchow), also near Arnstadt and Balingen, up to at least 550 m a.s.l.
Remane (1987, and pers. comm.), Wagner (1955), Nickel & Remane (1996), Nickel (1997), HN; see also
Lauterer (1996)

Kybos populi (Edwards, 1908)

B VI – E X; egg, 2 gen.
On poplars (Populus tremula, P. nigra, P. alba, also hybrids) in floodplains of rivers and
streams, in forests (usually along margins and roads, as well as in clearings), also in
 Typhlocybinae Kirschbaum, 1868 119

urban settlements. Widespread in Germany, in intermediate to high frequency on the

hosts, found up to at least 800 m a.s.l. in the Mittelgebirge and the foothills of the Alps.
Niedringhaus (1991, 1997), Reimer (1992), Remane (1987), Remane & Reimer (1989), Schiemenz (1990),
Wagner (1935, 1955), Nickel & Remane (1996), HN

Kybos lindbergi (Linnavuori, 1951) and K. betulicola (W. Wagner, 1955)

Empoasca betulicola W. Wagner, 1955
Dworakowska (1976), Nast (1972, 1987) and Ossiannilsson (1981) treat Kybos lindbergi (Lnv.) and
K. betulicola (W.Wg.) as two distinct species. According to the first author, both hybridise in central
Europe (although laboratory rearing has not been carried out). German specimens can only rarely
be clearly assigned to either of the two. Transitional specimens have recently also been recorded
from the Czech Republic (Lauterer 1996). The drawings of Dworakowska (1976) show consider-
able intraspecific variation in genitalic morphology and even appear contradictory in comparison
with those of Ossiannilsson (1981) regarding the shape of the pygofer appendages. Furthermore,
there are no differences in host preferences at least in central Europe. Hence, both forms are likely
to belong to a single species with pronounced variability of male genitalic structures. In this case
Linnavuori’s name has priority. In addition, there is a nomenclatural problem: according to Hamil-
ton (1983a), K. betulicola (W.Wg.) is a junior synonym of Empoasca luda Davidson & De Long, 1938
described from North America. However, De Long’s drawing shows a species without lateral
appendages of the aedeagus, which may belong to the butleri or limpidus group, but is clearly
distinct from European birch-dwelling specimens. Hence, this opinion is refused, and Linnavuo-
ri’s name will be used here until more detailed biosystematic research and further investigations
of type material have been carried out. Nevertheless, the European birch-feeding species occurs
on introduced Betula pendula in Canada and the United States (I. Dworakowska, pers. comm.).
B VI – E X; egg, 2 gen.
On birches (Betula pendula and B. pubescens) in wet to moderately dry sites, on solitary
trees and along alleys, as well as in open forests. In the western half of Germany, includ-
ing Thuringia, specimens with more or less intermediate genitalic morphology are wide-
spread, although not yet recorded to the south of the Danube. Specimens from Berlin and
Saxony increasingly show characters of the lindbergi type. Usually occurs in intermediate
to high frequency and intermediate individual numbers on both species of birch. Highest
localities are at 800 m a.s.l. in the German Mittelgebirge, at 1485m a.s.l. in the Swiss Alps.
Niedringhaus (1991, 1997), Reimer (1992), Remane & Reimer (1989), Schiemenz (1990), Wagner (1955),
Nickel & Remane (1996), HN; see also Günthart (1987a)

Kybos digitatus (Ribaut, 1936)

In France VIII (della Giustina 1989); egg, 2 gen.
Recorded in 2000 in the extreme southwest of Germany: Weil am Rhein, in a sand pit, prob-
ably on Salix purpurea or S. eleagnos. Otherwise only known from a few localities in southern
and southwestern France; most of these records are from undetermined species of Salix.
Mühlethaler (2001, and pers. comm.); see also Dworakowska (1977), della Giustina (1989)

Kybos smaragdula (Fallén, 1806)

E V – E X; egg, 2 gen.
120 Review of species

On Alnus glutinosa and A. incana (also reported from A. alnobetula from Italian parts of
the Alps), mainly along rivers, streams and lake shores, as well as on seepy slopes.
Widespread and common in Germany and in high frequency on the host plants at least
in lowland areas (up to c. 600 m a.s.l.), frequently, but less common in the Mittelgebirge
and the Alps up to 1000 m a.s.l., recorded at 1600 m a.s.l. in the Engadine (Switzerland).
Achtziger (1991), Fischer (1972), Heller (1987a), Niedringhaus (1991, 1997), Reimer (1992), Remane & Reimer
(1989), Schiemenz (1990), Wagner (1935, 1939a, 1951a), HN; see also Günthart (1987a), Vidano & Arzone (1987a)

Kybos calyculus (Cerutti, 1939)

So far E VI – E VIII (Schiemenz 1990); egg, 2(?) gen.
At present this species is only known from Poland, England, Switzerland and eastern
parts of Germany, where it was recorded in two sites in the 1980s: Dubringer Moor
(near Hoyerswerda), E VIII 1987, and Berlin (Biesdorf), E VI – E VIII 1983. It is reported
to live on Betula pubescens and is apparently confined to lower altitudes. The highest
locality is the locus typicus in Switzerland (Chippis, Valais, c. 600 m a.s.l.).
Schiemenz (1990); see also Cerutti (1939a), Dworakowska (1976)

Kybos strigilifer (Ossiannilsson, 1941)

K. perplexus (Ribaut, 1952), showing very similar genitalic structures, has been described from the
French side of the upper Rhine plain and has been reported from various mountainous regions of
eastern and central Europe, as well as from central Asia. Like K. strigilifer (Oss.), it lives on Salix
caprea and can be distinguished only by irregular spines on the aedeagus, which are more numer-
ous and more strongly developed. However, German specimens of K. strigilifer (Oss.) show great
variability of this character, even within the same populations and on the same host individual. It
should also be mentioned that the few published notes on this problem are controversial. Günthart
(1974), who reared the species under laboratory conditions, reported variability in the shape of the
aedeagus and the anal tube appendages. However, if one follows Dworakowska (1976), Günthart’s
drawings in fact indicate K. perplexus (Rib.). Hence, there appears to be no character clearly distin-
guishing these forms, which may in fact be conspecific. Furthermore, K. paraltaicus (Orosz, 1996)
described after a single vagrant † from Hungary is likely to belong to this group.
M VI – E IX; egg, 2 gen.
On Salix caprea, S. cinerea and S. myrsinifolia, mainly in forest glades, clearings, mining
areas, fens, intermediate bogs, and along streams and ditches. Widespread in Germany,
usually in intermediate frequency on the host plants, recorded up to 1200 m a.s.l. in the
Allgäu Alps. Individuals clearly bearing characters of K. perplexus (Rib.) have been found
in southeastern parts, notably Lower Bavaria.
Niedringhaus & Olthoff (1993), Post-Plangg & Hoffmann (1982), Reimer (1992), Schiemenz (1990), Wagner
(1955), Weis & Schönitzer (2001), Nickel & Remane (1996), HN; see also the drawings of della Giustina
(1989), Dworakowska (1976, 1977a), Günthart (1974)

Kybos mucronatus (Ribaut, 1933)

Has been treated as specifically distinct from Kybos mucronatus verbae Zachvatkin, 1953 by Dwora-
kowska (1976) and later, Nast (1987) and della Giustina (1989). However, all figured specimens
show an enormous intraspecific variation of the shape of aedeagus, anal tube appendages, ster-
 Typhlocybinae Kirschbaum, 1868 121

nal and tergal apodemes. The Bavarian specimen (see below) showed sternite apodemes as
described for verbae, but tergite apodemes as shown for mucronatus (cf. della Giustina 1989).
Thus, at least central European populations of both taxa are considered here as conspecific.
Doubts on the distinctiveness have also been expressed by Günthart (1997).
In Poland and Switzerland VII – IX; egg, 1(?) gen. (Dworakowska 1976, Günthart 1987b).
Largely confined to the region of the Alps and the Carpathians. Reported to live on
Alnus glutinosa, but also on Salix. In Germany only known from three trap catches in the
Rhineland and the Bavarian Alps: Krefeld, c. 40 m a.s.l., c. 1970, Burscheid, 200 m a.s.l.,
3 individuals, July 1986, and Garmisch-Partenkirchen, Friedergries, c. 900 m a.s.l., 1 †,
September 2000. Found between 450 and 1475m a.s.l. in the Swiss Alps.
Kolbe & Bruns (1988, and pers. comm.), Coll. Munk. (Remane det.), Nickel & Voith (unpublished data); see
also Dworakowska (1976), Günthart (1987b)

Kybos strobli (W. Wagner, 1949)

In Switzerland VI – IX (Günthart 1987b); egg, 2(?) gen.
This is apparently an endemic species of the Alps and the Carpathians, which is report-
ed to live on Alnus incana. At present there are only two records from Germany, both
comprising single †† only: Ulm (Swabian foreland of the Alps), c. 500 m a.s.l.,
03.VIII.1908, and Hindelang (Allgäu Alps), 1200 m a.s.l., 02.VIII.1995. Recorded from
altitudes between 460 and 1560 m a.s.l. in Switzerland.
Dworakowska (1976) (see this paper also for identification), Nickel (1999b); see also Günthart (1987b)

Kybos virgator (Ribaut, 1933)

Kybos volgensis Vilbaste, 1961, described from the Astrakhan region (southern Russia) on the
basis of differences in body size and adeagus shape, has been reported from Sweden, Switzer-
land and eastern Germany (Ossiannilsson 1981; Günthart 1987a; Sander et al. 1999). However,
due to the high variability and incongruent descriptions of Russian and Swedish specimens the
specific identity is in need of further clarification.
B VI – E X; egg, 2 gen.
On Salix alba and S. fragilis in floodplains of rivers and streams, locally also on S. vimina-
lis, S. aurita, S. cinerea and S. babylonica (also far from water), few specimens found on S.
pentandra. Widespread and common in the lowlands of Germany, usually in high fre-
quency and large numbers particularly on Salix alba and S. fragilis, but recorded only up
to 600 m a.s.l. in the Mittelgebirge and the foreland of the Alps.
Fischer (1972), Lauterer & Malenovsky (1995), Mölleken & Topp (1997), Niedringhaus (1991, 1997), Reimer
(1992), Wagner (1935, 1939a, 1951a), HN, and others

Empoasca affinis Nast, 1937

†† E VII – B XI, ‡‡ probably until V/VI, but these cannot be identified with certainty;
‡‡(?), 1(?) gen.
Along margins of shrubby and herbaceous vegetation in sunny, damp to dry sites, mainly
dry grassland, ruderal sites, abandoned vineyards, forest margins, hedges, etc.; most
122 Review of species

individuals are swept from deciduous shrubs and trees (Rubus, Ulmus, Carpinus, Cory-
lus, Populus, Salix and others), and in the herb layer, often in low numbers only. In Ger-
many at the edge of the range, mainly found in the warmer regions of southern and
eastern parts, extending northward at least to Koblenz, Marburg, Gotha, Sondershau-
sen, Berlin and Frankfurt an der Oder; single individuals have been found near Cologne,
Helmstedt, Lauenburg and Kaufbeuren, c. 650 m a.s.l., but usually below 400 m a.s.l.
Fischer (1972), Frommer (1996), Reimer (1992), Remane (1987), Remane (pers. comm.), Schiemenz (1990),
Schönitzer & Oesterling (1998b), Wagner (1951a, 1952), HN

Empoasca apicalis (Flor, 1861)

E VIII – M V; adult, 1 gen. (see Günthart 1987a; Lauterer 1980).
This species‘ range essentially includes Siberia and Scandinavia, with isolated popula-
tions in central Europe. From Germany there are only three records from southern parts:
Illasberg near Füssen, c. 800 m a.s.l., 13.V.1951; upper Neckar valley near Rottweil,
23.IX.1956, and Breitbrunn am Ammersee, 23.VIII.1959, in both sites on Sambucus ebu-
lus. In the Engadine (Switzerland), nymphs and adults were found on Lonicera xylos-
teum at c. 1100 m a.s.l. in spruce forests with luxuriant undergrowth. In Moravia (Czech
Republic), single individuals were recorded on Lonicera nigra between 290 and 620 m
a.s.l. in shady forests. Finnish populations have been reported to migrate from L. xylos-
teum to Alnus incana, Juniperus and Picea in autumn.
Fischer (1972), Remane & Fröhlich (1994b); see also Günthart (1987a), Lauterer (1980), Nuorteva (1948, 1952a)

Empoasca decipiens Paoli, 1930

Adults I – XII, mainly M VII – E V; adult (including ††!), 2(?) gen.
In various, damp to dry, sunny to moderately shady sites, usually along margins of
herbaceous and shrubby vegetation, in ruderal sites, gardens and fields. Mainly on weeds
in the latter, but locally causing feeding damage on cucumbers and sweet pepper in
south German greenhouses. Also reported to be noxious in various cultures of vegeta-
bles and fruits in more southern European countries, although species identification
may not be accurate in some cases. Host plants are mainly dicotyledonous herbs be-
longing to many families (Rosaceae, Fabaceae, Solanaceae, Caprifoliaceae, Asteraceae
and others) and – particularly in autumn – also on shrubs (Rosa, Rubus, Cornus, Corylus,
Sambucus and others), during hibernation presumably again on herbs. Widespread and
common in the lowlands of middle and southern parts of Germany, more scattered and
increasingly preferring synanthropic habitats towards the north. Usually confined to
altitudes below 500 m a.s.l., but locally found up to 800 m a.s.l. along the edge of the
Bavarian Alps. Reported from 1300 m a.s.l. in the Valais (Switzerland). From Scandina-
via there are only two known records from southern Norway and southern Sweden;
likewise, there is only a single record from Denmark.
Achtziger (1991), Frommer (1996), Niedringhaus & Olthoff (1993), Post-Plangg & Hoffmann (1982), Rau-
pach (pers. comm.), Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1951a), HN, and
others; see also Cerutti (1939a), della Giustina (1989), Gillerfors (2002), Günthart (1971a, 1987a), Koblet-
Günthardt (1975), Ossiannilsson (1981), Wagner & Franz (1961)
 Typhlocybinae Kirschbaum, 1868 123

Empoasca pteridis (Dahlbom, 1850)

Adults I – XII, mainly M VIII – E X, according to laboratory and field studies of Günthart
(pers. comm.) †† and ‡‡ hibernate (the former only in very low numbers), according to
Remane (1987) hibernation takes place mainly in the egg stage, 2(?) gen.
In moist to damp, occasionally also wet or dry, sunny to moderately shady, usually
disturbed sites, mainly fertilized meadows and pastures, herbaceous margins, fields,
ruderal sites, waysides, etc., but found in low numbers and perhaps only as vagrants in
most other types of non-forested terrestrial habitats (e.g. dry grassland, wet meadows,
also on various deciduous shrubs and trees). Host plants are various herbs including
cultivated potato. Widespread and common at lower altitudes of Germany, locally found
up to c. 1000 m a.s.l. in the Mittelgebirge and along the edge of the Alps.
Achtziger (1991), Reimer (1992), Remane (1987), Schiemenz (1990), Wagner (1935, 1939a), Wais (1990), Nickel
& Achtziger (1999), HN, and others; see also Günthart (1971b)

Empoasca ossiannilssoni Nuorteva, 1948

This taxon is likely to be conspecific with E. dealbata Cerutti, 1939, which was described after a
single † from the Valais (Switzerland). Only one further † from Styria (Austria, on Prunus padus)
has been recorded since (Wagner & Franz 1961). A number of very similar taxa has been de-
scribed also from Siberia (see Anufriev & Emelyanov 1988). However, the shape of the pygofer
appendages, which has been used as the most important distinguishing character in this genus,
has been found to be much variable (e.g. Günthart 1974; Nickel, unpublished data). Thus, a
revision of Palearctic species would be much welcome.
In Sweden and Finland E VII – B IV; adult, 1 gen. (see Ossiannilsson 1981).
Until recently, this taxon has been known only from the coniferous forest zone of Eur-
asia and the Ukrainean Carpathians. A more recent record is from the Bavarian foothills
of the Alps: Hackensee near Dietramszell, 660 m a.s.l., 09.IX.1994, 1 †, along a forest
margin near a lake shore with dominating Picea abies, Prunus padus and Corylus avellana.
Data on the biology are scarce. In Sweden and Latvia, the species has been found on
Prunus padus and in a birch forest; hibernation is suspected to take place on Picea.
HN; see also Anufriev & Emelyanov (1988), Nast (1972), Ossiannilsson (1981)

Empoasca vitis (Göthe, 1875)

The use of the name E. flavescens (F.) for this species until the second half of the 20th century is
caused by misinterpretation of Fabricius‘ description, which was finally clarified by Wagner
(1961) after a study of the type material.
Adults I – XII, mainly B VII – M V; adult (also ††), 1 gen.
Among all central European arboricolous leafhoppers, this species shows the widest
range of host plants, living extremely polyphagously on various woody plants: Fagus,
Quercus, Betula, Alnus, Carpinus, Corylus, Ulmus, Rosa, Rubus, Malus, Sorbus, Prunus, Acer,
Frangula, Rhamnus, Cornus, Vitis (in southern Europe and southwestern Germany caus-
ing feeding damage in some years), Populus, Salix, Tilia, Viburnum, Lonicera, Fraxinus,
even on introduced species (Platanus, Aesculus, etc.), frequently also in the herb layer (at
least adults on Solanum tuberosum, Althaea, Helianthus and Petasites). In late summer
124 Review of species

most individuals migrate to evergreen plants for hibernation (mainly Picea, also Pinus
and Hedera, often extremely abundant on Taxus in gardens). In many parts of central
Europe this species is, along with Fagocyba cruenta (H.-S.), the most abundant typhlocy-
bine leafhopper, frequently occurring up to the subalpine tree line, found up to at least
1500 m a.s.l. in Bavaria, up to 2000 m a.s.l. in Switzerland.
Achtziger (1991), Fischer (1972), Herrmann et al. (1999), Louis & Schirra (1997), Reimer (1992), Remane &
Wachmann (1993), Schiemenz (1990), Schruft & Wegner-Kiß (1999), Wagner (1935), Wais (1990), HN and
others; see also Cerutti (1939a), Günthart (1987a)

Austroasca vittata (Lethierry, 1884)

E V – B X; egg, 2 gen.
On Artemisia absinthium in moderately dry, usually disturbed sites (ruderal sites, way-
sides, abandoned vineyards, etc.), occasionally also on A. maritima in inland saltmarshes.
In Germany, at the northwestern edge of the range, this species has been recorded since
the 1920s, occurring only locally and rather scattered in warm-summer regions, with strong-
holds in Saxony, the rain shadow area to the east of the Harz Mountains, and the Rhine
valley between Mannheim and Cologne, single records are from the Wendland (eastern
Niedersachsen), Mecklenburg, the Uckermark, northern Hessen and Mainfranken, usu-
ally below 300 m a.s.l., but reported from 1500 m a.s.l. in the Engadine (Switzerland).
Fröhlich (1996a), Frommer (1996), Schiemenz (1990), Walter (pers. comm.), Nickel (1997), HN; see also
Günthart (1987a)

Kyboasca bipunctata (Oshanin, 1871)

B VI – E IX; egg, 2 gen.
On Ulmus minor and U. x hollandica, perhaps also U. laevis, mainly on low-growing indi-
viduals (coppices, saplings), usually in sunny or moderately shady sites. Has been found
on Cannabis and Glycyrrhiza in Poland and the Czech Republic. In Germany only re-
corded in very scattered localities, almost exclusively in urban habitats in river valleys
(Monheim near Düsseldorf, Cologne, Berlin, Halle, Bitterfeld, Göttingen, Arnstadt, Jena,
Görlitz and Badberg/Kaiserstuhl, also near Stettin, Poland), up to 400 m a.s.l.; is also
known from a few localities to the west and north (southern France, Netherlands, En-
gland and southern Finland).
Frommer (pers. comm.), Kolbe & Bruns (pers. comm.), Schiemenz (1990), MTD, Nickel (1994), HN; see also
della Giustina (1989), Dworakowska (1973), Haupt & Hedicke (1934), Lauterer (1984), Nast (1987)

Chlorita dumosa (Ribaut, 1933)

This species and the following belong to a subgroup, which was named Xerochlorita Zachvatkin,
1953. Most authors treat it as a subgenus of Chlorita Fieb. However, Zachvatkin did not provide
a description, and consequently, the name is not valid and has to be treated as a nomen nudum.
M V – M X; egg, at least 2 gen.
In cushions of thyme (Thymus praecox, Th. serpyllum, probably Th. pulegioides) in sunny,
dry to moderately dry, low-vegetated, usually grazed sites, preferentially on sand and
 Typhlocybinae Kirschbaum, 1868 125

limestone, but also on other well-drained substrates. In Germany at the northwestern

edge of the range, with strongholds in warmer regions of southern and eastern parts.
Most records are from the rain shadow area to the east of the Harz Mountains, Thurin-
gia and Mainfranken; rather scattered localities are near Cologne, Mainz, Heidelberg,
Regensburg, Göttingen, Frankfurt an der Oder and Bautzen, up to 450 m a.s.l.
Frommer (pers. comm.), Reimer (1992), Remane (1987), Schiemenz (1990), Wagner (1941a), Nickel (1994), HN

Chlorita pusilla (Matsumura, 1906)

See taxonomic remark above!
B VI – E IX; egg, at least 2 gen.
This species is altogether known only from 7 localities in dune regions of western Eu-
rope: Brodnica (previously Strasburg, to the northeast of Warsaw/Poland), “zahlreiche
Exemplare (...) auf niedrigen krautartigen Pflanzen”; Vieille-St.-Girons, 26.VI.1996, Vieux-
Boucau-les Bains, 06.IX.1997 (both south-western France); Gartz near Angermünde,
12.VIII.1937, on Thymus cushions; Weinberg near Perleberg, 1965 and 1966, altogether
9 ††, 2 ‡‡ on a “Schafschwingel-Schillergras-Sandtrockenrasen”; Boizenburg, 15.VI.
1998, 9 ††, 50 ‡‡, as well as Gerwisch near Magdeburg, 01.VI.2000, 2 ††, 3 ‡‡, the two
latter records on inland dunes on Thymus serpyllum.
Schiemenz (1969), Wagner (1941a), HN; see also della Giustina & Remane (2001), Matsumura (1906)

Chlorita paolii (Ossiannilsson, 1939)

B V – M X; egg, at least 2 gen.
In sunny and oligotrophic, damp to dry sites on various substrates, mainly dry grassland,
low-productivity meadows and pastures, ruderal sites and roadside embankments. Host
plants are Achillea millefolium and Artemisia campestris, locally A. absinthium and A. vulgar-
is. Widespread and common in many places of eastern and southern Germany, but be-
coming rather scarce towards the northwest, where living at the edge of the range, and
towards the edge of the Alps. Border localities are known from near Koblenz, Marburg,
Göttingen, Braunschweig, Uelzen and Hamburg; frequently recorded at 1100 m a.s.l. in
higher parts of the Mittelgebirge. Not reported from Denmark and southern Sweden, al-
though widespread in Mecklenburg. Also common on the German side of the upper Rhine
plain, although in France known only from the western Alps (up to 1700 m a.s.l.), Burgun-
dy and Auvergne. Furthermore, there is a single record from the Netherlands.
Büttner (1964), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schiemenz (1990), Schwo-
erbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), Nickel (1994), HN; also Cobben & Gravest-
ein (1958), della Giustina (1989, and pers. comm.)

Fagocyba carri (Edwards, 1914)

So far E V – B IX; egg, 2 gen.
On oaks (Quercus robur and Qu. petraea) in forests and along their margins. Probably
widespread in the lowlands of Germany, but only poorly documented and apparently
126 Review of species

in rather low frequency and individual numbers; neither recorded in northeastern parts
(Mecklenburg-Vorpommern, Saxony-Anhalt) nor in the foreland of the Alps. The high-
est localities are at 500 m a.s.l. in the Mittelgebirge.
Achtziger (1991), Günthart (1988), Niedringhaus (1991, 1997), Reimer (1992), Remane (1987), Schiemenz
(1990), Wagner (1935), HN

Fagocyba cruenta (Herrich-Schäffer, 1838)

Typhlocyba douglasi Edwards, 1878; Typhlocyba inquinata Ribaut, 1936
This taxon is considered here as a single species comprising three morphs differing only in
characters of colouration, but not in habitat preferences, host plants, phenology and distribu-
tion, although the more strongly pigmented reddish (= cruenta) morph apparently occurs only
in the summer generation. Some authors, however, treat the yellowish white F. douglasi (Edw.),
the reddish F. cruenta (H.-S.), and F. inquinata (Ribaut, 1936), which shows a fumose clavus, as
distinct species. Striking intraspecific differences in colouration are also found within other spe-
cies of Typhlocybini, e.g. Eurhadina and Eupteryx.
M V – E X; egg, 2 gen., perhaps 1 at higher altitudes.
The more widespread yellowish white douglasi morph is eurytopic and polyphagous on
various deciduous woody plants, in closed forests as well as on solitary trees. Preferen-
tially on Fagus and Carpinus (where locally extremely abundant), but also – roughly in
order of decreasing densities – on Acer, Tilia, Ulmus, Prunus, Sorbus, Alnus, Betula, Cory-
lus, Populus, Salix, Aesculus, Quercus, Rosa, Crataegus, Fraxinus and others, even Juglans.
Very common and widespread all over Germany, locally with a low proportion of fe-
male individuals of var. inquinata (Rib.). Frequently found up to 1800 m a.s.l. in the
Bavarian and Allgäu Alps, and perhaps reproducing on subalpine shrubs. The more
scattered and uncommon reddish cruenta morph shows the same host preferences and
geographic distribution, although it has not been recorded in the north German plain
(but in Denmark and southern Sweden, as well as near the alpine tree line). Most indi-
viduals are found among large numbers of the douglasi morph, but the earliest central
European specimens date from the first decade of July.
Achtziger (1991), Bittner & Remane (1977), Frommer (1996), Reimer (1992), Remane & Wachmann (1993),
Schiemenz (1990), Schönitzer & Oesterling (1998b), SMNS, Wagner (1935, 1939a), HN, and others; see also
Claridge & Wilson (1976, 1981), Nast (1987), Ossiannilsson (1981)

Ossiannilssonola callosa (Then, 1886)

B VI – M IX; egg, 1(?) gen.
On Acer pseudoplatanus in forests, parks and on solitary trees, usually in cool sites (val-
ley bottoms, shady slopes, near water). Widespread in Germany and usually in inter-
mediate frequency on the host, but only poorly documented, frequently occurring up to
at least 1300 m a.s.l. in the Bavarian and Allgäu Alps, although single specimens were
found at 1500 m a.s.l.
Reimer (1992), Remane & Reimer (1989), Schiemenz (1990), Wagner (1935), Nickel (1999b), HN; see also
Whittaker (1984)
 Typhlocybinae Kirschbaum, 1868 127

Edwardsiana alnicola (Edwards, 1924)

So far E VI – B VIII; egg, 2(?) gen.
On alders (Alnus glutinosa, A. incana) in moist to wet sites, often near water. The distri-
bution in Germany is rather poorly documented; the species is known only from scat-
tered records in the western half, between the North Sea coast and the Alps, where it
occurs at least in low frequency and individual numbers. Highest localities are at 1000 m
a.s.l. in Bavaria, but at 1660 m a.s.l. in the Engadine (Switzerland). The species is not
reported from France, Belgium and the Netherlands; thus, the western edge of its range
may run through Germany.
Achtziger (1991), Frommer (pers. comm.), Kolbe & Bruns (1988), Niedringhaus & Olthoff (1993), Wagner
(1935), Nickel (1997), HN; see also Günthart (1987a)

Edwardsiana avellanae (Edwards, 1888)

Typhlocyba staminata Ribaut, 1931
Many authors consider E. staminata (Rib.) to be a distinct species (e.g. Nast 1972, 1987; Ossian-
nilsson 1981). Compared to E. avellanae (Edw.), two tiny appendages of much variable length on
the aedeagus tip are the only distinctive characters. Moreover, most, if not all of these individu-
als are pipunculized or otherwise abnormal. Body size, colouration, host plant, habitat and dis-
tribution (also within Germany) are almost identical to E. avellanae (Edw.). Therefore, Lauterer
(pers. comm.), della Giustina (1989) and Le Quesne & Payne (1981) consider them conspecific.
This interpretation is adopted here.
B VI – M X; egg, 2 gen.
Monophagous on Corylus avellana; most individuals are swept along forest margins and
roads, generally in more shady and cool sites than E. spinigera (Edw.). Widespread in
Germany, fairly common and in intermediate frequency, found up to at least 850 m a.s.l.
in the Mittelgebirge and the Bavarian Alps, up to 1400 m a.s.l. in the Valais (Switzerland).
Achtziger (1991), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1990), Trümbach (1959), Wag-
ner (1935, 1939a), HN; see also Cerutti (1939a)

Edwardsiana bergmani (Tullgren, 1916)

B VI – B IX; egg, 2 gen., probably 1 at higher altitudes.
On birches (Betula pendula, B. pubescens) and alders (Alnus glutinosa, A. alnobetula), usu-
ally in moist, cool or shady habitats (e.g. bogs, forests, subalpine alder shrub, near
streams). Widespread in Germany, but rather scattered and in low frequency and indi-
vidual numbers, found up to at least 1500 m a.s.l. in the Bavarian Alps.
Achtziger (1991), Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1990), HN

Edwardsiana candidula (Kirschbaum, 1868)

E. candidula martigniaca (Cerutti, 1939), described from Switzerland is distinguished only by
slight differences in shape and length of the aedeagus appendages, but apparently not in life
history. Recently, it has been treated as a distinct species by various authors (e.g. Nast 1987;
Lauterer, pers. comm.). Both forms occur in central Europe, although only the nominate one has
128 Review of species

been recorded from Germany. The drawings of Ribaut (1936) show the latter, those of Ossian-
nilsson (1981) show E. candidula martigniaca (Cer.), which is preliminarily treated here only as a
variety until further evidence has been obtained.
E V – M X; egg, 2 gen.
Monophagous on Populus alba, usually on solitary trees and along forest margins in
river floodplains and parks. Widespread in the lowlands of Germany and in high fre-
quency on the host in most regions, but apparently absent from northwestern parts.
Localities at the edge of the range include Cologne, Göttingen, Lauenburg and Lübeck;
found only up to 500 m a.s.l. in the foreland of the Alps.
Frommer (pers. comm.), Remane (1987, and pers. comm.), Schiemenz (1990), Wagner (1935, 1939a), Nickel
& Remane (1996), Nickel (1997), HN

Edwardsiana crataegi (Douglas, 1876)

A morph with different shape and length of the aedeagus appendages has been described as
Typhlocyba froggatti Baker, 1925. However, according to laboratory rearings of Günthart (1971b)
the offspring of a single ‡ showed both the aedeagus type of the nominate form and of froggatti
Bak.; the latter is generally considered to be a variety. Regarding individual numbers, it domi-
nates in most parts of the study area.
E V – M X; egg, 2 gen.
On woody species of Rosaceae, mainly Malus, Crateagus, Prunus domestica, P. avium, P.
spinosa; is also reported from Mespilus and Sorbus from Switzerland and Wales. Has
been reported to cause damage on apple trees in southern Europe and elsewhere. Wide-
spread and common in Germany, found up to at least 700 m a.s.l. in the Mittelgebirge, up
to 1400 m a.s.l. in the Swiss Alps.
Achtziger (1991), Frommer (1996), Lehmann (1973a), Müller (1956), Niedringhaus (1991), Reimer (1992),
Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), HN, and others; see also Clar-
idge & Wilson (1981), Günthart (1971b, 1987a)

Edwardsiana diversa (Edwards, 1914)

According to China (1943), the name Anomia tridentata Edwards, 1928, which had been created
for a parasitized specimen, is a junior synonym of E. diversa (Edw.). Wagner (1935) published
records of parasitized individuals from Bremen, the aedeagus of which resembled the one of A.
tridentata Edw., but he argued that its specific status may be uncertain. China’s opinion is appar-
ently based only on the fact that A. tridentata Edw. (only 1 †!) had been taken from Cornus. Thus,
its status is doubtful, since parasitized Edwardsiana males are often impossible to identify. Hence,
the occurrence of E. diversa (Edw.) in northern Germany is not proven.
B VI – E X; egg, 2 gen.
On dogwood (Cornus sanguinea, C. mas) in damp to dry, usually eutrophic or basic sites,
mainly dry grassland, along hedges, forest margins and roads, as well as in parks. In
Germany at the northern edge of the range, only known from widely-separated locali-
ties, but apparently widespread in middle parts and at least locally in intermediate to
high frequency on the hosts (up to at least 450 m a.s.l.). Not found in the north German
plain and to the south of the Danube; the northernmost localities are near Cologne,
Göttingen and Magdeburg.
 Typhlocybinae Kirschbaum, 1868 129

Frommer (pers. comm.), Lehmann (1973a, 1973b), Müller (1978), Nicolaus (1957), Reimer (1992), SMNS,
Nickel & Remane (1996), Nickel (1994), HN

Edwardsiana flavescens (Fabricius, 1794)

For a long time, most authors distinguished E. fratercula (Edwards, 1908) and E. sororcula (Os-
siannilsson, 1936) after differences in shape and length of the lateral aedeagus appendages.
However, since Wagner & Franz (1961) both names are treated as junior synonyms of E. flave-
scens (F.).
E V – E X; egg, 2 gen.
Preferentially on Carpinus betulus and Fagus sylvatica in forests, parks and along hedges,
occasionally also on other species of deciduous woody plants (e.g. Salix, Quercus, Acer,
Ulmus and Betula). Widespread in Germany and usually common on beech and horn-
beam, frequently found up to at least 900 m a.s.l. in the Mittelgebirge and the Bavarian
Alps, single specimens at 1200 m a.s.l.
Bittner & Remane (1977), Förster (1961), Niedringhaus & Olthoff (1993), Nikusch (1976), Post-Plangg &
Hoffmann (1982), Reimer (1992), Schiemenz (1990), Wagner (1935, 1939a), Nickel & Remane (1996), HN

Edwardsiana rhodophila (Cerutti, 1937)

E VI – B X; egg, 2 gen.
Previously, this species was only known from the Mediterranean region, notably south-
ern France, Switzerland (Valais), Greece, northern Italy and eastern Spain, reported from
„ ... speziellen Rosa-Arten in mehr oder weniger montanen Bereichen“. Recently also
recorded in the Saale-Unstrut region and Mainfranken, where it lives on Rosa rubiginosa
in xerothermic sites, mainly in upper slope and plateau situations on limestone and
gypsum. At present there are altogether 9 German localities from the surroundings of
Nordhausen, Artern, Freyburg an der Unstrut, Ilmenau, Gotha, as well as Karlstadt am
Main, some of them holding high individual numbers. Found up to 1530 m a.s.l. in
Spain, but only up to 410 m a.s.l. in Germany.
Nickel (1998); see also Cerutti (1937), Remane & Fröhlich (1994b)

Edwardsiana sociabilis (Ossiannilsson, 1936)

M VI – B X; egg, 2 gen.
This species is reported from northern and central parts of Europe, but also from Geor-
gia. In Germany it occurs mainly in urban areas and along hedges on ornamental roses,
mainly Rosa rugosa. Probably it is more widespread, but is only known from scattered
localities (East Frisian Islands, Bremen, Hamburg, Göttingen, Marburg and surround-
ings, Hof, Görlitz, Steingaden/Upper Bavaria, up to at most 800 m a.s.l.). Furthermore,
populations have been recorded in near-natural habitats on Filipendula ulmaria in tall
herb stands of rather wet or cool sites (northeastern Harz Mountains, Elbe valley near
Dresden, Vogelsberg, Thuringian Forest, Bad Königshofen im Grabfeld). The species is
reported to be common on cultivated Rosa sp. and Rubus idaeus in gardens in southern
130 Review of species

Finland, and from Filipendula ulmaria in Sweden. This is one of the rare documented
cases of a stenophagous native insect species infesting an introduced plant.
Achtziger (1991), Emmrich (1975), Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Wagner
(1937g), HN; see also Linnavuori (1952), Ossiannilsson (1981)

Edwardsiana frustrator (Edwards, 1908)

E V – E X; egg, 2 gen.
On various deciduous woody plants, without any apparent host preference: Corylus, Acer,
Quercus, Aesculus (based on laboratory rearings or nymphal records in Wales and Swit-
zerland), also occasionally found on Carpinus, Tilia, Ulmus, Fagus, Betula, Malus, Prunus,
Rosa, Crateagus, and probably additional species. Widespread in the lowlands of Germa-
ny, but usually in rather low numbers, found only up to 600 m a.s.l. in the Mittelgebirge.
Not recorded from Mecklenburg-Vorpommern, where probably overlooked.
Achtziger (1991), Frommer (1996), Niedringhaus & Olthoff (1993), Post-Plangg & Hoffmann (1982), Reimer
(1992), Schiemenz (1990), Wagner (1935, 1939a), HN; see also Claridge & Wilson (1976, 1981), Günthart (1974)

Edwardsiana ampliata (W. Wagner, 1948)

Edwardsiana solearis rossica Zachvatkin, 1948
Resembling E. frustrator (Edw.) in genital morphology and life habits. Both are considered only
as distinct subspecies by Zachvatkin (1948) and Anufriev (pers. comm.).
B VI – B X; egg, 2 gen.
Described from Silesia (Poland) and since then mainly reported from the deciduous
forest region of eastern Europe. Recently it was recorded in Mainfranken, where it is
currently known from altogether 6 localities near Schweinfurt and Bad Königshofen,
along forest margins on various deciduous trees (Acer campestre, A. pseudoplatanus, Corylus
avellana, Quercus robur), between 200 and 300 m a.s.l.; all records comprised only 1 or
2 †† respectively. Apparently polyphagous, reported to live on Quercus and Tilia in
Moravia (Czech Republic), mainly on Tilia cordata in Poland, on Quercus, Corylus and
Cerasus in middle parts of Russia.
Nickel & Remane (1996), HN; see also Anufriev & Kirillova (1998), Chudzicka (1986a), Lauterer (pers.
comm.), Zachvatkin (1948)

Edwardsiana geometrica (Schrank, 1801)

E VI – M X; egg, 2 gen., perhaps 1 at higher altitudes.
On alders in various damp to wet sites, usually on Alnus glutinosa in lowland regions,
preferentially on A. incana in the Mittelgebirge and the Alps. Widespread in Germany
and, compared to other Edwardsiana species, well documented due to the striking co-
louration. Frequently found up to at least 1000 m a.s.l. in the Mittelgebirge and the Ba-
varian and Allgäu Alps, single specimens up to 1800 m a.s.l. Also found in the subal-
pine belt in the Engadine and Valais (both Switzerland), where presumably living on
Alnus alnobetula.
 Typhlocybinae Kirschbaum, 1868 131

Achtziger (1991), Fischer (1972), Frommer (pers. comm.), Niedringhaus (1991), Nikusch (1976), Reimer
(1992), Remane & Reimer (1989), Schiemenz (1990), Wagner (1935, 1939a), HN; see also Cerutti (1939a),
Günthart (1987a), Vidano & Arzone (1987a)

Edwardsiana gratiosa (Boheman, 1852)

B VII – M X; egg, 2 gen.
Monophagous on Alnus glutinosa, usually near standing or running water. Known from
sporadic localities in the lowlands of Germany, but probably under-recorded (e.g. not
yet found to the south of the Danube, nor in Rhineland-Palatinate and Mecklenburg-
Vorpommern, but from adjacent regions of Poland and France). Usually in intermediate
frequency, but low individual numbers on the host; highest localities are in the Mittelge-
birge at 600 m a.s.l. only.
Frommer (1996), Heller (1987a), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer (1992), Schiemenz
(1990), Wagner (1935, 1939a), Nickel & Remane (1996), HN; see also Nast (1976a), Ribaut (1936)

Edwardsiana lamellaris (Ribaut, 1931)

This species is only known from a few countries in central and southern Europe and is
apparently rare in most parts of its range. Specimens published from the surroundings
of Halle turned out to belong to E. prunicola (Edw.) after a revision. The only definitive
record from Germany is from a moderately dry grassland site with scattered oaks near
Stuttgart: Eichenhain near Riedenberg, c. 400 m a.s.l., 08.VI.1994, 2 ††, 1 ‡, on Rosa can-
ina. This plant is also likely to be the host in Moravia (Czech Republic). From Italy,
however, the species is reported to live monophagously on Quercus petraea and Qu.
robur. Perhaps there is a host shift between the first and the second generation.
Lehmann (1973a), SMNS (Heller leg. et det.); see also Lauterer (1984), Vidano & Arzone (1987b)

Edwardsiana lethierryi (Edwards, 1881)

B VI – B X; egg, 2 gen.
On Acer campestre, Tilia cordata and T. platyphyllos (in England and Wales also on Aesculus
hippocastanum), in lower numbers and without evidence for breeding also on other decidu-
ous trees (Ulmus, Acer platanoides, Corylus and others). Widespread at least in the lowlands
of Germany (including the East Frisian Islands, Heligoland, Holstein and Mecklenburg),
but poorly documented; highest known localities are at only 550 m a.s.l. in the Mittelgebirge.
Recorded in northern Europe only from the island of Gotland and southern Norway.
Fischer (1972), Frommer (1996), Niedringhaus (1991, 1994), Nikusch (1976), Reimer (1992), Schiemenz (1990),
Schwoerbel (1957), Wagner (1939a), HN; see also Claridge & Wilson (1981), Günthart (1974), Ossiannilsson (1981)

Edwardsiana plurispinosa (W. Wagner, 1935)

The recent revision of the E. lethierryi group presented by Wilson & Claridge (1999) could only
partially be taken into consideration for this book. Populations hitherto treated as E. hippoc-
astani (Edw.) sensu Ribaut were split into two species, but the name was shown to be a junior
132 Review of species

synonym of E. lethierryi (Edw.). Thus, almost all published records of E. hippocastani (Edw.) sensu
Ribaut are in need of revision (e.g. Achtziger 1991; Schiemenz 1990) and probably refer to E.
plurispinosa (W.Wg.) or E. ulmiphagus Wils. & Clar.
B VI – E X; egg, 2 gen.
On hazel (Corylus avellana) and alders (Alnus glutinosa, A. incana, cultivated A. alnobetu-
la) in forests and in semi-open habitats; single individuals (probably without reproduc-
tion) are found on most native deciduous tree species. Widespread in the lowlands of
Germany, but poorly documented; usually in intermediate to high frequency on the
hosts, up to at least 700 m a.s.l., although E. hippocastani (Edw.) is mentioned from Alnus
at 1600 m a.s.l. in the Swiss Alps.
Niedringhaus & Olthoff (1993), Wagner (1935), HN; see also Cerutti (1939a), Claridge & Wilson (1976,
1981); for identification see Wilson & Claridge (1999)

Edwardsiana ulmiphagus Wilson & Claridge, 1999

E V – B X; egg, 2 gen.
On elms (Ulmus minor, U. glabra, U. laevis) in forests, parks and on solitary trees; single
vagrants also on other deciduous trees. Widespread in the lowlands of Germany, but poorly
documented, usually in intermediate to high frequency and individual numbers on all
three elm species. Highest localities are at 700 m a.s.l. in the Mittelgebirge, but E. hippoc-
astani (Edw.) has been reported from Ulmus at 1230 m a.s.l. in the Engadine (Switzerland).
HN; see also Claridge & Wilson (1976, 1981), Günthart (1987a); for identification see Wilson & Claridge (1999)

Edwardsiana ishidai (Matsumura, 1932)

Opinions on the status of this taxon and the following are contradictory: according to Dwora-
kowska (1982), Nast (1972, 1987) and Ossiannilsson (1981) Typhlocyba lanternae W.Wg. is a junior
synonym of T. ishidai Mats.; but Lauterer (1984) and Le Quesne & Payne (1981) argue that there
are differences in the shape of the aedeagus and in host choice. Evidence from German popula-
tions supports the latter view, although the total number of records of E. lanternae (W.Wg.) is
extremely low. Until the problem has been resolved, both shall be treated as separate taxa.
E V – M X; egg, 2 gen.
E. ishidai (Mats.) is apparently widespread in middle and southern parts of Germany,
extending northward at least to Cologne, Göttingen, Nordhausen and Eberswalde. Is
usually swept along forest margins and roads, in parks, and from solitary trees, record-
ed up to 350 m a.s.l. only. Host plants are Ulmus minor, U. glabra, U. laevis, and perhaps
Corylus avellana. Not recorded from France, but from the Baden and Palatinate part of
the upper Rhine plain.
Frommer (pers. comm.), Nicolaus (1957), SMNS, Nickel & Sander (1996), Nickel (1994, 1999b), HN; see also
Günthart (1987b)

Edwardsiana lanternae (W. Wagner, 1937)

See taxonomic remarks above.
So far E VI – B IX; egg, 2 gen.
 Typhlocybinae Kirschbaum, 1868 133

Described after 2 †† collected in street lamp domes in Hamburg-Altona, shortly after-

wards 1 † was recorded in the Duvenstedter Brook to the north of Hamburg. There are
two more recent records from the Emsland near Lingen „1 † in einem Alnus-domini-
erten Feuchtgebüsch, 04.VII.89, 1 † in einer Mischgehölzhecke, 04.IX.89“. Reported from
Alnus glutinosa and Sorbus aucuparia in England and Wales. Otherwise the distribution
of this taxon is poorly known, since it is usually considered to be conspecific with the
preceding one (see above).
Wagner (1937d, 1941b), Niedringhaus & Olthoff (1993); see also Claridge & Wilson (1976), Le Quesne &
Payne (1981)

Edwardsiana soror (Linnavuori, 1950)

In the Czech Republic and Slovakia B VI – B X (Lauterer 1980); egg, 2 gen.
The range of this species, which is usually reported from Alnus incana, extends from
eastern Siberia to the eastern half of central Europe. There are only two German records:
Oberlausitz: Quoos, c. 150 m a.s.l., 02.IX.1958, 17 †† „auf Teichdämmen an Alnus“, and
Mittelfranken: Langensendelbach, 340 m a.s.l., 09.VII.2001, 1 †, on Alnus incana in the
undergrowth of an open pine forest. Has been found at 930 m a.s.l. in Carinthia (Aus-
tria).
Förster (1961), HN; see also Anufriev (1975), Anufriev & Emelyanov (1988), Holzinger (1999b), Lauterer
(1980)

Edwardsiana nigriloba (Edwards, 1924)

So far M VI – M VIII; egg, 2 gen.
Monophagous on Acer pseudoplatanus in mountain forests, river floodplains and parks,
generally preferring rather cool sites. Widespread in Germany (even found on Heligo-
land), but only in low frequency, probably overlooked, frequently found up to at least
800 m a.s.l. in the Mittelgebirge, but single specimens at 1700 m a.s.l. in the Bavarian
Alps. Not recorded from Baden-Württemberg, Brandenburg and Mecklenburg-Vorpom-
mern, but from France, Switzerland and Poland. In more northern parts of Europe, the
species is only known from a few records from southern Sweden and Scotland.
Niedringhaus & Olthoff (1993), Remane & Reimer (1989), Schiemenz (1990), Wagner (1935, 1939a), Walter
(1998), Nickel (1999b), HN; see also Le Quesne & Payne (1981), Nast (1976a, 1987), Ossiannilsson (1981)

Edwardsiana plebeja (Edwards, 1914)

M VI – M X; egg, 2 gen.
Monophagous on elms (Ulmus minor, U. laevis, U. glabra); usually swept along forest
margins and alleys and on solitary trees. Widespread in the lowlands of Germany, in
intermediate frequency on the hosts, but only scattered and with strongholds in river
valleys, the highest localities are at only c. 450 m a.s.l. in the German Mittelgebirge, but
at 800 m a.s.l. in the Engadine (Switzerland).
Fischer (1972), Schiemenz (1990), Wagner (1935, 1939a), Nickel (1999b), HN; see also Günthart (1987a)
134 Review of species

Edwardsiana prunicola (Edwards, 1914)

Typhlocyba barbata Ribaut, 1931
E. prunicola (Edw.) was described from Prunus, E. barbata (Rib.) from Salix. Most authors treat
them as conspecific due to the apparent lack of morphological differences (e.g. Nast 1972, 1987;
Ossiannilsson 1981).
B VI – M X; egg, 2 gen.
Usually on hairy-leaved willows (Salix cinerea, S. aurita, S. viminalis, occasionally also S. caprea)
in various wetlands, as well as on species of Prunus (mainly P. spinosa and P. domestica), along
hedges and forest margins and in fruit-tree plantations, a few records also on ornamental
Viburnum lantana. Widespread in Germany, usually in intermediate frequency and individual
numbers, but poorly documented (e.g. not recorded from Bavaria to the south of the Danube);
highest localities are at 600 m a.s.l. in the Mittelgebirge, at 1250 m a.s.l. in the Swiss Alps.
Achtziger (1991, 1994), Frommer (1996), Günthart (1988), Niedringhaus (1991), Reimer (1992), Remane
(1987), Ribaut (1936), Schiemenz (1990), SMNS, Wagner (1935, 1939a), HN; see also Günthart (1987b)

Edwardsiana rosae (Linnaeus, 1758)

B V – E X, occasionally XI/XII; egg, 2 gen., perhaps only 1 at higher altitudes.
On various roses (Rosa canina, R. corymbifera, R. rubiginosa, R. rugosa and others) in moder-
ately shady to sunny, damp to dry sites; in lower numbers and preferentially (perhaps ex-
clusively) in the second generation also on other woody species of Rosaceae (Prunus spinosa,
species of Rubus and Sorbus, cultivated cherry, plum, peach and apple, in Wales also record-
ed from cultivated strawberries). Widespread and common all over Germany, frequently
found up to at least 1500 m a.s.l. in the Bavarian Alps. Locally this species is extremely
abundant on ornamental roses in gardens and parks and may cause feeding damage.
Achtziger (1991), Lehmann (1973a, 1973b), Müller (1956), Niedringhaus (1991), Reimer (1992), Remane
(1987), Schiemenz (1990), Schwoerbel (1957), Trümbach (1959), Wagner (1935), HN; see also Claridge &
Wilson (1978a), Günthart (1971b)

Edwardsiana rosaesugans (Cerutti, 1939)

M VII – E IX; egg, 1(?) gen.
On wild roses (mainly Rosa pendulina, a few individuals have been found on R. canina
agg.), in open forests, shrubs and pastures from the montane to the subalpine belt. In
Germany only known from 8 localities in the Bavarian and Allgäu Alps (Ponten above
Hindelang, Schachen above Garmisch-Partenkirchen, Sojernspitze above Mittenwald,
Brauneck above Lenggries, as well as Roßfeld, Ahornbüchsenkopf, Jenner and Wim-
bach valley, all above Berchtesgaden) between 900 and 1800 m a.s.l.
Remane & Fröhlich (1994b), Nickel (1999b), HN; see also Cerutti (1939a), Claridge & Wilson (1978b), Lau-
terer (1983)

Edwardsiana salicicola (Edwards, 1885)

M VI – E X; egg, 2 gen., perhaps only 1 at higher altitudes.
 Typhlocybinae Kirschbaum, 1868 135

On hairy-leaved willows (Salix cinerea, S. aurita, S. caprea, occasionally also S. viminalis

and S. appendiculata) in rather wet or cool sites, e.g. along margins of bogs and fens, in
wet meadows, forest glades, and among subalpine scrub. Probably widespread in Ger-
many, although poorly documented. Usually in intermediate to high frequency and
rather high individual numbers on the hosts, recorded at least up to 900 m a.s.l. in the
Mittelgebirge, up to 1500 m a.s.l. in the Bavarian Alps.
Frommer (1996), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Remane & Reimer (1989), Schiemenz
(1990), Trümbach (1959), Wagner (1935, 1939a), Nickel (1999b), HN

Edwardsiana spinigera (Edwards, 1924)

M VI – M IX; egg, 2 gen.
On Corylus avellana; locally syntopic with E. avellanae (Edw.), but generally preferring
dryer and warmer sites, reaching the edge of the range in the north German plain. At
least scattered and in low frequency in southern and middle parts of Germany, appar-
ently scarce in the north (recorded near Bremen and Lüchow). Not reported from Den-
mark and Scandinavia. Highest localities are at 850 m a.s.l. in the Mittelgebirge, at 1350 m
a.s.l. in the Swiss Alps.
Achtziger (1991), Frommer (1996), Niedringhaus & Olthoff (1993), Post-Plangg & Hoffmann (1982), Reimer
(1992), Schiemenz (1990), SMNS, Trümbach (1959), Nickel (1997), HN; see also Günthart (1987b)

Edwardsiana smreczynskii Dworakowska, 1971

So far M VI – M IX, in Krakow according to Dworakowska (1971) until M X; egg, 2(?) gen.
So far altogether known only from 8 localities: Crakow (locus typicus), Paris (described
almost synchronously as E. guentharti Dlabola, 1971!), Caen (Normandy), Warsaw, Ba-
jtava (Slovakia), Basel (Switzerland), Main valley near Schweinfurt (Unterfranken), 220 m
a.s.l., 12.IX.1994, 8 ††, on Ulmus spec., and Oberbergen (Kaiserstuhl) near Freiburg,
14.VI.2002, 2 ††, on Ulmus minor. Nymphal development takes place on elm (perhaps
Ulmus laevis), but in Crakow a few adults have also been found on other shrubs and
trees (Cornus, Crataegus, Betula, Malus, Rosa, Salix).
Malenovsky (pers. comm.), Nickel (1999b); see also Chudzicka (1982), Dworakowska (1971), della Giustina
(1999), Günthart (1974), Lauterer (1980), Mühlethaler (2001)

Edwardsiana stehliki Lauterer, 1958

M VI – B IX; egg, 2 gen., in Slovakia until M XI (Lauterer 1984).
On hazel (Corylus avellana, in Slovakia also C. maxima, but not C. colurna) in damp to dry
sites. From Germany there are only two records, both from the Oberlausitz: „Umge-
bung Bautzen, 1957-61 (häufig)“, as well as Ebersbach near Görlitz, 01.IX.1998, 1 †, on
C. avellana. Is reported from Sorbus aria and Viburnum at 600 m a.s.l. in Switzerland.
Otherwise only known from Sweden, Poland and Austria.
Förster (pers. comm. to Lauterer 1984), HN; see also Günthart (1987b)
136 Review of species

Edwardsiana tersa (Edwards, 1914)

E V – E IX; egg, 2 gen.
On Salix viminalis, usually in valleys of rivers and streams, perhaps also on other spe-
cies of willows (S. triandra, S. pentandra). Widespread in the lowlands of Germany, up to
at least 650 m a.s.l., often in high frequency and individual numbers on the host, but not
recorded to the south of the Danube. In Austria the species is only reported from a
single locality in Styria (Judenburg, 680 m a.s.l., on Salix daphnoides), in France south-
ward to Paris; recently found in Basel (Switzerland).
Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1990), Schwoerbel (1957),
Wagner (1935), HN; see also Holzinger (1995a), della Giustina (1989)

Eupterycyba jucunda (Herrich-Schäffer, 1837)

B VII – E X; egg, 1 gen.
Monophagous on Alnus glutinosa, mainly near standing or running water, also in forests
of moist to wet sites and mining areas. Widespread in the lowlands of Germany, in
intermediate to high frequency, but usually low individual numbers, recorded only up
to 600 m a.s.l. in the Mittelgebirge and the foothills of the Alps.
Achtziger (1991), Fischer (1972), Frommer (pers. comm.), Heller (1987a), Niedringhaus (1991, 1997), Re-
imer (1992), Remane & Wachmann (1993), Schiemenz (1990), Wagner (1935, 1939a, 1951a), HN

Linnavuoriana decempunctata (Fallén, 1806)

Adults I – XII, mainly B VIII – VI; adult, 1 gen.
On birches (Betula pendula and B. pubescens), usually in cool sites, e.g. river basins (also
in urban parks), fens and bogs; hibernating individuals were found on Picea abies. In
Germany probably much under-recorded due to late emergence, but apparently wide-
spread and usually in low to intermediate frequency, up to at least 1000 m a.s.l.
Frommer (1996), Niedringhaus & Olthoff (1993), Schiemenz (1990), Schönitzer & Oesterling (1998b), SMNS,
Wagner (1935), HN

Linnavuoriana intercedens (Linnavuori, 1949)

Has been treated as a subspecies of L. decempunctata (Fall.), but is likely to be specifically distinct
due to constant differences in the morphology of the aedeagus and different host plant (Lauter-
er & Novotný 1991).
So far B VII, in Moravia at least until M X (Lauterer & Novotný 1991); ad.(?), 1 gen.
So far only known from Norway, Sweden, Finland, northern Russia, Latvia and Cze-
chia, but is perhaps more widespread, and published records of L. decempunctata (Fall.)
from Alnus in Poland, Switzerland and Italy should be revised. There is a single recent
record from southeastern parts of Germany: Langensendelbach near Forchheim, 340 m
a.s.l., 09.VII.2001, 2 ††, 1 ‡ (freshly emerged), on Alnus incana in a forest clearing. Is
apparently monophagous on A. incana. The altitudes of Bohemian localities range from
 Typhlocybinae Kirschbaum, 1868 137

200 to 730 m a.s.l. Specimens reported from the Engadine and the Valais (both Switzer-
land) at altitudes between 1500 and 1700m a.s.l. may also belong to this species.
HN; see also Ossiannilsson (1981), Lauterer & Novotný (1991), Günthart (1987a)

Linnavuoriana sexmaculata (Hardy, 1950)

E VII – X, single ‡‡ frequently until M VI; mainly egg(?), 1 gen.
On hairy-leaved species of willows in various wetland habitats (along rivers and streams,
in bog areas, near lakes and ponds, in mining areas, etc.) as well as in forest glades;
usually in high frequency on Salix viminalis, less frequently on S. cinerea, S. caprea, S.
aurita, as well as S. alba, in the Alps and their foothills also on S. eleagnos. Has been taken
from Pinus and Picea in winter. Widespread and fairly common in Germany, although
usually found in low numbers only; the highest localities are at 700 m a.s.l. in the Mittel-
gebirge, at 1000 m a.s.l. in the Bavarian Alps, at 1520 m a.s.l. in the Engadine (Switzer-
land).
Frommer (1996), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987), Schiemenz (1990), Schwo-
erbel (1957), Wagner (1939a), Nickel (1999b), HN; see also Günthart (1987a)

Lindbergina aurovittata (Douglas, 1875)

According to Le Quesne (1977) this taxon shows pronounced intraspecific variation in morphol-
ogy of the aedeagus. Accordingly, he considers Lindbergina pandellei (Lethierry, 1878) to be a
junior synonym of L. aurovittata (Dgl.). Several authors, however, have treated Typhlocyba pan-
dellei spoliata Horváth, 1907, originally described only as a variety of the latter, as a distinct
species (e.g. Drosopoulos et al. 1986; D’Urso 1995). Compared to L. aurovittata (Dgl.), differences
in life history are apparently weak, if present at all (Vidano & Arzone 1987b; Vidano et al. 1990;
Bosco & Arzone 1991).
E V – E X; egg, 2 gen.
The range of this taxon includes the Mediterranean region and parts of western Europe.
Since the beginning of the 1990s it has been found in 6 localities in the extreme west of
Germany: middle Rhine valley near Cologne, upper Rhine valley near Bruchsal, Nahe
valley near Schloßböckelheim, Neckar valley near Heilbronn (large populations on cul-
tivated Rubus fruticosus and perhaps noxious), as well as Palatinate, Haardtrand near
Wachenheim and Annweiler (on Quercus petraea), all sites between 50 and 200 m a.s.l.;
also reported from the French side of the southern upper Rhine plain near Colmar. From
southern and western Europe a remarkable host shift has been described: in Wales,
overwintering eggs are laid in leaves of evergreen Rubus (presumably mainly R. frutico-
sus), adults of the spring generation migrate to other woody species for oviposition
(mainly Quercus robur and Fagus sylvatica, also Alnus glutinosa, Corylus avellana, Betula
pubescens, Quercus cerris, Qu. petraea); the summer generation emerging from these eggs
return to Rubus for laying the winter eggs. In Piedmont (Italy), however, winter eggs are
laid on evergreen oaks (Quercus ilex and Qu. suber); the spring generation migrates to
deciduous species (Quercus pubescens, Qu. cerris) and other trees.
Schrameyer (pers. comm.), Frommer (pers. comm.), Fröhlich & Nickel (unpublished data), HN; see also
Claridge & Wilson (1978a), della Giustina & Balasse (1999), Vidano & Arzone (1987b), Vidano et al. (1990)
138 Review of species

Ribautiana alces (Ribaut, 1931)

So far M VI – B IX, unidentified ‡‡ of either this or the following species until B X; egg,
2(?) gen.
On oak (Quercus robur, perhaps also Qu. petraea) in rather warm sites, preferentially in
upper slope or plateau situations and along sunny forest margins. In Germany appar-
ently at the northeastern edge of the range, only recorded in middle parts (Donners-
berg/Palatinate, Mainzer Sand, middle Rhine valley near Bacharach, Ahr valley to the
south of Bonn, middle Hessen, surrounding plates of the Thuringian Basin near Artern
and Arnstadt), up 450 m a.s.l., but up to 1300 m a.s.l. in the Valais (Switzerland). Not
recorded from Denmark, Scandinavia and Poland.
Remane (1987, and pers. comm.), Wagner (1939a), HN; see also Cerutti (1939a), Nast (1987)

Ribautiana scalaris (Ribaut, 1931)

So far mainly M VII – M IX, single records B VI and B X; egg, 2(?) gen.
Like the preceding species, on oak in rather warm sites, but preferring Quercus petraea
instead of Qu. robur; adults being usually swept along sun-exposed forest margins, oc-
casionally in alleys or groups of trees. Widespread at least in middle and southern parts
of Germany, but sporadic, in low frequency and low individual numbers; not recorded
in the area to the south of the Danube. Northernmost localities are near Herne, Ham-
burg, Göttingen and Eberswalde, but the species has been found in southern Scandina-
via and Poland. The highest German sites are at 400 m a.s.l. only, but at 1300 m a.s.l. in
the Valais (Switzerland).
Köhler (pers. comm.), Niedringhaus & Olthoff (1993), Schwoerbel (1957), Trümbach (1959), Wagner (1939a),
Wagner & Franz (1961), Nickel (1997), HN, see also Cerutti (1939a), Chudzicka (1982), Claridge & Wilson
(1981), Ossiannilsson (1981)

Ribautiana debilis (Douglas,1876)

E V – B XII; egg (presumably also single ‡‡), 2 gen.
The first German report of this species dates only from 1994. Shortly afterwards it was
found in numerous sites in the valleys of the upper and middle Rhine, the Nahe, Ne-
ckar and Main, as well as in middle Hessen. Most specimens were sampled on Rubus
fruticosus and R. caesius, along hedges and forest margins and in the undergrowth of
parks and open forests. The northeastern edge of the range follows a line from Cologne
to Marburg, Würzburg, Heilbronn, Stuttgart and Freiburg; highest localities are at 300 m
a.s.l. only. In France and Britain the species is reported from various woody plants,
preferentially members of the Rosaceae (Malus, Pyrus, Prunus, Cydonia, Rosa, Rubus),
Quercus, Alnus, also Corylus, Betula, Tilia, Ulmus and others.
Frommer (1996), Heller (1994), Remane (1995), Schrameyer (pers. comm.), HN; see also Le Quesne & Payne
(1981), Ribaut (1936)
 Typhlocybinae Kirschbaum, 1868 139

Ribautiana tenerrima (Herrich-Schäffer, 1834)

E V – E X, occasionally until III; egg, 2 gen.
On Rubus fruticosus, R. idaeus and R. caesius in semi-open cultivated areas (forest mar-
gins and clearings, gardens, hedges, etc.), in low numbers also on other woody plants,
mainly Quercus, Salix, Acer, Prunus and Corylus. In Italy and Wales, breeding is reported
to take place on Rubus and Corylus only. Widespread and common in the lowlands of
Germany, but probably under-recorded; only single specimens found to the south of the
Danube and above c. 600 m a.s.l. (in the Mittelgebirge), but reported from 900 m a.s.l. in
the Valais (Switzerland).
Achtziger (1991), Frommer (1996), Heller (1994), Niedringhaus (1991), Post-Plangg & Hoffmann (1982),
Reimer (1992), Remane (1987), Schiemenz (1990), Wagner (1935, 1939a), HN; see also Arzone & Vidano
(1987a), Claridge & Wilson (1981), Raine (1960)

Ribautiana ognevi (Zachvatkin, 1948)

So far E V – M IX; egg, 2 gen.
This species’ range essentially includes the eastern part of the European deciduous forest
zone, extending westward to Bohemia (Czech Republic) and eastern parts of Austria and
Germany, where it is currently known from 5 localities up to 200 m a.s.l. only: Main flood-
plains near Schweinfurt, Oder valley near Wriezen and Angermünde, and Elbe floodplains
near Gorleben and Burg. Lives on Ulmus laevis usually in river floodplains, apparently in
intermediate frequency and low to intermediate individual numbers. Is perhaps much un-
der-recorded, but has not been found in host stands in the valleys of the Rhine and Leine.
Nickel (1999b), HN; see also Dlabola (1958, 1977), Emelyanov (1964a) (see this paper also for identification)

Ribautiana ulmi (Linnaeus, 1758)

E V – B XI; egg, 2 gen.
On elms – in the lowlands usually Ulmus minor and U. laevis, in more mountainous
regions mainly U glabra – in forests and open stands as well as on solitary trees. Wide-
spread and fairly common in Germany, in high frequency and individual numbers on
the hosts, occurring up to at least 1000 m a.s.l. in the Bavarian Alps, up to at least 1350 m
a.s.l. in the Engadine (Switzerland).
Achtziger (1991), Bittner & Remane (1977), Fischer (1972), Reimer (1992), Remane (1987), Schiemenz (1990),
Wagner (1935, 1951a), HN; see also Günthart (1987a)

Typhlocyba quercus (Fabricius, 1777)

Mainly M VI – M IX, occasionally until IV; egg, 1(?) gen.
In semi-open cultivated areas, mainly along forest margins, in river floodplains, fruit-
tree plantations and along hedges, usually on species of Prunus and Quercus (P. padus, P.
avium, P. spinosa, P. domestica, Qu. robur, Qu. petraea); in low numbers also on other woody
species (Corylus, Carpinus, Fagus and others); in Wales breeding has been found to take
140 Review of species

place on these (as well as on Castanea and ornamental Nothofagus). Widespread in Ger-
many, but often in intermediate frequency and low individual numbers only. Highest
localities are at 900 m a.s.l. in the Mittelgebirge and Bavarian Alps, at 1240 m a.s.l. in the
Engadine (Switzerland).
Achtziger (1991), Fischer (1972), Lehmann (1973a), Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992),
Remane (1987), Schiemenz (1990), Schwoerbel (1957), Trümbach (1959), Wagner (1935, 1939a), HN; see also
Claridge & Wilson (1976, 1981), Günthart (1987a)

Zonocyba bifasciata (Boheman, 1851)

E VI – M X; egg, 1(?) gen.
On Carpinus betulus, Ulmus minor and U. glabra, in closed forests as well as in semi-open
tree stands, occasionally even along hedges. Single specimens found on Acer, Tilia, Al-
nus, and other deciduous trees, but probably without breeding. Widespread in the low-
lands of Germany, but often in low individual numbers only, usually in high frequency
on Carpinus, but less frequently on Ulmus; highest localities are at 600 m a.s.l.
Frommer (1996), Günthart (1988), Niedringhaus (1997), Post-Plangg & Hoffmann (1982), Reimer (1992),
Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), HN, and others; see also Clar-
idge & Wilson (1981)

Eurhadina concinna (Germar, 1831)

B VI – B X; egg, 1 gen.
Preferentially on Quercus robur and Qu. petraea in forests and semi-open stands, in lower
numbers on Fagus, Betula, Alnus (in Wales breeding was found to take place on these),
Tilia and others. Widespread in the German lowlands, usually in intermediate to high
frequency on both oak species, but recorded only up to 700 m a.s.l. in the Mittelgebirge
and the foothills of the Alps.
Achtziger (1991), Frommer (1996), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Remane (1987),
Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), HN; see also Claridge & Wilson (1976, 1981)

Eurhadina kirschbaumi W. Wagner, 1937

So far M VII – B IX; egg, 1 gen.
On Quercus petraea; adults are usually swept along forests roads and margins, mainly in
acidic, occasionally also basic, usually damp to moderately dry sites. In Germany rath-
er sporadic and in low frequency, but perhaps overlooked. Recorded from the surround-
ings of Hamburg (locus typicus), Berlin, Göttingen, Artern, Ahrweiler, Koblenz, Mainz,
Saarbrücken, Aschaffenburg, Würzburg, Schweinfurt, Arnstadt, Niesky and Ludwigs-
burg. Highest localities are at 450 m a.s.l. only. Is also reported to live on Quercus pube-
scens, Qu. cerris and Castanea sativa in the Mediterranean region; these potential host
species have not yet been sampled in Germany, however.
Post-Plangg & Hoffmann (1982), Remane (pers. comm.), SMNS, Wagner (1937a, 1939a, 1951a), ZMB, Nick-
el (1994), HN; see also Drosopoulos (1999), Vidano & Arzone (1987b)
 Typhlocybinae Kirschbaum, 1868 141

Eurhadina loewii (Then, 1886)

Eurhadina untica Dlabola, 1967
After Dlabola (1967a) north and central European populations with unforked lateral appendag-
es of the aedeagus are conspecific with Eurhadina untica Dlabola, 1967 described from Ulmus
pumila in Mongolia. The author also suggested that individuals with forked appendages mainly
reported from the southern half of Europe belong to a distinct species (E. loewii s.str.). This
opinion is rejected here. According to Lauterer (pers. comm.) the type of E. loewii (Then) shows
an anomalous spinulation of the aedeagus branches.
M VI – E IX; egg, 1 gen.
On Acer pseudoplatanus, often in shady or cool situations (preferentially on slopes and in
river floodplains, but also in parks and gardens). Widespread in Germany, in low to
moderate individual numbers and in intermediate frequency on the host, found to at
least 900 m a.s.l. in the Bavarian Alps, up to 1290 m a.s.l. in the Engadine (Switzerland).
Bittner & Remane (1977), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer (1992), Schiemenz (1990),
SMNS, Wagner (1935, 1939a), Nickel & Remane (1996), HN; see also Günthart (1987a, 1987b)

Eurhadina pulchella (Fallén, 1806)

B VI – B X; egg, 1 gen.
On Quercus robur and Qu. petraea, usually in forests and semi-open stands, occasionally
on solitary trees; in low numbers also on Betula, Fagus, Carpinus and others, but proba-
bly without breeding. Widespread in the lowlands of Germany, usually in high frequen-
cy and intermediate individual numbers on both host species, found up to at least 750 m
a.s.l. in the Allgäu, but reported from 1300 m a.s.l. in the Valais (Switzerland).
Achtziger (1991), Fischer (1972), Frommer (1996), Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992),
Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), HN; see also Cerutti (1939a),
Claridge & Wilson (1981), Vidano & Arzone (1987b)

Eurhadina ribauti W. Wagner, 1935

E V – M X; egg, 2(?) gen.
Like the preceding species, mainly on Quercus robur and Qu. petraea, occasionally Fagus,
Carpinus, Tilia, Alnus and others, but breeding apparently only on Quercus. Widespread
in Germany, but apparently confined to lowland regions (recorded up to 450 m a.s.l.
only), usually in low to intermediate frequency and individual numbers. Not recorded
to the south of the Danube, except from a single locality in the Lake Constance Basin.
Niedringhaus & Olthoff (1993), Remane (1987), Schiemenz (1990), Wagner (1935, 1939a), Nickel & Remane
(1996), HN; also Claridge & Wilson (1981)

Eurhadina saageri W. Wagner, 1937

E VI – B X; egg, 1 gen.
On oak, mainly (perhaps exclusively) on Quercus robur. Adults are usually swept along
forest margins and in semi-open host stands in lowland areas and river basins. The few
142 Review of species

host records from other European countries also refer exclusively to Qu. robur. In Ger-
many only very sporadic, in low frequency and individual numbers, at the northern
edge of the range. Border localities are near Wilhelmshaven, Hamburg (locus typicus),
Lübeck, Wolfsburg, Gera and Kamenz, up to at most 550 m a.s.l.
Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Schiemenz (1990), Wagner (1937a, 1951a), HN;
also Anufriev & Kirillova (1998), Lauterer (pers. comm.)

Eupteryx atropunctata (Goeze, 1778)

M V – E X; egg, at least 2 gen.
Polyphagous in rather tall herb stands in wet to dry, usually sunny sites (low-input
grassland, waysides, ruderal habitats, abandoned fields, fens and moderately saline
sites, also synanthropic in gardens and fields). Host plants are mainly Lamiaceae (Men-
tha, Salvia, Clinopodium, Origanum, Ballota, Melissa and others), also Verbascum thapsus,
Thalictrum flavum, Solanum tuberosum, Pastinaca, presumably also Cirsium and Urtica
dioica. Furthermore, Lavatera, Althaea, Nepeta, Lamium, Beta vulgaris and Vicia faba are
mentioned in the literature. Widespread and common in Germany, found up to at least
900 m a.s.l. in the Mittelgebirge and the Bavarian Alps. Is reported from 1500 m a.s.l. in
Switzerland and 1700 m a.s.l. in Austria, but without information on breeding at these
altitudes.
Fischer (1972), Fröhlich (1996a), Niedringhaus (1991, 1997), Post-Plangg & Hoffmann (1982), Reimer (1992),
Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), Wais (1990), Nickel & Acht-
ziger (1999), HN, and others; see also Günthart (1987a), Stewart (1988), Wagner & Franz (1961)

Eupteryx aurata (Linnaeus, 1758)

M V – E X; egg, 2 gen.
Like E. atropunctata (Goeze) in tall herb stands, but usually in cool or moderately shady,
moist to temporarily flooded sites, often near water, along forest roads and among sub-
alpine scrub, in lower numbers in gardens and ruderal sites. Host plants are tall herbs,
in the first generation Urtica dioica, in the second also various species of Asteraceae
(mainly Petasites, Arctium, Senecio, Cirsium, Dahlia), Lamiaceae (mainly Lamium, Men-
tha), Apiaceae (Heracleum sphondylium, also H. mantegazzianum), Malvaceae (Althaea ro-
sea) and others. Literature records include Humulus lupulus and Solanum tuberosum, as
well as additional species of the plant families mentioned above. Widespread and very
common all over Germany, also in higher parts of the Mittelgebirge, frequently found up
to 1700 m a.s.l. in the Bavarian and Allgäu Alps.
Fischer (1972), Niedringhaus (1991, 1997), Reimer (1992), Remane (1987), Schiemenz (1990), Wagner (1935,
1939a), Nickel & Achtziger (1999), HN; see also Stewart (1988)

Eupteryx austriaca (Metcalf, 1968)

Published data on life history and distribution appear to be contradictory, perhaps due to taxo-
nomic problems. The status of north German lowland populations, in particular, is in need of
revision.
 Typhlocybinae Kirschbaum, 1868 143

M VI – B X; egg, 1(?) gen.

Mainly on Knautia dipsacifolia in damp to moist, moderately shady sites of the montane
and subalpine belt, usually along sunny forest margins and roads, as well as in subal-
pine scrub. In Germany largely confined to the Alps and the southern Black Forest,
between 750 and 1850 m a.s.l., in intermediate frequency on the host. On the other hand,
there are older records from Succisa pratensis in wetland habitats of northern Germany
(Albersdorf/Holstein, 24.VII.1933, and Duvenstedter Brook near Hamburg, 22.VIII.1937).
Fischer (1972), Wagner (1935, 1941b), HN

Eupteryx heydenii (Kirschbaum, 1868)

According to Nast (1987), this species has been described after two syntypes, one † from Swit-
zerland and one ‡ from Germany. However, according to Fieber (1872a, 1872b), who studied the
type material, the ‡ (collected near Bad Rippoldsau, Black Forest) belongs to the preceding spe-
cies.
So far E VI – E IX; egg, 1(?) gen.
In tall herb stands in the submontane to the subalpine belt, usually along forest margins
and roads, along streams and ditches, and in montane pastures. The host plant is Chaero-
phyllum hirsutum, perhaps also Ch. aureum. Furthermore, Petasites has been reported
from the Austrian and Swiss Alps, but these feeding records are probably erroneous.
There are only few published data from Germany, although the species is widespread
and rather common at least in the Alps and their foothills; there is also a number of
localities in the Mittelgebirge (Harz Mountains, Rhön Mountains, Thuringian Forest,
Frankenwald, Vogtland, Erzgebirge, Franconian Jura and Oberpfälzer Wald, probably
also in other hilly regions). According to existing data, these are at the northern edge of
the range. Found between 400 and 1850 m a.s.l. in Germany, up to 2000 m a.s.l. in Styria
(Austria).
Fischer (1972), Reimer (1992), Schiemenz (1990), Schönitzer & Oesterling (1998b), Walter (pers. comm.),
HN; see also Cerutti (1939a), Günthart (1974), Wagner & Franz (1961)

Eupteryx lelievrei (Lethierry, 1874)

B VI – E X; egg, 2 gen.
On Betonica officinalis in temporarily wet to temporarily dry, usually moderately shady
sites, mainly along forest margins, near shrubs, and in low-input meadows, preferen-
tially on clayey soils. In Germany only very sporadic, in low frequency and low densi-
ties; at present there are only 10 known localities: southern Niedersachsen (near Göttin-
gen), Thuringian Basin (near Erfurt), middle Hessen (Wermertshausen), Taunus (Nie-
derreifenberg), Franconia (Sulzheim and Gesees), Kaiserstuhl (Vogtsburg) and Bavari-
an Alps (Berchtesgaden, Ramsau, and Nußdorf am Inn, c. 1300 m a.s.l.). Otherwise only
known from a few records from Belgium, France, Switzerland, Austria, the Czech Re-
public, Slovakia, Bulgaria and middle Russia.
Fechtler (pers. comm.), Remane & Fröhlich (1994b), Nickel & Voith (unpublished data), Nickel (1994), HN;
see also Lauterer (1983, 1984), Nast (1987)
144 Review of species

Eupteryx origani Zachvatkin, 1948

B VI – B X; egg, 2 gen., probably 1 at higher altitudes.
On Origanum vulgare, in oligotrophic, moderately dry to damp, moderately shady sites;
mainly along herbaceous margins in dry grassland, montane meadows and pastures,
preferably near trees or shrubs. In Germany only very sporadic and known from scat-
tered localities in the Bavarian Alps (up to at least 1600 m a.s.l.), the Franconian and
Swabian Jura, the Frankenwald, the Thuringian Forest, the middle Rhine and its tribu-
taries (basins of Ahr, Lahn and Dill) and the north German plain (near Hamburg and
Berlin). To the west of this, there are records from Scotland, England, the Netherlands,
Switzerland and south-eastern France (Savoie). Has been reported from Veronica
chamaedrys, but these data require confirmation.
Remane (1961b), Remane & Fröhlich (1994b), Post-Plangg & Hoffmann (1982), Schiemenz (1990), SMNS,
HN; see also della Giustina & Remane (2001), Günthart (1997), Le Quesne & Payne (1981), Nast (1987)

Eupteryx signatipennis (Boheman, 1847)

E VI – B X; egg, 2 gen.
On Filipendula ulmaria, usually in wet and cool sites (in fens, along shores and ditches, in
pastures and abandoned meadows, apparently absent from habitats mown more than
once a year). From Germany there are only very few published records, but the species
has been found in most regions except the northeast (Mecklenburg-Vorpommern, Bran-
denburg, Saxony-Anhalt), where it is probably overlooked. The highest localities are at
1200 m a.s.l. in the Allgäu Alps and the southern Black Forest.
Bittner & Remane (1977), Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer
(1992), Remane (1962), Remane & Fröhlich (1994b), Rombach (pers. comm.), Schiemenz (1990), Nickel &
Achtziger (1999), Nickel & Remane (1996), HN

Eupteryx adspersa (Herrich-Schäffer, 1838)

So far M VI – E IX, in Poland according to Dworakowska (1970a) V – X; egg, 2 gen.
Along dry waysides and in ruderal sites on various substrates (mainly sand, limestone
and gypsum), on Artemisia absinthium. The German localities are at the northwestern
edge of the range; there are only c. 15 records along the middle Rhine and its tributaries,
since 1890 (St. Goarshausen, Bacharach, Limburg an der Lahn), in the rain shadow area
to the east of the Harz Mountains (near Eisleben, Freyburg an der Unstrut, Artern), near
Lüchow, Berlin, Göttingen and Meißen. The occurrence at the locus typicus near Regens-
burg (Bavaria) is not confirmed due to formerly diverging interpretations of the spe-
cies’ identity, and because Herrich-Schäffer’s collection has been lost during the last
war, although the most closely related species, E. artemisiae (Kbm.) is not known from
southern Germany to the south of the Main river (see below). All German localities of E.
adspersa (H.-S.) are below 250 m a.s.l., but the species is reported from up to 1650 m a.s.l.
in the Engadine and Valais (both Switzerland).
Dworakowska (1970a; 1972a), Herrich-Schäffer (1838), Remane (pers. comm.), Schiemenz (1990), Walter &
Emmrich (1995), Nickel (1997), HN; see also Cerutti (1939a), Günthart (1987a)
 Typhlocybinae Kirschbaum, 1868 145

Eupteryx artemisiae (Kirschbaum, 1868)

E V – M X; egg, 2 gen.
In salt marshes along the coasts of the North and Baltic Sea, also locally inland in saline
sites and old gardens. Reports from dry grassland habitats probably refer to the preced-
ing species. Usually on Artemisia maritima, but on A. abrotanum in gardens. Known in
Germany from the East and North Frisian Islands (where locally abundant), the Jade
Bay, the coasts of Schleswig-Holstein, and the island of Poel; further populations occur
in the rain shadow area to the east of the Harz Mountains (Solgraben Artern), in Ham-
burg, Göttingen and Marburg (botanical gardens), as well as in Frankfurt am Main (lo-
cus typicus, no habitat given). All localities are below 200 m a.s.l.
Niedringhaus & Olthoff (1993), Remane (pers. comm.), Schiemenz (1990), Wagner (1935, and unpublished
data), Nickel (1997), HN; for identification of nymphs see Stewart (1986a)

Eupteryx calcarata Ossiannilsson, 1936

B V – E X; egg, 2 gen.
On Urtica dioica, in sunny, damp to moist sites; generally preferring more sunny and dry
habitats than the other Urtica-dwelling congeners, e.g. along ditches and streams in
open fields, in ruderal sites and mining areas, occasionally also in pastures. Widespread
in the lowlands of Germany, but only localized, rarely found above 400 m a.s.l., a single
record is from c. 750 m a.s.l. in the Allgäu.
Fischer (1972), Heller (1987a), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1962), Schiemenz
(1990), Schwoerbel (1957), Wagner (1939a, 1951a), Nickel (1994), HN

Eupteryx cyclops Matsumura, 1906

E V – E X; egg, 2 gen.
On Urtica dioica, mainly near standing or running water (also along ditches within fields),
herbaceous and woody margins, and in open forests. In comparison to other species of
Eupteryx on this host, with preference for moderately sunny, moist and higher-altitude
sites. Widespread and common all over Germany, frequently found up to at least 1000 m
a.s.l. in the Mittelgebirge, the Bavarian and Allgäu Alps (single individuals up to 1600 m
a.s.l.), up to 2000 m a.s.l. in the Valais (Switzerland).
Achtziger (1991), Fischer (1972), Heller (1987a), Mölleken & Topp (1997), Niedringhaus & Olthoff (1993),
Reimer (1992), Schiemenz (1990), HN, and others; see also Cerutti (1939a), Günthart (1987a), Stewart (1988),
Stiling (1980a); for identification of nymphs see Stewart (1986b)

Eupteryx immaculatifrons (Kirschbaum, 1868)

M VI – E X; egg, 2 gen.
Formerly, this species was often overlooked or misinterpreted. It lives on Lamium macu-
latum in moist to damp, usually shady sites, mainly in floodplain forests along rivers
and streams as well as on seeping slopes. In Germany recorded only from middle and
southern parts (locus typicus: Mombach near Mainz), apparently at the northern edge of
146 Review of species

the range, but it may be more widespread, together with its host. Border localities are
near Bonn, Marburg, Göttingen, Bad Frankenhausen, Dresden and Görlitz. Has been
found at 700 m a.s.l. in the Bavarian foothills of the Alps, but to the north of the Danube
it is usually confined to locations below 500 m a.s.l.
Büchs (1988), Emmrich (1975), Wagner (1939a), Remane & Fröhlich (1994b), Nickel (1994, 1999b), HN

Eupteryx urticae (Fabricius, 1803)

E V – E X; egg, 2 gen.
On Urtica dioica in damp to moderately wet, usually moderately shady to shady sites,
preferentially in various types of forests (often along roads and margins) and along
shores of standing or running water (also ditches). In Wales and England individuals of
the second generation may occur in numbers on Urtica urens and Parietaria judaica, al-
though overwintering eggs are found on U. dioica only. Widespread and common in
Germany, found up to at least 900 m a.s.l. in the Bavarian Alps, up to 1500 m a.s.l. in
Switzerland.
Achtziger (1991), Mölleken & Topp (1997), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz
(1990), Schwoerbel (1957), Wagner (1935, 1939a), HN, and others; see also Günthart (1987a), Stewart (1988,
and pers. comm.), Stiling (1980a); for identification of nymphs see Stewart (1986b)

Eupteryx collina (Flor, 1861)

This species has often been misinterpreted; thus, many older reports refer to other species. After
a revision of Flor’s collection Vilbaste (1973) concluded that Eupteryx alticola Ribaut, 1936 is a
junior synonym of E. collina (Fl.), and that E. florida Rib., which had originally been described
only as a variety, is a distinct species. However, there is some evidence that records published
even recently as E. collina (Fl.) may refer to a complex of several species.
So far M VI – E IX; egg, 1 gen., probably 2 in lower altitudes.
On Mentha longifolia along streams and ditches, as well as in seepy pastures. Is also
reported from Acinos alpinus in the Valais (Switzerland) and from various species of
Lamiaceae in Italian gardens. In Germany the species is largely confined to the Bavari-
an and Allgäu Alps and their foothills, mainly between 700 and 1700 m a.s.l. A few
isolated localities are near Augsburg (c. 500 m a.s.l.) and in the vicinity of Berlin (c. 60 m
a.s.l.). Most if not all other lowland reports refer to E. florida Rib. or ‡‡ which are diffi-
cult to identify.
Fischer (1972), Remane (1961b), Schiemenz (1990), ZMB, HN; see also Arzone (pers. comm.), Günthart
(1987a), Vidano & Arzone (1978)

Eupteryx stachydearum (Hardy, 1850) sensu Dworakowska (1972a)

This taxon and the following have long been considered to be conspecific, until Dworakowska
(1972a) pointed out that there are differences in head colouration and particularly in the shape
of the aedeagus (see also Vilbaste 1973). This opinion has been followed by some authors, e.g.
della Giustina (1989) and Remane & Fröhlich (1994a), but not by others, e.g. Nast (1987) and
Ossiannilsson (1981). Until further studies have been done, both shall be treated here as differ-
ent species, although differences are only slight and transitional stages may occur. Apparently,
 Typhlocybinae Kirschbaum, 1868 147

they both live oligophagously on various species of Lamiaceae in the forest herb layer, but have
never been found to occur syntopically, despite overlap of their host and geographical ranges.
The latter overlap in the region between the Danube and the northern edge of the Mittelgebirge,
as well as in northwestern Poland (see Nast 1976a). In Germany E. curtisii (Fl.) lives at the north-
ern edge of its range, whereas E. stachydearum (Hardy) is also reported from Denmark and south-
ern Sweden.
M V – E X; egg, 2 gen.
E. stachydearum (Hardy) prefers moist to damp, usually shady sites (notably beech and
oak forests) and lives on Stachys sylvatica and Lamium galeobdolon, but also on additional
species of Lamiaceae. It is widespread and fairly common in most parts of Germany,
although apparently less frequent in the north German plain, recorded up to at least
1200 m a.s.l. in the Black Forest and the Bavarian Alps. Many published records from
eastern Germany may also refer to the following taxon and are in need of revision.
Frommer (1996), Dworakowska (1972a), Reimer (1992), Remane (1987), Nickel (1997), HN; see also Ossian-
nilsson (1981); for identification see Dworakowska (1972a), della Giustina (1989)

Eupteryx curtisii (Flor, 1861)

See taxonomic note on the preceding species.
E V – M X; egg, 2 gen.
Generally more in warmer and less shady sites than E. stachydearum (Hardy) and ap-
parently not syntopic. Preferentially on Teucrium scorodonia in open oak and pine forests
and along their margins, in acidic, usually moderately dry sites, perhaps also on Stachys
sylvatica and Clinopodium vulgare as well as on other species of Lamiaceae. The distribu-
tion in Germany is poorly known. There are confirmed records from the area to the
north of a line from Lörrach to Singen am Hohentwiel and Passau, and to the south of a
line from Düsseldorf to Lengerich, Hannoversch Münden and Herzberg am Harz, usu-
ally below 600 m a.s.l., but also from sun-exposed slopes up to 1100 m a.s.l. in the south-
ern Black Forest.
Büchs (1988), Kolbe & Bruns (pers. comm.), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987),
Nickel & Sander (1996), Nickel (1997), HN

Eupteryx florida Ribaut, 1952

Mainly M V – E X, occasionnally until V; egg, in low numbers also adult, at least 2 gen.
In moist to moderately dry, usually moderately shady sites, often on Ballota nigra along
nitrophilous herbaceous and woody margins, on Stachys sylvatica in open forests and
along their margins, as well as in gardens on Salvia officinalis and Melissa officinalis; fur-
ther populations have been found on Clinopodium vulgare, Lamium album, Mentha longi-
folia, Stachys palustris and Glechoma hederacea. In Germany widespread and fairly com-
mon between the Danube and the north edge of the Alps (up to at least 600 m a.s.l.,
although single specimens were found at 1300 m a.s.l.), but uncommon in the north
German plain, where increasingly synanthropic and at the edge of the range. Border
localities are near Lingen (Ems), Bremen, Kiel, Rostock and Eberswalde.
148 Review of species

Frommer (1996), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987, and pers. comm.), Schie-
menz (1990), Nickel & Remane (1996), HN; see also Stewart (1988); for identification of nymphs see Stew-
art (1986a)

Eupteryx salviae Arzone & Vidano, 1994

So far only VIII; egg, 2(?) gen.
So far only known from northeastern Italy and the former Yugoslawia (Serbia, Croatia,
Montenegro), where reported from various species of Lamiaceae (Salvia officinalis, Teu-
crium polium, Thymus pulegioides, Th. praecox). Recently, a population was found on Salvia
officinalis in a garden in Stuttgart.
Remane (pers. comm.), Nickel & Remane (2002); see also Arzone & Vidano (1994)

Eupteryx decemnotata Rey, 1891

M V – E X; egg, 2 gen.
Until recently this species was only known from the Mediterranean region (southern
France, Italy) and Switzerland. Apparently, there is a dramatic recent range expansion.
The first German record is from Malaise traps from near Speyer in 1989; since then it
has been found in more than 30 sites in gardens, urban green and abandoned vineyards
in the upper Rhine plain (Weil am Rhein, Annweiler, Neustadt an der Weinstraße,
Wachenheim, Lorsch, Wiesbaden), in the basins of the Neckar, Nahe, Main and Regnitz
(Stuttgart, Schloßböckelheim, Würzburg, Bamberg, Forchheim), in Cologne, Münster,
Marburg, Oldenburg, Hildesheim, Göttingen, Erfurt and Berlin. Host plants are vari-
ous species of ornamental and officinal Lamiaceae (mainly Salvia officinalis, also Melissa
officinalis, Nepeta cataria and Thymus); all localities are below 300 m a.s.l. Recently it was
also recorded from the French and Swiss side of the southern upper Rhine plain near
Colmar and urban areas of Basel, from Vienna, Graz (both eastern Austria) and Nova
Gorica (western Slowenia).
Fechtler (pers. comm.), Frommer (pers. comm.), Remane (1995, and pers. comm.), Remane & Fröhlich (1994b),
Nickel & Remane (1996), HN; see also della Giustina & Balasse (1999), Günthart (1987b), Mühlethaler
(2001), Nast (1987), Holzinger & Seljak (2001), Vidano & Arzone (1978)

Eupteryx melissae Curtis, 1837

Mainly E V – B X, some adults and nymphs until V (see also Stewart 1988); (mainly?)
egg, 2(?) gen.
This species occurs mainly in southern and western parts of Europe. In Germany it
appears to prefer synanthropic sites and is known only from a few locations: Elend
(Harz Mountains), near Eisleben (recorded in 1910) and Quedlinburg (on Mentha and
Marrubium vulgare), Göttingen (botanical gardens as well as private gardens and parks),
also found in urban areas and gardens of Weil am Rhein, Stuttgart, Cologne, Münster,
Oldenburg, Marburg, Hildesheim, Erfurt and Berlin. Most recent records were from
ornamental Nepeta cataria and Salvia officinalis, a few also from Leonurus cardiaca and
Althaea officinalis, the latter in the second generation only. All localities are situated be-
 Typhlocybinae Kirschbaum, 1868 149

low 250 m a.s.l. except the first (a single specimen at c. 500 m a.s.l.). There is some evi-
dence of a recent range expansion; on the other hand the first German records date from
1910. From Britain and other countries, Lavatera arborea, Althaea rosea and Melissa offici-
nalis have been reported as additional hosts.
Fechtler (pers. comm.), Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Remane (pers. comm.), Schi-
emenz (1990), Nickel (1994), HN; see also Stewart (1988); for identification of nymphs see Stewart (1986a)

Eupteryx thoulessi Edwards, 1926

So far E VI – B X, according to Wagner (1941b) rarely until B XI; egg, 2 gen.
In sunny, wet to temporarily flooded sites, mainly tall-sedge swamps and reeds, locally
also in sand pits and moderately saline sites. Host plants are Mentha aquatica and occa-
sionally Lycopus europaeus; the latter probably only in the second generation. Further-
more, M. spicata and M. arvensis have been reported. In Germany the species occurs
rather locally, with strongholds in the northern half. Most records are from the north
German plain and southern Niedersachsen (up to at most 300 m a.s.l.); localities in the
Thuringian Basin (near Erfurt) and the southern upper Rhine plain (near Freiburg) ap-
pear to be rather isolated.
Kuntze (1937), Niedringhaus & Olthoff (1993), Schiemenz (1990), Wagner (1941b), Nickel (1997), HN; see
also Ribaut (1936), Stewart (1988); for identification of nymphs see Stewart (1986a)

Eupteryx tenella (Fallén, 1806)

B V – E X; egg, 2 gen.
On Achillea millefolium along herbaceous margins in moderately shady, damp to dry,
basic to acidic sites, mainly in dry grassland, along hedges and forest edges, occasional-
ly also in old gardens; apparently absent from mown meadows. Widespread in the mid-
dle parts of Germany, but rather uncommon, most populations being confined to very
small sites. Not recorded to the south of the Danube; very few localities are in the north
German plain (near Hamburg, Bad Schwartau, Rathenow and Seelow). Found up to c.
700 m a.s.l. in the Mittelgebirge.
Achtziger (1991), Frommer (pers. comm.), Nikusch (1976), Reimer (1992), Remane (1987), Schiemenz (1990),
Wagner (1939a), HN; for identification of nymphs see Stewart (1986a)

Eupteryx vittata (Linnaeus, 1758)

M V – E X; egg, 2 gen.
On various dicotyledons in sunny to shady, wet to moderately moist sites, mainly low-
input meadows and deciduous forests. Favoured host plants are Ranunculus repens and
Glechoma hederacea, but the species also lives on Ajuga reptans, Valeriana dioica and oth-
ers. Furthermore, from Britain it is reported to live on Veronica montana, Prunella vulgar-
is, Plantago major and Crepis paludosa, although some host species may be utilized in the
second generation only. Widespread in Germany and fairly common, extending to at
least 1350 m a.s.l. in the Mittelgebirge and the Alps.
150 Review of species

Achtziger (1991), Bittner & Remane (1977), Fischer (1972), Haas (1975), Niedringhaus (1997), Reimer (1992),
Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others;
see also Stewart (1988); for identification of nymphs see Stewart (1986a)

Eupteryx notata Curtis, 1837

M V – E X; egg, 2 gen., at higher altitudes probably 1.
Among low-growing herbs in oligotrophic and sunny, basic to acidic, dry to damp, oc-
casionally also rather wet sites, mainly low-productivity meadows and pastures, dry
grassland, heaths and roadside embankments, occasionally straw meadows and calcar-
eous sedge fens. Usually on Hieracium pilosella and Leontodon hispidus, but also reported
from Prunella, Thymus, Hypochaeris and Plantago. Widespread in Germany and fairly
common, with strongholds in regions with less intensive agriculture (e.g. at higher alti-
tudes, on sandy soils or in rather dry climates). Frequently found up to 1700 m a.s.l. in
the Bavarian and Allgäu Alps, up to 2100 m a.s.l. in North Tyrol (Austria), and even at
2250 m a.s.l. in Switzerland.
Achtziger (1991), Fischer (1972), Kuntze (1937), Reimer (1992), Remane (1987), Schiemenz (1969, 1990),
Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN; see also Günthart (1987a), Leis-
ing (1977), Morcos (1953), Payne (1981), Stewart (1988); for identification of nymphs see Stewart (1986a)

Eupteryx filicum (Newman, 1853)

B VI – E X; egg, 2 gen.
In damp to moderately dry, often acidic, more or less shady and rocky sites, usually in open
forests (often on roadside embankments), locally in gardens and on walls. Host plants are
various species of ferns, mainly Polypodium; in England and Wales P. vulgare and P. interjec-
tum in the first generation, additionally Polystichum, Dryopteris, Phyllitis, Asplenium, Athyri-
um and Pteridium in the second. The range essentially includes the Mediterranean region
and western Europe (northward to England, Wales and Ireland). From Germany it was first
published only recently and is known mainly from western parts, usually below 550 m
a.s.l., where it is locally common and occurs in high frequency. Localities near Remscheid,
Cologne, Göttingen, Ilmenau, Gotha, Tauberbischofsheim, Bad Dürkheim and Freiburg may
be situated at the edge of the range, but the species is probably under-recorded. A recent
eastward range expansion, as in other western European species, is unlikely; because it is
known from the Polish Beskid and Tatra Mountains since 1911 and 1925, respectively.
Fechtler (pers. comm.), Frommer (pers. comm.), Remane & Fröhlich (1994b), HN; see also Badmin (1991,
1992), Smreczynski (1954), Stewart (1988); for identification of nymphs see Stewart (1986a)

Wagneripteryx germari (Zetterstedt, 1840)

M VI – M X, occasionally until IV/V; egg, 1 gen.
On Pinus sylvestris and P. mugo, reported also from P. nigra and P. cembra in the Swiss
Alps; eurytopic in forests as well as on solitary trees, in bogs as well as among subalpine
scrub. Widespread in Germany and in intermediate to high frequency, found up to 1950 m
a.s.l. in the Bavarian and Allgäu Alps, up to 2150 m a.s.l. in Switzerland.
 Typhlocybinae Kirschbaum, 1868 151

Fischer (1972), Frommer (1996), Niedringhaus (1997), Reimer (1992), Remane & Reimer (1989), Schwoerbel
(1957), Trümbach (1959), Wagner (1935, 1939a), HN, and others; see also Cerutti (1939a), Günthart & Günthart
(1983)

Aguriahana pictilis (Stål, 1853)

In Latvia E VI (Vilbaste 1974), in Austria M IX (Wagner & Franz (1961), egg(?), 1 gen.(?)
This species is often listed for Germany, even in more recent literature, probably based
upon an old statement of Typhlocyba picta F. (= Eupteryx atropunctata Goeze) which was
copied by later authors. However, the only definite record has been gathered only very
recently in the southern Black Forest: Föhrenmoos near Ibach, 1000 m a.s.l., 18.VIII.2001,
10 ‡‡, in a shady spruce bog. All individuals were swept from Vaccinium myrtillus (which
is also reported to be the host plant in other European countries) growing under Betula
pubescens, but the densities were extremely low. Thus it is likely that these individuals
had fallen down from the tree canopies and that B. pubescens is the true host plant.
Moreover, most, if not all species of the genus are purely arboricolous. Otherwise the
species is also reported to occur in other parts of central Europe, but most localities
appear to be rather isolated.
Kirschbaum (1868), Sahlberg (1871), HN; see also Dworakowska (1972b), Metcalf (1968), Nast (1987), Os-
siannilsson (1981)

Aguriahana stellulata (Burmeister, 1841)

E VI – B X; egg, 1(?) gen.
Eurytopic in semi-open cultivated areas (river floodplains, parks, fruit-tree plantations,
forest margins, etc.), usually on Tilia cordata, Prunus avium (including fruit trees), P. spi-
nosa, Populus nigra, Betula pendula, Acer platanoides and A. campestre, also reported from
Aesculus hippocastanum in Switzerland. Widespread in the lowlands of Germany, but
usually in low to intermediate frequency and low individual numbers only, highest
localities are at 550 m a.s.l. in the Bavarian foreland of the Alps, but at 1200 m a.s.l. in
the Engadine (Switzerland).
Carl & Schönitzer (1995), Frommer (1996), Lehmann (1973a), Niedringhaus (1997), Olthoff (1986), Reimer
(1992), Schwoerbel (1957), Wagner (1935, 1939a), Nickel (1997), HN; see also Günthart (1987a)

Zyginella pulchra P. Löw, 1885

Adults I – XII, mainly M VIII – B VI; adult, 1 gen.
Mainly on Acer pseudoplatanus, less frequently on A. campestre, A. platanoides and A. mon-
spessulanum, also occasionally (but probably not breeding) on Quercus, Alnus and Cratae-
gus; overwintering individuals were found on Picea. Adults are usually swept along more
or less shady forest margins as well as in parks and on roadside trees in urban areas. In the
1930s the only known German localities were near Mainz, but during the past two de-
cades it was recorded from scattered sites over most of the middle parts of Germany
(roughly between the Danube and a line from Cologne to Marburg, Göttingen, Erfurt and
Dresden). Since 1985 some populations, which appear to be rather isolated, have also
152 Review of species

been reported from urban areas of Hamburg and Bremen. The species is frequently found
in the submontane and montane belt of the Mittelgebirge, up to at least 950 m a.s.l. in the
southern Black Forest and Bavarian Forest. It has been suggested that there was a recent
northeastward range expansion, with anthropogenic habitats being colonised first, but
the evidence is not convincing. Although there are many records from urban habitats,
most populations apparently live in forested areas away from human settlements. Fur-
thermore, the species is known from the Polish Carpathians since the beginning of the last
century. Increasing numbers of records may simply have been caused by increasing aware-
ness of the species in late summer, when collecting activity on trees is generally low.
Frommer (1996), Niedringhaus & Olthoff (1986, 1993), Wagner (1939a), Remane & Fröhlich (1994b), SMNS,
Nickel (1994, 1999b), HN; see also Dworakowska (1969), Nast (1976a)

Alnetoidia alneti (Dahlbom, 1850)

Some authors argued that this might be a complex of several species, particularly based on the
smaller body size, specific host association, and reduced yellowish pigmentation both in A.
alneti f. coryli (Tollin, 1851) and A. alneti f. mali (Edwards, 1915). After a detailed analysis of
morphometrics and colouration, Gillham & Claridge (1994) concluded that alder and hazel pop-
ulations formed the extremes of a considerable variation, but are clearly conspecific with the
nominate morph. They suggested that differences in morphology may be host-induced.
E V – M X; egg, 2 gen.
Polyphagous on deciduous trees and shrubs in forests, in semi-open cultivated areas,
and on solitary trees, preferentially in damp to wet sites, often abundant and in high
frequency on Alnus glutinosa and A. incana, common also on Acer, Carpinus, Prunus,
Corylus, Tilia, Cornus, Aesculus, in lower numbers on Quercus, Ulmus, Crataegus, Betula,
Salix and Fagus. Widespread and common in the lowlands of Germany, frequently found
up to 900 m a.s.l. in the Mittelgebirge and the foothills of the Alps, up to 1400 m a.s.l. in
the Engadine (Switzerland).
Achtziger (1991), Fischer (1972), Lehmann (1973a), Niedringhaus (1991, 1997), Reimer (1992), Remane (1987),
Schiemenz (1990), HN, and others; see also Claridge & Wilson (1976, 1981), Gillham & Claridge (1994),
Günthart (1987a)

Hauptidia distinguenda (Kirschbaum, 1868)

Adults I – XII, mainly B IX – M VI; adult, 1 gen.
Habitat requirements imperfectly known; usually found in xerothermic sites among
ruderal herb stands and along herbaceous margins, often in vineyard situations; Gera-
nium robertianum and G. pusillum are reported as food plants. Presumably described
from the surroundings of Mainz, although the holotype label is no longer readable.
From Germany known only from 6 localities (all below 300 m a.s.l.) along the upper and
middle Rhine and its tributaries: Ahr near Altenahr, Nahe near Schloßböckelheim and
Bad Münster am Stein, Neckar near Heilbronn (on Lagenaria siceraria), Rhine valley near
Bacharach, and Wachenheim an der Weinstraße.
Fröhlich & Nickel (unpublished data), Remane (1961b, and pers. comm), Schrameyer (pers. comm.), Wag-
ner (1939a); see also Cerutti (1939a), Günthart (1987a), Ossiannilsson (1981)
 Typhlocybinae Kirschbaum, 1868 153

Hauptidia provincialis (Ribaut, 1931)

So far VIII – I, ad.(?), 1(?) gen.
This is a Mediterranean species so far only known from southern France, Italy, Croatia,
Greece and Cyprus. From summer 2000 until spring 2002, a large, synanthropic popula-
tion caused feeding damage on cultivated, perennial Ocimum basilicum and Valerianella
locusta, in a greenhouse in southern Germany: Ulm, c. 500 m a.s.l. From southern coun-
tries, it is reported to feed on Parietaria, Urtica, Geranium and Solanum melangena and is
apparently polyphagous on a number of plants avoided by most other leafhoppers.
Coll. R. Dengler (det. HN); see also Dworakowska (1970d), Drosopoulos et al. (1986), della Giustina (1989)

Zyginidia franzi (W. Wagner, 1944)

According to Dworakowska (1970b) this species has already been described as Erythroneura
silvicola var. alpicola Cerutti, 1939. This interpretation was adopted by Nast (1972, 1987), della
Giustina (1989) and others, who used the name Z. alpicola (Cer.). However, this name was creat-
ed on the basis of a single †, the description of which included only two short notes on wing
colouration and shape of the anal tube appendages, which only slightly diverged from typical
Z. mocsaryi (Horv.). A type revision (P. Mazzoglio, pers. comm.) confirmed the conspecifity with
the latter and Cerutti’s correct identification.
So far E VII – B X, according to Wagner & Franz (1961) in Styria (Austria) also E VI;
egg(?), 1 gen.
Confined to the Alps and Carpathians; in Germany found only in 5 localities between
1250 and 1800 m a.s.l.: Ponten above Hindelang, Deutenhausen near Füssen, Frieder-
spitz above Garmisch-Partenkirchen, Sojernspitze above Mittenwald, and Königs-
bachalm above Berchtesgaden. Lives in open pine forests and alpine grassland, usually
on limestone, presumably on Sesleria spp. Has been found above 2400 m a.s.l. in the
Engadine (Switzerland).
Fischer (1972), Nickel & Voith (unpublished data); see also Günthart (1987a, 1997), Lauterer (1984), Wagner
& Franz (1961)

Zyginidia mocsaryi (Horváth, 1910)

B VI – E IX, occasionally until M XI; egg, 2(?) gen.
On Sesleria albicans in open pine and spruce forests, usually in warm situations on sun-
exposed calcareous or gypsum slopes, locally also on Festuca altissima in acidic and
rather cool habitats. Rather localized in Germany and at the northern edge of the range,
but abundant in favourable sites, with strongholds in the Bavarian Alps, the Franconian
Jura, and the Triassic limestone and gypsum regions of Thuringia. Localities at the edge
of the range include Bad Münstereifel, Höxter, Göttingen, Nordhausen, Jena and Dres-
den. Found at 900 m a.s.l. in the southern Black Forest and up to at least 1200 m a.s.l. in
the Allgäu Alps, although single individuals were recorded at 1600 m a.s.l.
Müller (1978), Rabeler (1962), Remane & Fröhlich (1994b), Rombach (1999b), Schiemenz (1990), Nickel
(1994), HN
154 Review of species

Zyginidia pullula (Boheman, 1845)

In Italy I – XII; ad, 4 gen. (A. Arzone pers. comm.).
Has been reported from Germany in older papers, but most cases are likely to refer to the
following species. On the other hand Z. pullula (Boh.) is reported to live in Denmark, the
Czech Republic and Italy. From Germany, however, there is only a single confirmed record
from eastern Bavaria: Oberpfälzer Jura, Naab valley near Kallmünz, 09.IX.1960, 1 †, which
is likely to have been a vagrant. From Italy this species is reported to live on various
grasses such as Elymus, Cynodon, Echinochloa, Hordeum, Sorghum, Triticum, Zea, Bromus,
Digitaria, Setaria, Molinia and others (locally causing feeding damage to cultivated maize
and other cereals), usually in disturbed sites such as fallow fields and waysides.
Remane & Fröhlich (1994b); also Lauterer (pers. comm.), Metcalf (1968), Ossiannilsson (1981), Vidano &
Arzone (1985)

Zyginidia scutellaris (Herrich-Schäffer, 1838) sensu Ribaut (1936)

It is uncertain whether Ribaut’s interpretation, which is now commonly accepted, refers to the
species described by Herrich-Schäffer. The holotype (a single ‡ from near Regensburg) is lost. At
present, resident populations of this species are apparently absent from this region, but it may
have occurred there during the 19th century, at least as a vagrant.
Adults I – XII, mainly M VIII – M VI; adult, 1(?) gen.
Mainly in ruderal sites, disturbed dry grassland, abandoned vineyards, etc., undertaking flights
in summer, and often found in large numbers in abandoned fields, fallows, winter cereals,
maize fields, fertilized meadows, and along waysides. Host plants are various grasses
(Festuca, Poa, Dactylis and others). The range includes the western Mediterranean region as
well as France, Germany, Belgium, the Netherlands and England. In Germany not mentioned
in older papers, and apparently present only since the 1950s (common in the Kaiserstuhl area
in 1953, in middle Hessen since the 1960s). At present, permanent populations (which may
show extremely high abundances) are known mainly from the southwestern half (Lake Con-
stance basin, upper and middle Rhine, valleys of the Neckar, Main and Regnitz, Eifel Moun-
tains, middle Hessen, up to at least 400 m a.s.l.), and very recently also in the Saale valley
between Rudolstadt and Naumburg. In the latter localities, the species was definitely not
present during the 1960s and 1970s. Furthermore, there are records of single individuals, which
are probably vagrants, from the Kyffhäuser, the Thuringian Forest, the Altmühl and Neckar
valley, and the Bavarian Alps and their foreland. Is reported to cause feeding damage on
maize in France. In southwestern Germany, damage has been observed mainly on sweet corn;
the economic importance, however, has not been quantified, and is perhaps negligible.
Bornholdt & Remane (1993), Frommer (pers. comm.), Günthart (1988), Post-Plangg & Hoffmann (1982),
Reimer (1992), Remane (pers. comm.), Rombach (1999b), Schiemenz (1990), Schrameyer (pers. comm.),
Schwoerbel (1957), Walter (pers. comm.), ZIMH, HN; see also Della Giustina (1989), Kirschbaum (1868),
Marion-Poll et al. (1987), Remane (1987), Wagner (1939a, 1951a), Wonn (1956)

Zyginidia viaduensis (W. Wagner, 1941)

Has recently been treated as conspecific with Z. cornicula Meusnier, 1982, which lives in the
subalpine belt of the western Alps and the Pyrenees (Mazzoglio 1989). This opinion is chal-
 Typhlocybinae Kirschbaum, 1868 155

lenged here. There are clear differences in the morphology of the aedeagus and the anal tube
appendages, as well as in life history and habitat choice.
So far B VI – B X; egg(?), 2(?) gen.
Known only from a few localities in the northeastern German plain, Poland, the Baltic
states and middle Russia, living in sandy xerothermic sites on Koeleria glauca. In Germa-
ny there are 7 known localities: Boizenburg an der Elbe, Gerwisch near Magdeburg,
Groß Machnow near Zossen, Niederlehme near Königs Wusterhausen, Oderberg near
Eberswalde, Mescherin (locus typicus) and Gartz, both near Angermünde.
Schiemenz (1990), Wagner (1941a, 1944), HN; for identification see Wagner (1941a)

Zygina nivea (Mulsant & Rey, 1855)

So far B VIII; adult(?), 1(?) gen.
On Populus alba, in France also reported from P. nigra, Salix eleagnos and S. alba. From
Germany there are only 4 records from the upper and middle Rhine between 50 and
220 m a.s.l.: Burkheim, 02.VIII.1997, Buggingen, 06.VIII.1997, small populations on P.
alba in river floodplains, as well as in urban areas of Cologne, two records between 1993
and 1995. Presumably, this species is more widespread in western parts of Germany. On
the other hand older reports either refer to Edwardsiana candidula (Kbm.) or originate
from areas outside present-day Germany.
Frommer (pers. comm.), HN; see also Ribaut (1936)

Zygina tithide Ferrari, 1882

So far only M IX; in the Valais (Switzerland) according to Cerutti (1939a) hibernation as
adults, 1(?) gen.
After a revision a single † collected near Jena (Thuringia, Leuchtenburg near Kahla,
400 m a.s.l., 17.IX. 1967), which had formerly been published as Z. nivea (M. & R.), turned
out to belong to Z. tithide Ferr. However, due to the absence of the host plant at the
collecting locality, a permanent occurrence is not proven. The nearest known localities
are in Poland (near Poznan), Bohemia (Czech Republic), eastern Austria and the Valais
(Switzerland). From the latter region this species is reported to live on Populus alba and
other deciduous trees and shrubs, migrating to coniferous trees, Juniperus, and Rubus
fruticosus for hibernation.
Coll. Müller, Schiemenz (1990); see also Cerutti (1939a), Dworakowska (1977b), Giordano et al. (2001), Nast
(1976a), Remane & Fröhlich (1994b)

Zygina lunaris (Mulsant et Rey, 1855)

In Strasbourg (France) VII – III; adult, 1 gen. (Lauterer & Malenovsky 1995).
This species was recently found in parks in the upper Rhine floodplains in Kehl and
Weil am Rhein and adjacent suburbs (Baden, c. 140 and 260 m a.s.l.), as well in Stras-
bourg (Alsace, France) and Basel (Switzerland), where it is rather common on Salix alba,
S. fragilis, S. babylonica and S. purpurea, migrating to Acer and Rosa in October and to
156 Review of species

various coniferous trees for hibernation. Otherwise it is only known to occur in the
western Mediterranean region.
Lauterer & Malenovsky (1995), Mühlethaler (2001, and pers. comm.), HN

Zygina angusta Lethierry, 1874

Populations living on Rosaceae differ slightly in colouration from those on Fagaceae and may
belong to a distinct species (Remane 1994). Moreover, since the description of Z. nigritarsis Rem.,
most published records are in need of revision.
E VII – B VI; adult, 1 gen.
In semi-open stands of shrubs and trees, usually in dry to damp sites; on the one hand
on hedges and shrubs such as Crataegus, Rosa and Prunus spinosa, on the other hand
along forest margins on Quercus robur, in lower numbers also on Qu. petraea, Fagus,
Carpinus, Prunus padus and others; found on coniferous trees (Picea, Pinus, Juniperus)
during hibernation, on Ribes in spring. The species is poorly documented, but apparent-
ly widespread in Germany between the foothills of the Alps and the coast, common at
least in middle and southern parts; the highest localities are at 700 m a.s.l.
Achtziger (1995a), Frommer (1996), Niedringhaus (1997), Reimer (1992), Remane (1987, and pers. comm.),
Nickel (1997), HN; for identification see Remane (1994)

Zygina flammigera (Geoffroy, 1785)

M VII – B VI; adult, 1 gen.
On various woody plants, in summer mainly on Rosaceae; the main hosts are several
species of Prunus, Malus, Crataegus, Sorbus, and perhaps Rosa. Also recorded on Fagus,
Quercus and others, usually on Picea in winter, occasionally on Pinus, Buxus, Juniperus,
Rubus and Hedera. Widespread in the lowlands of Germany and common, found up to
950 m a.s.l. in the Mittelgebirge and the Bavarian Alps, at 1280 m a.s.l. in the Engadine
(Switzerland), at 1400 m a.s.l. in the Trentino (northern Italy).
Achtziger (1995a), Frommer (1996), Niedringhaus (1991, 1997), Post-Plangg & Hoffmann (1982), Reimer
(1992), Remane (1987, 1994), Schiemenz (1990), HN; see also Günthart (1987a)

Zygina ordinaria (Ribaut, 1936)

This taxon is often confused with Z. tiliae (Fall.) and other species of Zygina. According to Re-
mane (1994) it may comprise a complex of several distinct species.
Adults I – XII, mainly M VII – E V; adult, 1 gen.
On narrow-leaved species of willows, often near running water, as well as in heath and
bog habitats, usually on willows, notably Salix alba, S. triandra, S. viminalis, S. fragilis, S.
purpurea, S. repens and S. pentandra, occasionally also on S. eleagnos and additional spe-
cies. The overwintering sites are unknown. Widespread in Germany, but poorly docu-
mented; usually in intermediate frequency, but low individual numbers on the hosts,
found up to at least 950 m a.s.l. in the Bavarian Alps.
Frommer (pers. comm.), Reimer (1992), Remane (pers. comm.), Nickel (1997, 1999b); for identification see
Remane (1994), Günthart (1974)
 Typhlocybinae Kirschbaum, 1868 157

Zygina rosea (Flor, 1861)

According to Remane & Fröhlich (1994b) the name Z. roseipennis (Tollin, 1851), which is used by
many authors for this taxon (e.g. Nast 1972, 1987), is doubtful and should be rejected. Further-
more, literature records from central and southern Europe probably refer to other species and
should be revised.
So far B VIII – M II, in Sweden according to Ossiannilsson (1981) until B VI; adult, 1 gen.
In raised and intermediate bogs, and along their margins. Nymphal development and
most adults on Betula pubescens, hibernation takes place on conifers, mainly Pinus sylvestris
(although there is one record from Pseudotsuga douglasi). In Germany largely restricted
to the remnant bog areas of the northern plains, recorded near Meppen, Lingen (Ems),
Oldenburg, Hamburg, Hanover and Ratzeburg (probably overlooked in Mecklenburg
and Brandenburg), also found near Fulda and Munich (550 m a.s.l.).
Niedringhaus (1997), Remane (1962), Remane & Fröhlich (1994b), Wagner (1941b), Weis (pers. comm.); see
also Linnavuori (1952), Nuorteva (1952a)

Zygina rubrovittata (Lethierry, 1869)

Adults I – XII; adult, 2(?) gen.
In sunny, usually dry and sandy sites (mainly heaths, edges of pine forests, drained
raised bogs) on Calluna vulgaris, which is also the winter food plant. In Germany locally
common in the northern plains, but rather scattered and restricted to regions of sand
and sandstone in middle parts (middle Hessen, Thuringian Forest, Erzgebirge, Pfälzer
Wald, northern upper Rhine plain, Mittelfranken, Danube valley); apparently absent
from the Alps, their foothills and foreland. The highest localities are at 450 m a.s.l.
Frommer (pers. comm.), Remane (pers. comm.), Schiemenz (1990), Struve (1939), Trümbach (1959), Wagner
(1935), HN; see also Badmin (1995)

Zygina suavis Rey, 1891 sensu Ossiannilsson (1981)

This is a rather critical taxon, the identity and distinction of which are subject to diverging
opinions in the literature (see Vidano 1959b, 1961). Perhaps there is another distinct species in
central Europe, showing a very similar colouration (see Remane 1994), but living on deciduous
trees (probably Quercus or Tilia). However, until further studies have been done, Vidano’s opin-
ion should be followed, treating Zygina rhamnicola Horváth, 1903 and Erythroneura inconstans
Ribaut, 1936 as junior synonyms of Z. suavis R. Another problem is caused by the fact that Z.
schneideri Günth., which is apparently widespread in Europe, was described only in 1974. Thus
most data published before that are in need of revision.
M VII – E V; adult, 1(?) gen., according to Günthart (1974) in Switzerland 2 gen.
In moist to dry, basic as well as acidic sites, mainly open forests, pastures with scattered
shrubs, and various margins (notably hedges, forest and bog edges), preferentially on
Frangula alnus, at least in low numbers also on Rhamnus cathartica, on Picea abies during
hibernation. In Germany only poorly documented and often confused, but widespread
in the lowlands, where occurring in intermediate frequency at least on Frangula alnus.
Northernmost localities are near Papenburg, Kiel and Königs Wusterhausen, but the
species is also reported from southern parts of Sweden and Finland and from north-
158 Review of species

western Poland. Found up to at least 600 m a.s.l. in the German Mittelgebirge, up to

1200 m a.s.l. in the Engadine (Switzerland).
Niedringhaus (1997), Reimer (1992), Remane (1987, and pers. comm.), Schiemenz (1990), Nickel (1994), HN;
see also Günthart (1974, 1987a), Nast (1976a), Ossiannilsson (1981); for identification see Remane (1994)

Zygina schneideri (Günthart, 1974)

See taxonomic notes on Z. suavis R.
Adults I – XII, mainly M VII – B VI; adult, 1(?) gen.
On woody species of Rosaceae in rather warm sites, mainly on sunny hillsides and
along hedges on Prunus spinosa, Rosa canina and R. rubiginosa, as well as in urban settle-
ments on Rosa rugosa, Potentilla fruticosa, Amelanchier, Cotoneaster, and other ornamental
shrubs; hibernation probably takes place in the litter layer. Widespread in middle parts
of Germany, between the Danube and a line from Cologne to Braunschweig, Halle and
Frankfurt an der Oder, particularly common and in high frequency in warm regions,
found up to 900 m a.s.l. on the Swabian Jura. Is also reported from England, southern
Sweden, southern Norway.
Achtziger (1995a), Bornholdt & Remane (1993), Frommer (pers. comm.), Post-Plangg & Hoffmann (1982),
Reimer (1992), Remane (1987, 1994), Remane & Fröhlich (1994b), Rombach (1999a), Nickel (1994), HN; see
also Le Quesne & Payne (1981), Ossiannilsson (1981); for identification see Günthart (1974), Remane (1994)

Zygina tiliae (Fallén, 1806)

Adults I – XII, mainly E VII – B VI; adult, 1(?) gen.
In open woodland and along forest margins, in wet to damp, occasionally also rather dry
sites. The main host is Alnus glutinosa. Lower numbers are also swept from Alnus incana,
Tilia cordata (where breeding has been found to take place at least locally) and other woody
species (notably Prunus padus, Salix and Betula); at least occasionally found on Picea abies
in winter. Widespread and locally common in Germany, at least to the north of the Danube,
although poorly documented due to identification problems. Recorded below 450 m a.s.l.
only, but reported from 1190 m a.s.l. in the Engadine (Switzerland).
Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987, and pers. comm.),
Nickel & Remane (1996), HN; see also Günthart (1987a); for identification see Remane (1994)

Zygina rosincola (Cerutti, 1939)

Mainly E VII – V; adult, 1(?) gen.
Usually in rather warm and sun-exposed sites on Rosa (from which nymphal develop-
ment is reported), Prunus spinosa and Crataegus; adults occasionally also on Quercus and
Betula; in winter found on coniferous trees (Pinus, Picea) and Juniperus. The distribution
in Germany is poorly documented due to difficulties in determination. Apparently wide-
spread, but uncommon at least in the western half (Bavaria, Hessen, southern Nieder-
sachsen, Westphalia) between the Danube and the northern edge of the Mittelgebirge; not
recorded from eastern parts (from Mecklenburg-Vorpommern to Saxony and Thuring-
 Typhlocybinae Kirschbaum, 1868 159

ia), where probably overlooked. The highest localities are at only 500 m a.s.l. From Swit-
zerland this species is reported from Rosa glauca, R. pendulina, R. montana, Sorbus aucu-
paria, Prunus padus and Malus, in altitudes up to 1520 m a.s.l.
Remane & Fröhlich (1994b), Nickel & Remane (1996), Nickel (1994), HN; see also Günthart (1987b, and
pers. comm.); for identification see Günthart (1974), Remane (1994)

Zygina griseombra Remane, 1994

B VII – B V; adult, 1 gen.
Nymphal development takes place on Carpinus betulus, usually along margins of forests
and in their interior; after emergence adults migrate to other woody plants and are
recorded on coniferous trees during winter, frequently on Crataegus in more open sites
in spring. In Germany the species is only known from the region between the Main and
the northern edge of the Mittelgebirge, where it occurs in rather low frequency and indi-
vidual numbers (although probably under-recorded). Northernmost localities are near
Altenahr, Marburg (locus typicus: Elnhausen), Göttingen, Bad Frankenhausen and Dres-
den, at least up to 500 m a.s.l. Outside Germany it has been recorded so far only from
northern France, Moravia (Czech Republic), Styria (Austria), the canton of Zurich (Swit-
zerland) and southern Italy.
Remane (1994, and pers. comm.), Nickel & Remane (1996), Nickel (1994), HN; see also Günthart (2000),
Holzinger & Remane (1994); see also della Giustina & Remane (2001), Lauterer (2000), for identification see
Remane (1994)

Zygina nigritarsis Remane, 1994

This species has been described only recently and is perhaps widespread in the European decid-
uous forest zone. Thus, a revision of older material labelled as Z. angusta Leth. or Z. tiliae (Fall.)
is likely to reveal more records.
M VIII – B VI; adult, 1 gen.
Nymphs on Prunus padus, mainly in floodplains of rivers and streams, adults migrating to
coniferous trees (Picea, Pinus) for hibernation; found on Crataegus, Ribes and other woody
plants in spring (also in parks, gardens, dry grassland and other habitats). The distribution
in Germany is only poorly documented, but the species is widespread at least between the
foreland of the Alps and the northern edge of the Mittelgebirge, although apparently scat-
tered and in low frequency; northernmost localities are near Oldenburg, Hamburg, Plön,
Bad Frankenhausen and Bitterfeld, recorded up to 550 m a.s.l. Outside Germany only known
from North Tyrol (Inn valley near Mils, Austria), but probably more widespread.
Frommer (pers. comm.), Neumann (1997), Remane (1994), ZIMH, ZIMH, HN; for identification see Re-
mane (1994)

Zygina hyperici (Herrich-Schäffer, 1836)

M V – M X; egg, 2 gen.
Monophagous on Hypericum perforatum in sunny, damp to dry, usually disturbed sites (rud-
eral habitats, waysides, railway embankments, mining areas, grazed or abandoned grass-
160 Review of species

land, etc.). Widespread in the lowlands of Germany, in intermediate to high frequency on

the host, but rather scattered in the northwestern plain and to the south of the Danube,
usually found below 500 m a.s.l., locally up to 1000 m a.s.l. in the southern Black Forest.
Achtziger (1991), Frommer (pers. comm.), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987),
Schiemenz (1990), Wagner (1935, 1939a), HN

Zygina hypermaculata Remane & Holzinger, 1995

So far E VII – E IX; egg, 1 gen.
This species has only recently been described from Upper Bavaria and Styria (Austria).
It is known only from the montane and subalpine belt (between 1300 and 1900 m a.s.l.)
of the northeastern Alps, living on Hypericum maculatum. In this region, however, it
appears to be widespread and fairly common. The locus typicus is Brauneck above Bad
Tölz (1500 m a.s.l., 07.VIII.1961), the paratypes are from three sites in Styria. Further
Austrian records are from Carinthia and Upper Austria. In Germany the species has
been found in the Berchtesgaden and Karwendel Alps (Roßfeld, Schneibstein and Rei-
teralm, all above Berchtesgaden, Sojernspitze near Mittenwald), in herb stands on sun-
ny hillsides as well as in open montane coniferous forests.
Remane & Holzinger (1995), HN; see also Holzinger (1999b)

Arboridia kratochvili (Lang, 1945)

So far M VI – B X; egg(?), 2(?) gen.
Until recently this species was only known from a few localities in southern Poland, the
Czech Republic and Slovakia. Since 1993 it has been recorded from altogether 6 sites on
Triassic limestone plateaus in the Main-Saale region of Franconia (Thüngersheim, Gam-
bach, Wiesenfeld, Aschfeld, Gössenheim and Machtilshausen, all c. 300 m a.s.l.). In these
sites it is locally abundant on Potentilla tabernaemontani in xerothermic grassland with
low-growing and incomplete cover of vegetation (Trinio-Caricetum humilis).
Nickel & Remane (1996), HN; see also Dlabola (1977), Lauterer (pers. comm.), Nast (1976a); for identifica-
tion see Lang (1945)

Arboridia erecta (Ribaut, 1931)

So far B VIII – B VI, adult, 1(?) gen.
On deciduous trees and shrubs in warm situations; reported to live on Acer campestre
and occasionally Castanea sativa in Italy, also on Prunus, Quercus, Corylus and Tilia in
Switzerland and France. In Germany the species lives at the northern edge of its range
and is only known from 4 records in Baden-Württemberg and Bavaria: Markgröningen
near Stuttgart, 230 m a.s.l., 17.IX.1980, 1 †, at light; Schwörstadt near Lörrach, 290 m
a.s.l., 05.IV.1987, in forest litter, and 02.VI. 1987, on Malus and Prunus avium, altogether
7 ††, 5 ‡‡; Freiburg, municipal cemetery, 250 m a.s.l., 04.VIII.1997, 4 ††, 2 ‡‡, on Acer
platanoides; Schönhofen near Regensburg, 450 m a.s.l., 16.IX.1997. Recorded at 1530 m
a.s.l. in the Engadine (Switzerland).
 Typhlocybinae Kirschbaum, 1868 161

Günthart (1988, and pers. comm.), Heller (1996), Remane (pers. comm.), HN; see also Günthart (1987b),
Ribaut (1936), Vidano & Arzone (1987b)

Arboridia parvula (Boheman, 1845)

A. potentillae (Moravskaja, 1948) was described from Potentilla arenaria (= P. incana) in southern
Russia. Apparently, it can be distinguished from A. parvula (Boh.) only by the narrowing distal half
of the aedeagus stem. Specimens found on Potentilla incana in several xerothermic localities in
eastern parts of Germany may belong to this taxon. Provisionally, however, they shall be treated as
A. parvula (Boh.), but the problem requires further studies of morphology and ecology.
Adults I – XII, mainly M VII – B VI; adult, 1(?) gen.
Usually along woody margins in dry to damp (occasionally also moist) sites, often in
rather warm situations, notably dry grassland with scattered shrubs and trees, along
hedges and forest margins. Adults on herbaceous and shrubby species of Rosaceae (Po-
tentilla incana, Filipendula ulmaria, Rubus idaeus and others), in lower numbers also on
various deciduous trees (Quercus, Prunus padus and others); most winter records are
from herbaceous vegetation. Widespread in Germany and locally common, particularly
in warmer regions, but rather sporadic and uncommon in most parts of the Mittelgebirge
and the north German plain, not reported from Mecklenburg-Vorpommern. The high-
est localities are at 1400 m a.s.l. in the Bavarian Alps, but the species is reported to occur
frequently in the subalpine belt up to 1800 m a.s.l. in Switzerland.
Niedringhaus & Olthoff (1993), Post-Plangg & Hoffmann (1982), Reimer (1992), Schwoerbel (1957), Schie-
menz (1990), Nickel & Remane (1996), HN; see also Cerutti (1939a), Günthart (1987a)

Arboridia ribauti (Ossiannilsson, 1937)

M VII – B VI; adult, 1 gen.
On deciduous trees (mainly Carpinus betulus, but also Tilia cordata and Quercus), usually
along forest margins and roads, sometimes also along hedges. Winter specimens have
been found on Calluna vulgaris and Hedera helix. In Germany recorded only between the
edge of the Alps and a line from Düsseldorf to Göttingen, Halle and Dresden, where
usually occurring in intermediate to low frequency and rather low numbers. Highest
localities are at 750 m a.s.l. in the Black Forest, but at 1400 m a.s.l. in the Trentino (north-
ern Italy) and at 1300 m a.s.l. in the Valais (Switzerland).
Frommer (1996), Kolbe & Bruns (pers. comm.), Reimer (1992), Schiemenz (1990), Wagner (1951a), HN; see
also Cerutti (1939a); for identification see Wagner (1939a)

Arboridia simillima (W. Wagner, 1939)

Recently A. loginovae (Emelyanov, 1964), known from the Altai Mountains, Siberia and Kazakh-
stan, has been reported from Moravia (Czech Republic) by Lauterer (2000). Morphology and
ecology are very similar as in A. simillima (W.Wg.). Southwest German populations of the latter
have been found to show a considerable morphological variation in the shape of the aedeagus.
Thus, the distinction of these two taxa is in need of clarification.
Adults I – XII, mainly M VII – M V; adult, 1(?) gen.
162 Review of species

On sun-exposed shrubs and in open xerothermic woodland, mainly in slope and pla-
teau situations on limestone, gypsum and porphyry. Host plants are Rosa spinosissima
and additional roses (probably R. rubiginosa, perhaps R. canina). In Germany largely
confined to the southern half, with strongholds in viticultural regions of the upper Rhine
valley (locus typicus: Wöllstein), Mainfranken and Saale-Unstrut region (including Kyff-
häuser, Hainleite, and valleys of Unstrut, Saale and Ilm). Further localities are situat-
ed in the Eifel Mountains and along the upper course of the Isar (Vorderriß, c. 1000 m
a.s.l.). The species is probably under-recorded.
Post-Plangg & Hoffmann (1982), Schiemenz (1990), Wagner (1939a), Nickel & Remane (1996), HN; for
identification see Wagner (1939a)

Arboridia velata (Ribaut, 1952)

E VII – M V; adult, 1 gen.
Along sunny margins of oak forests, usually on rather warm upper slopes. At present
known from Germany only from the Lake Constance Basin, the valleys of the Rhine
(between Waldshut and Koblenz), Nahe and Neckar, the slopes along the Danube be-
tween Kelheim and Passau, the Kyffhäuser, the upper course of the Saale near Bad
Blankenburg, and the Niederlausitz. The majority of individuals has been swept from
oaks (Quercus petraea and Qu. robur, perhaps also Qu. pubescens), a few – probably va-
grants – also from neighbouring woody species. Highest localities are at c. 500 m a.s.l.,
but at 1200 m a.s.l. in the Valais (Switzerland).
Günthart (1988), Schiemenz (1990), SMNS, Nickel (1999b), HN; see also Cerutti (1939a), Ribaut (1931, 1936),
Vidano & Arzone (1987b)

Arboridia spathulata (Ribaut, 1931)

According to Dworakowska (1970c) this is a junior synonym of A. versuta (Melichar, 1897). How-
ever, the latter was described after a ‡, and most ‡‡ in this genus cannot be identified with
certainty. In contrast, Ribaut’s description contains exact and unambiguous drawings of † gen-
italia. Hence, the identity of A. versuta (Mel.) is considered here as dubious.
So far only E X; adult(?), 1(?) gen.
In central Europe this species has only rarely been found. The only German record is
from Franconia: Bad Windsheim, c. 320 m a.s.l., 20.X.1937. From Italy, it is reported to
live monophagously on Quercus petraea, Qu. robur, Qu. pubescens and Qu. cerris. Has
been found at 1250 m a.s.l. in the Valais (Switzerland).
Wagner (1939a); see also Cerutti (1939a), Vidano & Arzone (1987b)

Arboridia pusilla (Ribaut, 1936)

Adults I – XII, mainly B VII – M VI, adult, 2(?) gen.
On Geranium sanguineum, along woody and herbaceous margins in rather warm situa-
tions, occasionally also in open pine and oak forests and in abandoned vineyards. Often
syntopic with A. simillima (W.Wg.), but in different layers of the vegetation. In Germany
 Deltocephalinae Fieber, 1869 163

the species is restricted to warm regions of the southern half, at the northern edge of the
range. So far, it was found in the Saale-Unstrut region (Kyffhäuser, valleys of Unstrut,
Saale and Gera), Elbe valley near Dresden, Eifel Mountains (near Prüm), Nahe valley,
Rheinhessen, Mainfranken (Main valley between Karlstadt and Haßfurt, Saale valley
between Hammelburg and Bad Königshofen), Swabian and Franconian Jura (near Tut-
tlingen and Regensburg) and Kaiserstuhl. The highest localities are at 900 m a.s.l.
Bornholdt & Remane (1993), Krause & Emmrich (1996), Remane (1987), Schiemenz (1990), Wagner (1939a),
Nickel & Remane (1996), HN

Fruticidia bisignata (Mulsant et Rey, 1855)

E VIII – B VII; adult, 1 gen.
The range of this species essentially includes the Mediterranean region, but extends
northeastward to the upper and middle parts of the Rhine valley and its tributaries.
Border localities are near Cologne, Koblenz, Bad Nauheim, Lohr am Main, Heidelberg
and Freiburg. Lives in various types of dry grassland with scattered shrubs or trees and
along sunny forest margins, usually on woody species of Rosaceae (mainly Crataegus,
but also Malus, Amelanchier and Quercus); winter specimens have been found on Calluna
and coniferous trees. The highest localities are at c. 300 m a.s.l. only.
Frommer (pers. comm.), Günthart (1988), Post-Plangg & Hoffmann (1982), Remane (pers. comm.), SMNS,
Wagner (1939a), Nickel (1999b), HN

Fruticidia sanguinosa (Rey, 1891)

A single †, which may have been a vagrant, was found near Heidelberg (Sandhausen),
27.IX.1982, on Crataegus, along the margin of a xerothermic inland dune. Otherwise,
this species is poorly known and only reported to occur in Mediterranean regions of
France and Italy, feeding on Crataegus, Rosa and Prunus, usually syntopically with the
preceding one.
Heller (1996); see also D’Urso (1995), Ribaut (1936)

4.2.4.13 Deltocephalinae Fieber, 1869

Fieberiella florii (Stål, 1864)

M VII – M X, single individuals B VI; egg, 1 gen.
A polyphagous species living on woody plants in open shrubland and along various
margins, mainly in dry grassland with scattered shrubs, but also in ruderal sites, along
rivers, streams and hedges, in parks and gardens. Host plants are usually Rosa, Prunus
spinosa, Ligustrum vulgare and Rubus idaeus (found only synanthropic on the two latter
species), at least single adults also on Quercus, Ulmus, Cytisus, Salix, Syringa, Elaeagnus
and others. In Germany at the northern edge of the range, with strongholds in the warmer
regions (mainly the basins of the Rhine, Main and Neckar). Northernmost records are
164 Review of species

from near Cologne, Bad Nauheim, Schlüchtern, Sondershausen, Berlin and Frankfurt
an der Oder, up to at least 400 m a.s.l., but reported from 880 m a.s.l. in Switzerland.
Cobben (1956), Frommer (pers. comm.), Meyer-Arndt & Remane (1992), Schiemenz et al. (1996), Wagner
(1963), Nickel & Remane (1996), HN; see also Günthart (1987b)

Fieberiella septentrionalis W. Wagner, 1963

B VII – E X (in southwestern Germany from E V onwards); egg, 1 gen.
On Prunus spinosa and Rosa species in dry grassland, ruderal sites, along forest margins
and hedges, but also synanthropic in gardens and parks, where usually found on Rosa
rugosa, Ligustrum vulgare, Rubus idaeus, R. fruticosus, Rhododendron, Spiraea, Cotoneaster,
Solidago canadensis and S. gigantea. Generally in similar sites as the preceding species
and syntopical in some places, but with a different distribution pattern. A tongue-shaped
part of the range extends into central parts of Germany from the southeast. According
to existing data, border localities include Regensburg, Ansbach, Landau (Pfalz), Bad
Kreuznach, Koblenz, Cologne, Minden, Quedlinburg, Halle and Eberswalde. Further-
more, the species occurs locally in urban settlements near the coast and in the foreland
of the Alps (Bremen, Hamburg, Rostock, Munich), where it is perhaps extending its
range. The highest localities are at c. 550 m a.s.l., the locus typicus is Jena.
Frommer (1996), Jöst (1966), Meyer-Arndt & Remane (1992), Niedringhaus & Olthoff (1986, 1993), Schie-
menz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1963), HN

Grypotes puncticollis (Herrich-Schäffer, 1834)

E VI – M X, according to Schiemenz et al. (1996) also M V; egg, 1 gen.
On pines in forests and along their edges as well as on solitary trees, in Germany re-
corded on Pinus sylvestris only, in Styria (Austria) also on P. nigra. Single individuals on
dispersal flight are frequently swept from herbaceous vegetation of various habitats
(meadows, dry grassland, etc.). Widespread in Germany, in high frequency and at inter-
mediate to high densities on the host, found up to at least 900 m a.s.l. in the Mittelgebirge
and the Bavarian Alps; recorded at 1400 m a.s.l. in Switzerland.
Fischer (1972), Niedringhaus (1991), Remane (1987), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b),
Schwoerbel (1957), Wagner (1935, 1939a), HN; see also Günthart (1987b), Holzinger (pers. comm.)

Goniagnathus brevis (Herrich-Schäffer, 1835)

B VIII – E VI, occasionally also VII; adult, 1 gen.
In sunny, moderately dry to dry and sparsely vegetated sites on various substrates (lime-
stone, gypsum, porphyry, diabas, sandstone, sand, etc.). In several occasions, Thymus
pulegioides and Th. praecox were recorded as food plant. In Germany confined to the
southern half and at the edge of the range; northernmost localities are on a line from
Koblenz to Marburg, Göttingen, Quedlinburg, Halle, Leipzig and Bautzen. Found up to
800 m a.s.l on the Swabian Jura and in the foothills of the Alps.
 Deltocephalinae Fieber, 1869 165

Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (pers. comm.), Schiemenz et al. (1996), Schönitzer
& Oesterling (1998b), Wagner (1939, 1951), Nickel (1994), HN

Japananus hyalinus (Osborn, 1900)

Japananus meridionalis Bonfils, 1981
B VIII – M X; egg, 1 gen.
Described from North America, where it is probably introduced; the main range in-
cludes the deciduous forest zones of eastern Asia and Europe. Several authors have
suggested that it was also introduced to Europe, based on the fact that it has been found
on ornamental trees (notably Acer japonicum and A. palmatum, both originating from
Japan) and on the late discovery, which happened only in 1942 in Styria (Austria). Since
then it was also found in Catalonia (Spain), southern France, Switzerland, southwest
Germany, northern Italy, the Czech Republic, Slovakia, Hungary, Serbia, Romania, Bul-
garia and southern Russia. However, many, if not most European records are from na-
tive maples in habitats which suffered little anthropogenic influence, although this is
only partially true for the German localities, most of which are in urban areas: Freiburg
(Günterstal, 2001, on Acer campestre), Stuttgart (Hofen, 1985, on A. campestre, and Rosen-
steinpark, 1985, on A. pseudoplatanus), Neckarsulm, Heilbronn (both 2001, on
A. platanoides, A. palmatum and A. campestre), Munich (2001, on ornamental Acer) and
Cologne (1994), up to 550 m a.s.l. The species is monophagous on maple; Acer campestre
is apparently the main host in Europe.
Billen & Schrameyer (pers. comm., det. Achtziger), Frommer (1996), Heller (1987b), Weis (pers. comm.); see
also Arno et al. (1988), Arzone et al. (1987), della Giustina (1989), Lauterer (1980, 1984, 1989), Mühlethaler
(2001), Remane & Fröhlich (1994b)

Opsius stactogalus Fieber, 1866

E VI – B X; egg, 1 gen.
In central Europe this species was originally restricted to gravel banks along rivers in the
Alps and their foothills. At least since the middle of the 19th century it has been reported
from Tamarix species in gardens and parks of the lowlands and is now widespread and
common all over Germany (for instance in northern Germany recorded from the East Fri-
sian Islands, Kiel and Rostock). Since the beginning of the 20th century it is also known from
North America, where it was probably introduced. From natural habitats, it is at present
only known from the Isar between Mittenwald and Wolfratshausen (formerly also along
the Saalach, Lech and upper Rhine), where it is locally common on Myricaria germanica.
Fischer (1972), Frommer (1996), Kirschbaum (1868), Kuntze (1937), Lauterborn (1923), Niedringhaus (1991),
Remane (pers. comm.), Schönitzer & Oesterling (1998b), Wagner (1939a), Nickel (1999a), HN; see also Hamil-
ton (1983a)

Neoaliturus fenestratus (Herrich-Schäffer, 1834) group

The taxonomy of this group is unclear and in need of a thorough revision. In Germany alone
there may be two, or perhaps even three morphologically and ecologically distinct forms.
166 Review of species

Adults I – XII, mainly E VI – VIII and B IX – V; adult, 2 gen., probably 1 at higher altitudes.
In sunny, moderately dry to dry, sparsely vegetated, usually disturbed sites on various
substrates (basic to acidic, gravelly to loamy), mainly grazed low-productivity grass-
land, inland dunes, rocky slopes, abandoned fields and vineyards, waysides and min-
ing areas. The main hosts are various species of Asteraceae, notably Leontodon spp.,
perhaps also additional dicotyledonous herbs. In Germany the species is widespread at
least between the Danube and a line from Koblenz to Marburg, Göttingen, Stendal and
Neubrandenburg; and there is a record of a single † from Hamburg. It is locally com-
mon in rather warm situations mainly below c. 400 m a.s.l., but uncommon in western
parts of the north German plain as well as in the Bavarian and Allgäu Alps, where
frequently recorded up to 1800 m a.s.l. Also reported from alpine habitats of Austria
and Italy, but apparently absent from Denmark, Norway and Sweden. From Israel it is
reported as a vector of Safflower phyllody in fields of Carthamus tinctorius, although it is
uncertain whether these populations are conspecific with the central European ones.
Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Schwoerbel (1957),
Wagner (1951a), Nickel (1994), HN; see also Klein (1992), Leising (1977), Ossiannilsson (1983)

Circulifer opacipennis (Lethierry, 1876) sensu Young & Frazier (1954)

Belongs to a taxonomically critical species group, which lives mainly in rather warm and dry
regions of the Old and New World. Some taxa are suspected to transmit various virus and MLO
diseases (e.g. Klein 1992). The identity of populations found in Germany is not quite clear. Ac-
cording to Nast (1972, 1987) Circulifer is a junior synonym of Neoaliturus, and Cicadula opacipen-
nis Leth. and Jassus haagii Kirschbaum, 1868 (locus typicus near Mainz, described after 2 ‡‡) are
junior synonyms of C. haematoceps (Mulsant & Rey, 1855). According to Young & Frazier (1954)
and Oman (1970), however, Circulifer is a distinct genus and C. opacipennis (Leth.) a distinct
species. This interpretation is followed here until further research has been done.
This taxon is reported from various open habitats in the Mediterranean region and the
Near East where it probably lives polyphagously on dicotyledonous herbs and dwarf
shrubs (adults found on Atriplex, Cistus, Rosmarinus, Marrubium, Salicornia, Portulaca,
Thymus, Beta and others, often on succulents). Populations in central Europe may be
only unstable. Presumably 2 ‡‡ collected near Mainz and described as Jassus haagii Kirsch-
baum, 1868 („Mombach, Sandhügel, zwischen niederen Pflanzen, Ende April“) belong
to this taxon. Recently, it was also found in the Oberlausitz (Lauta, 1994, 1 † in a yellow
tray), in mining areas near Halle, as well as on flat roofs of Basel (Switzerland).
Funke (pers. comm.), Al Hussein et al. (1999), Kirschbaum (1868), Wagner (1939a), Walter & Emmrich
(1995); see also della Giustina (1989), Mühlethaler (2001), Remane (1987), Young & Frazier (1954)

Coryphaelus gyllenhalii (Fallén, 1826)

So far B VII – B IX; egg, 1 gen.
Along shores of mesotrophic lakes and ponds, in dense, often slightly shaded stands of
Schoenoplectus lacustris. In Germany at the western and southern edge of the range and
confind to the eastern half, mainly in glacial lakes of the northeast German plain and
Upper Bavaria, as well as in fish pond areas of the Oberlausitz, Thuringian Forest and
southern Harz Mountains. Western- and southernmost localities, which appear to be
 Deltocephalinae Fieber, 1869 167

strongly isolated from each other, are near Berlin, Bad Sachsa, Arnstadt, Ilmenau, Sees-
haupt and Penzberg (600 m a.s.l.).
Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Nickel (1997), HN

Balclutha boica W. Wagner, 1950

For this taxon the name B. lineolata (Horváth, 1904) was commonly used for several decades.
However, Webb & Vilbaste (1994) recently studied Horváth’s type material collected in central
Asia and concluded that it is a junior synonym of B. punctata (F.). This opinion is also supported
by Emelyanov (pers. comm.).
In Latvia and Lithuania according to Vilbaste (1974) M VIII – M V; adult(?), 1(?) gen.
Reported mainly from the eastern half of Europe. The only German record is from the
type locality in the Bavarian foreland of the Alps: Landsberg am Lech, 575m a.s.l.,
02.IX.1941, 3 †† „im hohen Gras der Auen am linken Lechufer ..., der Fundort wurde
noch 1941 durch Baumaßnahmen zerstört“. Reported from forests and forest glades in
Latvia and Lithuania, from Picea abies in spring in Sweden. Otherwise only published
from Estonia, Poland, the Czech Republic, Slovakia, Bulgaria and Greece. Reports from
other countries may refer to other species.
Fischer (1972), Wagner (1950); see also Anufriev & Emalyanov (1988), Drosopoulos et al. (1986), Nast (1987),
Ossiannilsson (1983), Vilbaste (1974), Webb & Vilbaste (1994)

Balclutha calamagrostis Ossiannilsson, 1961

Adults I – XII, mainly M VIII – E V; adult, 1 gen.
In scattered stands of Calamagrostis epigejos, occasionally also on C. pseudophragmites, in sun-
ny to moderately shady, moderately dry to moderately moist sites, mainly in mining areas,
ruderal sites, forest glades, as well as on gravel banks of alpine rivers. Hibernation presum-
ably takes place in the grass layer. In Germany only poorly documented and often misiden-
tified, but apparently widespread and common in favourable sites, e.g. in southern and
eastern Niedersachsen, in parts of Saxony-Anhalt, Thuringia, Saxony, northern Baden, as
well as in Upper Bavaria, found up to at least c. 800 m a.s.l. in the Rhön Mountains. So far
there are only few further records from northern and central parts of Europe (Sweden,
Finland, Baltic states, Poland, the Czech Republic, Austria, eastern France), but the range is
imperfectly known. German localities may be near its southern and western border.
Fischer (1972), Funke & Witsack (1998), Remane (1987), Remane & Reimer (1989), Rombach (1999a), Schie-
menz et al. (1996), Walter (pers. comm.), Nickel (1997), HN; see also della Giustina & Remane (2001), Holz-
inger (1996b), Nast (1987), Ossiannilsson (1983), Szwedo et al. (1996)

Balclutha punctata (Fabricius, 1775) sensu Wagner (1939)

Balclutha lineolata (Horváth, 1904) sec. Webb & Vilbaste (1994)
The taxonomic situation of European species of Balclutha is critical due to strong morphological
similarities. Further complications arose by the installation of a neotype for the name B. punctata
(F.) by Blocker (1967), who chose a specimen of the following species, which in turn had been
described as B. rhenana W. Wagner 1939 (see below). Following Blocker, Nast (1986, 1987) decid-
ed that B. tricolor (Gmelin, 1790) should be the valid name for B. punctata (F.) sensu Wagner.
168 Review of species

However, there is no evidence at all that Blocker’s neotype is conspecific with the species de-
scribed by Fabricius. Moreover, Blocker’s interpretation would cause serious confusion in Old
World leafhopper taxonomy. Therefore, it is rejected here in accordance with della Giustina (1989),
Holzinger et al. (1997), Ossiannilsson (1983), Remane & Fröhlich (1994a) and Webb & Vilbaste
(1994), who all follow Wagner’s opinion.
Adults I – XII, mainly E VII – E VI; adult, 1 gen.
In damp to wet, usually cool or moderately shady sites. Development of nymphs takes
place on grasses (Deschampsia flexuosa, Agrostis capillaris, Calamagrostis epigejos, Holcus
mollis and others), mainly in meadows and pastures (often near forest), along alleys and
hedges, in clearings, ruderal sites and open forests; adults often migrate to other habi-
tats and are found on coniferous trees in winter (notably Picea, Pinus, Taxus). The spe-
cies is widespread and common all over Germany, found up to at least 1300 m a.s.l. in
the Mittelgebirge and the Bavarian and Allgäu Alps. Single specimens, which may have
been vagrants, were collected up to 2000 m a.s.l. (also in Austria and Switzerland).
Achtziger (1991), Fischer (1972), Frommer (1996), Niedringhaus (1991), Reimer (1992), Remane (1987), Schie-
menz et al. (1996), Schwoerbel (1957), Wagner (1935, 1939a), HN; see also Günthart (1987a), Wagner &
Franz (1961)

Balclutha rhenana W. Wagner, 1939

Described from near Mainz and considered to be conspecific with B. punctata (F.) in a revision of
Nearctic Balclutha species by Blocker (1967), declaring a specimen of B. rhenana (W.Wg.) from the
Baltic Sea island of Rügen as neotype. This opinion is rejected here (see above).
Adults I – XII, mainly M VII – M VI; adult, 1 gen.
On Phalaris arundinacea usually in wet and rather eutrophic sites, usually near standing
or running water (including ditches) as well as in abandoned grassland, probably hi-
bernating in the grass layer. Widespread in Germany, in high frequency on the host (e.g.
along the edge of the Bavarian Alps, up to at least 750 m a.s.l., in the valleys of the
Danube, Neckar and Main, in Thuringia, southern Niedersachsen, the lower Elbe val-
ley), but often overlooked or confused with the preceding species. Thus, the number of
published records is low. From western parts of central Europe the species is only known
from the Netherlands and Switzerland, but not from Belgium and France. Hence, local-
ities along the upper and middle Rhine between Freiburg and Cologne may be at the
edge of the range. The locus typicus is Heidesheim near Mainz.
Fischer (1972), Frommer (pers. comm.), Reimer (1992), Schiemenz et al. (1996), SMNS, HN; see also Nast (1987)

Balclutha saltuella (Kirschbaum, 1868)

So far only M VIII – E VIII; adult(?), 1(?) gen.
In most parts of central Europe this species probably occurs as a temporary stray only,
living on grasses on various sunny, usually more or less disturbed sites. Although de-
scribed from the surroundings of Wiesbaden 130 years ago (from a single ‡), there are
only three further and very recent German records, from Upper Bavaria, southern Nied-
ersachsen and the Lake Constance Basin: Vorderriß, Isar floodplains, 800 m a.s.l.,
27.VIII.1996, 1 †, 1 ‡, on gravel banks; Ebergötzen, 170 m a.s.l., 19.VIII.1996, 1 ‡, and
 Deltocephalinae Fieber, 1869 169

Radolfzell, 400 m a.s.l., 21.VIII.1998, 1 ‡, both along waysides. The nearest records which
probably refer to more permanent populations are from southern Moravia (Czech Re-
public) and the middle Rhone valley (France).
Kirschbaum (1868), Nickel (1999a); see also della Giustina (pers. comm.), Lauterer (pers. comm.), Ribaut
(1952)

Macrosteles alpinus (Zetterstedt, 1828)

M VII – E IX; egg, 1 gen.
In central Europe, this species is confined to stands of tall sedges, in usually sunny, wet
to boggy sites of the upper montane and subalpine belt of the Alps and Sudetes. In
Germany it has only been recorded in 4 localities in the Allgäu and Bavarian Alps (Ober-
stdorf, Wannenkopf, 1650 m a.s.l., 1936; Oberstdorf, Geißalpsee, c. 1600 m a.s.l.,
06.VIII.1995; Berchtesgaden, Priesberger Moos, 1350 m a.s.l., 10.VIII.1997, and Berchtes-
gaden, Funtensee, 1600 m a.s.l., August and September 1992, in Malaise traps). Densi-
ties in these sites were usually high. Has been found between 1160 and 2380 m a.s.l. in
North Tyrol (Austria) and the Engadine (Switzerland). Otherwise it is reported from
large parts of the arctic and boreal zone of Europe, Asia and North America.
Wagner (1939a), Nickel (1999b), HN; see also Dlabola (1970), Günthart (1987a), Leising (1977), Nast (1972),
Vilbaste (1980)

Macrosteles cristatus (Ribaut, 1927)

E V – M X; egg, 2 gen.
A pioneer species in fallow fields, ruderal sites, mining areas, along field paths and
forest roads, on dry pond bottoms, in cereal crops, etc.; low numbers of individuals
(some of which may be vagrants) are also found in moderately moist to moderately dry
fertilized meadows and pastures and other habitats. Host plants are grasses and herbs,
perhaps also species of Cyperaceae. The original habitats presumably include gravel
and mud banks along rivers. Widespread all over Germany and particularly common
in southern and middle parts. Breeding takes place up to at least 800 m a.s.l., but single
individuals are frequently recorded at 1800 m a.s.l. in the Bavarian and Allgäu Alps.
Has been shown to transmit some yellows diseases in various parts of Europe, but prob-
ably without economic relevance.
Achtziger (1991), Fischer (1972), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Tharsen (1987),
Wagner (1935), Nickel & Achtziger (1999), HN; see also Brčák (1979)

Macrosteles fieberi (Edwards, 1889)

E V – B X; egg, 2 gen.
In hollows and on floating mats in raised and intermediate bogs, nowadays mainly in
old and overgrown peat diggings. The host plant is Eriophorum angustifolium. Literature
records from other types of habitats such as salt marshes, sand pits, etc. should be re-
vised. In Germany apparently confined to the northern half; at present, there are only
170 Review of species

6 verified records from the Weser-Ems region, North Friesland, eastern Holstein, the
Dübener Heide and the Solling, up to 450 m a.s.l.
Hildebrandt et al. (1998), Remane (1958), Schiemenz et al. (1996), Nickel (1997), HN

Macrosteles frontalis (Scott, 1875)

E V – B X; egg, 2 gen., perhaps 1 at higher altitudes.
In stands of horsetail in wet to moderately dry sites, usually along field paths and mar-
gins, railway embankments, in mining areas, but also in fens, intermediate bogs and
along lake shores. The main host plants are Equisetum arvense and E. palustre, but the
species is also reported to live on E. sylvaticum. Widespread in Germany and fairly com-
mon in the Alps and their foothills, rather sporadic elsewhere. The highest localities are
at 1350 m a.s.l. in the Berchtesgaden Alps.
Fischer (1972), Remane (1958), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1935,
1939a, 1951a), HN

Macrosteles horvathi (W. Wagner, 1935)

Among populations of northern Germany and the Bavarian Alps, single specimens have been
found which resemble M. nubilus Ossiannilsson, 1936 described from Sweden (Wagner 1939a,
HN). Until further evidence has been presented, these are considered to be abnormal individu-
als of M. horvathi (W.Wg.). Ossiannilsson (1954) himself was not convinced of the specific dis-
tinctiveness of M. nubilus Oss.
E V – M X; egg, 2 gen., according to Leising (1977) 1 in the North Tyrolian Alps (Austria).
A hygrophilous pioneer species of disturbed and wet, usually waterlogged, seepy or slight-
ly saline sites, often in gravel, sand and clay pits, clearings, along temporarily wet shores
of lakes and ponds, also in spring mires with predominating rushes and moderately sa-
line wet meadows and pastures (near the coast of the North and Baltic Sea as well as
inland). Host plants are rushes (Juncus articulatus, J. gerardii, probably also J. bufonius and
additional species). Widespread in Germany, but only scattered, recorded up to 1500 m
a.s.l. in the Bavarian Alps. In North Tyrol (Austria) and the Swiss Alps large populations
have been found in the subalpine belt, low numbers even up to 2250 m a.s.l.
Fischer (1972), Niedringhaus (1991), Reimer (1992), Schiemenz et al. (1996), Wagner (1935), HN; see also
Günthart (1987a), Leising (1977)

Macrosteles laevis (Ribaut, 1927)

M V – M X; egg, 2 gen., perhaps 1 at higher altitudes.
Another pioneer species of sparsely vegetated mud banks along rivers and lakes, sec-
ondarily, and often in high abundances, in fertilized meadows, ruderal sites, on tempo-
rarily dry ponds bottoms, in cereal crops and mining areas. Lives polyphagously on
grasses, rushes, sedges, and also dicotyledonous herbs. Widespread in Germany and
particularly abundant in the southern half (although with strong annual fluctuations of
numbers), but with decreasing density towards the north. Breeding takes place at least
up to 900 m a.s.l. in the Mittelgebirge, single individuals have been found at 1800 m a.s.l.
 Deltocephalinae Fieber, 1869 171

in the Allgäu Alps. In North Tyrol (Austria), large numbers have been reported from
1940 m a.s.l. Is known as a potential vector of plant viruses and mycoplasma-like or-
ganisms from various European countries, notably European aster yellows (EAY), stol-
bur, Clover phyllody (CP), Clover dwarf (CD) and Oats blue dwarf virus (OBDV).
Achtziger (1991), Fischer (1972), Marchand (1953), Müller (1956), Niedringhaus (1997), Remane (1987),
Schiemenz al. (1996), Tharsen (1987), Wagner (1935, 1951a), Nickel & Achtziger (1999), HN; see also An-
drzejewska (1962), Brčák (1979), Leising (1977), Ossiannilsson (1983)

Macrosteles lividus (Edwards, 1894)

B VI – E IX; egg, 2 gen.
In temporarily or permanently flooded, mesotrophic to moderately eutrophic, usually
sunny sites, notably shores of ponds, sand and gravel pits, as well as moderately saline
marshes, inland and near the coast. The host plant is Eleocharis palustris (probably in-
cluding E. uniglumis). Rather scattered in Germany, apparently at the southwestern edge
of the range, although probably under-recorded. Altogether there are c. 20 sites, mainly
along the coasts of the North and Baltic Sea and in the eastern half; westernmost local-
ities, most of which appear to be rather isolated, are on the island of Borkum, near
Hanover, Göttingen, Bad Brückenau (Rhön Mountains) and Vorderriß (upper Isar, 800 m
a.s.l.). Is also reported from England, the Netherlands and northern France, but not
from Belgium and Switzerland.
Niedringhaus & Olthoff (1993), Remane (pers. comm.), Schiemenz et al. (1996), Wagner (1939a, 1941b), Walter
(pers. comm.), Nickel & Remane (1996), Nickel (1997, 1999a), HN; see also della Giustina & Remane (2001)

Macrosteles maculosus (Then, 1897)

E V – B X; egg, 2 gen.
In sunny, damp to moderately dry, usually sandy or gravelly sites of early successional
stages with sparse or low-growing vegetation cover, mainly trampled patches, ruderal
sites, gravel and paved sites (even within cities). The host plant is Polygonum aviculare.
In Germany apparently at the northwestern edge of the range, known from a few records
in the southeastern half only. Border localities are near Balingen (Swabian Jura), Stutt-
gart, Wetzlar, Marburg, Göttingen, Salzgitter, Helmstedt, Quedlinburg, Halle, Leipzig
and Meißen; is also reported from near Stettin (Poland). The highest localities are at
700 m a.s.l. in the Rhön Mountains, at 1000 m a.s.l. in the Engadine (Switzerland).
Fröhlich (1996a), Rabeler (1952), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Wagner (1941a), Nickel
(1994), HN; see also Günthart (1987a)

Macrosteles oshanini Razviazkina, 1957

So far M VIII – M IX, in England according to Le Quesne (1969) VI; egg, 2(?) gen.
Most records of this species are from the eastern half of Europe. It lives in wet sites,
preferentially fens with strongly fluctuating water tables or moderate salinity, also near
shady forest margins. The host plants are unknown, but may include species of Cyper-
172 Review of species

aceae or Poaceae. In Germany found only in 6 localities in Holstein (Steinbek), Meck-

lenburg (Hagenow, Leussow, Neukloster), Brandenburg (Trechwitz) and Thuringia (Jena),
all below c. 150 m a.s.l.
Kuntze (1937), Müller (pers. comm.), ZIMH, HN; see also Ossiannilsson (1983)

Macrosteles ossiannilssoni Lindberg, 1954

B VI – B X; egg, 1 - 2 gen.
In wet, usually peaty sites (intermediate bogs, spring mires, peaty meadows and pas-
tures). The host plants are probably Carex species and Rhynchospora alba, perhaps also
Juncus spp. and grasses. Vagrants may be encountered far away from typical habitats.
In Germany autochthonous populations are known from scattered sites in the north-
western plains as well as in the Erzgebirge, Frankenwald, Bavarian and Black Forest,
and the Bavarian Alps. Found up to at least 1500 m a.s.l. in Upper Bavaria, up to 2370 m
a.s.l. in the Engadine (Switzerland).
Niedringhaus & Olthoff (1993), Reimer (1992), Walter (1998), Nickel & Achtziger (1999), Nickel & Sander
(1996), Nickel (1999b), HN; see also della Giustina (1989), Günthart (1987a, 1987b), Leising (1977), Nast (1976a)

Macrosteles quadripunctulatus (Kirschbaum, 1868)

E V – B X; egg, 2 gen.
In sunny, disturbed, usually sandy and sparsely vegetated, moderately dry to dry sites
(sand pits, abandoned fields, military training areas, waysides, trampled patches in
dry grassland, etc.), in warmer regions of southern Germany also on loamy substrates
(notably in vineyards). Setaria and Panicum, as well as Corispermum have been reported
as host plants. The species is uncommon in Germany, rather localized, largely confined
to sand and viticultural regions (up to at most 300 m a.s.l.); it has been recorded in the
north German plain, middle Hessen, the Thuringian and Leipzig Basin, along the up-
per and middle Rhine, in the Main valley and Mittelfranken. The locus typicus is Mom-
bach near Mainz. Is reported as a potential vector of yellows diseases in other countries,
but is unlikely to cause major effects in central Europe due to the scattered occurrence.
Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Remane (1987), Schiemenz et al. (1996), Trümbach
(1959), Wagner (1935), HN

Macrosteles sardus Ribaut, 1948

So far E V – E IX; egg, 2 gen.
Usually in moist, moderately shady sites in river floodplains and along lake shores. Is
reported to live on grasses in Kazakhstan, but no further details are known on habitat
and host plants. In Germany known only from a few records from eastern Holstein
(near Plön and Lübeck), the lower course of the Elbe (near Lüneburg) and the upper
Rhine plain (near Freiburg, Speyer, Heidelberg and Darmstadt), all below 200 m a.s.l.,
but found at 700 m a.s.l. in Trentino (northern Italy).
Remane (1962), Remane & Fröhlich (1994b), Nickel (1997), HN; see also Mityaev (1971)
 Deltocephalinae Fieber, 1869 173

Macrosteles septemnotatus (Fallén, 1806)

B VI – E X; egg, 2 gen.
In wet, sunny to moderately shady sites, mainly abandoned grassland, fens, shores of
running and standing water, and fen woods. The host plant is Filipendula ulmaria. Wide-
spread in Germany, fairly common, in intermediate to high frequency on the host, re-
corded up to 1200 m a.s.l. in the Allgäu Alps.
Fischer (1972), Heller (1987a), Nikusch (1976), Reimer (1992), Schiemenz et al. (1996), Wagner (1935), Nick-
el & Achtziger (1999), HN

Macrosteles sexnotatus (Fallén, 1806)

M V – M X; egg, 2 gen. (see also Witsack 1985).
The life history is very much like in M. cristatus (Rib.) and M. laevis (Rib.). M. sexnotatus
(Fall.) frequently occurs syntopically with these as a pioneer species in fertilized mead-
ows and pastures, as well as in cultivated crops, but in general, it is more hygrophilous
and often dominant in moist, eutrophic meadows, moist ruderal sites, and on mud banks
with sparse vegetation, also in moderately saline grassland. Migrating individuals are
sometimes found in large numbers in other habitats. Host plants are Poaceae, Cyper-
aceae, and probably also dicotyledonous herbs. Widespread in Germany and very com-
mon; frequently up to at least 1200 m a.s.l. in the Bavarian and Allgäu Alps, at least
occasionally up to 1600 m a.s.l.
Achtziger (1991), Fischer (1972), Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992), Schiemenz et
al. (1996), Wagner (1935), Walter (1996, 1998), Nickel & Achtziger (1999), HN, and others

Macrosteles sordidipennis (Stål, 1858)

E V – E IX; egg, 2 gen.
In salt marshes along the coast and inland. The main host plant is Puccinellia distans, but
P. maritima and Juncus gerardii have also been reported. In Germany known from the
East Frisian Islands, the North Sea coast between Emden and Cuxhaven, the Baltic Sea
coast near Heiligenhafen, as well as from inland salt marshes in Holstein, Saxony-An-
halt, Thuringia, middle Hessen and eastern Niedersachsen, also from sites heavily af-
fected by phosphate immissions near Jena. The highest localities are at c. 250 m a.s.l.
Fröhlich (1996a, and pers. comm.), Hildebrandt (1995), Müller (1985), Niedringhaus & Olthoff (1993), Schi-
emenz et al. (1996), Wagner (1937a), HN; see also Linnavuori (1952)

Macrosteles variatus (Fallén, 1806)

B VI – E X; egg, 2 gen.
In eutrophic, moderately moist to moderately wet, usually moderately shady sites with
lush herb layer, mainly along shores and margins in floodplains of rivers and streams,
along forest roads, shaded ditches, etc., preferentially on Urtica dioica, perhaps also on
other species of dicotyledonous herbs. Widespread in Germany, fairly common in most
174 Review of species

valley bottoms, but often overlooked; recorded up to at least 800 m a.s.l. in the Mittelge-
birge and at the edge of the Alps.
Fischer (1972), Heller (1987a), Mölleken & Topp (1997), Niedringhaus (1997), Reimer (1992), Remane (pers.
comm.), Lauterborn (1920), Schiemenz et al. (1996), Trümbach (1959), Wagner (1935), HN

Macrosteles viridigriseus (Edwards, 1922)

E V – M X; egg, 2 gen.
Mainly in temporarily dry or moderately saline pastures and meadows (often in river
floodplains or floodplain depressions, occasionally in intensively managed sites), also
in rather low-vegetated spring mires and fens and, as a pioneer species, on sparsely
vegetated banks of gravel, sand and mud. Records from other habitats probably refer to
vagrants. Host plants are various grasses, probably also Cyperaceae. Has been found in
most parts of Germany, but often in very low numbers only. Records of large and repro-
ducing populations are more localized, but have been found up to 850 m a.s.l. in the
Thuringian Forest, up to 1250 m a.s.l. in Austria. Has been reported as a vector of Clo-
ver phyllody (CP) in Britain, but apparently without economic importance.
Emmrich (1966), Marchand (1953), Niedringhaus & Olthoff (1993), Schiemenz et al. (1996), Walter (1996),
Nickel & Achtziger (1999), HN; see also Brčák (1979), Wagner & Franz (1961)

Erotettix cyane (Boheman, 1845)

So far M VII – M IX; egg, 1 gen.
On floating water plants, usually on small ponds and lakes in cool and moderately
shady situations. Host plants are Potamogeton natans, Nuphar lutea, Nymphaea alba, and
probably additional species of floating plants. Has been reported from Trapa natans and
Marsilea quadrifolia in Austria. In Germany only very sporadic, known from about 20
localities, mostly to the east of a line running from Schwerin to Kassel, Würzburg and
Munich, recorded up to 600 m a.s.l. Further localities, which appear to be isolated, are
situated in the Weser-Ems region and the southern upper Rhine plain, although the
species is also reported from the Netherlands, Belgium and France.
Funke & Witsack (1998), Coll. Förster (Niedringhaus pers. comm.), Schiemenz et al. (1996), Walter (1996),
Nickel & Remane (1996), HN; see also Holzinger (1995b)

Sonronius binotatus (J. Sahlberg, 1871)

M VII – B IX; egg, 1 gen.
This species is widespread in the Siberian and north European taiga zone, with rather
isolated populations in central European mountain regions. It lives in forest clearings
and along roads on Epilobium angustifolium; at least adults are also found on other spe-
cies of Epilobium. In Germany only known from altogether 4 localities in the Bavarian
Alps (Mittenwald), the Vogtland (near Plauen) and the Rhön Mountains (to the east of
Fulda), between 500 and 1000 m a.s.l.
Bittner & Remane (1977), Haupt (1925), Reimer (1992), Walter & Emmrich (1995), Nickel (1999b)
 Deltocephalinae Fieber, 1869 175

Sonronius dahlbomi (Zetterstedt, 1840)

M VII – B IX; egg, 1 gen.
Known from Germany only from altogether 6 records in the Allgäu and Bavarian Alps
(in the vicinity of Oberstdorf, Mittenwald and Lenggries, between 700 and 1100 m a.s.l.).
Found on Epilobium angustifolium in cool and moderately shady sites, usually in or near
woodland; from Sweden Rubus idaeus, Alchemilla spec. and Filipendula ulmaria have been
reported as additional food plants.
Haupt (1925), Remane (pers. comm.), Wagner (1939a); see also Ossiannilsson (1983)

Sagatus punctifrons (Fallén, 1826)

M VI – M IX; egg, 1 gen.
Along rivers and streams, on gravel banks, dunes and in mining areas, on narrow-leaved
species of willows; most individuals (including nymphs) found on Salix purpurea, S.
triandra and S. viminalis, low numbers also on other species (S. pentandra, S. fragilis, S.
alba). Furthermore, S. repens is the main host in coastal areas, S. eleagnos in the Alps and
their foothills. Widespread at lower and middle altitudes of Germany (up to at least
950 m a.s.l.), in intermediate to high frequency, locally abundant in favourable sites.
Fischer (1972), Frommer (pers. comm.), Niedringhaus (1991), Reimer (1992), Schiemenz et al. (1996),
Schönitzer & Oesterling (1998b), Wagner (1935, 1939a), HN

Deltocephalus maculiceps Boheman, 1847

E VI – M IX; egg, 1 gen.
This is a western European species of intermediate and heathland bogs. Eriophorum
vaginatum has been reported as host plant, but there is more evidence that the species
lives on Molinia caerulea. It is rare in Germany and restricted to northern parts, recorded
from the Weser-Ems area, the Lüneburger Heide, Holstein, Mecklenburg, the Nieder-
lausitz and the vicinity of Halle, up to c. 100 m a.s.l. Otherwise it is only reported from
the island of Gotland (Sweden), Denmark, Ireland, England, the Netherlands, north-
western Poland, France and Spain; records from other countries are doubtful.
Remane (1958, and pers. comm.), Schiemenz et al. (1996), Wagner (1935), HN; see also Le Quesne (1969),
Nast (1976a, 1987), Ossiannilsson (1983)

Deltocephalus pulicaris (Fallén, 1806)

M V – E X; egg, 2 gen., according to Leising (1977) 1 in the North Tyrolian Alps (Austria).
In sunny to moderately shady, wet to moderately dry sites, often extremely abundant in
low-vegetated sheep and cattle pastures, fertilized meadows, in lower numbers also
along forest roads, in fens, moderately saline habitats and lawns in parks, gardens and
even sports grounds. Host plants are various grasses (Agrostis, Poa, Dactylis, Elymus,
Lolium and others). Widespread in Germany, very common in most parts, also in the
Mittelgebirge and the Alps, where frequently found up to at least 1500 m a.s.l. In Austria
176 Review of species

and Switzerland large populations occur at 2100 m a.s.l., at least single individuals were
recorded at 2510 m a.s.l.
Fischer (1972), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Schwoerbel
(1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others; see also Günthart (1984, 1987a),
Leising (1977), Wagner & Franz (1961)

Recilia coronifer (Marshall, 1866)

E VI – M X; egg, 1 gen.
In moderately shady, occasionally also sunny, wet to damp, usually acidic sites, mainly
open oak and pine forests (often in clearings, along margins and roads), bog margins
and straw meadows. Host plants are Holcus mollis and, less frequently, Molinia caerulea.
Widespread in the lowlands of Germany (up to at least 600 m a.s.l.), but usually in low
individual numbers, at the northern edge of the range in Schleswig-Holstein; northern-
most localities are near Heide, Rendsburg and Eckernförde.
Achtziger (1991), K. Heller (pers. comm.), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1958,
and pers. comm.), Schiemenz et al. (1996), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger
(1999), HN

Recilia horvathi (Then, 1896)

So far E VIII – E IX; egg(?), 1(?) gen.
This species’ range extends from Mongolia to central Europe, where it occurs in rather
isolated populations in xerothermic sites. In Germany there are only two known local-
ities in the northern upper Rhine plain near Heidelberg: Pferdstrieb and Pflege Schönau,
both near Sandhausen, c. 110 m a.s.l., 26.VIII.1981 and 27.IX.1982, several ††, on inland
sand dunes on Corynephorus canescens. There are further records from Moravia (Czech
Republic), Styria (Austria), Slowenia, northern Italy and Ticino (Switzerland).
Heller (1996); also Lauterer (pers. comm.), Then (1896), Remane & Fröhlich (1994b)

Recilia schmidtgeni (W. Wagner, 1939)

So far E VIII – X; egg(?), 1(?) gen.
Like the preceding species, known in Germany only from two localities in sand areas of
the northern upper Rhine plain: Mainz, 24.VIII.1951, 2 ††, „in einem Pflanzgarten bei
Uhlerborn“, and Dudenhofen near Speyer, Oktober 1989, „in Anzahl in Cynodon-Be-
ständen in feuchteren Senken am Rand eines Dünengebietes“. Has been recorded in
moderately saline sites in eastern parts of Austria. In the Mediterranean region the spe-
cies lives in various open habitats, often in disturbed patches.
Remane & Fröhlich (1994b), Wonn (1956); see also Fröhlich (1996a)

Endria nebulosa (Ball, 1900)

B VII – M IX; egg, 1 gen.
 Deltocephalinae Fieber, 1869 177

In stands of tall grass in sunny to moderately shady, usually rather acidic, moist to
damp sites, mainly in forest clearings, coal mining areas and abandoned fields. Lives on
Calamagrostis epigejos, presumably also on C. canescens. First recorded in Germany in
1959, probably at the western and southern edge of the range, known only from a few
localities in the eastern half, reaching westward to Greifswald, Lüchow, Goslar, Halle
and Dresden, also found to the south of Munich (c. 600 m a.s.l.); not reported from the
Netherlands, Belgium, France, Switzerland and Austria. It has been proposed that this
species had recently been introduced from the Nearctic region. This opinion does not
seem plausible considering the strong association with a native grass species. More-
over, it is very difficult to collect due to its epigeic life habits; thus it is probably much
under-recorded in most parts of its range.
Funke & Witsack (1998), Remane (1961c), Schiemenz et al. (1996), Nickel (1997), HN; see also Arzone et al.
(1987), Lauterer (1980)

Doratura exilis Horváth, 1903

B VI – E IX; egg, 1 gen., perhaps 2 in southern Germany or in warm years.
On usually grey-leaved subspecies of Festuca ovina in oligotrophic, sunny, low-vegetat-
ed, often strongly grazed sites on dry, basic as well as acidic substrates. Widespread in
the lowlands of Germany, but only scattered, locally dominant on sandy dry grassland
of the north German plain and on calcareous hillsides in the middle parts. Not recorded
to the south of the Danube, but found up to at least 750 m a.s.l. on the Swabian Jura. The
northernmost localities are near Hamburg and Lauenburg; the distribution border ap-
parently runs through southern parts of Holstein. Not reported from Schleswig, Den-
mark and Scandinavia, except southern Sweden.
Reimer (1992), Remane (1958, 1987), Schiemenz et al. (1996), SMNS, Trümbach (1959), Wagner (1935, 1951a),
HN; see also Ossiannilsson (1983)

Doratura stylata (Boheman, 1847)

B VI – M X, in warmer regions M V – XI; egg, 1 gen., perhaps 2 in southwestern Germany.
In oligotrophic and sunny, dry to damp, occasionally also moist sites, mainly low-pro-
ductivity grassland, waysides and ruderal sites, often dominant in meadows and pas-
tures on nutrient-poor substrates of the submontane belt. The main host plants are Fes-
tuca rubra, F. ovina and Agrostis capillaris, probably also additional species of fine-leaved
grasses, e.g. Poa pratensis angustifolia and Nardus stricta. Widespread in Germany and
particularly common in regions with less intensive agriculture, found up to at least
1200 m a.s.l. in the Black Forest, but frequently in the alpine belt (up to 2200 m a.s.l.) of
the Austrian and Swiss Alps.
Achtziger (1991), Fischer (1972), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Remane (1987), Schie-
menz (1969), Schiemenz et al. (1996), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others; see
also Günthart (1987a), Leising (1977), Wagner & Franz (1961)
178 Review of species

Doratura horvathi W. Wagner, 1939

M VI – B X; egg, 1 gen.
In sunny, moderately dry to dry, basic (occasionally also acidic), usually grazed sites
with low-growing and sparse grass cover. The host plant is Helictotrichon pratensis. In
Germany at the northern edge of the range, only scattered, but locally dominant, nota-
bly in regions of limestone, gypsum and porphyry of middle parts, between the foot-
hills of the Alps and the northern edge of the Mittelgebirge (Eifel Mountains, southern
Niedersachsen, eastern Harz Mountains, Thuringia, Rhine-Nahe region, Main and Rhön
region, Tauber valley and Franconian Jura, also foreland of the Alps and Kaiserstuhl).
The northernmost localities are near Mechernich, Brilon, Göttingen, Halberstadt, Halle
and Jena, up to at least 700 m a.s.l. Otherwise the species is reported only from central
Poland, Moravia (Czech Republic), Slovakia, Austria and Italy.
Reimer (1992), Rombach (1999a, 1999b), Schiemenz et al. (1996), SMNS, Wagner (1939a, 1951a), ZIMH,
Nickel (1994), HN; see also D’Urso (1995), Holzinger & Remane (1994), Lauterer (1984), Nast (1976a)

Doratura impudica Horváth, 1897

M VI – E IX; egg, 1 gen.
Among sparse stands of tall grasses, usually on acidic, sandy to gravelly soils in moder-
ately dry to dry, often moderately shady sites, mainly in clearings of pine forests, in the
vicinity of inland dunes and in sandy ruderal sites. The host plant is usually Calama-
grostis epigejos, perhaps also Elymus spp. (in northern Italy Elymus athericus). In Germa-
ny with strongholds in the northeastern plains, where locally common in favourable
sites, but perhaps at the western edge of the range. Westernmost localities are on the
island of Amrum, near Lüneburg, Halle, Leipzig, Dresden and Görlitz, up to at most
200 m a.s.l. Published records from the Mittelgebirge refer to stray or misidentified indi-
viduals. Is also reported to occur in northern and eastern parts of France and in the
Netherlands.
Bartels et al. (1992), Funke & Witsack (1998), Remane (pers. comm.), Schiemenz (1969), Schiemenz et al.
(1996), Walter (pers. comm.), Nickel (1997), HN; see also Della Giustina (1989), Gravestein (1976)

Doratura homophyla (Flor, 1861)

M V – M X; egg, 2 gen.
In sunny, sparsely vegetated grass stands in damp to dry, moderately eutrophic, often
disturbed sites, usually on sandy, occasionally also loamy substrates (disturbed patches
in dry grassland, abandoned fields, low-productivity pastures of cattle or sheep, way-
sides, military training areas, etc.). Host plants are grasses (Agrostis capillaris, probably
also Elymus repens, Festuca rubra, Poa compressa and others). Locally common in the low-
lands of Germany (up to at most 400 m a.s.l.) and largely confined to sand and gypsum
regions (north German plain, upper and middle Rhine, valleys of the Main, Danube
and Regnitz, Thuringian and Leipzig Basin, Oberlausitz).
Frommer (1996), Niedringhaus (1997), Remane (1987), Schiemenz et al. (1996), Trümbach (1959), Wagner
(1935, 1939a, 1951a), Nickel & Achtziger (1999), HN
 Deltocephalinae Fieber, 1869 179

Doratura littoralis Kuntze, 1937

Was originally described as a variety of D. homophyla (Fl.), after specimens from near Rostock,
but treated as a distinct species by Wagner (1941b), and subsequently, Dworakowska (1968).
However, the morphological differences appear to be very small; after a revision of more mate-
rial they may turn out to fall within the range of intraspecific variation of D. homophyla (Fl.).
So far VI – VIII; egg, 1(?) gen.
Altogether known only from very few records from Heligoland, eastern Holstein (Hoh-
wachter Bucht), Mecklenburg (between Rostock and Ahrenshoop) and the lower Oder
valley (to the south of Stettin, Poland), mainly on white and grey dunes near the coast;
the Polish locality, however, is an inland dune. Ammophila arenaria has been suspected
to be the host plant.
Kuntze (1937), Niedringhaus & Olthoff (1993), Schiemenz et al. (1996), Wagner (1941b); for identification
see Dworakowska (1968)

Platymetopius undatus (De Geer, 1773) and Platymetopius henribauti Dlabola, 1961
P. henribauti Dlab. was described after highly variable characters (shape of forehead and 7th
sternite of ‡‡), with contradictory information on food plants, without revising the type materi-
al of P. undatus (De G.), and without citing the preliminary revision of the P. undatus group by
Wagner (1939a). Despite all these uncertainties, P. henribauti Dlab. has since been reported from
most parts of the Palearctic (see Nast 1972, 1987). Published data on life history also appear
contradictory (see Dlabola 1961b; Ossiannilsson 1983; Ribaut 1952; Wagner 1939a). In conclu-
sion, it is uncertain whether P. undatus (De G.) and P. henribauti Dlab. are distinct species.
So far VII – IX; egg, 1 gen.
Most German records of †† with pygophor appendages of the P. undatus type are from
the rain shadow region to the east of the Harz Mountains (near Bad Frankenhausen,
Eisleben, Querfurt, Dessau, Grimma, Bautzen, Guttau, the latest record dating from
1965); more scattered localities are near Hamburg, Rostock, Hanover, in the northern
upper Rhine plain (Mainz, Bad Dürkheim, the latter dating from 1995). In most cases,
there were only single individuals swept from low-growing xerothermophilic vegeta-
tion (Helianthemum and other species) or from oak and birch. Apparently, there was a
dramatic decline during recent decades.
Kirschbaum (1868), Remane (1987), Schiemenz et al. (1996), Wagner (1935, 1939a), ZIMH, HN

Platymetopius major (Kirschbaum, 1868)

B VI – M X; egg, 1 gen., 2 in southern Germany.
Along sunny woodland margins in damp to dry, basic to acidic sites (dry grassland
with scattered trees and shrubs, spatially diverse forest margins, etc.). Nymphs are found
among low vegetation (locally in cushions of Helianthemum nummularium, but also on
other plants), adults ascend various species of deciduous trees and shrubs (mainly Quer-
cus, Betula, Acer, also Crataegus, Prunus, Fagus and others). Widespread in middle and
southern parts of Germany and fairly common in warmer regions, but rather localized
in the north, where recorded from Holstein, the Wendland of Niedersachsen and Meck-
lenburg (although apparently absent from the Weser-Ems area). To the north of it, there
180 Review of species

are only a few old reports from Jutland and the Baltic Sea island of Öland. The highest
localities are at 500 m a.s.l. in the Mittelgebirge, at 900 m a.s.l. on sunny calcareous hill-
sides of the Bavarian Alps.
Fischer (1972), Post-Plangg & Hoffmann (1982), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oester-
ling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN; see also Ossiannilsson (1983)

Platymetopius guttatus Fieber, 1869

So far B VII – E VIII; egg, 1 gen.
In open xerothermic forests and along their margins, mainly in sun-exposed slope or
plateau situations. Adults are usually found on Betula pendula and Quercus; the food
plants of nymphs are unknown, but are likely to include low-growing herbs. From Ger-
many known only from about 15 scattered localities in the valleys of the Nahe, Main
and Franconian Saale, and in warmer regions of Thuringia and Saxony-Anhalt (near
Nordhausen, Bad Frankenhausen, Querfurt, Jena, Gera and Rudolstadt), up to at most
400 m a.s.l., but the species may be under-recorded. Not reported from north Germany,
but from southern parts of Norway and Sweden.
Perner (1997), Schiemenz et al. (1996), Wagner (1939a), Wagner & Franz (1961), Nickel & Remane (1996),
HN; see also Anufriev & Kirillova (1998), Ossiannilsson (1983)

Idiodonus cruentatus (Panzer, 1799)

B VII – M X; egg, 1 gen.
In temporarily moist to moderately dry, basic as well as acidic sites, usually in open
stands of trees or shrubs with varying proportions of evergreens. Nymphs are found
among low-growing vegetation, most adults have been swept from Tilia, Betula, Fagus,
Rubus, Vaccinium, Calluna, and additional species of woody plants and dwarf shrubs.
Widespread and locally common in the southern half of Germany, mainly in open pine
and spruce forests in areas of Jurassic and Triassic limestone, and among subalpine
scrub, up to at least 1750 m a.s.l. in the Bavarian Alps, up to 2200 m a.s.l. in alpine dwarf
shrub heaths of North Tyrol (Austria) and the Engadine (Switzerland); rather scattered
in the northern half, where mainly confined to raised bog margins, occasionally in for-
est clearings and glades.
Fischer (1972), Reimer (1992), Rombach (1999a), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b),
Wagner (1935, 1939a), HN; see also Günthart (1987a), Leising (1977)

Colladonus torneellus (Zetterstedt, 1828)

B V – E VIII, in the Czech Republic and Slovakia according to Lauterer (1980) M IV – B
VII; nymph, 1 gen.
In mixed and coniferous forests (usually under Picea or Pinus) in cool sites with shrubby
undergrowth, also reported from deciduous forests in the Czech Republic and Slovakia.
Not much is known on the species’ life history; perhaps it is a vertical migrant which quick-
ly ascends into the canopy layer after emergence. Recorded in larger numbers from Melica
 Deltocephalinae Fieber, 1869 181

uniflora in Moravia (Czech Republic), also on various shrubs and dwarf shrubs (Salix, Betu-
la, Alnus, Rubus, Vaccinium, Calluna) in northern Europe, the Swiss Alps and Mecklenburg.
In Germany only known from 15 records usually comprising single specimens, probably
living near the western edge of the range; border localities are near Plön, Waren an der
Müritz, Ilmenau, Augsburg and Mittenwald, found up to 1000 m a.s.l. in the Bavarian Alps.
Furthermore, there are a few French records, mostly from eastern parts.
Haupt (1925), Fischer (1972), Peter & Roth (1996), Remane & Fröhlich (1994b), Schiemenz (1977), Schie-
menz et al. (1996), Schönitzer & Oesterling (1998b), HN; see also Günthart (1987a), Lauterer (pers. comm.),
Ossiannilsson (1983), Ribaut (1952)

Lamprotettix nitidulus (Fabricius, 1787)

B VII – E X; egg, 1 gen.
In deciduous forests in damp to moist, usually eutrophic sites, mainly flood-plain and
fen woods, as well as oak-hornbeam forests. Nymphs are found in low-growing herba-
ceous vegetation (often among Poaceae, Lamium or Urtica), adults after emergence quickly
ascend various deciduous trees (Quercus, Alnus, Betula, Ulmus, Tilia and others). Wide-
spread in the lowlands of Germany, but seldom found (although easily recorded in
numbers with suitable methods such as tree eclectors, light traps, and sweeping of
nymphs on the forest floor). Highest localities are at 600 m a.s.l. in the Mittelgebirge and
the foothills of the Alps.
Büchs (1988), Frommer (1996), Kupka (1925), Niedringhaus & Olthoff (1993), Reimer (1992), Schönitzer &
Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a), HN

Allygus communis (Ferrari, 1882)

Was originally described as a variety of the following species, and for a long time considered to
be conspecific with it, until Ossiannilsson (1983) pointed out that there are considerable differ-
ences in genitalic structures in both sexes. Thus, a number of older records of A. mixtus (F.) may
refer to A. communis (Ferr.).
M VI – M X; egg, 1 gen.
Along sunny forest margins and in semi-open stands of trees; in damp to dry sites,
locally syntopic with A. mixtus (F.), but generally preferring warmer and drier habitats.
Nymphs are found among low-growing herbaceous vegetation, presumably on grasses
and dicotyledons, adults usually on Quercus and Betula. Widespread in the lowlands of
Germany, but less common than the following species, and with strongholds in rather
warm regions of low altitudes, recorded only up to 600 m a.s.l. in the Mittelgebirge and
the foothills of the Alps.
Günthart (1988), Kolbe & Bruns (1988), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987),
Schönitzer & Oesterling (1998b), SMNS, ZIMH, Nickel & Remane (1996), Nickel (1997), HN

Allygus mixtus (Fabricius, 1794)

Most older records are in need of revision and may refer to A. communis (Ferr.) (see above).
E VI – M X; egg, 1 gen.
182 Review of species

In closed to semi-open stands of trees in moist to moderately dry sites (deciduous and
mixed forests, forest margins, alleys, etc.). Nymphal development takes place among
low-growing herbaceous vegetation, presumably on grasses and also dicotyledons; most
adults are swept from various trees (Betula, Populus, Salix, Quercus, Acer, Alnus, Frangu-
la, Pinus and others). Widespread in Germany and fairly common in many places, fre-
quently found up to 1300 m a.s.l. in the Mittelgebirge and the Bavarian Alps.
Achtziger (1991), Frommer (1996), Günthart (1988), Niedringhaus (1991, 1997), Reimer (1992), Remane
(1987), Schönitzer & Oesterling (1998b), SMNS, Nickel (1997), HN

Allygus maculatus Ribaut, 1952

B VII – B X; egg, 1 gen.
In oak forests in basic to acidic, temporarily moist to dry sites of lower altitudes (recorded
up to 450 m a.s.l. only). Nymphal development takes place in the herb layer (mainly on
grasses), most adults are swept from oaks (Quercus robur and Qu. petraea). In Germany wide-
spread, but seldom found (at present c. 30 localities only), mainly between the Danube and
the northern edge of the Mittelgebirge, also near Berlin and Ratzeburg. To the north of it,
there are only two old reports from Jutland and southern Sweden. Most sweep net records
comprise only single individuals, larger numbers have been collected in light traps.
Fischer (1972), Rabeler (1962), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Schwoerbel (1957), HN;
see also Ossiannilsson (1983)

Allygus modestus Scott, 1876

M VI – B X; egg, 1 gen.
In open deciduous forests and semi-open cultivated habitats (along forest margins, alleys,
etc.), usually in moderately wet to damp, occasionally also moderately dry sites. Nymphs
feed on grasses (but perhaps also on dicotyledonous herbs), adults on Quercus, Ulmus, Al-
nus, Betula, Acer, Prunus and other deciduous trees. Widespread in Germany, including the
northern plains, but apparently confined to the lowlands (up to at least 550 m a.s.l. in the
foreland of the Alps). North German localities are near the northern border of the range,
which runs through England, Jutland, southern Sweden and northwestern Poland.
Frommer (1996), Günthart (1988), Niedringhaus & Bröring (1988), Remane (1987), Schiemenz al. (1996),
SMNS, Trümbach (1959), Wagner (1935, 1939a, 1951a), HN; see also Gillerfors (2002), Le Quesne (1969),
Nast (1976a), Ossiannilsson (1983)

Allygidius commutatus (Fieber, 1872)

M VI – B X; egg, 1 gen.
Along sunny forest margins and in open stands of deciduous and coniferous trees of
moderately moist to dry, basic to acidic sites, preferentially in warm situations. Host
plants are mainly tall grasses (Brachypodium pinnatum, Calamagrostis arundinacea, C. epige-
jos, Melica uniflora and others), at least some adults are found on deciduous trees (Quer-
cus, Ulmus, Betula and others). Widespread and fairly common in middle and southern
 Deltocephalinae Fieber, 1869 183

parts of Germany (although usually collected in small numbers only), up to at least

800 m a.s.l. in the Mittelgebirge and the foothills of the Alps, with decreasing density of
localities in the north German plain; apparently absent from the Weser-Ems area. How-
ever, the species is reported to be widespread in southern Scandinavia.
Achtziger (1991), Fischer (1972), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schönitzer
& Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN; see also Ossiannilsson (1983)

Allygidius abbreviatus (Lethierry, 1878)

M VI – B X; egg, 1 gen.
In basic to acidic, xerothermic sites, usually in the vicinity of shrubs or trees. Nymphs
are found in low-growing herbaceous vegetation; at least some adults ascend various
deciduous trees. In Germany confined to southwestern parts. The status in Germany
has long been uncertain, since there was only a single proved but inaccurate record
(„Rheinhessen“). Further records dating before 1940 are not verified or were corrected
later. Has recently been found in 5 localities in the northern Eifel Mountains and along
the Palatinate Haardt (near Bad Münstereifel, Grünstadt and Neustadt an der Wein-
straße, all between 150 and 350 m a.s.l.). Otherwise only reported from Belgium, France,
the Czech Republic, Slovakia, Hungary, Austria and Italy.
Rombach (1999a), Wagner & Franz (1961), HN; see also Puton (1886), Wagner (1935, 1941b), Hegab et al.
(1980) (see this paper also for identification of nymphs), Lauterer (pers. comm.), Nast (1987)

Allygidius atomarius (Fabricius, 1794)

Mainly B VI – M IX, according to Schiemenz et al. (1996) M V – E X; egg, 1 gen.
In warm, moderately moist to moderately dry sites with semi-open stands of trees, mainly
in river floodplains, along forest margins, in coppice-with-standards, etc. The nymphs
live among herbaceous vegetation, probably on tall grasses, most adults are swept from
Ulmus, Quercus and other deciduous trees. In Germany at the northern edge of the range,
found only sporadically and in low numbers, at lower altitudes (up to at least 600 m
a.s.l.), mainly in the larger river valleys (Rhine, Main, Elbe, Saale, Unstrut, Havel, Oder
and others). Northernmost localities are near Cologne, Göttingen, Boizenburg, Stendal,
Oranienburg and Frankfurt an der Oder. Is perhaps under-recorded.
Achtziger (1991), Frommer (1996), Remane (1987), Wagner (1935, 1939a), Walter (1998), Nickel (1997), HN

Graphocraerus ventralis (Fallén, 1806)

M V – B IX; egg, 1 gen.
In sunny, usually oligotrophic, moderately dry to damp, occasionally also moist or sa-
line sites, mainly low-input meadows and pastures, ruderal sites, grey and inland dunes,
abandoned vineyards and waysides. Feeds on various grasses (probably Anthoxanthum
odoratum, Poa pratensis and others. In field choice experiments in Britain, adults and
nymphs fed on Arrhenatherum elatius, Helictotrichon pubescens and Trisetum flavescens).
Widespread in Germany and locally common, but rather scattered in regions of inten-
184 Review of species

sive agriculture. Found up to at least 900 m a.s.l. in the Mittelgebirge, up to 2000 m a.s.l.
in the French Alps.
Achtziger (1991), Fischer (1972), Fröhlich (1996a), Kuntze (1937), Niedringhaus (1991), Nikusch (1976), Reimer
(1992), Remane (1987), Schaefer (1973), Schiemenz (1969), Schiemenz et al. (1996), Schwoerbel (1957), Wagner
(1935, 1939a, 1951a), Nickel & Achtziger (1999), HN, and others; see also Cook (1996), della Giustina (1989)

Mimallygus lacteinervis (Kirschbaum, 1868)

So far M VIII – M IX, according to Moosbrugger (1946) in Vorarlberg (Austria) E VII;
egg, 1 gen.
On low-growing willows (Salix purpurea, probably also S. eleagnos) on almost bare grav-
el banks of alpine rivers. Older reports of Hippophae rhamnoides as host plant presum-
ably refer to Salix eleagnos. Known from Germany only from the Iller near Oberstdorf,
the Isar between Mittenwald and Fall, and the Friedergries near Garmisch-Partenkir-
chen; all sites between 750 and 900 m a.s.l. The Iller population, which has been record-
ed in 1937, is probably extinct due to drainage.
Franz (1943), MTD, Nickel & Voith (unpublished data), Nickel (1999a); see also Kirschbaum (1868), Moos-
brugger (1946)

Phlepsius intricatus (Herrich-Schäffer, 1838)

B VIII – B V; adult (only ‡‡?), 1 gen.
This species is reported to occur from central Asia to the Mediterranean region, known from
Germany only from several records in the Kaiserstuhl (Badberg near Oberbergen, c. 400 m
a.s.l.), the latest one dating from 1977. Lives in xerothermic grassland sites with scattered
shrubs; adults are at least occasionally swept from shrubs, but the food plants are unknown.
Remane (pers. comm.), Wagner (1963), ZIMH

Phlepsius ornatus (Perris, 1857)

So far E VII – M IV; adult, 1 gen.
In Germany known only from altogether 4 localities in the Nahe valley (Lemberg, Bad
Münster, Ebernburg, Schloßböckelheim, all between c. 150 and 300 m a.s.l.) in xerother-
mic, acidic porphyry sites with scattered shrubs. Most adults were swept in low-grow-
ing herbaceous vegetation, a few also from shrubs. The food plants are unknown.
Jöst (1966), Wagner (1963), HN

Rhytistylus proceps (Kirschbaum, 1868)

E VI – M X, according to Schiemenz et al. (1996) from B VI onwards; egg, 1 gen.
In sparse and low-growing stands of grasses, in sunny, moderately dry to dry sites (main-
ly sandy dry grassland, heaths, open margins of pine forests, occasionally chalk and
sandstone hillsides, etc.). The main host plant is Festuca ovina; additional species report-
ed in the literature include Corynephorus canescens, Nardus stricta and Carex arenaria, but
 Deltocephalinae Fieber, 1869 185

these are probably erroneous. The species is widespread and fairly common in sandy
areas of the north German plain, but more localized to the south of it, being largely
restricted to areas of aeolian sand in larger river valleys in southern parts (along the
upper Rhine, the Main and the Danube); the highest localities are at c. 500 m a.s.l. only.
Niedringhaus (1991, 1997), Post-Plangg & Hoffmann (1982), Remane (1987), Schiemenz (1969), Schiemenz
et al. (1996), Wagner (1935, 1939a), Nickel & Remane (1996), HN; see also Ossiannilsson (1983)

Hardya melanopsis (Hardy, 1850)

Is reported from mountainous regions of western and southern European countries (Nast 1987),
also from arctic and subarctic parts of Siberia (Anufriev & Emelyanov 1988), although those
population have been described as H. taimyrica Vilbaste, 1969. In general, the taxonomy of Euro-
pean populations may be critical.
So far only B VIII, in England according to Le Quesne (1969) VII – V; adult, 1 gen.
Known in Germany only from a single site in the upper montane belt of the southern
Black Forest: Schauinsland, Steinwasen, 1100 m a.s.l., 05.VIII.1997, a large population
(including nymphs) in a pasture with scattered rocks and spruce trees. An older pub-
lished record from Rheinhessen refers to the following species. In Britain the species has
been recorded on heathland sites with dominating Festuca ovina and F. rubra.
HN; see also Le Quesne & Morris (1971), Wagner (1939a, 1955)

Hardya signifer (Then, 1897)

B VII – B V; adult, 1 gen.
On xerothermic hillsides with open grassy vegetation and scattered trees or shrubs,
usually (perhaps exclusively) on species of fescue belonging to the Festuca ovina group.
Known from Germany only from a few isolated sites, on gypsum, Jurassic limestone
and porphyry in warm areas of southern and eastern parts, all between 150 and 450 m
a.s.l. (Kyffhäuser, valleys of the Altmühl and Nahe), although locally in high abun-
dance. Originally described from alpine habitats of Salzburg state (Austria), and also
reported from the alpine belt of Switzerland and France (Alps and Pyrenees). Other-
wise published from Lithuania, Switzerland, Spain, Italy, the former Yugoslawia and
Bulgaria. Some of these records, however, should be confirmed.
Remane (1961b), Schiemenz (1969), Wagner (1955), HN; see also della Giustina (1989), Günthart (2000),
Then (1897)

Hardya tenuis (Germar, 1821)

Specimens currently assigned to this taxon in Germany belong to two different species, one of
which is of unknown identity. The latter is mainly found in southwestern limestone regions.
The characters of ecology and distribution given below only refer to H. tenuis (Germ.) sensu
Ribaut (1952).
Mainly M VII – X, ‡‡ occasionally VI; adult(?), 1 gen.
Among sparse vegetation in moderately shady, moderately dry to dry, usually acidic
and sandy, rarely basic sites. Mainly found along forest margins and under solitary
186 Review of species

trees (notably Pinus and Quercus) in heathland, occasionally in abandoned vineyards

and dry grassland, often confined to small patches. Host plants are grasses (Festuca
ovina, probably also Poa nemoralis, Agrostis capillaris and others). In Germany only very
scattered, mainly in large pine forest areas of northern parts, as well as in inland dune
areas (northern upper Rhine plain, Danube, Main and Regnitz valley, Mittelfranken),
locally also on sandstone and limestone in warm situations (valleys of the Fränkische
Saale, Main and Naab), recorded up to 400 m a.s.l. Has apparently vanished from some
former localities; most records date back several decades. In addition, some published
records are based on misidentified specimens.
Reimer (1992), Schiemenz et al. (1996), Trümbach (1959), Wagner (1935, 1939a), HN; see also Niedringhaus
(1991), Wagner (1941b)

Sardius argus (Marshall, 1866)

Thamnotettix aphrodoides Haupt, 1935 described from near Bonn is likely to be a junior synonym,
although the type material could not be found during a revision of Haupt’s collection.
E V – B X, egg(?), 2 gen.; according to Le Quesne (1969) in England III, IV and VII – XI.
This species’ range includes the Mediterranean region and parts of western Europe. It
lives in oligotrophic and acidic, usually moderately dry sites, preferentially in grazed
heathland. Host plants are grasses, perhaps Agrostis capillaris or Danthonia decumbens.
Uncommon in Germany, at the eastern edge of the range, only recorded from Rhein-
land-Pfalz and Hessen, as well as near Aschaffenburg and probably Bonn (from the
latter location published as Th. aphrodoides Haupt, 1935). Highest localities are at c. 350 m
a.s.l. only. Has apparently vanished from a number of sites during recent decades.
Haupt (1935), Kirschbaum (1868), Post-Plangg & Hoffmann (1982), Remane & Fröhlich (1994b), Wagner
(1951a), HN; see also Cobben & Rozeboom (1983), Le Quesne (1969)

Paluda flaveola (Boheman, 1845)

E VI – M X; egg, 1 gen.
Among tall grasses in sunny to moderately shady, peaty to damp, usually acidic and
rather cool sites (mainly clearings, open forests, fens, intermediate bogs, margins of
raised bogs, abandoned meadows and ruderal sites; apparently absent from mown grass-
land). Host plants are species of small-reed (Calamagrostis epigejos, C. canescens, C. vil-
losa, C. arundinacea), but also further grasses (perhaps Deschampsia cespitosa, Phalaris
arundinacea, Molinia caerulea and others). Widespread in the northeastern half of Germa-
ny and fairly common in favourable sites (up to at least 900 m a.s.l.), but apparently at
the edge of the range; southwesternmost localities are known from the Eifel and Rhön
Mountains, Mainfranken, the Black and Bavarian Forest. Is not reported from Austria,
Switzerland and France.
Niedringhaus & Olthoff (1993), Reimer (1992), Remane (pers. comm.), Remane & Fröhlich (1994b), Schie-
menz (1971a), Schiemenz et al. (1996), Wagner (1941a), Nickel & Remane (1996), HN; see also della Giusti-
na & Remane (2001)
 Deltocephalinae Fieber, 1869 187

Rhopalopyx adumbrata (C. Sahlberg, 1842) sensu Vilbaste (1962)

Was considered to be conspecific with Rh. preyssleri (H.-S.) for a long time, until Vilbaste (1962)
pointed out that there are pronounced differences in the male genitalia and in habitat preferenc-
es, although without studying the type material. Concerning life history, both species are simi-
lar in phenology, distribution and habitat requirements. However, there are differences in mois-
ture requirements (despite some overlap) and host choice (see below). Thus they may co-occur
in damp sites, but on different plants.
B VII – B X; egg, 1 gen.
In usually low-growing stands of grasses in oligotrophic, moderately wet to moderate-
ly dry, sunny to moderately shady, acidic to basic sites, mainly low-productivity mead-
ows and pastures, heathland and calcareous hillsides. Definite host plant records all
refer to Festuca rubra and, less commonly, F. ovina. Widespread in Germany, but at the
edge of the range; westernmost localities are on the East Frisian Islands, in the Eifel
Mountains and the Black Forest. The species is also reported from England and the
French Alps, where it is found up to 2000 m a.s.l., although highest localities in the
German Mittelgebirge and the Bavarian Alps are only at 1500 m a.s.l.
Niedringhaus & Olthoff (1993), Reimer (1992), Remane & Fröhlich (1994b), Rombach (1999b), Nickel &
Achtziger (1999), Nickel (1997), HN; see also Le Quesne (1969)

Rhopalopyx preyssleri (Herrich-Schäffer, 1838) sensu Vilbaste (1962)

See taxonomic notes on the preceding species.
B VI – M X; egg, 1 gen.
Usually in sunny, occasionally also slightly shaded grass stands in damp to dry, basic as well
as acidic sites, mainly in disturbed habitats (ruderal sites, abandoned fields and vineyards,
waysides, forest clearings, dry grassland, etc.), locally syntopic with the preceding species. Is
often found among Calamagrostis epigejos, Brachypodium pinnatum and Elymus repens, which
have been suggested to be the host plants, but apparently, the species lives exclusively on Poa
pratensis angustifolia. Widespread in Germany, but often found in low numbers only. Highest
localities are at 800 m a.s.l. in the foothills of the Alps and on the Swabian Jura.
Achtziger (1991), Frommer (1996), Niedringhaus (1991), Reimer (1992), Schiemenz et al. (1996), Nickel
(1997), HN

Rhopalopyx elongata W. Wagner, 1952

This species, bearing a diagnostic pygophor, was already shown by Ribaut (1936) as Thamnotet-
tix vitripennis (Fl.), after specimens collected in the French Roussillon region. The type series of
Jassus vitripennis Flor, 1861, however, included specimens both from Latvia and southern France,
which later turned out to belong to different species. Thus, after strict interpretation of nomen-
clatural rules, the species’ name would have been fixed by Ribaut. Later Wagner (1952b), who
was not aware of that, and who used the name vitripennis also for the central and north Europe-
an populations, found out that there were two distinct species and described Rh. elongatus W.Wg.
(locus typicus in Spain). In order to avoid further complicating of the nomenclatural situation,
Ribaut (1959) did not insist on the priority of his interpretation. This opinion is adopted here.
The grammatical gender of Rhopalopyx is female (see Ribaut 1939).
So far E VI – M X; egg, 2(?) gen.
188 Review of species

Occurs mainly in southwestern parts of Europe, but has recently been found in 4 local-
ities in the northern Eifel Mountains and the lower Rhine region (near Mechernich, Bad
Münstereifel and Cologne, all between 50 and 350 m a.s.l.) in rather dry and sun-ex-
posed sites on limestone and sand. The host plants are apparently grasses. No further
details are known on the life history.
Frommer (pers. comm.), Rombach (1999a); see also Nast (1987)

Rhopalopyx vitripennis (Flor, 1861)

Is usually considered to be a much variable taxon comprising a number of varieties, which were
formerly treated as distinct species. These varieties include Rh. parvispinus W. Wagner, 1948, Rh.
monticola Ribaut, 1939, Thamnotettix graminis Haupt, 1935, and Th. andropogonis Haupt, 1924 (see
Wagner 1967). See also taxonomic notes on the preceding species.
E V – M X; egg, 2 gen.
In oligotrophic, sunny, moderately dry to dry, usually short or sparsely vegetated sites
on various substrates. Host plants are fine-leaved species of fescue (Festuca ovina group,
perhaps also F. rubra). Widespread in the lowlands of Germany, often among the domi-
nant species in the sand areas of the north German plain and in warmer regions of
southern and eastern parts, but rather sporadic elsewhere. Essentially a species of low
altitudes, but occasionally found up to 700 m a.s.l. in the Mittelgebirge and the foreland
of the Alps, up to 1050 m a.s.l. in Switzerland. However, small individual numbers are
highly vagile; thus reproduction should be ascertained in submontane and montane
localities.
Fischer (1972), Marchand (1953), Reimer (1992), Remane (1958, 1987), Schiemenz et al. (1996), Struve (1939),
Trümbach (1959), Wagner (1935, 1939a, 1951a), Walter (1998), HN; see also Günthart (1987b)

Elymana kozhevnikovi (Zachvatkin, 1938)

M VII – E IX; egg, 1 gen.
Among tall grassy undergrowth of open forests, mainly under beech, pine, spruce and oak;
host plants are several species of small-reed (Calamagrostis arundinacea, C. varia, C. canescens,
but not C. epigejos). This is an eastern species reaching its distribution limit in Germany. So
far it has been found in mountain forests and pine heaths of the Bavarian Alps and their
foothills, the Harz Mountains, Elbsandsteingebirge and Erzgebirge, in low-input forests
and coppice-with-standards in Thuringia and Unterfranken, as well as in an alder fen near
the southwestern edge of the Harz Mountains, between 250 and 1400 m a.s.l.
Emmrich (1986), Fischer (1972), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Nickel (1994), HN; for
identification of nymphs see Dmitriev (1999), Walter (1978)

Elymana sulphurella (Zetterstedt, 1828)

E VI – M X; egg, 1 gen. (see also Witsack 1985).
Eurytopic in grass stands of wet to moderately dry, sunny to moderately shady sites,
mainly in low-input meadows and pastures, abandoned fields, clearings, shady patch-
 Deltocephalinae Fieber, 1869 189

es in dry grassland and heaths, open forests, waysides, and moderately saline sites
along the coast and inland. Often among the dominant species in low-input grassland
of the submontane belt, but in regions of more intensive agriculture of lower altitudes,
it is largely restricted to margins along ditches and waysides. Lives on various grasses,
notably Calamagrostis epigejos, C. arundinacea, C. varia, Holcus mollis, H. lanatus, Elymus
repens and Brachypodium pinnatum. Widespread and common in Germany, frequently
recorded up to 1500 m a.s.l. in the Bavarian Alps, at least single specimens were found
at 2000 m a.s.l.
Achtziger (1991), Fischer (1972), Fröhlich (1996a), Nikusch (1976), Reimer (1992), Remane (1987), Schie-
menz et al. (1996), Wagner (1935), Nickel & Achtziger (1999), HN, and others; see also Günthart (1987a),
Morcos (1953); for identification of nymphs see Dmitriev (1999), Walter (1978)

Cicadula albingensis W. Wagner, 1940

Mainly M VII – B X, occasionally B VI – E X; egg, 1 gen.
In wet to peaty, meso- to moderately eutrophic, sunny to moderately shady sites (main-
ly in spring mires and along headwaters of streams, also in fens and intermediate bogs,
straw and peaty meadows). The preferred host plant is Scirpus sylvaticus, but there are
also a few records from tall-sedge stands (Carex acuta, C. acutiformis and others). In Germa-
ny near the western and southern edge of the range; westernmost localities are near
Freiburg, Pirmasens, Koblenz, Marburg, Göttingen and Hamburg (locus typicus). Has
been found in France and Belgium, but is not reported from the Netherlands, Switzer-
land and Italy. Widespread in Germany, locally common at submontane altitudes, but
rather scattered in the lowlands. Highest localities are at 1100 m a.s.l. in the Bavarian
Alps and the Black Forest.
Fischer (1972), Fröhlich (1996a), Heller (1987a), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer
(1992), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Trümbach (1959), Wag-
ner (1940b, 1951a), Nickel & Achtziger (1999), Nickel (1997), HN

Cicadula rubroflava Linnavuori, 1952

Some authors (e.g. Ossiannilsson 1983) use the name C. longiventris (J. Sahlberg, 1871) for this
species. However, this name is based on the description of ‡‡, which could not be identified
with certainty until Remane & Fründ (1986). Therefore, Linnavuori’s name should be kept until
the type material has been revised (see Remane & Fröhlich 1994b).
B VIII – M X; egg, 1 gen.
On Carex brizoides in forests and forest glades on seepy or waterlogged substrates, usu-
ally in stands of spruce and pine in cool situations. In Germany only very sporadic, at
the western and southern edge of the range; at present there are only 15 known locali-
ties, mostly in Saxony (Elbsandsteingebirge, Oberlausitz, Dresdener Heide, Vogtland)
and the foothills of the Alps (Ammersee area, near Kelheim and Biberach), also in the
Bavarian Forest (near Freyung), Vogelsberg (Waldensberg) and Main valley (Volkach),
between 200 and 750 m a.s.l.
Emmrich (1989), Remane & Fröhlich (1994b), Remane & Fründ (1986), Schönitzer & Oesterling (1998b),
Walter (pers. comm.), HN
190 Review of species

Cicadula persimilis (Edwards, 1920)

B VI – E X; egg, 2 gen., probably 1 at higher altitudes.
On Dactylis glomerata in sunny to slightly shady, moist to moderately dry sites, notably
moderately eutrophic meadows and pastures, abandoned fields, ruderal sites, way-
sides, etc. In Germany widespread and common in most regions, although less com-
mon and confined to field margins in regions of more intensive agriculture. Frequently
occurs up to at least 1800 m a.s.l. in the Bavarian and Allgäu Alps.
Achtziger (1991), Fischer (1972), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz et al. (1996),
Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others; see also Morcos (1953)

Cicadula quinquenotata (Boheman, 1845)

E VII – E X; egg, 1 gen.
In intermediate bogs of dune valleys and moraine depressions. The host plant is not
known, most authors suggest an unidentified species of Carex or Eriophorum. In Germa-
ny the species is confined to the northern plains; southernmost localities are on the East
Frisian Islands, near Hamburg, Lauenburg, Waren an der Müritz, and Luckenwalde.
All German sites are below 100 m a.s.l., but the species is reported from altitudes above
1500 m a.s.l. in the Austrian Alps.
Kuntze (1937), Niedringhaus (1991), Remane & Fründ (1986), Schiemenz et al. (1996), Wagner (1935, 1937e);
see also Holzinger (1999b), Ossiannilsson (1983), Wagner & Franz (1961)

Cicadula saturata (Edwards, 1915)

B VII – M X; egg, 1 gen.
In wet, usually peaty and acidic sites (intermediate bogs, fens, floating mats of Sphag-
num, peaty meadows); host plants are sedges, mainly Carex nigra, perhaps also C. rostra-
ta. In Germany at the western and southern edge of the range, with strongholds in
rather cool areas of the north German plain, the Mittelgebirge and the Bavarian and
Swabian foothills of the Alps, found up to 1500 m a.s.l. in Upper Bavaria. Not reported
from Austria, Switzerland, Italy and Belgium, but from the Netherlands and eastern
parts of France.
Bittner & Remane (1977), Fröhlich (1996a), Hoffmann (1980), Nikusch (1976), Reimer (1992), Schiemenz et
al. (1996), SMNS, Wagner (1935, 1946), Nickel & Achtziger (1999), Nickel (1997, 1999b, 2002); see also Cobben
& Gravestein (1958), della Giustina (pers. comm.), den Bieman (pers. comm.), Nast (1987), Ribaut (1952)

Cicadula flori (J. Sahlberg, 1871)

Mainly E VI – E X, in some years from E V onwards; egg, 2 gen.
In stands of tall sedges in wet to temporarily flooded, usually moderately eutrophic sites,
mainly fens and straw meadows, occasionally also in fen woods, often syntopic with C.
quadrinotata (F.) and locally dominant, but more stenotopic and restricted to rather warm
situations. Host plants are tall sedges (preferentially Carex acuta, perhaps also C. acutifor-
mis, C. elata and additional species). Published records from central Europe are sparse,
 Deltocephalinae Fieber, 1869 191

but the species is apparently widespread in most lowland areas of northern and middle
Germany (e.g. Holstein, southern and eastern Niedersachsen, Thuringia, Hessen, Franco-
nia). Highest localities are at 500 m a.s.l. in the Bavarian foreland of the Alps.
Hildebrandt (1995), Niedringhaus (1991), Reimer (1992), Remane (1958), Schiemenz et al. (1996), Trümbach
(1959), Wagner (1946), Weis & Schönitzer (2001), Nickel & Achtziger (1999), HN; see also Lauterer (1986)

Cicadula quadrinotata (Fabricius, 1794)

B VI – E X; egg, 2 gen., 1 at higher altitudes.
Eurytopic in peaty (also temporarily flooded) to moist, locally also moderately dry, sunny
to moderately shady sites, often dominant in fens, intermediate bogs, low-input mead-
ows, along ditches, and in peat diggings in raised bogs, also found in ruderal sites, mod-
erately saline marshes and fen woods. In regions of more intensive agriculture, however,
the species is often restricted to ditches and other margins. Host plants are Carex acutifor-
mis, C. acuta, C. rostrata, C. nigra, C. elata, C. disticha, C. brizoides, C. hirta and others; few
individuals have also been found on Bolboschoenus maritimus and Eriophorum angustifoli-
um. Widespread in Germany and among the dominant species in many wetland habitats,
although population sizes are likely to have decreased due to draining and intensification
of grassland management. Occurs up to at least 1500 m a.s.l. in the Allgäu Alps, but up to
2400 m a.s.l. in North Tyrol (Austria) and the Engadine (Switzerland).
Bittner & Remane (1977), Fischer (1972), Marchand (1953), Reimer (1992), Schiemenz (1971a), Schiemenz et al.
(1996), Wagner (1935), Nickel & Achtziger (1999), HN; see also Günthart (1987a), Leising (1977), Novotný (1995)

Cicadula frontalis (Herrich-Schäffer, 1835)

Mainly M VII – M X, in southern parts and in warm years from B VI onwards; egg, 1
(-2?) gen.
In sunny to moderately shady, wet to temporarily flooded, usually moderately eutrophic
sites, mainly fens and shores of ponds and streams, occasionally in fen woods; appar-
ently absent from mown sites. Host plants are tall-growing sedges, mainly Carex acuti-
formis and C. riparia, perhaps also C. pseudocyperus and C. acuta. Widespread in the low-
lands of Germany, but uncommon, usually in lower abundances than the remaining
species of Cicadula. Occurs up to at least 700 m a.s.l. in the foothills of the Alps, but
apparently only below 500 m a.s.l. in regions to the north of the Danube.
Frommer (1996), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oester-
ling (1998b), Wagner (1935, 1951a), HN

Cicadula ornata (Melichar, 1900)

So far only B VII – M VII, in Scandinavia VII – IX (Ossiannilsson 1983); egg, 1(?) gen.
This species’ range includes the whole boreal zone of the Palearctic and the Nearctic. In
Europe it was hitherto only known from England, Sweden, Finland, Estonia, Latvia and
Poland. Is reported to live on tall sedges in river floodplains, but no further details are
known on habitat requirements. From Germany there is only a single record from suburban
192 Review of species

areas of Berlin: Biesdorf, c. 50 m a.s.l., July 1983, 6 ††, 4 ‡‡, at light, formerly published as C.
intermedia (Boheman, 1845). During a recent revision, these specimens turned out to belong
to the closely related C. ornata (Mel.). Further old records of C. intermedia (Boh.) from north-
western parts of Germany referred to C. quinquenotata (Boh.) and have been corrected later.
Hence, the occurrence of C. intermedia (Boh.) in Germany is not proven.
Emmrich & Nickel (unpublished data), Schiemenz et al. (1996), Wagner (1935, 1937e); see also Flint (1990),
Nast (1987), Ossiannilsson (1983), Vilbaste (1974)

Mocydia crocea (Herrich-Schäffer, 1837)

Adults I – XII, mainly B VIII – B VI; adult, 1 gen. (see also Müller 1976).
In sunny to moderately shady, dry to temporarily moist sites on basic to acidic sub-
strates, usually in dry grassland, ruderal sites, and along waysides, locally also in straw
meadows, temporarily dry salt marshes and open, thermophilous forests of oak or pine.
Host plants are Brachypodium pinnatum, Bromus erectus, Calamagrostis epigejos, Elymus
repens and other tall grasses. In Germany at the northern edge of the range; widespread
and locally common in middle and southern parts, but rather localized in the north
German plain; recorded northward to Cologne, Hanover, Wolfsburg, Magdeburg and
Neubrandenburg. Usually found below 500 m a.s.l., except in the higher limestone re-
gions (Bavarian Alps, Swabian Jura) where occurring up to at least 1000 m a.s.l.
Achtziger (1991), Fischer (1972), Fröhlich (1996a), Frommer (pers. comm.), Post-Plangg & Hoffmann (1982),
Reimer (1992), Remane (1987), Rombach (1999a), Schiemenz et al. (1996), Schwoerbel (1957), Nickel &
Achtziger (1999), Nickel (1994), HN, and others

Mocydiopsis attenuata (Germar, 1821)

E VII – V; adult, 1 gen.
On species of fescue (Festuca ovina group, F. rubra, F. heterophylla) usually in moderately
shady, damp to moderately dry, acidic as well as basic sites; mainly along margins of
forests of pine, oak and beech (occasionally also in their interior), as well as in low-pro-
ductivity grassland with scattered trees or shrubs. The distribution in Germany (as well
as in other parts of Europe) is poorly documented due to identification problems; howev-
er, the species is apparently widespread (although not yet recorded from Mecklenburg-
Vorpommern and Baden-Württemberg), but densities are usually low. Most localities are
below 400 m a.s.l., but there is a single record from the foothills of the Alps at 750 m a.s.l.
Frommer (1996), Niedringhaus (1991), Nikusch (1976), Remane (1961a, 1987), Schiemenz et al. (1996), HN

Mocydiopsis intermedia Remane, 1961

E VII – V; adult, 1 gen.
In moderately dry to dry, moderately shady sites on basic to moderately acidic sub-
strates, usually in xerothermic grassland with scattered trees or shrubs and along sun-
ny forest margins, probably associated with Poa pratensis angustifolia. In Germany this
species occurs at the northern edge of the range and is largely confined to warmer low-
 Deltocephalinae Fieber, 1869 193

land regions between the Danube and the northern edge of the Mittelgebirge (mainly
along the upper and middle Rhine, Franconian Jura, Mainfranken, Saale-Unstrut re-
gion), up to at least 550 m a.s.l. Border localities are near Bonn, Marburg, Bad Neustadt
an der Saale, Bad Frankenhausen, Quedlinburg, Halle and Leipzig. It is not reported
from Belgium, the Netherlands, the British Islands, northern Europe and Poland. Has
apparently disappeared in a number of localities.
Funke & Witsack (1998), Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1961a,
1987), Remane & Fröhlich (1994b), Schiemenz et al. (1996), HN

Mocydiopsis longicauda Remane, 1961

B VII – V; adult, 1 gen.
In moderately dry to dry, basic to acidic sites, usually in the vicinity of trees or shrubs
(mainly dry grassland, meadows with scattered fruit trees, forest margins, etc.). Host
plants are fine-leaved species of fescue (Festuca ovina group). A tongue-shaped part of
the range extends from southeastern Europe into Germany, with strongholds in Thurin-
gia, Mainfranken and the Franconian Jura. Border localities are near Regensburg, Eich-
stätt, Würzburg, Bad Neustadt an der Saale, Gotha, Bad Frankenhausen, Halberstadt,
Haldensleben, Dessau and Gera, at least up to 500 m a.s.l.
Müller (1978), Reimer (1992), Remane (1961a), Remane & Fröhlich (1994b), Schiemenz et al. (1996), HN

Mocydiopsis monticola Remane, 1961

E VII – V; adult, 1 gen.
On Holcus mollis in damp to moist, moderately shady, usually acidic sites, mainly open
oak forests, clearings, coppice-with-standards, forest margins, etc. In Germany only
known from the middle parts, between the Danube and the northern edge of the Mittel-
gebirge, up to at least 400 m a.s.l., although probably under-recorded. Northernmost
localities are near Cologne, Frankenberg an der Eder, Artern and Görlitz. The species is
neither reported from more northern parts of Europe, nor from Belgium, the Nether-
lands, Poland and the Czech Republic, but it is probably overlooked or misidentified.
Frommer (pers. comm.), Remane (pers. comm.), Remane & Fröhlich (1994b), HN; for identification see
Remane (1961a)

Mocydiopsis parvicauda Ribaut, 1939

E VII – IV; adult, 1 gen.
On Agrostis capillaris in oligotrophic and acidic, sunny to moderately shady, moderately
moist to dry sites (mainly pastures and heaths, also clearings, forest margins, coppice-
with-standards, etc.). Widespread in western parts of Germany, but usually at low den-
sities, more scattered towards the east, where apparently at the northeastern edge of the
range. Border localities are on a line from Kiel to Hamburg, Stendal, Leipzig and Hoy-
erswerda. The species is found up to at least 600 m a.s.l. in the Mittelgebirge and the
foreland of the Alps.
194 Review of species

Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer (1992), Remane (1961a.),
Schiemenz et al. (1996), HN

Speudotettix subfusculus (Fallén, 1806)

This is a rather polymorphic species with much varying body size, wing length and colouration
intensity. Variation is particularly pronounced in upper montane and subalpine populations of
the Alps. Speudotettix montanus Gȩbicki & Szwedo, 1991, described on the basis of merely two
specimens from a mountain site in southeastern Poland, is likely to be within this range of
variation and should be revised.
Mainly E IV – M VIII, at higher altitudes until M IX; nymph, 1 gen.
Eurytopic in deciduous and coniferous forests and along their margins, in wet to mod-
erately dry, basic to acidic sites, also among subalpine scrub and in alpine grassland.
Nymphs are found among low vegetation (often on sedges and grasses); most adults
feed on various deciduous trees and shrubs (Betula, Corylus, Alnus, Populus, Prunus and
others), probably on dwarf shrubs above the alpine tree line. Widespread and common
in Germany, also in higher parts of the Mittelgebirge and the Bavarian and Allgäu Alps,
where frequently encountered up to 2000 m a.s.l.; in Switzerland even found up to 2400
m a.s.l.
Achtziger (1991), Fischer (1972), Frommer (1996), Niedringhaus (1997), Reimer (1992), Remane (1987), Schi-
emenz et al. (1996), Schwoerbel (1957), Wagner (1935, 1939a), HN, and others; see also Günthart (1987a)

Hesium domino (Reuter, 1880)

E VI – B X; egg, 1 gen.
In wet to moderately dry, basic to acidic sites with semi-open stands of deciduous trees
(fens, intermediate bogs, low-productivity meadows and pastures, dry grassland, for-
est margins and glades, clearings, etc.). Adults usually on Betula pendula and B. pubes-
cens, occasionally on Alnus, Ulmus and other trees, nymphs in low-growing herbaceous
vegetation, probably feeding on grasses (Festuca rubra, Holcus mollis and others). Wide-
spread in Germany, except in northwestern parts, and in intermediate frequency, but
low individual numbers. Apparently absent from the northwest German plain, the Neth-
erlands, Denmark and the British Isles; border localities are near Bonn, Siegen, War-
burg, Northeim, Goslar, Hamburg and Rostock. Recorded up to at least 1200 m a.s.l. in
the Mittelgebirge and Bavarian Alps, up to 1750 m a.s.l. in the French Alps.
Achtziger (1991), Fischer (1972), Jöst (1966), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Wagner
(1935, 1939a), HN, and others; see also della Giustina (1989)

Thamnotettix confinis (Zetterstedt, 1828)

Mainly M V – M VIII, at montane and subalpine altitudes VI – IX; nymph, 1 gen.
Polyphagous in deciduous and coniferous forests of damp to moist, acidic to basic sites,
as well as among subalpine scrub. Nymphal development takes place in low-growing
herbaceous vegetation; at least some adults ascend various deciduous trees or shrubs
(Quercus, Betula, Rubus and others). Widespread all over Germany, found up to at least
 Deltocephalinae Fieber, 1869 195

1800 m a.s.l. in the Bavarian and Allgäu Alps, up to 2200 m a.s.l. in North Tyrol (Aus-
tria) and the Engadine (Switzerland).
Fischer (1972), Frommer (1996), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Schwoerbel (1957),
Wagner (1935, 1939a), HN; see also Günthart (1987a), Leising (1977)

Thamnotettix dilutior (Kirschbaum, 1868)

B V – B X; nymph, 1 gen.
In open oak forests and along their margins, in moderately dry to damp, acidic as well
as basic sites. Nymphs are found among low-growing herbaceous vegetation (on Des-
champsia flexuosa and other grasses), occasionally on shrubs, most adults on oaks (Quer-
cus robur and Qu. petraea). Widespread in Germany, but restricted to rather low altitudes
(found up to 400 m a.s.l. only, but reported to occur at 800 m a.s.l. in Ticino, Switzer-
land). Locally common in warmer regions in southern and eastern parts (often abun-
dant in Malaise traps), but more localized and less common towards the north, not
recorded to the south of the Danube.
Post-Plangg & Hoffmann (1982), Niedringhaus (1997), Remane (1987), Schiemenz et al. (1996), Wagner
(1935, 1939a, 1951a), HN; see also Günthart (1987b)

Pithyotettix abietinus (Fallén, 1806)

E V – M IX, single individuals until XI; nymph, 1 gen.
In coniferous and mixed forests of higher altitudes, secondarily also in lowland conifer
plantations. Nymphs and adults on Picea abies, perhaps also on Abies alba. In Germany
originally confined to the natural host range in the Alps and at higher altitudes of the
Mittelgebirge, where locally common, but nowadays also widespread in the lowlands,
even found in parks and cemeteries. However, the species is seldom recorded, since
adults usually stay in the canopy layer. Highest localities are at 1700 m a.s.l. near the
alpine tree limit of Upper Bavaria.
Fischer (1972), Frommer (1996), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b),
Schwoerbel (1957), Trümbach (1959), Wagner (1935, 1939a, 1951a), HN, and others

Perotettix pictus (Lethierry, 1880)

E V – E VII, occasionally until X; nymph, 1 gen.
Like P. abietinus (Fall.) on Picea abies and perhaps Abies alba, mainly in the canopy layer
of open forests, along forest roads and edges. Less common than the preceding species
and more restricted to forest areas of higher altitudes; in Germany only found between
300 and 1700 m a.s.l., at the northern edge of the range; recorded from the Alps and
their foreland, the Black Forest, the Erzgebirge, the Rhön Mountains, middle Hessen,
Thuringia, the Rothaargebirge and the Harz Mountains.
Reimer (1992), Remane (pers. comm.), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Schönitzer &
Oesterling (1998b), SMNS, Nickel & Remane (1996), HN
196 Review of species

Colobotettix morbillosus (Melichar, 1896)

M V – E VIII, single individuals until IX; nymph, 1 gen.
Another species living on Picea abies and perhaps Abies alba, usually in higher situations;
most adults are swept in open forests and along their margins. In Germany probably
under-recorded, but apparently more or less restricted to the alpine foothills and the
Mittelgebirge (up to at least 1200 m a.s.l.), but occasionally found in the north German
plain. According to present data, the range is largely confined to central Europe and
includes southern Finland, France (only Savoie), Poland, Switzerland, Austria and the
Czech Republic; the species is not reported from Russia.
Fischer (1972), Reimer (1992), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Schönitzer & Oesterling
(1998b), Trümbach (1959), Wagner (1935, 1939a), HN; see also della Giustina (1989), Emelyanov (pers. comm.),
Günthart (pers. comm.), Holzinger (1996a), Lauterer (pers. comm.), Nast (1976a, 1987), Ossiannilsson (1983)

Macustus grisescens (Zetterstedt, 1828)

E IV – M VIII, at higher altitudes occasionally until IX; nymph, 1 gen.
In tall-growing stands of sedges and grasses in peaty to damp, sunny to moderately
shady sites, mainly in fens, intermediate and raised bogs, straw meadows, fen woods,
along ditches, as well as in forest clearings, abandoned fields and open forests. Host
plants are sedges (Carex acutiformis, C. paniculata, probably also C. acuta, C. rostrata, C.
nigra, and others), grasses (Calamagrostis arundinacea, C. epigejos, Festuca arundinacea,
Dactylis glomerata, Elymus repens and others) and woodrushes (Luzula spp.). Widespread
in Germany and fairly common in favourable sites, found up to at least 1900 m a.s.l. in
the Bavarian and Allgäu Alps, up to 2170 m a.s.l. in the Engadine (Switzerland).
Fischer (1972), Niedringhaus (1991), Reimer (1992), Schiemenz et al. (1996), Schwoerbel (1957), Wagner
(1935, 1939a), Nickel & Achtziger (1999), HN; see also Günthart (1987a)

Doliotettix lunulatus (Zetterstedt, 1838)

B V – B VIII; nymph, 1 gen.
In moderately shaded stands of grasses, usually in wet to moderately moist, acidic to
moderately basic sites, mainly at submontane altitudes, mainly in deciduous and conif-
erous forests and along their margins, probably associated with Agrostis stolonifera. In
Germany only localized, but fairly common in favourable habitats, with strongholds in
the Mittelgebirge (Eifel Mountains, Rothaargebirge, Vogelsberg, hilly regions of Thurin-
gia, Harz Mountains, Swabian Jura and others) and the foothills of the Alps, occasional-
ly also found in the lowlands (near Hamburg, Berlin, Wiesbaden and Landau/Pfalz).
Highest localities are at 1000 m a.s.l.
Fischer (1972), Fröhlich (1996a), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Schönitzer & Oester-
ling (1998b), Wagner (1935, 1939a), HN

Athysanus argentarius Metcalf, 1955

B VI – E IX, according to Schiemenz et al. (1996) B V – B XI; egg, 1 gen.
 Deltocephalinae Fieber, 1869 197

In stands of tall grasses in sunny, wet to moderately dry sites, mainly fens, low-input
meadows, abandoned fields, forest clearings and heaths, but also waysides and moder-
ately saline sites. Host plants are various grasses (Calamagrostis spp., Elymus repens,
Dactylis glomerata, Arrhenatherum elatius, Deschampsia cespitosa, Festuca arundinacea, Hol-
cus spp. and others). Widespread and fairly common in the lowlands of Germany, found
up to 900 m a.s.l. in the Mittelgebirge and the foothills of the Alps.
Achtziger (1991), Fröhlich (1996a), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz et al. (1996),
Schönitzer & Oesterling (1998b), Wagner (1935, 1939a), Walter (pers. comm.), Nickel & Achtziger (1999), HN

Athysanus quadrum Boheman, 1845

B VII – M X; egg, 1 gen.
In sunny, moist to wet, usually mesotrophic sites, mainly calcareous sedge fens and
non-fertilized peat and straw meadows. The host plants are unknown, but may include
dicotyledonous herbs (perhaps Lathyrus, Inula, or Filipendula). Has been found in large
parts of Germany, but most records date before 1970. Highest sites are at 600 m a.s.l. in
the Bavarian foothills of the Alps. German localities are apparently near the southern
and southwestern edge of the range. The species is reported from the Netherlands, Bel-
gium and Austria, but not from Switzerland; the French records are uncertain. Howev-
er, it has been found in the Kaiserstuhl, close to the French and Swiss border.
Fischer (1972), Marchand (1953), Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Schiemenz et
al. (1996), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN;
also della Giustina (pers. comm.), Nast (1987)

Handianus ignoscus (Melichar, 1896)

Species identities within this genus are obscured by the loss of some type material, notably of H.
flavovarius (Herrich-Schäffer, 1835) and H. procerus (Herrich-Schäffer, 1835). The species inter-
pretation in this book is adopted from Dlabola (1954) and Emelyanov (1964a).
So far B VII – B IX; egg, 1 gen.
Has been published from eastern and southeastern Europe and from several Asian coun-
tries, but the total range is unknown due to taxonomic uncertainties. Is reported to live
on dry sites on woody species of Fabaceae, mainly Cytisus (s.l.), and is known from
Germany only from 5 acidic and sandy localities: Wahrburg (near Stendal), 1925, Fal-
kenberg (to the west of Torgau), 1966, 1967, and 1973, as well as Erlangen, between 1954
and 1957. Furthermore reported from „Brandenburg“, but without details. Recently, the
species was found in another site near Erlangen: Schießplatz Tennenlohe, 09.VII.2001,
10 ††, 11 ‡‡, 04.IX.2001, 1 †, 2 ‡‡, on Cytisus scoparius in a former military training area
on aeolian sand. The highest localities are at c. 300 m a.s.l.
Schiemenz et al. (1996), Trümbach (1959), Wagner & Franz (1961), HN; see also Emelyanov (1964a), Her-
rich-Schäffer (1835b)

Stictocoris picturatus (C. Sahlberg, 1871)

E VI – E X; egg, 1 gen.
198 Review of species

In sunny and low-vegetated grassland of dry to temporarily dry, usually basic, occa-
sionally also acidic sites (sheep pastures, rocky hillsides, locally also dry waysides and
embankments, as well as straw meadows). Main host plants are woody species of Fa-
baceae, notably Ononis spinosa and Genista tinctoria, occasionally Chamaespartium sagit-
tale, perhaps also herbaceous species (Trifolium?). In Germany only scattered, at the north-
western edge of the range, with strongholds in warm areas of middle and southern
parts up to 800 m a.s.l., but recorded at 1500 m a.s.l. in Carinthia (Austria) and the Val-
ais (Switzerland). Border localities include Koblenz, Marburg, Bad Lauterberg, Quedlin-
burg, Lauenburg and Neubrandenburg.
Kupka (1925), Nikusch (1976), Reimer (1992), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Schönitzer
& Oesterling (1998b), Schwoerbel (1957), Wagner (1951a), HN; see also Prohaska (1923)

The following taxon Ophiola Edwards, 1922 has recently been placed into the genus Limotettix J.
Sahlberg, 1871 s.l. as a subgenus by Hamilton (1994). Instead of Ophiola Edw., this author prefers
Scleroracus Van Duzee, 1894 as name. The latter, however, is rejected by most European authors
as a nomen nudum (e.g. Nast 1987; Ossiannilsson 1983; Remane & Fröhlich 1994a, and others).

Ophiola cornicula (Marshall, 1866)

This species was misinterpreted for a long time, and various names have been used (notably
Jassus orichalceus Thomson, 1869, and Limotettix intractabilis Kontkanen, 1949). Kontkanen (1948)
showed that in central and northern Europe there are at least three species with distinct genita-
lia and morphometrics. Later Le Quesne (1962) studied Marshall’s type material and clarified
the nomenclatural situation.
E VI – E IX; egg, 1 gen.
In acidic, moderately dry to peaty, sunny to moderately shady sites, notably in heaths,
raised bogs (preferentially in drained patches) and open pine forests. Feeds on Calluna
vulgaris, probably also on species of Vaccinium. Widespread in Germany, but largely
restricted to areas of sand, sandstone, bogs and other acidic substrates, mainly in the
north German plain, the submontane and montane belt of the Mittelgebirge and along
the edge of the Alps; the highest localities are at 1200 m a.s.l.
Bittner & Remane (1977), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987), Schiemenz et al.
(1996), HN

Ophiola decumana (Kontkanen, 1949)

Cicada striatula Fallén, 1806 (p. H.)
Identity and species differentiation in this taxon are not satisfactorily studied. Perhaps in cen-
tral Europe there is a complex of several distinct species. Furthermore, the identity of O. ocellaris
(Lethierry, 1880) is uncertain; it has been described from France and Austria and is reported
from large parts of the western Palearctic by Nast (1972, 1987), but has not been mentioned by
Hamilton (1994) in his generic review.
E V – B X; egg, 2 gen.
In sunny, moderately dry to dry, acidic to basic sites of early successional stages, usually
on gravelly to sandy, locally also on loamy substrates, mainly mining areas, trampled
patches (preferring middle strips of dirt roads), military training areas, abandoned fields,
low-input vineyards, as well as gravel banks of alpine rivers. Host plants are Polygonum
 Deltocephalinae Fieber, 1869 199

aviculare, Rumex acetosella, and perhaps further dicotyledonous herbs; from Sweden
Vaccinium myrtillus and Calluna vulgaris have been reported. Widespread in Germany,
but only localized; most older records are in need of revision. There are recent verified
records from various parts of the north German plain, the warmer regions in eastern
and southern parts (mainly Saale-Unstrut region, valleys of Main and Regnitz, upper
Rhine plain), and the Alps (upper course of the Isar, Berchtesgaden); usually found
below 300 m a.s.l., but up to at least 900 m a.s.l. along alpine rivers.
Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Nickel (1999a), HN;
see also Ossiannilsson (1983)

Ophiola russeola (Fallén, 1826)

Has been treated by some authors as conspecific with O. plutonia (Uhler, 1877). According to
Hamilton (1994), the latter is a distinct species confined to the New World.
E VI – B X; egg, 1 gen.
On dwarf shrubs (mainly Calluna vulgaris, but also Erica carnea, Vaccinium oxycoccus and
others) in raised bogs, heaths and open pine forests, usually in acidic sites, but also on
limestone gravel along alpine rivers. Widespread in Germany and locally common in
heath and bog areas of the north German plain and the Mittelgebirge, in sandstone and
inland dune areas, as well as along the edge of the Alps. Highest localities are at 1200 m
a.s.l. in the Allgäu Alps and the Black Forest; small numbers are also reported from the
alpine belt of the Engadine (Switzerland) and Styria (Austria), but reproduction is not
proven in these altitudes.
Niedringhaus & Olthoff (1993), Reimer (1992), Schönitzer & Oesterling (1998b), Strübing (1955), HN; see
also Günthart (1987a)

Ophiola transversa (Fallén, 1826)

E VI – M IX, according to Schiemenz et al. (1996) from B VI onwards; egg, 1 gen.
The range of this species extends from Mongolia to central Europe, apparently with
isolated populations in the Alps and the Pyrenees. It inhabits oligotrophic grassland
usually in moderately dry or temporarily dry sites (mainly heaths, locally also calcare-
ous hillsides). In Germany there are only sporadic populations, mainly in sand areas of
the northeastern half; border localities are on the island of Borkum, near Fallingbostel,
Halle, Leipzig and Görlitz. Another locality in southern Thuringia, at 550 m a.s.l., ap-
pears to be isolated. Some populations from Germany (as well as from the Italian Alps
and the Pyrenees) were taken from Achillea millefolium. Recorded at 1900 m a.s.l. in the
Slovakian Tatra Mountains.
Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Wagner (1935); see
also Lauterer (1984)

Limotettix atricapillus (Boheman, 1845)

E VI – E IX; egg, 1 gen.
200 Review of species

In intermediate and raised bogs. In Germany only known from 10 sites, mostly in bog
areas of northern parts (near Plön, Lauenburg, Neubrandenburg, Berlin, Leipzig, Dres-
den), a few isolated sites are in the Lake Constance area (Bodanrück) and the Bavarian
foothills of the Alps (Oberstdorf, 850 m a.s.l., and Inzell, 700 m a.s.l.). Has vanished
from some former localities. The host plants are poorly known; Carex has been stated in
the literature. The Inzell record comprised a large population living on Rhynchospora
alba in an overgrown peat digging.
MTD, Schiemenz et al. (1996), Wagner (1941a, 1943), Wagner & Wagner (1938), Remane (1962), Nickel (1999b)

Limotettix striola (Fallén, 1806)

B VI – E X; egg, 2 gen., probably 1 in bog habitats and at higher altitudes.
In various habitats in sunny, peaty or temporarily flooded sites: in moderately saline mead-
ows, floodplain depressions, raised and intermediate bogs, also as a pioneer species in sand
and gravel pits. Host plants are various species of Cyperaceae (mainly Eleocharis, probably
also Schoenoplectus and Trichophorum). Published records from rushes are probably based on
host misidentification. Widespread in Germany and locally fairly common and abundant,
mostly recorded along the coasts of the North and Baltic Sea, in inland salt areas (notably in
Thuringia and Saxony-Anhalt), along larger river valleys, and in bog areas, rather scattered
elsewhere; found up to at least 1350 m a.s.l. in the Allgäu and Bavarian Alps.
Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Remane (1958), Schiemenz et al. (1996), Wagner (1935,
1939a), Nickel & Achtziger (1999), HN

Laburrus impictifrons (Boheman, 1852)

B VII – B X; egg, 1 gen.
This is a steppe species living monophagously on Artemisia campestris in sunny, moder-
ately dry to dry sites, usually on acidic and sandy substrates. In Germany largely con-
fined to the northeastern half, at the edge of the range, with strongholds in sand areas of
Mecklenburg-Vorpommern, Brandenburg, Saxony-Anhalt and Saxony; otherwise only
known from isolated sites along the lower course of the Elbe, on gypsum hillsides in
Thuringia, and inland sand dunes along the Rhine (Mainz, Darmstadt, Heidelberg),
Main (Aschaffenburg, Karlstadt, Volkach, Bamberg) and Danube (Kelheim); the highest
localities are at 350 m a.s.l. only. Furthermore, there are a few 19th century records from
southwestern France (Gironde, Landes).
Remane (1987), Schiemenz et al. (1996), SMNS, Wagner (1935, 1939a, 1951a), Walter (pers. comm.), Nickel
(1997), HN; see also Ribaut (1952)

Laburrus pellax (Horváth, 1903)

B VI – B X; egg, 1 gen.
Monophagous on Aster linosyris in dry grassland in sunny, usually basic sites, mainly
on limestone and gypsum, occasionally porphyry. This is another xerothermophilous
steppe species, its range extends from central Asia to southern parts of Germany. From
 Deltocephalinae Fieber, 1869 201

there it is known from scattered localities in the Nahe and upper Rhine valley (also on
the French side), southern Palatinate, Mainfranken, the Tauber valley, southern parts of
the Franconian Jura, and the Kyffhäuser, up to 400 m a.s.l. only.
Remane & Wachmann (1993), Schönitzer & Oesterling (1998b), SMSN, Wagner (1939a, 1951a), Nickel (1999b),
HN; see also della Giustina & Remane (2001)

Euscelidius schenckii (Kirschbaum, 1868)

B VII – M X, according to Schiemenz et al. (1996) from M VI onwards; egg, 1 gen.
In disturbed, moist to moderately dry, sunny to moderately shady sites, mainly ruderal
habitats, tall herb stands (preferentially Urtica dioica), and disturbed patches in dry grass-
land and vineyards, apparently polyphagous on various dicotyledonous herbs. Wide-
spread in Germany, but seldom recorded due to secretive life habits close to the ground;
the highest localities are at 500 m a.s.l. only. The locus typicus is Weilburg an der Lahn.
Achtziger (1991), Hildebrandt (1986), Reimer (1992), Schiemenz et al. (1996), Trümbach (1959), Wagner
(1951a), HN

Euscelidius variegatus (Kirschbaum, 1858)

B VII – M X; egg, 1 gen.
Like the preceding species in disturbed sites, but – despite broad overlap – preferring
warmer and drier situations and more sandy substrates, mainly ruderal habitats, aban-
doned fields, sand and gravel pits, waysides, vineyards, and even gardens. Many records
comprise only single individuals, which may have been on dispersal flight; thus, some
published data on habitat requirements and food plants are perhaps uncertain. The
host plants are unknown; presumably, the species lives polyphagously on various di-
cotyledonous herbs. In Germany it is largely confined to warm regions in southern and
eastern parts (mainly upper and middle Rhine valley and its larger tributaries, the rain
shadow area to the east of the Harz Mountains), at the northwestern edge of the range.
Border localities are on a line from Cologne to Gießen, Bad Frankenhausen, Halberstadt
and Berlin; the highest sites are at 350 m a.s.l. only. The locus typicus is Mombach near
Mainz. Is reported as a vector of Clover phyllody (CP) from France.
Frommer (1996), Remane (1987, and pers. comm.), Rombach (1999a), Schiemenz et al. (1996), SMNS, Wag-
ner (1939a, 1951a), HN; see also Brčák (1979)

Conosanus obsoletus (Kirschbaum, 1858)

B VII – M X, according to Schiemenz et al. (1996) M V – M XI; egg, 1 gen.
In sunny, peaty to damp or temporarily waterlogged sites, usually in fens, low-input
meadows and pastures, and salt marshes, also found in intermediate bogs, forest clear-
ings, ruderal sites, and along waysides and ditches. Host plants are rushes (Juncus ef-
fusus, J. filiformis, J. inflexus, J. gerardii and others) and tall grasses (Festuca arundinacea,
perhaps Elymus repens, Arrhenatherum elatius and others). Widespread in Germany and
fairly common, the highest localities are at 1200 m a.s.l. in the Black Forest.
202 Review of species

Bittner & Remane (1977), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Reimer (1992), Remane &
Wachmann (1993), Schiemenz (1971a), Schiemenz et al. (1996), Wagner (1935, 1939a), Nickel & Achtziger
(1999), HN, and others

Euscelis distinguendus (Kirschbaum, 1858)

E VI – B X; egg, 1 gen.
In sunny, oligotrophic, basic to acidic sites; in the Bavarian and Allgäu Alps and the
Black Forest mainly found in temporarily moist to moderately dry low-productivity
pastures and meadows up to at least 1600 m a.s.l. (up to 2100 m a.s.l. in North Tyrol,
Austria), in the lowlands preferring moderately dry grassland on limestone or siliceous
substrates (also on sand), particularly on abandoned or disturbed pastures and fields.
Lives on species of Asteraceae, probably Taraxacum or Picris hieracioides, perhaps also
Hieracium and others. The German sites are at the northwestern edge of the range; the
species is widespread and fairly common in the Alps and their foothills, but rather
sporadic elsewhere, mainly in regions of limestone and sand. Border localities are near
Mainz (locus typicus), Göttingen and Schleswig.
Achtziger (1991), Remane (1987), Remane (pers. comm.), Schiemenz et al. (1996), Strübing (1966, and pers.
comm.), Wagner (1939a, 1951a), Nickel (1997), HN; see also Leising (1977)

Euscelis incisus (Kirschbaum, 1858)

E III – XI; nymph, usually 2 gen., at higher altitudes egg, 1 gen. (Müller 1954); 3rd and
4th instar nymphs and eggs, at higher altitudes only the latter, overwinter in thermic
quiescence; all other stages and adults usually die in autumn (Müller 1981; Witsack
1981).
In usually sunny, moist to moderately dry, eutrophic to oligotrophic sites, mainly in
meadows, pastures, abandoned fields, along waysides and ditches, and even on lawns
in gardens and parks; often abundant in low-vegetated, fertilized pastures and mead-
ows in sun-exposed situations. Host plants are various species of Fabaceae (mainly Tri-
folium spp.) and grasses (in England Trisetum flavescens, Briza media, Helictotrichon pube-
scens, Arrhenatherum elatius, Bromus erectus and others). Widespread in Germany and
very common, frequently found up to 1000 m a.s.l. in the Alps and on the Swabian Jura,
but usually below 700 m a.s.l. elsewhere. Records from higher altitudes probably refer
to stray individuals. Has been identified as a vector of various mycoplasma-like organ-
isms, such as Clover phyllody (CP), Clover dwarf (CD), Stolbur and Parastolbur in
various European countries.
Fischer (1972), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Wagner (1935,
1939a), Nickel & Achtziger (1999), HN; see also Brčák (1979), Cook (1996)

Euscelis lineolatus Brullé, 1832

E V – B XI; egg, 1 gen.
This species is widespread in the Mediterranean region and western Europe. In Germa-
ny it is restricted to the extreme western parts. Border localities are near Prüm and
 Deltocephalinae Fieber, 1869 203

Daun (both Eifel Mountains), Bonn, Münster, Leer and on the island of Norderney, up
to 450 m a.s.l. Furthermore, there is a single individual record from the vicinity of Bre-
men. Lives on eutrophic meadows and pastures, often in diked coastal grassland, also
in clover and alfalfa fields; host plants are species of Fabaceae (reported to live among
Trifolium, Medicago and Lotus in western and southwestern Europe) and grasses (in Eng-
land Lolium, Holcus, Festuca, Poa, Dactylis). Was identified as a vector of Clover phyllody
(CP) and other clover diseases in England.
Bornholdt & Remane (1993), Hildebrandt (1995), Hildebrandt et al. (1998), Müller (1957), Niedringhaus
(1991), Remane (pers. comm.), Strübing (1966, 1978, and pers. comm.), HN; see also Brčák (1979), Prestidge
& McNeill (1983)

Euscelis ohausi W. Wagner, 1939

E VI – M X; egg, 1 gen.
This species is confined to western parts of Europe; it is only reported from Portugal,
France, Switzerland, Germany, Belgium, the Netherlands, Britain (northward to Scot-
land), Jutland and northwestern Poland. It lives on woody species of Fabaceae in sunny,
occasionally also in moderately shady, damp to moderately dry sites, on acidic sub-
strates (mainly sand and sandstone). There are two morphs in Germany, which are com-
monly regarded as subspecies: the ssp. ohausi W. Wagner, 1939 is found on Genista angli-
ca, in heath and dune areas of northern Germany (Lüneburger Heide, in the vicinity of
Hamburg and Ludwigslust, and on the north Frisian islands of Amrum and Sylt); the
ssp. singeri W. Wagner, 1951 lives on Cytisus scoparius along margins of low-productivi-
ty grassland and forest glades, mainly in middle and southern parts. Border localities,
some of which appear to be rather isolated, are near Cologne, Marburg, Nordhausen,
Torgau an der Elbe, Niesky, Aschaffenburg, Erlangen and Oberstdorf im Allgäu (800 m
a.s.l.). Is apparently absent from the Czech Republic and Austria.
Frommer (1996), Remane (1967, and pers. comm.), Schiemenz et al. (1996), Strübing & Schwarz-Mittels-
taedt (1988), Trümbach (1959), Wagner (1951a), Walter (pers. comm.), HN

Euscelis venosus (Kirschbaum, 1868)

M VII – E IX; egg, 1 gen.
In oligotrophic, basic, damp to moderately dry (also temporarily dry), usually short-
vegetated pastures (occasionally meadows), in the Alps mainly on sun-exposed hill-
sides, in lower altitudes also on north-facing slopes. The host plants are unknown, but
are likely to include species of Asteraceae (perhaps Carlina acaulis and C. vulgaris). In
Germany the species is restricted to the southern half and occurs at the northern edge of
the range, with strongholds in the Bavarian and Allgäu Alps and their foothills, where
it is locally fairly common up to at least 1500 m a.s.l., but very sporadic in middle parts.
Border localities are near Mainz (locus typicus), in the Vogelsberg and Meißner, near
Herzberg am Harz, Stendal and Potsdam.
Fischer (1972), Nikusch (1976), Remane (pers. comm.), Schiemenz et al. (1996), Wagner (1939a, 1951a),
Nickel (1994), HN
204 Review of species

Ederranus discolor (J. Sahlberg, 1871)

So far B VI – B IX; probably egg, 1(?) gen.
At present, this species is only known from northern Russia (Karelia), southern Finland
and Brandenburg; its habitat requirements are poorly known. Phragmites australis and
Glyceria maxima have been reported as host plants; thus, the species is likely to live in
rather wet sites. The two German record are from inland salt marshes with Phragmites
australis and Juncus gerardii near the cities of Brandenburg and Potsdam: Trechwitz,
Netzener See, c. 30m a.s.l., 05.VI.1993, 1 †, 09.IX.1993, 4 ††, 1 ‡, and Gröben, Gröbener
See, c. 30m a.s.l., 10.IX.1993, 6 ††, 4 ‡‡.
Fröhlich (1996a), Remane & Fröhlich (1994b); see also Emelyanov (1964a), Lindberg (1947), Ossiannilsson
(1983), Sahlberg (1871)

Sotanus thenii (P. Löw, 1885)

So far B VII – B VIII, in Tyrol (Austria) E VI – E IX; nymph, 1 gen. (Dobler 1985; Leising 1977)
This species is confined to grasslands of damp, well-drained sites of the subalpine to
the upper alpine belt of the Alps; probably it lives on an unidentified species of Poace-
ae. From Germany it is only known from 5 localities in the Allgäu near Oberstdorf,
between 1700 and 2050 m a.s.l.: Hochvogel, Rotspitz, Nebelhorn, Rotkopf and Mindel-
heimer Hütte (below Liechelkopf), but it is probably more widespread in Bavaria. Oth-
erwise only reported from Austria, Switzerland and northern Italy, where found be-
tween 1500 and 3050 m a.s.l.
Fischer (1972), Schönitzer & Oesterling (1998b), HN; see also Dobler (1985), Günthart (1987a), Leising (1977),
Nast (1987), Thaler (pers. comm.), Wagner & Franz (1961)

Streptanus aemulans (Kirschbaum, 1868)

M VI – E X; egg, 2(?) gen.
In grass stands in sunny to moderately shady, usually damp to moist, occasionally also
moderately dry or wet sites (mainly meadows, pastures, forest glades, ruderal habitats,
waysides, also white and grey dunes near the coast and moderately saline sites). Host
plants are various grasses, often Holcus lanatus, H. mollis and Elymus repens. Widespread
in Germany and common in most regions, but usually below 900 m a.s.l.
Achtziger (1991), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Nikusch (1976), Reimer (1992),
Wagner (1935, 1951a), Walter (pers. comm.), Nickel & Achtziger (1999), HN, and others

Streptanus confinis (Reuter, 1880)

E VI – B X; egg, 1 gen.
Among tall grasses in sunny to shady, peaty to moist (preferentially waterlogged) sites,
mainly abandoned or low-input meadows and pastures, fens and fen woods, often in rath-
er cool situations, locally also in moderately saline sites. Usually (perhaps exclusively) deep
in tussocks of Deschampsia cespitosa; seldom recorded due to secretive life habits. In Germa-
 Deltocephalinae Fieber, 1869 205

ny perhaps near the western border of the range, currently known from most regions except
the northeastern states (Mecklenburg-Vorpommern, Brandenburg, Berlin, Saxony-Anhalt).
Most records comprise single or few individuals only. Not reported from Belgium, the Neth-
erlands, Poland and Austria; in France only known from a single locality in the Jura, but
probably under-recorded. The highest localities are at 800 m a.s.l. in the Rhön Mountains,
but single specimens have been found at 1800 m a.s.l. in the Bavarian Alps.
Bittner & Remane (1977), Niedringhaus & Olthoff (1993), Reimer (1992), Remane & Fröhlich (1994b), Walter
(1998), Nickel & Achtziger (1999), HN; see also della Giustina & Remane (2001), Nast (1987)

Streptanus marginatus (Kirschbaum, 1858)

M IV – E VIII; nymph, 1 gen.
In moderately shady, moderately moist to moderately dry sites, mainly in open decidu-
ous and coniferous forests, also along forest margins in dry grassland, heaths and mon-
tane meadows. Host plants are fine-leaved grasses, mainly Deschampsia flexuosa and
Festuca ovina, perhaps also Nardus stricta and further species. Widespread in Germany
and fairly common, found up to at least 1000 m a.s.l. in the Mittelgebirge and the Alps.
Achtziger (1991), Bornholdt (1996), Fischer (1972), Nikusch (1976), Reimer (1992), Remane (1987), Schie-
menz (1971a), Schiemenz et al. (1996), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN

Streptanus okaensis Zachvatkin, 1948

This taxon is perhaps conspecific with S. ogumae (Matsumura, 1911) described from Sakhalin.
The question is left open here until the type material has been revised.
E VI – B X; egg, 1 gen.
In Europe known only from Russia, Scandinavia, central Spain and the north German
plain, where recorded in c. 20 localities in Niedersachsen, Holstein, Brandenburg and
the Oberlausitz, up to at most 150 m a.s.l. It lives on Calamagrostis canescens in acidic,
wet, occasionally peaty, sunny to moderately shady sites (mainly fens, intermediate
bogs and fen woods). Is probably under-recorded due to secretive life habits.
Remane (1995), Remane & Fröhlich (1994b), Niedringhaus & Olthoff (1993), Walter (pers. comm.), HN

Streptanus sordidus (Zetterstedt, 1828)

M VI – M X, according to Schiemenz et al. (1996) M V – M XI; egg, 2(?) gen.
In moist to peaty or temporarily flooded, sunny to moderately shady sites, mainly low-
input pastures and meadows, fens, intermediate bogs and salt marshes, but also open
forests and clearings, rarely in conventionally managed grassland. Host plants are var-
ious grasses, preferentially Agrostis stolonifera and A. capillaris, perhaps also Deschamp-
sia cespitosa and others. The species is widespread in Germany and fairly common; it
occurs up to at least 1200 m a.s.l. in the Allgäu and Black Forest, but has been found at
1700 m a.s.l. in Styria (Austria).
Achtziger (1991), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Schie-
menz et al. (1996), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN; see also Wagner & Franz (1961)
206 Review of species

Artianus interstitialis (Germar, 1821)

E VI – B X; egg, 1 gen.
In scattered stands of grasses in moderately dry to dry sites, usually on sandy, locally
also loamy substrates, notably in ruderal sites, abandoned fields, along waysides and
in disturbed patches in dry grassland, occasionally in temporarily dry salt marshes.
Lives on various grasses, often Elymus repens, perhaps also Agrostis capillaris, Festuca
rubra and Holcus spp. Widespread and locally common in southern and eastern parts of
lowland Germany (frequently up to c. 400 m a.s.l., occasionally up to 700 m a.s.l.), at the
northwestern edge of the range. Border localities are on a line from Cologne to Mar-
burg, Gotha, Nordhausen, Quedlinburg, Wolfsburg and Lauenburg. Records published
from the Netherlands are erroneous.
Fröhlich (1996a), Frommer (1996), Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane
(1987), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Thüs (1996), Trümbach (1959), Wagner
(1935), HN; see also Metcalf (1967), Nast (1987), Reclaire (1944)

Paramesus obtusifrons (Stål, 1853) and P. major Haupt, 1927

Paramesus major Haupt, 1927 has been described from Israel and reported from most parts of the Pale-
arctic since (from eastern Siberia to the eastern Mediterranean region, Poland, the Czech Republic and
Hungary). Differences between eastern European populations and P. obtusifrons (Stål), which is con-
fined to central and western Europe, are weak and only concern the shape of the terminal aedeagus.
Thus, the specific distinctiveness of these two taxa may be uncertain, although no transitional individ-
uals have been found so far. †† of all German inland populations are not distinct from those living in
eastern Europe, which are currently treated as P. major Hpt. (see Anufriev & Emelyanov 1988). From
Poland Nast (1976a) mentions P. major Hpt. from inland sites and P. obtusifrons (Stål) from the coast.
However, it is unknown whether the ranges overlap along the Mediterranean and the Baltic Sea.
B VII – B X; egg, 1 gen.
Monophagous and halobiotic on Bolboschoenus maritimus, usually in reeds in brackish
and temporarily flooded sites along the coast and inland. Recorded from all East Fri-
sian Islands, the Weser estuary near Bremerhaven, the island of Sylt, and the coast of
the Baltic Sea between Schleswig and Hiddensee (P. obtusifrons Stål), as well as inland
salt marshes (rarely in freshwater sites with strongly fluctuating water tables) in the
rain shadow area to the east of the Harz Mountains (roughly between Magdeburg, Halle,
Erfurt and Sondershausen), in eastern Niedersachsen (Schreyahn), the Werra valley
(Ettmarshausen), the Wetterau (Wisselsheim, Münzenberg), and in the northern upper
Rhine valley near Speyer (P. major Hpt.). The highest localities are at 250 m a.s.l. only.
Published feeding records on Schoenoplectus are probably erroneous.
Fröhlich (1996a), Kuntze (1937), Hildebrandt (1995), MTD, Niedringhaus (1991), Schaefer (1973), Schie-
menz et al. (1996), Wagner (1935), HN

Parapotes reticulatus (Horváth, 1897)

M VII – B X; egg, 1 gen.
In tussocks of Schoenoplectus lacustris, usually in permanently flooded reeds along shores
of fishponds, glacial and oxbow lakes, a single individual also found in a brackish spring
 Deltocephalinae Fieber, 1869 207

mire on Sch. tabernaemontani. In Germany only known from altogether 8 records, all
from northeastern parts (up to at least 450 m a.s.l.) and apparently at the western edge
of the range. Border localities are near Waren an der Müritz, Dannenberg, Bad Lauter-
berg, Ilmenau and Görlitz. Is probably under-recorded. Has been reported from salt
marshes of eastern Austria and Hungary.
Höhne (1921), Sander et al. (1999), Schiemenz et al. (1996), Nickel (1997), HN; see also Fröhlich (1996a),

Paralimnus phragmitis (Boheman, 1847)

B VII – B X; 1 gen., in southwestern Germany from E V onwards, 2(?) gen.
Monophagous on Phragmites australis in mesotrophic to moderately eutrophic, tempo-
rarily or non-flooded sites, usually along shores of lakes and ponds, in fens, mining
areas, coastal and inland salt marshes. Widespread in Germany, but only scattered; of-
ten sampled in low numbers only. The highest localities are at 700 m a.s.l. in the Bavar-
ian foothills of the Alps.
Fröhlich (1996a), Heller (1987a), Hoffmann (1980), Niedringhaus (1991), Schiemenz et al. (1996), Schönitzer
& Oesterling (1998b), Wagner (1939a), Nickel (1999b), HN

Paralimnus rotundiceps (Lethierry, 1885)

So far only IX, in Austria M VI (Moosbrugger 1946), egg(?), 1(?) gen.
This is a rare species of European distribution, living on Phragmites australis. In Germany it
has only recently been discovered in two widely separated localities: coal mine of Mücheln
near Merseburg, c. 150 m a.s.l., several records in low-growing and sparse pioneer stands of
reed on rather dry, sandy soil, e.g. 03.IX.2001, 2 ††, 1 ‡, and Griesen near Garmisch-Parten-
kirchen, c. 900 m a.s.l, IX 2000, 1 †, in a Malaise trap on a gravel bank. Further known
localities are in Vorarlberg (Austria), the Valais (Switzerland) and southern Scandinavia.
Funke (pers. comm.), Al Hussein et al. (1999), Nickel & Voith (unpublished data); see also Cerut-
ti (1939b), Fröhlich (1996a), Moosbrugger (1946), Ossiannilsson (1983)

Metalimnus formosus (Boheman, 1845)

Mainly M VI – E X, according to Fischer (1972) also E V; egg, 1 gen., perhaps 2 in low-
land sites.
Among tall sedges in wet to temporarily flooded, often peaty sites, usually in fens,
straw meadows, along shores of oligotrophic to moderately eutrophic lakes and ponds,
as well as in floodplain depressions. Host plants are tall-growing species of Carex (nota-
bly C. acuta and C. elata, perhaps also C. nigra). The species is widespread in Germany,
but occurs only localized, with highest frequencies in river valleys from the Elbe east-
ward as well as along the edge of the Alps (up to at least 900 m a.s.l., but found at
1500 m a.s.l. in North Tyrol, Austria).
Fischer (1972), Fröhlich (1996a), Hoffmann (1980), Remane (1962), Schiemenz et al. (1996), Schönitzer &
Oesterling (1998b), SMNS, Wagner (1939a), Wagner & Wagner (1938), HN; see also Cobben & Gravestein
(1958), Ribaut (1952)
208 Review of species

Metalimnus steini (Fieber, 1869)

Has been described as a new variety of the preceding taxon, from the surroundings of Berlin,
but the whereabouts of the type material are unknown. Later it was treated as a distinct species,
and M. tredecimpunctatus (Lindberg, 1929), described from Japan, as a junior synonym. It has
been reported from large parts of the Palearctic, including Hungary, Ukraine, Austria, Kazakh-
stan and far-eastern Russia (Anufriev & Emelyanov 1988; Holzinger et al. 1997; Nast 1972; Re-
mane & Fröhlich 1994a). However, the taxonomic situation is rather critical, since the identity of
M. steini (Fieb.) is not clear, and since it is likely to comprise a complex of species.
So far M VI – M X; egg, 2(?) gen.
The locus typicus is near Berlin, but there are no more verified records since, although 2
dark-coloured ‡‡ from the collection of Haupt, taken near Aken an der Elbe, 23.VII.1911,
are likely to belong to this taxon. Furthermore, there are three very recent records from
the upper Rhine plain between Freiburg and Darmstadt (100 – 250 m a.s.l.), in stands of
Carex hirta in sunny to moderately shady, moderately wet to moderately dry ruderal
habitats and meadows. However, in the Rhine populations conspecificity with the type
material has yet to be confirmed.
Fieber (1869), MTD, HN

Arocephalus longiceps (Kirschbaum, 1868)

M V – M X; egg, 2 gen., perhaps 1 at higher altitudes.
In sunny to moderately shady, moderately moist to moderately dry sites, mainly clear-
ings, forest glades, dry grassland, heaths, ruderal sites and abandoned fields, occasional-
ly also open forests and low-input-meadows. Single individuals on dispersal flight are
frequently encountered in other habitats. Host plants are various species of Poaceae, no-
tably Holcus mollis, H. lanatus and Bromus erectus. Widespread in Germany and fairly com-
mon in middle and southern parts, but more scattered in the north German plain, where
living at the edge of the range. Northernmost localities are on the East Frisian Islands,
near Pinneberg, Plön and Greifswald (but has recently been found in Skåne, southern
Sweden). Found up to at least 1100 m a.s.l. (but single specimens at 1600 m a.s.l.) in the
Mittelgebirge and Bavarian Alps, at 1730 m a.s.l. in the Engadine (Switzerland).
Achtziger (1991), Fischer (1972), Niedringhaus & Olthoff (1993), Nikusch (1976), Post-Plangg & Hoffmann
(1982), Reimer (1992), Remane (1987), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1939a),
Nickel & Achtziger (1999), HN; see also Gillerfors (2002), Günthart (1987a)

Arocephalus languidus (Flor, 1861)

M V – B X; egg, 2 gen., 1 at higher altitudes.
In moderately dry to dry, sunny to moderately shady sites on basic as well as acidic,
gravelly to loamy substrates, usually in various types of xerothermic grassland (also
with open stands of pine or oak) as well as in pastures and meadows of the subalpine
and alpine belt. Host plants are Sesleria albicans and Stipa spp., probably also Koeleria
glauca and other grasses. Rather localized in Germany, at the northwestern edge of the
range, and with strongholds in the Alps and their foothills, as well as in warmer regions
of southern and eastern parts, often among the dominant species in favourable sites.
 Deltocephalinae Fieber, 1869 209

Border localities are on a line from Koblenz to Schlüchtern, Eschwege, Nordhausen,

Magdeburg and Rügen. Frequently found up to at least 2000 m a.s.l. in the Bavarian
and Allgäu Alps.
Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987, and pers. comm.), Schie-
menz et al. (1996), Wagner (1939a, 1951), Nickel (1994), HN

Arocephalus punctum (Flor, 1861)

B VI – E X; egg, 2 gen., perhaps 1 at higher altitudes.
In oligotrophic, sunny to slightly shaded, low-vegetated, dry to moist (rarely wet), acidic
as well as basic sites; in the north German plain favouring sandy dry grassland, heaths
and forest margins, in the region of the Mittelgebirge mainly on meadows and pastures
of the submontane and montane belt, locally also in dry grassland on limestone, sand
and other dry substrates at low altitudes. Lives in tussocks of fine-leaved grasses, main-
ly Festuca ovina, perhaps also F. rubra, Nardus stricta and Corynephorus canescens. Wide-
spread in northern and middle parts of Germany and locally common, but not recorded
to the south of the Danube; the highest localities are at 1250 m a.s.l. in the Black Forest.
Niedringhaus (1991), Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Schiemenz et al.
(1996), Trümbach (1959), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others

Arocephalus sagittarius Ribaut, 1952

So far only M VI, in France also B IX (della Giustina 1989); egg, 2(?) gen.
Has very recently been found in the Kaiserstuhl in extreme southwestern Germany:
Badberg, 14.VI.2002, small populations in several patches of xerothermic loess grass-
land, apparently among fine-leaved grasses (perhaps Festuca ovina). Was already re-
ported from adjacent parts of Alsace (France), but is otherwise only known from Swit-
zerland, Spain and Portugal.
Lauterer & Malenovsky (pers. comm.), Remane (pers. comm.), Nickel & Remane (2002), HN; see also della
Giustina (1989), Remane & Fröhlich (1994b)

Psammotettix maritimus (Perris, 1857)

E VI – M IX; egg, 1(?) gen.
This species is an endemic of western European coasts and lives monophagously on Am-
mophila arenaria in calcareous primary and white dunes close to the shoreline. From Ger-
many it is known only from the East and North Frisian Islands (Borkum, Memmert, Juist,
Norderney, Baltrum, Langeoog, Spiekeroog, Wangerooge, Amrum and Sylt). Otherwise it
is reported only from England, the Netherlands, Belgium, France and Portugal.
Niedringhaus (1991), Niedringhaus & Olthoff (1993), Remane (pers. comm.), Wagner (1935); see also Nast (1987)

Psammotettix poecilus (Flor, 1861)

M V – B X; egg, 2 gen.
210 Review of species

In sandy to gravelly, usually sunny, acidic as well as basic sites with very scattered
stands of grass, mainly inland dunes, mining areas, forest clearings and almost bare
gravel banks of alpine rivers. Lives on Calamagrostis epigejos in lowland sites, but on C.
pseudophragmites in the Alps and their foothills. In Germany at the western edge of the
range, widespread and locally fairly common in the sand areas of northeastern parts
(westward at least to Greifswald, Brandenburg, Halle, Leipzig and Bautzen), but rather
sporadic and in strongly isolated populations elsewhere, notably in the Rhine-Main
region, Mittelfranken, and along the upper course of the Isar between Mittenwald and
Lenggries, where found up to 900 m a.s.l. There are also single localities in the Kyff-
häuser and northern Hessen. Not recorded from Denmark, the Netherlands and Bel-
gium; in France known only from the western Alps (Savoie).
Funke & Witsack (1998), Remane (1987), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Wagner (1939a,
1951a), Nickel (1999b), HN; see also della Giustina & Remane (2001)

Psammotettix unciger Ribaut, 1938

So far only B VIII – E VIII; 1(?) gen.
This is probably the rarest species treated in this book. Since the original description, it
has been recorded only twice. The locus typicus is near Oberstdorf im Allgäu: Iller flood-
plains; 04. and 07.VIII.1936; also North Tyrol (Austria): Lech valley to the northwest of
Reutte, VIII 1973; and Upper Bavaria: Isar floodplains near Vorderriß, 26.VIII.1997, 6 ††,
8 ‡‡. It lives on very scattered grass stems on almost bare gravel banks of unchanneled
alpine rivers; specimens taken near Vorderriß were swept from Calamagrostis pseudo-
phragmites. All localities are at c. 800 m a.s.l. The Iller population is probably extinct due
to draining of the river bed.
Remane & Fröhlich (1994b), Ribaut (1938) (see this paper also for identification), Nickel (1999a)

Psammotettix kolosvarensis (Matsumura, 1908)

E V – E IX; egg, 2 gen.
Usually in low-growing grass stands in sunny, moderately wet to moderately dry sites
with strongly fluctuating levels of fresh or moderately saline water, mainly inland salt
marshes, sandy abandoned fields, as well as temporarily dry meadows and pastures,
also in more intensively used sites. Single individuals on dispersal flight are frequently
encountered in other habitats. Is often found on Puccinellia distans and Elymus repens,
but probably feeds also on additional grasses. In Germany largely confined to eastern
parts, probably at the edge of the range, with strongholds in the basins of the Elbe and
Oder, where locally abundant in riverine grassland, which is frequently flooded during
winter, but may become rather dry during summer. Border localities are near Schwedt,
Rathenow, Lauenburg, Wolfsburg, Sondershausen, Meißen and Görlitz, up to 200 m
a.s.l. To the west of Germany there is only a single record from the Gironde area (France).
Fröhlich (1996a), Schiemenz et al. (1996), Rothenbücher (pers. comm.), Walter (1996), Nickel & Achtziger
(1999), Nickel (1997), HN; see also Ribaut (1959)
 Deltocephalinae Fieber, 1869 211

Psammotettix angulatus (Then, 1899)

Regarding genital morphology, this taxon closely resembles Ps. koeleriae Zachvatkin, 1948, which
is reported from middle Russia to Mongolia. There are differences, however, in body coloura-
tion: Ps. angulatus Then is straw-coloured, Ps. koeleriae Zachv. is green. According to Remane
(1965), they should preliminarily be treated as distinct species, until further evidence has been
provided.
So far M VI – E VII; egg, 2(?) gen.
Known from Germany only from a single site in Brandenburg, on a xerothermic, sandy,
south-facing moraine slope: Groß Machnower Weinberg near Zossen, 70 m a.s.l. („Schil-
lergras-Schafschwingelrasen mit Elementen der Silbergrasflur auf einer Waldlichtung“),
12.VI.1964, 1 †, 1 ‡, 27.VII.1964, 1 ‡. Koeleria glauca may have been the host plant. In
1998 the site was studied again, but without a positive record. Hence, the species is
perhaps extinct in Germany. Otherwise it is reported only from single localities in Bohe-
mia (Czech Republic), central Poland and northern Italy.
Schiemenz (1969) (Remane det.), Schiemenz et al. (1996); see also Nast (1972, 1987); for identification see
Remane (1965)

Psammotettix nardeti Remane, 1965

M VII – E IX; egg, 1 gen.
This is an endemic species of the Alps, where it is apparently confined to western parts.
Lives in meadows and pastures of the subalpine to the upper alpine belt, probably on a
specific grass (Nardus stricta has been suggested). In Germany it is known only from
4 records from the Allgäu, the Ammergebirge, and the Berchtesgaden Alps, between
1600 and 2000 m a.s.l. (Untere Gottesackerwände above Oberstdorf, Hochplatte above
Füssen, Friederspitz above Garmisch-Partenkirchen, and Funtensee above Berchtes-
gaden). Otherwise it is reported only from Austria, France and Italy, between 1550 and
2600 m a.s.l.
Remane (1965), HN; see also Günthart (1984, 1987a), Leising (1977), Nast (1987); for identification see
Remane (1965)

Psammotettix inexpectatus Remane, 1965

So far M VII – B X; egg, 1(?) gen.
For a long time, this species was known only from the locus typicus in Thuringia, until
there was a second record from Mount Olymp in Greece. In Germany it is restricted to
southwest-facing, xerothermic gypsum hillsides in the Kyffhäuser near Bad Franken-
hausen, c. 200 to 270 m a.s.l. From this locality there are altogether 5 records: 29.VIII.1956,
1 †, 3 ‡‡; 10.X.1965, 1 †, 2 ‡‡; 17.VII.1991, 1 †, 1 ‡; 06.IX.1996, 2 ††, 1 ‡; 02.IX.2000,
4 ††, 5 ‡‡. No details are known on the biology; the host plant is probably a non-iden-
tified species of grass.
Fröhlich (pers. comm.), Remane (1965), Schiemenz et al. (1996), Nickel et al. (2001), HN; see also Drosopo-
ulos et al. (1986); for identification see Remane (1965)
212 Review of species

Psammotettix alienus (Dahlbom, 1850)

Mainly E V – M X, occasionally B V – B XI; egg, 2 gen.
This is a eurytopic pioneer species in sunny, dry to wet sites, mainly in fallow and cereal
fields, meadows, mining areas and slightly saline sites. Invades all types of tree-less
habitats and is among the very few leafhopper species of fertilized meadows and pas-
tures, although individual numbers may fluctuate strongly, and breeding success may
be reduced in these places. Host plants are various grasses. Widespread in Germany
and particularly common in the southern half, below 900 m a.s.l., but less frequent in
the north German plain. Was recorded as a vector of Band mosaic of wheat and rye
(BMWR), Wheat dwarf virus (WDV), and perhaps additional plant diseases in various
countries. WDV on cultivated barley has recently become a problem in parts of Saxony-
Anhalt and also Sweden.
Fischer (1972), Müller (1956), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Wagner (1935, 1939a),
Manurung et al. (2001), Nickel & Achtziger (1999), HN, and others; see also Brčák (1979), Lindblad & Arenö
(2002); for nymphal morphology see Guglielmino & Virla (1997)

Psammotettix sabulicola (Curtis, 1837)

E V – E IX; egg, 2 gen.
On tall but scattered grasses on coastal dunes and in sandy, sun-exposed and open
habitats inland. Ammophila arenaria and Elymus spp. are possible host plants, although
some specimens have been found among Leymus arenarius and Calamagrostis epigejos. In
Germany the species is confined to the northern plains, with strongholds in coastal
areas near the North and Baltic Sea, where it is locally fairly common on white, grey
and brown dunes, but there is also a number of localities up to c. 70 km inland, notably
near Oldenburg, Winsen, Boizenburg and Neuhaus. The range is apparently confined
to northern and western European coastal lowlands; records from Italy and Bulgaria
may be doubtful.
Niedringhaus (1991), Remane (1958, and pers. comm.), Schaefer (1973), Schiemenz et al. (1996), Wagner
(1935), HN; see also della Giustina & Remane (2001), Gravestein (1965)

Psammotettix pallidinervis (Dahlbom, 1850)

E V – M X; egg, 2 gen.
In sparsely vegetated, xerothermic grassland, in northern Germany mainly on sand, in
middle and southern parts also on (preferentially grazed) gypsum and limestone sites .
Lives on fine-leaved species of fescue (Festuca ovina group), but has been reported from
Corynephorus canescens. In Germany the species occurs at the southwestern edge of the
range, only very localized, and usually in low abundances, mainly in the north German
plain and the rain shadow area to the east of the Harz Mountains; further localities,
which appear to be rather isolated, are near Lingen (Ems), Celle, Bad Wildungen and
Regensburg, up to at most 400 m a.s.l. It is reported from the Netherlands and Austria,
but not from Belgium, France, Switzerland and Italy.
Remane & Fröhlich (1994b), Schaefer (1973), Schiemenz et al. (1996), Wagner (1937a), HN; see also Nast (1987)
 Deltocephalinae Fieber, 1869 213

Psammotettix cephalotes (Herrich-Schäffer, 1834)

E V – M X; egg, 2 gen., at higher altitudes probably only 1.
On Briza media in oligotrophic and sunny sites; at lower altitudes the species is largely
restricted to grazed, dry grassland on basic soils; at submontane and higher altitudes it
is more widespread and common, favouring peaty to moderately dry low-productivity
meadows and pastures (also on acidic substrates). Widespread in Germany, but appar-
ently absent from the northwest German plain to the west of the Elbe. Lives up to at
least 1500 m a.s.l. in the Bavarian and Allgäu Alps, up to 2200 m a.s.l. in the Engadine
(Switzerland) and North Tyrol (Austria).
Reimer (1992), Rombach (1999b), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Nickel & Acht-
ziger (1999), Nickel (1994), HN; see also Günthart (1987a), Leising (1977)

Psammotettix helvolus (Kirschbaum, 1868) group

According to Remane & Fröhlich (1994a), this is a „Formengruppe mit ungeklärter taxonomi-
scher Wertigkeit der verschiedenen Formen, z.B. P. rhombifer (Fieber, 1869), P. substriatus (Then,
1902), P. obtusiceps (Kirschbaum, 1868)“. Remane (1987) has provisionally named two forms
occurring in Germany which are morphologically and ecologically distinct. Accordingly, „helvo-
lus basic“ is macropterous and lives on various grasses, mainly in neutral to basic, and dry sites
of lower altitudes (usually below 800 m a.s.l.), whereas „helvolus acidic“ is subbrachypterous
and is usually found in acidic sites of higher altitudes (usually above 300 m a.s.l.). The latter
prefers Deschampsia flexuosa as host plant. See also Günthart (1987a), Leising (1977), Wagner &
Franz (1961). Biosystematic studies are needed to clarify the taxonomic situation.
B V – M X; egg, 2 gen., at higher altitudes only 1.
In sunny, occasionally slightly shady, dry to moist sites on basic to acidic substrates.
Particularly abundant in low-input meadows, dry grassland and abandoned fields, also
found in pastures, forest clearings, open forests, and along waysides, occasionally in
fertilized meadows. Widespread and common in southern and middle parts of Germa-
ny, but known only from a few localities in the north German plain (recorded near
Lüchow, Zossen and Greifswald). It lives up to at least 1900 m a.s.l. in the Allgäu Alps,
up to 2600 m a.s.l. in the Engadine (Switzerland). Records from even higher altitudes
probably not refer to resident populations, but only to stray individuals.
Fischer (1972), Reimer (1992), Remane (1987), Rombach (1999b), Schiemenz et al. (1996), Wagner (1939a,
1951a, 1958), Walter (1996), Nickel & Achtziger (1999), Nickel (1994), HN; see also Dobler (1985), Günthart
(1984, 1987a), Leising (1977), Wagner & Franz (1961)

Taxa belonging to the Ps. nodosus group, comprising Ps. nodosus (Rib.), Ps. albomarginatus W.Wg.,
Ps. putoni (Then), Ps. excisus (Mats.) and Ps. dubius Oss., are morphologically little distinct (in
particular considering the aedeagus shape), but show specific habitat requirements and – at
least in some cases – also host preferences. Their taxonomic status is not satisfactorily clarified.

Psammotettix albomarginatus W. Wagner, 1941

E V – E X; egg, 2 gen.
In oligotrophic, sunny and sparsely vegetated sites, usually with a high proportion of
lichens, probably on Agrostis vinealis or Corynephorus canescens, preferentially on sandy
substrates, but also found on porphyry. So far known from Germany only from a few
214 Review of species

localities in the north German plain and near Halle, Leipzig, Dresden and Speyer, all
below 200 m a.s.l. Apparently confined to northwestern and central parts of Europe,
otherwise reported only from Ireland, Wales, England, southern Sweden, Finland, Den-
mark, the Netherlands and Poland.
Remane (1958, and pers. comm.), Schaefer (1973), Schiemenz (1969), Schiemenz et al. (1996), Wagner (1941a),
HN; see also Kirby (1992), Nast (1987)

Psammotettix putoni (Then, 1898)

E V – B X; egg, 2 gen.
Halobiotic in salt marshes; often abundant in salt meadows near the North and Baltic Sea
(eastward at least to Kiel), also found in two inland sites of northern Germany (Barnstorf
near Braunschweig, and Sülldorf near Magdeburg). Is reported to live on Puccinellia marit-
ima and perhaps P. distans. All localities are below 150 m a.s.l. Otherwise the species is only
published from Norway, Sweden, northern Russia, Ireland, Britain, Denmark, the Nether-
lands, Belgium and France. Records from southeastern Europe require verification.
Fröhlich (1996a), Niedringhaus (1991), Remane & Fröhlich (1994b), Tulowitzki (1990), Wagner (1941a), HN;
see also Schiemenz et al. (1996),

Psammotettix dubius Ossiannilsson, 1974

In North Tyrol (Austria) according to Leising (1977) M VIII – M IX; egg, 1 gen.
So far only known from Norway and Sweden (northward to c. 65o N), middle Russia, North
Tyrol and Bavaria. Reported to live in Scandinavia „ ... on grasses on dry, sandy soils, ... also
in heather marshes and dry moors“, in North Tyrol in alpine peaty meadows and bogs,
between 1900 and 2100 m a.s.l. Has recently been found in the Allgäu Alps: Oberjoch above
Hindelang, 1200 m a.s.l., 05.VIII.1995, 4 ††, 2 ‡‡, at the edge of a raised bog.
Nickel (1999b), HN; see also Anufriev & Kirillova (1998), Leising (1977), Ossiannilsson (1983)

Psammotettix excisus (Matsumura, 1906)

M V – E X; egg, 2 gen.
In sandy, xerothermic, sparsely-vegetated and acidic sites, mainly inland dunes, heaths,
military training areas and sand pits, preferentially on Corynephorus canescens, perhaps
also on other grasses. Widespread and fairly common in the north German plain and
locally dominant in favourable sites, but apparently absent or rare on coastal dunes of
all kinds; also found in scattered localities in middle and southern parts of Germany,
notably in inland dune areas in the northern upper Rhine plain (between Speyer and
Darmstadt), in the valleys of the Main (near Karlstadt, Kitzingen, Volkach), Regnitz
(near Bamberg, Forchheim, Erlangen) and Danube (Siegenburg near Kelheim), as well
as in the basin of Mittelfranken (Altdorf near Nuremberg), up to 450 m a.s.l. Otherwise
only reported from Estonia, Lithuania, Poland, Denmark, the Netherlands, France, Por-
tugal, the Czech Republic and Hungary.
Remane (1958, 1965), Schiemenz et al. (1996), Trümbach (1959), Wagner (1941a), Nickel (1999b), HN
 Deltocephalinae Fieber, 1869 215

Psammotettix nodosus (Ribaut, 1925)

M V – M X; egg, 2 gen.
This is a pioneer species of sunny, dry to damp and sparsely vegetated sites on various
substrates, mainly in strongly grazed and oligotrophic pastures, in dunes and ruderal
sites, but also in forest clearings, in dried peat diggings, on roadside embankments, etc.;
host plants are various, usually low-growing grasses, often Festuca ovina. Widespread
in the lowlands of Germany, breeding takes place up to at least 800 m a.s.l., single indi-
viduals have been found up to 1700 m a.s.l. in the Bavarian Alps. Widespread in Ger-
many, locally abundant in sandy sites of the north German plain, but more scattered
and in lower densities elsewhere.
Fischer (1972), Niedringhaus (1991), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schi-
emenz et al. (1996), Wagner (1939a, 1941a), HN, and others

Psammotettix notatus (Melichar, 1896)

Has been misinterpreted for a long time. In fact, almost all published records (see Metcalf 1967;
Nast 1987) are in need of revision.
So far only B IX – M IX; egg(?), 1(?) gen.
This species is apparently very rare, and almost nothing is known on its life history. There
are two German records: Lower Bavaria, Saal an der Donau, c. 400 m a.s.l., 02.IX.1960,
1 †, and Mainfranken, Klotz near Retzbach, c. 250 m a.s.l., 15.IX.1994, 2 ‡‡, both on xero-
thermic calcareous hillsides. The host plants are presumably grasses. Otherwise, only the
locus typicus is known (“Nieder-Oesterreich, am Eichkogel bei Mödling, im August und
September”). Most, if not all, other published records are based upon misinterpretation.
Remane (1961b, and pers. comm.), HN; see also Melichar (1896), Wagner (1939a) (see this paper also for
identification)

Psammotettix confinis (Dahlbom, 1850)

Psammotettix remanei Orosz, 1999 (see Nickel & Remane 2002)
This taxon shows a considerable variation of colouration, body size and aedeagus shape. Re-
cently the original description of Psammotettix remanei Orosz, 1999 was published, with paratypes
from the Oberlausitz. However, the type specimens were clearly individuals whose genital ap-
paratus was not fully developed (perhaps parasitized), notably regarding the genital plates,
lateral lobes of the pygophor, as well as the base and shaft of the aedeagus. Moreover, the con-
nective was rather abnormal and did not even show the typical Paralimnini shape. Such indi-
viduals of Ps. confinis (Dhlb.) with rudimentary genital apparatus, reduced body size and pig-
mentation are more or less frequently found among central European populations. Only some
of them show visible external or internal signs of parasitation, but their morphology is clearly
abnormal.
M V – M X; egg, 2 gen.
Eurytopic in grass stands of sunny, dry to moderately wet, preferentially neutral to
acidic sites, often abundant in pastures, fertilized meadows and ruderal sites, but also
on inland dunes, in wet meadows and moderately brackish sites. Host plants are vari-
ous grasses, probably Lolium, Poa, Festuca and Agrostis. Widespread all over Germany
216 Review of species

and very common, found up to at least 1200 m a.s.l. in the Bavarian Alps, up to at least
1900 m a.s.l. in subalpine meadows of North Tyrol (Austria).
Bornholdt (1996), Emmrich (1966), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Reimer (1992),
Remane (1958, 1987), Schiemenz (1969), Schiemenz et al. (1996), Wagner (1935, 1939a, 1951a), Nickel &
Achtziger (1999), HN, and others; see also Leising (1977), Prestidge & McNeill (1983a, 1983b)

Ebarrius interstinctus (Fieber, 1869)

The identity of this taxons remained unclear for a long time, until Nast (1977) published a ge-
neric revision. Specimens from the French Alps with slight differences compared to the nomi-
nate form were described as E. interstinctus sofiae della Giustina, 1989.
M VI – E IX; egg, 2(?) gen.
Reported from alpine dwarf shrub heaths of the French and Swiss Alps, between 2100
and 2500 m a.s.l. In Germany, however, there are only two known lowland localities:
northwestern slopes of the Swabian Jura: Balingen, c. 500 m a.s.l., 23.VI.1955, a popula-
tion on a sun-exposed roadside embankment, as well as Nahe valley near Schloßböck-
elheim, c. 250 m a.s.l., between mid June and end of September 1995, several specimens
in Malaise traps, in a xerothermic, abandoned vineyard. Probably lives on grasses. Oth-
erwise reported only from France, Switzerland, Austria, Hungary, northern Italy, former
Yugoslavia, Bulgaria and Greece.
Remane & Fröhlich (1994b), Fröhlich & Nickel (unpublished data); see also Drosopoulos et al. (1986), della
Giustina (1989), Günthart (1987a), Nast (1987)

Ebarrius cognatus (Fieber, 1869)

M VII – E IX; egg, 1 gen.
In oligotrophic, sunny, short-vegetated sites of the montane to the alpine belt, locally also at
lower altitudes along alpine rivers; mainly in grassland with predominating Sesleria albi-
cans, Carex ferruginea or dwarf shrubs. Host plants are grasses. Has been taken on Festuca
vivipara in Scotland, which does not occur in central Europe, however. In Germany confined
to the Bavarian and Allgäu Alps, where apparently widespread between 1500 and 1850 m
a.s.l., occasionally on riverine gravel banks of lower altitudes (Wimbach valley near Ram-
sau, between 800 and 1400 m a.s.l.). Other lowland records are in need of revision. Was
reported from the upper alpine belt (up to 2800 m a.s.l.) of the Tyrolean Alps (Austria).
Remane & Fröhlich (1994b), HN; see also Günthart (1984), Leising (1977), Woodroffe (1971)

Adarrus multinotatus (Boheman, 1847)

M V – M X, occasionally until XII; egg, 2 gen.
On Brachypodium pinnatum in dry to temporarily moist, sunny to moderately shady
sites, usually on basic substrates, mainly in various types of rather dry grassland as
well as in open forests of pine and oak, occasionally in rather moist straw meadows.
Absent or scarce in most parts of the north German plain, but widespread to the south
of it, particularly common and abundant in limestone regions. Frequently found above
950 m a.s.l. on the Swabian Jura, up to 1500 m a.s.l. in the Bavarian Alps.
 Deltocephalinae Fieber, 1869 217

Achtziger (1991), Fischer (1972), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Rombach
(1999b), Schiemenz (1969), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957),
Wagner (1939a), Nickel (1994), HN, and others; see also Cook (1996)

Adarrus bellevoyei (Puton, 1877)

So far E VII – B X; egg, 1(?) gen.
On Brachypodium pinnatum, usually in temporarily dry to damp, basic sites. In Germany it
is found only along the upper course of the Isar and in the Eifel Mountains: near Vorder-
riß, Wallgau and Krün, between 800 and 900 m a.s.l., in open, cattle-grazed pine and
spruce forests on alluvial gravel; near Prüm and Bad Münstereifel, 400 to 500 m a.s.l., in
calcareous grassland. Is reported to live in forest clearings between 1300 and 1500 m a.s.l.
in the Valais (Switzerland), between 1300 and 1800 m a.s.l. in the French Alps.
Bornholdt & Remane (1993), Remane (1961b), Rombach (1999b), HN; see also Cerutti (1938), della Giustina (1989)

Errastunus leucophaeus (Kirschbaum, 1868)

Deltocephalus antennalis Haupt, 1924 (see Nickel & Remane 2002)
In the whole range so far found B VI – B IX; egg, 2(?) gen.
According to present data this species is a rare endemic of the northern Alps and their
foothills, living on almost bare gravel banks of unchanneled rivers. Outside Germany it is
only known from the Swiss and Austrian Rhine and the North Tyrolean Lech valley: Rhine
near Ragaz (type locality), Rhine delta near Bregenz, 400 m a.s.l., 03.VI.1917, and Lech near
Reutte, 800 m a.s.l., VIII 1973. From Germany there are only three records: Iller floodplains
near Oberstdorf, 11.VIII.1934 and 20.VIII.1935; Isar floodplains near Ascholding, 10.VII.1959,
and near Geretsried, 10.IX.1994, between 600 and 800 m a.s.l. Probably lives on an uniden-
tified grass. The Iller and Rhine populations may be extinct due to channeling.
Haupt (1924), Remane (1961b), Remane & Fröhlich (1994b), Wagner (1939a), Nickel (1999a); see also Kir-
schbaum (1868); for identification see Wagner (1939a)

Errastunus ocellaris (Fallén, 1806)

Subalpine populations tend to be more strongly pigmented and have been described as E. ocel-
laris tatraensis (Heller, 1975) from the Tatra Mountains. However, there is apparently clinal vari-
ation towards the nominate form, since specimens from the Bavarian Alps show characters of
both taxa. The late Heller (pers. comm.) himself had serious doubts on the distinctiveness.
M V – E X; egg, 2 gen.
Eurytopic in sunny to slightly shaded stands of grasses in moderately dry to wet, mesotrophic
to eutrophic sites. Usually among the dominant species in intensively managed meadows
and pastures of lower altitudes, but also in abandoned fields, ruderal habitats, forest clear-
ings, along waysides, in straw meadows, fens and even brackish sites. Host plants are var-
ious, usually taller-growing grasses (species of Holcus and Calamagrostis, Elymus repens,
Dactylis glomerata and others). Widespread and very common in Germany, frequently ex-
tending up to 1000 m a.s.l.; small populations have also been found in the upper montane
and subalpine belt of the Berchtesgaden and Allgäu Alps between 1500 and 2000 m a.s.l.
218 Review of species

Achtziger (1991), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992),
Remane (1987), Remane & Fröhlich (1994b), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a, 1951a),
Nickel & Achtziger (1999), HN, and others

Turrutus socialis (Flor, 1861)

M V – E X; egg, 2 gen. (see also Witsack 1985).
Usually in sunny and oligotrophic, peaty to moderately dry, basic to acidic sites on
various substrates, preferentially low-productivity meadows and pastures, locally rud-
eral sites, waysides, inland dunes, etc., often abundant in calcareous grassland and
submontane meadows. Lives on grasses (Festuca rubra, Bromus erectus, probably Nardus
stricta and others; also reared from Helictotrichon pubescens and Brachypodium pinnatum
taken from British calcareous grassland). Apparently absent from the northwest Ger-
man plain, but widespread and common in most other parts of Germany, found up to at
least 900 m a.s.l. in the Mittelgebirge and the Bavarian Alps, up to 1500 m a.s.l. in the
Engadine (Switzerland).
Achtziger (1991), Fischer (1972), Kuntze (1937), Nikusch (1976), Reimer (1992), Remane (1987), Rombach
(1999b), Schiemenz (1969), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1939a, 1951a),
Nickel & Achtziger (1999), HN, and others; see also Cook (1996), Günthart (1987a)

Jassargus pseudocellaris (Flor, 1861)

Jassus distinguendus Flor, 1861
Flor (1861) created the name J. distinguendus (Fl.) as a replacement name in a chapter of addenda
in the same book, based on the erroneous opinion that the name Jassus distinguendus was preoc-
cupied by Athysanus (= Euscelis) distinguendus Kirschbaum, 1858.
E V – E X; egg, 2 gen.
In low-growing grass stands in moist to moderately dry, usually oligotrophic, more or
less acidic sites, mainly low-productivity meadows and pastures, heaths and abandoned
fields, occasionally also ruderal sites, forest margins and clearings. Host plants are grasses
(Agrostis capillaris and Festuca rubra, perhaps also Nardus stricta and Holcus spp.). Wide-
spread in Germany, often among the dominant species of low-productivity grassland of
submontane altitudes and in the north German plain, particularly on sand, sandstone,
granite and other siliceous substrates, but largely absent from limestone regions. Fre-
quently found up to at least 1400 m a.s.l. in the Alps and in higher parts of the Mittelge-
birge, single specimens have been recorded up to 1800 m a.s.l.
Achtziger (1991), Fischer (1972), Niedringhaus (1991, 1997), Nikusch (1976), Post-Plangg & Hoffmann (1982),
Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935,
1939a), Nickel & Achtziger (1999), HN, and others

Jassargus obtusivalvis (Kirschbaum, 1868)

B V – E X; egg, 2 gen. (see also Witsack 1985).
In sunny, moderately dry to dry, basic to acidic sites on various substrates, mainly dry
grassland, abandoned vineyards, inland dunes, low-productivity meadows, waysides,
 Deltocephalinae Fieber, 1869 219

etc., preferentially in taller grass stands. Lives oligophagously on grasses and is often
abundant in swards of Bromus erectus, Brachypodium pinnatum and others. In Germany
confined to warmer regions of southern and eastern parts between the Danube and the
north edge of the Mittelgebirge (mainly basins of Rhine, Main, Neckar and Nahe, and
the Saale-Unstrut region, up to at most 650 m a.s.l.) and apparently with sharp northern
distribution border along a line from Cologne to Marburg, Nordhausen, Halle and
Bautzen. Has been found up to 1500 m a.s.l. in the French Alps.
Frommer (1996), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Rombach (1995), Schie-
menz et al. (1996), Schulz (1976), Schwoerbel (1957), Wagner (1939a, 1951a), HN

Jassargus repletus (Fieber, 1869)

So far M VII – M IX, in the Austrian Alps from M V onwards (Wagner & Franz 1961);
egg, 1(?) gen.
On sun-exposed calcareous hillsides and riverine gravel banks, usually with open stands
of pine and other trees, probably monophagously on an unidentified grass species. In
Germany it is only known from a few localities along the edge of the Alps (Illasberg
near Füssen, c. 800 m a.s.l., and upper course of the Isar between Wolfratshausen and
Krün, 600 to 1000 m a.s.l.), and the upper Rhine plain (inland dunes near Speyer, c.
100 m a.s.l.). At least small numbers have been found at 1300 m a.s.l. in southern parts
of Switzerland.
Fischer (1972), Remane & Fröhlich (1994b), Schulz (1976), HN; see also Günthart (1987a, 1997)

Jassargus flori (Fieber, 1869)

B VI – B X; egg, 2 gen., perhaps 1 at higher altitudes.
Usually in moderately shady, moderately moist to moderately dry sites, preferentially
open deciduous and coniferous forests, and clearings of lower altitudes (below c. 700 m
a.s.l.), also in sunny and open pastures in the submontane and montane belt of the Alps,
found up to at least 1500 m a.s.l. in Upper Bavaria. Host plants are narrow-leaved grasses
(perhaps Poa pratensis angustifolia, Agrostis capillaris, Deschampsia flexuosa or Festuca hetero-
phylla). Widespread in most parts of Germany, but uncommon and in low densities only.
Fischer (1972), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Schulz (1976), Schwoerbel (1957),
Wagner (1935, 1939a), Trümbach (1959), HN

Jassargus alpinus (Then, 1896)

M VII – E IX, according to Schiemenz et al. (1996) from E V onwards; egg, 1 gen.
In damp to moist, moderately shady to sunny sites, mainly at higher altitudes of the
Mittelgebirge and the Alps; usually in open forests and clearings of the submontane to
the upper montane belt, higher up also in pastures and meadows. Host plants are var-
ious grasses (locally Calamagrostis villosa, probably also C. arundinacea, Deschampsia flex-
uosa and others). In Germany confined to the Alps and higher parts of the Mittelgebirge,
and with distinct subspecies: ssp. alemannicus W.Wg. has been found in the Allgäu Alps
220 Review of species

and the Black Forest, ssp. alpinus Then lives in the Bavarian Alps, Rhön Mountains,
Thuringian Forest, Solling and Harz Mountains, between 450 and at least 1900 m a.s.l.;
small numbers have been collected along the edge of the Harz Mountains between 150
and 300 m a.s.l., but breeding in these sites may be uncertain.
Fischer (1972), Reimer (1992), Schulz (1976), Wagner (1958), Schiemenz et al. (1996), HN

Jassargus allobrogicus (Ribaut, 1936)

Deltocephalus bavaricus Ribaut, 1936
M VI – M X, after Schiemenz et al. (1996) from M V onwards; egg, 1 gen., perhaps 2 at
low altitudes.
Preferentially in open deciduous and coniferous forests on siliceous substrates, and of-
ten abundant in stands of Deschampsia flexuosa under beech, spruce and pine; locally
and in lower numbers also on basic substrates where probably feeding on Festuca ovina,
F. heterophylla and F. rubra. Widespread in Germany and common, also in higher parts of
the Mittelgebirge and in the upper montane belt of the Alps, where found up to at least
1800 m a.s.l. Reported from 2200 m a.s.l. in Switzerland.
Achtziger (1991), Fischer (1972), Reimer (1992), Schulz (1976), Schiemenz et al. (1996), Wagner (1935, 1939a),
HN; see also Günthart (1987a)

Jassargus sursumflexus (Then, 1902)

M VI – E X; egg, 1 - 2 gen.
Monophagous on Molinia caerulea s.l. in peaty to temporarily moist, sunny to moderate-
ly shady sites, mainly in intermediate bogs, straw meadows and open forests. Wide-
spread in Germany, but scattered, with strongholds in the Alps and their foothills, the
Mittelgebirge, and remnant bog areas of the northern plains, where locally common and
abundant in favourable sites, but absent or rare elsewhere. Recorded up to 1350 m a.s.l.
in the Allgäu, at 2000 m a.s.l. in the French Alps.
Fischer (1972), Kuntze (1937), Reimer (1992), Remane (1958), Wagner (1935), Schiemenz (1971a, 1975, 1976),
Schiemenz et al. (1996), Schulz (1976), Schwoerbel (1957), Nickel & Achtziger (1999), HN

Mendrausus pauxillus (Fieber, 1869)

B VII – M X; egg, 1 gen.
This species is widespread in the steppe zone between Mongolia and eastern parts of
central Europe, with rather isolated populations in eastern Germany. Lives on sunny
and sparsely vegetated xerothermic sites with relic steppe vegetation (mainly on gyp-
sum, locally also on limestone), on greyish-green subspecies of Festuca ovina. In Germa-
ny it is confined to the rain shadow area to the east of the Harz Mountains, currently
known only from altogether 6 localities in the vicinity of Bad Frankenhausen, Artern
and Halle, between 100 and 300 m a.s.l.
Schiemenz (1969), Schiemenz et al. (1996), HN
 Deltocephalinae Fieber, 1869 221

Pinumius areatus (Stål, 1858)

Hamilton (1983a) proposed – without presenting further arguments – that European popula-
tions belonged to Pinumius nebulicola Emelyanov, 1972, described from high alpine steppes of
the Pamir mountains. From a biogeographical point of view, this would seem extremely unlike-
ly. Moreover, specimens recently collected in the Lausitz as well as populations from the Euro-
pean part of Russia and Ukraine clearly belong to P. areatus (Stål) (Emelyanov pers. comm.;
Walter pers. comm.).
B VI – M X; egg, 2 gen.
This is another eastern steppe species with isolated populations in central Europe. Lives
on sunny and sparsely vegetated dry grassland, usually on aeolian or moraine sand on
an unidentified grass species, perhaps Festuca ovina or Koeleria spp. In Germany there is
only a single recent site in a large coal mining area in the Lausitz (Innenkippe Nochten,
09.VI. 1995, 3 ††, and 27.VI.1995, 1 ‡). Altogether 5 further populations found in the
1930s and 1960s, some of which showed extremely high abundances, are presumably
extinct (Mainz, Neustrelitz, as well as three sites on the Oder slopes near Eberswalde
and Seelow). All localities are below 150 m a.s.l.
Kuntze (1937), Schiemenz (1969, and unpublished data), Wagner (1939a), Walter (pers. comm.), Remane (1987)

Diplocolenus bohemani (Zetterstedt, 1838)

Mainly M V – M VII, at higher altitudes VII – X; egg, 1 gen.
In tall, but usually scattered stands of grasses in temporarily moist to dry, sunny to mod-
erately shady sites of almost all altitudinal belts. In the lowlands and the Mittelgebirge the
species is scattered and largely confined to calcareous and gypsum hillsides as well as
forest clearings of various substrates; in the Alps and their foothills it is more eurytopic
and widespread in various types of mesotrophic grassland, particularly in montane pas-
tures, and in open forests. Preferentially feeds on species of small-reed (Calamagrostis varia,
C. villosa, C. epigejos), but also on other grasses; the main host plant in calcareous grass-
land of lower altitudes is probably Bromus erectus. Locally common and abundant in the
Bavarian and Allgäu Alps (up to at least 1900 m a.s.l., but found up to 2250 m a.s.l. in
North Tyrol, Austria), in most limestone regions of southern and eastern Germany (nota-
bly Swabian and Franconian Jura, Triassic limestone regions of Baden, Franconia, Thurin-
gia), as well as in the montane belt of most Mittelgebirge). Despite its wide distribution in
Scandinavia there are no records from the north German plain; the northernmost German
localities are on a line from Siegen to Kassel, Hildesheim, Halberstadt and Cottbus.
Fischer (1972), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957),
Wagner (1951a), Nickel (1994), HN; see also Günthart (1987a), Leising (1977), Ossiannilsson (1983)

Diplocolenus penthopitta (Walker, 1851)

Some authors treat D. sudeticus (Kolenati, 1860) which shows slight differences in genital mor-
phology, as a distinct species and consider the identity of Walker’s type material as dubious
(Dlabola 1980; Nast 1977, 1987). On the other hand Fieber (1869) has redescribed D. penthopitta
(Walk.), his material was revised by Wagner (1939a, 1948a) and declared conspecific with D.
sudeticus (Kol.). This opinion is followed by Knight (1974), della Giustina (1989) and Holzinger
222 Review of species

et al. (1997). Specimens from Bavaria do neither fully correspond with the French nor with the
Czech ones (see Wagner 1939a; Dlabola 1954), but largely with a single individual from Vorarl-
berg, Austria (see Holzinger 1999a). Like in other cases of isolated mountain populations, for
instance in the genus Jassargus, these differences are likely to be only clinal variations on subspe-
cific level (e.g. Schulz 1976).
So far M VII – M VIII, egg(?), 1 gen.
This species is hitherto known only from the French Alps, Vorarlberg (Austria) and the
Polish and Czech Sudetic Mountains. In France it lives between 1650 and 1800 m a.s.l.;
the only known specimen from Austria is from c. 1600 m a.s.l. More recent records are
from the Bavarian Alps (Mangfallgebirge) near Bayrischzell: Hochmiesing, c. 1800 m
a.s.l., 14.VII. and 12.VIII.1998, altogether 7 ††, 4 ‡‡, in Malaise traps of two different
sites, and Rotwand, c. 1750 m a.s.l., 18.VIII.2002, several populations in subalpine pas-
tures among Deschampsia cespitosa and Nardus stricta (but probably feeding on another,
rather inconspicuous grass). In the Czech Sudetes it prefers avalanche tracks of the
upper montane forest belt.
Nickel & Voith (unpublished data), HN; see also della Giustina (1989), Holzinger (1999a), Lauterer (pers. comm.)

Verdanus abdominalis (Fabricius, 1803)

In the lowlands mainly M V – VIII, at higher altitudes VII – B X; egg, 1 gen.
In sunny, damp to moist, occasionally also wet or moderately dry sites (low-input mead-
ows and pastures, fields, clearings, etc.), on Holcus lanatus and other grasses (notably
Festuca rubra, Agrostis capillaris, Deschampsia flexuosa). Widespread in Germany, particu-
larly common in the northern plains and the submontane and montane belt of the Mit-
telgebirge and the Alps, but absent or uncommon in warmer regions of southern parts
(e.g. upper Rhine plain, Main valley). Recorded up to 2050 m a.s.l. in the Bavarian Alps
and the Allgäu, up to 2600 m a.s.l. in North Tyrol (Austria) and Switzerland.
Achtziger (1991), Fischer (1972), Nikusch (1976), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oester-
ling (1998b), Nickel & Achtziger (1999), HN, and others; see also Dobler (1985), Günthart (1987a), Leising
(1977)

Verdanus bensoni (China, 1933)

Deltocephalus obenbergeri Dlabola, 1945
So far E VI – B VIII, egg, 1 gen.
This species’ range is disjunct and restricted to European mountains. Until recently it
was only known from England and Scotland, the Swiss and French Jura, the Bohemian
Forest (Czech Republic), the Giant Mountains (Poland) and the Ural Mountains (Rus-
sia). During recent years large populations were also recorded in the Bavarian Forest
(Falkenstein near Zwiesel, 1130-1150 m a.s.l., in pastures, perhaps living on Nardus stricta
and Deschampsia flexuosa), as well as in alpine grassland of the Allgäu and Upper Bavar-
ia (Hochplatte above Füssen, 1850 m a.s.l., and Friederspitz above Garmisch-Parten-
kirchen, c. 1900 m a.s.l.).
Biedermann (1998), Nickel & Voith (unpublished data); see also Emelyanov (2000), Lauterer (pers. comm.),
Nast (1976a, 1987), Remane & Fröhlich (1994b), Waloff (1979, 1980)
 Deltocephalinae Fieber, 1869 223

Arthaldeus arenarius Remane, 1960

Until the generic revision of Remane (1960) was published, this species was commonly misiden-
tified as A. striifrons (Kbm.). Thus, a considerable part of these records from central Europe –
even after Remane’s paper – refer to A. arenarius Rem., which is much more widespread.
E VI – B IX; egg, 1 gen.
In sunny to moderately shady, moist to moderately dry sites, mainly in forest clearings,
mining areas and various other ruderal habitats, occasionally also in open coppice-with-
standards and along forest roads, monophagous on Calamagrostis epigejos. The distribu-
tion is only poorly known due to former identification problems (see above). However,
the species is widespread in Germany and common in many places, at least to the east
and south of a line from Düsseldorf to Hanover and Kiel (but not recorded from Mecklen-
burg-Vorpommern), up to at least 800 m a.s.l. Otherwise known only from southeastern
Poland, the Czech Republic, northern France, Austria, southern Russia and Kazakhstan,
although probably more widespread in Europe. The locus typicus is Gauting near Munich.
Frommer (pers. comm.), Kolbe & Bruns (pers. comm.), Reimer (1992), Remane (1960), Remane & Fröhlich
(1994b), Schiemenz et al. (1996), SMNS, Nickel (1994), HN; see also Emelyanov (1964a), della Giustina &
Remane (2001), Lauterer (pers. comm.), Mityaev (1975), Nast (1976a)

Arthaldeus striifrons (Kirschbaum, 1868)

B VI – E X; egg, 2 gen.
Locally fairly common along the coasts of the North and Baltic Sea, where usually found
in sunny, moderately saline and temporarily wet to damp sites, mainly meadows and
pastures, but also abandoned fields and ruderal sites. Inland, however, the species is
rather localized. Many older records (mainly those from forest and bog habitats, as well
as from higher altitudes) refer to the preceding species and are in need of revision. The
only verified inland localities are in saltmarshes of Thuringia and Saxony-Anhalt, the
lower course of the Oder and Isar, the Lake Constance basin, the upper Rhine plain and
the Nahe valley, up to 400 m a.s.l. In these sites, the species lives on moderately saline or
compacted substrates on Festuca arundinacea, probably also on F. pratensis or Lolium per-
enne. The locus typicus is near the salt-works of Bad Dürkheim.
Fröhlich (1996a), Niedringhaus (1991), Niedringhaus & Olthoff (1993), Remane (1960), ZIMH, Nickel &
Achtziger (1999), Nickel (1999b), HN

Arthaldeus pascuellus (Fallén, 1826)

M V – E X; egg, 2 gen. (see also Witsack 1985).
In peaty to damp, usually sunny sites, mainly meadows and pastures, also fens, abandoned
fields, ruderal sites, clearings, waysides, moderately saline marshes inland and near the
coast. Host plants are various grasses (Festuca, Lolium, Poa, Calamagrostis and others ). Wide-
spread and common all over Germany, often dominant in fertilized grassland, frequently
found in higher parts of the Mittelgebirge, up to 1500 m a.s.l. in the Bavarian Alps.
Achtziger (1991), Emmrich (1966), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Reimer (1992), Remane
(1958), Schaefer (1981), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others
224 Review of species

Sorhoanus assimilis (Fallén, 1806)

E VI – M X; egg, 1 gen.
In acidic to basic, usually peaty and permanently wet, sunny to moderately shady sites,
mainly intermediate and spring mires, calcareous sedge fens and straw meadows. Host
plants are sedges (perhaps Carex panicea, C. nigra or C. rostrata). Widespread in Germa-
ny, but scattered, locally dominant in favourable sites, with strongholds in the Mittelge-
birge, the Alps and their foothills; the highest localities are at 1000 m a.s.l.
Fischer (1972), Reimer (1992), Schiemenz et al. (1996), Trümbach (1959), Wagner (1935, 1939a, 1941b, 1951a),
Nickel & Achtziger (1999), HN

Sorhoanus schmidti (W. Wagner, 1939)

B VII – E IX; egg, 1 gen.
In peaty to temporarily wet, basic to acidic sites, mainly straw meadows, calcareous sedge
fens and intermediate bogs, occasionally syntopically with S. assimilis (Fall.) or S. xantho-
neurus (Fieb.). Appears to be associated with open stands of Molinia caerulea, which may
be the host plant. This is an endemic species of the Alps and adjacent areas only known
from Austria, Switzerland and southern Germany. In Germany it is largely confined to
the Alps and their foothills (mainly Upper Bavaria and the Allgäu), where it occurs local-
ly between 600 and 1200 m a.s.l.; it is also found in the southern Black Forest and the Main
valley near Schweinfurt, the latter site is situated at only 210 m a.s.l. The locus typicus is
Oberstdorf im Allgäu. Otherwise the species is only reported from altogether 8 localities
in the northern and central Alps of Styria (Austria) and the Engadine (Switzerland), where
it is found up to 1680 m a.s.l., but it is probably more widespread in the Alps.
Remane & Fröhlich (1994b), Fischer (1972), HN; see also Günthart (1987a), Wagner & Franz (1961); for
identification see Wagner (1939a)

Sorhoanus xanthoneurus (Fieber, 1869)

E VII – M X; egg, 1 gen.
On Eriophorum vaginatum in raised and intermediate bogs, usually (but not exclusively)
in treeless and sunny sites. Widespread in Germany, but very scattered, although often
dominant in remnant bogs of the north German plain, the Mittelgebirge (Harz Moun-
tains, Solling, Thuringian Forest, Vogtland, Erzgebirge, Black Forest), and along the edge
of the Bavarian and Allgäu Alps, up to 1350 m a.s.l. Has been found on Trichophorum
cespitosum in the Czech Republic, but this is probably not the host plant.
Niedringhaus & Olthoff (1993), Schiemenz (1971a, 1975, 1976), Schiemenz et al. (1996), Wagner (1941b),
Walter (pers. comm.), Nickel (1999b, 2002), HN; see also Novotný (1995)

Lebradea calamagrostidis Remane, 1959

Is considered as the only leafhopper species apparently endemic to Germany, distinguished
from the Holarctic Lebradea flavovirens (Gillette & Baker, 1895) only by the length of the aedeagus
appendages. However, in German populations of L. calamagrostidis Rem. there is some variabil-
 Deltocephalinae Fieber, 1869 225

ity in this character. Furthermore, from a biogeographical point of view, the occurrence of a
north German endemic is unlikely. L. flavovirens (Gill. & Bak.) is also reported from Calamagrostis
“in dry as well as in marshy habitats” and has very recently been found in central and southern
Sweden. Thus, the relationship between the two taxa should be revised.
E VI – VIII; egg, 1 gen.
So far only reported from the north German plain. Altogether 6 localities are known, all
in southern parts of Schleswig-Holstein below 50 m a.s.l.: Appen, Wennebek, Megger-
dorf, Nettelsee, Plön and Lebrade (locus typicus), on Calamagrostis canescens in fens and
intermediate bogs.
Remane (1959), Remane & Fröhlich (1994b), Niedringhaus & Olthoff (1993), Schröder & K. Heller (pers.
comm.); for identification see Remane (1959); see also Gillerfors (2002, and pers. comm.), Ossiannilsson
(1983), Vilbaste (1980)

Cosmotettix caudatus (Flor, 1861)

B VI – B IX; egg, 1 gen.
On Carex hirta in temporarily moist to wet, usually moderately eutrophic, often dis-
turbed sites on various substrates (low-input pastures and meadows, along ditches and
waysides, ruderal habitats). In Germany apparently near the western edge of the range,
in low to intermediate frequency on the host. Mainly found to the northeast of a line
from the island of Wangerooge to Cologne, Göttingen, Marburg, Aalen, Forchheim and
Dippoldiswalde (Erzgebirge, c. 700 m a.s.l.); isolated localities are known from England,
western France, Switzerland and the Czech Republic, but there are no records from the
Netherlands, Belgium, Austria and Poland. The species is probably much under-re-
corded due to its secretive and specific life habits.
Frommer (1996), Hempel et al. (1971), Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Schie-
menz et al. (1996), SMNS, Wagner (1937f), Nickel & Achtziger (1999), HN; see also della Giustína & Re-
mane (2001), Kirby (1992), Nast (1987)

Cosmotettix panzeri (Flor, 1861)

B VII – E X; egg, 1 gen.
This is a tyrphophilous species which is restricted to northern and central parts of Europe. It
lives in intermediate and raised bogs on cottongrass (Eriophorum angustifolium, perhaps also
E. vaginatum). In Germany only sporadic, with strongholds in the remnant bog areas of the
north German plain; also found in a few sites in the Mittelgebirge and near the edge of the
Bavarian Alps up to 900 m a.s.l. Southernmost records are from Wolfratshausen (to the south
of Munich) and Titisee-Neustadt (southern Black Forest). Border localities are known from the
French Alps, the Czech Republic and Slovakia. Not reported from Austria and Switzerland.
Reimer (1992), Remane (1958, and pers. comm.), Schiemenz et al. (1996), Wagner (1935), Nickel & Remane
(1996), Nickel (1997), HN; see also della Giustina (1989), Jansky & Okali (1993), Lauterer (1984)

Cosmotettix costalis (Fallén, 1826)

E VI – B X, occasionally B VI; egg, 1 gen.
226 Review of species

In wet to temporarily flooded, usually mesotrophic to moderately eutrophic, treeless sites,

mainly in fens, straw meadows and depressions of river floodplains. Lives on sedges (Carex
acuta, probably also C. elata and C. nigra). In Germany at the southwestern edge of the range,
rather scattered and in low individual numbers, although perhaps under-recorded due to
secretive life habits. Border localities are on the island of Borkum, near Oldenburg, Göttin-
gen, Gießen, Schweinfurt, Lenzkirch (Black Forest), Immenstadt (Allgäu) and Garmisch-
Partenkirchen, up to 750 m a.s.l. There are only very few isolated records to the west and
south of this line, notably from Norfolk (England), near Apeldoorn (Netherlands), Zurich
(Switzerland) and Klagenfurt (Austria). Not reported from France.
Fröhlich (1996a), Niedringhaus (1991), Nikusch (1976), Remane (1962), Remane & Fröhlich (1994b), Nickel
& Achtziger (1999), Nickel & Remane (1996), Nickel (1997), HN; see also Bieman & Rozeboom (1993),
Günthart (1987b, and pers. comm.), Holzinger (1995c), Kirby (1992)

Cosmotettix aurantiacus (Forel, 1859)

So far M VI – E VIII; egg, 1 gen.
In wet, usually peaty sites, mainly straw meadows and fens of the foothills of the Alps,
presumably on an unidentified species of Carex. Known in Germany from only 10 locali-
ties, in Upper and Lower Bavaria (locally extending northward to the Danube valley), the
Allgäu and the southern upper Rhine plain (Burkheim near Freiburg), between 150 and
800 m a.s.l. Otherwise the species is reported from Lithuania, Poland, Switzerland, Aus-
tria, eastern France, European and eastern Siberian parts of Russia, as well as Mongolia.
Remane (1961d), Remane & Fröhlich (1994b), HN; see also Anufriev & Emelyanov (1988), della Giustina &
Remane (2001), Nast (1972, 1987)

Calamotettix taeniatus (Horváth, 1911)

So far B VII – B IX; egg, 1 gen.
This is another monophagous species on Phragmites australis, preferentially occurring in
temporarily flooded freshwater sites as well as inland salt marshes. In Germany at the
edge of the range and was recorded only from 9 sites in the southern half: upper Rhine
valley (Speyer and Landau), Bodanrück (Konstanz), near Stuttgart (Horrheim), Wetter-
au (Rockenberg and Selters), Grabfeld (Bad Königshofen), Oberlausitz (Königshain and
Nochten) and Geiseltal near Halle, between 100 and 450 m a.s.l. It is probably under-
recorded as it is rather wary and thus, difficult to sweep.
Fröhlich (1996a), Heller (1987b), Funke (pers. comm.), Remane (1995), Remane & Fröhlich (1994b), Nickel
(1999b), HN

Praganus hofferi (Dlabola, 1947)

So far B VII – M IX, in Bohemia (Czech Republic) according to Dlabola (1947) also E V;
egg, 2 gen.
This species is widespread in the Asian steppe zone, with relic populations in central
Europe. In Germany it is only known from a single site in the Thuringian Basin:
 Deltocephalinae Fieber, 1869 227

Schwellenburg near Kühnhausen, 210 m a.s.l., 15.IX.1963, 36 specimens, in a dry, south-

sloping pasture on Triassic gypsum with dominating Stipa capillata and Festuca ovina.
From this site there are also two recent records: 13.VII.1997, 1 †, 6 ‡‡, and 03.VII.1998,
4 ‡‡. The species is probably associated with Stipa.
Schiemenz (1973), HN; also Emelyanov (pers. comm.); for identification see Dlabola (1947, 1954), Mityaev
(1971)

Enantiocephalus cornutus (Herrich-Schäffer, 1838)

E VI – E IX; egg, 1 gen.
In sunny and scarcely vegetated, usually disturbed stands of grasses, mainly in moder-
ately dry to dry (occasionally also temporarily moist and moderately saline), sandy to
loamy sites, e.g. waysides, fallows, inland salt marshes and disturbed patches of dry
grassland. The main host is Elymus repens, but the species probably lives also on Agrostis
capillaris and Festuca rubra. In Germany only locally in lowland areas below 400 m a.s.l.,
near the northern limit of the range, recorded from the northern upper Rhine plain, the
valleys of the Nahe and Tauber, Franconia (locally common in the valleys of Main, Saale
and Regnitz, Grabfeld), the rain shadow area to the east of the Harz Mountains, the Thu-
ringian Basin and southeastern Niedersachsen (vicinity of Helmstedt and Wolfenbüttel).
Fröhlich (1996a), Reimer (1992), Remane & Fröhlich (1994b), Schiemenz et al. (1996), SMNS, HN

Mocuellus collinus (Boheman, 1850)

M V – M X; egg, 2 gen.
Often syntopic with the preceding species, but more common and also occurring in the
northern plains, with more pronounced preference of sandy soils. In sunny, moderately
dry to dry sites with incomplete vegetation cover, mainly abandoned fields, waysides
and pastures. Lives on various grasses (Elymus repens, low-growing species of Festuca,
Poa compressa and others). In Germany widespread and locally common, particularly in
the lowlands of southern and eastern parts, but absent from most parts of the Mittelge-
birge as well as to south of the Danube. Highest localities are at 450 m a.s.l. only.
Fischer (1972), Post-Plangg & Hoffmann (1982), Remane (1987), Schiemenz (1969), Schiemenz et al. (1996),
Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN, and others

Erzaleus metrius (Flor, 1861)

B VI – E X; egg, 2 gen., probably 1 in cooler sites.
On Phalaris arundinacea in moderately wet to temporarily flooded, often eutrophic sites,
mainly along ditches and shores of rivers and lakes, and in abandoned grassland, occa-
sionally on meadows cut at most once a year. Widespread in Germany, usually in inter-
mediate to high frequency and high abundance on the host, found up to at least 1000 m
a.s.l. in the Mittelgebirge and the Bavarian Alps.
Fischer (1972), Haupt (1925), Mölleken & Topp (1997), Niedringhaus (1991), Reimer (1992), Remane (1958),
Schiemenz et al. (1996), Trümbach (1959), Wagner (1935, 1939a), Witsack (1985), HN; see also Lauterer (1986)
228 Review of species

4.3 Dubious records

Ribautodelphax pallens (Stål, 1854)

Most records of this species are from northern Europe, the Alps (between 1450 and
2250 m a.s.l.) and the Pyrenees; furthermore, it is reported from Poland, the former
Yugoslavia and Bulgaria (Nast 1987; Ossiannilsson 1978; Remane & Fröhlich 1994b).
From Germany there is only a single record from Thuringia: Sulza near Jena, 26.V.1977,
1 †, 1 ‡ (Coll. Sander, Bieman det.). This is the only central European lowland site. Thus,
there may be a possibility of exchanged labels. The species lives monophagously on
Festuca ovina (Bieman 1987b).

Tettigometra concolor Fieber, 1865

Listed by Nast (1972, but not 1987) for West and East Germany. Varieties bearing the
name concolor Fieb. have been described for several taxa of the genus Tettigometra. Iden-
tity, taxonomic status, as well as occurrence in Germany must be regarded as dubious.

Micrometrina longicornis Signoret, 1866

Listed by Nast (1972, 1987) for West Germany, presumably referring to Melichar (1896),
Hüeber (1904) and Haupt (1935), but no specimens have been found in any of the re-
cently revised collections. Therefore, these statements most likely refer to localities out-
side present-day Germany.

Paradorydium paradoxum (Herrich-Schäffer, 1837)

This taxon is widespread from central Asia to the Mediterranean region, but perhaps
described from Franconia („von Dr. Hahn angeblich im Herbste bei Nürnberg gefunden“,
see Herrich-Schäffer 1837), but never recorded again in Germany since. Its occurrence
in central Europe may be only erratic. The food plants are grasses in steppe-like habi-
tats (Emelyanov 1964a; Mitjaev 1971; see also D’Urso 1992).

Chlorita viridula (Fallén, 1806)

Has been listed for Germany in older papers, but was usually confused with Ch. paolii
(Oss.). A more recent record was published by Schiemenz (1990): Strausberg near Frank-
furt an der Oder, 06.IX.1979, 1 †. However, the specimen is apparently lost. Occasional-
ly, specimens of Ch. paolii (Oss.) showing abnormal genital morphology have been found.
Hence, this record is regarded here as dubious. From England Ch. viridula (Fall.) is re-
ported to live in coastal saltmarshes on Artemisia maritima (Le Quesne & Payne 1981;
Kirby 1992).
 Dubious records 229

Edwardsiana kemneri (Ossiannilsson, 1942)

The identity of this taxon is not quite clear. It has been described after a single † from
central Sweden. After that, further records from Czechia, Slovakia, Romania, Cyprus
and Mongolia have been published (Dlabola 1965, 1967b, 1977). It is, furthermore, re-
ported from the Spitzberg near Tübingen without any further comments (Schwoerbel
1957). However, this record is not found in Wagner’s unpublished files, who has seen
Schwoerbel’s material. Due to the widespread occurrence of genitalic abnormalities
caused by parasitation in Edwardsiana specimens, this taxon should be omitted from the
German list. Populations of E. tersa (Edw.), in particular, have been found to contain ††
of the kemneri aedeagus type.

Selenocephalus obsoletus (Germar, 1817)

There are perhaps two specimens from Germany, the exact origin of which is dubious:
Bavaria, Main valley, Retzbach, 09.VI.1964, 1 † (ZMH, leg. H. Lindberg) and “Württem-
berg, v. Roser, 1872-76” (SMNS). The nearest confirmed localities are in eastern Austria,
Slovakia, South Tyrol (Italy) and southern France. In Mediterranean regions the species
has been swept from tall herbs, woody species of Fabaceae, and Cistus.

Handianus flavovarius (Herrich-Schäffer, 1835) sensu Emelyanov (1964a)

This is another species perhaps described from Regensburg, without any further records
from Germany. The original diagnosis does not include a description of the genitalia,
the type material is lost, and no type locality is given. The species is not mentioned in
the ‘Fauna Ratisbonensis’ (Herrich-Schäffer 1935b, 1840). Hence, origin and identity of
the holotype as well as the occurrence in Germany must be considered as dubious.
According to Nast (1972, 1987), the species is reported from the Altai westward to east-
ern parts of central Europe. Nearest localities are in southern Poland and Moravia (Nast
1976a; Lauterer 1983). According to various authors, it lives on grasses on sunny, damp
to dry sites; in Latvia and Lithuania adults have been found from mid June until mid
August (Emelyanov 1964a; Vilbaste 1974).

Handianus procerus (Herrich-Schäffer, 1835) sensu Emelyanov (1964a)

Often cited as having been described from Regensburg (e.g. Metcalf 1967; Nast 1987).
However, Herrich-Schäffer (1835b, 1840) neither gave any information on the locus typi-
cus, nor listed the species in his ‘Fauna Ratisbonensis’ (Herrich-Schäffer 1840); further-
more, the type material is lost. Thus, the occurrence in Germany is not confirmed. Ac-
cording to Nast (1987), the species is widespread from central Asia to western Europe
(see also Ribaut 1952), although the identity may be unclear. Senecio and Echinops are
reported as food plants from southern Russia (Emelyanov 1964a); in Moravia it has
been found on Genista pilosa in xerothermic grassland (P. Lauterer, pers. comm.).
230 Review of species

4.4 Short-term changes

4.4.1 Declines
Due to the lack of accurate data, long-term population declines are difficult to substan-
tiate. On the other hand, local extinctions and a decline of the whole range can often be
documented relatively easily by checking host plants or specific habitat structures. In the
Red Data list of German Auchenorrhyncha (Remane et al. 1998), only three species are
referred to as ‘extinct’ or ‘possibly extinct’, namely Cercopis arcuata Fieb., Paradorydium par-
adoxum (H.-S.) and Balclutha boica W.Wg. However, field evidence for a real extinction is
only substantial for the former, which has been recorded 4 times in the foothills of the Alps,
probably before 1950. After that, it has not been found any more, despite its striking appear-
ance and frequent sampling in the area. The two latter species, however, have been record-
ed only in single occasions, respectively. Regarding P. paradoxum (H.-S.), the origin of the
type material has been questioned by Herrich-Schäffer (1837) himself (see section 4.3). Fur-
ther, there is evidence that both Tettigometra laeta H.-S. and Psammotettix angulatus (Then)
have disappeared. In both cases, there is only a single substantiated record dating back 170
and 40 years, respectively. A conservative list of declining species is presented in Table 3,
based either on substantiated extinction in at least 3 sites during recent decades, or on large-
scale and long-termed decrease of records, or on large-scale habitat changes. Possible fac-
tors are taken mostly from Nickel et al. (1999); the present status is mostly taken from Re-
mane et al. (1998). Population declines are particularly obvious in the following groups:
(i) Tettigometridae
There is an overall and dramatic decline of tettigometrid species without apparent
reasons, although habitat destruction is involved in a few cases. Most older collections
include considerable numbers of several species (Kirschbaum 1868; Lindberg 1948; MTD;
Schiemenz 1969; Schwoerbel 1957; SMNS; Wagner 1935, 1939a, 1951; Schönitzer & Oester-
ling 1998a; ZMB). Tettigometra obliqua (Panz.) was particularly numerous and even re-
ported to cause cereal damages, but the last German record dates back to the 1970s.
There is also substantial evidence that Tettigometra fusca Fieb., T. impressopunctata Duf.,
T. macrocephala (Fieb.) and others have vanished from many sites without obvious hab-
itat changes. The causes are unknown, but all taxa prefer complex habitats, usually
comprising mosaics of sun-exposed, open woodland margins, patches of bare ground
and probably certain ant species.
(ii) Peatland species
A dramatic decline has also occurred in most peatland species after the draining of most
bogs of the north German plain and elsewhere, although habitat destruction was less severe
in the foothills of the Alps. Examples include Cixius similis Kbm., Ommatidiotus dissimilis
(Fall.), Deltocephalus maculiceps Boh., Cicadula quinquenotata (Boh.) and Sorhoanus xanthoneu-
rus (Fieb.). In some cases of substantiated local extinctions, notably in Nothodelphax albocar-
inata (Stål) and Limotettix atricapillus (Boh.), the habitats do still exist, but have become sub-
ject to heavy eutrophication, resulting in a change of the vegetation. The present occurrence
of some highly tyrphobiotic species on floating mats of old and overgrown turbaries creat-
ed by local farmers is particularly noteworthy. These species originally prefer the vicinity of
hollows, which have disappeared from many bogs after draining. In some localities, the
 Declines 231

Table 3. Declining Auchenorrhyncha species in Germany. Present status mainly after Remane et al. (1998),
possible factors after Nickel et al. (1999).

Species Region of decline Present status Possible factors

Cixius stigmaticus (Germ.) Most parts Endangered Forestry

Cixius similis Kbm. North Germany, Thuringia Vulnerable Draining of peatland
Myndus musivus (Germ.) Rhine, Main, Weser, Elbe Critical Channeling of rivers
Nothodelphax albocarinata (Stål) Eastern half of Germany Critical* Peatland
eutrophication
Xanthodelphax flaveola (Fl.) Most parts Endangered Grassland
improvement
Pseudodelphacodes flaviceps (Fieb.) Lech floodplains Critical Channeling of rivers
Javesella salina (Hpt.) Eastern Germany Endangered Grassland
improvement
Tettigometra macrocephala (Fieb.) Thuringia, Franconia Critical Unknown
Tettigometra atra Hag. Most parts Critical Unknown
Tettigometra fusca Fieb. Thuringia, Bavaria, Baden Endangered Unknown
Tettigometra impressopunctata Duf. Thuringia, Bavaria Endangered Unknown
Tettigometra leucophaea (Preyssl.) Most parts Probably extinct* Unknown
Ommatidiotus dissimilis (Fall.) Northern Germany Endangered Draining of peatland
Tibicina haematodes (Scop.) Baden-Württemberg Critical Viticulture, habitat
destruction
Macropsis impura (Boh.) Most parts Vulnerable Wetland improvement
Macropsidius sahlbergi (Fl.) Eastern Germany Critical Habitat destruction
Anoscopus histrionicus (F.) Eastern half of Germany Endangered Unknown
Coryphaelus gyllenhalii (Fall.) Thuringia, Bavaria Critical Habitat destruction
Deltocephalus maculiceps Boh. Northern Germany Critical Draining of peatland
Doratura horvathi W.Wg. Southern Niedersachsen Vulnerable Cessation of grazing
Platymetopius undatus (De G.) Most parts Endangered Unknown
Mimallygus lacteinervis (Kbm.) Iller floodplains Critical Channeling of rivers
Hardya tenuis (Germ.) Most parts Endangered Unknown
Sardius argus (Marsh.) Western parts Critical Unknown
Cicadula quinquenotata (Boh.) Northern Germany Endangered Draining of peatland
Mocydiopsis intermedia Rem. Thuringia Vulnerable Unknown
Athysanus quadrum Boh. Most parts of Germany Endangered Wetland improvement
Limotettix atricapillus (Boh.) Eastern half of Germany Critical Peatland
eutrophication
Laburrus impictifrons (Boh.) Eastern half of Germany Endangered* Habitat destruction
Psammotettix unciger Rib. Iller floodplains Critical Channeling of rivers
Psammotettix pallidinervis (Dhlb.) Most parts Endangered Unknown
Errastunus leucophaeus (Kbm.) Iller floodplains Critical Channeling of rivers
Pinumius areatus (Stål) Brandenburg and Mainz Critical Unknown
Sorhoanus assimilis (Fall.) Most parts Vulnerable* Wetland improvement
Sorhoanus xanthoneurus (Fieb.) Northern Germany Endangered Draining of peatland

* = not identical with status suggested by Remane et al. (1998)

232 Review of species

only populations of Macrosteles fieberi (Edw.) and Limotettix atricapillus (Boh.) were confined
to such turbaries, but were absent from other parts of the bog (Nickel 1997, 1999b). Further,
the improvement of fen habitats during recent decades was probably equally severe and
affected a large number of species, although evidence is only indirect in most cases, except
in Macropsis impura (Boh.), Athysanus quadrum Boh. and Sorhoanus assimilis (Fall.).
(iii) Species of river banks
The regulation of rivers, altering their natural flooding dynamics, has brought some
species to the brink of extinction. This is particularly true for Psammotettix unciger Rib.
and Errastunus leucophaeus (Kbm.), which are both endemics of the northern Alps, and
Pseudodelphacodes flaviceps (Fieb.) and Mimallygus lacteinervis (Kbm.). Direct evidence of
decline is lacking for Pentastiridius beieri (W.Wg.), which has probably been overlooked by
earlier collectors. All these species live on almost bare gravel banks with extremely scat-
tered vegetation (see Nickel 1999a). Myndus musivus (Germ.) shows similar habitat re-
quirements. To the north of the Alps, however, it is apparently confined to sand banks
along lowland rivers. Until 1950, the species was found in a number of sites along the
Rhine, Weser, Elbe and some of their tributaries, but, despite intensive search, the only
recent records are from the Elbe valley near Meißen. None of these species has been found
to colonize secondary anthropogenic habitats, like sand pits, field margins and fallow
fields, except Pentastiridius beieri (W.Wg.) and Javesella stali (Metc.) (Nickel et al. 2002a).
(iv) Heliophilous grassland species
There is a decline of heliophilous species living on dry calcareous or siliceous grass-
land formerly subject to grazing. Sheep keeping has decreased over large parts of central
Europe during recent decades, and sward height has increased on former pastures with
short grass and some bare ground. Mostly the evidence for decrease of Auchenorrhyncha
is only indirect, but in some substantiated cases, species associated with Festuca ovina,
Helictotrichon pratense and Thymus spp., such as Kosswigianella exigua (Boh.), Doratura hor-
vathi W.Wg. and Goniagnathus brevis (H.-S.) have vanished from formerly grazed sites in
southern Niedersachsen and elsewhere.
(v) Psammobiotic and psammophilous species
Some species associated with sandy soils suffered a dramatic decline during recent
decades, particularly Macropsidius sahlbergi (Fl.), Handianus ignoscus (Mel.) and Pinu-
mius areatus (Stål). Furthermore, the only known German population of Psammotettix
angulatus (Then) could not be confirmed during a recent visit and may have become
extinct. The reasons for this decline are not certainly known. Most habitats are still pre-
served, but the vegetation cover is likely to have become more dense and tall, perhaps
due to nitrogen immission, cessation of grazing, or even restriction of public access.
(vi) Elm-dwelling species
Dutch elm disease is caused by the ascomycete fungi Ophiostoma ulmi (Buisman) Nann-
feldt and O. novo-ulmi Brazier, and is transmitted by bark beetles of the genus Scolytes.
Due to the pathogen’s rapid evolutionary changes, several waves of attack have occurred
throughout the temperate deciduous forest zone of the northern hemisphere, and future
attacks have been predicted (Brasier & Buck 2001). Ulmus minor, U. glabra and their hy-
brid U. x hollandica are particularly susceptible, whereas U. laevis is apparently less con-
cerned. Altogether, the density of elms in central Europe may have declined to less than
10% during the past century, and many millions of trees have died (Karnosky 1979; Röh-
 Range expansions 233

rig 1996; U. Gruber, pers. comm.). Although the total distribution of the three central Eu-
ropean elm species is probably not affected (see Haeupler & Schönfelder 1989; Benkert et
al. 1996), the population density has decreased dramatically, and in many localities only
coppices or young trees are left, most of which become re-infested by the fungus as soon
as their stems are thick enough to harbour the beetle. Thus, a considerable decline of
specialized elm-dwelling leafhoppers, comprising at least 9 monophages, must have oc-
curred. On the other hand, many of the remaining elm stands still show a considerable
infestation of several leafhopper species, and Kyboasca bipunctata (Osh.) even seems to
prefer coppices growing after the chopping of dying trees.
(vii) Woodland species
Cixius stigmaticus (Germ.) and Platymetopius undatus (De G.) suffered a severe decline
at least since 1970. The reasons are unknown. They both ascend to the canopy layer
after emergence and may require a more complex habitat structure.

4.4.2 Range expansions

Statements on range expansions must be drawn with care due to the shortage of
faunistic information. Records of new species, which either have been misidentified
formerly or which require special habitats or host plants, are constantly being added to
many faunal lists even in central Europe. In most cases, this is not caused by range
expansions, but rather by the increase of taxonomic and ecological knowledge. Usually,
this phenomenon can be definitely proven by the study of older collections. For in-
stance, Kelisia sima Rib., Florodelphax paryphasma (Fl.), Macropsis najas Nast, Alebra coryli
Le Q., A. viridis R., Zygina griseombra Rem., Z. nirgitarsis Rem., Balclutha calamagrostis
Oss., Streptanus confinis (Reut.) and Arthaldeus arenarius Rem. have all been found to be
common and widespread in most parts of central Europe shortly after publication of
the relevant information, and most of them also turned up in museum collections. In
other cases, however, there are good arguments and sufficient data for a real range
expansion. The following assessment is also based on a careful evaluation of negative
records in collections and older publications, notably MTD, SMNS, ZIMH, ZMB, ZSM,
Haupt (1935), Kirschbaum (1868), Kuntze (1937) and Wagner (1935, 1939, 1941a, 1951).
Mere expansions within central Europe are documented for Eurybregma nigrolineata Scott
(but see 4.1.2.4), Haematoloma dorsatum (Ahr.), Viridicerus ustulatus (M. & R.), Liguropia juni-
peri (Leth.), Ribautiana debilis (Dgl.), Zyginella pulchra P. Löw, Zyginidia scutellaris (H.-S.) and
Fieberiella septentrionalis W.Wg. (section 4; see also Niedringhaus & Olthoff 1986; Remane
1995; Remane & Fröhlich 1994b). Some of these species prefer anthropogenic or otherwise
disturbed habitats, often with introduced plants. E. nigrolineata Scott and Z. scutellaris (H.-
S.) mainly live on grasses along waysides and on leys, V. ustulatus (M. & R.), R. debilis (Dgl.),
Z. pulchra P. Löw and F. septentrionalis W.Wg. occur in natural habitats in southern parts of
Germany, but prefer urban sites along the expansion front line. H. dorsatum (Ahr.) is the only
expanding species, which avoids anthropogenic sites and prefers closed forests. The Medi-
terranean Liguropia juniperi (Leth.) lives on the ornamental Chamaecyparis lawsoniana origi-
nating from North America and must have invaded only after the host’s introduction.
Range expansions have also been suggested for Macropsis megerlei (Fieb.) and Balca-
nocerus larvatus (H.-S.), which both live on Rosa species and Prunus spinosa. Their host
234 Review of species

plants, however, are very difficult to sample with the sweep net. Moreover, there are old
records of both species from southern Norway and Mecklenburg, respectively (Kuntze
1937; Ossiannilsson 1981; Remane & Fröhlich 1994b; Nickel 1997). Edwardsiana sociabilis
(Oss.) is now common and widespread in urban habitats on ornamental Rosa rugosa and
has even been reported as a pest in Finland. Originally, it was probably confined to
Filipendula ulmaria in rather moist habitats (Nuorteva 1955; Remane & Fröhlich 1994b).
It is not clear, however, whether its host shift caused a real expansion of the geographic
range or merely an increase of abundance.

4.4.3 Introductions
There is no doubt that Stictocephala bisonia Kopp & Yonke and Graphocephala fennahi
Young have been introduced from North America, and that Macropsis elaeagni Em. has
been introduced from central Asia. Another interesting example is Opsius stactogalus
Fieb., which was formerly confined to Myricaria germanica along banks of unregulated
alpine rivers. At least since the second half of the 19th century, it has spread over most
lowland areas, living on ornamental Tamarix in urban habitats.
Introduction, at least range expansion, is also likely in some Eupteryx species living
on archaeophytic spice and medicinal herbs. E. decemnotata R., which was only known
from Mediterranean regions of France and Italy a few years ago, has appeared in Swit-
zerland, most parts of Germany and eastern parts of Austria. There is further evidence
of introduction for E. melissae Curt., which is another Mediterranean species, for
E. artemisiae (Kbm.), which occurs mainly on salt meadows of the European coastline,
as well as Austroasca vittata (Leth.) and E. adspersa (H.-S.), both probably originating
from central Asia, although earliest records from interior central Europe of these spe-
cies date far back (section 4.2.4.12). Another Mediterranean species, E. rostrata Rib., has
recently been collected in the Netherlands, without details being known (Bieman &
Rozeboom 1993). Very recently, a population of Eupteryx salviae Arz. & Vid., which is
otherwise only known from the Adriatic region, has been found in a garden in south
Germany. In 2001, the Mediterranean typhlocybid Hauptidia provincialis (Rib.) was found
in greenhouses in Ulm feeding on corn salad (Valerianella) and perennial basil (Oci-
mum). It is not yet clear whether these populations will be stable.
Various authors consider Endria nebulosa (Ball) and Japananus hyalinus (Osb.) to be
introduced from the Nearctic, the latter originally having come from Japan (see Arzone
et al. 1987). They argue that their systematic position in Europe is rather isolated, and
that their discovery happened to be rather late. However, these species are both mono-
phagous on native European plants in more or less natural habitats, and their abun-
dances are usually low. E. nebulosa (Ball) has a rather cryptic way of life, preferring the
very base of its hostplant. J. hyalinus (Osb.) was first recorded in Europe in 1942 (Wag-
ner & Franz 1961) and has since been found mostly on the native Acer campestre, al-
though some records are from ornamental Acer (section 4.2.4.13). Incidentally, there is
quite a number of species with most relatives living in the Nearctic region, e.g. Empoas-
ca, Ossiannilssonola, Zonocyba, Deltocephalus, Colladonus and Limotettix, without being
assumed to be introduced. Thus, the evidence for introduction in these two species is
considered here as inconclusive.
 Occasional influxes 235

Further Nearctic species may invade in the future, notably the flatid Metcalfa pruinosa
(Say) and the deltocephaline Scaphoideus titanus Ball. Both are now common and wide-
spread in parts of France and Italy and in some other south European countries, and both
have a considerable potential for being noxious to cultivated plants (section 5.5). The typhlo-
cybine leafhopper Kyboasca maligna (Walsh) was found in Alsace (France), close to the Ger-
man border in 1997 and in the French Ardennes in 2000 (della Giustina & Remane 2001). In
recent years, the eastern Palearctic typhlocybine Vilbasteana oculata (Ldb.) has appeared in
Moscow on ornamental Syringa (Tishechkin 1989) and may spread westward.

4.4.4 Occasional influxes

The Mediterranean issid species Agalmatium bilobum (Fieb.) was common in dune
habitats of the East Friesian island of Wangerooge at least in 1960, but was not found
afterwards (Harz 1965, 1988; Niedringhaus 1991). Balclutha saltuella (Kbm.) was described
from a single female from the vicinity of Wiesbaden in the second half of the 19th centu-
ry. Only 130 years later, it happened to be recorded again in Germany; but despite three
records from widely separated localities, all specimens were apparently vagrants with-
out evidence for reproduction (Nickel 1999a). This species is also known to occur errat-
ically in other parts of Europe. Likewise, an apparently single specimen of Zyginidia
pullula (Boh.) found in eastern Bavaria may have immigrated from more southern or
eastern parts of Europe. Toya propinqua (Fieb.) breeds in the upper Rhine plain at least in
some years, but single specimens have also been found in southern Bavaria. Single indi-
viduals of the Mediterranean Cicada orni L. outside their known range were also record-
ed, but not sampled (P. Dynort, pers. comm.; Heller 1987c). Back (1976) and Hess (pers.
comm.) observed single individuals of Tibicina haematodes (Scop.) on dispersal flight in
southern Germany. Some older, unconfirmed records of further cicada species (Duffels
& van der Laan 1985) may refer to migrating specimens.
Autochthonous occurrence in Germany is also uncertain in Zygina tithide Ferr., Fruti-
cidia sanguinosa (R.) and Circulifer opacipennis (Leth.), which may occur only temporarily
as vagrants. In contrast, Achorotile albosignata (Dhlb.), Megadelphax haglundi (J. Shlb.)
and Megamelodes lequesnei W.Wg., are probably native, although there are only single
specimens. Most of these were brachypterous, and in addition, these species either live
among thorny scrub or show a hidden way of life, their host plants being unknown.

4.5 General remarks on the German Auchenorrhyncha fauna

At present, altogether 620 Auchenorrhyncha species are known and verified from
Germany, although in a few cases, there are uncertainties regarding the correct nomen-
clature. Considering the fact that more than 40 new records, 4 of which were new orig-
inal descriptions, have been added just during the past decade, a reasonable estimate of
the real species number should certainly exceed 650 and may even approach 700. Dubi-
ous species were not included in the total number. Such a list could be easily extended
by checking old literature (e.g. Hüeber 1904; Haupt 1935; Melichar 1896). Even Nast
(1972, 1987) listed a number of species, which could not be verified and which are likely
 236 Review of species

to refer to an uncritical adoption of old records. Some of these dubious records are
discussed in section 4.3.
Five species, Toya propinqua (Fieb.), Agalmatium bilobum (Fieb.), Cicada orni L., Zy-
ginidia pullula (Boh.) and Balclutha saltuella (Kbm.), are apparently irregular immigrants,
with reproductive success at most in favourable years. At least for three further species,
namely Stictocephala bisonia Kopp & Yonke, Macropsis elaeagni Em. and Graphocephala
fennahi Young, an introduction over long distances by human activities is documented
(section 4.4.3). In other cases, such as Eupteryx decemnotata R., E. salviae Arz. & Vid. and
Hauptidia provincialis (Rib.), all or most known populations are synanthropic. The near-
est populations in natural habitats live in the Mediterranean region. Thus, an active
introduction cannot be documented, but is likely.
An overview of the verified species and their systematic affiliations is given in Fig. 2.
Accordingly, 6 families of Fulgoromorpha and 4 families of Cicadomorpha are repre-
sented (with cicadas and froghoppers interpreted as families, respectively). The Cicadel-
lidae are the largest subgroup, with 452 species and 13 subfamilies. Altogether, species
numbers are highest in the Deltocephalinae, Typhlocybinae and Delphacidae. These
three groups mainly comprise species feeding on graminoids and deciduous trees, i.e.
the most apparent plants in central Europe, and alone account for almost 75% of the
Auchenorrhyncha species total. Some more tropical subgroups, which are present in
southern Europe, notably the Meenoplidae, Derbidae, Ricaniidae, Flatidae, Tropi-
duchidae and Stegelytrinae, are entirely absent from Germany. Likewise, some groups,
which are rather diverse in southern Europe, e.g. the Issidae, Cicadidae, and – to a
lesser extent – the Dictyopharidae, are represented by few species only.
250

Fulgoromorpha Cicadomorpha
(S = 145) (S = 475) 194
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Species number

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 General remarks and problems 237

5 Utilization of plant resources

5.1 Plant taxa

5.1.1 General remarks and problems

In view of more than 600 species of Auchenorrhyncha and roughly 3000 plants, re-
sulting in 1.8 million possible associations, the approach of assessing the overall rela-
tionships between a rather diverse insect group and a whole flora in a large geograph-
ical area such as Germany may seem out of reach. But in fact, only a relatively small
proportion of the taxonomic plant diversity is exploited, and the data base on the more
relevant groups is often sufficient or at least moderate, though not in all parts of the
study area. Nevertheless, there are substantial gaps in the knowledge of plant prefer-
ences of many polyphages and of all species spending the nymphal stage in the soil,
such as the Cixiidae, Cicadidae and Cercopinae. Furthermore, there is a general scarcity
of specific data from plant species confined to southern parts of Germany, notably the
alpine and subalpine belt as well as viticultural regions of the Rhine valley and its trib-
utaries. For instance, alpine grasses, sedges and willows, as well as extensive forests of
Castanea sativa along the upper and middle Rhine may provide ample resources, which
are not yet studied. Further sampling is also needed on a number of more widespread
plants, such as Abies alba, Ulmus laevis and even Betula pubescens. In contrast, the lack of
Auchenorrhyncha data from rare or localized plants is probably of minor significance,
since almost all species live on more widespread and abundant plants. Exceptions from
this rule are few, and comprise, for instance, the delphacids Kelisia minima Rib., living
on Carex distans, and Nothodelphax albocarinata (Stål), found on Carex limosa.
Another difficulty is the evaluation and classification of food plant records of phy-
tophagous insects in general and the resulting problems for analysis, which were dis-
cussed by Ward (1988) and, more extensively, by Schoonhoven et al. (1998). The major
problems will be discussed in the following sections.
(i) Uncertainties regarding the feeding substrate
Although the majority of central European Auchenorrhyncha species are rather ho-
mogeneous in feeding, there are some exceptions. The Achilidae are considered to suck
on fungi mycelium. At least in Empoasca decipiens Paoli feeding is known on phloem as
well as on mesophyll. Prestidge (1982), without experimental studies, reported xylem-
feeding in Euscelis incisus (Kbm.). However, deltocephaline leafhoppers and Euscelis
spp. in particular, which were reared by Müller, Strübing and Witsack for decades, are
238 Utilization of plant resources

widely considered as phloem-feeders. In fact, systematic experimental studies on the

type of substrate have been done only in a few species, most of which are of economic
importance, although generally, there is no reason to question classifications made by
Müller and co-workers, who determined food substrates after the type of excrements:
greenish (due to chlorophyll) in mesophyll feeders, watery both in xylem and phloem
feeders, but viscous due to the surplus of sugar in the latter (Müller 1956).
(ii) Dependence on developmental stage and season
Some species utilize different plants for oviposition, larval development, adult feed-
ing, wintering, or even in the first and the second generation. These shifts can be either
facultative or obligate. Muellerianella fairmairei (Perr.) develops on Holcus species, where
at least some individuals of the first generation lay their eggs. Oviposition of the second
generation, however, obligatorily takes place on Juncus effusus (Drosopoulos 1977, and
pers. comm.). Many vertical migrants feed on grasses and other herbaceous plants as
nymphs, and on various trees and shrubs as adults, e.g. Haematoloma dorsatum (Ahr.),
Stictocephala bisonia Kopp & Yonke, Allygus spp., Allygidius spp. and Hesium domino (Reut.).
Many Zygina species, being monophagous as nymphs on deciduous woody plants, feed
on conifers as wintering adults and on various woody Rosaceae in spring (mainly Cratae-
gus spp., Prunus spinosa and P. padus). Also, Balclutha punctata (F.), Empoasca vitis (Göthe)
and probably some species of Idiocerinae overwinter on conifers. B. punctata (F.) develops
on Poaceae, the others on deciduous trees and shrubs of various taxa. Claridge & Wilson
(1978a) and Vidano et al. (1990) demonstrated food plant shifts between the first and the
second generation, which may be due to the plant’s suitability for egg overwintering. In
Wales (and probably in central Europe, too), Lindbergina aurovittata (Dgl.) migrates from
evergreen Rubus species to Quercus and other deciduous trees, and back. In some species
of Eupteryx, the first generation is monophagous, the second, however, is oligophagous or
even polyphagous. Thus, nymphs of the first generation of Eupteryx aurata (L.) are appar-
ently confined to Urtica dioica, whereas the emerging adults colonize dicotyledonous plants
of numerous families. Perennial populations of Eupteryx filicum (Newm.) only occur on
the evergreen fern Polypodium, but adults of the first generation may found a second gen-
eration also on hemicrytophytic fern species. In Eupteryx thoulessi Edw., most populations
are found on Mentha aquatica, but at least summer individuals (including nymphs) also
occur on Lycopus europaeus, where both plants grow together. A possible explanation for
this host shift (or more precise: diet width expansion) may be differing suitability of plant
species for egg overwintering. This seems rather evident in Eupteryx filicum (Newm.), for
which hemicryptophytic ferns can not provide substrate for winter eggs.
(iii) Subterranean feeding
With few exceptions, e.g. Haematoloma dorsatum (Ahr.) – see above – the host plants of
subterranean nymphs of Cixiidae, Cicadidae, and Cercopinae are completely unknown.
There is evidence, that at least some species feed on woody plants. But if this is true,
more tight host relations are possible or even likely, simply because of the limited abil-
ity of small insects to move through the soil for longer distances and into another tree’s
rhizosphere. So far, most of these species are tentatively considered as polyphagous.
(iv) ‘Vagrants’, ‘tourists’, ‘migrants’
Migrating and drifted individuals, as well as those fallen down from the canopy
above have caused many erroneous food plant records in the literature. For a long time,
 General remarks and problems 239

such records concealed the monophagy of a number of species, for instance in most
idiocerine leafhoppers.
(v) Geographic differences
Many examples of stenophagous insects feeding on different plant species in differ-
ent parts of their range have been described. For Auchenorrhyncha, however, the most
widespread phenomenon seems to be a change of the diet breadth. In some cases, it has
been shown that the host range becomes narrower towards the north, thus causing a
‘geographical monophagy’ in regions, where only one out of several potential host spe-
cies occurs. For instance, Eurhadina kirschbaumi W.Wg. is strictly monophagous on Quer-
cus petraea in most parts of central Europe. In Mediterranean regions, however, it feeds
additionally on various endemic oak species and Castanea sativa (Drosopoulos 1999;
Vidano & Arzone 1987b). Thus, it would be classified here only as 1st degree oligopha-
gous. Extensive field data on the delphacid planthopper Hyledelphax elegantula (Boh.)
revealed a decrease in diet width along a northward gradient through central Europe.
In the lowlands of southern and middle parts of Germany host plants include various
grasses (Calamagrostis arundinacea, Brachypodium pinnatum, Holcus mollis, Molinia caer-
ulea, Deschampsia flexuosa and others). At higher altitudes and in the north German plain,
however, the latter is the only host. Some species of the genera Chloriona and Euides,
which live strictly monophagous on Phragmites australis in central Europe, have been
reported from Arundo donax in Greece (Drosopoulos et al. 1983). Adults of Haematoloma
dorsatum (Ahr.), which has been found exclusively on Pinus sylvestris in Germany, live
on Pinus nigra and other conifers including Juniperus in southern and western Europe
(Moraal 1996). Examples of geographic monophagy may be more widespread, but re-
quire a broad data base, which rarely exists. Other insect groups have been reviewed by
Schoonhoven et al. (1998).
(vi) Host shifts
In central Europe, there are only few reported cases of host shifts of stenophagous
Auchenorrhyncha species correlated with anthropogenic changes, although this phe-
nomenon has been documented in other insect groups (see Strong et al. 1984). Myricaria
germanica, a native plant on gravel banks of alpine rivers, is the original host of Opsius
stactogalus Fieb. At least from the second half of the 19th century onwards, it was found
to live on ornamental species of Tamarix in parks and gardens (e.g. Kirschbaum 1868).
Edwardsiana sociabilis (Oss.) is widespread on ornamental Rosa rugosa, which was intro-
duced from eastern Asia. The original host is Filipendula ulmaria and perhaps Rubus
idaeus. Macropsis megerlei (Fieb.), a monophage of native roses, is another species which
accepted Rosa rugosa as a further host. The situation of Graphocephala fennahi Young,
which has been introduced from the Nearctic, is not quite clear, but a host shift needs
not necessarily be involved. Although the species is reported to live on cultivated Rhodo-
dendron ponticum, a native to southern Europe and Asia Minor, many ornamental plants
in central and western Europe show introgression towards North American species or
are grafted onto North American stems (H. Bruelheide, pers. comm.). The typhlocybid
leafhopper Eupteryx artemisiae (Kbm.) is another case. It mainly lives on Artemisia marit-
ima on salt meadows near the coast, but occasionally on A. abrotanum in gardens far
inland. However, it is also reported from southeastern Europe, which is the geographi-
cal origin of A. abrotanum. Liguropia juniperi (Leth.), originally living on scaly Cupres-
240 Utilization of plant resources

saceae in the Mediterranean region (R. Remane, pers. comm.), has been found on the
Nearctic Chamaecyparis lawsoniana in cemeteries and parks of southwestern Germany,
where none of the natural host plants occur.
(vii) Differing evidence from field and laboratory studies
Laboratory rearings of insects may be successful on plants never utilized in the field.
For planthoppers, this has been demonstrated in the genera Muellerianella und Ribautodel-
phax (Booij 1982; Bieman 1987b). Most species of both genera could be reared on various
Poaceae species over several generations, although rates of survival and reproduction
were reduced, and larval development was prolonged in most, though not all cases.
(viii) Exceptional field records
For a few monophagous Auchenorrhyncha species, there are definite records of feed-
ing or even development on plants other than their usual hosts. For instance, there is an
October record of numerous adults and nymphs of Muellerianella extrusa (Scott) in pure
stands of Deschampsia flexuosa near a peat bog, where large populations fed on the usual
host Molinia caerulea, and another record of the same species, with nymphs, on Agrostis
stolonifera, away from Molinia (Nickel, unpublished data). Near the very border of the
range, triploid adults and nymphs of the closely related M. fairmairei (Perr.), usually
being confined to Holcus, were found on D. flexuosa (R. Remane, pers. comm.). Howev-
er, both species are widely reported to be strictly monophagous (Drosopoulos 1977;
Booij 1982), and from Germany alone, there are more than 50 field records of both spe-
cies, respectively, on the substantiated host. Nevertheless, these single occasions may
be more widespread and deserve further attention. In combination with results of labo-
ratory rearings (Booij 1982; Bieman 1987b – see above), they may provide evidence for
an ‘evolutionary back door’ that potentially enables monophagous species to escape
the drawbacks of specialization. On the other hand, differences in host preference be-
tween populations and even individuals are known in some Lepidoptera species
(Schoonhoven et al. (1998).
(ix) Sampling bias caused by plant apparency
Abundant and widespread plants are more strongly utilized by phytophagous in-
sects than uncommon and scattered ones. But plant apparency also affects collecting
habits of entomologists: common and large plants are more likely to be encountered,
sampled and identified. Rare plant species are in turn little studied, because they are
less known and less likely to be found by collectors.
(x) Study subject and students’ experience
Some plant taxa, notably Poaceae and Cyperaceae, are poorly known to most ento-
mologists. In most studies of taxonomy and even ecology of Auchenorrhyncha, trophic
relationships with plants are simply ignored. Many records refer only to plant genera or
even family, which rarely means that the insect has been found on several plant species
of one genus, but rather that the host plant was only identified to generic or family
level, respectively. In this study, for instance, a number of Auchenorrhyncha species,
which have hitherto been considered as broadly monophagous on various species of
the genus Carex, have been found to be confined to single species.
(xi) Plant taxonomy and diet breadth categories
Every classification of died breadth of non-monophages has to face inconsistencies
caused by plant taxonomy. As a consequence, purely nomenclatural changes in plant
 General remarks and problems 241

taxonomy can move a phytophage from one category of diet breadth into another. For
instance, depending on the nomenclatural interpretation of alder buckthorn (Frangula
alnus Mill. or Rhamnus frangula L.), the typhlocybine leafhopper Zygina suavis R. may be
either treated as 1st degree oligophagous or 2nd degree monophagous. Further, speci-
ficity on a certain level in one plant group is necessarily not the same as in another
group. These problems were treated by Symons & Beccaloni (1999), who discuss phylo-
genetic indices to measure diet breadth. These indices, however, are only practicable, if
plant phylogeny is well known.
Most authors distinguish between ‘food plants’ and ‘hosts’. The first refer to those
species, from which food is taken, the latter are species on which nymphal or larval
development, and usually also oviposition take place. For the majority of central Euro-
pean Auchenorrhyncha, both terms can be used as synonyms, except for those migrat-
ing to other plants after emergence. The term „guild“ is originally defined “as a group
of species that exploit the same class of environmental resources in a similar way …
without regard to taxonomic position” (Root 1967). Commonly used classifications of
phytphagous insects mainly distinguish between leaf-chewing, sap-sucking, leaf-min-
ing and stem-boring guilds (Root 1967; Strong et al. 1984), although there are many
other connotations (see Simberloff & Dayan 1991). In this book, it is applied for various
species utilizing the same plant taxon, regardless of the utilized type of plant sap.
The major part of the food plant data basis for the present work, altogether compris-
ing almost 9,000 single records, was gathered by the author. With few exceptions (e.g.
Biedermann 1998a; Weber & Maixner 1998; Witsack 1985), literature records were checked
critically (see above) and included only after verification. Also, vague remarks like
‘polyphagous on Gramineae’ have been ignored when no evidence was provided. Ex-
tensive suction sampling and direct observations in the grass layer during recent years
have shown or confirmed that a large proportion of species is strictly associated with
one or few species of Poaceae or Cyperaceae, although it can not be ruled out that adults
may occasionally feed on plants other than their hosts. However, Booij (1982) and Bie-
man (1987b) demonstrated for the delphacid genera Muellerianella and Ribautodelphax,
that monophagous species may be reared on other plants, but with reduced rates of
survival and reproduction. Furthermore, in choice experiments, the field host was al-
ways preferred. But in general, there is still a considerable deficit in understanding
food plant relationships of phytophagous insects, in particular of those feeding on Poace-
ae and Cyperaceae, which are considered to be poor in secondary compounds, but rich
in silicate content (Hegnauer 1963; Tscharntke & Greiler 1995).
In the following chapters, the important plant groups exploited by central European
Auchenorrhyncha are enumerated, with introduction and discussion of their associat-
ed guilds. Informations on distribution and ecology of plant species were combined
after Benkert et al. (1996), Ellenberg (1996), Haeupler & Schönfelder (1989) and Ober-
dorfer (2001). Species groups (“agg.” sensu Wisskirchen & Haeupler 1998) were treated
as a single taxon, e.g. Ranunculus auricomus, Alchemilla spp. and Rubus fruticosus. Taxon-
omy and systematic order follow Wisskirchen & Haeupler (1998), although phyloge-
netic aspects will be discussed on the basis of Judd et al. (1999) (see chapter 5.1.3). The
classification of the diet breadth applied here is shown in Table 4 and follows Schaefer
(1992, slightly modified). Like other classifications, it is unable to solve the problem of
242 Utilization of plant resources

Table 4. Classification of diet breadth applied in this book, after Schaefer (1992), modified

Diet breadth Classification Abbreviation

1 host species 1st degree (= strictly) monophagous m1

1 host genus 2nd degree monophagous m2
1 host family 1st degree oligophagous o1
2 food plant families or up to 4 species 2nd degree oligophagous o2
which belong to no more than 4 plant families
More Polyphagous po

inadequate definitions of supraspecific plant taxa. A host record is defined here as at

least three nymphs or at least 10 adults, respectively, on an identified plant species.
Records are treated as uncertain, if they have been gathered under unfavourable condi-
tions, particularly after windy weather or late in the season, or if they largely consisted
of parasitized individuals. An insect – plant relationship is assumed here, if there are at
least 3 host records (or, alternatively, food plant records in vertical migrants), marked
with “x” (or “X”, for at least 10 records) in the following tables. Less than 3 records are
treated as uncertain relationships, marked with “?”. Parantheses denote host relations
during the nymphal stage only (e.g. in species leaving the nymphal host after emer-
gence). Feeding records from outside Germany were also treated as uncertain due to
the possibility of regional changes. The assumption of an insect to be monophagous on
the basis of three records may appear critical, but it should be noted that for the major-
ity of species the record numbers are much higher, and that during recent years, almost
all species, which were only rarely reported in the literature, turned out to be mono-
phagous on little-studied or neglected plants.

5.1.2 Plant groups and their associated Auchenorrhyncha guilds

5.1.2.1 Pteridophyta
The pteridophytes represent a geologically very old group and occur in considerable
species numbers in many forest ecosystems worldwide. Ferns, in particular, can reach a
high dominance in the herbaceous layer of central European forests, notably where air
humidity is high. Their Auchenorrhyncha guilds, however, comprise only very few spe-
cies (Table 5). Apart from 4 specialists, non of which occurs in high frequency, utiliza-
tion of Pteridophyta by Auchenorrhyncha is a rather scarce phenomenon; there are only
accidental records of some polyphagous species. From an evolutionary point of view,
host relations between pteridophytes and Auchenorrhyncha specialists clearly repre-
sent a derived trait, since the latter belong to groups which mainly live on grasses,
sedges and other more advanced herb families. These examples can be considered as a
host shift back to more primitive plants.
The knowledge on Auchenorrhyncha living on pteridophytes in central Europe seems
sufficient for characterizing existing guilds, but their geographic distribution is only
imperfectly known. For instance, the delphacid planthopper Javesella stali (Metc.) and
 Pteridophyta 243

Table 5. Auchenorrhyncha species utilizing Pteridophyta in Germany. X = more than 10 host records (see
chapter 5.1.1), x = at least 3 host records, A = adult records only, W = winter records only, S = spring records
only, ? = feeding uncertain. Classification of diet breadth see Table 4, parentheses indicate host associations
during the nymphal stage only. Substrate: P = phloem, X = xylem, M = mesophyll, F = fungi mycelium;
Overwintering stage: eg = egg, ny = nymph, ad = adult. Asterisks indicate literature records ( x* = from
Germany, ?* = from adjacent countries).

At hy rium filix-femina

Overw intering stage

Pt eridium aquilinum

Poly podium v ulgare

Equiset um arv ense

Poly st ichum spp.

A. scolopendrium

Dry opt eris spp.

Asplenium spp.

P. int erject um
E. sy lv at icum

D iet breadth
E. palust re

Substrate
Species
Javesella stali (Metc.) X m 1 P ny
M acrosteles frontalis (Scott) X x* x* m 2 P eg Monophages
Ditropis pteridis (Spin.) X m 1 P ny
Eupteryx filicum (N ew m .) ?* ?* ?* ?* ?* ?* X ?* o1 M eg Oligophages
Lepyronia coleoptrata (L.) ?* x* po X eg Polyphages
Cercopis vulnerata Rossi A po X ny Vertical
Centrotus cornutus (L.) A po P ny m igrants
Javesella pellucida (F.) ? po P ny.
Philaenus spumarius (L.) ? po X eg Un-
Empoasca decipiens Paoli ? po M? ad certain
M acrosteles sexnotatus (Fall.) ? po P eg
Total 2 2 1 3 0 0 0 0 0 1 0
* = after Badmin (1991, 1992), Nickel (1979), Ossiannilsson (1981), Schiemenz et al. (1996), Stewart (1988)

the typhlocybid leafhopper Eupteryx filicum (Newm.) have both only recently been found
to be widespread in large parts of southern and middle Germany.
Two Auchenorrhyncha species are monophagous on Equisetum, namely Javesella stali
(Metc.) and Macrosteles frontalis (Scott). The former is strictly monophagous on field horsetail
(Equisetum arvense) in disturbed habitats, the latter is also found on marsh and wood
horsetail (E. palustre and E. sylvaticum). Bracken (Pteridium aquilinum) is infested by the
monophagous planthopper Ditropis pteridis (Spin.). Eupteryx filicum (Newm.) is oligopha-
gous on various species of Polypodiaceae in the second generation, although winter eggs
can apparently survive only on common polypody (Polypodium vulgare), in Britain also on
western polypody (P. interjectum), both of which are evergreen. Generalists include Cerco-
pis vulnerata Rossi, Centrotus cornutus (L.) and probably further species. In addition, there
are records of a few polyphages, such as Philaenus spumarius (L.), Lepyronia coleoptrata (L.)
and Empoasca decipiens Paoli, but this group might be underrecorded.
The poverty of herbivores on pteridophytes is a general phenomenon also documented
in other insect groups and has been explained by generally high concentrations of de-
fence compounds (Ottosson & Anderson 1983). Thus diverse insect guilds on single
species such as bracken (Lawton 1976, 1982) are exceptional. Ferns, and Polypodium
spp. in particular, are known to be rich in phytoecdysones, which may affect insect
moult, although the present evidence is still limited (Lafont et al. 1991).
244 Utilization of plant resources

5.1.2.2 Gymnospermae
Conifers, and in particular the Pinaceae, cover large areas of central Europe, although
only Norway spruce (Picea abies), Scots pine (Pinus sylvestris), and locally, silver fir (Abies
alba) play a major role as dominating tree species. Before man fundamentally altered the
vegetation cover, these species were largely confined to the montane belt of the Alps and
the higher Mittelgebirge as well as to extremely dry or sandy or peaty sites, where decidu-
ous trees cannot grow. Nowadays, all three have been planted on more favourable soils
throughout most of the lowland regions. Mountain pine (P. mugo) is the dominating wood
in subalpine scrub (also named “krummholz”) as well as in bogs of the foothills of the
Alps and in some of the Mittelgebirge. Black pine (P. nigra), originating from dry mountain
regions of southern Europe and southwestern Asia, has been planted all over Germany.
Stone pine (P. cembra) and European larch (Larix decidua) are both native only to the upper
montane belt of the Alps, but have been commonly planted in the lowlands.
Yew (Taxus baccata) is restricted to steep slopes in rather cool sites of the submontane
and montane belt; common juniper (Juniperus communis) is locally common in pastures,
heaths and open forests, whereas tamarix juniper (J. sabina) is only found in the alpine
belt of the Chiemgau and Allgäu (although also planted in gardens). Many other spe-
cies of gymnosperms have been introduced and are usually grown as ornamental trees
in gardens and parks, except the western North American douglas fir (Pseudotsuga men-
ziesii), which locally plays an important economic role in forestry. The German data
base on the Auchenorrhyncha fauna is moderate only for Picea abies and Pinus sylvestris.
Most of the remaining species have not or only rarely been sampled. An extensive over-
view and discussion of the Hemiptera on the Palearctic conifers, based on a large amount
of references, has been presented by Reuter (1909).
Auchenorrhyncha guilds on coniferous trees (in Germany represented only by Pi-
naceae, Cupressaceae, and Taxaceae) are shown in Table 6. Ten species are obligatorily
associated with this group; 4 of them are mainly, perhaps exclusively, associated with
Picea abies, including one vertical migrant. All of these are also mentioned for Abies alba
in the older literature, but these records require confirmation. Pinus sylvestris has 5 spe-
cialists, two of them – Cixidia confinis (Zett.) and Aphrophora corticea Germ. – show typ-
ical pine bark colouration. C. confinis (Zett.) does not feed on phloem, but on undeter-
mined fungi associated with pine. A. corticea Germ. and Haematoloma dorsatum (Ahr.)
are obligate vertical migrants, their nymphs developing also on dwarf shrubs or grass
roots, respectively. Thus, these species’ association with pine is obligate only through
adult feeding or oviposition. Wagneripteryx germari (Zett.) and Grypotes puncticollis (H.-
S.) live permanently in the canopy. The 4 latter species have also been found on P. nigra
in eastern Austria (W. Holzinger, pers. comm.). W. germari (Zett.) is locally abundant in
subalpine stands of P. mugo in Germany and is reported to live also on P. cembra in Swit-
zerland. Taxus baccata and Juniperus communis are both only utilized as winter food plants.
Liguropia juniperi (Leth.), which has been found only recently in southwest Germany,
lives on the Nearctic ornamental Chamaecyparis lawsoniana.
Most coniferous trees are important winter food plants for many species hibernating
as adults, e.g. Zygina spp., Empoasca spp. and probably some Idiocerinae. However, the
diet width of some of these species is narrow at least in the nymphal stage. Further
 Gymnospermae 245

Table 6. Auchenorrhyncha species utilizing Gymnospermae in Germany. Explanations see Table 5.

Chamaecy paris law soniana

Pseudot suga menz iesii

Overw intering stage

Juniperus communis
Pinus sy lv est ris

Taxus baccat a
Larix decidua

D iet breadth
Picea abies
Abies alba

P. cembra

Substrate
P. mugo
P. nigra
Species
Liguropia juniperi (Leth.) X m 1? M? ad
Pithyotettix abietinus (Fall.) ? X m 1? P ny
Perotettix pictus (Leth.) ? X m 1? P ny
Colobotettix morbillosus (Mel.) ? X m 1? P ny Mono-
Cixidia confinis (Zett.) X* m 1? F ny phages
Grypotes puncticollis (H .-S.) X ?** m1 P eg
W agneripteryx germari (Zett.) X ?** X ?** m2 M eg?
Cixius beieri W.Wg. ? A m 1? P ny
Haematoloma dorsatum (Ahr.) A ?** (o1) X ny Vertical
A phrophora corticea Germ . A ?** o2 X eg m igrants
Cixius similis Kbm . A ? o2? P ny
Balclutha punctata (F.) W ? W W (o1) P ad
Empoasca vitis (Göthe) ? W ? W ? ? W po P ad
M etidiocerus rutilans (Kbm .) W m2
(m2) P ad
Tremulicerus fulgidus (F.) W m1
(m1) P ad
Z ygina lunaris (M. & R.) W W m2
(m2) M ad Overw in-
Z ygina nigritarsis Rem . W W (m 1) M ad tering
Z ygina angusta Leth. W W W o2 M ad
Z ygina flammigera (Geoffr.) W W W o2? M ad
Z ygina rosincola (Cer.) W W W? o2? M ad
Z ygina rosea (Fl.) ?** W m 2?
(m2?) M ad
Tettigometra impressopunctata Duf. ? ? po P ad
Tettigometra virescens (Panz.) ?** ?** po P ad
Issus coleoptratus (F.) ? W W po P ny
N eophilaenus campestris (Fall.) ? o1 X eg
Dryodurgades reticulatus (H .-S.) ? m 1? P ad
Emposaca apicalis (Fl.) ?** ?** o1? M? ad Un-
Linnavuoriana decempunctata (Fall.) W? m2 P ad certain
Z yginella pulchra P. Löw W? m2 P ad
Z ygina suavis R. W? o1 M ad
Z ygina tiliae (Fall.) W? m2 M ad
A llygus mixtus (F.) ? po P eg
Thamnotettix confinis (Zett.) ? ? po P ny
Total (excl. overw intering species) 0 0 4 0 6 0 1 0 0 1 0
* = Nymphs feed on fungi mycelium; ** = after Günthart (1987b), Günthart & Günthart (1983), Holzinger (pers.
comm.), Nuorteva (1952b), Smreczynski (1954), Wagner (1941b)
246 Utilization of plant resources

winter food plants are perhaps ivy (Hedera helix), privet (Ligustrum vulgare) and other
evergreen broad-leaved woody plants, but they have only rarely been sampled. In ad-
dition, three species, notably Pithyotettix abietinus (Fall.), Perotettix pictus (Leth.), and
Colobotettix morbillosus (Mel.) – all living on Picea abies – are the only purely arboricolous
species of central Europe overwintering in the nymphal stage. It should also be men-
tioned that a number of species may escape drought on hot summer days by migrating
up into the tree layer. Thus, Neophilaenus campestris (Fall.) and Dryodurgades reticulatus
(H.-S.) have occasionally been collected on Pinus sylvestris.
Regarding the huge biomass provided by this plant group in many parts of the north-
ern hemisphere, the associated Auchenorrhyncha guilds seem rather poor in species.
As in the Pteridophyta, monophagous Auchenorrhyncha species taxonomically belong
to groups largely feeding on more advanced plants. Thus, a secondary host shift from
angiosperms to conifers must be postulated. The same conclusion was already drawn
by Reuter (1909) who found the same pattern in Palearctic Hemiptera, although partly
on the basis of obsolete literature. He also argued that most species feeding on conifers
probably evolved only during the Pleistocene and mentions two specific mirids with
close relatives living on Salix. However, the fact that a number of Auchenorrhyncha
specialists on Pinaceae and Cupressaceae such as Liguropia juniperi (Leth.), Pithyotettix
abietinus (Fall.) and others show a derived morphology, and that genera are either mo-
notypic or comprise exclusively conifer-feeding species, may provide evidence that these
groups are perhaps older.

5.1.2.3 Ranunculaceae
81 species of the buttercup family are reported from Germany. The majority of these
are perennial hemicryptophytes or geophytes, whereas therophytes, hydrophytes and
phanerophytes (sensu Raunkiaer 1907) are few. Only 7 species of Auchenorrhyncha have
been found to exploit this group, plus perhaps three uncertain ones, and only 4 plant
genera are utilized (see Table 7). Lesser meadow rue (Thalictrum minus) harbours Cercopis
sanguinolenta (Scop.) and Micantulina micantula (Zett.), the latter being the only host spe-
cialist on Ranunculaceae. Most feeding data are from Italy, where it is also reported from
further Thalictrum species (Vidano 1965). In most grassland sites creeping buttercup (Ra-
nunculus repens) is the main host of Eupteryx vittata (L.). Members of Ranunculus were also
found to be exploited by the polyphages Hyalesthes obsoletus Sign., Philaenus spumarius
(L.) and Erythria manderstjernii (Kbm.). Thus, host generalists and mesophyll feeders pre-
dominate. In general, the Ranunculaceae play only a minor role as resources for Auchen-
orrhyncha, despite their large species number and local dominance in biomass.

5.1.2.4 Ulmaceae
Only three species plus one common hybrid of the elm family occur in Germany, all
belonging to a single genus. Due to Dutch elm disease, caused by ascomycete fungi of
the genus Ophiostoma, all have declined dramatically in many parts of their range since
the 1920s. The current situation may be stable, though on a low population level, but
future waves of pathogen attack have been predicted (Brazier & Buck 2001).
 Ulmaceae 247

Table 7. Auchenorrhyncha species utilizing Ranunculaceae in Germany. Explanations see Table 5.

Overw intering stage

Ranunculus repens

Thalict rum minus

Clemat is v it alba
Calt ha palust ris

D iet breadth
R. mont anus
R. bulbosus

Th. flav um

Substrate
Species
M icantulina micantula (Zett.) ? m 1? M ad Monophages
Eupteryx vittata (L.) X o2 M eg Oligophages
Issus coleoptratus (F.) S po P ny
Philaenus spumarius (L.) x X po X eg Polyphages
Erythria manderstjernii (Kbm .) ? ? po M ad
Hyalesthes obsoletus Sign. x* po P ny Vertical
Cercopis sanguinolenta (Scop.) x* po X ny m igrants
A phrophora alni (Fall.) po X eg
Centrotus cornutus (L.) ? po P ny Un-
Empoasca decipiens Paoli ? po M? ad certain
Eupteryx atropunctata (Goeze) ? po M eg
Total 1 1 2 1 0 1 0
* = after Biedermann (1998a), Weber (pers. comm.)

Wych elm (Ulmus glabra) is locally common on moist but well-drained soils on shady
slopes of the colline and montane belt, although in many places only young trees are
found, which can not yet be attacked by elm bark beetles (Scolytes spp.), the common
vectors of the disease. Native stands of both smooth-leaved elm (U. minor) and Europe-
an white elm (U. laevis) are mainly confined to valley bottoms of rivers and streams.
The former is widespread all over cental Europe, the latter has two disjunct strongholds
in eastern Germany and the Rhine valley. All three species were commonly planted
along waysides and in parks, but many of these trees have died. The same is true for
Dutch elm (U. x hollandica), a formerly widespread hybrid between U. minor and U. glabra,
which was even more common than pure U. minor. However, its discrimination from
U. minor is very difficult, and many – if not most – food plant records from the latter
may in fact refer to the hybrid.
The data base of Auchenorrhyncha is moderate, although most published records
refer only to host genus level due to identification problems. The elm guild comprises
17 confirmed, plus 8 unconfirmed species (Table 8). With 6, 15 and 9 feeders on U. laevis,
U. minor and U. glabra, respectively, and 17 species altogether, the overall diversity on
Ulmaceae is fairly high. The ratio species number of Auchenorrhyncha : plants is very
high, surpassed only by the Fagaceae and Betulaceae. As on many other trees, the pro-
portion of mesophyll-feeders is significant, and most species overwinter in the egg stage.
A few species are apparently 1st degree monophagous, notably Ribautiana ognevi
(Zachv.) on U. laevis, and Macropsis glandacea (Fieb.) and Iassus scutellaris (Fieb.) on
U. minor (but probably also on the hybrid). Furthermore, there are 6 monophagous spe-
cies in the 2nd degree, namely Edwardsiana ulmiphagus Wils. & Clar., E. ishidai (Mats.),
248 Utilization of plant resources

Table 8. Auchenorrhyncha species utilizing Ulmaceae in Germany. Explanations see Table 5.

Ulmus laev is

D iet breadth

Overw inter-
U. minor*

Substrate
U. glabra

ing stage
Species
Ribautiana ognevi (Zachv.) X m1 M eg
M acropsis glandacea (Fieb.) ? X m 1? P eg
Iassus scutellaris (Fieb.) X m 1? P eg
Edwardsiana smreczynskii Dw or. ? ? m 2? M eg
Kyboasca bipunctata (Osh.) ? X ? m2
m2? M eg Monophages
Edwardsiana ulmiphagus Wils. & Clar. X X X m2 M eg
Edwardsiana ishidai (Mats.) X X X m2 M eg
Edwardsiana plebeja (Ed w .) X X X m2 M eg
Ribautiana ulmi (L.) X X X m2 M eg
Z onocyba bifasciata (Boh.) X x o2 M eg Oligophages
Issus coleoptratus (F.) x ? po P ny
A lebra wahlbergi (Boh.) x X x po M eg
Empoasca vitis (Göthe) ? x x po P ad Polyphages
Fagocyba cruenta (H .-S.) ? x x po M eg
A lnetoidia alneti (Dhlb.) ? x x po M eg
Lamprotettix nitidulus (F.) ? A po P eg Vertical
A llygidius atomarius (F.) A o2? P eg m igrants
Cixius nervosus (L.) ? po P eg
Empoasca affinis N ast ? po M? ad
Edwardsiana frustrator (Ed w .) ? po M eg
Edwardsiana lethierryi (Ed w .) ? o2 M eg Uncon-
Fieberiella florii (Stal)
å ? po P eg firm ed
A llygus modestus Scott ? po P eg
A llygidius commutatus (Fieb.) ? ? po P eg
Hesium domino (Reut.) ? po P eg
Total 6 15 9
* = Not distinguished from Ulmus x hollandica

E. plebeja (Edw.), Ribautiana ulmi (L.), and probably Edwardsiana smreczynskii Dwor. and
Kyboasca bipunctata (Osh.), although the latter is also reported from Cannabis and Gly-
cyrrhiza in eastern parts of Europe (Dworakowska 1973; Lauterer 1984). Zonocyba bifas-
ciata (Boh.) prefers Carpinus betulus, but also breeds on Ulmus minor and U. glabra. Issus
coleoptratus (F.), Alebra wahlbergi (Boh.), Empoasca vitis (Göthe), Fagocyba cruenta (H.-S.)
and Alnetoidia alneti (Dhlb.) are polyphagous. Vertical migrants only include Lamprotet-
tix nitidulus (F.) and Allygidius atomarius (F.).

5.1.2.5 Urticaceae
Five native or at least archaeophyte species of the nettle family occur in Germany.
Both stinging and small nettle (Urtica dioica and U. urens) are widespread and very com-
mon, whereas marsh nettle (U. kioviensis) is rare and confined to the Havel basin. Erect
and spreading pellitory-of-the-wall (Parietaria officinalis and P. judaica) are both rather
 Urticaceae 249

localized, the latter being confined to the Rhine valley. Only Urtica dioica, which is per-
haps one of the most abundant herbaceous plants in central Europe, is known to be
utilized in Germany, and it is of major importance as an Auchenorrhyncha host (see
Table 25). Five species have been found to be 1st degree monophagous, notably Macrop-
sis scutellata (Boh.), Eupteryx cyclops Mats., E. calcarata Oss., E. urticae (F.) and Macrosteles
variatus (Fall.). The three latter, however, may feed on more plant species, at least in
other parts of their range. According to Davis & Lawrence (1973), Eupteryx urticae (F.)
has been found in large numbers on Urtica urens and Parietaria judaica in British church-
yards, indicating a broader diet width (at least for the second generation) wherever
further suitable host species occur. On the other hand, U. urens alone as an annual plant
is unlikely to be capable of supporting permanent insect populations. Urtica dioica is
also the main host for wintering eggs and the first generation of the otherwise polyph-
agous Eupteryx aurata (L.). Furthermore, it harbours breeding populations of the broad-
ly polyphagous species Philaenus spumarius (L.), Aphrodes makarovi Zachv. and Evacan-
thus interruptus (L.). Finally, nymphal development is documented for the vertical mi-
grants Hyalesthes obsoletus Sign. and Centrotus cornutus (L.).
The ratio Auchenorrhyncha species number : plant species number in the Urticaceae is
moderately high (Fig. 6), but considering the fact that only Uritica dioica is known to be
utilized, this species is certainly outstanding among all non-woody and non-graminoid
plants in central Europe. The extremely wide geographical as well as altitudinal distri-
bution (e.g. Haeupler & Schönfelder 1989; Benkert et al. 1996) combined with the high
abundance, both rendering U. dioica probably one of the biomass-richest herbaceous
plants, may offer an important clue for the understanding of the heavy infestation, which
is also found in other insect groups (Davis 1973, 1983). Thus the efficiency of plant
defence mechanisms, notably the stinging hairs, is pronounced regarding grazing mam-
mals, but apparently low regarding phytophagous insects. It should also be noted here
that the proportion of specialist and generalist feeders are roughly equal (see Table 25).
Insect communities on nettles have been subject to numerous ecological studies. An
overview of guilds is presented by Davis (1973, 1983). Typhlocybine leafhoppers of the
genus Eupteryx on nettles were used as a model system for several studies of communi-
ty ecology. Different, although broadly overlapping preferences of light and humidity
as well as oviposition sites on the plant were interpreted as evidence for interspecific
competition (Le Quesne 1972; Stewart 1988; Stiling 1980b; see also Denno et al. 1995).
Stiling (1980b) pointed out, that Urtica dioica, with overwintering stems, is the only suit-
able host for the spring generation of Eupteryx aurata (L.), because the additional sum-
mer hosts do not offer above-ground components through the winter. Stewart (1988)
studied field distribution in Wales, laboratory food and host plant choice and nymphal
survival of Eupteryx aurata (L.), E. cyclops Mats. and E. urticae (F.) on Urtica dioica, U. urens
and Parietaria judaica. He found that for feeding and oviposition, U. dioica was favoured
most by Eupteryx aurata (L.) and E. cyclops Mats., but not by E. urticae (F.), which pre-
ferred Parietaria judaica. Nymphal survival was optimal on U. dioica for all three species,
but on U. urens and P. judaica it was reduced in Eupteryx aurata (L.) and E. cyclops Mats.
Egg parasitism of Eupteryx urticae (F.) by a myrmarid wasp (Anagrus spec.) was found
to be similarly high both on U. dioica and P. judaica, although Stiling (1980b) had argued
that leafhoppers may escape heavy parasitation by host alternation. Stewart (1988) sug-
250 Utilization of plant resources

gested that leafhopper habitat preferences (which are in turn affected by interspecific
competition), plant phenology and biochemistry were the major causes for a differen-
tial utilization of the Urticaceae. Finally, it is noteworthy that Zabel & Tscharntke (1998),
who studied Auchenorrhyncha, Heteroptera and Coleoptera on Urtica dioica, could dem-
onstrate that habitat fragmentation affected species numbers of monophages more neg-
atively than the total species number.

5.1.2.6 Fagaceae
The oak family plays the most important role in many forests of temperate latitudes,
certainly being one of the biomass-richest plant taxa all over Europe. In central Europe,
however, only 4 native species occur, and another one, sweet chestnut (Castanea sativa),
is believed to be introduced by the Romans into viticultural regions of southwest Ger-
many. Downy oak (Quercus pubescens) is also confined to warm habitats, mainly in the
southwest. The remaining three species, common oak (Quercus robur), durmast oak (Qu.
petraea), and in particular, beech (Fagus sylvatica) are all climax trees in lower and sub-
montane altitudes on most soils of intermediate humidity, the latter even reaching up to
the alpine tree line. In addition, the North American red oak (Quercus rubra) is locally
planted, and turkey oak (Qu. cerris), originating from southeastern Europe, is an orna-
mental tree in some parks. Most of the following remarks only refer to F. sylvatica, Qu.
robur and Qu. petraea, the Auchenorrhyncha fauna of which is well known in Germany;
but for the remaining species, data are available only for southern Europe and Wales.
Reviews on utilization of Fagaceae in southern and western Europe, with special em-
phasis on typhlocybid leafhoppers, have been published by Vidano & Arzone (1987b)
and Claridge & Wilson (1976, 1981), but see also Lauterer (1983, 1984). It should be
noted that some of the following results must be considered preliminary, since hybrids
within the genus Quercus are difficult to identify.
Fagaceae in Germany are frequently utilized by at least 42 confirmed plus 14 uncon-
firmed Auchenorrhyncha species (Table 9). The ratio Auchenorrhyncha species number
: plant species number is by far the highest among all plants. This is even more striking
since feeding data almost exclusively refer to Quercus robur and Qu. petraea.
Surprisingly, differences between the guilds of Quercus and Fagus are much pro-
nounced, and similarities are small. Species numbers on oak are extremely high, but
individual numbers are often low, whereas the opposite is true for beech, where mass
catches usually comprise Fagocyba cruenta (H.-S.) only.
Fagus sylvatica, which is the most abundant and dominant central European decidu-
ous tree, is infested by only 5 species (plus a few uncertain ones), none of which is
monophagous. This seems rather paradox, in particular regarding the similarities with
Quercus robur and Qu. petraea in plant architecture, geographic range and abundance.
The most important clue may be plant chemistry or vegetation history. The strong dom-
inance of beech in central Europe is a very young phenomenon only found for a few
thousand years, and beech records from earlier interglacial periods are scarce or con-
fined to southern Europe (Lang 1994). Important aspects of this question could be an-
swered by studying the phytophagous fauna in countries, where beech has occurred
through the glacial ages, e.g. in southwestern Europe and the Caucasus.
 Fagaceae 251

Table 9. Auchenorrhyncha species utilizing Fagaceae in Germany. Explanations see Table 5.

Cast anea sat iv a

Fagus sy lv at ica

Overw intering
Q uercus robur

Q u. pubescens

D iet breadth
Q u. pet raea

Substrate

stage
Species
Eurhadina saageri W.Wg. X m 1? M eg
A lebra albostriella (Fall.) X ? ?* ?* m 1? M eg 1st d egree
A lebra viridis R. ? X ?* m 1? M eg m onophages
Eurhadina kirschbaumi W.Wg. X ?* ?* m1 M eg
Iassus lanio (L.) X X m2 P eg
Fagocyba carri (Ed w .) X X m2 M eg
Eurhadina ribauti W.Wg. X X m2 M eg
Eurhadina pulchella (Fall.) ? X X m2 M eg 2nd d egree
Ribautiana alces (Rib.) X ? ?* m 2? M eg m onophages
Ribautiana scalaris (Rib.) x X ?* m 2? M eg
A rboridia velata (Rib.) X X ? m2 M ad
A rboridia spathulata (Rib.) ?* ?* ?* m2 M ad
Typhlocyba quercus (F.) ? x x ?* o2? M eg
Eurhadina concinna (Germ .) ? X X ?* ?* o2? M eg Oligo-
Z ygina angusta Leth. x x x o2 M ad phages
Edwardsiana flavescens (F.) x ? ? ?* o2 M eg
Fagocyba cruenta (H .-S.) X ? ? ?* po M eg
A lnetoidia alneti (Dhlb.) x ? ? po M eg
Empoasca vitis (Göthe) x x x ?* ?* po P ad
Lindbergina aurovittata (Dgl.) ?* ?* ?* ?* ? po M eg
Issus coleoptratus (F.) x x po P ny Polyphages
Issus muscaeformis (Schrk.) ?* ?* po? P ny
Penthimia nigra (Goeze) x x po P ny
Ledra aurita (L.) ? x x po P ny
Edwardsiana frustrator (Ed w .) ? x x po M eg
Cixius distinguendus Kbm . ? A ? po P ny
Cixius dubius W.Wg. A ? po P ny
Cixius stigmaticus (Germ .) A A po P ny
Tachycixius pilosus (Ol.) A A po P ny
Centrotus cornutus (L.) A ? po P ny
Platymetopius major (Kbm .) A ? po P eg
Platymetopius guttatus Fieb. A ? po? P eg
Lamprotettix nitidulus (F.) ? ? po P eg
A llygus communis (Ferr.) A A o2? P eg Vertical
A llygus mixtus (F.) A A po? P eg m igrants
A llygus maculatus Rib. A A o2 P eg
A llygus modestus Scott A A po? P eg
A llygidius commutatus (Fieb.) A A po P eg
A llygidius atomarius (F.) A ? po P eg
Thamnotettix confinis (Zett.) A A po P ny
Thamnotettix dilutior (Kbm .) A A po P ny
Cixidia pilatoi D’Urso & Gugl. ? ? po? F ny
Tettigometra impressopunctata Duf. ? po? P ad Uncon-
Edwardsiana ampliata (W.Wg.) ? o2? M eg firm ed
252 Utilization of plant resources

Table 9. (continued):

Cast anea sat iv a

Fagus sy lv at ica

Overw intering
Q uercus robur

Q u. pubescens

D iet breadth
Q u. pet raea

Substrate

stage
Species
Edwardsiana lamellaris (Rib.) ?* ?* o2? M eg
Ribautiana debilis (Dgl.) ?* po? M eg
Z yginella pulchra P. Löw ? o2 M ad
A rboridia parvula (Boh.) ? o2 M ad
A rboridia ribauti (Oss.) ? o2 M ad
Ribautiana tenerrima (H .-S.) ? ? o2? M eg Uncon-
Z ygina rosincola (Cer.) ? ? o1? M ad firm ed
Platymetopius undatus (De G.) ? ? po? P eg
Fruticidia bisignata (M. & R.) ? ? o2 M ad
Z ygina flammigera (Geoffr.) ? ? ? o1? M ad
A lebra wahlbergi (Boh.) ? ? ?* ?* po M eg
A rboridia erecta (Rib.) ?* o2? M ad
Total substantiated 5 33 25 0 0
Total unconfirm ed 8 22 23 10 11
* = after Claridge & Wilson (1976, 1978a, 1981), Ribaut (1936), Vidano & Arzone (1987b), Wagner (1935)

In contrast, the guilds on oak are extremely diverse, with only minor differences be-
tween Qu. robur and Qu. petraea, although the latter has been sampled less intensively.
With 33 and 25 Auchenorrhyncha species, respectively, plus more than 20 unconfirmed
feeders, the genus Quercus is clearly among the most heavily infested plant taxa in central
Europe, rivalled only by silver birch and the grasses Festuca and Calamagrostis (Fig. 14).
Remarkably, Fagocyba cruenta (H.-S.), Alebra wahlbergi (Boh.) and Alnetoidia alneti
(Dhlb.), which are among the most polyphagous typhlocybids, have been found in very
small numbers only and are probably not breeding on oaks. A similar phenomenon has
also been documented in polyphagous aphids (Eastop 1973). Four leafhopper species
are likely to be 1st degree monophagous: Alebra albostriella (Fall.) and Eurhadina saageri
W.Wg., both on Quercus robur, and A. viridis R. and E. kirschbaumi W.Wg., both on Qu.
petraea. However, this statement is not valid for southern and western Europe: with the
exception of Eurhadina saageri W.Wg., which is generally rare and little known, these
species have all been found to reproduce also on other oak species and even Castanea
sativa (Demichelis & Bosco 1995; Drosopoulos 1999; Gillham 1991; Vidano & Arzone
1987b), their diet breadth narrowing near the northern edge of the range.
Further 8 species are monophagous in the 2nd degree: both Ribautiana alces (Rib.)
and R. scalaris (Rib.) show a preference for Quercus robur or Qu. petraea, respectively.
The remaining species breed to the same extent on both oaks, notably Iassus lanio (L.),
Fagocyba carri (Edw.), Eurhadina pulchella (Fall.), E. ribauti W.Wg., Arboridia velata (Rib.)
and probably A. spathulata (Rib.). Edwardsiana lamellaris (Rib.) is reported to live on
both Quercus robur and Qu. petraea in Italy, but perhaps migrates to Rosa in the first
generation.
 Betulaceae (incl. Corylaceae) 253

There are no 1st degree oligophages, i.e. species exclusively common to both Quercus and
Fagus. Edwardsiana flavescens (F.), Typhlocyba quercus (F.), Zygina angusta Leth. and Eurhadina
concinna (Germ.) are less specific and breed also on plants of other families, although the
latter clearly prefers oaks. Further, there are 9 polyphagous species, and at least 17 vertical
migrants. The latter mainly include species of Cixiidae, Platymetopius, Allygus, Allygidius
and Thamnotettix. This high number of vertical migrants is only surpassed in the birch fam-
ily, although there is a considerable overlap. In general, most mesophyll feeders are monoph-
agous and overwinter as egg, whereas phloem feeders tend to be host generalists and com-
prise about equal numbers of species wintering in the egg stage and as nymphs, respectively.

5.1.2.7 Betulaceae (incl. Corylaceae)

The birch family includes 4 native genera with altogether 9 species, all of which are
woody. Hornbeam (Carpinus betulus), hazel (Coylus avellana), black alder (Alnus gluti-
nosa) and silver birch (Betula pendula) are all widespread and common in low and sub-
montane altitudes, the latter species reaching up to the alpine tree line. Carpinus betulus
is a climax species on locations of intermediate humidity and fertility. Corylus avellana is
essentially a heliophilous underwood shrub of open forests. Betula pendula is a pioneer
tree occupying a wide ecological amplitude on soils with extremely varying supply of
humidity and nutrients. In contrast, Alnus glutinosa is confined to wet and temporarily
flooded locations, often along streams and rivers. Downy birch (B. pubescens) is wide-
spread in the north German plain; in central and southern parts, however, it prefers
higher and cooler sites on wet and acidic soils. Grey and green alder (Alnus incana and
A. alnobetula) grow in higher altitudes, the former mainly along streams and on seepy
mountain slopes (although now often planted in lower altitudes). The latter is a domi-
nant shrub of the subalpine „krummholz“ belt, with scattered occurrence as a pioneer
species in lower altitudes and in the Black Forest. Tree hazel and filbert (Corylus colurna
and C. maxima), both of southwest European and western Asian origin, are commonly
planted in parks and gardens. Finally, dwarf and shrub birch (Betula nana and B. humilis)
occur very rarely in bogs of north Germany and the foothills of the Alps.
The data base of Auchenorrhyncha is quite extensive for Carpinus betulus, Corylus
avellana, Alnus glutinosa and Betula pendula, but only moderate and incomplete for
B. pubescens, Alnus incana and A. alnobetula. There are a few samples from Corylus colur-
na and C. maxima, but none from Betula nana and B. humilis.
Altogether, 61 species of Auchenorrhyncha are confirmed feeders, further 27 are un-
confirmed (Table 10). Thus, the total number is among the highest of all central Europe-
an plant families, surpassed only by the Salicaceae, Cyperaceae and Poaceae. Similarly,
the ratio Auchenorrhyncha species number : plant species number is the second highest
(6.8 vs. 8.4 in the Fagaceae). Betula pendula, Alnus glutinosa and Corylus avellana, with 28,
19 and 16 species, respectively, are among the most-favoured plants. On the other hand,
low species numbers are found on the localized Alnus alnobetula as well as on the intro-
duced Corylus colurna and C. maxima. Low species numbers can also be predicted for
Betula nana and B. humilis. The ratio specialists : generalists is < 1 (except in A. alnobetula
and the introduced hazels, which are insufficiently studied) indicating that plant de-
fence mechanisms may either be less effective or less specific. As in other tree families,
254 Utilization of plant resources

Table 10. Auchenorrhyncha species utilizing Betulaceae in Germany. Explanations see Table 5.

Cory lus av ellana

Carpinus bet ulus

Alnus glut inosa

Overw intering
Bet ula pendula

A. alnobet ula

D iet breadth
B. pubescens
C. maxima
C. colurna

Substrate
A. incana

stage
Species
Oncopsis carpini (J. Shlb.) X . . . . . . . . m1 P eg
Z ygina griseombra Rem . X . . . . . . . . (m 1) M ad
Oncopsis avellanae Ed w . . X . . . . ?* . . m1 P eg
Edwardsiana avellanae (Ed w .) . X . . . . . . . m1 M eg
Edwardsiana spinigera (Ed w .) . X . . . . . . . m1 M eg
Edwardsiana stehliki Laut. . X . . . . . . . m 1? M eg
A lebra coryli Le Q. . X x x . . . . . m2 M eg
Oncopsis appendiculata W.Wg. . . . . X ? . . . m 1? P eg
Kybos calyculus (Cer.) . . . . . ?* . . . m 1? M eg
Z ygina rosea (Fl.) . . . . . x . . . m 1? M ad
Oncopsis tristis (Zett.) . . . . X X . . . m2 P eg
Oncopsis flavicollis (L.) . . . . X X . . . m2 P eg
Oncopsis subangulata (J. Shlb.) . . . . X x*
?* . . . m 2? P eg Monophages
Kybos lindbergi (Lnv.)** . . . . X X . . . m2 M eg
Linnavuoriana decempunctata (Fall.) . . . . X X . . . m2 M ad
Kybos mucronatus (Rib.) . . . . . . ?* . . m 1? M eg
Edwardsiana gratiosa (Boh.) . . . . . . X . . m1 M eg
Eupterycyba jucunda (H .-S.) . . . . . . X . . m1 M eg
Kybos strobli (W.Wg.) . . . . . . . ?* . m 1? M eg
Edwardsiana soror (Lnv.) . . . . . . . x ? m 1? M eg
Linnavuoriana intercedens (Lnv.) . . . . . . . x ?* m 1? M ad
Oncopsis alni (Schrk.) . . . . . . X X . m2 P eg
Kybos smaragdula (Fall.) . . . . . . X X ?* m2 M eg
Edwardsiana alnicola (Ed w .) . . . . . . x X . m2 M eg
Edwardsiana geometrica (Schrk.) . . . . . . X X ? m2 M eg
Z ygina tiliae (Fall.) . . . . . . X x . o2 M ad
Edwardsiana plurispinosa (W.Wg.) . X . . . . X ? x o1 M eg
Edwardsiana bergmani (Tull.) . . . . X X x . x o1 M eg
A lebra neglecta W.Wg. X . . . . . . . . o2 M eg
Edwardsiana flavescens (F.) X . . . ? . . . . o2 M eg Oligo-
Z onocyba bifasciata (Boh.) X . . . . . ? . . o2 M eg phages
A rboridia ribauti (Oss.) X . . . . . . . . o2 M ad
Eurhadina concinna (Germ .) ? . . . ?* ? ?* . . o2? M eg
Typhlocyba quercus (F.) ?* ?* . . . . . . . o2? M eg
Edwardsiana lanternae (W.Wg.) . . . . . . ?* . . o2? M eg
Empoasca vitis (Göthe) x x ? . x x x x ? po P ad
Fagocyba cruenta (H .-S.) X x ? . x ? x ? ? po M eg
A lnetoidia alneti (Dhlb.) X X x . ?* ?* X X ? po M eg
A lebra wahlbergi (Boh.) X ? . . x . x . . po M eg Poly-
Edwardsiana frustrator (Ed w .) ? x . . ? . . . . po M eg phages
Issus coleoptratus (F.) . x . . x . . . . po P ny
Ledra aurita (L.) . ? . . x . ? . . po P ny
A guriahana stellulata (Burm .) . . . . x . . . . po M eg
Cixius nervosus (L.) . A . . A . A A ? po P ny Vertical
Cixius cunicularius (L.) . A . . A . A . . po P ny m igrants
Cixius dubius W.Wg. . A . . . . . . . po P ny
 Betulaceae (incl. Corylaceae) 255

Table 10. (continued):

Cory lus av ellana

Carpinus bet ulus

Alnus glut inosa

Overw intering
Bet ula pendula

A. alnobet ula

D iet breadth
B. pubescens
C. maxima
C. colurna

Substrate
A. incana

stage
Species
Cixius stigmaticus (Germ .) . . . . A . ? . . po P ny
Cixius similis Kbm . . . . . . A . . . o2? P ny
Tachycixius pilosus (Ol.) . . . . A . . . . po P ny
A phrophora alni (Fall.) . A . . A A A A ? po X eg
A phrophora major Uhler . . . . A A . . . po X eg
Platymetopius major (Kbm .) . . . . A . . . . po P eg
Platymetopius guttatus Fieb. . . . . A . . . . po P eg Vertical
Lamprotettix nitidulus (F.) . . . . ? . A . . po P eg m igrants
A llygus communis (Ferr.) . . . . A . . . . o2? P eg
A llygus mixtus (F.) ? . . . A ? A ? . po? P eg
A llygus modestus Scott . . . . A . A . . po? P eg
A llygidius commutatus (Fieb.) . . . . A . . . . po? P eg
Speudotettix subfusculus (Fall.) A A . . A . ? . ? po P ny
Hesium domino (Reut.) . . . . A A ? . . o2? P eg
Thamnotettix confinis (Zett.) ? ? . . A . ? . . po P eg
Cixius cambricus China . . . . . . . . ?* po? P ny
Cixius heydenii Kbm . . . . . . . . . ?* po? P ny
Cixius simplex (H .-S.) . . . . ? ? ? ? . po P ny
Pentastiridius beieri (W.Wg.) . . . . . . . ?* . o2? P ny
Issus muscaeformis (Schrk.) . ? . . . . . . . po? P ny
Lepyronia coleoptrata (L.) . ?* . . . ?* . . . po X eg
Philaenus spumarius (L.) . ?* . . ?* ?* ?* . . po X eg
Centrotus cornutus (L.) . ? . . ? . ? . . po P ny
A lebra albostriella (Fall.) . . . . ? . ? . . m 1? M eg
A lebra viridis R. . . . . . . . . ? m 1? M eg
Empoasca affinis N ast ? . . . . . . . . po M? ad
Empoasca decipiens Paoli . ? . . ? . . . . po M? ad
Edwardsiana ampliata (W.Wg.) . ? . . . . . . . o2? M eg
Edwardsiana ishidai (Mats.) . ? . . . . . . . m 2? M eg Unconfirmed
Lindbergina aurovittata (Dgl.) . ?* . . . ?* ?* . . po M eg Unconfirm ed
Ribautiana debilis (Dgl.) . ?* . . ?* . ?* . . po? M eg
Ribautiana tenerrima (H .-S.) . ?* . . ? . ? . . o2? M eg
Eurhadina ribauti W.Wg. ? . . . . . ? . . m2 M eg
Eurhadina pulchella (Fall.) ? . . . ? . . . . m2 M eg
A guriahana pictilis (Stal)
å . . . . . ? . . . m 1? M eg?
Z yginella pulchra P. Löw . . . . . . ? . ? m 2? M ad
Z ygina angusta Leth. ? . . . . . . . . o2 M ad
Z ygina rosincola (Cer.) . . . . ? . . . . o1? M ad
A rboridia erecta (Rib.) . ?* . . . . . . . o2? M ad
Platymetopius undatus (De G.) . . . . ? . . . . o2? P eg
Idiodonus cruentatus (Panz.) . ? . . ? . . . . po P eg
Colladonus torneellus (Zett.) . . . . ? . ? . . po P ny
Total substantiated 11 16 2 1 28 12 19 10 2
Total unconfirm ed 9 16 2 0 17 11 20 7 14
* = after Claridge & Wilson (1978b, 1981), Claridge et al. (1977), Dworakowska (1976), Freese (pers. comm.),
Grissemann (1983), Günthart (1987a), Lauterer (1980), Le Quesne & Payne (1981), Ossiannilsson (1950),
Ribaut (1936), Vidano & Arzone (1987a); ** = incl. K. betulicola (W.Wg.)
256 Utilization of plant resources

most Auchenorrhyncha species on Betulaceae winter in the egg stage, but the preva-
lence of mesophyll feeders is less pronounced.
Carpinus betulus is infested by two host specialists, notably Oncopsis carpini (J. Shlb.)
and Zygina griseombra Rem., although the latter migrates to coniferous trees for hiberna-
tion and to shrub species of Rosaceae in spring. Alebra neglecta W.Wg., Edwardsiana flave-
scens (F.), Zonocyba bifasciata (Boh.) and Arboridia ribauti (Oss.) prefer C. betulus at least
locally, but also breed on members of other plant families. Polyphages include Empoasca
vitis (Göthe), Fagocyba cruenta (H.-S.), Alnetoidia alneti (Dhlb.), Alebra wahlbergi (Boh.) and
adults of Speudotettix subfusculus (Fall.). F. cruenta (H.-S.) may be extremely abundant.
Oncopsis avellanae Edw., Edwardsiana avellanae (Edw.), E. spinigera (Edw.), E. stehliki
Laut. and Alebra coryli Le Q. are all specific to Corylus avellana, although the latter infests
also ornamental C. colurna and C. maxima. In Wales Oncopsis avellanae Edw. was also
found to breed on Alnus glutinosa. The only oligophagous species is Edwardsiana pluris-
pinosa (W.Wg.), which reproduces both on Corylus avellana and Alnus spp. Among poly-
phages, Alnetoidia alneti (Dhlb.) is clearly the most abundant species, accompanied by
smaller numbers of Empoasca vitis (Göthe), Fagocyba cruenta (H.-S.), Edwardsiana frustra-
tor (Edw.), Issus coleoptratus (F.) and the vertically migranting Cixius nervosus (L.),
C. cunicularius (L.), C. dubius W.Wg. and Speudotettix subfusculus (Fall.).
Birches are among the most important food plants for Auchenorrhyncha. Betula pen-
dula alone holds 28 feeding species, which is one of the highest numbers among all
central European plants, only exceeded by Quercus robur (see Fig. 14). The relatively
low species number, mainly of polyphages, on B. pubescens might be the result of less
extensive sampling. Oncopsis appendiculata W.Wg. is perhaps specific to B. pendula (al-
though there is a single record on B. pubescens). Kybos calyculus (Cer.) and Zygina rosea (Fl.)
are both probably specific to B. pubescens (but the latter wintering on coniferous trees).
Oncopsis tristis (Zett.), O. flavicollis (L.), Kybos lindbergi (Lnv.), Linnavuoriana decem-
punctata (Fall.) and probably O. subangulata (J. Shlb.) breed exclusively on both birches.
Edwardsiana bergmani (Tull.) mainly breeds on both birches, but locally also on alders.
Polyphages (mainly on B. pendula) include Issus coleoptratus (F.), Ledra aurita (L.), Alebra
wahlbergi (Boh.), Empoasca vitis (Göthe), Fagocyba cruenta (H.-S.) and Aguriahana stellula-
ta (Burm.). The number of vertical migrants on birches (again mainly on B. pendula) is
also high and comprises 17 species belonging to the genera Cixius, Aphrophora, Platyme-
topius, Allygus and others. Adults of Platymetopius major (Kbm.), P. guttatus Fieb., Ally-
gus communis (Ferr.) and Hesium domino (Reut.) all show clear preferences for B. pendula,
but usually spend the nymphal stage in the herb layer.
Alders are another preferred host group. Strictly monophages are confined to Alnus
glutinosa and A. incana. The former is exclusively utilized by Edwardsiana gratiosa (Boh.),
Eupterycyba jucunda (H.-S.) and probably Kybos mucronatus (Rib.), the latter probably by
Kybos strobli (W.Wg.), Linnavuoriana intercedens (Lnv.) and Edwardsiana soror (Lnv.). 2nd
degree monophages include Oncopsis alni (Schrk.), Edwardsiana alnicola (Edw.), Kybos
smaragdula (Fall.) and E. geometrica (Schrk.). The two latter perhaps breed also on
A. alnobetula. Zygina tiliae (Fall.) favours Alnus glutinosa, but has also been found to re-
produce on A. incana and Tilia cordata. Edwardsiana plurispinosa (W.Wg.) and E. bergmani
(Tull.) are oligophagous in the 1st degree on Betulaceae. Alebra wahlbergi (Boh.), Empoas-
ca vitis (Göthe), Fagocyba cruenta (H.-S.) and Alnetoidia alneti (Dhlb.) are polyphagous.
 Tiliaceae 257

Vertical migrants include Cixius nervosus (L.), Aphrophora alni (Fall.), Lamprotettix nitidu-
lus (F.), Allygus mixtus (F.) and A. modestus Scott. Altogether, black alder is attacked by
19 species, which is one of the highest numbers all over Germany (see Fig. 14).

5.1.2.8 Tiliaceae
The linden family comprises only two native species of a single genus. Small- and
large-leaved lime (Tilia cordata and T. platyphyllos) are both widespread and common in
most lowland and hill regions up to c. 1000 m a.s.l., although in the north German plain
most stands have been planted. The former species is rather eurytopic, preferring oak-
hornbeam and floodplain forests. The latter is more confined to shady slopes on seepy
and nutrient soils. The Auchenorrhyncha fauna on small-leaved lime is well studied, in
contrast to large-leaved lime. Many published records refer only generally to Tilia spp.
Apart from that, there are several ornamental species and hybrids (e.g. T. petiolaris)
planted in parks and alleys, with very little information on their phytophagous fauna.
Altogether, at least 9 Auchenorrhyncha species breed on Tilia in Germany, plus one verti-
cal migrant and 10 unconfirmed species (Table 11). The degree of specificity is low, without

Table 11. Auchenorrhyncha species utilizing Tiliaceae in Germany. Explanations see Table 5.

Overw intering stage

T. plat y phy llos
Tilia cordat a

D iet breadth

Substrate

Species
Pediopsis tiliae (Germ .) X X m2 P eg Monophages
Edwardsiana lethierryi (Ed w .) X x o2 M eg Oligophages
Z ygina tiliae (Fall.) x o2 M ad
Issus coleoptratus (F.) x po P ny
A lebra wahlbergi (Boh.) X x po M eg
Empoasca vitis (Göthe) X X po P ad Polyphages
Fagocyba cruenta (H .-S.) X x po M eg
A lnetoidia alneti (Dhlb.) X X po M eg
A guriahana stellulata (Burm .) X po M eg
Cixius nervosus (L.) A po P ny Vertical m igrants
Ledra aurita (L.) ? po P ny
Edwardsiana flavescens (F.) ? ? o2 M eg
Edwardsiana frustrator (Ed w .) ? ? po M eg
Eurhadina concinna (Germ .) ? o2 M eg
Edwardsiana ampliata (W.Wg.) ?* o2? M eg Uncon-
Typhlocyba quercus (F.) ?* o2? M eg firm ed
Z onocyba bifasciata (Boh.) ? o2 M eg
A rboridia ribauti (Oss.) ? ? o2 M ad
Lamprotettix nitidulus (F.) ? ? po P eg
A llygus mixtus (F.) ?* po P eg
Total 10 6
* = after Chudzicka (1986a, 1986b)
258 Utilization of plant resources

any 1st degree monophages, and Pediopsis tiliae (Germ.) as the only 2nd degree monophage.
As in elms and maples, 1st degree oligophages are identical with 2nd degree monophages
due to the existence of only a single host genus in the family. Among 2nd degree oligophag-
es, Edwardsiana lethierryi (Edw.) breeds mainly on Tilia, but also on Acer campestre. In a few
cases, Zygina tiliae (Fall.) has been found to reproduce on small-leaved lime, but Alnus glu-
tinosa is clearly the main host. The remaining species are broadly polyphagous and include
generalists like Issus coleoptratus (F.), Alebra wahlbergi (Boh.), Empoasca vitis (Göthe), Fagocyba
cruenta (H.-S.), Alnetoidia alneti (Dhlb.). Aguriahana stellulata (Burm.) is noteworthy in prefer-
ring single host genera belonging to at least 5 different plant families.

5.1.2.9 Salicaceae
The willow family is represented by only two genera comprising altogether 28 native
species. Almost half of them are confined to subalpine and alpine habitats, with only
few positive data available on their insect fauna. Most of these have never or only unin-
tentionally been sampled for their Auchenorrhyncha. Although most low-growing wil-
lows (e.g. Salix retusa, S. herbacea, S. reticulata) are supposed to be devoid of Auchenor-
rhyncha, shrubby species, like S. bicolor, S. hastata and S. daphnoides may well harbour
diverse guilds. Recent extensive collecting on S. eleagnos in Bavaria even revealed a
new leafhoppers species (Macropsis remanei Nick.). Most of the remaining species of
Salicaceae are widespread in lowland areas, mainly along rivers, streams and lakeshores.
Of particular importance are all three species of poplar, namely aspen, black and white
poplar (Populus tremula, P. nigra, P. alba) and the taller willow species, some of which are
commonly called sallows and osier, notably crack willow (Salix fragilis), white willow
(S. alba), almond willow (S. triandra), osier (S. viminalis), purple willow (S. purpurea),
eared willow (S. aurita), grey willow (S. cinerea) and goat willow (S. caprea). These spe-
cies are all common and often dominant in floodplain habitats, except P. tremula and
S. caprea, which both grow in early stages of woodland succession and along forest
margins. Creeping willow (Salix repens) is usually found in bogs and heaths. The data
base of all these species is sufficient, but not so in a number of more localized species
growing in the Alps, in riverine habitats of their foreland or in the eastern half of Ger-
many, e.g. large-leaved willow (S. appendiculata), violet willow (S. daphnoides), olive
willow (S. eleagnos), dark-leaved willow (S. myrsinifolia) and bay willow (S. pentandra).
There is also a large number of intrageneric hybrids within Salix and Populus, as well as
some introduced and ornamental species, which have not been sampled so far.
The guilds on poplars show only little overlap with those of other trees, even within
the family Salicaceae, with 18 monophagous species in the 1st degree, but only one in the
2nd degree. Macropsis fuscinervis (Boh.), Tremulicerus tremulae (Estl.), Populicerus laminatus
(Fl.) and P. populi (L.) live exclusively on P. tremula. Strictly monophagous feeders on P. alba
include Macropsis vicina (Horv.), Tremulicerus distinguendus (Kbm.), Viridicerus ustulatus
(M. & R.), Populicerus albicans (Kbm.), Edwardsiana candidula (Kbm.), Zygina nivea (M. &
R.) and probably Zygina tithide Ferr. In contrast, P. nigra shares its Auchenorrhyncha spe-
cialists with its hybrids and Lombard poplar (P. nigra italica). These include Stenidiocerus
poecilus (H.-S.), Kybos abstrusus (Lnv.), Tremulicerus fulgidus (F.), T. vitreus (F.), Macropsis
graminea (F.), Rhytidodus decimusquartus (Schrk.) and Populicerus nitidissimus (H.-S.). The
 Salicaceae 259

proportion of poplar-feeding species hibernating in the adult stage is relatively high, but
the winter food plants are largely unknown. T. fulgidus (F.) has repeatedly been taken
from Picea abies. Kybos populi (Edw.) is the only 2nd degree monophage, being frequently
found on all three poplars, though not in large numbers. There are no oligophages at all.
Polyphages include Empoasca vitis (Göthe) and Aguriahana stellulata (Burm.), the latter
being frequently found on P. nigra. Aphrophora alni (Fall.), Stictocephala bisonia Kopp &
Yonke and Allygus mixtus (F.) are vertical migrants, the latter also frequently on P. nigra.
The host range of Auchenorrhyncha on willows is somewhat different, including al-
together 12 monophages in the 1st degree and 24 in the 2nd degree. The following sec-
tion treats (i) 1st degree monophages and their hosts, (ii) monophages specific to nar-
row-leaved willows, (iii) monophages specific to broad- and hairy-leaved sallows, (iv)
Salix monophages common to most Salix species, and (v) oligo- and polyphages and
vertical migrants, which also utilize further plant families.
(i) 1st degree monophages and their hosts
Surprisingly, the occurrence of strict monophages among Salix appears to be rather
unequal. Macropsis marginata (H.-S.), M. haupti W.Wg., Kybos rufescens Mel., Idiocerus
similis Kbm., I. vicinus Mel. and Mimallygus lacteinervis (Kbm.) all live on S. purpurea
(the two latter perhaps also on S. eleagnos). Salix alba is quite remarkable in harbouring
three species, all of which belong to a single genus (Macropsis albae W.Wg., M. gravesteini
W.Wg. and M. najas Nast). The recently described Macropsis remanei Nick. is associated
with S. eleagnos, Macropsis viridinervis W.Wg. with S. triandra. However, the latter spe-
cies perhaps breeds also on S. pentandra. Finally, Kybos digitatus (Rib.) is probably strict-
ly monophagous either on S. purpurea or S. eleagnos (R. Mühlethaler, pers. comm.). Apart
from these strictly monophagous feeders, these willows are also infested by a number
of less specific leafhoppers belonging to various taxa (mainly Aphrophorinae, Macrop-
sis, Idiocerinae, Kybos, Edwardsiana). Thus, Salix purpurea, S. alba and S. triandra are among
those plant species with the highest Auchenorrhyncha diversity at all, ranging between
16 and 19 species (see also Fig. 14).
(ii) Monophages specific to narrow-leaved willows
Zygina ordinaria (Rib.) and Sagatus punctifrons (Fall.) are confined to narrow-leaved
‘willows’, notably S. pentandra, S. triandra, S. viminalis, S. eleagnos, S. purpurea, S. alba and
S. fragilis. Idiocerus herrichii (Kbm.) and Zygina lunaris (M. & R.) are only found on the two
latter species. Macropsis notata (Proh.) mainly breeds on S. triandra, but locally also on
S. fragilis. S. viminalis and S. repens are quite peculiar in sharing many species common to
willows and sallows, but also show a small group of leafhoppers being more or less spe-
cific. Thus, Metidiocerus impressifrons (Kbm.), Kybos limpidus (W.Wg.) and Edwardsiana ter-
sa (Edw.) all mainly attack S. viminalis, but these species have occasionally been found
also on S. triandra or others. S. repens is the main host of Macropsis impura (Boh.), which
has rarely been reported also from low-growing individuals of S. aurita. Kybos butleri (Edw.)
favours S. repens in coastal habitats, but S. triandra inland; locally it is also found on sal-
lows. Finally, it should be mentioned that Myndus musivus (Germ.) and Pentastiridius beieri
(W.Wg.), both of which are vertical migrants, were so far only sampled on two sallow
species in Germany, but on different species or even on members of different plant fami-
lies in the Italian and French Alps. Pure weeping willow (S. babylonica) is rarely found, but
hybrids with S. alba are commonly planted in urban areas. In a few cases, Kybos virgator
260 Utilization of plant resources

(Rib.), Macropsis albae W.Wg., Idiocerus stigmaticalis Lew., Zygina lunaris (M. & R.) and
Empoasca vitis (Göthe), have been found on these, but only the first species is a confirmed
breeder. Salix alba, S. triandra, S. purpurea and S. viminalis show the most diverse Auchen-
orrhyncha guilds, comprising between 16 and 19 species (see also Fig. 14).
(iii) Monophages specific to broad- and hairy-leaved sallows
A well distinct group of leafhoppers is restricted to broad-leaved willow species,
most of them commonly known as ‘sallows’ (S. aurita, S. cinerea, S. caprea). These in-
clude Idiocerus lituratus (Fall.), Kybos strigilifer (Oss.), Macropsis prasina (Boh.), Metidio-
cerus elegans (Fl.), Populicerus confusus (Fl.), Edwardsiana salicicola (Edw.) and Linnavuori-
ana sexmaculata (Hardy). Most of these species also breed on the narrow-leaved
S. viminalis, and at least some of them on the more localized and seldom-studied S.
appendiculata and S. myrsinifolia. The host preferences of M. infuscata (J. Shlb.) are some-
what different. It mainly lives on S. caprea, but, along the edge of the Alps, also on S.
myrsinifolia. Auchenorrhyncha species numbers are usually high, reaching 16 species
on S. aurita and 17 on S. cinerea and S. caprea, respectively.
(iv) Salix monophages common to both willows and sallows
Another group of 2nd degree monophages exploits a rather wide range of both wil-
low and sallow species. These include Aphrophora pectoralis Mats., A. salicina (Goeze),
Macropsis cerea (Germ.), Idiocerus stigmaticalis Lew., Metidiocerus rutilans (Kbm.) and Ky-
bos virgator (Rib.).
(v) Oligo- and polyphages, and vertical migrants
In contrast, numbers and proportions of oligophages and polyphages utilizing Sali-
caceae are very small. There is a striking lack of 1st degree oligophages (i.e. species
feeding both on Populus and Salix). 2nd degree oligophages are represented only by
Edwardsiana prunicola (Edw.) and perhaps Pentastiridius beieri (W.Wg.). Polyphages in-
clude Philaenus spumarius (L.), Empoasca vitis (Göthe) and Aguriahana stellulata (Burm.),
as well as some vertical migrants, notably Cixius nervosus (L.), Aphrophora alni (Fall.),
A. major Uhl. and Stictocephala bisonia Kopp & Yonke.
Altogether, the total species number of Auchenorrhyncha on Salicaceae is the highest
among all dicotyledonous plant families, rivalled only by the Betulaceae. Altogether, 66
confirmed plus 14 unconfirmed species have been found. The absolute numbers of host
specialists as well as the ratio host specialists : generalists, are both exceptionally high
in most of the tall-growing lowland species, notably Salix caprea, S. triandra, S. viminalis,
S. alba, S. cinerea, S. purpurea and S. aurita (Table 12). Numbers of 1st degree monophag-
es on Populus alba, P. nigra and Salix purpurea are among the highest of all central Euro-
pean plants (with 7, 7 and 6 respectively), being surpassed only by the grass Phragmites
australis. The lack of overlap between the guilds of Populus and Salix, but also between
those of Salicaceae and other plant families is striking. Despite the large total number of
associated species there is not a single 1st degree oligophage. Within the genus Salix,
there is also a remarkable contrast between the guilds of widespread and tall-growing
species of the lowlands and low-growing, rather localized species of the Alps. This overall
pattern may be explained by combined effects of plant availability (size, range, abun-
dance) and plant physiology (chemistry). Thus the high content of phenolic hetero-
sides, which are toxic to many insects and which are largely restricted to the Salicaceae,
is likely to play an important role.
 Salicaceae 261

Table 12. Auchenorrhyncha species utilizing Salicaceae in Germany. Explanations see Table 5.

Populus hybrids

S. appendiculat a
Populus t remula

Overw intering
S. my rsinifolia

D iet breadth
Salix fragilis

S. pent andra

S. v iminalis

S. purpurea
S. eleagnos
S. t riandra

Substrate
S. cinerea
S. caprea
S. repens
S. aurit a
P. nigra
P. alba

S. alba

stage
Species
M acropsis fuscinervis (Boh.) X . . . . . . . . . . . . . . . . m1 P eg
Tremulicerus tremulae (Estl.) X . . . . . . . . . . . . . . . . m1 P eg
Populicerus laminatus (Fl.) X . . . . . . . . . . . . . . . . m1 P eg
Populicerus populi (L.) X . . . . . . . . . . . . . . . . m1 P eg
M acropsis vicina (H orv.) . X . . . . . . . . . . . . . . . m1 P eg
Tremulicerus distinguendus (Kbm .) . X . . . . . . . . . . . . . . . m1 P eg
V iridicerus ustulatus (M. & R.) . X . . . . . . . . . . . . . . . m1 P ad
Populicerus albicans (Kbm .) . X . . . . . . . . . . . . . . . m1 P eg
Edwardsiana candidula (Kbm .) . X . . . . . . . . . . . . . . . m1 M eg
Z ygina nivea (M. & R.) . X . . ?* . . . . ?* . . . . . . . m 1? M ad
Z ygina tithide Ferr. . ? . . . . . . . . . . . . . . . m 1? M ad
Stenidiocerus poecilus (H .-S.) . . X . . . . . . . . . . . . . . m1 P ad
Kybos abstrusus (Lnv.) . . X . . . . . . . . . . . . . . m1 M eg
Tremulicerus fulgidus (F.) . . X ? . . . . . . . . . . . . . m1 P ad 1 st d e-
Tremulicerus vitreus (F.) . . X x . . . . . . . . . . . . . (m 1) P eg gree m o-
M acropsis graminea (F.) . . X x . . . . . . . . . . . . . (m 1) P eg nophages
Rhytidodus decimusquartus (Schrk.) . . X x . . . . . . . . . . . . . (m 1) P ad
Populicerus nitidissimus (H .-S.) . . X x . . . . . . . . . . . . . (m 1) P eg
M acropsis albae W.Wg. . . . . . X . . . . . . . . . . . m1 P eg
M acropsis gravesteini W.Wg. . . . . . X . . . . . . . . . . . m1 P eg
M acropsis najas N ast . . . . . X . . . . . . . . . . . m1 P eg
M acropsis viridinervis W.Wg. . . . . . . ? X . . . . . . . . . m 1? P eg
M acropsis remanei N ick. . . . . . . . . . X . . . . . . . m1 P eg
M acropsis marginata (H .-S.) . . . . . . . . . . X . . . . . . m1 P eg
M acropsis haupti W.Wg. . . . . . . . . . . X . . . . . . m1 P eg
Idiocerus similis Kbm . . . . . . . . . . . X . . . . . . m1 P eg
Kybos rufescens Mel. . . . . . . . . . . X . . . . . . m1 M eg
Idiocerus vicinus Mel. . . . . . . . . . ? X . . . . . . m 1? P ad
M imallygus lacteinervis (Kbm .) . . . . . . . . . ? X . . . . . . m 1? P eg
Kybos digitatus (Rib.) . . . . . . . . . ?* ?* . . . . . . m 1? M eg
Kybos populi (Ed w .) X X X x . . . . . . . . . . . . . m2 M eg
Edwardsiana tersa (Ed w .) . . . . . . ? ? X . . . . . . . . m 2? M eg
M etidiocerus impressifrons (Kbm .) . . . . . . . x X . x . . . . . . m2 P ad
Kybos limpidus (W.Wg.) . . . . . ? ? x X . . . . . . . . m2 M eg
Idiocerus herrichii (Kbm .) . . . . x X . . . . . . . . . . . m2 P ad
Z ygina lunaris (M. & R.) . . . . X X . . . . x . . . . . . m 2? M ad
Kybos virgator (Rib.) . . . . X X ? x x . . . x x . . . m2 M eg 2nd d e-
Idiocerus stigmaticalis Lew . . . . . x X x x ? x ? . ? ? x . . m2 P eg gree m o-
M etidiocerus rutilans (Kbm .) . . . . ? X . X X ? X . x x x . . m2 P ad nophages
M acropsis cerea (Germ .) . . . . X x . x X . X . X X X . x m2 P eg
A phrophora pectoralis Mats. . . . . . x . . x . X . X X X ? x m2 X eg
A phrophora salicina (Goeze) . . . . x X . x x . X x x x x . . m 2? X eg
Z ygina ordinaria (Rib.) . . . . x X x X X ? X x ? . . . . m2 M ad
Sagatus punctifrons (Fall.) . . . . ? ? ? X x X X X . . . . . m2 P eg
Kybos butleri (Ed w .) . . . . . . ? X . . . X . x . . . m2 M eg
 262 Utilization of plant resources

Table 12. (continued):

Populus hybrids

S. appendiculat a
Populus t remula

Overw intering
S. my rsinifolia
D iet breadth
Salix fragilis

S. pent andra

S. v iminalis

S. purpurea
S. eleagnos
S. t riandra

Substrate
S. cinerea
S. caprea
S. repens
S. aurit a
P. nigra
P. alba

S. alba

stage
Species
M acropsis impura (Boh.) . . . . . . . . . . . x* x . . . . m2 P eg
Idiocerus lituratus (Fall.) . . . . . . . . ? . . X X . X . . m2 P eg
Kybos strigilifer (Oss.) . . . . . . . . . . . . x X X . x m2 M eg
M acropsis prasina (Boh.) . . . . . . . . . . . . X X X . . m2 P eg 2nd
M etidiocerus elegans (Fl.) . . . . . . . . ? . ? . X X X . . m2 P eg? d egree
Populicerus confusus (Fl.) . . . . ? ? . ? X . . . X X X x x m2 P eg m ono-
Edwardsiana salicicola (Ed w .) . . . . . . . . ? . . . X X X x . m2 M eg phages
Linnavuoriana sexmaculata (H ard y) . . . . . ? . . X x . . X X X . . m2 M ad
M acropsis notata (Proh.) . . . . x . . X . . . . . . . . . m2 P eg
M acropsis infuscata (J. Shlb.) . . . . . . . . . . . . . . X . x m2 P eg
Edwardsiana prunicola (Ed w .) . . . . . . . . x . . . x x x . . o2 M eg Oligo-/
Empoasca vitis (Göthe) x x x ? . x ? x x x x . ? x x . . po P ad Poly-
A guriahana stellulata (Burm .) . . x . . . . . . . . . . . . . . po M eg phages
Cixius nervosus (L.) . . . . A A . A A . . . . A A . ? po P ny
M yndus musivus (Germ .) . . . . . . . A A . ? . . . . . . m 2? P ny
Pentastiridius beieri (W.Wg.) . . . . . . . . . . A . . . . . . o2? P ny
A phrophora alni (Fall.) A A A ? A A . A A A A . A A A . ? po X eg Vertical
A phrophora major Uhler . . . . . A . . . . A . . . . . . po X eg migrants
Centrotus cornutus (L.) ? . . . . . . . . . . . . . . . . po P ny
Stictocephala bisonita Kopp & Yonke . A A . . . . . . ? A . ? . . . . po P eg
A llygus mixtus (F.) . . A . ? . . . . . . . . ? . . . po P eg
Hyalesthes obsoletus Sign. . . . . . . . . . . . . . . . . . po P ny
Issus coleoptratus (F.) . . . . . . . . . . . . . . . . . po P ny
Lepyronia coleoptrata (L.) ?* . . . . . ?* . . . . ?* . ?* . . . po X eg
Philaenus spumarius (L.) ?* . . . . . . . ?* . . ?* . ?* ?* . . po X eg
Ledra aurita (L.) ? . . . . . . . . . . . . . . . . po P ny
Penthimia nigra (Goeze) ? . . . . . . . . . . . . . . . . po P ny
Speudotettix subfusculus (Fall.) ? . . . . . . . . . . . . . . . . po P ny Uncon-
Fagocyba cruenta (H .-S.) ? . ? . . . . . . . . . . . . . . po M eg firm ed
Empoasca affinis N ast . . ? . . . . . . . . . . ? . . . po M? ad
Edwardsiana flavescens (F.) . . . . . . . . . . . . . ? . . . o2 M eg
Ribautiana tenerrima (H .-S.) . . . . . . . . . . . . . . ? . . o2? M eg
Colladonus torneellus (Zett.) . . . . . . . . . ? . . . . ? . . po? P ny
A lnetoidia alneti (Dhlb.) . . . . . ? . . . . . . . . . . . po M eg
Fieberiella florii (Stål) . . . . . . . . . . ? . . . . . . po P eg
Total substantiated 7 10 12 5 10 16 2 16 18 7 19 6 16 17 17 2 5
Total unconfirm ed 7 1 3 3 5 5 8 2 5 8 5 2 4 6 4 1 2
* = after Ossiannilsson (1950), Ribaut (1936), Wagner (1950)

5.1.2.10 Ericaceae
The heath family s. str. (excl. the Pyrolaceae and Monotropaceae) comprises 17 native
species. Most of these are dwarf shrubs, with a small number of nanophanerophytes, usual-
ly confined to bogs, open conifer forests or subalpine scrub. The formerly vast Calluna-
 Ericaceae 263

dominated heathlands of the northwest German plain developed after extensive degrada-
tion by cutting, grazing and litter removal, but have largely been reforested during the past
century. Many species are cultivated in gardens and cemeteries due to their evergreen hab-
its. The genus Rhododendron, in particular, includes numerous ornamental species and vari-
eties originating from Mediterranean mountains, North America and Asia, many of which
have changed their original character by cross-breeding and grafting. The data base of phy-
tophagous insects is sufficient to moderate for Calluna, cultivated Rhododendron and most
species of Vaccinium and Erica, but there is no information for the remaining groups, namely
Arctostaphylos, Ledum, native Rhododendron, Kalmia, Rhodothamnus, Loiseleuria and Androme-
da, all of which are rather uncommon and localized. The Monotropaceae and Pyrolaceae,
both of which are now considered as subgroups of the Ericaceae, altogether comprise 9
species, which are apparently not utilized by Auchenorrhyncha.
Altogether, 12 Auchenorrhyncha species are known as feeders on Ericaceae in Germa-
ny (Table 13), plus 8 which need further confirmation. Heather (Calluna vulgaris), being

Table 13. Auchenorrhyncha species utilizing Ericaceae in Germany. Explanations see Table 5.
Rhododendron cultivated

Overw intering stage

Vaccinium my rt illus

Calluna v ulgaris
V. uliginosum

V. v it is-idaea

Erica t et ralix

D iet breadth
V. oxy coccos

Substrate
E. carnea

Species
Graphocephala fennahi Young X (m 2)? X eg
Ulopa carneae W.Wg. X m1 P ny/ ad Monophages
Ulopa reticulata (F.) X ? m1 P ny/ ad
Z ygina rubrovittata (Leth.) X m1 M ad
Ophiola cornicula (Marsh.) ? ? X o1? P eg
Ophiola russeola (Fall.) ? ? x* X x o1 P eg Oligophages
Erythria aureola (Fall.) X ? o2 M eg
Philaenus spumarius (L.) x ?* po X eg Polyphages
Erythria manderstjernii (Kbm .) x ? po M ad
Cixius similis Kbm . A ? ? o2 P ny Vertical
A phrophora corticea Germ . x ? X (po) X eg m igrants
Idiodonus cruentatus (Panz.) A ? ? po P eg
Fieberiella septentrionalis W.Wg. ? po P eg
Lepyronia coleoptrata (L.) ?* ?* po X eg
A guriahana pictilis (Stål) ?* m 1? M eg?
Colladonus torneellus (Zett.) ?* ?* ?* ?* po? P ny
Ommatidiotus dissimilis (Fall.) ? ? ? m 1? P eg Unconfirm ed
Planaphrodes trifasciata (Geoffr.) ? o2? P eg
Empoasca vitis (Göthe) ? po P ad
A rboridia ribauti (Oss.) W? o2 M ad
Ophiola decumana (Kontk.) ?* ?* o1? P eg
Total 1 5 0 0 1 6 0 2
* = after Günthart (1987a), Holgersen (1945), Ossiannilsson (1981), Strübing (1955)
264 Utilization of plant resources

widespread and particularly common on sandy soils and in bogs, is the most important
host species, with Ulopa reticulata (F.) and Zygina rubrovittata (Leth.) as strictly monopha-
gous species, and Ophiola russeola (Fall.), O. cornicula (Marsh.) and Erythria aureola (Fall.)
as oligophages. Locally, heather is also an important food plant for nymphs of Aphrophora
corticea Germ., the adults of which live in the tree layer of pine forests. Furthermore, there
is a number of dubious records, e.g. of Planaphrodes trifasciata (Geoffr.), which perhaps
favours heather in the north German plain and in bogs of the middle and southern parts.
Spring heath (Erica carnea) is a common plant in the understory of open forests of
pine and spruce, in the Alps and their foothills as well as in subalpine scrub, with spo-
radic stands in northeastern Bavaria and Saxony. The only feeding records refer to the
strictly monophagous Ulopa carneae W.Wg., which is probably an endemic of the Alps,
and the oligophagous Ophiola russeola (Fall.). No positive records were made on cross-
leaved heath (E. tetralix), which is widespread in boggy heathland of north Germany,
but it has been reported as food plant of Ulopa reticulata (F.) in older papers.
The Auchenorrhyncha guild of Vaccinium includes mostly generalists. Bilberry (V.
myrtillus) is widespread and common on most acidic substrates, often dominating in
the undergrowth of conifer forests, in bogs and mountain heathland. It is utilized by
Philaenus spumarius (L.), Erythria manderstjernii (Kbm.), Cixius similis Kbm., Aphrophora
corticea Germ., Idiodonus cruentatus (Panz.) and perhaps Aguriahana pictilis (Stål). Only
the latter is reported as monophagous. The remaining species of Vaccinium are not prop-
erly studied. The only feeding record is of Ophiola russeola (Fall.) on cranberry (V. oxy-
coccos). Finally, the Nearctic neozoon Graphocephala fennahi Young breeds on ornamental
Rhododendron, the taxonomy of which is rather puzzling due to multiple cross-breeding
of various species, although North American gene material is apparently widespread
in western Europe (H. Bruelheide, pers. comm.). Adult leafhoppers, however, are also
found on other deciduous woods (J. Badmin, pers. comm.).

5.1.2.11 Rosaceae
In Germany the rose family is represented by more than 160 species (excluding 140
subspecies of Rubus fruticosus and Alchemilla spp.), 50 of which are woody. These in-
clude a number of fruit or ornamental species, some of which have been cultivated
since the Neolithic Age. Altogether, the Rosaceae comprise host plants for at least 47
confirmed plus 23 unconfirmed Auchenorrhyncha species. 13 species are monopha-
gous (10 in the 1st degree, three in the 2nd). Specialists are only found on Rubus, Rosa,
Prunus, Filipendula ulmaria and Potentilla tabernaemontani. Numerous further species,
many of which are oligophagous mesophyll feeders, are found on Cotoneaster, Pyrus,
Malus, Sorbus and Crataegus (see Table 14).
Both Cotoneaster integerrimus and C. tomentosus are restricted to xerothermic and ba-
sic sites in the hilly regions and the Alps. Further species, like C. horizontalis, C. salicifolius
and C. dammeri are cultivated ornamentals. Most Auchenorrhyncha data are from the
latter, but usually without proper host identification. So far only oligo- and polypha-
gous species have been recorded, mostly individuals of Zygina schneideri (Günth.) as
well as the second generation of Edwardsiana rosae (L.). Only in these cases nymphal
development has been documented. Breeding is also likely in Typhlocyba quercus (F.),
 Rosaceae 265

Empoasca vitis (Göthe), Fieberiella septentrionalis W.Wg. and perhaps further oligo- and
polyphagous species. Occasional sweep-netting of Cotoneaster integerrimus on steep hill-
sides in southern Germany had no positive results, but this statement should be consid-
ered preliminary due to small sample size and difficult collecting.
Cultivated and wild pear (Pyrus communis) are both widespread. The first is grown as a
fruit tree in almost all regions, the latter is rather scattered, mainly in floodplain forests and
on xerothermic hillsides from the lowlands at least up to the submontane belt. The same is
true for cultivated and wild apple (Malus domestica). Systematic sampling of Auchenor-
rhyncha on cultivated apple has been done in eastern Germany (Lehmann 1973a) and in
Switzerland (Günthart 1971). Edwardsiana crataegi (Dgl.), f. froggatti (Bak.), clearly dominat-
ed in most sites, locally also E. rosae (L.). Both have been reported to be noxious in England,
Switzerland and other countries. Furthermore, Zygina flammigera (Geoffr.), Empoasca vitis
(Göthe), Alnetoidia alneti (Dhlb.) and Fagocyba cruenta (H.-S.) have been found to reproduce
frequently, although in low individual numbers. In Switzerland (probably also in southern
parts of Germany) these species are accompanied by Zygina schneideri (Günth.), Ribautiana
debilis (Dgl.) (at least in the second generation) and Fruticidia bisignata (M. & R.). Single
specimens of Ewardsiana prunicola (Edw.) and Zygina angusta Leth. should be treated as
temporary immigrants. In contrast, pear has been sampled only very few times. In one case
nymphs of Typhlocyba quercus (F.) were found (R. Remane, pers. comm.). Adult specimens of
Edwardsiana rosae (L.) and Ribautiana tenerrima (H.-S.) were perhaps vagrants. For France,
Ribaut (1936) mentions Pyrus as a food plant of Ribautiana debilis (Dgl.).
The genus Sorbus is represented by at least 5 native species (plus a number of hybrids
and subspecies), none of which is a typical climax tree. A higher dominance, however,
may occur in certain stages of woodland succession, in xerothermic habitats and in the
montane and subalpine belt. Rowan (S. aucuparia) is usually common in open and acidic
woodland sites, particularly along forest margins, in clearings and on pastures from the
lowlands to the upper montane belt. Whitebeam (S. aria) occurs in numerous subspecies,
mainly in xerothermic woodland and on rocky and sun-exposed hillsides in middle and
southern parts of the country, locally dominating along the subalpine tree limit of the
Black Forest and the Alps. Service tree (S. torminalis) often grows syntopically with S. aria,
but also in the interior of beech and oak-hornbeam forests; its strongholds are clearly in
the colline belt. Swedish whitebeam (Sorbus intermedia) is a hybridogenous endemic of
the southern Baltic region not growing in the wild in Germany, but often cultivated in
parks of the northern half. Dwarf whitebeam (S. chamaemespilus) shows similar habitat
requirements as S. aria, but is largely confined to limestone sites of the montane and sub-
alpine belt of the Alps. Finally, sorb (S. domestica) occurs rather localized in thermophil-
ous woodland in the basin of the upper and middle Rhine. As for Auchenorrhyncha, the
two latter species are not studied at all, the others only occasionally. Sorbus torminalis and
S. intermedia have only been sampled in two sites in southern Niedersachsen.
The Auchenorrhyncha fauna of Sorbus is poor in species and individuals. There are
no monophages, and only few oligophages. Polyphages, namely Empoasca vitis (Göthe),
Fagocyba cruenta (H.-S.), Alebra wahlbergi (Boh.) and Alnetoidia alneti (Dhlb.) are domi-
nating. Furthermore, there are some 1st degree oligophages: Edwardsiana rosae (L.) on
S. aria, in Wales according to Claridge & Wilson (1981) also on S. aucuparia, as well as
Zygina flammigera (Geoffr.) on S. intermedia and perhaps S. aria. From Switzerland, Zygi-
266 Utilization of plant resources

Table 14. Auchenorrhyncha species utilizing Rosaceae in Germany. Explanations see Table 5.

Cot oneast er cultivated

Py racant ha coccinea

Amelanchier spp.
Sorbus aucuparia
Malus domest ica
Py rus communis

Crat eagus spp.

Rubus idaeus
S. int ermedia

S. t orminalis

R. frut icosus
Spiraea spp.

R. caesius
S. aria
Species
M acropsis brabantica W.Wg. . . . . . . . . . . . . . X
M acropsis scotti Ed w . . . . . . . . . . . . . X .
Edwardsiana rosaesugans (Cer.) . . . . . . . . . . . . . .
Edwardsiana rhodophila (Cer.) . . . . . . . . . . . . . .
Balcanocerus larvatus (H .-S.) . . . . . . . . . . . . . .
Balcanocerus pruni (Rib.) . . . . . . . . . . . . . .
Z ygina nigritarsis Rem . . . . . . . . . S . . . . .
A rboridia kratochvili (Lang) . . . . . . . . . . . . . .
Eupteryx signatipennis (Boh.) . . . . . . . . . . . . . .
M acrosteles septemnotatus (Fall.) . . . . . . . . . . . . . .
M acropsis fuscula (Zett.) . . . . . . . . . . . X X X
M acropsis megerlei (Fieb.) . . . . . . . . . . . . . .
A rboridia simillima (W.Wg.) . . . . . . . . . . . . . .
Edwardsiana rosae (L.) . X ? X ?* x x . x . . ? ? .
Z ygina schneideri (Günth.) . x . ?* . . . . S x x . . .
Z ygina flammigera (Geoffr.) . . . X ? ? x . x . ? . . .
Edwardsiana crataegi (Dgl.) . . . X . . . ? X . . ? . .
Fruticidia bisignata (M. & R.) . . . ?* . . . . x . . . . .
Ribautiana debilis (Dgl.) . . ?* ?* . . . . . . . . X X
Z ygina rosincola (Cer.) . . . . ?* . . . S . . . . .
Edwardsiana sociabilis (Oss.) . . . . . . . . . . . . . .
Typhlocyba quercus (F.) . ? ? . . x . . ? . . . . .
Edwardsiana prunicola (Ed w .) . . . ? . . . . ? . . ? . .
Z ygina angusta Leth. . . . ? . . . . X . ? . . .
A lebra neglecta W.Wg. . . . . . . . . X . . . . .
Ribautiana tenerrima (H .-S.) . ? . . . . . . . . X X X
A rboridia parvula (Boh.) . . . . . . . . . . . ? . .
Fagocyba cruenta (H .-S.) . . . x x x x x x . . ? . ?
A lnetoidia alneti (Dhlb.) . . . X ?* x . ? x . . ? . .
Empoasca vitis (Göthe) . ? . X X ? X ? x . . X X X
Ledra aurita (L.) . . . x . . . . . . . . . .
A lebra wahlbergi (Boh.) . . . . ? x x x . . . . . .
Issus coleoptratus (F.) . . . . . x . . . . . . . .
Empoasca decipiens Paoli . . . . . . . . . . . ? ? ?
Philaenus spumarius (L.) . . . . . . . . . . . . . .
A guriahana stellulata (Burm .) . . . . . . . . . . . . . .
Fieberiella septentrionalis W.Wg. x ? . . . . . . . . . X x .
Fieberiella florii (Stål) . . . . . . . . . . . x ? .
Lindbergina aurovittata (Dgl.) . . . . . . . . . . . ?* ?* ?*
Cixius sticticus R. . . . . . . . . . . . . . .
Tachycixius pilosus (Ol.) . . . . . . . . ? . . . . .
Reptalus panzeri (P. Löw ) . . . . . . . . . . . . . .
 Rosaceae 267

Table 14. (continued):

Overw intering stage

Filipendula ulmaria
P. t abernaemont ani
Pot ent illa incana

R. spinosissima
R. cory mbifera

Prunus av ium
Fragaria spp.

R. rubiginosa

D iet breadth
P. domest ica
R. pendulina
Rosa canina
P. frut icosa

P. spinosa
P. cerasus

Substrate
R. rugosa
R. glauca

P. padus
. . . . . . . . . . . . . . . . . m1 P eg
. . . . . . . . . . . . . . . . . m1 P eg
. . . . . ? . . X . . . . . . . . m 1? M eg
. . . . . . . . . X . . . . . . . m1 M eg
. . . . . . . . . . . . . . . . X m1 P eg 1st d egree
. . . . . . . . . . . . . . . . X m1 P ad monophages
. . . . . . . . . . . . . . . X . (m 1) M ad
. X . . . . . . . . . . . . . . . m 1? M ad
. . . . X . . . . . . . . . . . . m1 M eg
. . . . X . . . . . . . . . . . . m1 P eg
. . . . . . . . . . . . . . . . . m2 P eg 2nd d egree
. . . . . ? . ? . X ? X . . . . . m2 P eg m ono-
. . . . . . . . . ? . X . . . . . m2 M ad phages
. . . . . X X ? . X X . x . x . ? o1 M eg
. . x . . ? . . . ? X ? ? . ? . X o1 M ad
. . . . ? . . . ? ? . X x* X X X o1? M ad
. . . . . . . . . . . . x ?* x . x o1 M eg 1st d egree
. . . . . . . . . . . . . . . . . o1 M ad oligophages
. . . . . ? . . . . . . . . . . . o1? M eg
. . . . . ? . . . . . . . . . ? ? o1? M ad
. . . . X . . . . . X . . . . . . o1 M eg
. . . . . ? . . . . ? . x x* x X x o2? M eg
. . . . . ? . . . ? . . ? . X . X o2? M eg
. . . . . x . . . ? . . ? ?* ? . x o2 M ad 2nd d egree
. . . . . . . . . . . . ? . . X . o2 M eg oligophages
. . . . . ? . . . . . . ? . . . ? o2? M eg
X . . . ? . . . . . . . . . . ? . o2? M ad
. . . . . ? . . ? ? ? ? x ?* ? x ? po M eg
. . . . . ? . . . . ? . X x* X X x po M eg
. . . . . x . . . x x x x . x x x po P ad
. . . . . . . . . . . . . . . . . po P ny
. . . . . . . . . . . . x ?* x . ? po M eg
. . . . . . . . . . . . . . . . ? po P ny Poly-
. . . . . . . . . . . . ? ?* ? ? ? po M? ad phages
. . . ?* X . . . . . . . . . . . . po X eg
. . . . . . . . . . . . x ?* . . x po M eg
. . ? . . x . . . x x ? . . . . X po P eg
. . . . . x . . . . . . . . . . x po P eg
. . . . . . . . . . . . . . . . . po M eg
. . . . . A . . . ? . . . . . . ? o2? P ny Vertical
. . . . . ? . . . . . . . . . . A po P ny m igrants
. . . . . A . . . ? . . . . . . A o2 P ny
268 Utilization of plant resources

Table 14. (continued):

Cot oneast er cultivated

Py racant ha coccinea

Amelanchier spp.
Sorbus aucuparia
Malus domest ica
Py rus communis

Crat eagus spp.

Rubus idaeus
S. int ermedia

S. t orminalis

R. frut icosus
Spiraea spp.

R. caesius
S. aria
Species
Tibicina haematodes (Scop.) . . . . . . . . . . . . . .
Cicadetta montana (Scop.) . . . . . . . . . . . . . .
A phrophora alni (Fall.) . . . . . . . . . . . . . .
Centrotus cornutus (L.) . . . . . . . . . . . A . .
Stictocephala bisonia K. & Y. . . ?* ?* . . . . ?* . . . . .
Cixius simplex (H .-S.) . . . . . . . . . . . . . .
Cixius stigmaticus (Germ .) . . . . . . . . . . . ? . .
Lepyronia coleoptrata (L.) . . . . . . . . . . . . . .
Evacanthus interruptus (L.) . . . . . . . . . . . ? . .
Emeljanoviana mollicula (Boh.) . . . . . . . . . . . . . .
Empoasca affinis N ast . . . . . . . . . . . . ? ?
Empoasca pteridis (Dhlb.) . . . . . . . . . . . . . .
Empoasca ossiannilssoni N uort. . . . . . . . . . . . . . .
Z yginella pulchra P. Löw . . . . . . . . ? . . . . .
Z ygina lunaris (M. & R.) . . . . . . . . . . . . .
Fruticidia sanguinosa (R.) . . . . . . . . ?* . . . . .
Z ygina tiliae (Fall.) . . . . . . . . . . . . . .
Edwardsiana frustrator (Ed w .) . . . . ?* ? ? . ? . . ? . .
Edwardsiana lamellaris (Rib.) . . . . . . . . . . . . . .
Edwardsiana lanternae (W.Wg.) . . . . ?* . . . . . . . . .
Platymetopius major (Kbm .) . . . . . . . . . . . . . .
Idiodonus cruentatus (Panz.) . . . . . . . . . . . ? . .
Colladonus torneellus (Zett.) . . . . . . . . . . . ? . .
Speudotettix subfusculus (Fall.) . . . . . . . . . . . . . .
Thamnotettix confinis (Zett.) . . . . . . . . . . . ? . .
Sonronius dahlbomi (Zett.) . . . . . . . . . . . ?* . .
A thysanus quadrum Boh. . . . . . . . . . . . . . .
Total substantiated 1 2 0 7 2 6 5 2 12 1 1 6 6 5
Total unconfirm ed 0 3 5 6 7 3 1 3 7 0 2 15 5 4
* = after Claridge & Wilson (1981), Günthart (1980, 1987b), Hoffrichter & Tröger (1973), Lauterer & Malen-
Vasarainen (1970), Remane (pers. comm.), Ribaut (1936), Vidano & Arzone (1987b)

na rosincola (Cer.) is reported from S. aucuparia (Günthart 1987b). Among 2nd degree
oligophages, only Typhlocyba quercus (F.) was found on S. aria. In Britain, Edwardsiana
frustrator (Edw.) and E. lanternae (W.Wg.) are reported from S. aucuparia (Claridge &
Wilson 1981; Le Quesne & Payne 1981), but their host range in central Europe is imper-
fectly known. Another lack of data exists for Alebra sorbi W. Wagner, 1949 described
from S. aria in the eastern Alps, but otherwise only recorded from ornamental trees in
urban areas of Warsaw (see Dworakowska 1993). A number of S. aria sampled in the
Bavarian Alps and in warm regions of southern and middle parts of Germany were
only infested by the polyphagous A. wahlbergi (Boh.).
 Rosaceae 269

Table 14. (continued):

Overw intering stage

Filipendula ulmaria
P. t abernaemont ani
Pot ent illa incana

R. spinosissima
R. cory mbifera

Prunus av ium
Fragaria spp.

R. rubiginosa

D iet breadth
P. domest ica
R. pendulina
Rosa canina
P. frut icosa

P. spinosa
P. cerasus

Substrate
R. rugosa
R. glauca

P. padus
. . . . . . . . . . . . . . . . A po X ny
. . . . . . . . . . . . . . . . A po X ny Vertical
. . . ?* ?* A . . . ? ? . . . . . A po X eg m igrants
. . . . . . . . . . . . . . . . ? po P ny
. . . . . ?* . . . . . . ? . . . ? po P eg
. . . . . . . . . . . . . . . . ? po P ny
. . . . . . . . . . . . . . . . . po P ny
. . . . ?* . . . . . . . . . . . . po X eg
. . . . . . . . . . . . . . . . . po X eg
. . . ?* . . . . . . . . . . . . . po M eg
. . . . . . . . . . . . . . . . ? po M? ad
. . . . . . . . . . . . ? ?* ? . ? po M? ad ?
. . . . . . . . . . . . . . . ?* . o2? M? ad
. . . . . . . . . . . . . . . . . m2 M ad
. . . . . ?* . . . . . . . . . . . m 2? M ad
. . . . . . . . . . . . . . ?* . . o1? M ad Uncon-
. . . . . . . . . . . . . . ? ? ? m 2? M ad firm ed
. . . . . . . . . . . . . . . . ? po M eg
. . . . . ? . . . . . . . . . . . o2? M eg
. . . . . . . . . . . . . . . . . o2 M eg
. . . . . . . . . . . . . . . . ? po P eg
. . . . . . . . . . . . . . . . . po P eg
. . . . . . . . . . . . . . . . . po? P ny
. . . . . . . . . . . . . . . . ? po P ny
. . . . . . . . . . . . . . . . . po P ny
. . . . ?* . . . . . . . . . . . . m 1? P eg
. . . . ? . . . . . . . . . . . . m 1? P eg
1 1 1 0 4 8 1 0 1 5 5 3 9 3 8 7 18
0 0 1 3 5 15 0 2 1 9 6 3 8 7 7 5 17
ovsky (1995), Lehmann (1973a), Le Quesne & Payne (1981), Ossiannilsson (1981, 1983), Raatikainen &

Hawthorn, mainly Crataegus monogyna and C. laevigata, is widely and densely spread
in damp to dry and sunny sites from the lowlands up to the montane belt. Species
identification may be difficult due to hybridization. Further ornamental species have
been planted in gardens and parks. Thus, the majority of data on Auchenorrhyncha
only generally refer to the genus Crataegus. Most species recorded belong to oligo- and
polyphagous, mesophyll-feeding typhlocybids. Hawthorn is one of the main hosts of
Zygina angusta Leth. (which is also found on other species of Rosaceae as well as on
Fagaceae), and for Alebra neglecta W.Wg. (also breeding on Prunus padus and Carpinus
betulus). Further oligophagous species include Edwardsiana rosae (L.), E. crataegi (Dgl.),
270 Utilization of plant resources

Zygina flammigera (Geoffr.) and Fruticidia bisignata (M. & R.). Fagocyba cruenta (H.-S.),
Alnetoidia alneti (Dhlb.) and Empoasca vitis (Göthe) are all polyphagous. As an early spring
food plant, Crataegus plays a role for adults of several species of Zygina, which prefer
fresh shooting leaves after hibernation (Remane 1994).
Fire thorn (Pyracantha coccinea), originating from the eastern Mediterranean region is
often grown in gardens. At least in Hessen it harbours reproducing populations of Zy-
gina schneideri (Günth.) (R. Remane, pers. comm.).
Among the genus serviceberry (Amelanchier) only three species are noteworthy, none
of which has been studied thoroughly. The only native species is A. ovalis, locally grow-
ing on basic, and usually, rocky xerothermic sites of southern Germany. A. lamarckii and
A. spicata are both cultivated, but have locally become well established in the under-
growth of forests in northern Germany. Altogether, only two alpine stands of A. ovalis
have been swept, yielding nymphs and freshly emerged individuals of an unidentified
species of Zygina. Locally Zygina schneideri (Günth.) is reproducing on cultivated servi-
ceberry (probably A. spicata). Altogether, because of the small sample size, no definite
statements can be given on the central European Auchenorrhyncha fauna on this plant
genus. At least some more polyphagous species should be expected to reproduce.
Five autochthonous species of Rubus are found in Germany. Cloudberry (R. chamaemorus)
is restricted to very few bog sites in the north. Stone bramble (R. saxatilis) is mainly found
in woodlands of limestone regions as well as in subalpine scrub, locally also in the north
German plain. Both species have not been studied so far, but are probably less suitable for
Auchenorrhyncha because they are much localized and herbaceous. The remaining spe-
cies of Rubus are all widespread and common and include low-growing or creeping shrubs
in open forests, along margins and on clearings. Raspberry (R. idaeus) is very common
from the plains up to the upper montane belt, usually in moist, acidic and loamy sites, but
is also cultivated in gardens. The latter is also true for bramble (R. fruticosus agg.), but its
strongholds are in sandy and damp sites of mild winter climates. More than 80 subspecies
are known to occur in Germany alone, many of which are autogamous and which can
only be distinguished by specialists. Dewberry (R. caesius) prefers valley sites on more
clayey and basic soils, which may even be subject to flooding.
Within the leafhopper genus Macropsis, a small tribe has become associated with Rubus.
Macropsis fuscula (Zett.) lives on all three species studied, M. scotti Edw. and M. braban-
tica W.Wg. are restricted to R. fruticosus and R. caesius, respectively. The two latter are
also the main hosts of Ribautiana debilis (Dgl.) and R. tenerrima (H.-S.). Further polypha-
gous species on Rubus include Empoasca vitis (Göthe), Fieberiella septentrionalis W.Wg., F.
florii (Stål) and probably a number of generalists. From Wales bramble is reported as the
host of the spring generation of Lindbergina aurovittata (Dgl.).
Further cultivated woody plants, such as quince (Cydonia oblonga), medlar (Mespilus
germanica), bridewort (Spiraea spp.), shrubby cinquefoil (Potentilla fruticosa) and others,
have only occasionally been sampled. The two latter are locally infested by some oli-
gophages and polyphages, like Empoasca vitis (Göthe), Zygina schneideri (Günth.) and
Fieberiella septentrionalis W.Wg.
Among herbaceous species, only meadowsweet (Filipendula ulmaria), cinquefoil (Po-
tentilla spp.) and perhaps strawberry (Fragaria spp.) are known to be attacked, the latter
only by generalists. Meadowsweet, which is very common and widespread all over the
 Rosaceae 271

country, holds the 1st degree monophages Eupteryx signatipennis (Boh.) and Macrosteles
septemnotatus (Fall.). It is further attacked by some generalists and is probably the na-
tive host of Edwardsiana sociabilis (Oss.), which is mainly found on the ornamental Rosa
rugosa originating from eastern Asia. Arboridia kratochvili (Lang) is a rare and localized
central European endemic of spring cinquefoil (Potentilla tabernaemontani) in extremely
xerothermic sites. The taxonomic situation of Arboridia populations on sand cinquefoil
(P. incana) in eastern Germany requires further studies, but they may belong to A. poten-
tillae (Mor.). From Moravia (Czech Republic), salad burnet (Sanguisorba minor) is known
to be the exclusive host of the typhlocybid Zygina frauenfeldi Leth. (Lauterer 1980), which
may also be expected to occur in southern Germany.
Roses (Rosa spp.) are difficult to sample and even more difficult to identify. Thus, most
published data on their Auchenorrhyncha fauna only generally refer to Rosa. Commonly,
more than 25 species are recognized in the wild in Germany, most of which grow in rather
dry and sunny sites; the number of ornamental species is difficult to assess, but is proba-
bly higher. Only 6 species, all of which are easy to identify, have been sampled for leaf-
hoppers. Sampling of less known roses may reveal further, perhaps even undescribed
species. Edwardsiana rhodophila (Cer.) and E. rosaesugans (Cer.) are both strictly monopha-
gous on sweet briar (Rosa rubiginosa) and alpine rose (R. pendulina), respectively, although
a few specimens of the latter have also been found on dog rose (R. canina). Burnet rose
(R. spinosissima) is an important, but not exclusive host of Macropsis megerlei (Fieb.) and
Arboridia simillima (W.Wg.). Furthermore, there are some oligophages, notably Edwardsi-
ana rosae (L.), E. sociabilis (Oss.) and Zygina schneideri (Günth.). E. sociabilis (Oss.) is rather
uncommon on Filipendula ulmaria in near-natural sites, but has infested ornamental Japa-
nese rose (Rosa rugosa) in many parts of central Europe. Dog rose (R. canina) which is
probably the most common and widespread among central European roses, is not known
to harbour any specialists, but it is utilized by numerous oligo- and polyphages, includ-
ing both species of Fieberiella as well as some vertical migrants such as Cixius sticticus R.,
Reptalus panzeri (P. Löw) and Aphrophora alni (Fall.). It is also likely to be the host of one
generation of the rare Edwardsiana lamellaris (Rib.). Corymb rose (Rosa corymbifera) is per-
haps equally common, but is probably often confused with the latter species.
The genus Prunus is represented by 5 native species. Wild cherry (P. avium, with the
cultivated ssp. juliana) and bird cherry (P. padus), both of which are widespread, but not
common, from the lowlands up to the montane belt, prefer deciduous forests and their
margins, the latter mainly in seepy or temporarily flooded sites. Sloe (Prunus spinosa) is
widespread and common in pastures, along waysides and margins of forests, fields and
meadows as well as in abandoned grassland, mainly in sunny, moderately dry or damp
sites. Dwarf and Mahaleb cherry (P. fruticosa, P. mahaleb) are both rare and restricted to
xerothermic sites of south Germany. There is also a number of introduced species, some of
which have been cultivated since the Roman or even Neolithic Age, like sour cherry (Pru-
nus cerasus), plum (P. domestica), peach (P. persica) and almond (P. dulcis), but also neo-
phytes like cherry laurel (P. laurocerasus) and American black cherry (P. serotina). Only the
first three species and plum have been sampled for Auchenorrhyncha.
Prunus is another important host group of central European leafhoppers, with alto-
gether three 1st degree monophages. Balcanocerus larvatus (H.-S.) and B. pruni (Rib.) are
both exclusively associated with sloe, whereas Zygina nigritarsis Rem. exclusively re-
272 Utilization of plant resources

produces on bird cherry, feeding on conifers in winter, and on Crataegus and other woody
species of Rosaceae in spring. Further, Prunus provides the main host plants for some
oligophages, such as Alebra neglecta W.Wg., Edwardsiana prunicola (Edw.), Typhlocyba quer-
cus (F.), Zygina flammigera (Geoffr.) and Z. schneideri (Günth.). Other oligophages and
polyphages include Edwardsiana rosae (L.), E. crataegi (Dgl.), Alebra wahlbergi (Boh.), Em-
poasca vitis (Göthe), Fagocyba cruenta (H.-S.), Aguriahana stellulata (Burm.) and Alnetoidia
alneti (Dhlb.). With altogether 18 Auchenorrhyncha species, sloe holds one of the most
diverse guilds of all German plants (Fig. 14).
Altogether, despite the predominance of herbaceous species within the rose family
(see above), trees and shrubs are clearly favoured by Auchenorrhyncha. With few ex-
ceptions, notably Potentilla and Filipendula, herbs remain almost completely uninfested.
There is also a clear tendency of native species harbouring more phytophages than in-
troduced ones (including archaeophytes). The high proportion of 1st degree oligophag-
es is certainly biased by the taxonomy of the Rosaceae, which – unlike e.g. Fagaceae and
Aceraceae – include numerous tree genera. In general, mesophyll feeders dominate,
even on herbaceous species, although among monophages, phloem feeding is also wide-
spread. Most species overwinter as egg, and those wintering as adults migrate to ever-
green plants in autumn.
For Britain, Leather (1985) has compiled information on phytophagous insects on
Prunus. He found (i) that species numbers of phytophagous insects tend to be higher on
native than on cultivated plants, (ii) that widespread plants hold more phytophages
than localized plants, and (iii) that P. padus had an unexpectedly low number of phy-
tophages. The Auchenorrhyncha data, which are mainly based on studies by Claridge
& Wilson (1976, 1981), seem roughly comparable to those from Germany. As shown in
Fig. 3, British species numbers are generally lower on all studied Prunus species (which

25

P. avium P. c erasus P. domestic a P. padus P. spinosa

20
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0

Fig. 3: Auchenorrhyncha species numbers on Prunus in Britain (left, dotted) and Germany (right, hatched/
black). Monophages are shown in black. British data according to Leather (1985), German data on P. cerasus
after Lehmann (1973a).
 Fabaceae 273

is a general pattern in arboricolous arthropods – see Brändle & Brandl 2001), with a
total lack of monophages, although the recently described Zygina nigritarsis Rem. may
have been overlooked. Noteworthy is also a marked decrease of species numbers on
P. spinosa and P. padus. These differences can be largely explained zoogeographically,
i.e. by the general absence of some insect species on the British Islands, e.g. Tibicina
haematodes (Scop.), Alebra neglecta W.Wg. and Fieberiella florii (Stål), by the higher taxo-
nomic resolution within the genus Zygina in Germany, and perhaps the focus of British
studies on mesophyll-feeding Typhlocybinae (Claridge & Wilson 1976, 1981). However,
among the genera Alebra, Fagocyba and Edwardsiana, there is some evidence that host
ranges become narrower towards the edge of the range, as was found in some gramini-
colous species within central Europe (see section 5.1.2.19).

5.1.2.12 Fabaceae
Fabaceae are among the dominating herbs in many grassland habitats, although Cytisus
and Genista, both of which are woody, are restricted to abandoned sites and woodland
margins. Only relatively few species, none of which is known to be utilized by Auchenor-
rhyncha, are found in the interior of forests. Regarding species richness, the Fabaceae are
among the most diverse plant families in Europe, with more than 160 species occurring in
Germany alone, including about 20 shrubs and dwarf shrubs. The data base for Auchen-
orrhyncha, however, is weak and rather unspecific, except in some woody species.
In general, this plant family is of moderate importance as food resource for
Auchenorrhyncha. Concerning host specialists, the few woody species are strongly
favoured, with broom as the most-frequented species, whereas most other species
are attacked by generalists only. Phloem feeding and overwintering in the egg stage
clearly prevail.
The Auchenorrhyncha fauna comprises 21 confirmed plus 8 unconfirmed feeders
(Table 15). A distinct guild of 5 species lives on common broom (Cytisus scoparius).
Dryodurgades antoniae (Mel.) is a 1st degree monophage, Handianus ignoscus (Mel.)
was also found on black broom (C. nigricans) at least in southern Europe, whereas
Gargara genistae (F.), Batracomorphus allionii (Turt.) and Euscelis ohausi W.Wg. are 1st
degree oligophagous, also found on whin (Genista spp.) and other predominantly
woody species. E. ohausi W.Wg. and H. ignoscus (Mel.) prefer low-growing plant indi-
viduals. Genista is furthermore favoured by Stictocoris picturatus (C. Shlb.), which also
lives on restharrow (Ononis spp.) and winged broom (Chamaespartium sagittale). Gar-
gara genistae (F.) is apparently a less specific feeder to various woody Fabaceae, in-
cluding restharrow Ononis spp., bladder senna (Colutea arborescens) and even herba-
ceous species, like crownvetch (Coronilla spp.) and perhaps sainfoin (Onobrychis spp.).
Dryodurgades reticulatus (H.-S.) is the only strictly monophagous feeder on herbaceous
Fabaceae, living on fine-leaved vetch (Vicia tenuifolia), although wintering may take
place on coniferous trees. Most of the remaining food plant records are from Medicago,
Trifolium, Lotus and Onobrychis, and refer to generic level only. Leafhoppers involved
belong to the taxa Megophthalmus, Agalliinae, Aphrodes and Euscelis, most of which
are oligophagous in the 1st or 2nd degree. Polyphages include Lepyronia coleoptrata
(L.), Philaenus spumarius (L.), Empoasca decipiens Paoli, E. pteridis (Dhlb.) and probably
 274 Utilization of plant resources

Table 15. Auchenorrhyncha species utilizing Fabaceae in Germany. Explanations see Table 5.

Chamaespart ium sagit t ale

Overw intering stage

Lupinus poly phy llus

Hippocrepis comosa
Colut ea arborescens
Lot us corniculat us
Cy t isus scoparius

Genist a t inct oria

O nobry chis spp.

O nonis spinosa

Vicia t enuifolia
Medicago spp.

Coronilla spp.
Trifolium spp.

spp.
Lat hy rus spp.

D iet breadth
C. nigricans

Vicia spp.

Substrate
Species
Dryodurgades reticulatus (H .-S.) . . . . . . . . . . . . . X . . m1 P ad Mono-
Dryodurgades antoniae (Mel.) . X . . . . . . . . . . . . . . m1 P eg phages
Handianus ignoscus (Mel.) . x ? . . . . . . . . . . . . . m 2? P eg
Batracomorphus allionii (Turt.) . X . x . . . . . . . . . . . . o1 P eg
Euscelis ohausi W.Wg. . X . x . . . . . . . . . . . . o1 P eg
Gargara genistae (F.) . X ?* x . x ?* ?* . x x . . . . . o1 P eg
Stictocoris picturatus (C. Shlb.) . . . X x X . ? . . . . . . . . o1 P eg
M egophthalmus scanicus (Fall.) . . . . . . x x x . . . . . . . o1 P eg
A phrodes bicincta (Schrk.) . ? . . . . ?* X . . . . . . . . o1? P eg Oligo-
A phrodes diminuta Rib. . . . . . . . ?* . . . . . . ?* . o1? P eg phages
Euscelis lineolatus Br. . . . . . . x* x* ?* . . . . . . . o2 P eg
Euscelis incisus (Kbm .) . . . . . . ? X . . . . ? . . . o2 P ny/ eg
A naceratagallia ribauti (Oss.) . . . . . . x* x* . . . . x* . . . o2? P ad
A gallia brachyptera (Boh.) . . . . . . . ? . . . . x* . . . o2? P eg
A naceratagallia venosa (Geoffr.) . . . . . . . . x . . x* . . . . o2? P eg
Empoasca decipiens Paoli . ? . . . . ?* . . . . . . . . . po M? ad Poly-
Empoasca pteridis (Dhlb.) . . . . . . x ? . . . . . . ? . po M? ad ? phages
Philaenus spumarius (L.) . . . . . . . ?* x* . . . . . ? ?* po X eg
Cercopis vulnerata Rossi A . . . . . . . . . . . . . . . po X ny Vertical
A phrophora alni (Fall.) . A . . . . . ?* ?* . . . . . . . po X eg m igrants
Stictocephala bisonia K. & Y. . . . . . . A . . . . . . . . . po P eg
Lepyronia coleoptrata (L.) . . . . . . . ?* . . . . . . ?* ?* po X eg
Utecha trivia (Germ .) . . . . . . . . . . . ?* . . . . po? P ad
M egophthalmus scabripennis Ed w . . . . . . . . . . . . . . . . . o1? P eg
A naceratagallia frisia (W.Wg.) . . . . . . . . . . . . . . . . o2? P ad Uncon-
Eupteryx atropunctata (Goeze) . . . . . . . . . . . . . . ?* . po P eg firm ed
A thysanus quadrum Boh. . . . . . . . . . . . . . . . ? o1? P eg
Handianus procerus (H .-S.) . . . ? . . . . . . . . . . . . o2? P eg
Euscelis venosus (Kbm .) . . . . . . . ? . . . . . . . . o2? P eg
Total 1 6 0 4 1 2 5 5 3 1 1 1 2 1 0 0

* = after Halkka et. al (1977), Nickel (1979), Ossiannilsson (1981), Schedl (1998), Schiemenz (1988), Stewart (1988),
Tishechkin (pers. comm.), Witsack (1985)

others. As to vertical migrants, the introduced Stictocephala bisonia Kopp & Yonke is
an important feeder on Medicago and may invade cultivated stands in the upper Rhine
plain. From Moravia Kyboasca bipunctata (Osh.), which is a monophagous elm feeder
in Germany, is known to live on licorice-root (Glycyrrhiza glabra), whereas Handianus
procerus (H.-S.), the status of which is uncertain in Germany, lives on Genista pilosa (P.
Lauterer, pers. comm.).
 Aceraceae 275

5.1.2.13 Aceraceae
In the study area the maple family is represented by only 5 native tree species of a
single genus. Thus, as in Ulmaceae and Tiliaceae, the groups of 2nd degree monophag-
es and 1st degree oligophages are identical. Introduced ornamentals include ashleaf,
sugar, silver and Japanese maple (Acer negundo, A. saccharum, A. saccharinum, A. palma-
tum) and some others. Only the latter is known to be infested by leafhoppers, although
the data base is small. As to native species, Italian Maple (Acer opalus) is confined to a
few sites in the extreme southwest near the Swiss border and has not been studied.
Montpellier maple (A. monspessulanum) is also confined to southwestern parts, but is
more widespread in xerothermic sites in the basins of the Moselle, Nahe and Main. The
only sampled stand is situated in Franconia at the very northeastern edge of the range.
The three remaining species are widespread and common over most of central Europe
and are often planted in parks and alleys. Their Auchenorrhyncha fauna is rather well-
studied. Sycamore (Acer pseudoplatanus) has its strongholds in rather cool, hilly to moun-
tainous situations, and is found up to the subalpine tree limit, whereas Norway and
field maple (Acer platanoides and A. campestre) occur in various types of deciduous for-
ests, usually below 1000 m a.s.l.
Aceraceae provide important leafhopper food plants, altogether harbouring 17 con-
firmed plus 10 unconfirmed species (Table 16). Acer pseudoplatanus is of particular
importance and holds the only 1st degree monophages, notably Ossiannilssonola cal-
losa (Then), Edwardsiana nigriloba (Edw.), Eurhadina loewii (Then) and perhaps Acericerus
heydenii (Kbm.). Along with Zyginella pulchra P. Löw, which also breeds on other ma-
ples, these species form a distinct guild, supplemented by Acericerus ribauti Nick. &
Rem., A. vittifrons (Kbm.) and several polyphages. The two latter species’ main host is
Acer campestre, which is also utilized by Japananus hyalinus (Osb.), Edwardsiana lethier-
ryi (Edw.) and a number of polyphages. E. lethierryi (Edw.) lives on Tilia cordata and
T. platyphyllos as alternative hosts. Guilds on Acer platanoides and A. monspessulanum
are less specific and less diverse. They both harbour only low densities of a few 2nd
degree monophages as well as of some polyphages. Polyphagous species common to
all or most species of maple include the mesophyll-feeding typhlocybids Alebra wahl-
bergi (Boh.), Fagocyba cruenta (H.-S.) and Alnetoidia alneti (Dhlb.), as well as the plan-
thopper Issus coleoptratus (F.), the phloem-feeding typhlocybid Empoasca vitis (Göthe)
and the vertical migrants Tachycixius pilosus (Ol.) and Allygus mixtus (F.). Apart from
that, Edwardsiana frustrator (Edw.) has been found to breed on Acer pseudoplatanus and
A. campestre in Wales (Claridge & Wilson 1981). Arboridia erecta (Rib.), the host plants
of which are unknown in central Europe, reproduces on A. campestre in Italy (Vidano
& Arzone 1987b). Finally, the Japanese Acer palmatum has been found to hold Japana-
nus hyalinus (Osb.) in at least one case.
In general, despite their low plant species number, the Aceraceae play a significant
role as Auchenorrhyncha food resources, with high values of absolute numbers of feed-
ing species as well as of the ratio host specialists : generalists, although differences within
the genus Acer are considerable. There is a slight dominance of mesophyll feeders over
phloem feeders. As in other arboricolous guilds, most species overwinter as egg, but,
compared to other deciduous trees, overwintering as adult is also common.
276 Utilization of plant resources

Table 16. Auchenorrhyncha species utilizing Aceraceae in Germany. Explanations see Table 5.

Overw intering stage

Acer pseudoplat anus

A. monspessulanum
A. plat anoides

D iet breadth
A. palmat um
A. campest re

Substrate
Species
Ossiannilssonola callosa (Then) X m1 M eg
Edwardsiana nigriloba (Ed w .) X m1 M eg
Eurhadina loewii (Then) X m1 M eg
A cericerus heydenii (Kbm .) X ? ? ? m 1? P ad Monophages
Z yginella pulchra P. Löw X x x x m2 M ad
A cericerus ribauti N ick. & Rem . x x X m2 P ad
A cericerus vittifrons (Kbm .) x ? X m2 P ad
Japananus hyalinus (Osb.) ? ? X ? m2 P eg
Edwardsiana lethierryi (Ed w .) ? X o2 M eg
Issus coleoptratus (F.) x x x x po P ny
A lebra wahlbergi (Boh.) x X X po M eg
Empoasca vitis (Göthe) x ? x ? po P ad Oligo-/ Polyphages
Fagocyba cruenta (H .-S.) X X x ? po M eg
A lnetoidia alneti (Dhlb.) X X x po M eg
A guriahana stellulata (Burm .) x ? po M eg
Tachycixius pilosus (Ol.) A po P ny Vertical
A llygus mixtus (F.) A A po P eg m igrants
A phrophora alni (Fall.) ? ? po X eg
Ledra aurita (L.) ? po P ny
Empoasca decipiens Paoli ? po M? ad
Edwardsiana flavescens (F.) ? o2 M eg
Edwardsiana ampliata (W.Wg.) ? ? o2? M eg Uncon-
Edwardsiana frustrator (Ed w .) ?* ?* po M eg firm ed
Ribautiana tenerrima (H .-S.) ? ? o2? M eg
A rboridia erecta (Rib.) ? ?* o2? M ad
Fieberiella septentrionalis W.Wg. ? ? po P eg
Platymetopius major (Kbm .) ? ? po P eg
Total 13 7 12 2 0
* = after Claridge & Wilson (1981), Vidano & Arzone (1987b)

5.1.2.14 Apiaceae
With 93 species in Germany, the carrot family is among the more diverse plant groups.
A large proportion of the native species, however, are annuals or biennials and are thus of
minor suitability as host plants. With few exceptions, their abundance and biomass is
certainly negligible in most ecosystems, although human activities have probably favoured
a number of heliophilous species. Umbellifer dominance is often restricted to sites, where
shading by trees is reduced and where supply of humidity and nitrogen is ample.
Ethereal oils are widespread, and as a consequence, many species are used as spices
and medical herbs (Frohne & Jensen 1998). Additionally, the group is unique in the
 Lamiaceae 277

Table 17. Auchenorrhyncha species utilizing Apiaceae in Germany. Explanations see Table 5.

Heracleum sphondy lium

Aegopodium podagraria
Chaerophy llum aureum

Ant hriscus sy lv est ris

Overw intering stage

H. mant egaz z ianum
Angelica sy lv est ris
Falcaria v ulgaris

Past inaca sat iv a

Ch. aromat icum

Daucus carot a

D iet breadth
Ch. hirsut um

Substrate
Species
Eupteryx heydenii (Kbm .) ? X m 2? M eg Monophage
Eupteryx atropunctata (Goeze) x po M eg
Eupteryx aurata (L.) x ? X x po M eg Polyphages
Philaenus spumarius (L.) x po X eg
Hyalesthes obsoletus Sign. A* po P ny Vertical
Cercopis sanguinolenta (Scop.) A* po X ny m igrants
Emelyanoviana mollicula (Boh.) ?* po M eg
Cercopis vulnerata Rossi ? po X ny Un-
Lepyronia coleoptrata (L.) ?* po X eg certain
A phrophora alni (Fall.) ?* po X eg
Total 0 2 0 1 0 1 1 0 1 1 1
* = after Biedermann (1998a), Halkka et al. (1977), Vidano (1965), Weber (pers. comm.)

abundance and structural diversity of substituted coumarins, and has been subject to
studies on the evolutionary ecology of specialized insects, providing arguments for the
classical idea of insect – plant coevolution (Berenbaum 1990).
Auchenorrhyncha species utilizing umbellifers almost exclusively belong either to the
typhlocybid genus Eupteryx, which is largely associated with aromatic herbs such as Lami-
aceae and Asteraceae, or to xylem-feeding cercopids (Table 17). Thus, only Philaenus spu-
marius (L.), Eupteryx aurata (L.), E. atropunctata (Goeze) and E. heydenii (Kbm.) are known
to breed frequently, and adults of two vertical migrants have been found. E. heydenii (Kbm.)
is the only host specialist, being confined to hairy chervil (Chaerophyllum hirsutum), which
is a very common plant growing along streams and forest tracks in mountain regions. It
seems tempting to presume that secondary plant compounds of umbellifers play an im-
portant role in deterring herbivores, but – as stated above – predictability and availability
of resources are also much reduced, and may provide alternative explanations. Finally, it
should be noted, that Eupteryx aurata (L.) is one of the few native herbivore species attack-
ing the aggressive neophytic giant hogweed (Heracleum mantegazzianum). Thus, there is
perhaps some potential for biological control, which may deserve further attention.

5.1.2.15 Lamiaceae
The mint family is represented in Germany by almost 100 species, the majority of
which grow in sun-exposed or semi-open sites (mainly pastures and non-managed grass-
land), as well as in the undergrowth of woodland. Most species are perennial herbs,
with few annuals and dwarf shrubs. Like the Apiaceae, many have been cultivated as
spices and medical plants in gardens, at least since the Roman Age, due to their content
278 Utilization of plant resources

of ethereal oils, which are usually stored in gland-headed hairs. Further compounds
include iridoids and phenolic glycosides (Judd et al. 1999).
Most genera support at least a few Auchenorrhyncha species (Table 18). The Auchen-
orrhyncha guild comprises at least 26 species, 15 of which belong to the genus Eupteryx,
plus 13 unconfirmed feeders. Altogether, 32 plant species have been found to be at-
tacked, although Scutellaria, Melittis, Galeopsis, Phlomis and Horminum have only rarely
been sampled. Dwarf shrubs are clearly favoured, with 6 species feeding on cultivated
garden sage (Salvia officinalis), occasionally causing damage, and altogether 5 species
on thyme (Thymus spp.). Further important host taxa include Teucrium, Lamium, Stachys
and Mentha. There are no positive feeding records on annuals, although temporary feed-
ing on Ocimum and Satureja in gardens by oligo- and polyphages is likely to occur, pro-
vided perennial hosts are nearby, where overwintering can take place, e.g. Salvia or
Melissa (see Stewart 1988). Among 1st degree monophages, Eupteryx immaculatifrons
(Kbm.) lives on spotted dead-nettle (Lamium maculatum), E. lelievrei (Leth.) on betony
(Betonica officinalis), E. origani Zachv. on marjoram (Origanum vulgare), E. collina (Fl.) on
horse mint (Mentha longifolia), and the rare Chlorita pusilla (Mats.) on breckland thyme
(Thymus serpyllum). 2nd degree monophages are only found on thyme, with Chlorita
dumosa (Rib.) and probably Goniagnathus brevis (H.-S.). Most oligo- and polyphages ex-
ploit a wider range of plant genera among the Lamiaceae, except Eupteryx thoulessi Edw.,
which in most sites is clearly monophagous on water mint (Mentha aquatica), but also
breeds on gipsywort (Lycopus europaeus), where both plants grow syntopically; the lat-
ter is probably host in the second generation only.
Among all plant families, the Lamiaceae are outstanding in showing the highest pro-
portion of mesophyll feeders, whereas exploitation of xylem and phloem is rare. Simi-
larly high values are only found in some further groups of dicotyledonous herbs, such
as the Ranunculaceae, Geraniaceae, Malvaceae, Clusiaceae, Scrophulariaceae and Dip-
sacaceae, although these groups are infested only by very few species.
Stewart (1988) has compiled the British feeding data of the genus Eupteryx and sug-
gested that niche separation by host species is much evident, and that overlap is small,
except on Urticaceae, where coexistence of several leafhopper species is facilitated by
more subtle mechanisms (see also section 6.1.4). Central European data on Lamiaceae –
leafhopper associations may support this view. In contrast to many other plant groups,
where Auchenorrhyncha infestation is much clumped on certain host species, e.g. with-
in the Salicaceae or the Poaceae, the utilization of Lamiaceae appears to be well-bal-
anced, mostly with no more than three leafhopper species per plant species. Even on
much-favoured hosts, like wood sage (Teucrium scorodonia), ground-ivy (Glechoma hed-
eracea), hedge woundwort (Stachys sylvatica) or horse mint (Mentha longifolia), there is
either only one dominating leafhopper species, or the leafhopper species belong to tax-
onomically distant groups, or they show distinct habitat preferences with only little
overlap (see Table 18). The only exception is sage, with at least 6 feeding species, but
this is an archaeophyte and should thus be excluded from comparison.
Interestingly, the many examples of rather diverse Auchenorrhyncha guilds on other
plants such as Phragmites, Populus, Betula, Salix, Calamagrostis and Festuca (see the rele-
vant sections of this chapter) largely encompass phloem feeders. Thus, the competitive
relationship among mesophyll-feeding species, the resources of which will become def-
 Asteraceae 279

initely exhausted after heavy infestation, is perhaps quite different from the situation in
other sap feeders, which simply tap an abundant resource without seriously injuring
the plant. These differences may be crucial for the understanding of Auchenorrhyncha
– plant relationships.
Although members of the genus Eupteryx have been found on 17 different plant fam-
ilies, the most frequently exploited group is clearly the Lamiaceae, where most host
specialists occur (Stewart 1988). Thus, like in paralimnine leafhoppers and graminoids,
or in the delphacid genus Kelisia and the Cyperaceae, there appears to be a particular
relationship with a certain plant group. Other plants attacked by Eupteryx include spe-
cies of Malvaceae, Apiaceae and Asteraceae. The common trait of these groups with the
Lamiaceae is the presence of ethereal oils (see Judd et al. 1999). A further interesting
aspect of Auchenorrhyncha on Lamiaceae is the invasion or at least range expansion of
Mediterranean species on medicinal or ornamental plants into gardens of central Eu-
rope, discussed in chapter 7.3.3.

5.1.2.16 Asteraceae
This is by far the most diverse vascular plant family in central Europe, with c. 80
genera and 457 species occurring in Germany. These include 170 species of Hieracium as
well as 11 sections of Taraxacum (counted here as species), the latter comprising more
than 370 apomictic subspecies. The majority is herbaceous, although wormwood (Arte-
misia spp.) and a few others do comprise dwarf shrubs. Asteraceae species are usually
common in many managed, abandoned, or near-natural grassland habitats and – to a
lesser extent – in various types of woodland.
30 confirmed plus 5 unconfirmed Auchenorrhyncha species have been found on
members of this plant family. Dwarf shrubs are clearly favoured, and the genus Arte-
misia has the highest species numbers, including some host specialists. Goldenrod
(Solidago), yarrow (Achillea), butterbur (Petasites), ragwort (Senecio), thistle (Carduus)
and dandelion (Taraxacum) are also of importance. The number of attacked Asterace-
ae species probably exceeds 30, but some feeding relations may be obscured by host
identification problems, particularly among difficult groups such as Crepis, Hieracium
and Senecio (see Table 19).
In view of the high plant species number, the number of Auchenorrhyncha species
attacking them is rather small. The proportion of host specialists is moderate, and feed-
ers of mesophyll and xylem are slightly overrepresented (see section 5.4).
Specialists among Auchenorrhyncha appear to prefer members of the subfamily As-
teroideae, with Laburrus pellax (Horv.) living on goldilocks (Aster linosyris), both Eupt-
eryx tenella (Fall.) and Ophiola transversa (Fall.) on yarrow (Achillea millefolium), both
Macropsidius sahlbergi (Fl.) and Laburrus impictifrons (Boh.) on field southernwood (Arte-
misia campestris), and Austroasca vittata (Leth.), Eupteryx adspersa (H.-S.) and E. artemisiae
(Kbm.) all on species of wormwood (Artemisia spp.). Hephathus nanus (H.-S.) is proba-
bly associated with an unidentified species or genus, perhaps dwarf thistle (Cirsium
acaule). The data base of the subfamily Cichorioideae is rather small, but Neoaliturus
fenestratus (H.-S.) is probably associated with hawkbit (Leontodon spp.), and Euscelis
distinguendus (Kbm.) has been found – unlike its congeners – on Taraxacum and perhaps
280 Utilization of plant resources

Table 18. Auchenorrhyncha species utilizing Lamiaceae in Germany. Explanations see Table 5.

Galeopsis angust ifolia

Teucrium mont anum

Glechoma hederacea
Marrubium v ulgare
O cimum basilicum

Leonurus cardiaca
Prunella v ulgaris

St achy s palust ris

Lamium album
L. galeobdolon
Nepet a cat aria
Ajuga rept ans

L. maculat um
T. scorodonia

Ballot a nigra

S. sy lv at ica
Species
Eupteryx immaculatifrons (Kbm .) . . . . . . . . . . . X . . . .
Eupteryx lelievrei (Leth.) . . . . . . . . . . . . . . . .
Eupteryx origani Zachv. . . . . . . . . . . . . . . . .
Eupteryx collina (Fl.) . . . . . . . . . . . . . . . .
Chlorita pusilla Mats. . . . . . . . . . . . . . . . .
Chlorita dumosa (Rib.) . . . . . . . . . . . . . . . .
Goniagnathus brevis (H .-S.) . . . . . . . . . . . . . . . .
A gallia consobrina Curt. . . . x . . x . . . ? ? . . . ?
Eupteryx stachydearum (H ard y) . . . . . . . . . . X ? . . . X
Eupteryx curtisii (Fl.) . . . X . . . . . . . . . . . ?
Eupteryx florida Rib. ?. . . ? . ? x . . X . . x X x X
Eupteryx decemnotata R. . . . . . x . . . . . . . . . .
Eupteryx salviae Arz. & Vid . . . . . . . . . . . . . . . . .
Eupteryx thoulessi Ed w . . . . . . . . . . . . . . . . .
Eupteryx melissae Curt. . . . . ?* x . . . . . . x . . .
Eupteryx vittata (L.) . x . . . . X ?* . . . . . . . .
Eupteryx notata Curt. . . . . . . . ?* . . . . . . . .
Erythria aureola (Fall.) . . ?* . . . . . . . . . . . . .
Evacanthus acuminatus (F.) . . . . . . ? . . . . ? . . . ?
Evacanthus interruptus (L.) . . . . . . . . . . . ? . . . .
Emelyanoviana mollicula (Boh.) . . . x . . . . . . . ?* . ?* . .
Eupteryx atropunctata (Goeze) . . . . ?* ?* . . . ? . . . ? x .
Eupteryx aurata (L.) . . . . . ?* . . . . . X ?* . . .
Hyalesthes obsoletus Sign. . . . . . . . . x* . . . . . . .
Cercopis sanguinolenta (Scop.) . . . . . . . . . . . . . . . .
Cercopis vulnerata Rossi . . . . . . . . . . . . . . . .
Lepyronia coleoptrata (L.) . . . . . . . . . . . . . . . .
Philaenus spumarius (L.) . . . . . . . . . . . . . . . .
A naceratagallia austriaca W.Wg. . . . . . . . . . . . . . . . .
A naceratagallia frisia (W.Wg.) . . . . . . . . . . . . . . . .
A naceratagallia ribauti (Oss.) . . . . . . . . . . . . . . . .
A naceratagallia venosa (Geoffr.) . . . . . . . . . . . . . . . .
Planaphrodes bifasciata (L.) . . . . . . . . . . ? ? . . . .
Planaphrodes trifasciata (Geoffr.) . . . . . . . . . . . . . . . .
Empoasca decipiens Paoli . . . . . . . . . . . . . . . .
Erythria manderstjernii (Kbm .) . . . . . . . ?* . . . . . . . .
Eupteryx calcarata Oss. . . . . . . . . . . . . . ? . .
Hauptidia provincialis (Rib.) ? . . . . . . . . . . . . . . .
Lamprotettix nitidulus (F.) . . . . . . . . . . . ? . . . .
Total substantiated 0 1 0 3 0 2 3 0 1 1 1 2 2 1 2 2
Total unconfirm ed 12 0 1 1 2 3 1 3 0 1 2 7 1 3 0 3
* = after Halkka et al. (1977), Remane (pers. comm.), Stewart (1988), Vidano & Arzone (1978)
 Asteraceae 281

Table 18. (continued):

Clinopodium v ulgare

Overw intering stage

Bet onica officinalis

Ly copus europaeus
Melissa officinalis

O riganum v ulgare

Ment ha aquat ica

Salv ia nemorosa

Thy mus praecox

Th. pulegioides

Th. serpy llum

S. v ert icillat a

D iet breadth
M. longifolia

M. x piperit a
S. officinalis

Th. v ulgaris
S. prat ensis

Substrate
. . . . . . . . . . . . . . . . m1 M eg
X . . . . . . . . . . . . . . . m1 M eg 1st d e-
. . . . . . . X . . . . . . . . m1 M eg gree m ono-
. . ?* . . ?* . . . . . . . . X . m 1? M eg phages
. . . . . . . . . . X . . . . . m1 M eg
. . . . . . . . x ? x . . . . . m2 M eg 2nd d egree
. . . . . . . . x x ? . . . . . m 2? P ad m onophages
. . . . . . . . . . . . . . . . o1? P ad
. . . . . . ? . . . . . . ? . . o1 M eg
. . . . . . ? . . . . . . . . . o1? M eg 1st d e-
. . X . . X X ? . . . . . . x . o1 M eg gree oligo-
. . X . . ? . ? . x . x . . . . o1 M eg phages
. . ?
?* . . . . . . . . . . . . . o1? M eg
. . . . . . . . . . x X . ?* o1 M eg
. . X . . ?* . . . . . . . . . ?* o2 M eg
. . . . . . . . . . . . . . . . o2 M eg 2nd d egree
. . . . . . . . ?* ?* . . . . . . o2 M eg oligophages
. . . . . . . . ? ? . . . . . . o2 M eg
. . . . . . . . . . . . . . . . po X eg
? . . . . . . . . . . . . . . . po X eg
? x X ? ? x ? . x ?* . . ?* ? . po M eg Polyphages
x x x . x x ? X . . . . . x . x po M eg
. . x . . . . . . . . . . ?* x . po M eg
. . . . . . . . . . . . . . . . po P ny Vertical
. . . x* . . . . . . . . . . . . po X ny m igrants
. . . . . . . . . . . . . . . . po X ny
. . . . . . . . . . . . ?* . . . po X eg
. . . . . . . . . . . . . ? ? . po X eg
. . . . . . . . ? ? . . . . . . o2? P ad
. . . . . . . . ? ? . . . . . . o2? P ad
. . . . . . . . . ? . . . . . . o2? P ad
. . . . . . . . . ? . . . . . . o2? P eg Uncon-
. . . . . . . . . . . . . . . . o1? P eg firm ed
. . . . . . . . . ? ? . . . . . o2? P eg
. . . . . ?* . . . . . . . . . . po M? ad
. . . . . . . . . ? . . . . . . po? M ad
. . . . . . . . . . . . . . . . m 1? M eg
. . . . . . . . . . . . . . . . po? M ad ?
. . . . . . . . . . . . . . . . po P eg
2 1 6 2 1 2 2 2 2 3 2 1 1 2 3 1
2 0 2 0 1 5 3 2 4 9 3 0 1 4 2 2
282 Utilization of plant resources

Table 19. Auchenorrhyncha species utilizing Asteraceae in Germany. Explanations see Table 5.

Eupat orium cannabinum

Chry sant hemum v ulgare

Helichry sum arenarium

Art emisia abrot anum

Achillea millefolium
Heliant hus annuus

Dahlia pinnat a
Ast er linosy ris

Pet asit es spp.

A. absint hium
Solidago spp.

A. campest ris

spp.
A. marit ima

Senecio spp.
A. v ulgaris
Species
Laburrus pellax (H orv.) . . X . . . . . . . . . . . .
Eupteryx tenella (Fall.) . . . . . . X . . . . . . . .
Ophiola transversa (Fall.) . . . . . . x
x* . . . . . . . .
M acropsidius sahlbergi (Fl.) . . . . . . . . . . . . X . .
Laburrus impictifrons (Boh.) . . . . . . . . . . . . X . .
Eupteryx adspersa (H .-S.) . . . . . . . . . . X . . . .
A ustroasca vittata (Leth.) . . . . . . . . . . X x . . .
Eupteryx artemisiae (Kbm .) . . . . . . . . x . . X . . .
Euscelis venosus (Kbm .) . . . . . . . . . . . . . . .
N eoaliturus fenestratus (H .-S.) . . . ?* . . . . . . . . . . .
Chlorita paolii (Oss.) . . . . . . X . . x x . X . .
Euscelis distinguendus (Kbm .) . . . . . . . . . . . . . . .
A gallia brachyptera (Boh.) . . . . . . ?* . . . . . . . .
Eupteryx notata Curt. . . . . . . . . . . . . . . .
Eupteryx vittata (L.) . . . . . . . . . . . . . . .
Lepyronia coleoptrata (L.) x* x* . . . . . ? . x* . . . . .
Philaenus spumarius (L.) . x* . . . . x x . x . . . ? ?
A phrodes makarovi Zachv. . ?* . . . . . . . . . . . . .
Evacanthus interruptus (L.) x . . . . . . . . . . . . x x
Erythria manderstjernii (Kbm .) ? . . . . . . . . . . . . . ?
Empoasca decipiens Paoli . ? . . . . . ? . . ?* . . . .
Empoasca pteridis (Dhlb.) . ?* . . . . . . . . . . . . .
Empoasca vitis (Göthe) . . . . ? . . . . . . . . ? .
Eupteryx atropunctata (Goeze) . . . . ?* . . ?* . . . . . ?* .
Eupteryx aurata (L.) ?* . . . ? x . . . . . . . X X
Hyalesthes obsoletus Sign. . . . . . . . x* . x* . . . . x
Cercopis sanguinolenta (Scop.) . . . . . . . . . . . . . . .
Cercopis vulnerata Rossi . A . . . . . ? . ? . . . . .
Stictocephala bisonia K. & Y. . A . . . . . . . ?* . . . . .
Centrotus cornutus (L.) . . . . . . . . . . . . . . .
A phrophora alni (Fall.) . . . . . . . . . . . . . . .
Hephathus nanus (H .-S.) . . . . . . . . . . . . . . .
Eupteryx heydenii (Kbm .) . . . . . . . . . . . . . ?* .
Fieberiella septentrionalis W.Wg. . ? . . . . . . . . . . . . .
A thysanus quadrum Boh. . . . . . . . . . . . . . . .
Total substantiated 2 4 1 0 0 1 4 2 1 4 3 2 3 2 3
Total unconfirm ed 2 4 0 1 3 0 1 4 0 2 1 0 0 4 2
* = after Biedermann (1998a), Bopp (1997), Kuntze (1937), Nickel (1979), Remane & Fröhlich (1994b), Stewart
 Asteraceae 283

Table 19. (continued):

O nopordum acant hium

Overw intering stage

Hieracium pilosella
Leont odon hispidus
Lapsana communis

Picris hieracioides
Hy pochaeris spp.

Taraxacum spp.

Hieracium spp.
Carlina acaulis

Cirsium acaule
Carduus spp.
Arct ium spp.

D iet breadth
C. oleraceum

Crepis spp.
C. v ulgaris

C. arv ense

Substrate
. . . . . . . . . . . . . . . . m1 P eg
. . . . . . . . . . . . . . . . m1 M eg
. . . . . . . . . . . . . . . . m1 P eg 1st d egree
. . . . . . . . . . . . . . . . m 1? P eg m onophages
. . . . . . . . . . . . . . . . m1 P eg
. . . . . . . . . . . . . . . . m 1? M eg
. . . . . . . . . . . . . . . . m2 M eg
. . . . . . . . . . . . . . . . m2 M eg 2nd d egree
? ? . . . . . . . . . . . . . . m 2? P eg m onophages
. . . . . . . . . . ? . ? . . . m 2? P ad
. . . . . . . . . . . . . . . . o1 M eg
. . . . . . . . . . . ? ? . ?* . o1? P eg
. . . . . . . . . . . . x . . . o2? P eg Oligophages
. . . . . . . . . ?* x . . X . . o2 M eg
. . . . . . ? . . . . . . . . x o2 M eg
. . . . . ?* . . . . . . ?* . . . po X eg
. . ? . . X . . . . . . ? . . x* po X eg
. . . x . ? . . . . . . X . . . po P eg
. . x . . . x . . . . . . . . . po X eg
. . . . . . . . . . . . . . . ?* po? M ad Polyphages
. . . . . . . . . . . . . . . . po M? ad
. . . . . ? . . . . . . ? . . . po M? ad ?
. . . . . . . . . . . . . . . . po P ad
. . ?* . . ? . . . . . . . . . . po M eg
. . X . . . x . x . . . . . . . po M eg
. . . . . . . . . . . . . . . . po P ny
. . . . . . . x* . . . . . . . . po X ny
. . . A . . . . . . . . . . . . po X ny Vertical
. . . . . . . . . . . . . . . . po P eg m igrants
. . . A . ? . . . . . . . . . . po P ny
. . . . . . . . . . . . . . . . po X eg
. . . . ? . . . . . . . . . . . m 1? P eg
. . . . . . . . . . . . . . . . m 2? M eg Unconfirm ed
. . . . . . . . . . . . . . . . po P eg
. . . . . . . . . . . . . . . . m 1? P eg
0 0 2 3 0 1 2 1 1 0 1 0 2 1 0 2
1 1 2 0 1 5 1 0 0 1 1 1 5 0 1 1
(1988), Strübing (pers. comm.), Vidano (1959a), Vidano & Arzone (1978)
284 Utilization of plant resources

other Asteraceae. Among oligophages, Chlorita paolii (Oss.) is confined to both worm-
wood and yarrow. Eupteryx vittata (L.) feeds on marsh hawk’s-beard (Crepis paludosa);
E. notata Curt. feeds on mouse-ear hawkweed (Hieacium pilosella) and other low-grow-
ing Asteraceae. The two latter species, however, are also found on members of other
plant families. Generalists include broadly polyphagous species such as Philaenus spuarius
(L.), Aphrodes makarovi Zachv., Empoasca spp., Eupteryx aurata (L.) and E. atropunctata
(Goeze).
According to Zwölfer (1988), who reviewed the insect fauna on thistles (but without
treating Hemiptera), the guild size of phytophages is a function of the geographical
distance from the evolutionary centre of the host. This pattern is probably also found in
leafhoppers living on Artemisia, which shows centres of radiation in steppe and moun-
tain regions of central and east Asia and North America (e.g. Hultén 1968). Likewise,
the guild size at least within the associated leafhopper groups decreases from central
Asia towards western Europe, as was shown for the Eupteryx artemisiae (Kbm.) group
(Dworakowska 1970a). The same is true for the genus Laburrus, most species of which
occur in Mongolia and Kazakhstan (see Mityaev1971; Nast 1972; Tishechkin 2002b).

5.1.2.17 Juncaceae
The rush family comprises only two genera and 40 herbaceous species in Germany,
many of which are difficult to identify. Among true rushes (Juncus spp.), in particular,
the data base on specific level is weak. A number of species is annual or restricted to
special habitats, like alpine grassland, salt marshes and temporarily flooded mud banks.
Even some of the more widespread species have never been identified by entomolo-
gists collecting Auchenorrhyncha. Thus, the majority of data refers to only a few wide-
spread fenland species, like jointed, soft and hard rush (Juncus articulatus, J. effusus, J.
inflexus). Most species of woodrushes (Luzula spp.) grow in forests or alpine grassland
on rather acidic substrates. Only a few have been sampled for Auchenorrhyncha on
specific level, with only a handful of positive records.
The Auchenorrhyncha guild comprises 9 confirmed plus 12 unconfirmed species (Table
20). There seem to be no 1st degree monophages, but Conomelus anceps (Germ.), Florodel-
phax leptosoma (Fl.), Macrosteles horvathi (W.Wg.) and probably Conomelus lorifer Rib. are
2nd degree monophagous on rushes, whereas Cicadella viridis (L.) and Conosanus obsol-
etus (Kbm.) are both generalists, which are frequently found to exploit also grasses and
other plants. Muellerianella fairmairei (Perr.) is strictly bound to stands of Juncus effusus
for the deposition of winter eggs, but apparently, the nymphs migrate to stands of vel-
vetgrass (Holcus spp.) immediately after hatching, where they usually build up a sum-
mer generation (Drosopoulos 1977, and pers. comm.). Woodrushes were only in a few
occasions found to be attacked by generalists such as Criomorphus albomarginatus Curt.,
Macustus grisescens (Zett.) and Forcipata forcipata (Fl.).

5.1.2.18 Cyperaceae
The sedge family comprises 132 exclusively herbaceous, mainly perennial species in
Germany, thus constituting a considerable proportion of the central European phytodi-
 Cyperaceae 285

Table 20. Auchenorrhyncha species utilizing Juncaceae in Germany. Explanations see Table 5.

Overw intering stage

J. conglomerat us
Juncus bufonius

J. subnodulosus

D iet breadth
J. art iculat us
J. acut iflorus
Luz ula spp.
J. gerardii
J. inflexus

Substrate
J. effusus
Species
Conomelus anceps (Germ .) ?* X x X ? X m2 P eg
Conomelus lorifer Rib. x m 2? P eg Monophages
Florodelphax leptosoma (Fl.) ? x x x X ? m2 P ny
M acrosteles horvathi (W.Wg.) ? X X ? m2 P eg
Criomorphus albomarginatus Curt. x* o2 P ny.
M acustus grisescens (Zett.) x o2 P ny Oligophages
Conosanus obsoletus (Kbm .) x X ? X ? o2 P eg
Cicadella viridis (L.) X x* X ? X po X eg Polyphages
Forcipata forcipata (Fl.) x po M eg
M egamelodes quadrimaculatus (Sign.) ? ? ? o2 P ad
M egamelodes lequesnei W.Wg. m 2? P ad
Javesella pellucida (F.) ?* ? ?* po P ny
Javesella obscurella (Boh.) ?* ?* po? P ny
Javesella salina (H pt.) o1? P ny
Lepyronia coleoptrata (L.) po X eg Uncon-
N eophilaenus lineatus (L.) ?* ?* po X eg firm ed
M acrosteles laevis (Rib.) ? ?* po P eg
M acrosteles ossiannilssoni Ld b. ?* po? P eg
M acrosteles sexnotatus (Fall.) ? ?* po P eg
M acrosteles sordidipennis (Stål) ? m 1? P eg
Limotettix striola (Fall.) ? ?* o1? P eg
Total 0 2 3 3 5 1 4 0 3
* = after Halkka et al. (1977), Novotný (1995), Remane (pers. comm.), Witsack (1985)

versity. Cyperaceae biomass and dominance are particularly high in open, moist to flood-
ed sites, notably bogs, fens and fen woods, marshes and lake shores. They are also com-
mon, though less abundant, in damp or even dry situations, such as deciduous forests,
dry hillsides and sand dunes.
For several reasons, published information on associated phytophagous insects is
rather poor. Firstly, identification problems are considerable, particularly among non-
flowering or withered Carex. Secondly, assigning sampled insects to a particular sedge
in mixed stands of graminoids is rather difficult and can be only done by careful obser-
vation. Finally, flooded or boggy habitats, where most sedges grow, are often difficult to
access. For this study, more than 700 stands of more than 40 species have been searched
for Auchenorrhyncha, revealing numerous insect-host associations, many of which
turned out to be highly specific. On the other hand, at least 30 species are probably too
scarce to harbour any phytophages, except a few generalists. In general, the data base
can now be considered to be at least moderate in many lowland regions of Germany,
but there is still a number of Cyperaceae species, which have never been sampled sep-
arately for any insect group. This is particularly true for subalpine and alpine habitats,
286 Utilization of plant resources

and for bogs and fens. Also, many species have been sampled only in a limited geo-
graphic area, with no information on their insect guilds elsewhere.
Altogether, 74 Auchenorrhyncha species have been found to feed and reproduce,
plus another 12 unconfirmed species. Hence, the importance of the Cyperaceae for
Auchenorrhyncha is only surpassed by the grass family. The majority shows a rather
specific association with the host, with 30 species being 1st and 2nd degree mono-
phagous, respectively. The remaining 14 species are oligophagous or polyphagous.
Thus, the proportion of monophages on Cyperaceae is by far the highest among all
plant families. In contrast, the ratio of species numbers Auchenorrhyncha : plants is
relatively low due to the high diversity of the Cyperaceae. Concerning the type of
substrate and the overwintering stage, phloem feeding and overwintering as egg clearly
predominate.
Cottongrasses (Eriophorum spp.) are locally dominant on wet, usually peaty sites,
and play a major role as Auchenorrhyncha hosts. Hare’s-tail cottongrass (E. vaginatum)
is a typical plant of raised bogs, particularly on hummocks, but also in drained and
otherwise disturbed stages. It is attacked by Nothodelphax distincta (Fl.), Ommatidiotus
dissimilis (Fall.) and Sorhoanus xanthoneurus (Fieb.), all of which are probably strictly
monophagous. Common cottongrass (E. angustifolium) prefers minerotrophic bogs. Spe-
cific Auchenorrhyncha include Macrosteles fieberi (Edw.), Cosmotettix panzeri (Fl.) and
Delphacodes capnodes (Scott). The latter, however, may also be found in places without
Eriophorum, although these populations are apparently triploid females, and live on
undetermined sedges. Broad-leaved cottongrass (E. latifolium) has only been found to
be exploited by Kelisia vittipennis (J. Shlb.), which also breeds on the two former cotton-
grasses and perhaps certain sedges.
Furthermore, there are some species with less specific or unknown host relationships.
Thus, Javesella simillima (Lnv.) lives mainly on common cottongrass, but is also reported
from beaked sedge (Carex rostrata). Spittle masses of Neophilaenus lineatus (L.), which is
rather polyphagous, are also found on hare’s-tail cottongrass. Slender and white cot-
tongrass (E. gracile and E. scheuchzeri) occur sporadically in bogs, mainly in the Alps or
the north German plain, but have not been sampled for their Auchenorrhyncha.
Deergrass (Trichophorum cespitosum) is another dominant Cyperaceae species of
raised bogs, but without any Auchenorrhyncha specialists. Locally it is attacked by
the generalist Neophilaenus lineatus (L.). It is also the most likely host of bog popula-
tions of Limotettix striola (Fall.), and has been reported to be utilized by Sorhoanus
xanthoneurus (Fieb.) in the Czech Republic (Novotný 1995). Alpine deergrass (Trichopho-
rum alpinum), which is rather widespread in south Germany, but almost extinct in the
northern plains, has not been studied so far.
5 native species of club-rush (Schoenoplectus spp.) occur in Germany, three of which
are rather localized and have not been sampled. Only bulrush (Sch. lacustris) and grey
club-rush (Sch. tabernaemontani), which are considered to belong to a single superspe-
cies, but can be easily distinguished in the field, have recently been shown to harbour a
specific Auchenorrhyncha guild. Collecting, however, can only be done by careful search
above the basis of the stems, which is usually flooded. Coryphaelus gyllenhalii (Fall.),
Parapotes reticulatus (Horv.) and Calligypona reyi (Fieb.) have all been found to breed on
bulrush. A single specimen of P. reticulatus (Horv.) and reproducing populations of C. reyi
 Cyperaceae 287

(Fieb.) were also collected on grey club-rush. Further species known to attack Schoeno-
plectus only include Macrosteles sexnotatus (Fall.) and perhaps Limotettix striola (Fall.).
Spike-rush (Eleocharis palustris s.l.) includes several subspecies, which have not been
clearly distinguished in this study. It is found in nutrient-rich, at least temporarily flooded
sites. Auchenorrhyncha data exist for common and slender spike-rush (E. palustris s.str.
and E. uniglumis). Euconomelus lepidus (Boh.) and Macrosteles lividus (Edw.) are specific
probably to both of them, whereas Limotettix striola (Fall.) also feeds on other species of
Cyperaceae, perhaps Trichophorum and Schoenoplectus.
Sea club-rush (Bolboschoenus maritimus) is infested by Paramesus obtusifrons (Stål) along
the coast, and by P. major Hpt. in inland salt marshes. Both are 1st degree monophagous,
but may in fact belong to a single species. Wood club-rush (Scirpus sylvaticus) is prefer-
ably utilized by Cicadula albingensis W.Wg., which probably also feeds on Carex. A single
stand of creeping club-rush (Sc. radicans) in Saxony has been found to harbour a small
population of the polyphagous Macrosteles sexnotatus (Fall.). Among Rhynchospora, only
white beak-sedge (Rh. alba) has been sampled in a few cases, revealing a large breeding
population of Macrosteles ossiannilssoni Ldb. and Limotettix atricapillus (Boh.), respec-
tively. The latter is perhaps strictly monophagous. Brown and black bog-rush (Schoenus
ferrugineus and Sch. nigricans) are perhaps the favoured host of Stiromella obliqua (W.Wg.).
Apart from Carex, none of the remaining genera of Cyperaceae have been studied so
far, but are perhaps not suitable as hosts, since most central European species are either
extremely rare or annuals. These genera include Cyperus, Isolepis, Scirpoides (= Holoschoe-
nus), Blysmus, Cladium and Kobresia. However, at least flat-sedge (Blysmus compressus ),
great fen-sedge (Cladium mariscus) and Bellard’s kobresia (Kobresia myosuroides) locally
occur in larger stands and should be checked.
Carex is among the most diverse plant genera all over the world, with more than 2000
species, dating back at least until the lower Oligocene (Mai 1995). Even after lumping
most subspecies, the species number in Germany is at least 93, with Auchenorrhyncha
data existing for 30 of them. In the following sections 4 groups will be distinguished:
Carex species with (i) verified and (ii) probable 1st degree monophages, species with
(iii) verified and (iv) probably 2nd degree monophages.
(i) Carex species with substantiated 1st degree monophages
Quaking sedge (Carex brizoides) is the only host of Cicadula rubroflava Lnv., and also
the main host of Kelisia praecox Hpt. The latter, however, is also found in places where
this plant is absent. Kelisia sabulicola W.Wg. is strictly associated with sand sedge
(C. arenaria). Florodelphax paryphasma (Fl.) has been found to live exclusively on brown
sedge (C. disticha), whereas Stenocranus longipennis (Curt.) is specific to greater tussock
sedge (C. paniculata). Glaucous sedge (C. flacca) and dwarf sedge (C. humilis) both har-
bour the highest numbers of specialists among central European Carex. The former is
exploited by Kelisia irregulata Hpt., K. guttula (Germ.) and Wagneriala sinuata (Then), all
of which are 1st degree monophagous, and Anakelisia perspicillata (Boh.), which addi-
tionally lives on pill sedge (C. pilulifera). Dwarf sedge is the exclusive host of the recent-
ly split group of Kelisia haupti W.Wg., including the nominate form, K. halpina Rem. &
Jung and K. hagemini Rem. & Jung., whereas Wagneriala minima (J. Shlb.) is also likely to
utilize white-flowered sedge (C. alba). Kelisia pallidula (Boh.) is exclusively associated
with C. panicea. Yellow sedge (C. flava) has been found to be the only host of Kelisia sima
288 Utilization of plant resources

Table 21. Auchenorrhyncha species utilizing Cyperaceae in Germany. Explanations see Table 5.

Trichophorum cespit osum

Bolboschoenus marit imus

Schoenoplect us lacust ris
Eriophorum v aginat um

Sch. t abernaemont ani

Schoenus ferrugineus
Eleocharis palust ris

Rhy nchospora alba

Scirpus sy lv at icus
E. angust ifolium

Carex muricat a
Sch. nigricans

C. paniculat a
E. lat ifolium

E. uniglumis

C. briz oides

C. elongat a
C. arenaria
S. radicans

C. dist icha
C. v ulpina

C. remot a
Species
N othodelphax distincta (Fl.) X . . . . . . . . . . . . . . . . . . . . .
Ommatidiotus dissimilis (Fall.) X . . . . . . . . . . . . . . . . . . . . .
Sorhoanus xanthoneurus (Fieb.) X . . ?* . . . . . . . . . . . . . . . . . .
Kelisia vittipennis (J. Shlb.) X X x . . . . . . . . . . . . . . . . . . .
M acrosteles fieberi (Ed w .) . X . . . . . . . . . . . . . . . . . . . .
Cosmotettix panzeri (Fl.) ?* X . . . . . . . . . . . . . . . . . . . .
Delphacodes capnodes (Scott) . X . . . . . . . . . . . . . . . . . . . .
Javesella simillima (Lnv.) . x . . . . . . . . . . . . . . . . . . . .
Coryphaelus gyllenhalii (Fall.) . . . . X . . . . . . . . . . . . . . . . .
Parapotes reticulatus (H orv.) . . . . X ? . . . . . . . . . . . . . . . .
Calligypona reyi (Fieb.) . . . . X x.
x . . . . . . . . . . . . . . . .
Paramesus major H pt. . . . . . . X . . . . . . . . . . . . . . .
Paramesus obtusifrons (Stål) . . . . . . X . . . . . . . . . . . . . . .
Cicadula albingensis W.Wg. . . . . . . . X . . . . . . . . . . . . . .
Euconomelus lepidus (Boh.) . . . . . . . . . x X . . . . . . . . . . .
M acrosteles lividus (Ed w .) . . . . . . . . . X ? . . . . . . . . . . .
Limotettix striola (Fall.) . . . ? . ? . . . x X . . . . . . . . . . .
Limotettix atricapillus (Boh.) . . . . . . . . . . . . . x . . . . . . . .
Kelisia monoceros Rib. . . . . . . . . . . . . . . x? X . . . . . .
Cicadula rubroflava Lnv. . . . . . . . . . . . . . . . . X . . . . .
Kelisia praecox H pt. . . . . . . . . . . . . . . . . X . . . . .
Kelisia sabulicola W.Wg. . . . . . . . . . . . . . . . . . X . . . .
Florodelphax paryphasma (Fl.) . . . . . . . . . . . . . . . . . . X . . .
Stenocranus longipennis (Curt.) . . . . . . . . . . . . . . . . . . . X . .
Oncodelphax pullula (Boh.) . . . . . . . . . . . . . . . . . . . . . .
Cicadula saturata (Ed w .) . . . . . . . . . . . . . . . . . . . . . .
Cicadella lasiocarpae Oss. . . . . . . . . . . . . . . . . . . . . . .
Cosmotettix costalis (Fall.) . . . . . . . . . . . . . . . . . . . . . .
M etalimnus formosus (Boh.) . . . . . . . . . . . . . . . . . . . . . .
Cicadula flori (J. Shlb.) . . . . . . . . . . . . . . . . . . ? . . .
Kelisia guttula (Germ .) . . . . . . . . . . . . . . . . . . . . . .
Kelisia irregulata H pt. . . . . . . . . . . . . . . . . . . . . . .
W agneriala sinuata (Then) . . . . . . . . . . . . . . . . . . . . . .
A nakelisia perspicillata (Boh.) . . . . . . . . . . . . . . . . . . . . . .
Kelisia haupti W.Wg. . . . . . . . . . . . . . . . . . . . . . .
Kelisia hagemini Rem . & Jung . . . . . . . . . . . . . . . . . . . . . .
Kelisia halpina Rem . & Jung . . . . . . . . . . . . . . . . . . . . . .
W agneriala minima (J. Shlb.) . . . . . . . . . . . . . . . . . . . . . .
Kelisia pallidula (Boh.) . . . . . . . . . . . . . . . . . . . . . .
Sorhoanus assimilis (Fall.) . . . . . . . . . . . . . . . . . . . . . .
Kelisia guttulifera (Kbm .) . . . . . . . . . . . . . . . ? . . . ? x ?
Kelisia minima Rib. . . . . . . . . . . . . . . . . . . . . . .
 Cyperaceae 289

Table 21. (continued):

Overw intering stage

C. pseudocy perus
C. semperv irens
C. cary ophy llea

C. acut iformis

D iet breadth
C. ferruginea

C. sy lv at ica
C. pilulifera

C. v esicaria
C. mont ana

C. rost rat a
C. panicea
C. humilis

C. dist ans

Substrate
C. riparia
C. limosa
C. pilosa
C. flacca
C. acut a

C. hirt a
C. nigra

C. flav a
C. elat a

C. alba

. . . . . . . . . . . . . . . . . . . . . . . m1 P ny
. . . . . . . . . . . . . . . . . . . . . . . m 1? P eg
. . . . . . . . . . . . . . . . . . . . . . . m 1? P eg
. . . . . . . . . . . . . . . . . . ? . . . . m2 P eg
. . . . . . . . . . . . . . . . . . . . . . . m1 P eg
. . . . . . . . . . . . . . . . . . . . . . . m 1? P eg
. ?* . . . . . . . . . . . . . . . . . . ?* ?* . o1 P ad
. . . . . . . . . . . . . . . . . . ?* . . . . o1? P ny
. . . . . . . . . . . . . . . . . . . . . . . m1 P eg
. . . . . . . . . . . . . . . . . . . . . . . m 1? P eg
. . . . . . . . . . . . . . . . . . . . . . . m2 P ny
. . . . . . . . . . . . . . . . . . . . . . . m1 P eg
. . . . . . . . . . . . . . . . . . . . . . . m1 P eg
? ? . . . . . . . . . . . . . . . . . . ? ?* . o1? P eg
. . . . . . . . . . . . . . . . . . . . . . . m 2? P eg
. . . . . . . . . . . . . . . . . . . . . . . m 2? P eg
. . . . . . . . . . . . . . . . . . . . . . . o1? P eg
. . . . . . . . . . . . . . . . . . . . . . . m 1? P eg
. . . . . . . . . . . . . ? . . . . . . . . . m2 P eg
. . . . . . . . . . . . . . . . . . . . . . . m1 P eg N arrow -
. . . . . ? . . . . . . . . . . . . . . . . . m2 P ad ly steno-
. . . . . . . . . . . . . . . . . . . . . . . m1 P ad phagous
. . . . . . . . . . . . . . . . . . . . . . . m1 P ny
. . . . . . . . . . . . . . . . . . . . . . . m1 P eg
X ? ? . . . . . . . . . . . . . . . ? . ? . . m 2? P ny
X . . . . . . . . . . . . . . . . . ? . . . . m 2? P eg
x . . . . . . . . . . . . . . . . . . . . . . m 2? X eg
? X ? . . . . . . . . . . . . . . . ?* . . ?* . m 2? P eg
?* X x . . . . . . . . . . . . . . . . . . . . m2 P eg
?* X ? . . . . . . . . . . . . . . . . . ? . . m 2? P eg
. . . . . X . . . . . . . . . . . . . . . . . m1 P eg
. . . . . X . . . . . . . . . . . . . . . . . m1 P eg
. . . . . x . . . . . . . . . . . . . . . . . m1 M eg?
. . . x . X . . . . . . . . . . . . . . . . . m2 P eg
. . . . . . . X . . . . . . . . . . . . . . . m1 P eg
. . . . . . . X . . . . . . . . . . . . . . . m1 P eg
. . . . . . . X . . . . . . . . . . . . . . . m1 P eg
. . . . . . . X ? . . . . . . . . . . . . . . m 2? M eg?
. . . . . . . . . . . X . . . . . . . . . . . m1 P eg
? . . . . . . . . . . ? . . . . . . ? . . . . m 2? P eg
. . . . . . . . . . . . . . X ? . . . . . . . m2 P eg
. . . . . . . . . . . . . . . x . . . . . . . m 1? P eg
290 Utilization of plant resources

Table 21. (continued):

Trichophorum cespit osum

Bolboschoenus marit imus

Schoenoplect us lacust ris
Eriophorum v aginat um

Sch. t abernaemont ani

Schoenus ferrugineus
Eleocharis palust ris

Rhy nchospora alba

Scirpus sy lv at icus
E. angust ifolium

Carex muricat a
Sch. nigricans

C. paniculat a
E. lat ifolium

E. uniglumis

C. briz oides

C. elongat a
C. arenaria
S. radicans

C. dist icha
C. v ulpina

C. remot a
Species
Kelisia sima Rib. . . . . . . . . . . . . . . . . . . . . . .
Kelisia punctulum (Kbm .) . . . . . . . . . . . . . . . . . . . . . .
Cicadula frontalis (H .-S.) . . . . . . . . . . . . . . . . . . . . . .
A nakelisia fasciata (Kbm .) . . . . . . . . . . . . . . . . . . . . . .
Cosmotettix caudatus (Fl.) . . . . . . . . . . . . . . . . . . . . . .
M etalimnus steini (Fieb.) . . . . . . . . . . . . . . . . . . . . . .
Stroggylocephalus agrestis (Fall.) . . . . . . ? . . . . . . . . . . . ? . . .
Stenocranus fuscovittatus (Stål) . . . . . . . . . . . . . . . . . . . x . .
M egamelus notula (Germ .) . . . . . . . . . . . . . . . . . . X . . .
N otus flavipennis (Zett.) . . . . . . ? . . . . . . . . . ?* . X x . .
Cicadula quadrinotata (F.) . ?* . . . . ? . . . . . . . . x x ? X x . .
Kelisia nervosa Vilb. . . . . . . . . . . . . . . . . . . . . . .
Kelisia confusa Lnv. . . . . . . . . . . . . . . . . . . . . . .
Kelisia ribauti W.Wg. . ? ? . . . . . . . . . . . . . . . . . . ?*
N othodelphax albocarinata (Stål) . . . . . . . . . . . . . . . . . . . . . .
Paradelphacodes paludosa (Fl.) . . . . . . . . . . . . . . . . . . . . . .
Stroggylocephalus livens (Zett.) . ? . . . . . . . . . . . . . . . . . . . .
W agneriala incisa (Then) . . . . . . . . . . . . . . ?* . . . . . . .
Forcipata citrinella (Zett.) . . . . . . . . . . . . . . . . . . . . . .
M acrosteles alpinus (Zett.) . . . . . . . . . . . . . . . . . . . . . .
Cicadula quinquenotata (Boh.) . ? ? . . . . . . . . . . . . . . . . . . .
Cicadula ornata (Mel.) . . . . . . . . . . . . . . . . . . . . . .
Cosmotettix aurantiacus (Forel) . . . . . . . . . . . . . . . . . . . . . .
M acrosteles ossiannilssoni Ld b. . . . . . . . . . . . . . x . . . . . . . .
M acustus grisescens (Zett.) . . . . . . . . . . . . . . . . ?* . . x . .
Cicadella viridis (L.) . . . . . . . . . . . . . . . . . . . x . .
N eophilaenus lineatus (L.) x . . x . ?* . . . ?* . . . . . . . . . . . .
Forcipata forcipata (Fl.) . . . . . . . . . . . . . . . . ? . . . . .
Speudotettix subfusculus (Fall.) . . . . . . . . . . . . . . . . . . . . . .
Lepyronia coleoptrata (L.) . . . . . ?* . . . ?* . . . . . . . . . . . .
Philaenus spumarius (L.) . . . . . ?* . . . . . . . . . . . . . . . .
M acrosteles sexnotatus (Fall.) . . . . x . . . ? x . . . . . . . . . . . .
Stiromella obliqua (W.Wg.) . . . . . . . . . . . ?* ? . . . . . . . . .
M egamelodes lequesnei W.Wg. . . . . . . . . . . . . . . . . . . . . . .
Javesella discolor (Boh.) . . . . . . . . . . . . . . . . ?* . . . . .
Javesella pellucida (F.) . . . . . . . . . . . . . . . . . . . . . .
Javesella obscurella (Boh.) . . . . . . . . . . . . . . . . . . . . . .
A phrodes diminuta Rib. . . . . . . . . . . . . . . . . . . . . . .
M acrosteles cristatus (Rib.) . . . . . . . . . . . . . . . . . . . . . .
M acrosteles laevis (Rib.) . . . . ? . . . . ? . . . . . . . . . . . .
 Cyperaceae 291

Table 21. (continued):

Overw intering stage

C. pseudocy perus
C. semperv irens
C. cary ophy llea

C. acut iformis

D iet breadth
C. ferruginea

C. sy lv at ica
C. pilulifera

C. v esicaria
C. mont ana

C. rost rat a
C. panicea
C. humilis

C. dist ans

Substrate
C. riparia
C. limosa
C. pilosa
C. flacca
C. acut a

C. hirt a
C. nigra

C. flav a
C. elat a

C. alba

. . . . . . . . . . . . . . . . X . . . . . . m1 P eg
. . . . . . . . . . . . . . . . . . . . X . . m 1? P eg N arrow -
. ? . . . . . . . . . . . . . . . ? . . X X . m2 P eg ly steno-
. ? . . . . . . . . . . . . . . . . . . ? X . m 2? P eg phagous
. . . . . . . . . . . . . . . . . . . . . . X m1 P eg
. . . . . . . . . . . . . . . . . . . . . . x m 1? P eg?
x X ? . . . . . . . . . . . . . . x . . X . . m 2? P eg
. ? x . . . . . . . . x . . . . . . . . ? . . m2 P ad Broad ly
x X x . . . . . . . . . . . . . . ? X x X ? . m2 P eg steno-
x X x . . . . . . . . . . . . . . ? ?* X X ? . o1 M eg phagous
X ? ? . . . . . . . . . . . . . . . X x X ? X m2 P eg
. . ? . . . . . . . . . . . . . . . . . . . . m 1? P eg?
. . . . . . . . . . . . . . . . . . . . ? ?* . m 2? P eg
? . . . . . . . . . . . . . . . . . ?* . . . . m 2? P eg
. . . . . . . . . . ? . . . . . . . . . . . . m 1? P ny
. . . . . . . . . . . ? . . . . . . ? . . . . m 2? P ny on un-
. . . . . . . . . . . . . . . . . . ?* . . . . m 2? P ad know n
. . . . ? . ?* . . . . . . . . . . . . . . . . m 2? Carex
M eg? Cypera-
? . . . . ? . . . . . . . . . . ? . . . . . . m 2? M eg ceae
?* ? . . . . . . . . . . . . . . . . . . ? . . m 2? P eg
. . . . . . . . . . . . . . . . . . . . . . . m 2? P eg
. . . . . . . . . . . . . . . . . . . . . . . m 2? P eg
. . . . . . . . . . . . . . . . . . . . . . . m 1? P eg
. . . . . . . . . . . . . . . . . . . . . . . po? P eg
? ? . . . . . . . . . . . . . . . . ? . x . . o2 P ny
x ? x . . . . . . . . . . . ? . . . x . ? x* x* po X eg
?* . x* . . . . . . . . . . . . . . . ?* . . . . po X eg Oligo-/
. . . . ? ? . x . . . . ? ? ? . . . . . ? . . po? M eg poly-
. . . . . . . . . x . . . . . . . . . . . . . po P ny phagous
?* . . . . . . . . . . . . . . . . . . . . . . po X eg
. . . . . . . . . . . . . . . . . . . . . . . po X eg
. . . . . . . . . . . . . . . . . . . . . . . po P eg
. . . . . . . . . . . . . . . . . . . . . . . m 2? P ny
. . . . . . . . . . . . . . . . . . . . . . . m 2? P ad
. . . . . . . . . . . . . . . . . . . . . . . po? P ny Uncon-
. . . . . . . . . . . . . . . . . . . . . . . po? P ny Uncon-
firm ed
. . . . . . . . . . . . . . . . . . . . . . . po? P ny firmed
. . . . . . . . . . . . . . . . . . . . . . . o1? P eg
. . . . . . . . . . . . . . . . . . . . . . . po P eg
. . . . . . . . . . . . . . . . . . . . . . . po P eg
292 Utilization of plant resources

Table 21. (continued):

Trichophorum cespit osum

Bolboschoenus marit imus

Schoenoplect us lacust ris
Eriophorum v aginat um

Sch. t abernaemont ani

Schoenus ferrugineus
Eleocharis palust ris

Rhy nchospora alba

Scirpus sy lv at icus
E. angust ifolium

Carex muricat a
Sch. nigricans

C. paniculat a
E. lat ifolium

E. uniglumis

C. briz oides

C. elongat a
C. arenaria
S. radicans

C. dist icha
C. v ulpina

C. remot a
Species
M acrosteles oshanini Razv. . . . . . . . . . . . . . . . . . . . . . .
M acrosteles viridigriseus (Ed w .) . . . . . . . . . ? . . . . . . . . . . . .
Deltocephalus maculiceps Boh. ? ? . . . . . . . . . . . . . . . . . . . .
Sorhoanus schmidti (W.Wg.) . . . . . . . . . . . . . . . . . . . . . .
Total substantiated 5 5 1 1 4 1 2 1 0 4 2 0 0 2 0 2 3 1 4 6 1 0
Total unconfirm ed 2 4 2 2 1 5 3 0 1 4 1 1 1 0 1 1 4 1 2 1 0 2
Substantiated plus unconfirm ed 7 9 3 3 5 6 5 1 1 8 3 1 1 2 1 3 7 2 6 7 1 2
* =after Drosopoulos et al. (1983), Halkka et al. (1977), Lauterer (1980, 1986), Leising (1977), Moosbrugger (1946),

Rib., shortly after Remane & Jung (1995) had clarified the taxonomic status. Finally,
Metalimnus steini (Fieb.) and Cosmotettix caudatus (Fl.) are strictly monophagous on hairy
sedge (C. hirta), although the former is taxonomically not clear.
(ii) Carex species probably with 1st degree monophages
There is another group of species with evidence for holding 1st degree monophages,
but this is based on insufficient data. This group includes common sedge (C. nigra),
soft-leaved sedge (C. montana), mud sedge (C. limosa), carnation sedge (C. panicea), dis-
tant sedge (C. distans), beaked sedge (C. rostrata), lesser pond sedge (C. acutiformis) and
great pond sedge (C. riparia). C. nigra is perhaps the most-favoured among central Eu-
ropean Cyperaceae, with 8 definitely and 11 possibly associated Auchenorrhyncha spe-
cies. Oncodelphax pullula (Boh.), Cicadella lasiocarpae Oss. and Cicadula saturata (Edw.)
show a clear preference and are perhaps strictly monophagous. Tufted sedge (C. elata)
was found to be infested by large numbers of K. nervosa Vilb. in one case, but further
sampling in a larger geographic area is necessary to confirm this relationship. It is also
one of the hosts of Metalimnus formosus (Boh.), and possibly a host of Cicadula flori (J.
Shlb.), Cosmotettix costalis (Fall.) and Oncodelphax pullula (Boh.), thus perhaps sharing
some feeders with Carex acuta and C. nigra. C. montana is the most likely host of Wagne-
riala incisa (Then), while C. limosa has been found to be infested by Nothodelphax albocar-
inata (Stål) in two cases. C. distans has in a few occasions been found to harbour popula-
tions of the rare Kelisia minima Rib. For C. rostrata only three rather euryphagous sedge-
feeding species are verified to reproduce, but there is evidence for further 13. Among
these, both Paradelphacodes paludosa (Fl.) and Sorhoanus assimilis (Fall.) are perhaps more
specific. C. acutiformis altogether harbours at least 7 confirmed plus 10 unconfirmed
species, but only Kelisia punctulum (Kbm.) is probably specific, whereas Cicadula fronta-
lis (H.-S.) also exploits the following sedge. C. riparia is the only verified host of Anakeli-
sia fasciata (Kbm.), although a few specimens have been taken on C. acuta and
 Cyperaceae 293

Table 21. (continued):

Overw intering stage

C. pseudocy perus
C. semperv irens
C. cary ophy llea

C. acut iformis

D iet breadth
C. ferruginea

C. sy lv at ica
C. pilulifera

C. v esicaria
C. mont ana

C. rost rat a
C. panicea
C. humilis

C. dist ans

Substrate
C. riparia
C. limosa
C. pilosa
C. flacca
C. acut a

C. hirt a
C. nigra

C. flav a
C. elat a

C. alba

. . . . . . . . . . . . . . . . . . . . . . . ? P eg Uncon-
. . . . . . . . . . . . . . . . . . . . . . . o2? P eg firm ed
Uncon-
. . . . . . . . . . . . . . . . . . . . . . . m 1? P eg firmed
. . . . . . . . . . . . . . . . . . . . . . . m 1? P eg
8 6 6 1 0 4 0 5 0 1 0 2 0 0 1 1 1 1 3 3 7 3 4
11 10 6 0 2 3 1 0 1 0 1 2 1 2 2 1 1 3 12 0 10 7 0
19 16 12 1 2 7 1 5 1 1 1 4 1 2 3 2 2 4 15 3 17 10 4
Nickel (1979), Novotný (1995), Schiemenz (1987), Vidano (1965), Wagner & Franz (1961), Witsack (1985)

C. acutiformis (Lauterer 1984). In Greece C. riparia is reported to be infested by Kelisia

confusa Lnv. (Drosopoulos et al. 1983).
(iii) Carex species with substantiated 2nd degree monophages
Further sedges are frequently attacked, but without showing exclusive associations.
False fox sedge (C. vulpina ssp. otrubae) and prickly sedge (C. muricata) are the main
hosts of Kelisia monoceros Rib. in the lowlands, but not in the Alps, where perhaps ever-
green sedge (C. sempervirens) is preferred. Kelisia guttulifera (Kbm.) usually lives on re-
mote and wood Sedge (C. remota and C. sylvatica), but on single occasions, reproducing
populations have also been found on elongated sedge (C. elongata), false fox sedge
(C. vulpina ssp. otrubae), greater tussock sedge (C. paniculata) and distant sedge (C. distans).
Cicadula flori (J. Shlb.), Metalimnus formosus (Boh.) and Cosmotettix costalis (Fall.) all favour
slender tufted sedge (C. acuta) in many lowland floodplain sites, along with some less
specific species, such as Megamelus notula (Germ.), Stroggylocephalus agrestis (Fall.), No-
tus flavipennis (Zett.) and Cicadula quadrinotata (F.). On acidic substrates Anakelisia per-
spicillata (Boh.) lives on pill sedge (C. pilulifera), but in more basic sites it is clearly asso-
ciated with glaucous sedge (C. flacca). Other infested, but little studied sedges include
cyperus and bladder sedge (C. pseudocyperus, C. vesicaria).
(iv) Carex species probably with 2nd degree monophages
There is also some field evidence, that spring sedge (C. caryophyllea), white-flowered
sedge (C. alba), rusty sedge (C. ferruginea), ciliated sedge (C. pilosa) and evergreen sedge
(C. sempervirens) may be attacked, but more samples are required for confirmation.
Furthermore, it should be noted, that in some Auchenorrhyncha species, the host
relationships of which are little known, an association with unidentified species of Car-
ex is highly likely. This is true in Kelisia confusa Lnv., K. ribauti W.Wg., Stroggylocephalus
livens (Zett.), Forcipata citrinella (Zett.), Cicadula quinquenotata (Boh.), C. ornata (Mel.),
294 Utilization of plant resources

Cosmotettix aurantiacus (For.) and perhaps Sorhoanus schmidti (W. Wg.), Macrosteles osha-
nini Razv. and a few more.

5.1.2.19 Poaceae
Although central Europe is essentially a woodland country, the grasses are by far the
most-favoured Auchenorrhyncha food plant family. Along with the Cyperaceae, they
were certainly the main ecological platform that facilitated the dramatic radiation of
the Delphacidae, Paralimnini and probably other taxa, which is likely to have occurred
not later than the Miocene (see Wilson et al. 1994; Jacobs et al. 1999). The number of
grass species in Germany exceeds 220, surpassed only by the Asteraceae with more
than double as many plant species, but only attacked by 30 Auchenorrhyncha species.
Most of these grasses are considered to belong to the subfamily Pooideae (with few
exceptions, e.g. Eragrostis, Digitaria, Setaria) and utilize the C3 pathway (except Cynodon
dactylon, and a handful of others) (Renvoize & Clayton 1992).
Regarding biomass, grasses are certainly the most important plant family in central
Europe today. But before man reduced the forests, grasslands or at least grassy patches
must have been confined to extreme sites, which were either too wet, too dry, saline, or
disturbed by grazing herbivores, storms or fire. At that time it was probably the Fagace-
ae, and Fagus and Quercus in particular, which dominated most terrestrial ecosystems.
In the undergrowth of central European primeval forests grasses may have been more
dominant than in most man-made monocultures of today (Körber-Grohne 1990), thus
offering more potential food plants for Auchenorrhyncha. Moreover, it is now broadly
discussed that megaherbivores, most of which are extinct today, played a role in creat-
ing and maintaining at least small patches of open grassland which in turn facilitated
the occurrence of heliophilous species (Bengtsson et al. 2000; Vera 2000). Livestock kept
from the Neolithic onwards initially did not reduce forest cover significantly, but at
least since the iron age, man was able to cut herbaceous plants with sickles, thus main-
taining open grassland, which was colonized by grasses and forbs. Selection pressure
by frequent cutting and grazing gave rise to meadow and pasture plant communities,
which in turn were colonized by further organisms, including phytophagous insects
(see Ellenberg 1996).
Natural habitats, where most meadow and pasture grasses of today (and probably
their phytophages) occurred before human impact, mainly include rocky hillsides, bogs
and fens, fen woods, lake shores, riverine gravel, mud banks and saltmarshes. Although
most grasses of central Europe are probably native, some species are suspected to have
invaded with the rise of anthropogenic mowing and grazing, notably Alopecurus praten-
sis, Arrhenatherum elatius, Phleum pratense and Cynosurus cristatus (Körber-Grohne 1990).
Altogether, grasses are utilized by at least 210 Auchenorrhyncha species, which is
roughly a third of the German species total, and the discovery of further feeding rela-
tionships is likely. 98 species, i.e. almost 50%, are 1st degree monophagous, further 20
species are 2nd degree monphagous, and 68 species are 1st degree oligophagous. Thus
89% are specific to grasses. This high degree of host specialization is only rivalled by
the Cyperaceae and Salicaceae (see sections 5.1.2.9, 5.1.2.18). An overview of Auchenor-
rhyncha species on grasses in Germany is given in Table 22.
 Poaceae 295

In the following sections, the guilds of the more important grasses are described and
discussed in order of their importance as host plants. Specific host records are given in
the appendix. Information on grass species and their habitat requirements, distribution
and response to management was taken from Conert (1998), Oberdorfer (2001) and
Petersen (1992).
(i) Fescue (Festuca)
Regarding Auchenorrhyncha species numbers, Festuca is the most important grass
genus in central Europe, although only the lowland species have been sampled specif-
ically. There is some evidence for alpine species (e.g. Festuca puccinellii, F. quadriflora,
F. pulchella, F. rupicaprina) harbouring little known leafhoppers such as Psammotettix
nardeti Rem. and Ebarrius cognatus (Fieb.), but this still requires confirmation. Altogeth-
er, 43 Auchenorrhyncha species are confirmed feeders on Festuca, with a high propor-
tion of host specialists, and another 32 species are unconfirmed. Therefore, in view of
the insufficient data base on this grass genus, the total species number may even reach
the maximum value of Carex (see chapter 5.1.2.18).
Sheep’s fescue (Festuca ovina) s.l. (including the grey-leaved F. valesiaca group) is a
dominant grass of dry low-productivity grassland of all kinds, notably heaths, pas-
tures, dunes, dry hillsides, etc. Its taxonomy is difficult, and more than 20 subspecies
and varieties are known from Germany alone. These have not been distinguished for
this study, but at least some Auchenorrhyncha apparently prefer the grey-leaved sub-
species. Literature records on feeding are scarce, and many merely refer to plant genus
level or are even erroneous. Thus it has long been overlooked, that Festuca ovina shows
one of the highest Auchenorrhyncha species numbers all over central Europe, with 27
confirmed and 18 unconfirmed feeders in Germany alone, and future sampling may
well reveal an even higher diversity. This value is surpassed so far only by Quercus robur
and Betula pendula (see Fig. 14). 12 species are classified here as strictly monophagous
on F. ovina, notably Jassidaeus lugubris (Sign.), Metropis latifrons (Kbm.), M. inermis W.Wg.,
Kosswigianella exigua (Boh.), Neophilaenus infumatus (Hpt.), Doratura exilis Horv., Rhyti-
stylus proceps (Kbm.), Hardya signifer (Then), Rhopalopyx vitripennis (Fl.), Mocydiopsis lon-
gicauda Rem., Psammotettix pallidinervis (Dhlb.), Mendrausus pauxillus (Fieb.) and per-
haps Achorotile albosignata (Dhlb.). Further species are 2nd degree monophagous and
additionally feed on red fescue (Festuca rubra), which replaces F. ovina on more damp
and wet sites. These include Delphacinus mesomelas (Boh.), Acanthodelphax spinosa (Fieb.),
Rhopalopyx adumbrata (C. Shlb.) and Mocydiopsis attenuata (Germ.), probably also Eu-
pelix cuspidata (F.) and Arocephalus punctum (Fl.). Dicranotropis divergens Kbm. is appar-
ently the only strict monophage on Festuca rubra. Both Festuca ovina and F. rubra are also
important hosts for a number of oligophages, including Criomorphus albomarginatus Curt.,
Neophilaenus campestris (Fall.), N. minor (Kbm.), Dikraneura variata Hardy, Streptanus
marginatus (Kbm.) and others. Increased fertilizing of grassland and cessation of sheep
grazing during recent decades have led to a decrease of these two grasses and their
complete Auchenorrhyncha guilds.
Ribautodelphax imitans (Rib.) is strictly monophagous on tall fescue (Festuca arundina-
cea), a plant mainly growing in temporarily wet or disturbed grassland such as fallows
and waysides. This plant is also the preferred host of Arthaldeus striifrons (Kbm.) as well
as a number of less specific feeders such as Delphacodes venosus (Germ.), Dicranotropis
 296 Utilization of plant resources

Table 22. Auchenorrhyncha species utilizing Poaceae in Germany. Explanations see Table 5.

A nthoxanthum odoratum

Helictotrichon pubescens

Corynephorus canescens
A rrhenatherum elatius

Deschampsia cespitosa

Calamagrostis villosa
Phalaris arundinacea
Phragmites australis

Trisetum flavescens
Cynodon dactylon

M olinia caerulea

A lopecurus spp.

A grostis canina
Koeleria glauca

Holcus lanatus

A . stolonifera
Phleum spp.
Setaria spp.

A . capillaris
H. pratense
A vena spp.

D. flexuosa

A . vinealis
Z ea mays

H. mollis
Pentastiridius leporinus (L.) . . . X . . . . . . . . . . . . . . . . . . . . .
Delphax crassicornis (Panz.) . . . X . . . . . . . . . . . . . . . . . . . . .
Delphax pulchellus (Cu rt.) . . . X . . . . . . . . . . . . . . . . . . . . .
Euides speciosa (Boh.) . . . X . . . . . . . . . . . . . . . . . . . . .
Chloriona unicolor (H .-S.) . . . X . . . . . . . . . . . . . . . . . . . . .
Chloriona sicula Mats. . . . X . . . . . . . . . . . . . . . . . . . . .
Chloriona dorsata Ed w . . . . X . . . . . . . . . . . . . . . . . . . . .
Chloriona glaucescens Fieb. . . . X . . . . . . . . . . . . . . . . . . . . .
Chloriona smaragdula (Stål) . . . X . . . . . . . . . . . . . . . . . . . . .
Chloriona stenoptera (Fl.) . . . X . . . . . . . . . . . . . . . . . . . . .
Chloriona vasconica Rib. . . . X . . . . . . . . . . . . . . . . . . . . .
Paralimnus phragmitis (Boh.) . . . X . . . . . . . . . . . . . . . . . . . . .
Paralimnus rotundiceps (Leth.) . . . X . . . . . . . . . . . . . . . . . . . . .
Calamotettix taeniatus (H orv.) . . . X . . . . . . . . . . . . . . . . . . . . .
M uellerianella extrusa (Scott) . . . . X . . . . . . . . . . . . . . . . . . . .
Litemixia pulchripennis Asche . . . . X . . . . . . . . . . . . . . . . . . . .
Deltocephalus maculiceps Boh. . . . . x . . . . . . . . . . . . . . . . . . . .
Jassargus sursumflexus (Then) . . . . X . . . . . . . . . . . . . . . . . . . .
X anthodelphax xantha Vilb. . . . . ? . . . . . . . . . . . . . . . . . . . .
Sorhoanus schmidti (W.Wg.) . . . . ? . . . . . . . . . . . . . . . . . . . .
Stenocranus major (Kbm .) . . . . . X . . . . . . . . . . . . . . . . . . .
Paraliburnia adela (Fl.) . . . . . X . . . . . . . . . . . . . . . . . . .
Balclutha rhenana W.Wg. . . . . . X . . . . . . . . . . . . . . . . . . .
Erzaleus metrius (Fl.) . . . . . X . . . . . . . . . . . . . . . . . . .
Ribautodelphax angulosa (Rib.) . . . . . . X . . . . . . . . . . . . . . . . . .
M egadelphax sordidula (Stål) . . . . . . . . . . X . . . . . . . . . . . . . .
Doratura horvathi W.Wg. . . . . . . . . . . . . X . . . . . . . . . . . .
Z yginidia viaduensis (W.Wg.) . . . . . . . . . . . . . X . . . . . . . . . . .
Psammotettix angulatus (Then) . . . . . . . . . . . . . ? . . . . . . . . . . .
A canthodelphax denticauda (Boh.) . . . . . . . . . . . . . . . X . . . . . . . . .
M uellerianella brevipennis (Boh.) . . . . . . . . . . . . . . . X . . . . . . . . .
Streptanus confinis (Reu t.) . . . . . . . . . . . . . . . X . . . . . . . . .
Psammotettix excisus (Mats.) . . . . . . . . . . . . . . . . . X . . . . . . .
Psammotettix albomarginatus W.Wg. . . . . . . . . . . . . . . . . . ? . . . ? . . .
Recilia horvathi (Then) . . . . . . . . . . . . . . . . . ? . . . . . . .
M uellerianella fairmairei (Perr.) . . . . . . . . . . . . . . . . . . X X . . . . .
M ocydiopsis monticola Rem . . . . . . . . . . . . . . . . . . . . X . . . . .
Ribautodelphax vinealis Biem . . . . . . . . . . . . . . . . . . . . . . X . . .
Doliotettix lunulatus (Zett.) . . . . . . . . . . . . . . . . . . . . . . . ? .
 Poaceae 297

Brachypodium pinnatum
C. pseudophragmites
A mmophila arenaria
Calamagrostis varia

Dactylis glomerata
Puccinellia distans

Leymus arenarius
Festuca altissima
Glyceria maxima

Sesleria albicans

Lolium perenne
M elica uniflora

Bromus erectus
C. arundinacea

F. arundinacea
F. heterophylla

Elymus repens

Hordeum spp.
N ardus stricta

Triticum spp.
Secale cereale
C. canescens

P. nemoralis
P. maritima
Briza media

P. pratensis
F. pratensis

Poa annua
C. epigejos

G. fluitans

P. trivialis
Stipa spp.

B. inermis
F. rubra
F. ovina
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . ? ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
 298 Utilization of plant resources

Table 22. (continued):

A nthoxanthum odoratum

Helictotrichon pubescens

Corynephorus canescens
A rrhenatherum elatius

Deschampsia cespitosa

Calamagrostis villosa
Phalaris arundinacea
Phragmites australis

Trisetum flavescens
Cynodon dactylon

M olinia caerulea

A lopecurus spp.

A grostis canina
Koeleria glauca

Holcus lanatus

A . stolonifera
Phleum spp.
Setaria spp.

A . capillaris
H. pratense
A vena spp.

D. flexuosa

A . vinealis
Z ea mays

H. mollis
Ribautodelphax collina (Boh.) . . . . . . . . . . . . . . . . . . . . . . X . .
M ocydiopsis parvicauda Rib. . . . . . . . . . . . . . . . . . . . . . . X . .
X anthodelphax straminea (Stål) . . . . . . . . . . . . . . . . ?* . . . . X X x .
Elymana kozhevnikovi (Zachv.) . . . . . . . . . . . . . . . . . . . . . . . . .
Paraliburnia clypealis (J. Shlb.) . . . . . . . . . . . . . . . . . . . . . . . . .
Criomorphus moestus (Boh.) . . . . . . . . . . . . . . . . . . . . . . . . .
Streptanus okaensis Zachv. . . . . . . . . . . . . . . . . . . . . . . . . .
Lebradea calamagrostidis Rem . . . . . . . . . . . . . . . . . . . . . . . . . .
Criomorphus borealis (J. Shlb.) . . . . . . . . . . . . . . . . . . . . . . . . X
Paluda flaveola (Boh.) . . . . ? ? . . . . . . . . . ? . . . . . . . . X
Eurysula lurida (Fieb.) . . . . . . . . . . . . . . . . . . . . . . . . .
M irabella albifrons (Fieb.) . . . . . . . . . . . . . . . . . . . . . . . . .
Endria nebulosa (Ball) . . . . . . . . . . . . . . . . . . . . . . . . .
Doratura impudica H orv. . . . . . . . . . . . . . . . . . . . . . . . . .
A rthaldeus arenarius Rem . . . . . . . . . . . . . . . . . . . . . . . . . .
Balclutha calamagrostis Oss. . . . . . . . . . . . . . . . . . . . . . . . . .
Psammotettix poecilus (Fl.) . . . . . . . . . . . . . . . . . . . . . . . . .
Pseudodelphacodes flaviceps (Fieb.) . . . . . . . . . . . . . . . . . . . . . . . . .
Psammotettix unciger Rib. . . . . . . . . . . . . . . . . . . . . . . . . .
Gravesteiniella boldi (Scott) . . . . . . . . . . . . . . . . . . . . . . . . .
Psammotettix maritimus (Perr.) . . . . . . . . . . . . . . . . . . . . . . . . .
Doratura littoralis Ku ntze . . . . . . . . . . . . . . . . . . . . . . . . .
Praganus hofferi (Dlab.) . . . . . . . . . . . . . . . . . . . . . . . . .
Psammotettix nardeti Rem . . . . . . . . . . . . . . . . . . . . . . . . . .
Struebingianella lugubrina (Boh.) . . . . . . . . . . . . . . . . . . . . . . . . .
Chlorionidea flava P. Löw . . . . . . . . . . . . . . . . . . . . . . . . .
Z yginidia mocsaryi (H orv.) . . . . . . . . . . . . . . . . . . . . . . . . .
Z yginidia franzi (W.Wg.) . . . . . . . . . . . . . . . . . . . . . . . . .
Psammotettix putoni (Then) . . . . . . . . . . . . . . . . . . . . . . . . .
M acrosteles sordidipennis (Stål) . . . . . . . . . . . . . . . . . . . . . . . . .
A noscopus limicola (Ed w .) . . . . . . . . . . . . . . . . . . . . . . . . .
Psammotettix cephalotes (H .-S.) . . . . . . . . . . . . . . . . . . . . . . . . .
Cicadula persimilis (Ed w .) . . . . . . . . . . . . . . . . . . . . . . . . .
Stenocranus minutus (F.) . . . . . . . . . . . . . . . . . . . . . . . . .
Ribautodelphax imitans (Rib.) . . . . . . . . . . . . . . . . . . . . . . . . .
A rthaldeus striifrons (Kbm .) . . . . . . . . . . . . . . . . . . . . . . . . .
M egamelodes quadrimaculatus (Sign.) . . . . . . . . . . . . . . . . . . . . . . . . .
Dicranotropis divergens Kbm . . . . . . . . . . . . . . . . . ?* . . . . . . . .
A canthodelphax spinosa (Fieb.) . . . . . . . . . . . . . . . . . . . . . . . . .
 Poaceae 299

Brachypodium pinnatum
C. pseudophragmites
A mmophila arenaria
Calamagrostis varia

Dactylis glomerata
Puccinellia distans

Leymus arenarius
Festuca altissima
Glyceria maxima

Sesleria albicans

Lolium perenne
M elica uniflora

Bromus erectus
C. arundinacea

F. arundinacea
F. heterophylla

Elymus repens

Hordeum spp.
N ardus stricta

Triticum spp.
Secale cereale
C. canescens

P. nemoralis
P. maritima
Briza media

P. pratensis
F. pratensis

Poa annua
C. epigejos

G. fluitans

P. trivialis
Stipa spp.

B. inermis
F. rubra
F. ovina
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
X X x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. x x X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . x X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . x X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . ? X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . X x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . X x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . ? . . . . . . . . . . . . . . ? . . . . . . . . . . . . .
. . . . . . . ? . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . X X . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . X . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . X . . . . x . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . ? . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . ? X . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . x ? . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . ? . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . X . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . X . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . X . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . ? X . ? . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . ? . . ? . . . . . . . . . . . . . .
. . . . . . . ? . . . . . . . . . . . . X . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . X x . . . . . . . . . . . . .
 300 Utilization of plant resources

Table 22. (continued):

A nthoxanthum odoratum

Helictotrichon pubescens

Corynephorus canescens
A rrhenatherum elatius

Deschampsia cespitosa

Calamagrostis villosa
Phalaris arundinacea
Phragmites australis

Trisetum flavescens
Cynodon dactylon

M olinia caerulea

A lopecurus spp.

A grostis canina
Koeleria glauca

Holcus lanatus

A . stolonifera
Phleum spp.
Setaria spp.

A . capillaris
H. pratense
A vena spp.

D. flexuosa

A . vinealis
Z ea mays

H. mollis
Delphacinus mesomelas (Boh.) . . . . . . . . . . . . . . . . . . . . . . . . .
Eupelix cuspidata (F.) . . . . . . . . . . . . . . . . . . . . . . . . .
Rhopalopyx adumbrata (C. Shlb.) . . . . . . . . . . . . . . . . . . . . . . . . .
M ocydiopsis attenuata (Germ .) . . . . . . . . . . . . . . . . . . . . . . . . .
A rocephalus punctum (Fl.) . . . . . . . . . . . . . . . . . ? . . . . . . .
Kosswigianella exigua (Boh.) . . . . . . . . . . . . . . . . . . . . . . . . .
Rhytistylus proceps (Kbm .) . . . . . . . . . . . . . . . . . . . . . . . . .
Rhopalopyx vitripennis (Fl.) . . . . . . . . . . . . . . . . . . . . . . . . .
Jassidaeus lugubris (Sign.) . . . . . . . . . . . . . . . . . . . . . . . . .
M etropis latifrons (Kbm .) . . . . . . . . . . . . . . . . . . . . . . . . .
M etropis inermis W.Wg. . . . . . . . . . . . . . . . . . . . . . . . . .
N eophilaenus infumatus (H pt.) . . . . . . . . . . . . . . . . . . . . . . . . .
Doratura exilis H orv. . . . . . . . . . . . . . . . . . . . . . . . . .
Hardya signifer (Then) . . . . . . . . . . . . . . . . . . . . . . . . .
M ocydiopsis longicauda Rem . . . . . . . . . . . . . . . . . . . . . . . . . .
Psammotettix pallidinervis (Dhlb.) . . . . . . . . . . . . . . . . . ? . . . . . . .
M endrausus pauxillus (Fieb.) . . . . . . . . . . . . . . . . . . . . . . . . .
A chorotile albosignata (Dhlb.) . . . . . . . . . . . . . . . . . . . . . . . . .
A rocephalus sagittarius Rib. . . . . . . . . . . . . . . . . . . . . . . . . .
Ebarrius cognatus (Fieb.) . . . . . . . . . . . . . . . . . . . . . . . . .
Ribautodelphax albostriata (Fieb.) . . . . . . . . . . . . . . . . . . . . . . . . .
X anthodelphax flaveola (Fl.) . . . . . . . . . . . . . . . . . . . . . . . . .
Rhopalopyx preyssleri (H .-S.) . . . . . . . . . . . . . . . . . . . . . . . . .
M ocydiopsis intermedia Rem . . . . . . . . . . . . . . . . . . . . . . . . . .
M uirodelphax aubei (Perr.) . . . . . . . . . . . . . . . . . . . . . . . . .
Jassargus flori (Fieb.) . . . . . . . . . . . . . . . . ? . . . . . ? . .
Criomorphus williamsi China . . . . . . . . . . . . . . . . . . . . . . . . .
Ditropsis flavipes (Sign.) . . . . . . . . . . . . . . . . . . . . . . . . .
Ribautodelphax pungens (Rib.) . . . . . . . . . . . . . . . . . . . . . . . . .
N eophilaenus albipennis (F.) . . . . . . . . . . . . . . . . . . . . . . . . .
A darrus multinotatus (Boh.) . . . . . . . . . . . . . . . . . . . . . . . . .
A darrus bellevoyei (Pu t.) . . . . . . . . . . . . . . . . . . . . . . . . .
Unkanodes excisa (Mel.) . . . . . . . . . . . . . . . . . . . . . . . . .
Eurysa lineata (Perr.) . . . . . . . . . . . . . . . . x . . . . . . . .
Eurysella brunnea (Mel.) . . . . . . . . . . ? . . . . . . . . . . . . . .
Eurybregma nigrolineata Scott . . . . . . . . . . . . . . . . . . x . . . . . .
Stiroma affinis Fieb. . . . . . . . . . . . . . . . ? x . . . . . . . .
Stiroma bicarinata (H .-S.) . . . . . . . . . . x . . . . X . . . X . . . . .
Laodelphax striatella (Fall.) . x . . . . . ?* ? . ? . . . . . . . . . . . . . .
 Poaceae 301

Brachypodium pinnatum
C. pseudophragmites
A mmophila arenaria
Calamagrostis varia

Dactylis glomerata
Puccinellia distans

Leymus arenarius
Festuca altissima
Glyceria maxima

Sesleria albicans

Lolium perenne

Bromus erectus
M elica uniflora
C. arundinacea

F. arundinacea
F. heterophylla

Elymus repens

Hordeum spp.
N ardus stricta

Triticum spp.
Secale cereale
C. canescens

P. nemoralis
P. maritima
Briza media

P. pratensis
F. pratensis

Poa annua
C. epigejos

G. fluitans

P. trivialis
Stipa spp.

B. inermis
F. rubra
F. ovina
. . . . . . . . . . . . . . . . . . . . X X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . ? X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . X x . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . x X X . . . . . . . . . . . . .
. . . . . . . ? . . . . . . . . . . . . ?* X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . ?* X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . ?* X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . ? X . . . . . . . . . . . . .
. . . . . . ? . . . . . . . . . . . . . . X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . x . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . x . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . x . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . ? . ? . . ? . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . X . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . X . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . X . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . X . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . X . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X . . .
. . . . . . . . . . x . . . . . . . . . . . . . . . X . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . x . . X . . . . .
. . . . . . . . . . . . . . . x . . . . . . . . . . . . . . X . . . .
. x . . . . . . . . x . . . . . x . . . . . . . . . x . . x . . . . .
. . . . . . . . ? . . . . . . x . . . . . . . . . . . . . . x . . . .
. . . . . . . . . . . . . . . ? . . . . . . ? ? ? . . . . . ? . ? ? x
 302 Utilization of plant resources

Table 22. (continued):

A nthoxanthum odoratum

Helictotrichon pubescens

Corynephorus canescens
A rrhenatherum elatius

Deschampsia cespitosa

Calamagrostis villosa
Phalaris arundinacea
Phragmites australis

Trisetum flavescens
Cynodon dactylon

M olinia caerulea

A lopecurus spp.

A grostis canina
Koeleria glauca

Holcus lanatus

A . stolonifera
Phleum spp.
Setaria spp.

A . capillaris
H. pratense
A vena spp.

D. flexuosa

A . vinealis
Z ea mays

H. mollis
Hyledelphax elegantula (Boh.) . . . . x . . . . . ?* ?* . . . . X . . x . . . . .
Delphacodes venosus (Germ .) . . . . x . . . . . . . . . . X . . . . x . . x .
Dicranotropis hamata (Boh.) . . . . . . . . . . ? . . . . . . . X X . . . . .
Dicranotropis montana (H orv.) . . . . . . . . . . . . . . . X . . . . . . . . .
Scottianella dalei (Scott) . . . . . . . . . . . . . . . . . . . . . . ?* . .
Criomorphus albomarginatus Cu rt. . . . . . . . . . . . . . . . x x . ? ? . . . . .
Toya propinqua (Fieb.) . . x . . . . . . . . . . . . . . . . . . . . . .
Javesella discolor (Boh.) . . . . . . . . . . . . . . . X X . . . . . . . .
Javesella pellucida (F.) . ? . . x ? . x x ?* X . . . ? X ? . X x . . x x .
Javesella dubia (Kbm .) . . . . . . . ? . . . . . . . . . . ? ? . . X X .
Javesella obscurella (Boh.) . . . . . . . X . . . . . . . . . . . . . . . ? .
Javesella salina (H pt.) . . . . . . . . . . . . . . . . . . . . . . . . .
Javesella forcipata (Boh.) . . . . . . . . . . . . . . . X ?* . . X . . . ? .
Cercopis sanguinolenta (Scop.) . . . . . . . . . . x . . . . . . . . . . . . . .
Cercopis vulnerata Rossi . . . . . ?* . . . . x . . . . . . . . . . . . . .
Haematoloma dorsatum (Ahr.) . . . . . . . . . . . . . . . . X . . . . . . . .
Lepyronia coleoptrata (L.) . . . x* ?* . . . . . . . . . . . . . . . . . . ?* .
N eophilaenus campestris (Fall.) . . . . . . . . . . ? . . . . . . . . . . . ? . .
N eophilaenus exclamationis (Thnbg.) . . . . . . . . . . . . . . . . ? . . . . . . . .
N eophilaenus lineatus (L.) . . . x* ? ? . . . . . . . . . X X . . x . . x ?* X
N eophilaenus minor (Kbm .) . . . . . . . . . . . . . x . . . X . . . . . . .
A phrophora major Uhl. . . . x* . . . . . . . . . . . . . . . . . . . . .
Philaenus spumarius (L.) . . . x* . . . . . . . . . . . . . . . ?* . . . . .
A noscopus albifrons (L.) . . . . . . . . . . ?* . . . ?* . X . . x . . ? . .
A noscopus flavostriatus (Don.) . . . . . . . . . . ? . . . . . X . . . . . . . .
A noscopus serratulae (F.) . . . . . . . . . . x . . . . . . . . x . . . . .
Cicadella viridis (L.) . . . ? . . . . . . . . . . . . . . . . . . . . .
Dikraneura variata H ard y . . . . . . . . . . . . . . . . X . . . . . . . .
Z yginidia pullula (Boh.) ?* ?* ?* . . . . . . . . . . . . . . . . . . . . . .
Z yginidia scutellaris (H .-S.) x ? . . ? . . . x . ? ?* . x . . . x ? ?* . . ?* . .
Balclutha punctata (F.) . . . . . . . . . . . . . . . ? X . . X . . X . ?
Balclutha saltuella (Kbm .) . . . . . . . . . . . . . . . . . . . . . . . . .
M acrosteles cristatus (Rib.) . . . . . . . ?* . ?* . . . . . . . . . . . . . . .
M acrosteles laevis (Rib.) . . . . . . . ?* ? . . . . . . . . . . . . . ?* . .
M acrosteles sexnotatus (Fall.) . . . ? . . . x ? . ?* . . . . . . . . . . . . . .
M acrosteles viridigriseus (Ed w .) . . . . . . . . . . . . . . . . . . . . x . ?* ?* .
Deltocephalus pulicaris (Fall.) . . . . . . . . . . . . . . . . . . ? . ? . x x .
Recilia coronifer (Marsh.) . . . . x . . . . . . . . . . . . . ?* X . . . . .
Recilia schmidtgeni (W.Wg.) . . x . . . . . . . . . . . . . . . . . . . . . .
 Poaceae 303

Brachypodium pinnatum
C. pseudophragmites
A mmophila arenaria
Calamagrostis varia

Dactylis glomerata
Puccinellia distans

Leymus arenarius
Festuca altissima
Glyceria maxima

Sesleria albicans

Lolium perenne
M elica uniflora

Bromus erectus
C. arundinacea

F. arundinacea
F. heterophylla

Elymus repens

Hordeum spp.
N ardus stricta

Triticum spp.
Secale cereale
C. canescens

P. nemoralis
P. maritima
Briza media

P. pratensis
F. pratensis

Poa annua
C. epigejos

G. fluitans

P. trivialis
Stipa spp.

B. inermis
F. rubra
F. ovina
. x . . . . . . . . . . . . ?* . . . . . x ?* . . . . x ?* . x . . . . .
. . . . . . . . . . . . . . . ? . ? x . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . X . . x . . . . . . . . . ? . X . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. x . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. x x x . ? . . . . . . x . . X . ?* X . x ? X X X X . ? ? . X . ?* ?* ?*
. . . . . . . . . . . . . . . . . . . . . . ? ? . . . . . . . . . . .
. . . . . . . . . x . . . . . . . . . . . . ? ? . . . . . . . . . . .
. . . . . . . . . . . . ? . ? . . . . . . . . . . . . . . . . . . . .
. . ? . . . . . . . . . . . . . . . . . . . . ?* . ? . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . x* . . x* . . . . x* . . . . .
. . . . . . . . . . . . . . . ?* . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . x . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . .
. . . . . . . . . . . . . . . . . . . ? . X . . . . . . . . . . . . .
x x x X . ? . x . . . . . . . ?* ? . . . X x . . . . . . . . . ? . . .
. . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . ?* . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . ?* ?* . . . . ?* ?* . . . . . . . x . . . . .
. . . ? . . . . . . . . . . . x . . ? ? ? . . . . . . . . . x . . . .
. . . . . . . . . . . . ? . . x . . ? . ? . ? ? . ? . . . . x . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . x x x . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . ?* . . . . . .
. . . ? . . . . . . . . . . ?* X . ?* ? . ? x ? ? ? . . ? ? ?* ? . . . x
? ? x X . . . . . . . . . . . . . . . . ?* . . . . . . . . . ? . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . ? ? . . . . . . . . ?* ?* ?*
. . . . . . . . . . . . . . . ?* . . . . ?* . ? ? . . . . . . . . ? ?* ?
. . . . . . . . . . . . x . . x* . . . . . . ? ? . . . . . . ? . ? ? ?
. . . . . . . . . . . . ? . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . x . . . . . . x x . ? . . . . x . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
 304 Utilization of plant resources

Table 22. (continued):

A nthoxanthum odoratum

Helictotrichon pubescens

Corynephorus canescens
A rrhenatherum elatius

Deschampsia cespitosa

Calamagrostis villosa
Phalaris arundinacea
Phragmites australis

Trisetum flavescens
Cynodon dactylon

M olinia caerulea

A lopecurus spp.

A grostis canina
Koeleria glauca

Holcus lanatus

A . stolonifera
Phleum spp.
Setaria spp.

A . capillaris
H. pratense
A vena spp.

D. flexuosa

A . vinealis
Z ea mays

H. mollis
Doratura stylata (Boh.) . . . . . . . . . . . . . . . . . . . . . . X . .
Doratura homophyla (Fl.) . . . . . . . . . . . . . . . . . . . . . . x . .
A llygus maculatus Rib. . . . . . . . . . . . . . . . . ? . . . . . . . .
A llygus modestus Scott . . . . . . . . . . . . . . . . . . . . . . . . .
A llygidius commutatus (Fieb.) . . . . . . . . . . . . . . . . . . . . . . . . .
A llygidius atomarius (F.) . . . . . . . . . . . . . . . . . . . . . . . . .
Graphocraerus ventralis (Fall.) . . . . . . ? . . . ?* ?* . . ?* . . . ? . . . ? . .
Hardya melanopsis (H ard y) . . . . . . . . . . . . . . . . ? . . . . . . . .
Hardya tenuis (Germ .) . . . . . . . . . . . . . . . . . . . . . . ? . .
Sardius argus (Marsh.) . . . . . . . . . . . . . . . . . . . . . . ? . .
Elymana sulphurella (Zett.) . . . . . . . . . . . . . . . . . . X X . . . ? ?*
M ocydia crocea (H .-S.) . . . . ? . . . . . ? . . . . . . . x . . . . . .
Thamnotettix dilutior (Kbm .) . . . . . . . . . . . . . . . . x . . . . . . . .
M acustus grisescens (Zett.) . . . . ? . . . . . . . . . . . . . ? ? . . . . .
A thysanus argentarius Metc. . . x* . . . . . . . X . . . . X . . x x . . . . .
Conosanus obsoletus (Kbm .) . . . . . . . . . . ? . . . . . . . . . . . . . .
Sotanus thenii (P. Löw ) . . . . . . . . . . . . . . . . . . . . . . . . .
Streptanus aemulans (Kbm .) . . . . . . . . . . x . . . . . . . X X . . . . .
Streptanus marginatus (Kbm .) . . . . . . . . . . . . . . . . X . . . . . . . .
Streptanus sordidus (Zett.) . . . . . . . . . . . . . . . ? . . . . . . x x .
A rtianus interstitialis (Germ .) . . . . . . . . . . . . . . . . . . ? . . . ? . .
A rocephalus longiceps (Kbm .) . . . . . . . . . . . . . . . . . . X X . . . . .
A rocephalus languidus (Fl.) . . . . . . . . . . . . . ? . . . . . . . . . . .
Psammotettix kolosvarensis (Mats.) . . . . . . . . . . . . . . . . . . . . . . . . .
Psammotettix alienus (Dhlb.) . x . . . . . ? ? . ? . . . . . . . . . . . ? ? .
Psammotettix sabulicola (Cu rt.) . . . . . . . . . . . . . . . . . . . . . . . . .
Psammotettix helvolus (Kbm .) bas. . . . . . . . . . . ? . . . . . ? . . . . . ? . .
Psammotettix helvolus (Kbm .) acid . . . . . . . . . . . . . . . . . X . . . . . . . .
Psammotettix nodosus (Rib.) . . . . . . . . . . . . . . . . x ? ? . . . ? . .
Psammotettix confinis (Dhlb.) . . . . . . . ?* . . . . . . . . ? ? . . . . x ?* .
Errastunus ocellaris (Fall.) . . . . . . . . . . ? . . . . . . . X X . . . ? .
Turrutus socialis (Fl.) . . . . . . . . . . . ?* . . . . . . . . . . . . .
Jassargus pseudocellaris (Fl.) . . . . . . . . . . . . . . . . . . . ? . . x . .
Jassargus obtusivalvis (Kbm .) . . . . . . . . . . . . . . . . . . . . . . . . .
Jassargus alpinus (Then) . . . . . . . . . . . . . . . . ? . . . . . . . X
Jassargus allobrogicus (Rib.) . . . . . . . . . . . . . . . . X . . . . . . . ?*
Diplocolenus bohemani (Zett.) . . . . . . . . . . . . . . . . . . . . . . . . X
V erdanus abdominalis (F.) . . . . . . . . . . . ?* . . . . ? . x ? . . ? . .
V erdanus bensoni (China) . . . . . . . . . . . . . . . . ?* . . ?* . . . . .
 Poaceae 305

Brachypodium pinnatum
C. pseudophragmites
A mmophila arenaria
Calamagrostis varia

Dactylis glomerata
Puccinellia distans

Leymus arenarius
Festuca altissima
Glyceria maxima

Sesleria albicans

Lolium perenne
M elica uniflora

Bromus erectus
C. arundinacea

F. arundinacea
F. heterophylla

Elymus repens

Hordeum spp.
N ardus stricta

Triticum spp.
Secale cereale
C. canescens

P. nemoralis
P. maritima
Briza media

P. pratensis
F. pratensis

Poa annua
C. epigejos

G. fluitans

P. trivialis
Stipa spp.

B. inermis
F. rubra
F. ovina
. . . . . . . ? . . . . . . . . . . . . X x . . ? . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . ? . . . .
. ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. x . x . . . . . . ? . . . . . . . . . . . . . . . . . . x . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . x . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . X . . . . ? . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
x x . X . . . . . . . . . . . . . . . . . . . . . . . . ? . x . . . .
? . . X . . . . . . . . . . . . . . . . . . . . . . . X ? X x . . . .
. . . . . . . . . . . . . . . . . . . ? . ? . . . . . . . . . . . . .
. x . x . . . . . . . . . . . x . . x . . . . . . . . . ? . x . . . .
. . x X . . . . . . . . . . . X . . x . . . . . . . . . ? . X . . . .
. . . . . . . . . . . . . . . . . . x . . . . . . . . . . . ? . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . X . . . .
. . . . . . . ? . . . . . . . . . . . . . X . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . X . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . X . . . . . . .
. . . . . . X . . . . X . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . X . . . . . . . . . . . . . . . . . x . . . .
. . . . . . . . . . . . ? . . . . . . . . . . ? ? . . . ? . ? . X ?* ?
. . . ? . ? . . . . . . . . . . . . . . . . . . . . . . . . ? ? . . .
. . . . . . . . . . . . . . . . . . . . . ? . . ? . . ? . ? ? . . . .
. . . . . . . ? . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . ? ? . . . . . . . . . . . . .
. . . . . . . . . . . . ? ? ?* . . ?* . . ? ? ? ? ? . . . . . . . . . .
. . x X . . . . . . . . . . . x . . . . . . . . . . . . ? . X . . . .
. . . . . . . ? . . . . . . . . . . . . X ?* . . . . . X . ?* . . . . .
. . . . . . . ?* . . . . . . . . . . . . x . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . ? . ? . . . . .
. ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . ? ? ? . . . . . . . . . . . . .
X ? . X . . . . . . . . . . . . ? . . . . . . . . . . x . . . . . . .
. . . . . . . . . . . . . . ?* . . . . . ? ?* . . . . . ?* . ?* . . . . .
. . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . . . . .
 306 Utilization of plant resources

Table 22. (continued):

A nthoxanthum odoratum

Helictotrichon pubescens

Corynephorus canescens
A rrhenatherum elatius

Deschampsia cespitosa

Calamagrostis villosa
Phalaris arundinacea
Phragmites australis

Trisetum flavescens
Cynodon dactylon

M olinia caerulea

A lopecurus spp.

A grostis canina
Koeleria glauca

Holcus lanatus

A . stolonifera
Phleum spp.
Setaria spp.

A . capillaris
H. pratense
A vena spp.

D. flexuosa

A . vinealis
Z ea mays

H. mollis
A rthaldeus pascuellus (Fall.) . . . . . . . . . . x . . . ? x . . X ? ? . ? x ?*
Enantiocephalus cornutus (H .-S.) . . . . . . . . . . . . . . . . . . . . . . ? . .
M ocuellus collinus (Boh.) . . . . . . . . . . . . . . . . . . . . . . ? . .
M egadelphax haglundi (J. Shlb.) . . . . . . . . . . . . . . . . . . . . . . . . .
Ederranus discolor (J. Shlb.) . . . ?* . . . . . . . . . . . . . . . . . . . . .
Psammotettix dubius Oss. . . . . . . . . . . . . . . . . . . . . . . . . .
Psammotettix inexpectatus Rem . . . . . . . . . . . . . . . . . . . . . . . . . .
Ebarrius interstinctus (Fieb.) . . . . . . . . . . . . . . . . . . . . . . . . .
Jassargus repletus (Fieb.) . . . . . . . . . . . . . . . . . . . . . . . . .
Pinumius areatus (Stål) . . . . . . . . . . . . . ? . . . . . . . . . . .
Balclutha boica W.Wg . . . . . . . . . . . . . . . . . . . . . . . . .
A llygidius abbreviatus (Leth.) . . . . . . . . . . . . . . . . . . . . . . . . .
Rhopalopyx elongata W.Wg. . . . . . . . . . . . . . . . . . . . . . . . . .
Psammotettix notatus (Mel.) . . . . . . . . . . . . . . . . . . . . . . . . .
Errastunus leucophaeus (Kbm .) . . . . . . . . . . . . . . . . . . . . . . . . .
Diplocolenus penthopitta (Walk.) . . . . . . . . . . . . . . . . . . . . . . . . .
A siraca clavicornis (F.) . . . . . . . . . . . . . . . . . . . . . . . . .
Dictyophara europaea (L.) . . . . . . . . . . . . . . . . . . . . . . . . .
Cercopis arcuata Fieb. . . . . . . . . . . . . . . . . . . . . . . . . .
A phrodes makarovi Zachv. . . . . . . . . . . . . . . . . . . . . . . . . .
Planaphrodes bifasciata (L.) . . . . . . . . . . ?* ?* . . ?* . . . . . . . . . .
Planaphrodes nigrita (Kbm .) . . . . . . . . . . . . . . . . . . . . . . . . .
A noscopus albiger (Germ .) . . . . . . . . . . . . . . . . . . . . . . . . .
A noscopus alpinus (W.Wg.) . . . . . . . . . . . . . . . . . . . . . . . . .
A noscopus histrionicus (F.) . . . . . . . . . . . . . . . . . . . . . . . . .
Forcipata forcipata (Fl.) . . . . . . . . . . . . . . . . . . . . . . . . .
M acrosteles ossiannilssoni Ld b. . . . . . . . . . . . . . . . . . . . . . . . . .
M acrosteles quadripunctulatus (Kbm .) . ? . . . . . . . . . . . . . . . . . . . . . . .
Colladonus torneellus (Zett.) . . . . . . . . . . . . . . . . . . . . . . . . .
Lamprotettix nitidulus (F.) . . . . . . . . . . . . . . . . . . . . . . . . .
A llygus communis (Ferr.) . . . . . . . . . . . . . . . . . . . . . . . . .
A llygus mixtus (F.) . . . . . . . . . . . . . . . . . . . . . . . . .
Hesium domino (Reu t.) . . . . . . . . . . . . . . . . . . . ? . . . . .
Thamnotettix confinis (Zett.) . . . . . . . . . . . . . . . . . . . . . . . . .
Euscelis incisus (Kbm .) . . . . . . . . . . ?* ?* . . ?* . . . ? . . . . . .
Euscelis lineolatus Br. . . . . . . . . . . . . . . . . . . ?* . . . . . .
Feeding substantiated 1 2 3 18 8 4 1 3 2 0 9 0 1 3 0 13 16 3 12 17 3 1 13 7 5
Feeding unconfirmed 1 4 1 3 8 4 1 6 4 2 17 7 0 3 6 4 13 6 11 10 2 1 18 10 4
Substantiated plus unconfirmed 2 6 4 21 16 8 2 9 6 2 26 7 1 6 6 17 29 9 23 27 5 2 31 17 9

x* = after literature from Germany, ?* = after literature from neighbouring countries.

 Poaceae 307

Brachypodium pinnatum
C. pseudophragmites
A mmophila arenaria
Calamagrostis varia

Dactylis glomerata
Puccinellia distans

Leymus arenarius
Festuca altissima
Glyceria maxima

Sesleria albicans

Lolium perenne
M elica uniflora

Bromus erectus
C. arundinacea

F. arundinacea
F. heterophylla

Elymus repens

Hordeum spp.
N ardus stricta

Triticum spp.
Secale cereale
C. canescens

P. nemoralis
P. maritima
Briza media

P. pratensis
F. pratensis

Poa annua
G. fluitans
C. epigejos

P. trivialis
Stipa spp.

B. inermis
F. rubra
F. ovina
? ? x X . . . ?* . . . . . . . X . x x . X . ? ? ? X . . . . X . . . .
. . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . X . . . .
. . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . X . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . ?* . . . . . ?* ?* . . . . . ?* . ?* . . . . .
. . ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . ? . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . ?* . . ?* . . ? . ? ? ? . . ?* . . . . . . .
. . . . . . . . . . . . . . . ?* . ?* . . . . ?* . ?* . . . . . . . . . .
4 10 16 19 3 2 1 1 1 2 2 3 4 1 1 15 2 1 10 2 14 27 2 2 7 2 4 5 0 11 18 1 1 0 2
3 5 4 5 1 4 1 10 2 0 2 1 7 3 8 8 2 8 3 5 24 18 12 13 11 5 1 8 10 8 10 2 5 6 5
7 15 20 24 4 6 2 11 3 2 4 4 11 4 9 23 4 9 13 7 38 45 14 15 18 7 5 13 10 19 28 3 6 6 7
308 Utilization of plant resources

hamata (Boh.), Macustus grisescens (Zett.), Athysanus argentarius Metc. and Conosanus obso-
letus (Kbm.). Various-leaved and wood fescue (F. heterophylla and F. altissima), both grow-
ing in forests, have been sampled only a few times, the former revealing populations of
Mocydiopsis attenuata (Germ.) and Dikraneura variata Hardy. The latter was found to be
exploited by Stiroma affinis Fieb. and Zyginidia mocsaryi (Horv.). M. attenuata (Germ.) is
2nd degree monophagous, feeding mainly on F. ovina and F. rubra, while the remaining
three species also feed on other grass genera. Giant fescue (F. gigantea), a typical grass of
wet forests on nutrient-rich substrates, is apparently not attacked at all, whereas meadow
fescue (F. pratensis), mainly growing in damp meadows and pastures, is suspected to hold
a few oligophagous and polyphagous species, like Javesella pellucida (F.), Zyginidia scute-
laris (H.-S.), Psammotettix confinis (Dhlb.), Arthaldeus pascuellus (Fall.), A. striifrons (Kbm.)
and Euscelis spp. Very recently, however, there were some records of both adults and
nymphs of the little known planthopper Megamelodes quadrimaculatus (Sign.).
(ii) Small-reed (Calamagrostis)
Most species of small-reed (Calamagrostis), notably purple, wood, rough, shaggy and
mountain small-reed (C. canescens, C. epigejos, C. arundinacea, C. villosa, C. varia), grow as
tall swards or tussocks in fenland or in forests and clearings, except bank small-reed (C.
pseudophragmites), which is a pioneer grass on bare gravel banks of alpine rivers. All
these species have been studied extensively during recent years. However, the insect
fauna of the rare bog small-reed (C. stricta), which is largely confined to bogs of north-
eastern Germany, is still almost unknown.
Altogether, Calamagrostis is attacked by 32 Auchenorrhyncha feeders, perhaps plus fur-
ther 12 unconfirmed species. The feeding guild of C. epigejos alone comprises 19 species,
and is one of the most diverse all over Germany (see Fig. 14). Host associations are usual-
ly easier to assess than in other grasses due to the conspicuous physiognomy. 15 of them
are classified here as monophagous in the 1st or 2nd degree. Paraliburnia clypealis (J. Shlb.),
Criomorphus moestus (Boh.) (the latter only after literature data), Streptanus okaensis Zachv.
as well as the only German endemic Lebradea calamagrostidis Rem. are all exclusively asso-
ciated with C. canescens, whereas Endria nebulosa (Ball), Arthaldeus arenarius Rem. and prob-
ably Doratura impudica Horv. are specific to C. epigejos. The latter is also the main host of
Balclutha calamagrostis Oss., Psammotettix poecilus (Fl.), Eurysula lurida (Fieb.) and Mirabella
albifrons (Fieb.). However, these species locally breed also on C. pseudophragmites or
C. canescens. Both Pseudodelphacodes flaviceps (Fieb.) and Psammotettix unciger Rib. are con-
fined to C. pseudophragmites and show a peculiar combination of life history traits of both
r- and K-strategists (see Nickel 1999a). Criomorphus borealis (J. Shlb.) is usually dominant
in stands of C. villosa in montane coniferous forests, but is also found on C. canescens in
lowland fens. Paluda flaveola (Boh.), Elymana kozhevnikovi (Zachv.) and perhaps Xanthodel-
phax xantha Vilb. utilize several Calamagrostis species. Further common, but less specific
feeders on Calamagrostis include Neophilaenus lineatus (L.), Balclutha punctata (F.), Allygid-
ius commutatus (Fieb.), Elymana sulphurella (Zett.), Mocydia crocea (H.-S.), Macustus grise-
scens (Zett.), Athysanus argentarius Metc., Errastunus ocellaris (Fall.), Jassargus alpinus (Then),
Diplocolenus bohemani (Zett.) and Arthaldeus pascuellus (Fall.).
(iii) Hair-grass (Deschampsia)
Hair-grass (Deschampsia), comprises 6 species in Germany, 4 of which are rather lo-
calized and rare, without any data on their insect herbivores. The western European
 Poaceae 309

bog hair-grass (D. setacea) grows on peaty substrates along shores of north German
ponds. In contrast, intermediate hair-grass (D. media) is essentially a Mediterranean
species with isolated populations in flood plain depressions in the northern upper Rhine
plain. Both Lake Constance and mud hair-grass (D. littoralis and D. wibeliana) are local-
ized endemics of Lake Constance and the lower Elbe, respectively. Only tufted and
wavy hair-grass (Deschampsia cespitosa and D. flexuosa) are both widespread and com-
mon. Their fauna has been sampled extensively in many areas.
Altogether, 27 Auchenorrhyncha species have been found to breed, plus another 13
uncertain records. Hence, regarding the biomass and wide distribution of the two grasses
and the usually high Auchenorrhyncha abundances, Deschampsia must be considered
as one of the major food plant groups in central Europe. However, the overlap of the
Auchenorrhyncha guilds between the two grasses is rather small. There are no 2nd
degree monophages at all, and even most 1st degree oligophages only exploit either of
the two grasses. Taxonomically, this situation is much better reflected in the former
treatment of D. flexuosa belonging to the dictinct genus Avenella.
Deschampsia cespitosa is common in various types of moderately rich, damp to wet
grassland, preferably on loamy or clayey substrates, and is also found in forests. It forms
large and conspicuous tussocks, the insect fauna of which is easy to study by direct
search. Acanthodelphax denticauda (Boh.), Muellerianella brevipennis (Boh.) and Streptanus
confinis (Reut.) are all strictly monophagous, the first two being widespread and fairly
common. The latter is more localized and confined to abandoned or low-productivity
grassland. 1st degree oligophages include Stiroma bicarinata (H.-S.), Delphacodes venosus
(Germ.), Javesella forcipata (Boh.), Athysanus argentarius Metc. and others.
In contrast, D. flexuosa forms monospecific turfs in many forests on acidic substrates
as well as in clearings, heaths and meadows. It is a preferred host for numerous Auchen-
orrhyncha species, but non of them is monophagous, although there is a phenomenon,
which can be decribed as ‚regional monophagy‘. For instance, Hyledelphax elegantula
(Boh.) lives on various grasses in middle and southern parts of Germany, e.g. Poa nem-
oralis, Holcus mollis and Calamagrostis arundinacea. Dikraneura variata Hardy locally breeds
on Festuca spp. at least in the region of the Mittelgebirge. In the north German plain,
however, these species are abundant and clearly dominant in monospecific stands of
Deschampsia flexuosa in most pine, oak and beech forests, where none of the other grass-
es occur. Furthermore, D. flexuosa is preferentially utilized by Javesella discolor (Boh.),
Neophilaenus lineatus (L.), Anoscopus albifrons (L.), A. flavostriatus (Don.), Balclutha punc-
tata (F.), Streptanus marginatus (Kbm.), Jassargus allobrogicus (Rib.), the acidophilic form
of Psammotettix helvolus (Kbm.) and some others, and nymphs of the cercopid Haemato-
loma dorsatum (Ahr.) live on its roots. Published records of further species, e.g. Dicrano-
tropis divergens Kbm., Xanthodelphax straminea (Stål) and Rhopalopyx adumbrata (C. Shlb.)
could not yet be confirmed and are perhaps based upon host misidentification. The
Auchenorrhyncha fauna of D. flexuosa in a pine forest near Hanover was recently stud-
ied in detail by Körner et al. (2001), see also Kuntze (1937) and Rabeler (1951, 1957).
(iv) Bent-grass (Agrostis)
Bent-grass (Agrostis) is probably one of the most ubiquitous grass genera in central
Europe, but unlike Festuca and Poa, it is usually absent from basic substrates. 23 Auchen-
orrhyncha species have been found to feed, but as on other low-growing grasses, the
310 Utilization of plant resources

number of unconfirmed feeders is also high and comprises further 16 species. Only the
common lowland species of Agrostis have been studied, but no data exist on the insect
fauna of alpine, rock and slender bent-grass (A. alpina, A. rupestris, A. agrostiflora), all of
which occur in the subalpine and alpine belt. Data are also lacking for some introduced
species, such as rough and Castilian bent-grass (A. scabra and A. castellana).
Common bent-grass (A. capillaris) is often dominant on siliceous low-productivity
grassland, including heaths, pastures, meadows as well as open woodland. It harbours
at least 13 breeders, and in further 18 species, records still need to be confirmed. Both
Ribautodelphax collina (Boh.) and Mocydiopsis parvicauda Rib. are strictly monophagous.
Xanthodelphax straminea (Stål) also breeds on other species of Agrostis. Other important,
but less specific feeders include Javesella dubia (Kbm.), Balclutha punctata (F.), Doratura
stylata (Boh.), D. homophyla (Fl.) and Streptanus sordidus (Zett.), as well as Javesella pellu-
cida (F.), Neophilaenus lineatus (L.), Deltocephalus pulicaris (Fall.), Streptanus sordidus (Zett.),
Psammotettix confinis (Dhlb.) and Jassargus pseudocellaris (Fl.). Noteworthy among un-
certain feeders is the western European Scottianella dalei (Scott), which is perhaps
monophagous.
Brown bent-grass (A. vinealis) is rather localized in sandy and xerothermic sites mainly
of the north German plain, but also on inland dunes and acidic rocky slopes of middle
and southern parts. It is often mistaken for the preceding species; thus, there is only
little information on its insect fauna. Morphological and ecological studies carried out
by Bieman (1987a, 1987b) revealed the existence of the planthopper species Ribautodel-
phax vinealis Biem., which was undescribed until then, and which is exclusively associ-
ated with this grass. Another monophagous feeder is perhaps the rare Psammotettix
albomarginatus W.Wg.
Creeping bent-grass (A. stolonifera) s.l. is a rather eurytopic plant on nutrient-rich
soils being subject to strong fluctuations of the water table, e.g. in ruderal sites, river
flood plains and in near-coastal grassland with moderate salinity, but also in open for-
ests and on clearings. A. gigantea, A. maritima and others are considered as subspecies
or varieties and have not been sampled separately. Specific sampling is particularly
difficult in mixed stands, where Alopecurus geniculatus and Poa spp. co-occur. Altogeth-
er, 17 Auchenorrhyncha species have been found, with confirmed breeding evidence
for only 6 of them. Doliotettix lunulatus (Zett.) is perhaps strictly monophagous, but
confined to cool and moderately shaded sites, avoiding frequently managed grassland.
Xanthodelphax straminea (Stål) prefers A. canina and A. capillaris, but has been found on
A. stolonifera in southern Germany. Further breeding species include Javesella dubia (Kbm.)
and Streptanus sordidus (Zett.) (both often occurring in high densities), as well as Delpha-
codes venosus (Germ.), Javesella pellucida (F.), Deltocephalus pulicaris (Fall.) and Arthaldeus
pascuellus (Fall.). Neophilaenus lineatus (L.), Macrosteles viridigriseus (Edw.), Psammotettix
alienus (Dhlb.), Ps. confinis (Dhlb.), Errastunus ocellaris (Fall.) and others are uncertain
feeders. Thus, in many flood plain meadows and pastures, A. stolonifera is an important
Auchenorrhyncha host.
Velvet bent-grass (A. canina) occurs in rather wet and acidic sites, notably along bog
margins, in fens and spring mires, usually on peaty soils. Its small tufts can be easily
searched for insects, but only three species have been recorded as breeding. At least
locally, it is the main host of the 2nd degree monophagous Xanthodelphax straminea (Stål).
 Poaceae 311

Less specialized feeders include Delphacodes venosus (Germ.), Macrosteles viridigriseus

(Edw.) and perhaps Deltocephalus pulicaris (Fall.) and Arthaldeus pascuellus (Fall.).
(v) Velvet-grass (Holcus)
This genus includes only two species, notably Yorkshire fog and creeping soft-grass
(H. lanatus and H. mollis). Both are widespread and common, the former is mainly found
on moist meadows and pastures of lower altitudes, the latter more on acidic substrates
and in open forests and on clearings. 36 Auchenorrhyncha species have been recorded
altogether, 21 of which definitely breed. With 18 species alone, H. mollis is among the
most-favoured plants in the study area (see Fig. 14).
Mocydiopsis monticola Rem. is apparently restricted to H. mollis, in moderately shad-
ed sites, and is absent from frequently managed grassland, where H. lanatus occurs.
Muellerianella fairmairei (Perr.) lives on both species of Holcus, mainly in meadows and
pastures, but needs Juncus effusus for the oviposition of winter eggs (Drosopoulos 1977).
Numbers of species and individuals of further less specific Auchenorrhyncha are often
high, with a marked overlap between the two grasses. For instance, Dicranotropis hama-
ta (Boh.), Javesella pellucida (F.), Elymana sulphurella (Zett.), Athysanus argentarius Metc.,
Streptanus aemulans (Kbm.), Arocephalus longiceps (Kbm.), Errastunus ocellaris (Fall.) and
Arthaldeus pascuellus (Fall.) all attack both species. Thus, most differences between the
two guilds may rather be explained by requirements of specific microclimatic condi-
tions than by host preferences. For instance, the more sciophilous or hygrophilous spe-
cies Stiroma bicarinata (H.-S.), Hyledelphax elegantula (Boh.), Javesella forcipata (Boh.),
Anoscopus albifrons (L.), Balclutha punctata (F.) and Recilia coronifer (Marsh.) have only
been found on H. mollis, whereas the more heliophilous or thermophilous species Eury-
bregma nigrolineata Scott, Mocydia crocea (H.-S.), Verdanus abdominalis (F.) and Arthaldeus
pascuellus (Fall.) clearly favour H. lanatus. The relationship of further species such as
Criomorphus albomarginatus Curt., Javesella dubia (Kbm.), Zyginidia scutellaris (H.-S.),
Macustus grisescens (Zett.) and Euscelis spp. has yet to be confirmed.
(vi) Common reed (Phragmites australis)
Phragmites australis forms large monospecific stands in temporarily or permanently
flooded sites, mainly near standing or slowly running water. It also occurs in saltmarsh-
es, straw meadows and fen woods. This plant is outstanding in harboring the maxi-
mum number of strictly monophagous Auchenorrhyncha species in central Europe, and
probably also on a larger geographical scale (see Fig. 14). In Germany alone these in-
clude the leafhoppers Paralimnus phragmitis (Boh.), P. rotundiceps (Leth.), Calamotettix
taeniatus (Horv.), and the planthoppers Delphax crassicornis (Panz.), D. pulchellus (Curt.),
Euides basilinea (Germ.), Chloriona unicolor (H.-S.), Ch. sicula Mats., Ch. dorsata Edw., Ch.
glaucescens Fieb., Ch. smaragdula (Stål), Ch. stenoptera (Fl.), Ch. vasconica Rib. and Pen-
tastiridius leporinus (L.). The latter species perhaps feeds on other plant species at least
in the nymphal stage. It has also been suggested that Ederranus discolor (J. Shlb.) lives on
reed, but host reports for this species are contradictory. Additional feeders occur in
more southern parts of Europe and Scandinavia (see Ossiannilsson 1978; Drosopoulos
et al. 1983; Nickel et al. 2002a). Most stands are infested by several species, with up to 7
monophages that have been recorded to live syntopically.
The decisive factors controlling the distribution of these species are probably water
table and salinity. For instance Chloriona glaucescens Fieb. is restricted to saltmarshes (see
312 Utilization of plant resources

also Hildebrandt 1999), whereas Ch. stenoptera (Fl.) was found to occur only on freshwa-
ter sites. Most of the remaining species on reed, however, apparently prefer a low or inter-
mediate salinity. The effects of water level and fooding regime are less evident, simply
because sampling can be done most easily in dry sites. On the other hand, occasional
sweep-netting from bridges has so far only revealed single individuals of Chloriona sma-
ragdula (Stål). At least for epigeic species (see below), such habitats are probably not suit-
able. Delphax crassicornis (Panz.) and D. pulchellus (Curt.) both prefer low and loosely grow-
ing reed stands often, but not exclusively, in salt marshes. The latter species is the only
reed specialist living in mown meadows. Finally, Delphax crassicornis (Panz.), Chloriona
stenoptera (Fl.), Ch. smaragdula (Stål) and Euides basilinea (Germ.) may all be found in shady
undergrowth of fen woods and floodplain forests. Therefore, reed dominance and height,
disturbances and shade are likely to play a further role in insect distribution.
Auchenorrhyncha communities on reed were subject to several studies on the biology
of closely related species. Thus Strübing (1960) found differences in oviposition sites and
life cycles between Euides basilinea (Germ.) and Chloriona smaragdula (Stål). The former ovi-
posits into the stem, the latter into the terminal leaf sheath. Vogel (1981) studied the distri-
bution of three Chloriona species and Euides basilinea (Germ.) in middle Hessen (Schweins-
berger Moor), and could not detect any differences concerning phenology and height of
feeding and oviposition sites. Moreover, all species equally preferred the interior part of
the reed patch. However, Chloriona spp. were found to stay on the upper parts of the stem,
preferably in fresh and scrolled leaves, their younger nymphs feeding on the flowers,
whereas Euides basilinea (Germ.) moved down to the ground after feeding. Gillham & de
Vrijer (1995) analysed the calling signals of 4 Chloriona species in the Netherlands and
could demonstrate species-specific differences, although female signals of Ch. smaragdula
(Stål) and Ch. dorsata Edw. largely overlapped. The authors discuss possible implications
on the evolution and coexistence of closely related species. In reeds of near-coastal sites of
northwestern Germany Hildebrandt (1999) found the halobiotic Chloriona glaucescens Fieb.
usually separate from the other species. He suggested that flooding and salinity may be
the most important factors for the distribution of reed-dwelling Auchenorrhyncha.
Tscharntke (1999) analysed the communities of endophagous insects attacking reed in
the upper Rhine plain near Karlsruhe. All 26 species found were monophagous. Their
occurrence and distribution were affected by stem diameter (which in turn depended on
nutrient and moisture content of the soil) and the occurrence of ‘primary attackers’, i.e.
species destroying internodes or the apical meristem (particularly Archanara geminipunc-
tata Haw., a noctuid moth), thus altering the resource availability for other groups, re-
ferred to as ‘secondary attackers’. Accordingly, since planthoppers of the genus Chloriona
spp. feed on reed flowers in the nymphal stage (see above), stem-boring moths are likely
to influence the abundance of planthoppers and perhaps other Auchenorrhyncha species.
Numerous studies have been made on the ecology of reed stands, focusing on pro-
duction and decomposition, herbivores and their predators, microclimatic conditions,
bird and mammal communities and ‘die-back’ of reeds in Europe (for references see
Ostendorp 1993).
(vii) Couch-grass (Elymus)
Couch-grass (Elymus) comprises 7 species in Germany, with occasional intrageneric
hybridization. Common couch (E. repens) is widespread and abundant in all kinds of
 Poaceae 313

non-managed grasslands on damp to moderately dry and nutrient-rich soils, from the
lowlands up to the submontane belt, notably along waysides and field margins, but
also in meadows. It is also of major significance as a weed in cereal fields. Bearded
couch (E. caninus) forms tall, but slender tufts in eutrophic and shady sites in river flood
plains. Only a few stands have been sampled, with no positive records of feeding Auchen-
orrhyncha. Hairy couch (E. hispidus), Mainz couch (E. arenosus) and the recently intro-
duced blunt-flowered couch (E. obtusiflorus) are all rare or localized. Sea couch (E. atheri-
cus) is a pioneer grass on open sand along the North Sea coast and in the upper Rhine
valley. Sand couch (E. farctus) is confined to primary dunes along the North and Baltic
Sea coast. None of these species has been specifically sampled for Auchenorrhyncha,
but at least the two latter are likely to be exploited. Thus, all feeding records of the
genus Elymus refer exclusively to E. repens, which has been studied extensively in many
parts of the country by both direct search as well as suction sampling.
28 Auchenorrhyncha species were found, 18 of which definitely breed. Thus, com-
mon couch is one of the most-favoured plant in central Europe (see Fig. 14), with many
stands revealing high insect densities. There is not a single monophage, but many spe-
cies show a clear preference for this grass, notably Eurybregma nigrolineata Scott, Dicra-
notropis hamata (Boh.), Athysanus argentarius Metc., Artianus interstitialis (Germ.), Erras-
tunus ocellaris (Fall.), Enantiocephalus cornutus (H.-S.) and Mocuellus collinus (Boh.). Apart
from these, Stiroma bicarinata (H.-S.), Javesella pellucida (F.), Anoscopus flavostriatus (Don.),
A. serratulae (F.), Deltocephalus pulicaris (Fall.), Elymana sulphurella (Zett.), Mocydia crocea
(H.-S.), Macustus grisescens (Zett.), Streptanus aemulans (Kbm.), Psammotettix kolosvaren-
sis (Mats.) and Arthaldeus pascuellus (Fall.) were found to breed. Among uncertain records,
the psammophilous Psammotettix sabulicola (Curt.) is noteworthy.
(viii) Cock’s-foot (Dactylis)
This genus comprises only two species in Germany, both preferring nutrient-rich and
damp substrates. Common cock’s-foot (D. glomerata) is one of the most important mead-
ow grasses, occurring up to the alpine tree line, whereas wood cock’s-foot (D. polygama)
is a plant of deciduous forests confined to lower altitudes. Sampling is easy due to the
tufty growth. The data base is extensive on D. glomerata, but only small on D. polygama.
Altogether, 15 confirmed plus 7 unconfirmed species have been recorded on the former,
but there is only a single possible feeder on the latter. Thus, at least D. glomerata is of
major importance as Auchenorrhyncha host in grassland sites.
The unconfirmed feeding record on D. polygama refers to Stenocranus minutus (F.),
which is otherwise monophagous on D. glomerata, as is Cicadula persimilis (Edw.). Other
feeders, most of which are 1st degree oligophagous, include Dicranotropis hamata (Boh.),
Javesella pellucida (F.), Zyginidia scutellaris (H.-S.), Athysanus argentarius Metc., Arthaldeus
pascuellus (Fall.), less commonly also Eurybregma nigrolineata Scott, Stiroma bicarinata
(H.-S.), Anoscopus flavostriatus (Don.), A. serratulae (F.), Deltocephalus pulicaris (Fall.),
Macustus grisescens (Zett.) and Errastunus ocellaris (Fall.).
(ix) Meadow-grass (Poa)
Meadow-grass (Poa) is a diverse and widespread group, which is present in most
grassland habitats of central Europe, but also in deciduous forests. Specific sampling is
often difficult since many species grow in loosely scattered stands among other grasses.
Thus, the overall data base is only moderate and includes a large proportion of uncon-
314 Utilization of plant resources

firmed feeding records. On the other hand, the total Auchenorrhyncha species number
is probably high, particularly since a number of more stenotopic Poa species have not
been studied at all, notably the montane or alpine broad-leaved, prostrate, alpine and
hybrid meadow-grass (P. chaixii, P. supina, P. alpina, P. hybrida), and the xerophilous Baden
meadow-grass (P. badensis).
Smooth meadow-grass (P. pratensis) s.l. is a common and widespread plant of pastures,
meadows, abandoned grassland, open forests and waysides, usually on moderate-
ly eutrophic, damp to moderately dry substrates, occurring in at least three subspecies,
which have only partially been distinguished for this study (see below). Due to its loose-
ly-cespitose growth, often scattered among taller grasses, sampled insects are difficult to
assign. Thus, only 6 out of the 17 recorded Auchenorrhyncha species can be considered as
confirmed feeders, notably the strictly monophagous species Ribautodelphax albostriata
(Fieb.), Xanthodelphax flaveola (Fl.), Rhopalopyx preyssleri (H.-S.) and Mocydiopsis intermedia
Rem., as well as the polyphagous Javesella pellucida (F.) and Cercopis sanguinolenta (Scop.),
the latter only in the adult stage. There is also evidence of Muirodelphax aubei (Perr.) and
Jassargus flori (Fieb.) to be 1st degree monophagous, but this needs to be confirmed by
more field data or laboratory rearing. Only R. albostriata (Fieb.) is fairly eurytopic and
occurs in high frequency, whereas the remaining species show specific requirements re-
garding moisture, insolation and perhaps soil (see chapter 4). Further less specific Auchen-
orrhyncha include Graphocraerus ventralis (Fall.), and probably Doratura stylata (Boh.),
Euscelis spp. and Psammotettix spp. It should be noted that most species, including those
suspected to be monophagous, were found on the ssp. P. p. angustifolia.
Annual meadow-grass (P. annua) prefers grazed or trodden habitats such as pastures
and ruderal sites, often along paths, where seeds can germinate easily. It is often peren-
nial; thus, hibernation of Auchenorrhyncha is possible, although not yet documented.
The situation of Auchenorrhyncha assessment is rather similar as in P. pratensis regard-
ing the total numbers, with altogether 15 species recorded. But only two of these, nota-
bly Javesella pellucida (F.) and Deltocephalus pulicaris (Fall.), are known to breed, and there
are no monophages.
Rough meadow-grass (P. trivialis), often occurring along with P. pratensis and P. annua,
but with a general preference for moist sites, and also tolerant of shade, is common on
meadows, along shores of fresh-water bodies and in fen forests. The data base includes
only a few suction samples from southern Lower Saxony. The eurytopic Javesella pelluci-
da (F.) and Arthaldeus pascuellus (Fall.) are the only breeding species found so far, al-
though there is evidence for Javesella forcipata (Boh.), Anoscopus serratulae (F.), Deltoceph-
alus pulicaris (Fall.) and Streptanus aemulans (Kbm.).
Wood meadow-grass (P. nemoralis) is essentially a plant of oak-hornbeam and beech
forests on damp, moderately eutrophic soils. It often forms monospecific stands in less
shaded sites, which are easy to sample. So far, only 4 planthopper species have been
found to breed, all of which are oligophagous grass-feeders, notably Eurysa lineata (Perr.),
Eurysella brunnea (Mel.), Stiroma affinis Fieb. and Hyledelphax elegantula (Boh.). Hardya
tenuis (Germ.) is perhaps a further associated species.
Flattened meadow-grass (Poa compressa) is a pioneer species on open, sandy to stony,
rather dry substrates, preferring gravel pits, waysides and walls. Only a single site with
dominant stands in southern Germany has been sampled, revealing large populations of
 Poaceae 315

three oligophagous grass-feeding species, namely Neophilaenus campestris (Fall.), Doratura

homophyla (Fl.) and Mocuellus collinus (Boh.), but a host association has yet to be confirmed.
Swamp meadow-grass (P. palustris), growing in wet and rather eutrophic sites, has
been sampled only in a few occasions. It is probably the host of the rarely found plant-
hopper Criomorphus williamsi China.
(x) Tor-grass (Brachypodium pinnatum)
Brachypodium pinnatum which forms dense, often monospecific swards on calcareous
hillsides and in open forests, is another grass heavily infested by Auchenorrhyncha,
with 11 confirmed plus 8 unconfirmed species. Ribautodelphax pungens (Rib.), Neophilae-
nus albipennis (F.), Adarrus multinotatus (Boh.) and A. bellevoyei (Put.) are all strictly
monophagous, and all except the localized A. bellevoyei (Put.) are widespread and fairly
abundant. Furthermore, tor-grass is an important host of Eurysella brunnea (Mel.), Stiro-
ma affinis Fieb., Mocydia crocea (H.-S.), Elymana sulphurella (Zett.), Allygidius spp. and
some others. Cook (1996) presented a detailed study of the guilds of Brachypodium pin-
natum and other grasses in a British chalk grassland, including choice experiments in
the field and in the greenhouse. Accordingly, additional species on this grass include
Zyginidia scutellaris (H.-S.), Turrutus socialis (Fl.), Verdanus abdominalis (F.) and also Planaph-
rodes bifasciata (L.).
Rock tor-grass (B. rupestre) is considered to belong to the same species group or as a
subspecies. It is confined to limestone regions of the Alps and the Swabian and Franco-
nian Jura, and has not been sampled separately. Surprisingly, the infestation of the closely
related false-brome (B. sylvaticum), growing in open deciduous woodland usually on
moist soils, is extremely low, despite extensive sampling. So far, not a single breeding
species has been found. There are only a few adult records of Mocydia crocea (H.-S.) and
Adarrus multinotatus (Boh.), which may have been vagrants. Recently, it has been dem-
onstrated, that the endophytic fungus Epichloë sylvatica Leuchtmann & Schardl (Asco-
mycota, Clavicipitaceae), which lives specifically in B. sylvaticum, may reduce noctuid
herbivory through fungal alkaloids (Brem & Leuchtmann 2001). In contrast, B. pinnatum
is only rarely infected by the closely related E. typhina (Pers.) Tul. & C. Tul. (Choke) (A.
Leuchtmann, pers. comm.). Further studies of this subject may reveal interesting in-
sights into host relationships of phytophages.
(xi) False Oatgrass (Arrhenatherum elatius)
Arrhenatherum elatius has been a dominant grass on many lowland meadows, but is
now less common due to intensive fertilization. Its occurrence in central Europe is per-
haps only caused by human influence, except on some montane rocky slopes in south-
ern parts (Körber-Grohne 1990). Much specific sampling has been done, but due to the
loosely tufted physiognomy, particularly after cutting, it is often difficult to decide if
sampled insects really feed on this grass. Thus, the total number of recorded Auchenor-
rhyncha species is 26, and includes results of field and greenhouse choice experiments
conducted in Britain (Cook 1996), but only 9 of these can be considered as confirmed
feeders or breeders in Germany.
Megadelphax sordidula (Stål) is the only species, which has been found exclusively on
this plant, although it has been reported from Scandinavia to transmit cereal viruses on
Triticum and Avena. The rare Criomorphus williamsi China is perhaps another host spe-
cialist, but more field data are needed to confirm its plant relationship. Oligo- and poly-
316 Utilization of plant resources

phages include Stiroma bicarinata (H.-S.), Javesella pellucida (F.), Anoscopus serratulae (F.),
Athysanus argentarius Metc., Streptanus aemulans (Kbm.) and Arthaldeus pascuellus (Fall.).
Furthermore, A. elatius is an important food plant of adults of both Cercopis sanguinolen-
ta (Scop.) and C. vulnerata Rossi; the latter has even been found to feed in the nymphal
stage (Biedermann 1998a; Mauri 1982). In England, according to Badmin (pers. comm.),
A. elatius is the main host of Neophilaenus campestris (Fall.) and Anoscopus duffieldi (Le
Q.), which is perhaps conspecific with A. alpinus (W.Wg.). Future field studies will prob-
ably reveal more breeding species. Thus records of adult Eurysella brunnea (Mel.), Lao-
delphax striatella (Fall.), Dicranotropis hamata (Boh.), further species of Anoscopus, Mocy-
dia crocea (H.-S.), Conosanus obsoletus (Kbm.), Psammotettix spp. and Errastunus ocellaris
(Fall.) need verification.
(xii) Purple Moor-grass (Molinia caerulea)
This is a widespread and dominant plant of temporarily wet, more or less acidic
soils, mainly straw meadows, drained or disturbed bogs, heaths and open forests. Spe-
cific sampling is facilitated by its tussock growth, so its Auchenorrhyncha fauna is well
known. However, M. arundinacea, which is usually considered as a subspecies, and pre-
fers more basic sites, such as pine forests and calcareous fens in the Alps and their
foreland, has not been distinguished. Altogether, 16 Auchenorrhyncha species have been
recorded, 8 of which are confirmed breeders.
Muellerianella extrusa (Scott), Jassargus sursumflexus (Then), Litemixia pulchripennis
Asche, Deltocephalus maculiceps Boh., and perhaps also Xanthodelphax xantha Vilb. and
Sorhoanus schmidti (W.Wg.), were found as monophagous feeders. The former two are
widespread and fairly common, but L. pulchripennis Asche is apparently restricted to
open pine forests of the upper Rhine plain, and is otherwise only known from France.
D. maculiceps Boh. is another western European species. In Germany it is confined to
heathland mires and intermediate bogs of the northern plains. Both X. xantha Vilb. and
S. schmidti (W.Wg.) are also very localized, but their host association is not yet proven.
Oligophagous species include Hyledelphax elegantula (Boh.), Delphacodes venosus (Germ.)
and Recilia coronifer (Marsh.). Javesella pellucida (F.) is the only polyphage; records of
Neophilaenus lineatus (L.), Paluda flaveola (Boh.), Mocydia crocea (H.-S.), Macustus grise-
scens (Zett.) and others need further confirmation.
(xiii) Brome (Bromus)
Bromus is a rather diverse group, but most central European species are introduced
or annual plants preferring cereal fields and ruderal habitats. None of these species is
apparently attacked by Auchenorrhyncha, except perhaps by Javesella pellucida (F.) and
some species of Macrosteles and Psammotettix. Upright brome (Bromus erectus), Hungar-
ian and hairy brome (B. inermis and B. ramosus, incl. B. benekenii) are the only native
perennials. The data base is only moderate and has been supported by field and green-
house data from Britain (Cook 1996). Altogether, 20 Auchenorrhyncha species have been
recorded on Bromus, 15 of which need further confirmation.
All 5 confirmed species records refer to Bromus erectus exclusively, which is a dominant
plant on dry or temporarily dry grassland, preferring basic and non-grazed sites. Ditrop-
sis flavipes (Sign.) is strictly monophagous, whereas Mocydia crocea (H.-S.), Arocephalus
longiceps (Kbm.), Turrutus socialis (Fl.) and Diplocolenus bohemani (Zett.) are oligophagous
grass feeders. In addition, sweep-net samples from almost monospecific swards contained
 Poaceae 317

numbers of Javesella pellucida (F.), Zyginidia scutellaris (H.-S.), Psammotettix helvolus (Kbm.)
and Jassargus obtusivalvis (Kbm.), but breeding of these species has yet to be confirmed.
Furthermore, Cook (1996) reared Hyledelphax elegantula (Boh.), Planaphrodes bifasciata (L.),
Euscelis incisus (Kbm.) and Verdanus abdominalis (F.) from this grass in Britain.
B. inermis is a common grass of waysides and disturbed grassland, mainly in regions
of pronounced summer drought in southern and eastern Germany. Feeding records of
Auchenorrhynha are all uncertain and include 10 oligo- or polyphagous species, e.g.
Javesella pellucida (F.), Zyginidia scutellaris (H.-S.), Psammotettix alienus (Dhlb.), Mocydia
crocea (H.-S.) and Athyanus argentarius Metc. From Russia the leafhopper Emeljanovianus
medius (M. & R.), which is widespread in southern and eastern parts of Europe, is re-
ported from this grass (Emelyanov 1964a). According to Mityaev (1971), the leafhopper
Mogangina bromi Em. lives on B. inermis in Kazakhstan.
B. ramosus is a grass of shady forest undergrowth on seepy and nutrient soils. Due to
its loose growth, it is probably a rather unfavourable insect host. Moreover, it is difficult
to sample. Auchenorrhyncha records so far were all negative.
(xiv) Saltmarsh-grass (Puccinellia)
Puccinellia is a group of exclusively halobiotic grasses mainly found in Asia, but also
in other continents. Reflexed saltmarsh-grass (P. distans) is a dominating plant on most
inland saltmarshes, but it is also found along the coast as well as in sites suffering heavy
anthropogenic immissions (e.g. along waysides and in the vicinity of east German phos-
phate works). Macrosteles sordidipennis (Stål) has been found to breed, being perhaps
strictly monophagous. Further feeders include Javesella pellucida (F.), Macrosteles sexno-
tatus (Fall.), Psammotettix kolosvarensis (Mats.) and perhaps Javesella salina (Hpt.), Anosco-
pus albiger (Germ.), A. serratulae (F.), Macrosteles viridigriseus (Edw.), Psammotettix alienus
(Dhlb.), Ps. putoni (Then) and Ps. confinis (Dhlb.). Common saltmarsh-grass (Puccinellia
maritima) is a dominating grass on coastal salt meadows near the North and Baltic Sea.
Unlike the preceding species, it is not found inland. Psammotettix putoni (Then) is the
most commonly associated leafhopper, although there are a few records from inland
salt marshes lacking this grass. Thus, Ps. putoni (Then) is probably 2nd degree monoph-
agous. Other host specialists, at least on generic level, perhaps include Anoscopus limico-
la (Edw.) and Macrosteles sordidipennis (Stål), whereas Psammotettix confinis (Dhlb.) is a
rather oligophagous grass feeder. Both mud and northern saltmarsh-grass (P. limosa and
P. capillaris) are rare and localized, the former on east German inland saltmarshes, the
latter along the coast. Their insect fauna has not been studied so far. Altogether, 4 con-
firmed plus 7 unconfirmed feeders are now known from the genus Puccinellia.
(xv) Reed-grass (Phalaris arundinacea)
Reed-grass (Phalaris arundinacea), which forms tall stands on wet or flooded, usually
eutrophic soils with strongly fluctuating water level, has a distinct Auchenorrhyncha
guild of 4 exclusive species, notably Paraliburnia adela (Fl.), Balclutha rhenana W.Wg.,
Erzaleus metrius (Fl.) and Stenocranus major (Kbm.), although the latter perhaps feeds
also on Calamagrostis epigejos during autumn migration. There are no further records of
breeding species, but adults of Neophilaenus lineatus (L.), Paluda flaveola (Boh.) and some
others have been found. Thus, the total species number on Ph. arundinacea is rather low,
but with a high proportion of 1st degree monophages. Unlike in many other plants, all
host specialists are equally widespread and often occur syntopically.
318 Utilization of plant resources

(xvi) Grass genera with at least one host specialist

Vernal-grass (Anthoxanthum) is represented by three species. Annual vernal-grass (A.
aristatum) is a psammophilous plant of the northern plains, growing in open pastures
and fallow fields. Alpine vernal-grass (A. alpinum) is restricted to alpine and subalpine
grassland of the Alps, the Bavarian Forest and the Harz Mountains. Both species have
not been sampled for Auchenorrhyncha. In contrast, sweet vernal-grass (A. odoratum) is
common and widespread, though rarely dominant, in nutrient-poor meadows, pastures
and open forests. It has been sampled in many locations, but was only found to be
attacked by the monophagous and rare Ribautodelphax angulosa (Rib.) and perhaps by
the oligophagous Graphocraerus ventralis (Fall.).
Oat-grass (Helictotrichon) (sensu Conert 1998) is a diverse group comprising c. 80 spe-
cies, most of which occur in temperate Eurasia, with 4 species found in Germany. Varie-
gated and alpine oat-grass (H. versicolor and H. parlatorei) are both rare and confined to
the subalpine and alpine belt of Upper Bavaria and the Allgäu. Downy oat-grass (H.
pubescens) is an inconspicuous, loosely tufted species of rather nutrient-poor meadows
and pastures, usually on damp to moderately dry soils. Specific sampling is very diffi-
cult due to its scattered and low growth. Thus, there are no specific Auchenorrhyncha
records from Germany, although many sites have been sampled where the grass oc-
curred. In Britain, however, field choice experiments as well as subsequent rearings
were conducted with some positive results (Cook 1996). Meadow oat-grass (H. prat-
ense) is a locally dominant plant of rather dry pastures (never found on meadows, as the
name suggests), with strongholds in limestone and basalt regions, although not actual-
ly calciphilous. Due to its low-growing tufts and isolated stems, insect records are diffi-
cult to assign. Direct sampling in some southern Lower Saxonian locations has revealed
large populations of the localized Doratura horvathi W.Wg., which is apparently strictly
monophagous. Further search may yield more species records.
Most hair-grasses (Koeleria) are xerophilous, low-growing and loosely tufted spe-
cies mainly of dry pastures and dunes. 5 species occur in Germany. Dune hair-grass
(K. arenaria) is a localized western European endemic usually found on grey dunes
of the East Frisian Islands. The only German location of rock hair-grass (K. vallesi-
ana) is a single xerothermic site in the upper Rhine plain. Pyramidal hair-grass (K.
pyramidata) is the most widespread and common representative of its genus in cen-
tral Europe, being absent only from most parts of the northern plains, but one of the
dominating grasses of grazed calcareous hillsides in middle and southern parts.
Occasionally, it is also found in open xerothermic pine forests, on inland dunes or in
temporarily dry meadows. Crested hair-grass (K. macrantha) shows similar habitat
requirements, but is more confined to dry and basic substrates in more continental
climates. For all 4 species there are no specific data on the associated Auchenor-
rhyncha fauna due to their low and inconspicuous growth. Glaucous hair-grass (Ko-
eleria glauca) forms small tufts on dry and open, usually basic sands of northeast
Germany and the northern upper Rhine plain. The few sampled sites have revealed
the strictly monophagous Zyginidia viaduensis (W.Wg.). Furthermore, the only Ger-
man record of the rare Psammotettix angulatus (Then) is from a site with dominating
K. glauca, and from Russia, the closely related, perhaps even conspecific Ps. koeleriae
Zachv. is reported to live on this grass. Further breeding species include Neophilae-
 Poaceae 319

nus minor (Kbm.), Zyginidia scutellaris (H.-S.), perhaps also Arocephalus languidus (Fl.)
and the rare Pinumius areatus (Stål).
Grey hair-grass (Corynephorus canescens) is the only central European representative
of its genus, growing as a perennial pioneer plant on open desalinated and decalcified
sand, mainly in the north German plain and in inland dune areas further south. It is
often dominant along with Festuca ovina s.l., forming low and compact tufts. In these
sites, the two grasses are usually the only Auchenorrhyncha hosts. Both the monopha-
gous Psammotettix excisus (Mats.) and the oligophagous Neophilaenus minor (Kbm.) of-
ten occur in high densities. It has also been proposed, that Ps. albomarginatus W.Wg. and
Recilia horvathi (Then) are associated with this grass, but this was based upon a few
records only and needs confirmation. Recent evidence suggests, that the former may
breed on Agrostis vinealis, the latter on Festuca ovina s.l. Furthermore, feeding has been
recorded in Zyginidia scutellaris (H.-S.) and perhaps Psammotettix nodosus (Rib.) and Ps.
confinis (Dhlb.). In contrast, most published records of Arocephalus punctum (Fl.) and Ps.
pallidinervis (Dhlb.) probably refer to grey-leaved subspecies of Festuca ovina.
Marram Grass (Ammophila arenaria) is the most important stabiliser on base-rich pri-
mary and secondary dunes along the coasts of the North and Baltic Sea. Locally, it has
also been sown on aeolian sand along the larger rivers inland. Although there are many
data on the insect fauna of coastal dunes (e.g. Niedringhaus 1991), the specific guild on
A. arenaria is only partially known due to syntopic occurrence of less conspicuous grasses,
such as Elymus spp., Festuca rubra and x Calammophila baltica. Both Gravesteiniella boldi
(Scott) and Psammotettix maritimus (Perr.) are monophagous on marram grass, and Do-
ratura littoralis Kuntze has also been reported, but both its taxonomic status and feeding
ecology require confirmation. Further abundant species of Ammophila-dominated dunes
include Javesella pellucida (F.), Neophilaenus lineatus (L.) and Psammotettix sabulicola (Curt.),
but definite feeding of these species is not yet proven. Hybrid marram (x Calammophila
baltica) is a locally common hybrid between Ammophila arenaria and Calamagrostis epige-
jos, growing on more stabilized coastal dunes. It has not yet been sampled, but may be
an interesting subject for the study of a plant hybrid with both parent species harbour-
ing specific and diverse insect guilds (see above).
Three species of moor-grass (Sesleria) are found in Germany, all with strongholds in
mountainous regions, usually on steep and rocky hillsides. Two-rowed moor-grass (S.
disticha) and moraine moor-grass (S. ovata) are both rare and confined to the high-alpine
belt of the Allgäu or Berchtesgaden Alps. There are no data available on their insect
fauna. Blue moor-grass (S. albicans) is the dominating plant of steep slopes on stony or
rocky substrates, mainly in the Alps and the limestone and gypsum regions. It occurs in
alpine grassland as well as in sun-exposed pine or beech forests of lower altitudes,
extending northward to the northern edge of the Mittelgebirge. The Auchenorrhyncha
data base is extensive, although inaccurate regarding host association. Chlorionidea flava
P. Löw is clearly monophagous. Zyginidia mocsaryi (Horv.) and Arocephalus languidus
(Fl.) are both often found to breed in large numbers, but the former lives additionally on
Festuca altissima at least in south Germany and the Czech Republic, whereas the latter
also lives on Stipa capillata and perhaps Koeleria spp. and others. The alpine Zyginidia
franzi (W.Wg.) is locally common in Sesleria swards, but its host plant is yet unknown.
Suction sampling may reveal further species.
320 Utilization of plant resources

Quaking grass (Briza media) is the only native representative of its genus. It is a rather
low-growing plant of open low-productivity grassland, mainly on grazed calcareous
hillsides and in peaty meadows. Specific sampling is difficult due to its scattered and
low growth and has seldom been done. Psammotettix cephalotes (H.-S.) is strictly mono-
phagous, and there is an unconfirmed record of Javesella salina (Hpt.). Further positive
records from field choice and rearing experiments in Britain include Anoscopus albifrons
(L.), Zyginidia scutellaris (H.-S.), Euscelis incisus (Kbm.), Psammotettix confinis (Dhlb.),
Turrutus socialis (Fl.) and Verdanus abdominalis (F.) (Cook 1996).
Sweet-grass (Glyceria) includes a number of hygrophilous species often found in nu-
trient-rich, temporarily flooded habitats. Both floating and reed sweet-grass (G. fluitans
and G. maxima) are infested by the 2nd degree monophagous Struebingianella lugubrina
(Boh.). The oligophagous Javesella obscurella (Boh.) occasionally lives on G. fluitans. It
has also been suggested, that Ederranus discolor (J. Shlb.) lives on G. maxima, but the host
plant in the only two known German locations has not been ascertained. None of the
remaining species of Glyceria has been sampled. At least plicate and small sweet-grass
(G. notata and G. declinata) are likely to be infested, because they are both widespread,
though more sporadic. Wood and striated sweet-grass (G. nemoralis and G. striata) are
both very rare, the latter is a neophyte originating from North America.
Lyme-grass (Leymus arenarius) is another important stabilizer of coastal dunes of the
North and Baltic Sea. It is often found along with Ammophila arenaria, but with prefer-
ence of more saline sites, where Ammophila is not yet established or dominant. Locally,
it is also sown on inland dunes. Although the dune communities of Auchenorrhyncha
are generally well known, specific sampling has only rarely been done. Unkanodes excisa
(Mel.) is clearly monophagous on Leymus arenarius, but records of Neophilaenus lineatus
(L.) and Psammotettix sabulicola (Curt.) need verification.
Mat-grass (Nardus stricta) is widespread and dominant in low-productivity grassland
on siliceous or peaty substrates, with strongholds in submontane to alpine altitudes, but
also growing in lowlands along bog margins and in peaty meadows. Its yield is extremely
poor. Thus, most sites have been converted into more productive meadows by fertilizing
during recent decades. Many Nardus-dominated grasslands have been studied, and much
is known on their Auchenorrhyncha species composition (e.g. Bornholdt 1996; Nikusch
1976; Walter 1998). But specific associations with the grass yet completely unknown, al-
though there are many uncertain records. Dicranotropis divergens Kbm., Xanthodelphax fla-
veola (Fl.), Neophilaenus lineatus (L.), Streptanus marginatus (Kbm.), Arocephalus punctum
(Fl.), the acidophilic form of Psammotettix helvolus (Kbm.), Ps. nardeti Rem., Turrutus socia-
lis (Fl.), Jassargus pseudocellaris (Fl.), Verdanus bensoni (China) and Arthaldeus pascuellus (Fall.)
are all locally common in Nardus grassland, but this does not necessarily imply a host
association, which, in many cases, is only documented for Festuca ovina and F. rubra. In
fact, most samples of monospecific Nardus patches did not yield a single individual of
plant- and leafhoppers. Thus, specific suction sampling should be carried out to estimate
the significance of Nardus as a hostplant.
The few German records of both Toya propinqua (Fieb.) and Recilia schmidtgeni (W.Wg.)
have all been gathered on Bermuda-grass (Cynodon dactylon). Along with Digitaria san-
guinalis and the cultivated Zea mays, this grass is one of the very few C4 plants of central
Europe and restricted to sunny and trodden or otherwise disturbed habitats mainly
 Poaceae 321

along the upper Rhine. T. propinqua (Fieb.) may emerge as a further example of regional
monophagy, since in Mediterranean regions it is rather eurytopic and reported to feed
on various grasses (Drosopoulos et al. 1983). In contrast, R. schmidtgeni (W.Wg.) is also
known as stenophagous from Sicily (Guglielmino 1991). Very recent suction sampling
of Cynodon near Darmstadt revealed further feeders on this grass, notably Athysanus
argentarius Metc. (M. Wallus, pers. comm.).
(xvii) Grasses exploited by oligophages and polyphages only
The remaining grasses do not harbour any host specialists. They include all the cere-
als, like maize (Zea mays), oats (Avena spp.), barley (Hordeum spp.), rye (Secale cereale)
and wheat (Triticum spp.). In most parts of central Europe, reported damage by virus
transmission and feeding is probably mainly caused by temporary influx of Javesella
pellucida (F.), Macrosteles spp. and Psammotettix spp., with reproduction at most in the
second generation or in winter cereals (see chapter 5.5). The same is probably true for
all the remaining annual grasses studied so far, notably bristle-grass (Setaria spp.), fin-
ger-grass (Digitaria spp.) and cockspur (Echinochloa crus-galli).
The number of indigenous, perennial grasses, which are apparently not attacked by
stenophagous Auchenorrhyncha is low. However, in almost all relevant cases the data
base is rather small, or the grasses are difficult to sample specifically due to their phys-
iognomy.
Heath grass (Danthonia decumbens), for instance, which is a widespread, but sporadic
and inconspicuous plant of rather dry and acidic grassland, has only rarely been sam-
pled, revealing only small numbers of host generalists such as Javesella pellucida (F.),
Macrosteles laevis (Rib.) and Deltocephalus pulicaris (Fall.). However, Sardius argus (Marsh.),
which is probably a specialist, has been suspected to live on this grass in the Nether-
lands (Cobben & Rozeboom 1983).
Eight native species of feathergrass (Stipa spp., incl. Achnatherum) occur in Germany,
most of which are confined to xerothermic habitats in southern or eastern parts. Only a
few Thuringian stands of hairy feathergrass (St. capillata) have been studied, where
adults of Jassidaeus lugubris (Sign.), Arocephalus languidus (Fl.) and perhaps Praganus
hofferi (Dlab.) were found. However, in the Baltic states, the Czech Republic, Hungary
or eastern Austria, several central Asian steppe species associated with Stipa show relic
populations, e.g. Dorycephalus baeri Kouch., Paradorydium paradoxum (H.-S.), Dudanus
pallidus Dlab., Mongolojassus sibiricus Horv., Chloothea zonata Em. and Henschia acuta P. Löw
(Emelyanov 1964a; Mityaev 1971; Nast 1987). Some of these may turn up in the eastern
half of Germany and may be found by suction sampling.
5 native species of melick (Melica) are known from Germany. Wood, mountain and
variegated melick (M. uniflora, M. nutans and M. picta) are mainly found in deciduous or
coniferous forests. Hairy and Transsylvanian melick (M. ciliata and M. transsilvanica) oc-
cur on rocky slopes and near old walls in dry grassland. The data base on members of this
group is small, and there are only a few records of oligophagous Auchenorrhyncha from
M. uniflora and M. ciliata. The former occasionally harbours Eurysa lineata (Perr.) and Stiroma
affinis Fieb., the latter revealed a population of Hardya tenuis (Germ.) in one occasion.
The last group comprises genera which include some of the most important meadow
grasses, namely meadow foxtail (Alopecurus pratensis), timothy (Phleum pratense), per-
ennial rye-grass (Lolium perenne) and yellow oat-grass (Trisetum flavescens). Most feed-
322 Utilization of plant resources

ing records only refer to the most abundant and eurytopic grassland Auchenorrhyncha
such as Javesella spp., Anoscopus serratulae (F.), Zyginidia scutellaris (H.-S.), Macrosteles
spp., Psammotettix spp., Errastunus ocellaris (Fall.) and Arthaldeus pascuellus (Fall.). Most
other members of these grass genera, like purple-stem, alpine and rough cat’s-tail (Phleum
phleoides, Ph. alpinum, Ph. hirsutum), orange foxtail and black grass (Alopecurus aequalis
and A. myosuroides) have not been specifically sampled. At least locally, marsh foxtail
(A. geniculatus) is the main host of Javesella obscurella (Boh.).
(xviii) Grasses without positive feeding records
Finally, there are no feeding records for the following grasses, all of which are either
annuals or rare or form only very loose stands:

Millet (Panicum spp.) Wood millet (Milium effusum)

Smooth finger-grass (Bothriochloa ischaemum) Sheath grass (Coleanthus subtilis)
Cut grass (Leersia oryzoides) Swamp grass (Scolochloa festucacea)
Love-grass (Eragrostis spp.) Whorl grass (Catabrosa aquatica)
Cord-grass (Spartina spp.) Crested dog’s-tail (Cynosurus cristatus)
Burdock grass (Tragus racemosus) Fern grass (Catapodium rigidum)
Holy grass (Hierochloe spp.) Tough grass (Sclerochloa dura)
Early sand-grass (Mibora minima) Gravel fescue (Micropyrum tenellum)
Soft-bearded oat-grass (Ventenata dubia) Vulpia fescue (Vulpia spp.)
Aira hair-grass (Aira spp.) Hard grass (Parapholis strigosa)
Silky-bent (Apera spp.) Wood barley (Hordelymus europaeus)

5.1.2.20 Further plant groups

The remaining plants exploited by Auchenorrhyncha shall be grouped in (i) woody
host plants harbouring breeding species, (ii) woody plants found to be utilized by adults
or in adjacent countries only, (iii) herbaceous plants harbouring specific feeders, and
(iv) herbaceous plants utilized by polyphages only. It should be noted, that this chapter
deals with some of the most diverse plant taxa in central Europe, such as the Brassicace-
ae, Caryophyllaceae and Scrophulariaceae, with more than 100 plant species each. Fur-
ther diverse groups almost neglected as food resources include the Chenopodiaceae,
Primulaceae, Geraniaceae, Onagraceae, Euphorbiaceae, Boraginaceae, Campanulace-
ae, Liliales, Asparagales and Orchidaceae.
(i) Remaining woody plants harbouring breeding species
Host specialists on the remaining woody plants are very few (Table 23). There are only
three monophages, notably Macropsis mulsanti (Fieb.) on sea-buckthorn (Hippophae rham-
noides), the central Asian neozoon Macropsis elaeagni Em. on cultivated Russian-olive (Elae-
agnus angustifolia) and Edwardsiana diversa (Edw.) on both dogwood (Cornus sanguinea)
and Cornelian cherry (C. mas). Oligophages include Opsius stactogalus Fieb. on both Ger-
man false tamarisk (Myricaria germanica) and cultivated tamarisks (Tamarix spp.), and
Zygina suavis R. on both common and alder buckthorn (Rhamnus cathartica and Frangula
alnus). In the last case the weak point of the classification of any diet breadth becomes
evident, since this species may be treated either as 2nd degree monophagous or 1st de-
gree oligophagous, depending on the question, whether alder buckthorn is placed into
 Further plant groups 323

Table 23. Auchenorrhyncha species utilizing further woody host plants in Germany. Explanations see Table 5.

Aesculus hippocast anum

Hippophae rhamnoides
Elaeagnus angust ifolia

Overw intering stage

My ricaria germanica
Rhamnus cat hart ica

Ligust rum v ulgare

Fraxinus excelsior
Cornus sanguinea

Sambucus nigra
Frangula alnus

Viburnum spp.
Plat anus spp.

Tamarix spp.
Vit is v inifera

D iet breadth

Substrate
C. mas
Species
M acropsis mulsanti (Fieb.) X m1 P eg Mono-
M acropsis elaeagni Em . X m1 P eg phages
Edwardsiana diversa (Ed w .) X X m2 P eg
Z ygina suavis R. x X o1 M ad Oligo-
Opsius stactogalus Fieb. X X o1 P eg phages
Edwardsiana lethierryi (Ed w .) x o2 M eg
Empoasca decipiens Paoli ? x ? ? x po M? ad
Empoasca vitis (Göthe) x x ? X X x ? x ? ? x po P ad
Fagocyba cruenta (H .-S.) x ? ? ? ? po M eg Poly-
A lnetoidia alneti (Dhlb.) ? X x ? ? po M eg phages
Fieberiella florii (Stål) ? x po P eg
Fieberiella septentrionalis W.Wg. ? x ? po P eg
Tachycixius pilosus (Ol.) A po P ny
Hyalesthes obsoletus Sign. A po P ny Vertical
A phrophora alni (Fall.) A A ? A po X eg m igrants
Centrotus cornutus (L.) A po P ny
Issus coleoptratus (F.) ? po P ny
Stictocephala bisonia K. & Y. ?* po P eg
A lebra wahlbergi (Boh.) ? ?* ? po M eg
Empoasca affinis N ast ? ? po M? ad Uncon-
Empoasca pteridis (Dhlb.) ? po M? ad ? firm ed
Edwardsiana frustrator (Ed w .) ?* ? po M eg
Edwardsiana prunicola (Ed w .) ? o2 M eg
A guriahana stellulata (Burm .) ?* po M eg
A rboridia ribauti (Oss.) ? o2 M ad
Total 1 4 2 2 2 3 1 1 1 4 1 1 3 2 1
* = after Claridge & Wilson (1981), Günthart (1980, 1987a)

the genus Frangula or Rhamnus. Edwardsiana lethierryi (Edw.) is the last species with a clear
host preference, usually favouring Acer campestre and Tilia spp., but it is occasionally also
found on the introduced horse-chestnut (Aesculus hippocastanum) originating form the
Balkan peninsula. The remaining woody hosts of Auchenorrhyncha are plane (Platanus),
grape vine (Vitis vinifera), privet (Ligustrum vulgare), elder (Sambucus nigra) and arrow-
wood (Viburnum), all only occasionally or locally exploited by host generalists, such as
Empoasca decipiens Paoli, E. vitis (Göthe), Fagocyba cruenta (H.-S.), Alnetoidia alneti (Dhlb.)
and Fieberiella spp. Finally, it should be stressed that ash (Fraxinus excelsior), a member of
the phylogenetically more derived Oleaceae is, along with beech (see Table 9), the only
climax tree of central European lowlands without any host specialist, even without 2nd
degree oligophages. Only Empoasca vitis (Göthe) nymphs have been found in single occa-
324 Utilization of plant resources

sions, and few adults of Alebra wahlbergi (Boh.) and Fagocyba cruenta (H.-S.); but these
species are among the most polyphagous typhlocybids in central Europe.
(ii) Woody plants not utilized for reproduction
Auchenorrhyncha records on the remaining woody plants are few and comprise adults
only (Table 24). Reproduction does probably not take place on any of these, and per-
haps most, if not all, records refer to vagrant individuals only, although ivy (Hedera
helix) may locally play a role as a winter food plant for Issus coleoptratus (F.) and Empoas-
ca vitis (Göthe), and currant (Ribes spp.) may be temporarily attacked by several Zygina
species in spring.
(iii) Remaining herbaceous plants utilized at least by oligophages
The remaining herbaceous plants which may constitute important host or food plants
are shown in Table 25. Members of the Urticaceae (Urtica dioica) have already been
discussed in chapter 5.1.2.5.
Yellow and white water-lily (Nuphar lutea and Nymphaea alba, both members of the
Nymphaeaceae) as well as broad-leaved pondweed (Potamogeton natans, a member of
the monocotyledonous Potamogetonaceae) have all been found to be attacked by Erotettix
cyane (Boh.). This species is remarkable, since it may feed on taxonomically much dif-
fering plant families, including both dicots and monocots. From other European coun-
tries it is furthermore reported from water-nut (Trapa natans, Trapaceae) and even from
the primitive clover fern Marsilea quadrifolia (Pteridophyta, Marsileaceae).

Table 24. Further woody plant species in Germany utilized by adult Auchenorrhyncha or in adjacent coun-
tries only. Explanations see Table 5.
Robinia pseudoacacia

Euony mus europaeus

Overw intering stage

Buxus semperv irens
Berberis v ulgaris

Sy ringa v ulgaris
Forsy t hia spp.
Lonicera spp.

D iet breadth
Juglans regia

Hedera helix
My rica gale
Ficus carica

Ribes spp.

Substrate

Species
Issus coleoptratus (F.) ? W po P ny
Empoasca vitis (Göthe) ? ? ? ? ? ? W? ? W ? po P ad Winter/
Z ygina flammigera (Geoffr.) S W? W? o1? M ad spring
Z ygina angusta Leth. S o2 M ad
Z ygina nigritarsis Rem . S (m 1) M ad
A phrophora alni (Fall.) ? po X eg
Lepyronia coleoptrata (L.) ?* po X eg
Philaenus spumarius (L.) ?* po X eg
Fagocyba cruenta (H .-S.) ? ? po M eg
A lnetoidia alneti (Dhlb.) ? ? po M eg Uncon-
Empoasca decipiens Paoli ? po M? ad firm ed
Typhlocyba quercus (F.) ? o2 M eg
Empoasca pteridis (Dhlb.) ? po M? ad ?
Fieberiella septentrionalis W.Wg. ? po P eg
Emposaca apicalis (Fl.) ?* o1? M? ad
Fieberiella florii (Stål) ? ? ? po P eg
* = after Günthart (1987a), Halkka et al. (1977), Lauterer (1980), Nuorteva (1952b), Smreczynski (1954)
Table 25. Auchenorrhyncha species utilizing further dicotyledonous herbs as main host or food plants in Germany. Explanations see Table 5.

Substrate
Overw intering stage

Epilobium angust ifolium

Hy pericum perforat um
Solanum t uberosum
Conv olv ulus arv ensis
Pot amoget on nat ans

H. maculat um
Verbascum ly chnit is

A. officinalis
Heliant hemum nummularium
Geranium sanguineum
Knaut ia dipsacifolia

P. lanceolat a
D iet breadth

Ny mphaea alba
Nuphar lut ea
Urt ica dioica
Alt haea rosea
Plant ago media

Rumex acet osella

Vincet oxicum hirundinaria

Poly gonum av iculare

P. major
Species
M acropsis scutellata (Boh.) . . X . . . . . . . . . . . . . . . . . . m1 P eg
Eupteryx calcarata Oss. . . X . . . . . . . . . . . . . . . . . . m 1? M eg
Eupteryx cyclops Mats. . . X . . . . . . . . . . . . . . . . . . m1 M eg
Eupteryx urticae (F.) . . X . . . . . . . . . . . . . . . . . . m 1? M eg
M acrosteles variatus (Fall.) . . X . . . . . . . . . . . . . . . . . . m 1? P eg
M acrosteles maculosus (Then) . . . . X . . . . . . . . . . . . . . . . m 1? P eg
Z ygina hypermaculata Rem . & H lzg. . . . . . X . . . . . . . . . . . . . . . m1 M eg Mono-
Z ygina hyperici (H .-S.) . . . . . . X . . . . . . . . . . . . . . m1 M eg phages
Batracomorphus irroratus Lew . . . . . . . . . . X . . . . . . . . . . . m 1? P eg
Sonronius binotatus (J. Shlb.) . . . . . . . . . . X . . . . . . . . . . m 2? P eg
Sonronius dahlbomi (Zett.) . . . . . . . . . . x* . . . . . . . . . . m 1? P eg
A rboridia pusilla (Rib.) . . . . . . . . . . . X . . . . . . . . . m1 M ad
M icantulina stigmatipennis (M. & R.) . . . . . . . . . . . . . . . . . . X . . m 1? M eg?
Eupteryx austriaca (Metc.) . . . . . . . . . . . . . . . . . . . X . m 1? M eg
Ophiola decumana (Kontk.) . . . X X . . . . . . . . . . . . . . . . o1 P eg
A gallia brachyptera (Boh.) . . . ?* . . . . . . . . . . . . . . . . . o2? P eg
Erotettix cyane (Boh.) x X . . . . . . . . . . . . . . . . . . X o2 P eg
Eupteryx vittata (L.) . . . . . . . . . . . . . . . . ?* . . . . o2 M eg Oligo-
Eupteryx melissae Curt. . . . . . . . x ?* . . . . . . . . . . . . o2 M eg phages
Further plant groups

Platymetopius undatus (De G.) . . . . . . . . . x* . . . . . . . . . . . o2? P eg

A naceratagallia ribauti (Oss.) . . . . . . . . . . . . . . . X x ? . . . o2? P ad
Eupteryx notata Curt. . . . . . . . . . . . . . . . ?* . ?* . . . o2 M eg
325
Table 25. (continued): 326

Epilobium angust ifolium

Substrate
Overw intering stage

Pot amoget on nat ans

Hy pericum perforat um
Geranium sanguineum
Solanum t uberosum
Conv olv ulus arv ensis

H. maculat um
Heliant hemum nummularium
Verbascum ly chnit is

A. officinalis
D iet breadth

Vincet oxicum hirundinaria

Knaut ia dipsacifolia

Ny mphaea alba
Alt haea rosea
P. lanceolat a
Plant ago media

Nuphar lut ea
Rumex acet osella
Urt ica dioica
Poly gonum av iculare
P. major
Species
Lepyronia coleoptrata (L.) . . . ?* . ?* ?* . . . ?* . . . . . ?* . . . . po X eg
Utilization of plant resources

Philaenus spumarius (L.) . . X . . . . x . . . . . . . . . . . . . po X eg

A phrodes makarovi Zachv. . . X . . . . . . . . . . . . . . . . . . po P eg
Evacanthus interruptus (L.) . . x . . . . . . . x . . . . . . . . . . po X eg Poly-
Erythria manderstjernii (Kbm .) . . . . . . . . . . . . . . . ?* . ?* . . . po M ad phages
Empoasca pteridis (Dhlb.) . . . . . . . . . . . . . X . . . . . . . po M? ad ?
Empoasca vitis (Göthe) . . . . . . . . x . . . . ? . . . . . . . po P ad
Eupteryx atropunctata (Goeze) . . ? . . . . . ?* . . . . X . . . . . . . po M eg
Eupteryx aurata (L.) . . X . . . . . x . . . . x* . . . . . . . po M eg
Hyalesthes obsoletus Sign. . . x* . . . . . . . . . . . X* . . . . . . po P ny
A phrophora alni (Fall.) . . . . . ?* ?* . . . . . . . . . . . . . . po X eg Vertical
Centrotus cornutus (L.) . . x* . . . . . . . A . x* . . . . . . . . po P ny m igrants
Platymetopius major (Kbm .) . . . . . . . . . x . . . . . . . . . . . po P eg
Utecha trivia (Germ .) . . . . . . . . . . . . . . . ?* . . . . . m 1? P ad
A gallia consobrina Curt. . . ? . . . . . . . . . . . . . . . . . . o1? P ad
A gallia brachyptera (Boh.) . . . ?* . . . . . . . . . . . . . . . . . o2? P eg
Emelyanoviana mollicula (Boh.) . . . . . . . . . . . . . . . . . . ? . . po M eg Uncon-
Empoasca decipiens Paoli . . . . . . . . ? . . . . . . . . . . . po M? ad firm ed
Lamprotettix nitidulus (F.) . . ? . . . . . . . . . . . . . . . . . . po P eg
Euscelidius schenckii (Kbm .) . . ? . . . . . . . . . . . . . . . . . . po P eg
* = after Le Quesne & Payne (1981), Morris (1972), Müller (1984a), Ossiannilsson (1981, 1983), Remane (1987, and pers. comm.), Stewart (1988), Vidano
(1959a), Weber (pers. comm.)
 Further plant groups 327

Among the Polygonaceae, both sheep’s sorrel (Rumex acetosella) and knotgrass
(Polygonum aviculare) are the main hosts of Ophiola decumana (Kontk.). The latter plant is
also the only host of the strictly monophagous Macrosteles maculosus (Then). The subge-
nus Hypericella of the otherwise arboricolous typhlocybid leafhopper genus Zygina has
specialized on herbs, notably on the Clusiaceae. In Germany there are two parapatric
species. Zygina hyperici (H.-S.) is widespread in the lowlands on perforate St John’s-
wort (Hypericum perforatum), whereas the recently described Zygina hypermaculata Rem.
& Hlzg. lives in subalpine meadows on imperforate St John’s-wort (Hypericum macula-
tum). Cultivated marsh mallow (Althaea officinalis), a member of the Malvaceae, has
been found to be the host of Eupteryx melissae Curt., at least in the second generation.
Common rock-rose (Helianthemum nummularium), belonging to the Cistaceae, is the only
known host of Batracomorphus irroratus Lew., although other species of Helianthemum
have not yet been studied. It may also be the host of nymphs of several species of Platyme-
topius, adults of which usually ascend up to the tree canopy. Among the Onagraceae,
only fireweed (Epilobium angustifolium) harbours Auchenorrhyncha feeders. Both na-
tive species of the leafhopper genus Sonronius were recorded, but their host range is not
sufficiently known. Likewise, Arboridia pusilla (Rib.) is the only leafhopper known to
attack species of the Geraniaceae in the study area, living monophagously on bloody
crane’s-bill (Geranium sanguineum), which is a locally common plant of open xerother-
mic woodland in the southern half of Germany. The typhlocybine leafhopper Hauptidia
distinguenda (Kbm.), reported from Sweden to infest other species of Geranium (Table
26) by Ossiannilsson (1981), could not yet be confirmed.
Among the Solanaceae, cultivated potato (Solanum tuberosum) is occasionally attacked
by the polyphagous typhlocybid leafhoppers Empoasca pteridis (Dhlb.), Eupteryx atro-
punctata (Goeze) and E. aurata (L.). Field bindweed (Convolvulus arvensis, Convolvu-
laceae) has been found to be an important food plant of the endogeic nymphs of the
cixiid planthopper Hyalesthes obsoletus Sign., whereas plantain (notably Plantago lan-
ceolata, and locally, P. major), a member of the Plantaginaceae, is the main (if not the
only) host of Anaceratagallia ribauti (Oss.), although in the laboratory, this leafhopper
has been reared on various species of Lamiaceae, Fabaceae and even Scrophulariaceae
(see Günthart 1987a). Further feeders on Plantago have been reported from other coun-
tries and include Utecha trivia (Germ.), Erythria manderstjernii (Kbm.), Eupteryx vittata
(L.) and E. notata Curt. White mullein (Verbascum lychnitis), utilized by Micantulina stig-
matipennis (M. & R.), is the only known example of a member of the Scrophulariaceae
being attacked by a monophagous leafhopper species. A few generalists, however, have
been found on various species of Verbascum. Finally, teasel-leaved knautia (Knautia dip-
sacifolia), a member of the Dipsacaceae, is apparently the only host of Eupteryx austriaca
(Metc.), at least in montane and subalpine altitudes of the Alps and the Black Forest.
(iv) Remaining herbs utilized only by generalists
Table 26 shows the remaining species accepted as host or food plants, although only
generalists or uncertain feeding records are involved. Remarkable is the fact, that Car-
damine pratensis and Silene flos-cuculi, the German names of which (‘Wiesen-
Schaumkraut’, ‘Kuckucks-Lichtnelke’) refer to the abundant spittle masses of Philaenus
spumarius (L.) in spring, are among the only members of their families (Brassicaceae,
Caryophyllaceae) exploited by Auchenorrhyncha.
Table 26. Auchenorrhyncha species utilizing further dicotyledonous herbs in Germany. Explanations see Table 5. 328

Overw intering stage

Substrate

Cardamine prat ensis

Verbascum t hapsus

Humulus lupulus
Ly simachia v ulgaris
Geranium robert ianum
Solanum nigrum
Caly st egia sepium
Succisa prat ensis

Bet a v ulgaris
G. pusillum
Sambucus ebulus
D iet breadth

Pariet aria judaica

Lagenaria siceraria
Cardaria draba
Valerianella locust a
Valeriana dioica

Silene flos-cuculi
Echium v ulgare

Viola spp.
Veronica spp.
Galium spp.
Species
Empoasca pteridis (Dhlb.) . . x . . . . . . . . . . . . . . . ? . . po M? ad ?
Eupteryx atropunctata (Goeze) . . x* . . . . . . . . . . . . ? . . . . . po M eg
Philaenus spumarius (L.) . . . X X . . . x . . . . . ? . X . . ? . po X eg
Utilization of plant resources

Emelyanoviana mollicula (Boh.) . ?* . . . . . . . . . . . . . ? . . . . . po M eg Oligo-/

Lepyronia coleoptrata (L.) . . . . . . . . . . . . . . . . ?* . . . . po X eg Polyphages
Eupteryx aurata (L.) ?* . . . . . . . . . . . . . . . . . . . . po M eg
A naceratagallia ribauti (Oss.) . . . . . . . . . . . . . . ?* . . . . . . o2? P ad
Eupteryx vittata (L.) . . . . . . . . . . . . . . ?* . . . . x . o2 M eg
Hyalesthes obsoletus Sign. . . . . . . . . . . . x* x* . . . . . . . . po P ny Vertical
Cercopis sanguinolenta (Scop.) . . . . . . . x* . . . . . . . . x* . . . . po X ny m igrants
A phrophora alni (Fall.) . . . . . ?* . . . . . . . . . . ?* . . . . po X eg
Utecha trivia (Germ .) . . . . . . . . . . . . . ?* . . . . . . . po? P ad
Hauptidia distinguenda (Kbm .) . . . . . . ? . . ?* ?* . . . . . . . . . . m 1? M ad
Hauptidia provincialis (Rib.) . ?* . . . . . . . . . . . . . ?* . . . . . po? M ad ?
Eupteryx urticae (F.) . ?* . . . . . . . . . . . . . . . . . . . m 1? M eg Uncon-
Eupteryx austriaca (Metc.) . . . . . . . . . . . . . . . . . . . . ?* m 1? M eg firm ed
Evacanthus acuminatus (F.) . . . . . . . . . . . . . . . ? . . . ? . po X eg
Eupteryx origani Zachv. . . . . . . . . . . . . . . ?* . . . . . . m 1? M eg
Emposaca apicalis (Fl.) . . . . . . . . . . . . . . . . . . ? . . o1? M? ad
* = after Biedermann (1998a), Billen (pers. comm.), Cerutti (1939a), Haupt (1935), Kuntze (1937), Le Quesne & Payne (1981), Ossiannilsson (1981),
Schrameyer (pers. comm.), Wagner (1935), Weber (pers. comm.)
 Major plant clades 329

5.1.3 Differences between plant taxa

5.1.3.1 Major plant clades

The distribution of the utilization of 7 major plant clades by Auchenorrhyncha in
Germany is highly uneven, with strong preferences of some groups (Table 27). In gener-
al, dicotyledonous and monocotyledonous angiosperms are by far the most-favoured
plants, holding more than 90% of the Auchenorrhyncha species total. Among these
groups, it is particularly the graminoid monocots (Poaceae, Cyperaceae and Juncaceae,
but not the Typhaceae) and the higher dicots, or tricolpates (eudicots) sensu Judd et al.
(1999), which account for almost all feeding records. Even within these, the distribution
is rather non-uniform (see below). In contrast, the primitive dicots (“non-monocot paleo-
herbs” and the “magnoliid complex”) and the non-graminoid monocots are only uti-
lized by very few broadly polyphagous species (included in row “more than one”).
Utilization of bryophytes (mosses) is not documented, although it has been suggest-
ed in a few cases of epigeic or tyrphobiotic species. Only 13 species (equalling 2%) are
specialised on fungi, pteridophytes or gymnosperms. In none of these, there is any ev-
idence of a phylogenetically old relationship, since all non-angiosperm feeders belong
to more advanced Auchenorrhyncha taxa, most of which predominantly feed on more
recent plant groups such as the Poaceae and Lamiaceae. In Achilidae, which largely
feed on fungi in decaying wood (i.e. organisms depending, in turn, on higher plants)
during the nymphal stage, adults are believed to suck on vascular plants. Likewise, the
primitive dicots, comprising non-monocot paleoherbs and magnoliids (see Judd et al.
1999) are neglected, except by the polyphagous Erotettix cyane (Boh.) (see below). On
the other hand, in Germany these plant groups include only the Nymphaeaceae, Cer-
atophyllaceae and Aristolochiaceae, with altogether only 8 species.
Fungi
Pteridophyta
Unknown

Several Gymnospermae

Graminoid
Higher dicots monocots

Fig. 4: Utilization of major plant clades by Auchenorrhyncha species in Germany (S Auchenorrhyncha = 620)
330 Utilization of plant resources

In the remaining 7% of the Auchenorrhyncha species, food plants belong to more

than one clade, or are unknown. Further species utilize non-angiosperms only (i) casu-
ally, e.g. some pronounced generalists such as Philaenus spumarius (L.), or (ii) in winter,
e.g. some idiocerine and typhlocybine leafhoppers, which breed on deciduous woody
dicots, or (iii) in the adult stage, e.g. Haematoloma dorsatum (Ahr.) and Aphrophora corti-
cea Germ., which are associated with Pinus, but frequently feed on angiosperms in the
nymphal stage. The deltocephaline leafhopper Erotettix cyane (Boh.) is remarkable in
feeding on a few plant species belonging to almost all major groups, notably the non-
monocot paleoherbs, non-graminoid monocots, pteridophytes and higher dicots (the
two latter in other countries).
Thus, it becomes evident that feeding on primitive plants is not likewise a primitive
trait of some Auchenorrhyncha groups, but has evolved secondarily and repeatedly by
host shift from both advanced dicots or monocots. Such host shifts have occurred in
single species, like Ditropis pteridis (Spin.), Javesella stali (Metc.) and Eupteryx filicum
(Newm.), the relatives of which live on grasses, sedges or Lamiaceae, and is also report-
ed from aphids (Eastop 1973). In other cases, such as the Achilidae, the nymphs of which
are mycophagous, host associations may have phylogenetic implications, since they are
shared only with the closely related Derbidae, and hence, can be interpreted as synapo-
morphic (see Bourgoin et al. 1997). Furthermore, host shifts between major plant clades
may perhaps be facilitated by initial acceptance of pteridophytes as additional hosts by
polyphages such as Philaenus spumarius (L.) or of coniferous winter hosts, for instance
in Zygina.
In general, sap-sucking on plants probably dates back as far as the Carboniferous
(Strong et al. 1984) and is perhaps even the oldest mode of herbivory, whereas chewing,
mining and gall-making are believed to have evolved later (Shear 1991). Hemiptera-
like insects are known from not later than the Permian (Hennig 1969). Thus, it is rather
surprising, that fungi and pteridophytes as primitive plants are not attacked by diversi-

Table 27. Species numbers of plants and Auchenorrhyncha of major plant clades in Germany.

Plants Auchenorrhyncha
Fulgorom orpha Cicad om orpha Total
Plant group n % n % n % n %
Fungi ? ? 2a 1.4 0 0 2 0.3
Pterid ophyta 76 2.6 2 1.4 2 0.4 4 0.6
Gym nosperm ae 12 0.4 1 0,7 6 1.3 7 1.1
N on-m onocot paleoherbs + m agnoliid s* 8 0.3 0 0 0 0 0 0
Gram inoid m onocots 424 14.6 108 74.5 169d 35.6 277 44.7
N on-gram inoid m onocots 222 7.7 0 0 0 0 0 0
H igher Dicotyled onae 2155 74.4 18 12.4 270c 56.8 289 46.6
More than one – – 4b 2.8 24e,f 5.1 27 4.4
Unknow n – – 10 6.9 4 0.8 14 2.3
Total 2897** 100** 145 100 475 100 620 100
a
= adults perhaps on vascular pants; b = incl. 2 occasional feeders on gymnosperms; c = incl. at least 13
species found on gymnosperms in winter; d = incl. 2 species found on gymnosperms in winter; e = incl. 2
species monophagous on gymnosperms as adults; f = incl. 3 polyphages also found on pteridophytes; * =
sensu Judd et al. (1999); ** = only vascular plants counted.
 Plant families 331

fied Auchenorrhyncha guilds, which in turn comprise primitive taxa. Instead, pterido-
phytes seem to have largely succeeded in getting rid of their herbivores, although a few
more derived species of both plant- and leafhoppers have managed to recolonize some
ferns and horsetails. Even in Achilidae, the host shift towards fungi must be interpreted
as a relatively recent phenomenon, since these organisms depend on dead parts of higher
plants. At least both species of Cixidia occurring in Germany are ecologically confined
to deciduous trees or pine, respectively. Details on their nymphal feeding, however, are
unknown.
This pattern of rare and only secondary utilization of non-angiosperms seems to
be of more general nature, and is supported by Wilson et al. (1994), who compiled
plant records of Fulgoromorpha groups. Accordingly, out of 1143 records gathered
worldwide, only 3% were from pteridophytes, and 4% from gymnosperms. In none
of the major planthopper clades there was any evidence for primitive feeding on
these groups. Moreover, the authors could not detect any correlation of planthopper
evolution with certain plant groups, except a slight tendency of primitive families
to feed more on monocots, whereas advanced families favour dicots. However, there
was also a tendency in primitive families to feed underground or under bark, which
often involves feeding on roots or fungi. This idea was adopted by Sorensen et al.
(1995), who concluded that “it seems probable that early fulgoromorphans initially
evolved to feed on roots and fungal hyphae, which exist in subterranean/semisub-
terranean niches”. On the other hand, it can be objected, that such life habits are
linked with a derived morphology of the female ovipositor typical for all fulgoro-
morphans except the Cixiidae and Delphacidae (see Bourgoin 1993). Altogether, in
view of the apparent potential of rapid host shift among strongly differing plant
groups and the occurrence of polyphagy in almost all major Auchenorrhyncha
groups, it seems uncertain, whether conclusions on higher phylogenetics can be
drawn from feeding relationships.
An only moderate correlation of plant and phytophage evolution, with frequent ac-
quisition of new host groups was also suggested for aphids (Eastop 1978). Evidence of
insect-plant relationships dating back as far as the late Cretaceous was recently provid-
ed by Sequeira & Farrell (2001), who studied DNA sequences of scolytid beetles living
on the Australian and South American conifer Araucaria.

5.1.3.2 Plant families

An overview of the Auchenorrhyncha of the most important vascular plant taxa in
Germany and some of their life history traits is presented in Table 28. Considered plant
taxa harbour at least 5 Auchenorrhyncha species or comprise at least 100 plant species.
Differences in selected parameters are shown in Fig. 5, Fig. 6, Fig. 7 and Fig. 8.
As in major plant clades, Auchenorrhyncha infestation is rather uneven (Fig. 5), with
roughly two third of the species total feeding on only 6 families, notably the Poaceae,
Cyperaceae, Fagaceae, Betulaceae, Salicaceae and Rosaceae. Thus, almost 70% of the
Auchenorrhyncha species attack less than 20% of the plant species. All the remaining
families are of negligible significance as food plants, and are – if at all – attacked only by
extremely polyphagous species such as Philaenus spumarius (L.), Lepyronia coleoptrata
Table 28. Auchenorrhyncha species numbers of important angiosperm taxa and selected characteristics of their life histories 332

Scrophulariaceae

Ranunculaceae
Fagaceae
Betulaceae
Caryophyllaceae
Fabaceae
Juncaceae

Pinaceae
Ulm aceae
Urticaceae
Tiliaceae
Salicaceae
Brassicaceae
Ericaceae
Rosaceae
Aceraceae
Apiaceae
Lam iaceae
Asteraceae
Cyperaceae
Poaceae

Param eter
Species total
Plants 8 81 4 6 6 9 119 2 32 139 29 124 126 6 93 82 130 457 47 144 225
Auchenorrhyncha 10a 7 17 11 42 61 1 10 66 2 12 47 21 17 4 26 4 30 9 74 211bc
Diet bread th
1st d egree m onophagous 6 1 4 5 4 15 0 0 30 0 3 10 2 5 0 5 1 6 0 30 98
Utilization of plant resources

2nd d egree m onophagous 1 0 5 0 8 9 0 1 26 0 1 3 1 3 1 2 0 4 4 30 20

1st d egree oligophagous 1 0 0 0 0 3 0 0 0 0 2 8 7 0 0 7 0 2 0 5 68
2nd d egree oligophagous 2 1 1 0 6 11 0 2 2 0 3 8 5 1 0 4 0 3 3 1 4
polyphagous 0 5 7 6 24 23 1 7 8 2 3 18 6 8 3 8 3 15 2 8 14
Ratios
S Auchenorrhyncha/ S Plants 1,3 0,1 4,3 1,8 7 6,8 0 5 2,1 0 0,4 0,4 0,2 2,8 0 0,3 1,3 0,1 0,2 0,5 0,9
S Specialists*/ S Generalists** 4 0,2 1,1 0,8 0,4 1 0 0,1 6 0 1 0,8 0,9 0,9 0,3 1 0,3 0,7 0,8 7,2 10
Substrate
Phloem 6 2 6 5 22 26 0 4 44 0 6 17 16 8 0 3 0 13 7 63 192
Xylem 2 2 0 2 0 2 1 0 4 2 3 4 3 0 1 3 1 5 1 5 13
Mesophyll 1 3 11 4 19 33 0 6 18 0 3 26 2 9 3 20 3 12 1 6 6
Fungi 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Overw intering stage
Egg 3 2 15 9 26 45 1 6 48 1 7 27 15 10 4 21 4 21 6 60 123
N ym ph 7 3 1 2 12 9 0 2 4 1 3 8 2 2 0 2 0 4 3 9 66
Ad ult 0 2 1 0 4 7 0 2 14 0 2 12 4 5 0 3 0 5 0 5 21
Vertical m igrants 4 2 2 2 17 18 0 1 8 1 3 8 3 2 0 2 1 5 0 0 0
a
* = 1st and 2nd degree monophages plus 1st degree oligophages, ** = 2nd degree oligophages plus polyphages; = excluding overwintering species; b =
diet breadth unknown in 7 species; c = 1 species probably semivoltine, overwintering both as nymph and adult
 Plant families 333

(L.), Aphrophora alni (Fall.) and Empoasca vitis (Göthe). These plant groups include, for
instance, the Papaveraceae, Caryophyllaceae, Chenopodiaceae, Amaranthaceae, Saxi-
fragaceae, Euphorbiaceae, Violaceae, Brassicaceae, Primulaceae, Boraginaceae, Genti-
anaceae, Campanulaceae, Liliaceae and Orchidaceae. Most of them are also largely avoid-
ed by Sternorryncha (Eastop 1978).
Relating the number of Auchenorrhyncha species to the number of plant species (Fig.
6), highest values are clearly found in the Fagaceae, Betulaceae, Tiliaceae, Ulmaceae
and Aceraceae. The Salicaceae, Urticaceae, and Pinaceae (the latter excluding overwin-
tering Auchenorrhyncha species) are in an intermediate position, whereas values are
lowest (> 1) in the remaining groups (including the Poaceae, Cyperaceae, Lamiaceae,
Rosaceae and Ericaceae). Thus, the ratio of species numbers Auchenorrhyncha : plants
is high in most families of woody plants, except the Rosaceae (which comprise numer-
ous herbs and introduced ornamental shrubs), Salicaceae (which comprise a number of
low-growing alpine shrubs) and Pinaceae (which comprise some little studied montane
species), but relatively low in the Poaceae and Cyperaceae. Moreover, large-sized and
structurally complex woody plants tend to harbour more congeneric insect herbivores,
whereas the guilds on more small-sized and simple graminoids rather consist of mem-
bers of different genera.
Reed (Phragmites australis), which occupies an intermediate position in size, and which
forms large, monospecific stands with distinct microclimatic conditions, is quite excep-
tional in holding the highest number of 1st degree monophages among all central Euro-
pean, and perhaps, western Palearctic plants (see section 5.1.2.19). A parallel is found in
the dicot Urtica dioica. Like reed, this species can be considered as an ‘ecosystem engi-
neer’ and is outstanding among herbaceous plants in being utilized by a diverse and
specific insect guild (see section 5.1.2.5). Regarding diversity, specificity and taxonomic
composition of their herbivore guilds, these species both rather resemble woody plants.
Presumably due to their structural complexity, they offer diverse spatial niches, includ-
ing leaves, stem and base, each of which is utilized by some insect species.
Thus, two major modes of radiation in Auchenorrhyncha can be postulated. The first
is facilitated by a small number of large-sized and structurally complex host species
(either individuals or stands), which promote speciation on the plant by specialising in
microniches such as leaves, twigs, bark, stem base, differentially sun-exposed parts,
etc.; the second mode implies speciation by a shift Setween numerous smaller host plants.
Examples for speciation on the same host are found in the genera Eurhadina on oak,
Oncopsis on birch, the Idiocerinae on willows and poplars, Chloriona on reed, Eupteryx
on nettle as well as the Nearctic species of Erythroneura on Platanus, which have been
subject to several studies of coexistence of closely related species (Ross 1957; McClure
& Price 1975, 1976). Examples for the second mode are many, such as Kelisia, Xanthodel-
phax, Muellerianella, Ribautodelphax, Wagneriala, Doratura, Rhopalopyx, Cicadula, Mocydi-
opsis, Streptanus, Psammotettix and Cosmotettix. It is mostly those genera in which spe-
cies can only be distinguished by the study of the genitalia, but many of which can be
readily identified in the field by their host plant. In these groups, species differentiation
must have occurred only relatively recently due to their close morphological similari-
ties. It should also be noted that parapatric speciation, i.e. speciation promoted by geo-
graphic isolation, is unlikely to play a role in many, if not most, of the genera men-
334 Utilization of plant resources

250

200
S Auchenorrhyncha

150

100

50

0
e

ae

ae
ae

oa e
ae
e

ae
e

r ia e
e
ae

e
e

ae
e

ae

n c ae
os ae
ric a e

ba e
ea

ea

ea

ea
yp e a
a
ea
a
a

Fa ea
ce

ce
ce
ce
ce
ce
ce

ce
hu c e
ce

J u ce
e
e
ce

ac

ac

ac

ac
ac
ac

ac

ac
ac
lia
la
ga
ca
la

ia
na

ra

er
yll

pi
ic

er
lm

ic
tu
cu

cr am
Ti

ce
rti

la

P
Fa
Pi

al

A
ss
ph

st
Be
U

R
un

A
S

A
ra
yo

C
an

op
B
ar
R

S
Fig. 5: Species numbers of Auchenorrhyncha guilds on the most important vascular plant families in Ger-
many (excluding overwintering species on Pinaceae). Only plant families with more than 5 Auchenorrhyn-
cha species or more than 100 plant species are shown.

7
S Auchenorrhyncha / S Plants

0
lia e
ae

ae
e
ae

ae

oa e
ae
i c ae

ria e
e
ae
e

e
e

ae
ae

n c ae
os ae
ric a e

ba e
a

ea

ea
a

yp e a
ea

ph c e a
ea

Fa ea
ce
ce

ce

ce
ce
ce

ce
ra a ce

hu c e

ce
J u ce
e
ce

ac

ac
c

ac
ac

ac

ac
ac
la
ga

a
ca

ra

ia
a

la

a
ul

er
pi
ic
lm

er
yl
in

cr am
cu

Ti

ce
rti

Fa

la

P
al
et

A
ss
P

st
U

R
un

A
S
B

A
yo

C
op
an

B
ar
R

Fig. 6: Ratios of Auchenorrhyncha species number : plant species number per plant family in the most
important plant families in Germany (excluding overwintering species on Pinaceae). Only plant families
with more than 5 Auchenorrhyncha species or more than 100 plant species are shown.
 Plant families 335

1 00

90

80
S Auchenorrhyncha (%)

70

60

50

40

30

20

10

ae
ae

e
al eae
e
lm a e

ca e

ae

ae
t u ae

ae

oa e
ae

e
o s ae
r ic a e

ba e
ga e
c u ae

ae
L a ce a
ea

ss ea

ea
J u c ea
a
ea
a

ce
ce
Be c e

ce

ce
ce

ce

ce
ce

e
e

C a ce
ac

c
c

ac
ac
y o l ac

ac
ac

ac

ri a
la

a
lia

ra
la

ia
pi

er
ic

er
ic
in

nc
yl

m
Ti

ce

la
rti

P
Fa

Fa

st
ph
P

yp
U

R
un

hu
U

A
S

A
ra

op
an

B
ar

cr
R

S
1st degree 2nd degree 1st degree 2nd degree poly-
monophagous monophagous oligophagous oligophagous phagous
Fig. 7: Diet breadth of Auchenorrhyncha guilds on vascular plant families in Germany (excluding overwin-
tering species on Pinaceae). Only plant families with more than 5 Auchenorrhyncha species or more than
100 plant species are shown.

100

90

80
S Auchenorrhyncha (%)

70

60

50

40

30

20

10

0
lia e
al e a e

ae
e

oa e
ae
lm a e

ae
i c ae

r ia e
e
t u ae

e
ca e

ga e

ae
c u ae

e
os ae
ric a e

ba e
ea

ea

ea
ea

yp e a
ea

ea
a

ea
a
ce

ce
ce

Be c e
ce

ce

ce
ce
e
e
u n ce

e
ac

ac

ac
c

ac
ac

y o lac

ac
ac

ac
ac
la

ia
ra
na

er
yll

pi
ic

nc
er
cr am
Ti

ce
rti

Fa

la

P
Fa
Pi

A
ss
ph

st

Ju
U

hu
U

A
S

A
ra

C
an

op
B
ar
R

Phloem Xylem Mesophyll Fungi

Fig. 8: Auchenorrhyncha food resources of vascular plant families in Germany (excluding overwintering
species on Pinaceae). Only plant families with more than 5 Auchenorrhyncha species or more than 100
plant species are shown.
336 Utilization of plant resources

tioned, since there are numerous examples of largely overlapping or even identical geo-
graphic ranges of closely related species, e.g. in Xanthodelphax, Oncopsis, Wagneriala,
Eurhadina, Doratura, Rhopalopyx, Cicadula and the Psammotettix nododus group. Thus, it
can be concluded that taxonomic plant diversity on the one hand, and plant size and
structural complexity on the other, may differentially enhance phytophage speciation.
Interestingly, such examples can be found both in Fulgoromorpha and Cicadomorpha.
Relatively high values (> 0.5) of the ratio Auchenorrhyncha species : plant species are
also found in the Hippocastanaceae, Dennstaedtiaceae, Taxaceae, Elaeagnaceae, Arali-
aceae, Platanaceae, Myricaceae, Tamaricaceae, Cupressaceae, Cornaceae, Rhamnaceae,
Oleaceae and Equisetaceae, but the overall importance of these families as Auchenor-
rhyncha hosts is much lower due to their low numbers of plant species.
Diet breadth of Auchenorrhyncha guilds of different plant families is shown in Fig. 7.
Accordingly, the proportions of monophages (1st and 2nd degree) are highest on the
Salicaceae, Cyperaceae and Pinaceae, and lowest (≤25%) in Ranunculaceae, Caryophyl-
laceae, Brassicaceae, Fabaceae, Apiaceae, Scrophulariaceae and Tiliaceae. Most remain-
ing groups of broad-leaved trees show intermediate values, notably the Ulmaceae, Fa-
gaceae, Betulaceae and Aceraceae. This is also true in the Poaceae, which are further
characterised by a large proportion of 1st degree oligophages. Thus, regarding specific-
ity up to plant family level, the grasses are inhabited by the highest proportion of spe-
cialists, surpassing even the Cyperaceae, Salicaceae and Pinaceae.
In the two latter groups, the driving force for phytophage specialisation may simply
be the more isolated systematic position, which is combined with distinct chemical com-
pounds such as phenolic heterosides (salicin, populin) in the Salicaceae, and specific
ethereal oils and resins in the Pinaceae. Similarly, the Poaceae and Cyperaceae are phy-
logenetically also remote from the remaining groups (see Judd et al. 1999), but are thought
to be largely defended physically by epidermal silica crystals (Frohne & Jensen 1998).
The degree of host specificity among ectophagous insect feeders on graminoids has
long been thought to be low, and statements such as ‘on grasses’ in most identification
books (e.g. Ossiannilsson 1978, 1983) directly or indirectly implied a rather broad host
range. Prestidge & McNeill (1983a, 1983b) accentuated this opinion after a case study
carried out on sown grassland plots of an early successional stage in Britain, arguing
that temporal and spatial variation of nitrogen content may be the main factor for host
choice in Auchenorrhyncha. Such habitats, however, a priori select for mobile and polyph-
agous generalists and are unlikely to be quickly colonized by specialists (Novotný 1994a,
1995). Despite the overall dominance of host specialists among graminicoles, the spe-
cies lists in these papers thus included only very few monophages (see Prestidge &
McNeill 1983a).
Cobben (1988) compiled data on host relations from western and northern Europe
and was the first to point out that even the limited reliable evidence suggests an almost
equally high proportion of 1st and 2nd degree monophages both in Delphacidae and
Cicadellidae (67 and 71%, respectively), which in turn comprise numerous graminoid
feeders. Later, Wilson et al. (1994) summarized that fulgoromorphans in general, and
Delphacidae, in particular, show a high degree of host specificity. Thus, in Auchenor-
rhyncha, and perhaps, in insects in general, the degree of host specificity of graminoid
feeders is clearly higher than previously thought.
 Plant families 337

In general, there is a positive correlation between species numbers of Auchenorrhyn-

cha and plants in the vascular plant families of Germany (Fig. 9), i.e. diverse plant fam-
ilies are exploited by more diverse guilds of Auchenorrhyncha than species-poor plant
families, but only 24% of the variation is explained (P < 0.01). There is a number of
species-rich plant families such as the Poaceae, Cyperaceae and Rosaceae, which har-
bour numerous Auchenorrhyncha species, but there are also many exceptions such as
the Brassicaceae, Caryophyllaceae, Scrophulariaceae, Asteraceae as well as the entire
non-graminoid monocots (e.g. the Orchidaceae, Liliaceae). These groups are altogether
attacked by only 31 Auchenorrhyncha species (equalling roughly 5% of the Auchenor-
rhyncha species total), although they account for almost 40% of the phytodiversity. In
conclusion, there is a general tendency for higher phytophage diversity on more di-
verse plant families, as was also shown in curculionoid and chrysomeloid beetles (e.g.
Farrell 1998). However, exceptions indicate that other factors must be strongly involved.
A second, more plausible correlation can be discerned when focusing on those plant
families, which show rather high (> 10) Auchenorrhyncha species numbers (Fig. 9). Their
common trait is a pronounced apparency in space and time, showing an exceptionally
high dominance among the vegetation all over central Europe. They include three groups:

Fig. 9: Relationship between species numbers of Auchenorrhyncha and plants in the plant families of Ger-
many. Only families with more than two Auchenorrhyncha species or more than 40 plant species are la-
belled: Ace = Aceraceae, Api = Apiaceae, Ast = Asteraceae, Bet = Betulaceae, Bor = Boraginaceae, Bra =
Brassicaceae, Cap = Caprifoliaceae, Car = Caryophyllaceae, Che = Chenopodiaceae, Cor = Cornaceae, Cup
= Cupressaceae, Cyp = Cyperaceae, D/T/E = Dennstaedtiaceae/Taxaceae/Elaeagnaceae, Eri = Ericaceae,
Fab = Fabaceae, Fag = Fagaceae, Hip = Hippocastanaceae, Jun = Juncaceae, Lam = Lamiaceae, Mal = Mal-
vaceae, Ole = Oleaceae, Ona = Onagraceae, Orc = Orchidaceae, Pol = Polygonaceae, Pin = Pinaceae, Poa =
Poaceae, Ran = Ranunculaceae, Rha = Rhamnaceae, Ros = Rosaceae, Rub = Rubiaceae, Sal = Salicaceae, Scr
= Scrophulariaceae, Sol = Solanaceae, Til = Tiliaceae, Ulm = Ulmaceae, Urt = Urticaceae.
338 Utilization of plant resources

(i) all important trees and shrubs of forests and late successional stages, notably the Sali-
caceae, Betulaceae, Rosaceae, Fagaceae, Ulmaceae, Aceraceae, Tiliaceae, Pinaceae and even
the non-native Hippocastanaceae, (ii) grasses, sedges and rushes as the dominating plants
of non-forested habitats, and (iii) the Urticaceae, with Urtica dioica being perhaps the most
apparent herbaceous dicotyledonous plant in most of central Europe and being wide-
spread all over the Holarctic region (Meusel et al. 1964).
Hence, woody plants and graminoids, as those plant groups richest in biomass, and
thus, as the most apparent ones, are the most-favoured resources for Auchenorrhyncha.
The most striking exceptions to this rule, however, can be easily explained: among grasses
and sedges, it is just the annual species, which do harbour only small numbers of in-
sects, without any host specialists, e.g. most species of Bromus, Vulpia and Aira. Similar-
ly, among the Urticaceae, most species are either annuals or rare or introduced, except
Urtica dioica, which is perennial, native and extremely widespread and abundant all
over the Holarctic, usually forming large and monospecific stands. It alone accounts for
the strong utilization of the Urticaceae by at least 11 species of Auchenorrhyncha, whereas
the remaining species of Urtica and Parietaria are largely neglected. Also regarding the
temporal aspect of apparency, these plants clearly offer the most predictable food re-
sources and habitats for insects, since trees and shrubs reach the oldest age among plants,
and most graminoids form persistent turfs or tussocks. Moreover, their dominance dates
back far into the Tertiary, although alternations of forests and grasslands have occurred
throughout the Pleistocene (Lang 1994; Mai 1995). However, this correlation between
plant apparency and herbivore guild diversity must be considered as vague, since there
are neither exact figures on the large-scale dominance of plant taxa in Europe, nor on
changes of plant dominance through evolutionary time.
Data on plant relationships of other diverse insect groups are scarce. Most studies ei-
ther treat only small regions or species-poor groups, or they utilize different classifica-
tions of diet breadth, or they refer to plant family or genus only, or they do not distinguish
between monophagous and polyphagous feeders, and vagrants. Eastop (1973, 1978) has
compiled host plants of host-specific Sternorrhyncha on a world-wide base. Among these,
only the Aphidoidea can be directly compared with the central European Auchenorrhyn-
cha due to their more northern temperate distribution. Accordingly, aphids share the gen-
eral pattern of a strong infestation of highly apparent plants, but with a pronounced pref-
erence of Asteraceae (605 species = 17.2%) and coniferous trees (363 species = 10.3%). In
contrast, the Poaceae and Cyperaceae are attacked by only 242 and 75 host specialists (=
6.9% and 2.1%) respectively. Eastop (1973) suggests that aphids evolved either on Hama-
melidae sensu Cronquist (1968), i.e. on Fagaceae, Betulaceae, Juglandaceae, Ulmaceae and
related families, or on some extinct group of gymnosperms. For 25 German tree genera,
Brändle & Brandl (2001) found positive correlations between herbivore species numbers
and present plant abundance (based on grid occupancy), pollen abundance, and tree height.

5.2 General conclusions on food plant choice

The following conclusions can be drawn from these patterns: sap-sucking herbivores
favour spatially and temporally highly apparent plant species, notably phanerophytes
 General conclusions on food plant choice 339

and perennial graminoids, which are tall or form large, contiguous stands. In contrast,
annuals, bienniels and geophytes are largely neglected, and infestation of chamaephytes
and most non-graminoid hemicryptophytes is low to moderate. Further, primitive plants,
such as pteridophytes and gymnosperms, have only secondarily been colonized by host
shift from more advanced plant groups. The same is true for fungi, whereas bryophytes
are not exploited at all. The large majority of species feed on a few plant taxa of higher
monocots and dicots, notably the clades of the Poales, Juncales, rosids and asterids (sensu
Judd et al. 1999). Apparently, this situation differs markedly from patterns found in Aphi-
doidea and scolytid beetles. In these groups, the host relationships with coniferous trees
may date back at least as far as the late Mesozoic (Eastop 1973; Sequeira & Farrell 2001).
Considering the classical idea of plant apparency (Feeny 1976; Rhoades & Cates 1976),
spatially and temporally apparent plants or at least plant parts are likely to be defended
by digestability-reducing substances, which in turn tend to occur in high concentrations
in the mesophyll tissue, but not in phloem and xylem sap due to their large molecular size
(Raven 1983). On the other hand, toxins, which are believed to be produced only tempo-
rarily and in low concentrations to defend vulnerable and reproductive tissues, may be
too costly for a permanent protection of the entire plant cormus. Thus, plant parasites
such as phloem- and xylem-feeders exploit a permanent, rather diffuse and little defend-
ed resource. Conversely, exploited plants seemingly do not suffer serious damage, since
they dominate most terrestrial habitats. Most, if not all known instances of catastrophic
plant damages caused by Auchenorrhyncha refer to anthropogenic, and thus, rather un-
stable systems. Similarly, feeders of mesophyll parenchyma ultimately destroy tissue, but
mostly in mature leaves. Buds and fresh leaves are usually present early in the season,
when leafhopper development has not yet much proceeded. In this period, small nymphs
demand only small quantities of food, and reproductive organs such as flowers and fruits
are not damaged (although this point may need re-examination in the Empoascini, some
of which feed on stem and fruit parenchyma). Further, infestation of saplings is usually
negligible due to their small size and stochastic constraints of colonization. Tissue feeders
must, however, deal with permanently high concentrations of digestability-reducing sub-
stances, the composition of which is usually plant-specific.
In sap feeders the significance of secondary plant compounds can be deduced from
numerous plant taxa which have never been found to be infested by any Auchenorrhyn-
cha species, except perhaps by single nymphs of highly polyphagous xylem-feeders, such
as Philaenus spumarius (L.). Thus, there is not a single feeding record in 87 out of 146 plant
families occurring in Germany, and only one broadly polyphagous generalist has been
found in further 15 families. Due to the restricted assessment of food plant records in this
study, some of these may in fact be more frequently utilized by a few generalists (though
certainly in low numbers only). Nevertheless, the general pattern is a preference of rela-
tively few plant groups and an avoidance of most of the existing taxonomic plant diversi-
ty. Further evidence for the susceptibility of sap-feeding Auchenorrhyncha to secondary
plant compounds is provided by the rather specific guilds on the Salicaceae, which are
avoided by most generalists found on other woody plants (see section 5.1.2.9).
Furthermore, seasonal development and temporal persistence of plants are of major im-
portance. Only perennial plants offer constantly available resources, although differences
between long-lived phanerophytes and more short-lived herbs should be minor, if the latter
340 Utilization of plant resources

grow in contiguous stands, where the loss of single host individuals will not result in herbi-
vore extinction. Conversely, any herbivore population on annual plants will inevitably loose
its resource after one or few generations. Thus, all host specialists of the Auchenorrhyncha
fauna of central Europe (1st and 2nd degree monophages) are exclusively associated with
perennials, and the same is true for most, if not all, oligophages. Geophytes, excluding rhi-
zome grasses, are another group of host plants offering rather adverse conditions, since
many species have green parts only in the first half of spring, when temperatures are still
relatively low. As a consequence, most of them are totally avoided.

5.3 Stratification and plant architecture

Auchenorrhyncha utilize all strata of central European terrestrial ecosystems. Four layers
will be distinguished here (see Table 29): (i) the soil, with the mineral layer and organic
litter, (ii) the herb layer, comprising the soil surface and the herbaceous vegetation (in-
cluding dwarf shrubs), (iii) the shrub layer, with basally branching woods mainly below
5 m height, including young trees and some creeping plants (e.g. Rubus, Hedera), and (iv)
the tree layer, with tall woods mainly branching apically and forming canopies, including
tall-growing willow species (Salix alba and S. fragilis). The two latter layers are also re-
ferred to as canopy layer here. Occasional specimens recorded on tree saplings were not
classified as shrub dwellers. Arboricolous species, which may be found both on trees and
shrubs, were not categorized as ‘multi-layer facultatively’, but according to their main
preferences. For example, Macropsis notata (Proh.), living both on Salix triandra and S. fragilis,
was assigned to the shrub layer, since the former species is the main host. Mimallygus
lacteinervis (Kbm.) living on very low willow shoots was assigned to the herb layer.
Altogether, the most-favoured stratum is the herb layer, which is utilized by 61% of
the species total through their whole life cycle. The bulk of these feed on Poaceae and
Cyperaceae (see section 5.1.2). Comparing the two main groups, the fulgoromorphans
are more strongly bound to the herbaceous vegetation than the cicadomorphans (77%
vs. 56%). This difference is mainly due to some subgroups, notably the largely arbori-
colous Macropsinae, Idiocerinae and Typhlocybinae on the one hand, and the vertically

Table 29. Utilization of strata of the German Auchenorrhyncha fauna

Auchenorrhyncha total Fulgorom orpha Cicad om orpha

Layer n % n % n %
Tree 107 17.3 0 0 107 22.6
Shrub 63 10.2 0 0 63 13.3
H erb 377 60.8 112 77.2 265 55.9
Soil 1 0.2 1 0.7 0 0
Facultative m ulti-layer 23 3.7 10 6.9 13 2.7
Obligate m ulti-layer 49 7.9 22 15.2 26 5.5
Total 620 100 145 100 469 100
 Stratification and plant architecture 341

migrating Cixiidae, most of which perform an obligate shift from the soil to the canopy,
on the other hand.
Whereas the evolution of arboricolous leafhoppers is certainly a relatively recent
development, which must be seen in connection with the Tertiary rise of angiosperm
trees (see section 5.4.3), the soil association of most planthopper groups is subject to a
controversial debate. After Wilson et al. (1994), there is a significant tendency for prim-
itve planthopper groups to live underground or under bark as nymphs, but there is also
an evolutionary trend from an orthopteroid type of ovipositor rather associated with
piercing or sawing into plants, towards an excavator type associated with oviposition
into or onto the soil. However, Bourgoin (1993), after a study of the female genitalia of
fulgoromorphans, argues, that each of these two types is partially based on differing
morphological structures, and that morphology, and thus evolution, must be carefully
separated from function. He further suggests an evolutionary scenario based on a com-
parative analysis of habitats (as well as ant-attendance) among Hemiptera, and con-
cludes that primitive groups rather lived on the ground, and that several independent
shifts into the ground occurred (notably in the Cixiidae and Kinnaridae–Meenoplidae)
and also towards ant-attendance (Bourgoin 1997; see also Emelyanov 1987).
Hence, the move towards the tree canopy in cixiids and other fulgoromorphan groups
remained partial, covering only the adult stage, whereas oviposition takes place on the
ground and nymphs are endogeic. This is also true in cicadas and Cercopinae, which
are considered as basal cicadomorphan clades (Bourgoin 1997; Dietrich et al. 2001). In
contrast, most species of Macropsinae, Idiocerinae, Typhlocybinae and some Deltoceph-
alinae, all of which oviposit into living plant tissue, evolved purely arboricolous life
habits.
Viewed against the background of anthropogenic changes of the central European
vegetation, many herb layer species have probably been strongly favoured by the change
of woodland into pastures and meadows. In some cases, it is likely that herb layer spe-
cies have expanded their range or even invaded central Europe. According to Körber-
Grohne (1990), several grass species appeared only after the rise of agriculture and the
reduction of forests. Most of these, however, are only utilized by generalists, except
Arrhenatherum elatius and Bromus erectus, which are the exclusive host plants of Megadel-
phax sordidula (Stål) and Ditropsis flavipes (Sign.), respectively. Assuming that there was
no host shift, these species must have invaded large parts of central Europe in company
with their hosts. The highest abundances of Auchenorrhyncha ever recorded in central
Europe (up to 5000 ind./m2) have not been found in natural habitats, but in grasslands
of anthropogenic origin. In most types of forests with a closed canopy, the cover of
herbaceous vegetation is either low or dominated by geophytes, only few of which can
be utilized by Auchenorrhyncha due to their transient apparency in spring. On the oth-
er hand, some open forests with a well-developed grass layer, e.g. pine forests, cop-
piced woods, clearings, glades, and mountain forests, can support rather high densities
(e.g. Körner et al. 2001).
The canopy layer is permanently utilized by altogether 168 cicadellid species, ac-
counting for 27% of the Auchenorrhyncha total, and 36% of the cicadomorphans (see
above). Reliable field data on canopy abundance in central Europe do not exist, but
there is some indirect evidence. In a single tree-eclector in a 30-year old monospecific,
342 Utilization of plant resources

uniform stand of Populus tremula in Thuringia, c. 33,000 individuals of the idiocerine

leafhopper Populicerus laminatus (Fl.) were collected within one season. The abundance
in this site was estimated to be more than 5000 ind./m2, which is the same order of
magnitude as in grasslands (see above). In this particular case, the real abundance was
probably even much higher, since P. laminatus (Fl.) is a permanently arboricolous spe-
cies, and only a certain proportion of the total number of individuals is likely to have
fallen down due to leaf-trembling caused by wind (which is certainly an aspen pecular-
ity). More exact figures of canopy abundance may be gained by tree fogging, but this
method has only rarely been applied in European forests (see section 3.5.2).
The diversity of arboricolous leafhoppers is positively correlated with the large amount
of biomass and the structural complexity of woody plants, which facilitated sympatric
speciation without host shift (see section 5.1.3.2). Among native species, Fraxinus excel-
sior is the only one, which is almost devoid of Auchenorrhyncha. Most of the remaining
tree species harbour diverse and specific guilds (see section 5.1.2). Among the most
important host families, the ratio of species numbers Auchenorrhyncha : plants is clear-
ly highest in trees, and declines rapidly in the following order: Fagaceae > Betulaceae >
Tiliaceae > Ulmaceae > Aceraceae > Salicaceae > Urticaceae > Pinaceae > Poaceae >
Cyperaceae (see section 5.1.3.2). This pattern of higher insect species numbers on struc-
turally complex plants is also evident within the herb layer. For instance, among grass-
es, the most diverse guilds are found on tall (e.g. Phragmites), tussock- forming (e.g.
Festuca ovina, Deschampsia cespitosa) or turf-forming (e.g. Elymus repens, Calamagrostis
epigejos) species, whereas loosely growing species such as Milium effusum, Festuca gi-
gantea, Bromus ramosus, Hordelymus europaeus and Helictotrichon pubescens, are only rare-
ly attacked. Further uninfested grasses are also found in the genera Bromus, Aira, Vul-
pia, Hordeum and Eragrostis, but in these cases, architectural simplicity and small size
are combined with a therophytic growth form, resulting in a reduced temporal persis-
tence (see below).
High Auchenorrhyncha species numbers on structurally complex plants are in accor-
dance with predictions made by Lawton (1983) and Strong & Levin (1979), who demon-
strated that plant architecture is a major determinant of insect phytophage diversity,
based on data sets on various insect groups mainly from Britain and North America.
The most important components of plant architecture include size and resource diversi-
ty. For Auchenorrhyncha, as well as other homogeneous feeding guilds, the latter may
be only of minor significance, since even the smallest vascular plant stems and leaves
provide the basic resources of phloem, xylem, mesophyll and oviposition sites. Other
resources are only utilized by very few species, such as Ledra aurita (L.) and some Idio-
cerinae, which are often found on older bark.
In the data base presented here, the statement of diverse herbivore guilds on struc-
turally complex plants is clearly corroborated concerning plant family level (see Fig. 6),
but must be modified when considering plant species level. Fig. 14 shows the most
diverse guilds in the study area, with at least 18 Auchenorrhyncha species per plant
species. Seven out of 12 of these guilds live on trees or shrubs, whereas the remaining 5
live on grasses. Although maximum species numbers are found on Quercus and Betula,
species numbers on Festuca ovina and Calamagrostis epigejos are also considerable. The
degree of host specificity on the grasses is higher than on the trees, despite their lower
 Stratification and plant architecture 343

architectural complexity. Moreover, data on insects feeding on grasses are fewer com-
pared to arboricolous species. It should also be noted that almost equally high numbers
of Auchenorrhyncha species per plant species are likely to occur on tall sedges, notably
Carex nigra, C. acuta and C. acutiformis, but this is partially inferred from the inclusion of
uncertain records, which still need to be confirmed (Table 21). The shared characteristic
of all these heavily attacked graminoids is their dense and contiguous growth, which
may be comparable to the structural complexity and biomass abundance of trees.
There are 49 species (8% of the species total), equally belonging to the Fulgoromor-
pha and the Cicadomorpha, which perform obligate vertical migrations from one stra-
tum to another between certain stages of their life cycle, and which will be referred to as
vertical migrants. The idea that Tettigometridae, which have long been thought to spend
the nymphal stage in ant nests below ground is too simplified (Bourgoin 1997), and
their vertical migrations are probably not obligate.
In the Cixiidae, Cicadoidea and the Cercopinae, these movements are performed
from the soil into the vegetation, whereas in vertically migrating species of Aphrophori-
nae, Membracidae and Cicadellidae, the nymphal stage is spent in the herb layer, and
the insects move up to the canopy after emergence (see Table 30). The latter group is
taxonomically less distinct and includes members of numerous subgroups such as
Platymetopius, Idiodonus, Lamprotettix, Allygus, Allygidius, Speudotettix, Hesium, Tham-
notettix and probably Colladonus. A few species not included here, notably Idiodonus
cruentatus (Panz.) and Thamnotettix confinis (Zett.), do perform movements in lowland
woodland habitats, but not above the alpine tree line, where adults probably live on
dwarf shrubs.
In most cases, the upward movement is combined either with a change of the food
plant species, or with a change of the diet width, or both. For instance, species of Allygus
and Allygidius feed on Poaceae (and perhaps other herbaceous plants) as nymphs, but on
trees (mainly Fagaceae and Betulaceae) as adults. Many species of Zygina are highly spe-
cific during oviposition and larval development, but become oligophagous or even poly-
phagous as adults. In contrast, nymphs of Haematoloma dorsatum (Ahr.) feed on Poaceae
roots of various genera, but adult feeding and mating take place only on Pinus. Similarly,
the spittle of Aphrophora corticea Germ. is found on low plants of various families, but
adults live exclusively on Pinus and even show the typical pine bark colouration. In gen-

Table 30. Species numbers of obligate vertical migrants and their utilization of strata

Auchenorrhyncha total Fulgorom orpha Cicad om orpha

Layers n % n % n %
Soil - H erb 5 10,2 2 9,1 3 11,5
Soil - Shrub 16 32,7 13 59,1 2 7,7
Soil - Tree 8 16,3 7 31,8 1 3,8
H erb - Shrub 3 6,1 0 0 3 11,5
H erb - Tree 17 34,7 0 0 17 65,4
Total 49 100 22 100 26 100
344 Utilization of plant resources

eral, nymphal diet width and other ecological traits of vertically moving species are poor-
ly known due to identification problems and cryptic life habits. Nymphs of Hyalesthes
obsoletus Sign. and Cercopis spp. have been shown to be clearly polyphagous, but almost
nothing is known about the host plants of most remaining species except some scattered
evidence from laboratory rearings (e.g. Müller 1942) and casual field records.
The ecological significance of obligate vertical migration is poorly understood. In the
Cixiidae, Cicadoidea and Cercopinae, these movements are associated either with
nymphal overwintering (all three groups), xylem-feeding (only the two latter), large
body size (cicadas and – to a lesser extent – most Cercopinae) or prolonged develop-
ment (mainly cicadas) – see section 5.4.2. Soil temperatures may facilitate feeding and
growth also during the cold season. Enemy-free space during nymphal development in
the soil and utilization of an alternative ample resource may provide clues for a better
understanding of this life history trait. The ecological and morphological differentia-
tion of nymphal or larval stages from adulthood has been associated with the explosion
of diversity in holometabolous insects and is subject to a long and controversial discus-
sion (Hennig 1969; Kristensen 1995; Truman & Riddiford 1999; Wheeler et al. 2001).
Thus, it is interesting to note, that in Auchenorrhyncha with a similar (although less
pronounced) distinction of life stages, radiation was less dramatic than in Holometabola.
Trigonocranus emmeae Fieb. is the only species living permanently in the soil, although
macropterous individuals undertake dispersal flights. However, at least partial utiliza-
tion of the upper soil perhaps also occurs in a number of epigeic taxa such as Eupelix,
Aphrodinae and Errhomenus, which may suck on fine roots in the top soil layer.

5.4 Utilized substrates

Table 31 shows the utilization of different types of food source. The type is largely
combined with phylogenetic affiliations (see below). It should be noted, however, that
most feeding experiments focused on relatively few species of economic importance, and
that opinions on whole subgroups gained by mere generalisation may have to be revised
in future. The differential exploitation of resources of the most important plant families in
the study area is shown in Fig. 8. Important reviews on these resources have been present-
ed by Raven (1983) and Tonkyn & Whitcomb (1987) and will be discussed below.

Table 31. Substrate utilization of the Auchenorrhyncha species of Germany

 Phloem 345

5.4.1 Phloem
Phloem-feeding is found in all the fulgoromorphans, except the mycophagous Achil-
idae and, in more southern countries, the Derbidae (Wilson et al. 1994). Phloem-feeding
is also widespread among cicadomorphans, with some exceptions (see below). Hence,
it has been considered as a primitive trait (Emelyanov 1987), an opinion which is also in
agreement with more modern views of phylogenetics (Campbell et al. 1995; Sorensen et
al. 1995). The shift both towards xylem- and mesophyll-feeding requires dramatic mor-
phological and physiological adaptations and is, thus, interpreted as an apomorphic
character which evolved in only one or few lineages, respectively. Phloem-feeding is
shared with most sternorrhynchans, notably aphids, psyllids, most coccids, as well as
some heteropterans (Tonkyn & Whitcomb 1987; Novotný & Wilson 1997; White 1993;
Jacobs & Renner 1998), although many subgroups are only poorly studied.
Phloem feeding is found in more than 70% of the German species (Table 31), including
all members of the Delphacidae, Deltocephalinae and some largely arboreal groups such
as the Macropsinae and Idiocerinae, and furthermore, the Cixiidae, Tettigometridae, Mem-
bracidae, Agalliinae, Aphrodinae. Moreover, phloem-feeding Auchenorrhyncha account
for a large proportion of the phytophage fauna in many terrestrial habitats. Hence, their
functional significance may be strongly underestimated, particularly in view of their con-
stant, evenly distributed, and thus, rather discrete production of honeydew.
Highest proportions of phloem-feeding species are found on graminoids, ranging
from 80 to 90% on the Poaceae, Cyperaceae and Juncaceae, respectively (Fig. 8). Thus,
in the former two families, maximum numbers of Auchenorrhyncha species per plant
family (see Fig. 5) are combined with strong preferences for phloem sap. Among dicot-
yledonous plants, phloem-feeding prevails in the guilds of the Fabaceae and Salicaceae
with 70 to 80% of all the feeding species. In contrast, it is rare in some herb groups such
as the Ranunculaceae, Scrophulariaceae, Caryophyllaceae, Brassicaceae and Lamiace-
ae. In the remaining groups, which include most woody plants plus the Ericaceae and
Urticaceae, the proportions are intermediate.

5.4.2 Xylem
Altogether, 26 species, accounting for about 5% of the German Auchenorrhyncha
fauna, suck on xylem vessels (Table 31), although the assignment of Errhomenus bra-
chypterus Fieb. must be considered as tentative due to its unclear systematic position.
Only the cicadas, Cercopidae and Cicadellinae are involved. Examples include Philae-
nus spumarius (L.), which is certainly one of the world’s most polyphagous insect herbi-
vores, as well as some other pronounced generalists such as Lepyronia coleoptrata (L.),
Aphrophora alni (Fall.) and Cercopis spp. Xylem feeders, notably the Cicadidae and Cer-
copinae, include the only cicadomorphans with endogeic nymphal life habits, and the
former group is peculiar among all insects for the prolonged nymphal development,
lasting up to 17 years in Nearctic species (e.g. Karban 1986). A comparison of the uti-
lized resources in different plant families (Fig. 8) shows that xylem-feeding species dom-
inate on the Ranunculaceae, Ericaceae, Apiaceae, Scrophulariaceae, Caryophyllaceae
and Brassicaceae. In these families, numbers of Auchenorrhyncha species are generally
346 Utilization of plant resources

low, and the two latter groups are in fact only exploited by the highly polyphagous
Philaenus spumarius (L.). The correlation between species numbers of xylem-feeding
Auchenorrhyncha species and plants (Fig. 10) explains more than half of the variation
and is, thus, much better than in the remaining groups (see Fig. 9).
Among all animals, cicadas, cercopids and cicadelline leafhoppers are the only known
taxa that have managed to tap xylem vessels, and apart from them, only a few parasitic
plants such as Orobanche (Scrophulariaceae sensu lato) and Viscum (Loranthaceae) share the
same resource (Raven 1983). Xylem sap is present in large quantities, but its nitrogen con-
tent is much unbalanced and lowest among all living plant resources (Mattson 1980; White
1993). Moreover, due to the negative tension, more energy is needed for exploitation. Thus,
all xylem-feeding Auchenorrhyncha have a bulbous front associated with powerful dilato-
ry cibarial muscles. Novotný & Wilson (1997) demonstrated that simply for physical rea-
sons, xylem-feeding is associated with a larger body size. They also suggested that xylem
feeders tend to be polyphagous because, due to the low nutritional value of their food, a
narrow host plant range would limit the choice of the momentarily most nutritious host.
However, this view is questionable since individual movement between different plant spe-
cies has neither been documented nor is it likely, because mobility in endogeic spittlebug
and cicada nymphs is greatly reduced. Further, tension is usually lower or even positive in
root xylem; thus, the costs of epi- and endogeic feeding are lower than in apical plant parts.
In the Auchenorrhyncha fauna studied here, monophagy in xylem feeders is less
common than in other feeding groups, being confined to less than 20% of the species,

14
Poa
12
y = 0,0219x + 0,0713
S xylem feeding species

2
10 R = 0,5396***

Ran Lam
6
Cyp Ast

4 Sal Ros Fab

Eri Rub Api
2 Pol Bra
Car Scr
0
0 100 200 300 400 500
S plan ts
Fig. 10: Species numbers of xylem-feeders on the vascular plant families in Germany. Ast = Asteraceae, Bra
= Brassicaceae, Car = Caryophyllaceae, Cyp = Cyperaceae, Eri = Ericaceae, Fab = Fabaceae, Lam = Lami-
aceae, Pol = Polygonaceae, Poa = Poaceae, Ran = Ranunculaceae, Rub = Rubiaceae, Sal = Salicaceae, Scr =
Scrophulariaceae.
 Xylem 347

notably Aphrophora pectoralis Mats., A. salicina (Goeze), Neophilaenus albipennis (F.) and
probably N. infumatus (Hpt.) and N. exclamationis (Thnbg.). These species are accociated
with willows or grasses, respectively, i.e. members of plant groups, which generally
hold high proportions of host specialists (Fig. 7).
Therefore, the dominance of xylem feeders on plants largely avoided by the remain-
ing Auchenorrhyncha (Fig. 8) is not explained by preferences, but rather by a pronounced
non-selectivity, since their species numbers per plant family are positively correlated
with phytodiversity. The more diverse a plant family is, the more likely is a particular
xylem feeder to utilize any of its members, no matter what its secondary compounds
are like (Fig. 10). Further, it is evident that woody and tall-growing species do not play
a role for xylem feeders, particularly in the nymphal stage, although after emergence
relatively many species perform an obligate stratum shift from the soil or herb layer up
to tall herbs or up the canopy, e.g. Cicadetta, Cercopis, Haematoloma and Aphrophora. There
are only few host records of woody plants, even of extremely polyphagous species like
Philaenus spumarius (L.), but this is certainly a sampling artefact. Ossiannilsson (1981)
mentions spittle records on adventitious shoots of numerous trees and shrubs, which is
a further argument for the pronounced polyphagy of xylem feeders, and also for the
assumption that this feeding mode is aggravated by plant height.
The fact that feeders of xylem have successfully tapped even the most poisonous
plants (although densities on these are generally low) may be either explained by its
low concentrations or even lack of chemoprotectants, or by a strong selective force to-
wards extreme polyphagy. The latter would require a powerful and rather universal
detoxification system. However, it seems difficult to conceive, why insects should evolve
enzymatic structures for dealing with plants which they only rarely ever encounter.
Given the large quantities of potential resources and the large number of potential
host species, it is quite surprising, that not more xylem-feeding species have evolved. In
fact, during the past decade, it has been discovered that the spittlebug taxon Philaenus is
subject to a dramatic sympatric speciation in the Mediterranean region, apparently strict-
ly associated with rather unusual and toxic host plants such as Asphodelus (Asphodelace-
ae) and Eryngium (Apiaceae) (Drosopoulos & Remane 2000; see also Abdul-Nour &
Lahoud 1995). Hence, it can be assumed that only peculiar plant compounds may exert
isolating mechanisms upon Philaenus populations eventually leading to speciation.
Finally, it should be noted that, despite their low species number in central Europe,
xylem feeders, and cercopids, in particular, can potentially occur on almost every plant,
although largest numbers are usually found on plant species also favoured by phloem-
feeders. Due to the much diluted and unbalanced soluables, larger body size and local-
ly high abundance (up to 1280 ind./m2 in Michigan, USA – see Wiegert 1964), the re-
moved sap quantity per day may exceed the fresh body weight by 100-1000 times (Tonkyn
& Whitcomb 1987). From a comparative study of Solidago altissima (Asteraceae) experi-
mentally infested by Philaenus spumarius (L.), a leaf-chewing chrysomelid beetle and a
phloem-feeding aphid, Meyer (1993) concluded that the plant damage in relation to
insect mass gain was 5-6 times greater in the spittlebug than in the beetle, but not mea-
surable in the aphid. Thus, the turnover of assimilates and the functional significance of
xylem-feeders may in fact be higher than previously thought.
348 Utilization of plant resources

5.4.3 Mesophyll
Mesophyll-feeding is found only in the Typhlocybinae, most of which are believed
to be nutritionally confined to this substrate. They produce greenish excrements con-
taining chlorophyll, and secondarily lack microbial symbiotes, mycetomes as well as
a filter chamber (H.-J. Müller 1956; J. Müller 1969; Ammar 1985). However, among the
Empoascini, Empoasca decipiens Paoli and the Nearctic E. fabae (Harris) apparently feed
on both parenchyma and phloem (Günthardt & Wanner 1981; Backus 1988). The latter
species is an important pest on alfalfa and other crops, and widely known as the cause
of leaf damages called ‘hopperburn’. Mesophyll-sucking has not been reported in Em-
poasca vitis (Göthe) and Jacobiasca libyca (Brg. & Zn.), both causing hopperburn-like
symptoms in European vineyards. They are regarded as pure phloem feeders since
the tissue studies of Vidano (1963c, 1967). Hence, feeding biology in these species
(and perhaps other Empoascini as well) is in need of re-examination with electronic
monitoring as well as morphological studies of the alimentary canal, in order to re-
evaluate applied aspects as well as the phylogenetic significance of mesophyll-feed-
ing in general. This feeding mode is certainly a derived trait evolved along with the
broad leaves of woody angiosperms since the late Cretaceous (see Mai 1995). Further
apomorphic traits in Typhlocybinae include the pronounced reduction of wing vena-
tion (see Wagner 1951b), which is combined with a permanent life in the tree canopy
and thus, with an improved flight capability. Finally, exploiting advanced herbaceous
angiosperms such as graminoids, Lamiaceae and Asteraceae by higher Typhlocybi-
nae is likely to constitute a more recent, though widespread evolutionary trend, which
has independently occurred within the Erythroneurini and Typhlocybini. On the oth-
er hand, herb feeding in the more primitve Empoascini and Dikraneurini (Wagner
1951b) may rather constitute an ancestral trait.
Altogether, the species number of mesophyll feeders is more than 150, equalling
one third of the Cicadomorpha and one fourth of the Auchenorrhyncha total number.
Among these, about two third are permanently arboricolous, and most of them feed
on the leaf parenchyma of advanced dicotyledonous woody plants. For non-hemi-
pterologists, the characteristic stippling of leaves, particularly visible late in the sea-
son, is perhaps the most conspicuous phenomenon caused by Auchenorrhyncha. The
remaining species live among lower vegetation, where the majority feeds on herb
mesophyll. Compared to phloem feeders, densities are generally lower. Suction sam-
pling in grassland and forest undergrowth rarely revealed more than 500 ind./m2,
e.g. in Dikraneura variata Hardy, Notus flavipennis (Zett.) and Eupteryx spp. (Körner et
al. 2001; M. Sayer, pers. comm.). Canopy abundance of Fagocyba cruenta (H.-S.) and
overwintering Empoasca vitis (Göthe) may locally appear to be enormous, but is al-
most impossible to relate to ground surface. Thus, the functional role of mesophyll
feeders in central Europe is perhaps rather insignificant, since they neither transmit
plant pathogens nor produce honeydew. Stippling of leaves is usually accumulating
only in late summer not long before fall. In Mediterranean countries, however, severe
leaf damage leading to early fall has been observed (Drosopoulos et al. 1987).
As to plant taxa, two striking features can be discerned (Fig. 8). The first is the dom-
inance of mesophyll-feeding species on some higher dicotyledonous herb families, no-
 Fungi 349

tably the Lamiaceae, Asteraceae, Ranunculaceae, Apiaceae and Scrophulariaceae, al-

though species numbers are generally low on the last three mentioned families. It should
also be noted here, that mesophyll-feeding typhlocybids have managed to perform some
rather spectacular host shifts. Examples include Eupteryx filicum (Newm.) on Polypodi-
um and other ferns, although most congeners live on Lamiaceae. Zygina, subgenus Hy-
pericella, is the only Auchenorrhyncha taxon specialised on Hypericum, a member of the
Clusiaceae, which is otherwise only (rarely) exploited by Philaenus spumarius (L.). Most
relatives live on woody trees or shrubs (see below for further examples). The second
feature of mesophyll-feeders is their high proportion on most woody plants, notably
the Ulmaceae, Fagaceae, Betulaceae, Tiliaceae, Rosaceae and Aceraceae, which offer
enormous quantities of resources.
Mesophyll-feeding is also found in the members of the Dikraneurini, all of which
live on leaves of sedges, grasses and forbs, except the Mediterranean Liguropia juniperi
(Leth.), which is found on ornamental Chamaecyparis (Gymnospermae, Cupressaceae).
Among the Empoascini, Chlorita, Austroasca and some species of Empoasca live on leaves
of herbaceous plants, and are considered to feed on mesophyll, although no experimen-
tal studies have been conducted. The Typhlocybini are largely arboricolous on broad-
leaved trees, except Wagneripteryx germari (Zett.), which has secondarily shifted to gym-
nosperms (Pinus spp.). Further exceptions are found in Eupteryx and Edwardsiana socia-
bilis (Oss.). The latter has colonized ornamental Rosa rugosa, but originally lives on the
herb Filipendula ulmaria. The genus Eupteryx is exclusively found on herbs and is a good
example of a secondary host shift towards the herb layer in an arboricolous group.
Preferred host plants mainly belong to the rather advanced Lamiaceae and Asteraceae,
many members of which originate from semi-arid parts of southern Europe and Asia.
Similarly, among the largely arboricolous Erythroneurini, the genus Zyginidia is purely
graminicolous, whereas Zygina species of the subgenus Hypericella are specialised on
herbaceous Hypericum spp. (see above), and Z. rubrovittata (Leth.) lives on the dwarf
shrub Calluna vulgaris. Further, among the genus Arboridia, arboricolous species are found
along with feeders on shrubs, as well as on the herbaceous Geranium and Potentilla.
Finally, Hauptidia comprises species reported from a number of rather unusual food
plants, including Silene (Caryophyllaceae), Primula (Primulaceae), Digitalis (Scrophu-
lariaceae), Geranium (Geraniaceae), Galium (Rubiaceae) and Ecballium (Cucurbitaceae)
(Dworakowska 1970d; della Giustina 1989).

5.4.4 Fungi
Among Auchenorrhyncha, feeding on fungal mycelium is rather exceptional, and only
found in nymphs of the fulgoromorphan groups of the Achilidae and Derbidae (Wilson et
al. 1994). Usually, they live on decaying wood, often under bark, but details on their biol-
ogy are unknown. Adults show morphological traits of typical corticoles, i.e. their body is
dorsoventrally flattened, and their colouration resembles the bark of their preferred tree
species. As in mesophyll and xylem feeding, this type of nutrition is highly likely to have
evolved from phloem feeding, because it is peculiar to two groups treated as adelphotaxa
by Bourgoin et al. (1997), and also because their food apparently depends on higher plants.
Both groups, however, are mainly found in more southern countries (Fennah 1950; O’Brien
350 Utilization of plant resources

& Wilson 1985), and only two species of Achilidae occur in Germany. Due to their life
habits, they escape most hemipterologists, and thus, only a handful of records is known
from both species, without any data on their feeding habits.

5.5 Pests on cultivated plants

Planthoppers and leafhoppers play an important role as pests on various cultivated

crops by transmitting plant diseases as well as by direct feeding. Transmitted patho-
gens include viruses, bacteria and mycoplasma-like organisms (MLO), some of which
have not yet been properly identified (Maramorosch & Harris 1979; Nault & Rodriguez
1985). Although much research has been done on etiology and vector control, knowl-
edge of causal connections is often poor. Thus, the capability of transmitting pathogens
has been demonstrated for many species under laboratory conditions, but the real amount
of resulting crop damage has rarely been quantified. There is no doubt that some spe-
cies are serious pests of rice, maize, legumes and other crops in tropical and subtropical
latitudes. Important planthopper pests include the delphacids Nilaparvata lugens (Stål)
and Sogatella furcifera (Horv.) on rice, Peregrinus maidis (Ashmead) on maize, and Perkin-
siella species on sugarcane. Deltocephaline leafhoppers of the genera Dalbulus, Nepho-
tettix, Circulifer and Macrosteles are vectors of diseases of maize, rice and other cereals as
well as beets (Müller 1956; Nault & Rodriguez 1985; Wilson & Claridge 1991). In central
Europe, however, there are only few recent cases where damage has been proved, most
of which has occurred in cultivations of perennial plants and in greenhouses.
In vineyards of southwestern Germany the cixiid planthopper Hyalesthes obsoletus (Sign.)
has been found to transmit an MLO-associated grapevine yellows disease locally known
as ‘Vergilbungskrankheit’ (Maixner et al. 1995). More recently, Psammotettix alienus (Dhlb.)
has been assumed to cause damage in barley fields of Sachsen-Anhalt by transmission of
wheat dwarf virus (WDV) (Manurung et al. 2001). Graphocephala fennahi Young is suspect-
ed to transmit Pycnostysanus azaleae (Peck.), a fungus causing bud blast on ornamental
Rhododendron, although the etiology is not properly known and evidence is ambiguous
(Vidano et al. 1987b). Scaphoideus titanus Ball is another Nearctic species, belonging to the
Deltocephalinae and potentially being noxious to grapevines (Maixner et al. 1993). First
recorded in the Gironde area, France, in 1958, it has spread over most French viticultural
areas and has invaded northern Italy and Ticino (della Giustina 1989; Maixner et al. 1993;
Bosco et al. 1997), but so far, it has not yet reached Germany.
Direct damage by sucking plays a more important role, with most species involved
belonging to the Typhlocybinae. During recent years, Empoasca decipiens Paoli has
caused losses of cucumber and sweet pepper yields in greenhouses in the Lake Con-
stance area (Schmidt & Rupp 1997; K. Raupach, pers. comm.). The closely related
E. vitis (Göthe) has periodically become noxious in vineyards of southwestern Ger-
many (Louis & Schirra 1997; Schruft 1983; see also Günthart & Günthart 1967). More
recently, Eupteryx decemnotata Rey, E. melissae Curt. and perhaps some congeners have
attacked various officinal herbs in gardens and fields, notably sage, marsh-mallow
and balm. However, the extent of damage is usually limited. Locally, Edwardsiana ro-
sae (L.), attacks ornamental roses and is able to cause damage, but can easily be con-
 Pests on cultivated plants 351

trolled by insecticides. At the beginning of the 20th century, Tettigometra obliqua (Panz.)
was reported to have invaded fields of rye and oats, with losses of at least some plants
(Torka 1905). Nowadays, this species appears to have completely vanished from most
parts of central Europe. Similarly, Empoasca pteridis (Dhlb.) and Eupteryx atropunctata
(Goeze) are both found on potatoes, but damages are apparently exceptional, although
leaf-rolling after feeding has been confused with symptoms of virus diseases (Müller
1956). Very recently, the Mediterranean typhlocybid Hauptidia provincialis (Rib.) has
turned up in a greenhouse of Bavarian Swabia causing feeding damage on corn salad
and perennial basil.
The introduced treehopper Stictocephala bisonia Kopp & Yonke has become a pest in
Italy on grapevine and various fruit trees. After oviposition into the bark of young twigs,
the phloem stream may become interrupted, resulting in necrosis (Vidano 1963a, 1963b;
Günthart 1980). From Germany, however, no damages have been reported yet, although
the species is locally common in southwestern parts of the country.
Apart from Graphocephala fennahi Young, xylem suckers have become noxious only in
forest cultivations. Schmutterer (1953) observed a mass attack of Cicadella viridis (L.) in
nurseries of black alder, with twigs and young trees dying after extensive oviposition.
In the Netherlands and the Mediterranean region, Haematoloma dorsatum (Ahr.) is caus-
ing problems in pine forests. While nymphs live on roots of grasses, adults ascend up to
the canopy and suck on pine needles, sometimes leading to necrosis (Moraal 1996).
Kurir (1967) describes a mass occurrence of Aphrophora salicina (Goeze) in plantations of
American osier in Burgenland (eastern Austria). After oviposition, young twigs were
spoilt. However, the species identity remains unclear, since the photograph presented
in the paper shows A. alni (Fall.).
A serious problem may be caused by Metcalfa pruinosa (Say, 1830), a flatid planthopper
originating from North America, and first found in northern Italy in 1979 (Dlabola 1981).
Since then, it has spread across much of Italy and southern France, invaded Slovenia,
Styria, Czechia and Ticino and may be expected in the upper Rhine valley soon. Single
individuals have been observed on wind screens of cars being transported for more than
20 kilometers, even through tunnels. In Italy and France, this species may occur in
extremely high abundance, causing considerable damage to various woody and herba-
ceous plants (Bonavia et al. 1998; della Giustina & Navarro 1993; Holzinger et al. 1996).
Finally, it should be noted that a number of central European species have been re-
ported as pests from other countries, but not from Germany, notably Megadelphax sor-
didula (Stål), Laodelphax striatella (Fall.), Javesella pellucida (F.), Macrosteles sexnotatus (Fall.)
and Euscelidius variegatus (Kbm.). In Finnish cereal fields, for instance, Javesella pellucida
(F.) and Macrosteles cristatus (Rib.) occurred in notable densities although numbers of
nymphs were very low; in general, the Auchenorrhyncha fauna in these fields com-
prised almost exclusively migrants (Raatikainen 1971, 1972). Moreover, a considerable
proportion was likely to feed on weeds, such as Phleum pratense, Elymus repens, De-
schampsia cespitosa and Poa spp. Müller (1956) lists further examples, for instance of
Cercopis vulnerata Rossi causing fruit damage on pears, and of Philaenus spumarius (L.)
attacking pine plantations in Anatolia, tobacco fields in Ukraine and potatoes in Wales.
352 Life strategies

6 Life strategies

6.1 Host specificity and its possible causes

Resource specialization is a central issue in evolutionary ecology (e.g. Begon et al.

1996; Futuyma & Moreno 1988) and is thought to be correlated with the dramatic
radiation of phytophagous insects and mites and their angiosperm hosts (Hutchinson
1959; Strong et al. 1984; Farrell 1998; Schoonhoven et al. 1998; Walter & Proctor 1999).
Concerning insects, the discussion on diet width for a long time focused on plant
defence and reciprocal evolutionary effects between insects and plants, initiated by
the classical paper of Ehrlich & Raven (1964) (see below). Crawley (1983) summa-
rized advantages of polyphagy and monophagy. Accordingly, food is easier to find
for polyphages, and its availability is temporally more constant. Energetic costs and
the risk of predation during search are low, and shortage of certain nutritious ele-
ments in a single plant species can be balanced. Moreover, toxic compounds of certain
host species may be tolerated in low concentrations. In contrast, monophages can
develop special adaptations in morphology, physiology and behaviour. Thus, their
feeding efficiency is high, and they may avoid interspecific competition. Therefore,
monophagy is likely to be advantageous in small and short-lived animals such as
insects, which need only a few plant individuals for feeding. The following sections
will present broad patterns found in the studied Auchenorrhyncha guilds and discuss
the relevant theories.

6.1.1 Diet width in Auchenorrhyncha

The classification of diet width used here is shown in Table 4 (see also Fig. 11). Alto-
gether 5 categories are distinguished, which must – at the present stage of knowledge –
deal with inconsistencies caused by different taxonomic categories of plants, and also,
if comparative studies are intended, with diverging classifications of diet width of oth-
er authors (see Symons & Beccaloni 1999). However, the data matrix presented here
(sections 4 and 5.1.2) should allow an alignment with categories of other authors. Spe-
cies were preliminarily assigned (marked with “?”) to one of the categories of diet width
after at least 3 host (i.e. breeding) records (or alternatively, feeding records in vertical
migrants and polyphages). Substantiated assignment is assumed here after at least 10
records. These criteria may seem rather speculative, but will be maintained here based
upon the following arguments:
 Diet width in Auchenorrhyncha 353

(i) In most species, the number of host records is clearly higher than three. The
total number of own insect – host records collected for the present study is 8,600.
Thus, the average number of positive plant records per Auchenorrhyncha species is
only 14, but these data are further supported by numerous negative plant records
and literature data.
(ii) Field experience has clearly shown that the bulk of previously only little known
species shows specific host requirements, and consequently, the occurrence is often rather
patchy, although the overall distribution may be wide. There are numerous examples of
species formerly thought to be rare, which turned out to be fairly common, as soon as
their hosts had been identified. This is particularly true for Kelisia sima Rib., K. guttu-
lifera (Kbm.), Calligypona reyi (Fieb.), Florodelphax paryphasma (Fl.), Elymana kozhevnikovi
(Zachv.), Streptanus confinis (Reut.) and Cosmotettix caudatus (Fl.), to name but a few.
Also, whenever changes of diet width classification of single species were necessary
due to new host records during the process of writing this book, most shifts were of
minor significance (for instance, from 1st to 2nd degree monophagous), or occurred
rather towards a narrower host range. Further, among species newly recorded for Ger-
many during recent decades, a disproportionately large part was monophagous. Thus,
the real proportion of specialists is rather likely to be higher than assumed here.
It should be stressed that these classifications are almost exclusively based on field
data. There is ample evidence that species, being monophagous in the field in one local-
ity or region, may show a broader host range in breeding experiments and in other
places. Thus, the term ‘monophagy’ needs not necessarily imply exclusive morpholog-
ical or physiological adaptations, but may simply involve a functional or regional spe-
cialization. The former case is likely to occur on biochemically rather peculiar plants
such as Salix, Hypericum and most species of Lamiaceae, whereas the latter case may
rather be found on little-defended plants such as graminoids.
All species of central European Auchenorrhyncha show at least some degree of host
selectivity, which may – depending on the geographic scale – ultimately result in local
monophagy. This is even true for the extremely polyphagous Philaenus spumarius (L.),
nymphs of which often appear to be clumped on certain plant species, though with a
high variation between populations. In general, the proportion of strict monophages
(including tentatively assigned species) is 39%, and 20% are 2nd degree monophagous,
i.e. restricted to a single plant genus, 18% are 1st degree oligophagous, i.e. restricted to
one plant family, and the remaining species utilize more than one plant family (Fig. 11).
Differences between Fulgoromorpha and Cicadomorpha (see Table 32) mainly occur in
the proportion of 1st degree monophages, which is higher in the former group. Howev-
er, the proportion of polyphages is also higher in fulgoromorphans, mainly due to the
cixiids, which are tentatively regarded as host generalists by most authors, although the
host range of nymphs may be reduced simply for reasons of the limited mobility in the
soil. Despite these uncertainties, figures presented here provide support for a more pro-
nounced host specificity in planthoppers compared to leafhoppers (e.g. Waloff 1980),
although it may seem paradox, that the proportion of polyphages is also higher.
Regarding plant families and their numbers of 1st and 2nd degree monophages (Fig.
12), the Poaceae show by far the highest values, with 115 species, accounting for almost
19% of the Auchenorrhyncha total of Germany. Cyperaceae and Salicaceae have about
354 Life strategies

?
po?
m1
po

o2?

o2

o1?

m1?

o1

m2? m2

Fig. 11: Diet width of the Auchenorrhyncha species known from Germany. m1 = 1st degree monophagous
(1 host species), m2 = 2nd degree monophagous (1 host genus), o1 = 1st degree oligophagous (1 host
family), o2 = 2nd degree oligophagous (2 food plant families or up to 4 species belonging to up to 4 plant
families), po = polyphagous. Question marks indicate tentative assignments.

Table 32. Diet width of the Auchenorrhyncha species of Germany. Explanation of categories see Fig. 11

Auchenorrhyncha total Fulgorom orpha Cicad om orpha

Diet w id th n % n % n %
1st d egree m onophagous 138 22.3 41 28.3 97 20.5
1st d egree m onophagous? 101 16.3 25 17.2 76 16.0
2nd d egree m onophagous 84 13.5 17 11.7 67 14.1
2nd d egree m onophagous? 44 7.1 11 7.6 33 7.0
1st d egree oligophagous 69 11.1 9 6.2 60 12.7
1st d egree oligophagous? 41 6.6 6 4.1 35 7.4
2nd d egree oligophagous 18 2.9 0 0 18 3.8
2nd d egree oligophagous? 22 3.5 4 2.8 18 3.8
Polyphagous 43 6.9 3 2.1 40 8.4
Polyphagous? 46 7.4 26 17.9 19 4.0
Unknow n 14 2.3 3 2.1 11 2.3
Total 620 100 145 100 475 100

half of this number, respectively. Higher numbers of monophages are also found on the
families of dominating woody plants, notably the Betulaceae, Rosaceae, Fagaceae, Ul-
maceae, Aceraceae and Pinaceae, most of which are represented by few species only,
and in the Asteraceae, Lamiaceae and Ericaceae, which are species-rich and contain
 Plant apparency 355

140

1 15
120
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ce

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et
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10
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Fig. 12: Species numbers of 1st and 2nd degree monophagous Auchenorrhyncha on the plant families of
Germany

toxic secondary compounds. Noteworthy are also 5 monophages on the Urticaceae,

which are restricted to Urtica dioica.
The situation is somewhat different when we regard plant genera (Fig. 13) Carex (true
sedges) is by far the most preferred taxon, with 46 specialists, followed by Salix (wil-
lows and sallows), Festuca (fescue), Populus (poplar), Calamagrostis (small-reed) and
Phragmites (common reed). The latter comprises only a single plant species and is note-
wothy in harbouring the highest number of 1st degree monophages all over central
Europe and probably all over the western Palearctic. Among the graminoids, Agrostis,
Eriophorum, Brachypodium, Molinia, Phalaris and Juncus are noteworthy. Among decidu-
ous trees, Quercus, Ulmus, Alnus, Betula, Acer, Corylus, Rosa and Picea are the most spe-
cies-rich. Only Urtica and Artemisia are noteworthy among the remaining genera. The
former is purely herbaceous, and only a single species is attacked. Among the latter,
only dwarf shrubs are exploited by specialists. The following sections will discuss pos-
sible explanations for these patterns against the background of more general theories.

6.1.2 Plant apparency

High absolute numbers of monophages on highly apparent trees, graminoids and
Urtica provide evidence, that first of all, plant apparency promotes herbivore special-
ization. Plants with many specialist herbivores are tall or grow in large stands, and they
are perennial and widespread (see Fig. 12, Fig. 13, Fig. 14).
A similar preference of apparent host plants is also found in Sternorrhyncha (based
on worldwide data). The majority of host-specific species of Aphidoidea, Psylloidea,
356 Life strategies

50
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Fig. 13: Species numbers of 1st and 2nd degree monophagous Auchenorrhyncha on the plant genera of
Germany

Aleyrodidae and Coccoidea lives on woody and graminoid plants, although in Aphi-
doidea, the Asteraceae, which are largely herbaceous, are the most favoured plant fam-
ily (Eastop 1978). On the other hand, Brändle & Brandl (2001) found that the proportion
of arthropod specialists on 25 German tree genera was negatively correlated with grid
occupancy, as a measure of tree abundance (though based on insufficient literature re-
garding the Auchenorrhyncha).
Spatial and temporal plant apparency have long been discussed as a major factor
determining herbivore feeding and specialization. The temporal component involves
short-term persistence, like plant age and succession, and long-term stability of cli-
mate and geology, whereas spatial plant apparency comprises range size, abundance,
as well as plant size and architectural complexity. Some of these factors are intercorre-
lated and difficult to disentangle. For instance, most persistent habitats of middle
latitudes are covered with spatially apparent plants, notably old-aged and large trees,
which in turn are abundant and widespread. In an undisturbed habitat even a single
host individual can persist for years, providing food and shelter and thus, rendering
costly dispersal unnecessary (see Denno et al. 1989). In contrast, early successional
stages and reduced food plant apparency favour polyphagy, as predicted by the clas-
sical r and K concept (see MacArthur 1962; Pianka 1970) and specifically confirmed
for Auchenorrhyncha in Bohemia and Slovakia (Novotný 1994a). Consequently,
monophagy should prevail in habitats dominated by perennial and large or abundant
plant species, like trees and grasses, whereas annuals and small or rare plants should
support mainly polyphages.
 Plant diversity 357

6.1.3 Plant diversity

Species-rich plant groups like the Poaceae, Cyperaceae and Salicaceae (and Carex, Salix,
Festuca in particular), tend to be attacked by more herbivore specialists than species-poor
groups (Fig. 13). Moreover, these specialists tend to be closely related and are often conge-
neric, for instance in Kelisia, Ribautodelphax, Macropsis, Idiocerinae, Kybos, Mocydiopsis, Psam-
motettix and Cosmotettix. Many of these hosts show a similar geographic distribution, and
thus, parapatric speciation is unlikely. However, the significance of secondary plant com-
pounds may be low at least on graminoids, which are more physically defended against
chewing herbivores. One of the best documented examples of sympatric host-associated
race formation with evidence for incipient speciation is found in the Nearctic membracid
treehopper Echenopa binotata Say (Wood & Guttman 1981, 1982; Wood 1987). In this spe-
cies complex, different host races are found on 7 plant genera belonging to 6 taxonomical-
ly rather distinct families (notably the Magnoliaceae, Rutaceae, Juglandaceae, Celastrace-
ae, Caprifoliaceae, Fabaceae). Interestingly, speciation appears to be mediated rather by
host-specific water contents causing differences in phenology of egg hatch, and perhaps
the impact of a myrmarid egg parasitoid, rather than by plant defence (Wood 1987, 1988).
Among central European Auchenorrhyncha, such examples of sympatric speciation on
different hosts are most likely to exist in guilds living on grasses, sedges, Salicaceae and
perhaps Lamiaceae.

40
33
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monophagous monophagous oligophagous oligophagous phagous

Fig. 14: Species numbers and diet width of the most diverse Auchenorrhyncha guilds (> 17 species) in
Germany
358 Life strategies

A further factor enhancing sympatric, though not syntopic, speciation may be inferred
from Fig. 14, showing the 12 most diverse Auchenorrhyncha guilds in the study area.
Accordingly, highest numbers and proportions of monophages are found on the grasses
Festuca ovina and Phragmites australis, both of which are highly eurytopic regarding soil
types and altitudinal distribution. Many of their associated Auchenorrhyncha species,
however, are restricted to rather special conditions, for instance calcareous grassland,
heaths, salt marshes, inundated or temporarily flooded sites. In contrast, guilds are less
specific on the more stenotopic grasses Holcus mollis and Elymus repens. In trees, most of
which support only moderate proportions of monophages, moisture, pH and other abiot-
ic parameters in the canopy may be less variable, irrespective of site conditions.

6.1.4 Plant defence

The evidence for surface defence in central European plants against Auchenorrhyn-
cha is rather limited. High contents of silica in epidermal tissues of Equisetum spp.,
which may deter cattle and other chewing herbivores, are apparently of little efficiency
against specialized sap-sucking insects, like Javesella stali (Metc.) and Macrosteles fronta-
lis (Scott) (see section 5.1.2.1). Similarly, trichomes are found in many dicotyledonous
plants heavily attacked by mesophyll-sucking typhlocybine leafhoppers, such as Urtica
dioica and numerous species of Lamiaceae. The same is true in some woody species
showing hairy leaf undersides, e.g. among the Salicaceae and Rosaceae. In fact, the
most heavily attacked plants are largely glabrous (notably most species of Poaceae,
Cyperaceae and Fagaceae) or lose pubescence after a while (at least on the upper leaf
surface, for instance some species of Salix, Populus and Betula). In plant families or gen-
era with both glabrous and hairy species, there appears to be no difference in species
richness of Auchenorrhyncha guilds, e.g. in Populus and Salix (see section 5.1.2.9).
As for chemical plant defence, the data presented here can only offer some general
clues. Overall, 75% of the Auchenorrhyncha species of the German fauna feed on phlo-
em or xylem. Unlike leaf parenchyma, these plant saps generally contain low levels of
digestability-reducing compounds, which are generally of a high molecular weight. In
contrast, concentrations and significance of toxins, which tend to consist of smaller
molecules, are only poorly studied (Raven 1983), but may be more important than pre-
viously thought (see Dixon 1998).
Noxious compounds are likely to play a role in some rather diverse plant taxa which
are entirely avoided by Auchenorrhyncha. This may be the case in most ferns (particu-
larly the Aspleniaceae and Dryopteridaceae), the Ranunculaceae, Chenopodiaceae,
Caryophyllaceae, Brassicaceae, Onagraceae, Euphorbiaceae, Apiaceae, Gentianaceae,
Boraginaceae, Scrophulariaceae, Campanulaceae, Rubiaceae, Asteraceae, the former
Liliaceae (now split into several families) and the Orchidaceae. In some of these groups,
however, for instance in the Chenopodiaceae, Caryophyllaceae, Brassicaceae and Ona-
graceae, reduced apparency may offer an alternative explanation because they com-
prise numerous annuals, biennials and rare species. Further groups, which are known
for their toxic compounds, such as Hypericum (Clusiaceae), Chaerophyllum (Apiaceae)
and Knautia (Dipsacaceae) are attacked only by a small number of specialised, monoph-
agous mesophyll feeders.
 Plant defence 359

The Salicaceae are probably the best example of a plant group showing chemical com-
pounds and Auchenorrhyncha guilds, both of which are highly specific. In addition, they
are avoided by numerous species, which are broadly polyphagous on many other trees
and shrubs (section 5.1.2.9). In this case, the defensive role of plant chemistry for host
selection is likely to dominate over other factors, permitting only specialist feeders. Sim-
ilarly, the Lamiaceae contain specific ethereal oils, many of which are known for their
antibiotic effects. Members of this family are the main and often exclusive hosts of the
typhlocybid leafhopper genus Eupteryx, for which mesophyll-feeding has been specifical-
ly studied (Pollard 1968, 1969). Thus, detoxification or sequestering mechanisms are like-
ly to occur in this taxon. Conversely, Fraxinus excelsior (Oleaceae), is the only central Euro-
pean tree belonging to the asterid clade, which is known for a wide occurrence of iridoids
(Frohne & Jensen 1998), and its infestation, even by polyphages, is very scarce.
In conclusion, against the background of 77% of the German Auchenorrhyncha species
being specific on plant family level, and 59% specific even on genus or species level, it is
tempting to assume that plant chemistry plays an important role for the selection of host
specificity. However, most patterns may also be explained by alternative mechanisms,
except in a few groups such as the Salicaceae and some dicotyledonous herbs. Thus, the
idea of an evolutionary impact of Auchenorrhyncha on their host’s chemistry must re-
main a speculation, until convincing experimental evidence has been provided.
Conventional plant defence theory attributes great importance to secondary plant
compounds, essentially based upon the idea of a coevolutionary ‘arms race’ between
plants and herbivores (Ehrlich & Raven 1964). The ecological literature abounds in well-
documented examples of detrimental effects on invertebrate as well as on vertebrate
herbivores (see Harborne 1972; Jacobson & Crosby 1971; Rosenthal & Janzen 1979).
However, secondary metabolites are also toxic to the plant itself and must be stored in
glands, vacuoles or as inactive forms. On the other hand, many host specialists have
evolved adaptations to cope with these substances and may even use them for host
recognition, feeding stimulation or sequester them as defence against predators (Craw-
ley 1983; Futuyma 1983). Principally, two categories of chemical defence have been dis-
tinguished (Feeny 1976; Rhoades & Cates 1976; see also Howe & Westley 1988). Quali-
tative defences, i.e. toxins, such as alkaloids, cyanogens, glucosinolates, non-protein
amino acids and terpenes, comprise small molecules and cause direct poisonous ef-
fects. Their amount in plants is usually low, with highest concentrations in buds, new
leaves or unripe fruits. They are most common in rare, short-lived, herbaceous or early
successional species, predominantly belonging to advanced angiosperms. In contrast,
quantitative defences comprise digestability-reducing substances, such as cellulose,
lignins, tannins and silica, which inhibit enzymic hydrolysis of proteins and assimila-
tion, and consequently, reduce nutritional value. They consist of complex polymers or
crystals usually occurring in high concentrations, mostly in woody tissues and mature
leaves of perennial species of ferns, gymnosperms and phylogenetically more primitive
angiosperms. These ideas, widely known as the plant apparency theory, have been broad-
ly discussed. A major point for criticism is the difficulty of a clear distinction between
qualitative and quantitative defence. Some authors also pointed out that further prop-
erties of secondary compounds may include repelling, growth inhibiting, reducing fe-
cundity and delaying of development (see Hartley & Jones 1997). Further ideas stressed
360 Life strategies

a possible synergistic interplay of different substances and their chemical distinctive-

ness, although a study of the British insect fauna on umbellifers did not provide posi-
tive evidence (Jones & Lawton 1991).
However, there is a number of shortcomings in explaining observed patterns by these
theories of plant defence. Thus, ideas stressing mutual effects on the ecology and evolution
of insect herbivores and plants have been challenged by the theory of sequential evolution,
which generally questions the impact of insects as a driving force in plant evolution, there-
by regarding most insect – plant relationships as rather asymmetric. Instead, climate, soil,
interspecific competition, pathogens and other factors are likely to be much more relevant
for plants, whilst the overall impact of insects on their hosts is negligible (Jermy 1976).
Concerning Auchenorrhyncha and other sap-sucking insects, published evidence of
specific plant defence mechanisms is rather scarce compared to leaf-chewing groups.
For instance, pubescence of cotton, soybean, alfalfa, potato and bean has been found to
act as an effective physical barrier against leafhopper probing and feeding, particularly
in the nymphs of Empoasca fabae (Harr.) (Tingey 1985). On the other hand, Lovinger et al.
(2000) also found evidence for negative effects of pubescence on parasitoid searching
behaviour (see also Eisner et al. 1998). Plant defence through protection by animals has
long been known mainly as plant-ant mutualism (Janzen 1966; Price et al. 1980). Since
the 1980s, however, much research work has focused on indirect plant defence through
various kinds of organisms, including parasitoids, predators and endophytic fungi (Sa-
belis et al. 1999; Clay 1996; Leuchtmann et al. 2000; Brem & Leuchtmann 2001). Effects
on Auchenorrhyncha have not been subject to study so far.
The significance of secondary plant metabolites for phloem and xylem feeders was
discussed by Raven (1983). He suggested that quantitative defences are of limited effi-
ciency due to their larger molecular size and reduced solubility compared to mostly mo-
nomeric and oligomeric nutrients. Moreover, they are toxic to the symplast and thus,
must be sequestered by the plant. In contrast, the toxicity of qualitative defences to plant
tissues is generally less severe, because they are often activated or released only where
and after tissue has been damaged by herbivores. Although toxins appear to be wide-
spread in the phloem sap of plants, their content is considerably lower than in other tis-
sues. As for xylem sap, the concentration of toxins may be potentially high, but the evi-
dence suggests that this is only exceptionally the case. Instead, the low and unbalanced
nutritional value combined with the strong negative pressure in itself renders xylem sap
a rather unattractive resource. Thus, Raven (1983) states, that there is no reason to assume
major negative effects of digestability-reducing compounds on sap-sucking Hemiptera,
while evidence for toxins as deterrents is inconclusive. Regarding Asian planthoppers of
rice and some aphids on plants of economic importance, there is some scattered evidence
of negative response to toxins and even more complex polymers, but the present data do
not suggest a major significance (Risebrow & Dixon 1987; Cook & Denno 1994).
In contrast, mesophyll-feeding typhlocybine leafhoppers ingest cells mainly of the pal-
isade and spongy parenchyma, and principally have to cope with secondary metabolites.
Their contact may be reduced, since glands and other storage organs are more common in
or on epidermic tissue, although vacuoles, which also contain secondary metabolites, are
mechanically broken before passing the food canal (Pollard 1968). Thus, we may con-
clude, that the relationship of mesophyll-feeding leafhoppers and chemical plant com-
 Nitrogen 361

pounds is largely unknown, but the dramatic relatively recent radiation of Typhlocybinae
suggests, that plant chemistry was not a major obstacle for detoxification. There is indi-
rect evidence, however, that digestability-reducing substances and also toxins enhance
host specialization. The effects of these substances need to be studied in the laboratory.

6.1.5 Nitrogen
Nitrogen is assumed to play an important role in plant-herbivore relationships (McNeill
& Southwood 1978; White 1993). Among insects, aphids in particular have been found to
show a synchrony of reproduction with spring and autumn peaks of available phloem ni-
trogen. During summer many species switch onto herbaceous plants (Dixon 1963, 1966,
1973, 1976). The mirid bug Leptopterna dolobrata (L.) moves between different plant parts,
notably the stem and the seeds, in order to exploit nitrogen more efficiently (McNeill 1973).
For leafhoppers on grasses, Prestidge (1982) found that maximum nitrogen utiliza-
tion efficiencies were reached at different nitrogen levels of the host and suggested that
they may be associated with particular nitrogen concentrations. Furthermore, Prestidge
& McNeill (1983a, 1983b) studied Auchenorrhyncha communities on monospecific plots
sown with 5 different grasses. They concluded that monophagy among grassland
Auchenorrhyncha is uncommon and that oligophagous species may perform plant
switching depending on the particular nitrogen content.
However, it must be objected, that sown grassland selects for a small group of eury-
topic, polyphagous and bivoltine species, which a priori show two annual peaks of abun-
dance (sections 6.1.7; 6.2.3), whereas the majority of grassland Auchenorrhyncha is clearly
monophagous (Fig. 11; see section 6.1.1). Moreover, most published curves of seasonal
abundance in central and northern Europe show a distinct midsummer peak, when
leaves are mature and nitrogen concentrations in plant sap are rather low (Marchand
1953; Schaefer 1973; Nikusch 1976; Waloff 1980; Peter 1981; Nickel, unpublished re-
sults). This phenology probably reflects the overall dominance of monovoltine Auchen-
orrhyncha species in central Europe (section 6.3.1), and may simply be a correlation
with temperature rather than plant nitrogen quantity.
Thus, small-scale responses to nitrogen may occur in some Auchenorrhyncha spe-
cies, which favour certain life strategies, but should not be generalized. Climate is like-
ly to play a much more important role for insect phenology than nitrogen content. Fur-
thermore, in phloem and xylem feeders, gain of nitrogen in summer can simply be
achieved by spending more time on sucking, even without moving to other plants or
feeding sites. An important clue to assess the significance of nitrogen for herbivore host
specificity may be provided by correlating the Auchenorrhyncha species numbers on
different grasses to phloem nitrogen content.

6.1.6 Interspecific competition

The role of interspecific competition on resource specialization has been a classical
topic in evolutionary ecology. Direct competition is most likely to occur, if two or more
species utilize a similar niche, and if densities are high in at least one of them. Central
European Auchenorrhyncha species tend to concentrate on relatively few plant fami-
362 Life strategies

lies and species (Fig. 9, Fig. 12, Fig. 13, Fig. 14). Thus, the potential for competition
should be rather high. However, on a closer look, niche parameters like host plant spe-
cies, feeding and oviposition sites on the plant, plant height, type of plant sap or tissue,
temperature, insolation, moisture, salinity, altitude, geographic range, phenology and
body size may allow alternative explanations of resource partitioning in most cases
(Bieman 1987a, 1987b; Booij 1981, 1982; Halkka et al. 1977; Le Quesne 1972; Müller
1978; Peter 1981; Waloff 1979). Rostrum length and acoustic interference, which are poorly
studied, deserve more attention and may provide further clues. On the other hand,
phloem and xylem sap are shared resources even to spatially separated sucking insects.
Thus feeding by one species results in a reduced quantity of resources available to all
other species and may even affect other guilds such as leaf-chewers and miners. As a
consequence, indirect interactions, including competition, are likely to be widespread
among terrestrial herbivore communities (e.g. Masters & Brown 1997).
Regarding direct interactions, however, only a few Auchenorrhyncha guilds in central
Europe must be seriously taken into consideration as potential competitors. Species groups,
which are closely related, feed on the same type of plant sap and share the same host or
the same phenology, are compiled in Table 33. Accordingly, high numbers of strictly
monophagous species are particularly remarkable on Populus nigra and P. tremula (5 and 3
species of idiocerine leafhoppers, respectively), Salix alba (three species of Macropsis) and
Phragmites australis (7 species of Chloriona). 5 species of Edwardsiana (4 of which are 1st
degree monophages) plus further 5 species of Typhlocybinae live on Corylus avellana. In
Eupteryx stachydearum (Hardy), E. curtisii (Fl.) and E. florida Rib., the distribution on Lami-
um galeobdolon is rather striking, since field evidence suggests, that a high proportion of
host stands are occupied, but each with a single Eupteryx species only.
Interspecific competition has long been assumed to be a major structuring force of
animal communities, mainly based upon Darwin’s (1859) idea of ‘the struggle for exist-

Table 33. Some potential model guilds for further studies of interspecific competition

Potential com petitors

H ost plant Taxon Species num ber Param eters potentially allow ing niche partitioning
Populus nigra Id iocerinae 5 Tim e, bod y size
Populus tremula Id iocerinae ≥3 Bod y size, sun exposure
Salix alba M acropsis 3 Geographical range, bod y size
Salix caprea Id iocerinae 5 Tim e, bod y size, alternative hosts
Salix spp. A phrophora spp. 2 Altitud e, alternative hosts
Corylus avellana Edwardsiana 5 Geographical range, sun exposure
Betula pendula Oncopsis ≥4 Altitud e, bod y size, m oisture
A lnus glutinosa Typhlocybinae ≥ 12 Moisture, alternative hosts?
Quercus robur Typhlocybinae ≥ 13 Moisture, alternative hosts
Quercus petraea Typhlocybinae ≥ 12 Alternative hosts, sun exposure?
Ulmus spp. Edwardsiana 4 Geographical range?, sun exposure?
A cer pseudoplatanus Typhlocybinae 8 Moisture
A cer campestre A cericerus ≥2 ?
Stachys sylvatica Eupteryx ≥3 Geographical range, m oisture, alternative hosts
Salvia officinalis Eupteryx ≥5 Geographical range
Juncus effusus Conomelus 2 Geographical range, m oisture
Phragmites australis Chloriona 7 Geographical range, m oisture, salinity
 Interspecific competition 363

ence’, and later, Gause’s (1936, 1937) principle of competitive exclusion. In the presence
of potential competitors, specialization would occur, eventually leading to character
displacement (Futuyma & Moreno 1988). Consequently, closely related phytophagous
insects should avoid competition by utilizing different host plants or parts of them.
During the second half of the 20th century serious doubts about the overall significance
of interspecific competition became evident, particularly for herbivore communities, and
alternative models were proposed (e.g. Hairston et al. 1960; Strong et al. 1984). These au-
thors argued that in most ecosystems only a small proportion of the living plant biomass is
consumed by herbivores, because low or strongly fluctuating food quality, plant defence
mechanisms, predators, and parasitoids were the main factors keeping their densities be-
low levels at which competitive interactions would occur. Thus, vertical forces (‘insects
between the devil and the deep blue sea’) were concluded to be more important in shaping
insect populations and communities than intraguild interactions (Lawton & McNeill 1979;
Strong et al. 1984). The short timespan of empirical studies, which makes access extremely
difficult, lead to the idea of ‘the ghost of competition past’ (Connell 1980).
More recently, interspecific competition among phytophagous insects has been re-
viewed (Denno et al. 1995; see also Stewart 1996). Accordingly, the bulk of experimental
studies carried out mainly through the 1980s revealed negative effects between poten-
tial competitors, particularly among closely related, introduced, sessile and aggrega-
tive species, as well as on species feeding on discrete resources such as plant sap and
marrow, forbs or grasses. However, the arguments presented in this book were not con-
clusive, since researchers are likely to select systems with a higher probability of posi-
tive evidence, whereas the study interest in cases of neutral coexistence is small, and
negative or equivocal results are less likely to be published. The most serious objection
is that experimental conditions do not reflect the situation in the field, where small-
scale migration and dispersal may reduce locally high densities. The significance of
competition in herbivores was further discussed by Stewart (1996) and Schoonhoven et
al. (1998), who also came to the conclusion, that the evidence is weak.
The following section reviews the most frequently cited experimental studies of in-
terspecific competition in Auchenorrhyncha, notably McClure & Price (1975, 1976) on
Erythroneura species on Platanus occidentalis, Stiling (1980) on Eupteryx species on Urtica
dioica, Hunter & Yeargan (1989) on Erythroneura species on Cercis and Settle & Wilson
(1990) on Erythroneura species on Vitis, all resulting in positive evidence for both intra-
and interspecific competition. However, densities in these experiments were high, rang-
ing from 1 to 32 individuals per leaf or between 1 and 16 individuals per 100 cm2 leaf
area, respectively. Moreover, due to caging, insects were prevented from migrating to
other less infested parts of the plant. For instance, Stiling (1980b) studied interactions
between Eupteryx urticae (F.) and E. cyclops Mats. in Wales, both feeding monophagous-
ly on Urtica. At densities between 1 and 8 caged individuals per 100 cm2 leaf area, the
author found evidence for both intra- and interspecific competition on potted nettles.
But when we assume a leaf area index of c. 5, which is typical for most central European
ecosystems (Ellenberg 1996), Eupteryx densities in the field are much lower and have
only exceptionally been found to exceed 500 ind./m2 ground surface (Zabel & Tscharn-
tke 1998; M. Sayer, pers. comm.; S. Zabel, pers. comm.). Accordingly, in Lower Saxony,
1 ind./100cm2 leaf area was the maximum field density, which was only rarely observed.
364 Life strategies

Moreover, in the field, non-caged insects may migrate to fresh nettle stems and leaves,
which frequently grow within or along the margin of the host stands until mid-summer,
thus further reducing densities. As for the classical example of Erythroneura species on
Platanus occidentalis (McClure & Price 1975, 1976), one of the authors later weakened his
original statement, arguing that many habitats were found to be totally unoccupied and
that any real effects in natural communities have yet to be demonstrated (Price 1983).
The most convincing laboratory experiments on interspecific competition among
Auchenorrhyncha species were conducted by Valle et al. (1989), who studied four south-
east Asian species of Nephotettix feeding on rice. Their design included single- and mixed
species experiments on rice seedlings running for 7 months to determine the system’s
carrying capacity. They also reared mixed-species populations starting at the same den-
sities. In most species pairs, either of the two showed reduced densities compared to
single-species conditions. In some cases the inferior species even became extinct after 7
months. Again, however, the question remains as to whether such simple experiments
can simulate field conditions.
A good field example against significant effects of intra- as well as interspecific com-
petition is found in xylem-feeding cercopids, which often form mono- and multispecif-
ic aggregations in shared spittle (Halkka et al. 1977; McEvoy 1986). Apparently, possi-
ble negative effects are outweighed by the advantages of aggregation. Unlike in leaf-
chewing insects, which ultimately remove biomass, plant sap will only become exhausted
after strong infestation.
Ross (1957, 1958), who was the first to study Erythroneura species on Platanus occiden-
talis in North America, argued, that in ten years of field work he has seen only a single
group of etiolated trees. He concluded, that competitive effects among Erythroneura on
American sycamore should be exceptional. Likewise, after almost 50 and 12 years of
intensive field work (Remane pers. comm.; Nickel unpublished data), evidence for se-
vere infestation of most central European deciduous trees by mesophyll-feeding leaf-
hoppers must be considered insufficient. Most students who carried out field studies
on this subject rather used the terms ‘coexistence’ and ‘resource partitioning’ (Halkka et
al. 1977; Le Quesne 1972; Müller 1978; Peter 1981; Waloff 1979). Thus, the evidence of an
evolutionary impact of negative intraguild interactions leading to diverging host selec-
tion in Auchenorrhyncha, must be considered as weak.
Whereas classical notions of interspecific competition usually referred to more ap-
parent resources, like space and food, there is now a growing awareness of indirect
effects. Thus, a number of more recent studies did not find direct competition between
species, but indirect effects mediated by parasitoids, host plants (including defence
mechanisms – see above), increased proportion of macropters and physical damage,
e.g. caused by stem borers (Denno & Roderick 1992; Masters & Brown 1997; Tscharntke
1999). Very recently, Alla et al. (2001a, 2001b) provided evidence for chemical interac-
tion between the aphid Rhopalosiphum padi (L.) and the leafhopper Psammotettix alienus
(Dhlb.), resulting in negative effects of an aphid extract on the leafhopper nymphs.
Clearly, this field deserves further attention. It should also be taken into consideration
that intra- and interspecific crowding effects by accumulating honeydew, e.g. of aphids,
and excrements of mesophyll feeders are likely to have the potential to reduce densities
by deteriorating physical conditions, e.g. adhesive properties of the plant surface, or by
 Resource predictability 365

favouring pathogens. Experimental studies should focus on more discrete interactions,

particularly diffuse competition (including inter-guild interactions), indirect effects of
predation and parasitoid attacks mediated by plant semiochemicals, and perhaps sound-
transmitting properties of the host, which may offer a considerable potential for ex-
plaining patterns in Auchenorrhyncha community structure.

6.1.7 Resource predictability

Predictable resources, involving habitat persistence as well as temporal and spatial
host plant apparency, are correlated with larger numbers of monophagous Auchenor-
rhyncha species, which prefer old-aged plants (notably trees) or plants forming large
stands (notably graminoids, Urtica dioica) (see chapters 5.1.3.2; 6.1.2). Conversely, most
unpredictable habitats select for host generalists (see the more detailed discussion in
section 6.2.3).

6.1.8 Neural constraints

Concerning the prevalence of monophagy in Auchenorrhyncha, many of these con-
cepts fail to offer a satisfactory explanation. While chemical plant defence is likely to play
a role in some broadly avoided groups, like Brassicaceae and Boraginaceae, the scarcity of
potentially noxious compounds in phloem and xylem sap suggest alternative mechanisms.
In particular, one of the major questions raised in this book is the high proportion of host
specialists on graminoid plants, which are generally thought to be poor in secondary
compounds (Frohne & Jensen 1998). None of the previous chapters offered the potential
to explain, why stems or leaves of grasses and sedges, which are almost indistinguishable
for humans, attract different and highly specific insect herbivores.
A potentially powerful explanation is the neural hypothesis proposed by Bernays
(1998, 1999a), although experimental confirmation for Auchenorrhyncha has not yet
been presented. Accordingly, polyphagous grasshoppers, whiteflies and aphids are sig-
nificantly less efficient in foraging, ovipositing and probably escaping from predators,
compared to specialists, since they have to cope with ambiguous stimuli from various
potential host plants (Bernays 1996, 1998, 1999b; Bernays & Funk 1999). Due to the
limited neural capacity of processing and analyzing these stimuli, the effects of plant
chemistry on host choice would rather be a facilitation and stimulation instead of a
restriction or inhibition. The principal autonomy of the evolution of ecological traits in
herbivorous insects from plant properties was also stressed by Schoonhoven et al. (1998).
This hypothesis is indirectly supported by experimental evidence from breeding of
monophagous species of the delphacid genera Muellerianella (Booij 1982) and Ribauto-
delphax (Bieman 1987b), which could successfully reproduce over several generations
on several grasses, which have never been found to be the host in the field.

6.1.9 Regional monophagy

Regional monophagy has been found in the delphacid Hyledelphax elegantula (Boh.),
which feeds on numerous grasses (including Calamagrostis arundinacea, Brachypodium pin-
366 Life strategies

natum, Holcus mollis, Molinia caerulea, Deschampsia flexuosa) in the lowlands of southern
and middle parts of Germany. At higher altitudes and in the north German plain, howev-
er, it is monophagous on the last mentioned species. A similar northward increase in host
specificity probably occurs in the deltocephaline leafhopper Mocydia crocea (H.-S.), which
is rather eurytopic and oligophagous in southern Germany, but thermophilic and largely
restricted to Calamagrostis epigejos near the border of its range in the northern half of Ger-
many. Conversely, moving southward towards the Mediterranean, Eurhadina kirschbaumi
W.Wg., which is a strictly monophagous feeder on Quercus petraea in Germany, also feeds
on Castanea and further oaks (Drosopoulos 1999). Likeweise, some central European spe-
cialists on Phragmites australis additionally attack the giant grass Arundo donax in south-
ern Europe (Drosopoulos et al. 1983; Vidano & Arzone 1987b). As a consequence, the
classification of diet width in these species would change from 1st degree monophagous
to 1st degree oligophagous. More examples are enumerated in section 5.1.1
In fact, regional monophagy may be a rather common phenomenon, since many
Auchenorrhyncha species are found near the limit of their geographic range in Germa-
ny. However, feeding data in southern Europe are rather scarce. The possible mecha-
nism is a reduced acceptance of potential host plants towards the border of the insect’s
range, ultimately allowing reproduction only on a single host species, whereas other
hosts grow in microclimatically and otherwise unsuitable sites. Regional monophagy is
also reported from British populations of the swallowtail butterfly Papilio machaon L.
(Strong et al. 1984).

6.1.10 Endophytic fungi

Finally, toxins produced by endophytic fungi might play a role for the unequal Auchen-
orrhyncha infestation of the closely related grasses Brachypodium sylvaticum and
B. pinnatum. The latter supports at least 11 substantiated feeders, 4 of which are mono-
phagous, whereas the former is entirely unattacked. Recently, Brem & Leuchtmann (2001)
found that an endophytic fungus specific to B. sylvaticum may reduce noctuid herbivory
through fungal alkaloids. In contrast, B. pinnatum is only rarely infected by another fun-
gus species (A. Leuchtmann, pers. comm.). These organisms are widespread, particularly
in grasses, and may turn out to offer a considerable potential in explaining differential
attack in sap-sucking insects on grasses (Clay 1990, 1996, 1997; Saikkonen et al. 1999).

6.2 Dispersal

6.2.1 Wing length

Wing length is a major determinant of flight capability, and hence, dispersal and
population dynamics (e.g. Denno & Roderick 1990; Johnson 1969; Taylor 1985). Mac-
roptery is considered to be the primitive condition in Auchenorrhyncha and other Hemi-
ptera (Hennig 1969), but wing dimorphism, subbrachyptery or brachyptery prevail in
several groups of both Fulgoromorpha and Cicadomorpha. Brachypterous individuals
in planthoppers and other insect groups have been shown to start reproducing earlier
 Wing length 367

and to have a higher fecundity than macropterous individuals, because they do not
spend resources for the development of the wing apparatus (e.g. Denno & Roderick
1990; Denno et al. 1991; Roff 1990; Andersen 1997).
Flight capability, but not necessarily flight activity, can be expressed as the relation of
fore and hind wing length, of hind wing and abdomen length or of wing surface area
and body weight. An overview of wing length distribution in central European Auchen-
orrhyncha, however, is complicated by the fact, that only few data are published, par-
ticularly in Deltocephalinae. Moreover, transitions are difficult to assess, proportions of
macropterous and brachypterous individuals within species are often variable, and
published classifications may be inhomogeneous. Thus, at the present state of know-
ledge, a simple distinction between macroptery and wing dimorphism is the most reli-
able classification of wing length condition of species.
In a detailed review, Roff (1990) discusses the ecological significance and the evolution
of secondary flightlessness in insects, concluding that spatial and temporal homogeneity
of the habitat is the most important condition for wing reduction, but habitat dimension-
ality, geographic distribution (altitude and latitude, in particular), parthenogenesis, gen-
der and paurometabolism play a further role. In general, this is likely to apply also for
central European Auchenorrhyncha, although the ranking may be somewhat different.
(i) Habitat dimensionality
The most conspicuous tendency is the presence of macroptery associated with the
utilization of the third spatial habitat dimension. Thus, all arboricolous Auchenorrhyn-
cha species as well as those living on tall herbs are monomorphic macropterous, in both
sexes and through all taxonomic groups. Further, all migrants between different layers
of vegetation as well as between food plant species are also long-winged. Hence, mac-
ropters include all species of Cixiidae, except the endogeic Trigonocranus emmeae Fieb.,
Achilidae, Tettigometridae, Issinae, Cicadidae, Cercopinae, Membracidae, Ledrinae,
Macropsinae, except the epigeic Macropsidius sahlbergi (Fl.), Idiocerinae, Penthimiinae,
and Typhlocybinae (including epigeic species). Finally, among central European Delto-
cephalinae, the genera Fieberiella, Grypotes, Japananus, Opsius, Balclutha, Sagatus, Platyme-
topius, Idiodonus, Colladonus, Lamprotettix, Allygus, Allygidius, Speudotettix, Hesium, Tham-
notettix, Pithyotettix, Perotettix and Colobotettix are exclusively long-winged.
Prevailing macroptery in arboreal Hemiptera was already demonstrated and dis-
cussed for the British fauna, and was explained by the fact that woody plants are archi-
tecturally more complex. They comprise a third spatial dimension, with widely spaced
leaves and branches, and moreover, they form discrete units spatially separated from
each other (Waloff 1983). Another argument is simply the risk of falling down in bra-
chypterous insects, and consequently, the possibility of heavy population losses due to
epigeic predators, and missing the proper host when climbing up again. The latter phe-
nomenon can be found in tree eclector catches containing numerous nymphs of species
being monophagous on other trees, perhaps due to nymphal inability of host recogni-
tion through the thick bark of basal trunks.
In contrast, wing dimorphism prevails in species living among herbaceous vegeta-
tion. In these cases, macroptery does occur at least in a low proportion of the population
and may increase, if densities are high or if host plant quality deteriorates (see Denno
1976; Denno et al. 1991). In general, males are often longer-winged than females, for
368 Life strategies

instance in many species of Deltocephalinae, where macroptery in male is often com-

bined with female subbrachyptery. Reduction is most pronounced in epigeic species,
with brachyptery prevailing in Delphacidae, except in Stenocranus, Laodelphax striatella
(Fall.) and Javesella pellucida (F.), as well as in males of reed-dwelling species, like Del-
phax, Euides and Chloriona. Females of the latter are the only brachypterous Auchenor-
rhyncha in central Europe, which spend their whole life cycle high above the ground.
Among Cicadomorpha, brachyptery is found in Ulopa, Errhomenus, Doratura and a few
others, all of which are usually found on the soil surface.
For the bulk of brachypterous Auchenorrhyncha species, it can be concluded that
they show epigeic habits. As phytophages, they are unlikely to be limited by shortage of
food, since much of the plant biomass in many ecosystems is left for consumption to
detritivores (e.g. Hairston et al. 1960; White 1993). Thus, mobility is less important,
except if habitat conditions deteriorate. Moreover, phloem- and xylem-feeding species
do not consume or destroy living plant biomass and may potentially spend their whole
life on a single plant stem. Thus, selective pressure for higher reproduction rates may
usually be stronger than for macroptery and dispersal ability. Among holometabolous
insect clades, only the Siphonaptera and the Mecoptera, notably the Boreidae, show a
high degree of secondary wing reduction (Roff 1990). Like in brachypterous Auchenor-
rhyncha, they live in a more or less two-dimensional habitat without food limitation. In
most other groups of terrestrial Holometabola, however, adults live in the tree or shrub
layer or among tall herbaceous vegetation. In those groups utilizing epigeic resources
(e.g. in saprophagous Diptera and predatory Coleoptera), food is often scattered, un-
predictable or mobile. Hence, macroptery in insects has been suggested to be the crucial
adaptation for the utilization of taller vegetation and predacious life habits (Hennig
1969; Snodgrass 1958; Swain 1978).
(ii) Habitat persistence
Pioneer species, like Laodelphax striatella (Fall.), Javesella pellucida (F.) and some spe-
cies of Macrosteles, are the most striking exception to the rule of prevailing brachyptery
in two-dimensional habitats (see section 6.2.3; Table 35). They show maximum densi-
ties in disturbed or unpredictable habitats, like high-intensity meadows and pastures,
ruderal sites and temporarily dry banks near standing or running water (Novotný 1994a;
Achtziger & Nickel 1997). Even in wing-dimorphic species, brachypters are rare, being
usually confined to habitats, where disturbance dates far back, e.g. old fallows or aban-
doned fields. Novotný (1995) discussed prevailing macroptery in temporary habitats,
combined with high dispersal ability, wide host plant range, wide geographic distribu-
tion and increased annual generation numbers in context with the r and K concept (see
Pianka 1970) and the C-S-R (competitive, stress-tolerant, ruderal) system (Grime 1979),
and suggested the term ‘colonization syndrome’ for the combination of these trails. It
should be noted, however, that Auchenorrhyncha communities on highly disturbed al-
pine river banks, which are subject to frequent flooding and translocation, are charac-
terized by a high proportion of host specialists, monovoltine species and European or
even Alpine endemics (Nickel 1999a).
(iii) Alpine habitats
Increased brachyptery in Auchenorrhyncha is evident above the alpine tree line, where
habitat dimensionality is reduced. The same should be true in higher geographic lati-
 Migration 369

tudes. For further conclusions on the relationship of wing reduction and geographic
altitude and latitude, a careful analysis of faunistic data (e.g. Günthart 1987a, 1997;
Leising 1977; Ossiannilsson 1978, 1981, 1983; Wagner & Franz 1961) along a continuous
gradient is required. Hence, the prevalence of brachyptery in alpine and tundra habi-
tats may simply be caused by treelessness. However, this is unlikely for Carabidae and
some other groups mentioned by Roff (1990).
(iv) Parthenogenesis
Parthenogenesis in central European Auchenorrhyncha is only known in the delpha-
cid planthoppers Muellerianella fairmairei (Perr.) (Booij 1981, 1982), Delphacodes capnodes
(Scott) (Bieman & de Vrijer 1987) and Ribautodelphax pungens (Rib.) (Bieman 1988). It
may be more widespread, but only a few species (all of which are predominantly bra-
chypterous) have been studied so far. Evidence for a correlation of parthenogenesis and
flightlessness was demonstrated and discussed by Bell (1982). As for Auchenorrhyn-
cha, more karyological studies are needed, but the current impression is, that partheno-
genesis is rare and insignificant, since exclusively female field populations appear to be
uncommon.
(v) Paurometabola
According to Roff (1990), secondary flightlessness is most widespread among pau-
rometabolous insects, e.g. Hemiptera and Orthopteroidea, although the causes are not
quite clear. In these groups, migration and macroptery may be not necessary, because
adults have largely the same feeding habits as nymphs and, thus, do not have to move to
different food plants for adult feeding, and then back for oviposition. In addition, small-
scale migration can be performed by the relatively mobile nymphs, and – if habitat condi-
tions deteriorate – by a small proportion of long-winged individuals. In contrast, in most
other hemimetabolous and holometabolous insects, immatures have completely different
life habits than adults, with a marked spatial and ecological segregation. Interestingly, all
Auchenorrhyncha groups for which this statement is also true, notably the Cixiidae, Cica-
didae and Cercopinae, are monomorphic macropterous (see above and section 5.3).

6.2.2 Migration
Much research has been done on the study of migration of Auchenorrhyncha pests of
cereals and other crops, focusing on a few species of economic importance, notably the
eastern Asian planthoppers Nilaparvata lugens (Stål) and Sogatella furcifera (Horv.) as
well as the Nearctic leafhoppers Circulifer tenellus (Bak.) and Macrosteles fascifrons (Stål)
(Kisimoto & Rosenberg 1994; Taylor 1985). Kisimoto (1976) was the first to correlate
long distance migration of delphacid planthoppers with large-scale air movements con-
trolled by synoptic weather conditions, thus creating the basis for predicting the occur-
rence of rice pests in Japan after formation of depressions over the Chinese continent.
Auchenorrhyncha flight strongly depends on the life history of species. Applying
Johnson’s (1969) categories, most individuals emigrate without returning, although the
latter is probably not true for Balclutha punctata (F.) (see Waloff 1980) and other species
overwintering in the adult stage. Malaise trap catches from viticultural areas of south-
western Germany revealed September and October peaks of Acericerus spp., Empoasca
spp., Zygina spp., Arboridia spp. as well as Hauptidia distinguenda (Kbm.), Zyginidia scutel-
370 Life strategies

laris (H.-S.), Fruticidia bisignata (M. & R.) and others (Fröhlich & Nickel, unpublished
data), suggesting enhanced pre-hibernation flight activity in these species. However,
return flight in spring is usually recorded for small numbers only. This may be explained
at least partially by winter mortality.
Waloff (1980) also pointed out, that most univoltine species wintering in the egg stage
show increased flight activity shortly after emergence, i.e. before reproduction, and that
some bivoltine species show different levels of activity between generations. Thus, Er-
rastunus ocellaris (Fall.) was found to fly only in the first generation, and also Hyledel-
phax elegantula (Boh.), Streptanus sordidus (Zett.), Jassargus pseudocellaris (Fl.) and Arthal-
deus pascuellus (Fall.) clearly showed a peak in the first generation. According to Lewis
& Taylor (1965), flight in Cicadellidae may be subject to diurnal periodicity, with most
of the species sampled showing peaks around midday, dawn or dusk, respectively. In
Finnish populations of Balclutha punctata (F.), Raatikainen & Vasarainen (1973) found a
maximum activity around midnight.
For numerous central European Auchenorrhyncha species, there is clear evidence for
mesoscale and probably also macroscale migration (sensu Kisimoto & Rosenberg 1994;
see also Pedgley et al. 1995), i.e. 1-100 km with a duration up to 48 hours as well as more
than 100 km, and lasting up to several days, respectively. Generally, the presence of
single immigrant specimens (inferred from host presence or habitat preference) is an
ubiquitous phenomenon in the field. For instance, in 4 rather small bog areas widely
isolated by woodland on the ridge of the Thuringian Forest, with a small range of po-
tential host plants, the total species number of immigrants recorded during 4 sampling
dates in a single season ranged from 3 to 11, equalling 10 and 24% of the resident spe-
cies (Nickel 2002). The extent of immigration becomes evident when we regard the
Auchenorrhyncha fauna of ruderal sites and mining areas, i.e. in habitats with com-
pletely altered vegetation. Funke & Witsack (1998) recorded altogether 141 species in
surface coal mines of eastern Germany, equalling one third of the regional species di-
versity. These included a considerable proportion of habitat and host specialists as well
as a number of predominantly brachypterous species, which must have immigrated as
macropters or by passive drift within a few years or decades.
Waloff (1973) and della Giustina & Balasse (1999) studied aerial trap catches sampled
at heights up to 12.2 m above ground from Silwood, England, and from 13 sites in north-
ern and eastern parts of France. Accordingly, a large proportion of the local fauna may
be found. In England, within 5 years 115 species were sampled, constituting more than
30% of the British species total. In France, during one year the species number was
altogether 137, equalling almost 17% of the French total. The most abundant species in
Silwood were Javesella pellucida (F.), Macrosteles sexnotatus (Fall.) and M. laevis (Rib.),
accounting for approximately half of the total catch. In France, the total catch included
almost 11,000 individuals. The most common species of both studies are shown in Table
34. Accordingly, most flight activity in Auchenorrhyncha is caused by very few species,
with considerable local differences. Javesella pellucida (F.) was the most common plan-
thopper, although almost absent from 4 sites, but accounting for more than 30% of the
total catch of all French sites, and for 22 and 13% of the Silwood catch in the two years,
respectively. Zyginidia scutellaris (H.-S.) was rather uncommon in Silwood, but abun-
dant in most of the French traps, accounting for 42% of the total catch of these sites.
 Migration 371

Surprisingly, both Macrosteles sexnotatus (Fall.) and M. laevis (Rib.), which are consid-
ered as pioneer species, were almost absent from all the French catches. In Silwood,
however, they accounted for a third of the total catch. These data suggest, that the aerial
trap catch not only depends on the species’ dispersal behaviour, but also on the local
fauna, since all the dominating species in Silwood were common in adjacent study plots.
Furthermore, Laodelphax striatella (Fall.), Javesella dubia (Kbm.), Empoasca pteridis (Dhlb.),
E. decipiens Paoli, E. affinis Nast and E. vitis (Göthe) were common in most of the French
sites. The latter was the only abundant arboricolous species, whereas most of the re-
maining species preferred grassland habitats.
From Germany, there is only a single small-scale study on Auchenorrhyncha dispers-
al by flight. Günthart (1988) sampled 294 individuals from May until September 1984,
using sticky mesh traps suspended on a tower of a meteorological station near Lörrach
(southern Baden) between 5 and 155 m above ground. Apparently, the catch comprised
mostly short-distance migrants. 86% of the specimens were found up to 30 m, i.e. the
height of surrounding trees. Allygus spp., all of which are vertical migrants, clearly dom-
inated (84%). Idiocerine leafhoppers associated with Populus and Salix accounted for
7%. Noteworthy were a few specimens of Cixius nervosus (L.), Laodelphax striatella (Fall.),
Aphrophora salicina (Goeze), Idiocerus stigmaticalis Lew., Stenidiocerus poecilus (H.-S.), Al-
lygus communis (Ferr.) and A. mixtus (F.) sampled in heights between 70 and 155 m.
Therefore, despite the small sample size, this study provides evidence that a consider-
able potential for dispersal is also likely for larger species of Cixiidae, Cercopidae, Idi-
ocerinae and Deltocephalinae.

Table 34. Dominance (%) of the most common Auchenorrhyncha species in aerial traps from 13 localities in
northern and eastern parts of France (after della Giustina & Balasse 1999) and Silwood, England (after
Waloff 1973)

Silw ood 1970

Silw ood 1971

Montpellier
Versailles

Auxerre

Valence
Orléans
Poitiers

Colm ar
Rennes

Angers

Reim s
Caen

Lens

Species
Laodelphax striatella (Fall.) 0.1 12.1 0 1.8 5.2 1.6 0 3.6 12.6 0.2 13.4 2.9 0 0
Toya propinqua (Fieb.) 0 6.3 0 0 0 0 0 0 23.8 0 4.7 0.3 0 0
Javesella pellucida (F.) 13.7 21.9 1.1 4.1 70.3 57.6 82.5 45 0 91.5 0.2 0.1 22.4 12.8
J. dubia (Kbm .) 3.3 18.7 0 1.5 1.8 0.3 0.1 1.2 0.5 0.3 0.3 0.2 -* -*
Z yginidia scutellaris (H .-S.) 69.3 10.7 57.1 65.2 10.4 19.9 12.1 28.1 12.8 2.1 56.7 73.4 -* -*
Empoasca spp. 7.9 6.9 20.9 15.5 1.8 3.5 2.6 8.4 10 4.4 10.6 9,8 -* -*
M acrosteles laevis (Rib.) 0 0 0 0 0 0 0 0 0.2 0 0.1 1 6.4 8.3
M . sexnotatus (Fall.) 0 0 0 0 0 0 0 0 0 0.1 0 0 27 25.5
Others 5.6 23.3 20.9 12 10.6 17.1 2.6 13.7 40.1 1.4 14.1 12.5 44.2 53.4
Total
Total (n individuals) 1860 347 177 950 444 316 876 666 429 1455 1892 1321 1276 1090
* = could not be related to the total
372 Life strategies

6.2.3 Colonization
Colonization of new habitats, i.e. in the state of incipient secondary succession, such as
ruderal sites, river banks, newly sown grassland, mining areas, abandoned fields etc., usu-
ally takes place within one or a few seasons, depending on the speed of plant succession
(Andrzejewska 1991; Hahn 1996; Hildebrandt 1986; Morris 1990a, 1990b; Schiemenz 1964).
Colonization takes also place on cereals and other crops (e.g. Afscharpour 1960; Wais 1989;
also Raatikainen 1972), but reproductive success in these habitats must be limited to one or
two generations at most, due to the removal of host plants by harvesting and ploughing.
Although nymphs of a number of species were found to immigrate a few meters into an
adjacent oatfield (Raatikainen 1972), longer distances can only be covered by flight of adults.
There is a striking similarity among early-successional Auchenorrhyncha communities
all over central Europe, regardless of moisture, pH and other habitat parameters (see Table
35). Pioneer species, like Javesella pellucida (F.), Macrosteles laevis (Rib.), M. sexnotatus (Fall.),
M. cristatus (Rib.), Psammotettix confinis (Dhlb.) and Ps. alienus (Dhlb.), are abundant in most
lowland sites, whereas Empoasca pteridis (Dhlb.), Macrosteles quadripunctulatus (Kbm.),
M. viridigriseus (Edw.) and Ophiola decumana (Kontk.) appear to be correlated with more
specific moisture conditions or soil properties. Laodelphax striatella (Fall.) and Zyginidia scutel-
laris (H.-S.) are both restricted to southern Germany, with occasional influxes into middle
and northern parts. These species show a combination of life history traits which has been
termed ‘colonization syndrome’ (Novotný 1995). In particular, there is a prevalence of high-
er generation numbers, macroptery, broad diet width and wide geographic distribution.
Similar patterns are found in anthropogenic grasslands, like meadows and pastures, which
are subject to frequent disturbance by mowing, grazing or fertilizing. Like ruderal sites, such
habitats can only support highly mobile and polyphagous species, many of which belong to
the genera Javesella, Macrosteles and Psammotettix, which are capable of recolonizing after mow-
ing (Andrzejewska 1979a; Novotný 1994a; Achtziger & Nickel 1997). Although exclosure ex-
periments are still to be done to asess survival rates and subsequent immigration, the herba-
ceous layer of most meadows is apparently almost devoid of insects after mowing. Nickel &
Hildebrandt (2003) speculated, that such habitats may form large population sinks for sur-
rounding habitats, because reproductive success is probably low due to frequent disturbance.
Thus, regarding their susceptibility to the invasion of new species, disturbed habitats show
similar characteristics as early-successional stages. If disturbance has caused extinction of
host plants, phytophagous insects will quickly respond and often become extinct, too. Then
selection will favour r-strategists, i.e. small, short-aged, mobile and fecund species.
A much different pattern is found on almost bare gravel banks of alpine rivers, which are
subject to frequent floodings and translocations. In a preliminary survey conducted mostly
along Bavarian and Tyrolian rivers, Nickel (1999a) found a number of highly stenotopic,
monophagous, wing-dimorphic and univoltine species, most of which show typical traits
of K-strategists, being confined to a small geographic range (Table 35). Pentastiridius beieri
(W.Wg.), Javesella stali (Metc.) and Opsius stactogalus Fieb. are the only species occurring
secondarily in anthropogenic habitats, like sand pits or gardens. Myndus musivus (Germ.) is
common along alpine rivers in northern Italy, but restricted to lowland sites in Germany.
The third group of species listed in Table 35 is also frequently found in early succes-
sional stages, but densities are usually higher in other habitats, notably grassland, rocky
 Colonization 373

Table 35: Auchenorrhyncha species of early successional and disturbed habitats and their life history traits.
Diet width: po = polyphagous, o2 = 2nd degree oligophagous, o1 = 1st degree oligophagous, m2 = 2nd
degree monophagous, m1 = 1st degree monophagous (see Table 4 for definitions). Substrate: P = phloem, X
= xylem, M = mesophyll. Wing length: d = wing-dimorphic, m = macropterous.

Species Diet Sub- Wing Overw inte- Volti- Geographic range

w id th strate length ring stage nism
Pioneer species
Laodelphax striatella (Fall.) po? P d nym ph 2 Palearctic
Javesella pellucida (F.) po? P d nym ph 2 Palearctic
Empoasca pteridis (Dhlb.) po M? m egg? 2? Western Palearctic
Z yginidia scutellaris (H .-S.) o1 M m ad ult 1? Med iterranean
M acrosteles cristatus (Rib.) po P m egg 2 Eurosiberian
M acrosteles laevis (Rib.) po P m egg (1-?)2 Palearctic?
M acrosteles quadripunctulatus (Kbm .) po? P m egg 2 Palearctic?
M acrosteles sexnotatus (Fall.) po P m egg 2 Palearctic
M acrosteles viridigriseus (Ed w .) o2? P m egg 2 European
Ophiola decumana (Kontk.) o1? P m egg 2 Eurosiberian?
Psammotettix alienus (Dhlb.) o1 P m egg 2 Palearctic
Psammotettix confinis (Dhlb.) o1 P d egg 2 Eurosiberian
Specialists of river banks
M yndus musivus (Germ .) m 2? P m nym ph 1 Western Palearctic
Pentastiridius beieri (W.Wg.) o2? P m nym ph 1 European-alpine
Pseudodelphacodes flaviceps (Fieb.) m1 P d nym ph 2 Western Palearctic
Javesella stali (Metc.) m1 P d nym ph 2 Siberian
Opsius stactogalus Fieb. o1 P m egg 1 Western Palearctic
M imallygus lacteinervis (Kbm .) m 1? P d egg 1 European-alpine
Psammotettix unciger Rib. m 1? P m? egg 1? Alpine
Errastunus leucophaeus (Kbm .) ? P d egg 2? Alpine
Species of d isturbed habitats
A siraca clavicornis (F.) po P d ad ult 1 Palearctic
Dicranotropis hamata (Boh.) o1 P d nym ph 1-2 Palearctic
Toya propinqua (Fieb.) o1? P d nym ph 2? Cosm opolitan
Javesella obscurella (Boh.) o1? P d nym ph 2 Palearctic
Ribautodelphax albostriata (Fieb.) m1 P d nym ph 2 Western Palearctic
Dictyophara europaea (L.) po P m egg 1 Western Palearctic
M egophthalmus scanicus (Fall.) o1 P d egg 1 Med iterranean
A naceratagallia venosa (Geoffr.) o2? P d egg 1 Eurosiberian?
A phrodes makarovi Zachv. po P d egg 1 European?
A noscopus albiger (Germ .) o1? P d egg 1 European?
N eoaliturus fenestratus (H .-S.) m 2? P d ad ult (1-?)2 Palearctic
M acrosteles frontalis (Scott) m2 P m egg (1-?)2 Eurosiberian
M acrosteles horvathi (W.Wg.) m2 P m egg 1-2 Eurosiberian
M acrosteles lividus (Ed w .) m 2? P m egg 2 Siberian
Deltocephalus pulicaris (Fall.) o1 P d egg 1-2 Palearctic
Doratura homophyla (Fl.) o1 P d egg 2 Palearctic
Limotettix striola (Fall.) o1? P m egg (1-?)2 Palearctic
Euscelidius schenckii (Kbm .) po P m egg 1 Western Palearctic
Euscelidius variegatus (Kbm .) po P m egg 1 Western Palearctic
Euscelis incisus (Kbm .) o2 P d nym ph/ egg 1-2 Palearctic
Euscelis lineolatus Br. o2 P d egg 1 Med iterranean?
Psammotettix nodosus (Rib.) o1 P m egg 2 European
A rthaldeus pascuellus (Fall.) o1 P d egg 2 Eurosiberian
374 Life strategies

slopes or salt marshes. Principally, however, this list is rather arbitrary, since any Auchen-
orrhyncha species may turn up in any habitat in small numbers, depending on the dis-
tance and position of the closest source populations. The life strategies in this group are
rather heterogenous, but there is at least a tendency towards a broader diet width and
wider distribution (see Table 35). Although the proportion of wing dimorphic species is
high, with predominating brachypters, most populations contain a small percentage of
macropters, which are often found away from typical habitats.
Thus, despite the general prevalence of r-selected life strategies in Auchenorrhyncha
of most early-successional and disturbed habitats, there is the striking exception of riv-
er banks, where just the opposite is true. A third group of species, which prefer dis-
turbed habitats, shows life history traits common both to specialists and generalists.
This prevalence of r-selected species, the dynamics of which must be subject to strong
stochasticity of environmental conditions, should be taken into account in field experi-
ments on newly established study plots.
After a single field experiment at Silwood (Britain), it has been suggested that selec-
tion in early successional habitats favours species overwintering in the adult or nymph-
al stage, which will be gradually replaced by species overwintering as egg (Hollier et al.
1994), and prevailing adult overwintering has also been postulated for Heteroptera
(Brown 1985, 1986). According to the present data, however, this should rather be a
generalization from a singular site. Among species listed in Table 35, the proportion of
egg-overwintering species is c. 66% and thus, almost identical with the proportion of
this group in the Auchenorrhyncha total of Germany (see Table 37). In contrast, species
overwintering as adults are almost absent from early-successional habitats, except Zy-
ginidia scutellaris (H.-S.), which may migrate to other sites in autumn. Asiraca clavicornis
(F.) and Neoaliturus fenestratus (H.-S.) are both localized and uncommon. Therefore, a
correlation of the overwintering stage with successional stage of the habitat is uncer-
tain, although there is weak evidence for species proportions shifting from overwinter-
ing as nymph towards overwintering as adult, with ongoing succession.

6.3 Life cycle

6.3.1 Voltinism
Annual generation numbers have been published for numerous species (Leising 1977;
Müller 1957, 1978, 1981, 1984a; Peter 1978, 1981; Remane 1958, 1987; Schiemenz 1969,
1987, 1988, 1990; Schiemenz et al. 1996; Strübing 1960; Witsack 1985 and others). The
current information on the voltinism of the Auchenorrhyncha fauna of Germany has
been compiled by Nickel & Remane (2002).
The most reliable data have been obtained by laboratory rearings with manipulation of
length of photoperiod and temperature. This tool, however, allows the identification of the
mechanisms, but not necessarily the voltinism in the field. The latter is best studied by
quantitative or semiquantitative sampling every 2 to 4 weeks throughout the season. This
allows a clear documentation of the sequential appearance of nymphs, †† and ‡‡ (in this
order) of each generation. Uncertainties remain, if late nymphs are found, which may be-
 Voltinism 375

long either to the second or a possible third generation, as well as in rare species. For reasons
of sampling methodology, such studies can usually be conducted only in grassland habitats
(see section 3.5), but not in woodland. Thus, some of the present information on the volt-
inism of arboricolous species is rather the result of intuition instead of analysis.
Most field workers have assumed one or two generations per year for central Euro-
pean species. In a few cases, e.g. Chlorita spp., Eupteryx spp., and Deltocephalus pulicaris
(Fall.), three generations have been suggested, but without presenting clear evidence,
usually only referring to late nymphal records. However, most nymphs found in Sep-
tember or even October are unlikely to be capable of reaching adulthood or wintering
successfully. For Macrosteles sexnotatus (Fall.) under laboratory conditions, Witsack (1985)
could clearly show that a photophase below 16 h per day, which is realized in central
Europe after the end of July, induced embryonic dormancy before oviposition. Most
bivoltine species, which overwinter as egg, however, do not reach their second maturi-
ty peak before August. Furthermore, hygric aestivation quiescence, extending the inter-
val between the first and second generation, was found in Euscelis incisus (Kbm.), and
was suspected in other bivoltine species (Witsack 1985). As a consequence, the chance
of an occurrence of a third generation remains – if at all – only after an early spring and
in more southern latitudes, perhaps in viticultural regions of southwestern Germany.
An overview of the generation numbers of the German Auchenorrhyncha species,
extracted from Nickel & Remane (2002), is given in Table 36. The majority, comprising
about 2/3 of the total, is monovoltine (although in some cases, data require confirma-
tion), while most of the remaining species are bivoltine. Apparently, differences between
planthoppers and leafhoppers are negligible. The voltinism of most species appears to
be constant and fixed under central European climates, indicating a strong genetic con-

Table 36. Annual generation numbers of the Auchenorrhyncha species of Germany (1?/2? = probably mono-
/bivoltine; 1(-2?)/2(-3?) = usually mono-/bivoltine, perhaps bi-/trivoltine after early spring or in southern
regions; 1-2 = mono- or bivoltine, depending on regional, altitudinal or weather conditions; (1-?)2 = usually
bivoltine, perhaps monovoltine in higher altitudes or in northern parts)

Generation Auchenorrhyncha total Fulgorom orpha Cicad om orpha

num ber n % n % n %
1 343 55,3 90 62,1 253 53,3
1? 55 8,9 8 5,5 47 9,9
1(-2?) 5 0,8 0 0 5 1,1
1-2 25 4,0 12 8,3 13 2,7
(1-?)2 23 3,7 4 2,8 19 4,0
2 126 20,3 29 20,0 97 20,4
2? 26 4,2 2 1,4 24 5,1
2(-3?) 8 1,3 0 0 8 1,7
<1 9 1,5 0 0 9 1,9
Total 620 100 145 100 475 100
376 Life strategies

trol and a conservative selective force, acting both on Fulgoromorpha and Cicadomor-
pha (see below).
However, there is evidence that at least 53 species may have either one or two gener-
ations a year, depending on local climate and annual weather conditions. It has been
clearly demonstrated in the laboratory, that in reed-dwelling delphacid species the an-
nual generation number is not constant, but may vary even within the same clutch,
depending on the onset of spring (Strübing 1960). Thus, after an early spring, the nymphs
produced by the spring generation reach adulthood in summer, but overwinter if spring
begins lately. There is good field evidence for similar conditions in some species of the
egg-wintering genus Cicadula in southern Lower Saxony. After an early spring, the first
adults appear at the beginning of June, and large numbers of nymphs can be found
again in August and September, thus clearly indicating a second generation. After a late
spring, adults may appear only at the end of June, and their eggs overwinter.
Moreover, there are phenological differences between southern and northern parts of
Germany, which may result in differences in generation numbers. In warm regions of
Thuringia, Franconia and Baden, Cicadella viridis (L.) has been shown to be bivoltine, but
in most other parts, there is only one generation (see Witsack 1985). The earliest adults of
Platymetopius major (Kbm.) frequently emerge at the beginning of June (probably with a
second generation in August) in the Kaiserstuhl, but not before mid-July in southern Lower
Saxony. Likewise, adults of Paralimnus phragmitis (Boh.) appear at the end of May in the
upper Rhine plain, but only around mid-July in northern parts of Germany.
Lowland populations of Euscelis incisus (Kbm.) are bivoltine, wintering as nymph, but
populations in the Erzgebirge studied by Müller (1981) turned out to be univoltine, winter-
ing in the embryonic stage. Witsack (1991) was able to identify the decisive mechanisms.
Macrosteles horvathi (W.Wg.), Cicadula persimilis (Edw.), Arocephalus languidus (Fl.) and other
species, which show a wide altitudinal distribution, are bivoltine in the lowlands, but uni-
voltine at least from the montane belt upwards (see Leising 1977). Variation in voltinism is
also influenced by geographical latitude. Kontkanen (1953) showed that Megadelphax sor-
didula (Stål), Javesella pellucida (F.), Macrosteles sexnotatus (Fall.), M. laevis (Rib.), Deltocephalus
pulicaris (Fall.) and Arthaldeus pascuellus (Fall.), all of which are bivoltine in central European
lowlands, are monovoltine in Finland. In contrast, Rhytidodus decimusquartus (Schrk.), Zy-
ginidia pullula (Boh.) and others have been reported to show up to 4 generations in Mediter-
ranean regions of Italy (Tromellini et al. 1988; Vidano & Arzone 1985).
Altogether, the number of species showing varying annual generation numbers is
probably underestimated, and many of those, which were classified as uncertain (‘prob-
ably monovoltine’ or ‘probably bivoltine’), may in fact be facultatively bivoltine. In the
Bavarian and Allgäu Alps there is no evidence for any species living above 1300 m a.s.l.
to have more than one generation a year. Towards northern Germany (Harz Mountains,
Rothaargebirge, Thuringian Forest), this limit drops to at most 900 m a.s.l. Unfortunate-
ly, field data from this critical altitude are scarce.
Nine species of Cicadomorpha, but none of the Fulgoromorpha, need more than one
year to complete their life cycle, wintering either at least twice as nymph or as nymph and
adult, respectively (see Melber 1989; Müller 1984a). These include the cicadas, which are
generally considered as perennial, although only a few species have been thoroughly
studied (Karban 1986). Development of Cicadetta montana (Scop.) in the vicinity of Mos-
 Dormancy 377

cow (Russia) lasts up to 6 years (Kudryasheva 1975), whereas Tibicina haematodes (Scop.)
takes 3 years (no locality given) (Schedl 2000). The remaining species are all semivoltine.
Both Centrotus cornutus (L.) and Ledra aurita (L.) overwinter twice as nymph, whereas
Ulopa reticulata (F.), U. carneae W.Wg., and probably Eupelix cuspidata (F.) and Errhomenus
brachypterus Fieb. spend their first winter as nymph, and their second as adult.
A common life history trait of some, but not all of these semivoltine and perennial
species is the larger body size and the epigeic or even endogeic habit, causing a reduc-
tion of sun-exposure. In cicadas, xylem-feeding has been suggested to play a further
role, as well as simple demographic factors, selecting prolonged development, if fecun-
dity increases and the mortality is low at least in older instars (Karban 1986). This nutri-
tional hypothesis may also be true for Errhomenus brachypterus Fieb., which is perhaps
another xylem feeder, but its systematic position within the Cicadellinae is uncertain
(Dietrich et al. 2001; R. Rakitov, pers. comm.). All the remaining species, however, prob-
ably feed on phloem sap.
Apart from geographic and climatic conditions, habitat disturbance is likely to be an-
other important factor for the selection of voltinism, favouring bi- and polyvoltine spe-
cies, as well as polyphagous, macropterous and more widespread ones. Thus, bivoltine
species (notably among the genera Javesella, Macrosteles and Psammotettix), have been shown
to dominate in heavily or recently disturbed habitats, such as high-input meadows and
dried pond bottoms. In contrast, they only play a minor role in less disturbed and near-
natural habitats, like fens and low-input meadows (Novotný 1994a, 1994b, 1995; Nickel
& Achtziger 1999). However, this assumption is only true for the herb layer, but not for
the tree canopy. The major part of arboricolous typhlocybine leafhopper species, many of
which are monophagous and confined to the European deciduous forest zone, are clearly
bivoltine, although their microhabitats are largely undisturbed.
A more detailed analysis of generation numbers of the whole central European
Auchenorrhyncha fauna should be based on a more complete data set, with particular
reference to geographical and temporal variation, and further groups of Hemiptera
should be considered. However, in a rather limited study area, and for purpose of com-
parison between different sites, a correlation between annual generation numbers and
certain habitat conditions (e.g. disturbance, successional stage and geographical posi-
tion) may offer interesting aspects of community ecology.

6.3.2 Dormancy
The overwintering stage is a central trait of the life history of all poikilothermic ani-
mals in temperate latitudes. It is of major significance not only for the escape from
adverse climatic conditions, but also for resource utilization and intraspecific synchro-
nization (e.g. Müller 1992; Tauber et al. 1986; Taylor & Karban 1986).
Much laboratory research has been done on the overwintering of central European
Auchenorrhyncha, identifying photoperiod, temperature and water supply as the main
factors controlling dormancy (notably Müller 1958, 1961, 1976, 1981, 1984a; Witsack
1971, 1973, 1981, 1985, 1991; also Drosopoulos 1977; Strübing 1960, 1963). Müller (1970,
1992) and Witsack (1981, 1985, 2002) developed the following classification scheme,
based on induction and termination (Fig. 15).
378 Life strategies

Emancipation
(Avoidance of adverse conditions)

Recession Dormancy
(Migration)

Consecutive dormancy Prospective dormancy

Quiescence Oligopause Parapause Eudiapause Hyperpause

Fig. 15: Dormancy phenomena in arthropods, after Müller (1970, 1992) and Witsack (1981, 1985, 2002)

Emancipation includes all kinds of responses to adverse environmental conditions.

The organism may migrate to more favourable localities (recession), or delay its devel-
opment (dormancy). For purpose of hibernation, dormancy plays the most important
role in central European Auchenorrhyncha. In addition, some species, e.g. Empoasca
spp., Zygina spp., Balclutha spp., show a small-scale migration to other food plants or
other strata (see section 6.2).
Consecutive dormancy is purely facultative, and is induced only after the onset of
adverse conditions. In quiescence, induction as well as termination happen immediate-
ly after environmental conditions, usually temperature or moisture, have changed. Ex-
amples include embryos and occasional adults of Anakelisia fasciata (Kbm.) and nymphs
and occasional embryos of Euscelis incisus (Kbm.), both species overwintering in ther-
mic quiescence. Further, thermic quiescence plays a major role – also among other in-
sect groups – as a secondary dormancy after the termination of oligopause, parapause
or eudiapause, e.g. in Philaenus spumarius (L.), Turrutus socialis (Fl.) and Muellerianella
brevipennis (Boh.). In oligopause, the response happens only gradually, often induced
and terminated cumulatively by the same factor. Examples include Euides basilinea
(Germ.) and Chloriona smaragdula (Stål). Noteworthy is the fact that there are transitions
towards quiescence (quiescent oligopause, not yet found in Auchenorrhyncha) and di-
apausic oligopause, found in Muellerianella brevipennis (Boh.).
Prospective dormancy starts before the onset of adverse conditions. Parapause is
ontogenetically triggered, but terminated by an external factor, e.g. cold, and often fol-
lowed by thermic quiescence, which in turn can be immediately terminated after the
onset of spring. Primary parapause, e.g. found in the oocytes of Stenocranus minutus (F.),
St. major (Kbm.) and Mocydia crocea (H.-S.), can be experimentally prevented by expo-
sure to terminating factors. Secondary parapause is rather fixed and will definitively
progress. It is often found in monovoltine species overwintering in the embryonic stage,
e.g. in Agallia brachyptera (Boh.), Anaceratagallia venosa (Geoffr.) and Elymana sulphurella
(Zett.), or potentially bivoltine, e.g. in Philaenus spumarius (L.), Cicadella viridis (L.) and
Erzaleus metrius (Fl.). Eudiapause is externally controlled and usually induced by pho-
toperiod, but terminated by low temperature. In 4 out of 5 cases studied, species were
bivoltine, and overwintering took place in the embryonic stage of eggs laid by the sec-
ond generation. Examples include Macrosteles sexnotatus (Fall.), Turrutus socialis (Fl.),
 Dormancy 379

Jassargus obtusivalvis (Kbm.) and Arthaldeus pascuellus (Fall.). The fifth species is of more
southern distribution: in eggs of Euscelis alsius Rib., an aestivation eudiapause induced
by long day and accompanied by hygric quiescence was found. This was interpreted as
an adaptation against summer drought in Mediterranean regions. Finally, hyperpause,
which has not yet been found in Auchenorrhyncha, is completely obligatory, and not at
all triggered by external factors.
Apart from these phenomena, which all aim at avoidance of winter cold, Witsack
(1985) found a hygric embryonic quiescence during summer in Euscelis alsius Rib. (see
above) and E. incisus (Kbm.). Later this was also experimentally induced in Macrosteles
sexnotatus (Fall.) and Euscelis lineolatus Br., as well as in the Moroccan E. marocisus Rem.
and the Madeiran E. ormaderensis Rem. (Schöpke 1996), providing further evidence for
aestivation as an important phenomenon even under humid climates. During a three-
year field study in the Saale valley near Jena, Peter (1978, 1981) found a considerable
delay of the second generation peak in a number of bivoltine species in the very dry
summer of 1976. However, this may be an artefact caused by increased mortality due to
drought. In general, aestivation is expected to play a major role in more arid regions, for
instance around the Mediterranean, but not in central Europe, except perhaps in dry
summer periods and in dry habitats.
Altogether, about 30 central European species, belonging to most known types of life
cycles, have been studied in the laboratory by Müller, Strübing and Witsack (references
see above). For most remaining species, field data can be used to assess the overwinter-
ing stage, but not for explaining the exact mechanisms (see Table 37). Roughly two third
(64%) of the total species number winter in the egg stage, 19% winter as nymph and
16% as adult. In 4 semivoltine species, ontogenesis includes one hibernation as nymph
and another as adult, Euscelis incisus (Kbm.) winters as nymph in the lowlands, but as
egg in higher altitudes.
A separate analysis of Fulgoromorpha and Cicadomorpha shows marked differences
(see Table 37). While the proportion of species wintering as egg is only 18.6% in plant-
hoppers, it is 78.4% in leafhoppers. Conversely, only 4.3% of leafhoppers, but 67.5% of

Table 37. Overwintering stages of the Auchenorrhyncha species of Germany

Overw intering Auchenorrhyncha total Fulgorom orpha Cicad om orpha

stage n % n % n %
Egg 379 61,1 27 18.6 352 74.1
Egg? 24 3.9 2 1.4 22 4.6
N ym ph 110 17.7 89 61.4 21 4.4
N ym ph? 7 1.1 7 4.8 0 0
Ad ult 82 13.2 18 12.4 64 13.5
Ad ult? 13 2.1 2 1.4 11 2.3
Mixed 5 0.8 0 0 5 1.1
Total 620 100 145 100 475 100
380 Life strategies

planthoppers, overwinter as nymphs. Differences are only slight in proportions of spe-

cies wintering as adults, whereas overwintering in more than one stage is found in the
Cicadomorpha only. The present data do not allow an explanation for this difference.
Thus, taxon-specific constraints may be involved.
There is a number of ambiguous cases among those species marked as uncertain (‘?’). For
instance, during mild and even moderately cold winters, adults of both Eupteryx melissae
Curt. and E. florida Rib. have been found to survive. The same is true for Anakelisia fasciata
(Kbm.), Megamelus notula (Germ.) and probably some other species. Ribautiana tenerrima
(H.-S.) and R. debilis (Dgl.) are frequently found until the onset of frost. The ecological signif-
icance of occasional overwintering as adults in these species has not been studied, although
most individuals probably die without reproduction. In Mediterranean regions, where oc-
casional warm winter days may allow feeding and other activities, overwintering as adults
may play a more important role. Thus, a number of Mediterranean species reaching central
Europe, clearly favour this overwintering stage, for instance most species of Tettigometridae,
Hauptidia, Arboridia and Mocydiopsis. Müller (1992) reviewed dormancy and other life cycle
traits in arthropods. He listed numerous examples of allopatric and sympatric variation of
life cycles and discussed possible implications on ecology and speciation.
Differences in overwintering stages between Fulgoromorpha and Cicadomorpha have
been interpreted as a temporal niche segregation. For nymphal development, which is sup-
posed to be the most crucial period with maximum demand of nutrients, the Fulgoromor-
pha utilize much more the autumn and early spring, whereas the Cicadomorpha rather
utilize the late spring and early summer. Müller (1978), Peter (1981) and Waloff (1979, 1980)
discussed niche partitioning among grassland Auchenorrhyncha, assuming that potential
interspecific competition should be markedly reduced by temporal segregation.
Other authors suggested an adaptation of Auchenorrhyncha phenology to seasonal
fluctuations of plant nitrogen content, which is highest in spring and autumn, but rela-
tively low in summer (Hill 1982; McNeill & Southwood 1978), as was found in aphids
(Dixon 1985; Risebrow & Dixon 1987; Wellings et al. 1980). In grassland communities of
Auchenorrhyncha in the Leutra valley (Thuringia) and Silwood (England), however,
peaks of total individual numbers were subject to conspicuous annual variation (Waloff
1979; Müller 1978). The reduction of densities during summer in some years were rath-
er explained by drought instead of response to nutrient conditions. Peter (1981) showed
that the maturity peaks of the dominating species in the Leutra valley were evenly
spread over the whole season, ranging from the beginning of April until the end of
August. As a consequence, total individual numbers reached their maximum in early
and high summer. The proportion of monovoltine species wintering in the egg stage,
and reaching their maturity peak in June and July was rather high.
Thus, phenological adaptations to nutrient contents of plants may occur in some species
of Auchenorrhyncha, but the majority shows maturity peaks in early and mid summer,
when nitrogen contents in above-ground parts of plants are rather low, suggesting that
temperature has a much stronger impact on phenology than nitrogen fluctuations. Hill (1982)
demonstrated that the leafhopper Recilia coronifer (Marsh.), which develops in summer, shows
a different nitrogen utilization strategy and uric acid storage, compared with two species
developing in spring. Further research in this field is needed to detect the mechanisms
enabling summer species to compensate the reduced nitrogen content of their hosts.
 Dormancy 381

7 Summary

This work presents an overview of the fauna of leafhoppers and planthoppers (Hemi-
ptera, Cicadomorpha and Fulgoromorpha = Auchenorrhyncha) of Germany and their
life strategies, with special reference to insect – plant relationships. The data base in-
cluded the available literature and the more important museum collections, as well as
extensive own material. During 13 years of field work, approximately 30,000 species
records were gathered, 8,600 of which could be referred to plant species. The number of
determined individuals comprised 300,000 from more than 500 localities in most parts
of Germany.
Part I (sections 1 to 4) gives a short description of the study area, material and meth-
ods, and of literature on identification. Brief accounts on 620 species are presented (145
Fulgoromorpha and 475 Cicadomorpha), with information on life cycles, phenology,
habitat requirements (with emphasis on moisture, sun-exposure and pH), host or food
plants, frequency on the host, abundance, seasonal migrations, geographic and altitu-
dinal distribution as well as economic importance. If necessary, annotations are made
on taxonomic and nomenclatural uncertainties and open questions. Dubious records
are omitted or discussed. Information on recent changes, notably range expansions,
declines, introductions and occasional influxes, is summarized. In part II (section 5), the
Auchenorrhyncha guilds on plant families are described and discussed, with further
discussion of responses to resource type and plant morphology. In part III (section 6),
patterns of resource utilization are described and analyzed. Major points include differ-
ences between main plant clades and families, between different vegetation layers and
different plant substrates. A short overview is given on pests on cultivated plants. Fur-
ther, host specificity, dispersal and life cycles are summarized and discussed.
The feeding preferences on plant groups are rather uneven. Among major clades,
both graminoids and advanced dicotyledonous plants support 45% or more of the total
Auchenorrhyncha species, respectively, although the number of plant species in the
former group is much smaller. Pteridophytes, gymnosperms, non-graminoid monocots
and primitive dicots are only exceptionally attacked. There is convincing evidence that
these groups were only secondarily colonized by host shift from higher plants.
Regarding plant families, highest Auchenorrhyncha species numbers are found on
graminoids, notably Poaceae and Cyperaceae, and woody plants. The former group
accounts for almost 300 species, i.e. half of the total species number, whereas Fagace-
ae, Betulaceae, Salicaceae and Rosaceae are attacked by approximately 50 species,
respectively. Some highly diverse plant groups are not or only rarely attacked, nota-
382 Summary

bly the Caryophyllaceae, Brassicaceae, Apiaceae and Scrophulariaceae. Due to much

differing plant species numbers per family, the ratio of Auchenorrhyncha species :
plant species per plant family is high on woody plants, but relatively low on grami-
noids. Thus, herbivore species richness is highest on plants, which are rich in species,
or biomass, or both. It can be concluded that apparent plants attract highest numbers
of herbivores.
Roughly two third of the total number of Auchenorrhyncha species live in the herb
layer, but only one third lives in the canopies of woody plants. Thus, despite the fact
that central Europe is a woodland country, where biomass is thought to be more con-
centrated in the canopy layer, speciation in Auchenorrhyncha was more pronounced in
the herbacous vegetation. Ten percent of the species are vertical migrants, i.e. they per-
form an obligate or facultative migration from one stratum to another. Some species
groups are particularly interesting due to their endogeic nympal life habits, raising the
question for the evolutionary significance of a diverging morphology and ecology of
immature and adult stages in hemimetabolous versus holometabolous insects.
More than 70% of the total number of Auchenorrhyncha species feed on phloem,
which is considered as the primitive substrate of Hemiptera. Secondarily, some groups
have tapped xylem, mesophyll and fungi mycelium. Species numbers of xylem feeders
on plant families are more strongly correlated with plant species number per family,
than all feeding groups together, providing evidence that plant chemistry of xylem may
play only a negligible role for host choice. Thus, the Caryophyllaceae and Brassicaceae,
which are avoided by phloem and mesophyll feeders, are attacked only by a few spe-
cies of xylem feeders, which generally tend to be more polyphagous. Mesophyll feed-
ers, which account for one fourth of the total fauna, show a high proportion of arbori-
colous species. Feeding on leaf parenchyma is only found in the Typhlocybinae and has
probably evolved along with the rise of angiosperm trees, but out of this group, a num-
ber of species has secondarily performed a host shift back to the herbaceous vegetation,
and colonized primitive or even toxic plants.
Despite widespread crop damage by feeding and transmission of plant diseases in
more southern countries, Auchenorrhyncha play only a minor role as pests in central
Europe, although a few recent cases are known, notably Psammotettix alienus (Dhlb.)
transmitting wheat dwarf virus (WDV), Hyalesthes obsoletus Sign. transmitting grape-
vine yellows (FD), and Empoasca vitis (Göthe) causing ‘hopperburn-like symptoms’ in
vineyards. Other examples are more localized and include Empoasca decipiens Paoli on
greenhouse vegetables, and Eupteryx spp. on medical herbs and spices. Further, the
Nearctic Graphocephala fennahi Young has been suspected to facilitate infection of orna-
mental Rhododendron by a noxious fungus causing bud burst. Invasion of further pest
species from Mediterranean countries, such as Metcalfa pruinosa (Say) and Scaphoideus
titanus Ball may occur in near future.
In general, the relationship between plants and Auchenorrhyncha appears to be rath-
er asymmetrical. Whereas plant apparency (and in particular, distribution, abundance
and size) and plant chemistry are likely to be crucial factors for Auchenorrhyncha oc-
currence, reverse effects are apparently discrete or weak. These effects, however, may
have been underestimated. On the one hand, vascular feeding does not remove living
plant tissue, but, on the other hand, constant removal of resources and transmission of
 Summary 383

pathogens (or at least facilitation of infection) certainly have the potential of causing
damages and persistent weakening of hosts. Thus, even if direct effects may be insignif-
icant, selective feeding may alter competitive relations among plant species. This field
clearly deserves future attention.
A central issue of this book is host specificity. The majority of the Auchenorrhyncha
species of Germany are restricted to one or few plant species. 39% were found to be 1st
degree monophagous (on 1 plant species), 20% are 2nd degree monophagous (on 1
plant genus), and 18% are 1st degree oligophagous (on 1 plant family). The remaining
species were found to feed on more than 1 plant family, or information was not suffi-
cient for a classification. Proportions of specialists were higher in Fulgoromorpha than
in Cicadomorpha. Comparing proportions of host specialists between plant families,
highest values are found in the Poaceae, Cyperaceae, Salicaceae, Pinaceae and Urti-
caceae. It is concluded that richness in both species numbers and biomass of plants not
only attracts herbivores, but also favours their speciation. However, it is further con-
cluded, that herbivore diversity on plants is differentially enhanced by these two plant
properties. On the one hand, there are plant groups comprising few, but tall-growing
species, such as most trees, which favour insect radiation in microhabitats on the same
plant species, whereas graminoids, which are small, but species-rich, favour insect ra-
diation by host shift from one grass to another.
Phragmites australis, the common reed, and Urtica dioica, the stinging nettle, are excep-
tional among herbaceous plants for their high Auchenorrhyncha species numbers, in-
cluding an exceptionally high proportion of monophages. It is suggested that the large
quantities of biomass, which are probably highest among all non-woody plants in central
Europe and elsewhere, favoured herbivore diversity and specialization on these plants.
The Poaceae alone account for at least 115 monophagous Auchenorrhyncha species;
further 61 are found on Cyperaceae. Thus, monophages on these two groups account
for almost 30% of the total number of all Auchenorrhyncha species in Germany. 56 spe-
cialists live on the Salicaceae. In the latter group, it is indirectly concluded that chemical
plant defence mechanisms are involved, since most generalists, which are frequently
encountered on other woody plants, avoid Salicaceae. In contrast, graminoids are known
for their low content of secondary compounds. Instead, they are thought to be more
physically defended by epidermal silica crystals, which are unlikely to be effective against
sap-sucking insects. Further, many grass species grow in mixed stands, particularly in
grassland of anthropogenic origin. Thus, the high proportion of monophages is rather
surprising. Possible explanations include regional monophagy, endophytic fungi and
neural constraints, whereas nitrogen fluctuations and interspecific competition may be
less important.
Concerning dispersal, wing-dimorphic species with prevailing brachyptery domi-
nate in herbaceous vegetation, whereas all arboricolous species and vertical migrants
are macropterous. However, one particular group of herb-dwelling species is also mono-
morphic macropterous, and is largely confined to disturbed habitats and early succes-
sional stages. In this group, most species are bivoltine and polyphagous, and their geo-
graphic ranges are large, whereas mesophyll-feeding and overwintering in the adult
stage are underrepresented.
384 Summary

Two third of the total species number are monovoltine, most of the remaining species
have two annual generations. In only a handful of species, development takes two or
more years. Some species are monovoltine in montane or alpine altitudes, and bivoltine
in the lowlands, but in general, transitions are apparently uncommon. Likewise, the
majority of species (two third) overwinter in the egg stage, whereas the remaining over-
winter as nymph or adult. In very few cases, overwintering may occur in more than one
stage, usually depending on the wheather or on altitude. The prevalence of constant life
cycles indicate a strong genetic control of both voltinism and dormancy.
Future studies in insect – plant relationships should not alone focus on laboratory
experiments on a few widespread generalist species, which can be easily reared, but all
show a similar pattern of life strategies, leading to a biased interpretation. Instead, the
prevalence of host specificity and stenoecious habits in many insect taxa should be tak-
en into account. Life history traits in these groups may diverge distinctly from those of
most generalists. Furthermore, the role of secondary plant compounds may appear in a
different light after a more thorough study of graminoid feeders, which are highly di-
verse not only in Auchenorrhyncha, but also in other insect taxa such as Orthoptera,
Diptera, Hymenoptera and Lepidoptera.
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233-242.
Woodroffe, G.E. (1971): Hemiptera from the Braemer area (Aberdeenshire), including the first
British record of Dikraneura contraria Ribaut (Hem., Cicadellidae). – Ent. month. Mag. 107:
172-173.
Yoshizawa K, Saigusa T. (2001): Phylogenetic analysis of paraneopteran orders (Insecta: Neoptera)
based on forewing base structure, with comments on monophyly of Auchenorrhyncha (Hemi-
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Young, D.A., Frazier, N.W. (1954): A study of the leafhopper genus Circulifer Zachvatkin (Ho-
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Zabel, J., Tscharntke, T. (1998): Does fragmentation of Urtica habitats affect phytophagous and
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Zachvatkin, A.A. (1948): Novye cikady (Homoptera – Cicadina) Srednerusskoy fauny. – Nauch-
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430 Appendix

Appendix

Index of Auchenorrhyncha genera and species

Page numbers are given only for chapter 4.

abbreviatus (Leth.), Allygidius 183 albomarginatus W.Wg., Psammotettix 213

abdominalis (F.), Verdanus 222 albosignata (Dhlb.), Achorotile 42
abietinus (Fall.), Pithyotettix 195 albostriata (Fieb.), Ribautodelphax 62
abstrusus (Lnv.), Kybos 118 albostriella (Fall.), Alebra 112
Acanthodelphax 53 alces (Rib.), Ribautiana 138
Acericerus 100 Alebra 111-113
achilleae Mity., Hephathus 91 alemannicus W.Wg., Jassargus alpinus 219
Achorotile 42 alienus (Dhlb.), Psammotettix 212
acuminatus (F.), Evacanthus 109 allionii (Turt.), Batracomorphus 101
Adarrus 216 f. allobrogicus (Rib.), Jassargus 220
adela (Fl.), Paraliburnia 47 Allygidius 182 f.
adspersa (H.-S.), Eupteryx 144 Allygus 181 f.
adumbrata (C. Shlb.), Rhopalopyx 187 alneti (Dhlb.), Alnetoidia 152
aemulans (Kbm.), Streptanus 204 Alnetoidia 152
aestuarina (Edw.), Aphrodes 103 alni (Fall.), Aphrophora 76
affinis Fieb., Stiroma 41 alni (Schrk.), Oncopsis 82
affinis Nast, Empoasca 121 alnicola (Edw.), Edwardsiana 127
Agallia 91 f. alpestris W.Wg., Cixius 24
Agalmatium 70 alpicola (Cer.), Zyginidia 153
agrestis (Fall.), Stroggylocephalus 108 alpicola W.Wg., Neophilaenus exclamationis 76
Aguriahana 151 alpina Mel., Aphrophora 76
albae W.Wg., Macropsis 84 alpinus (Then), Jassargus 219
albicans (Kbm.), Populicerus 99 alpinus (W.Wg.), Anoscopus 107
albifrons (Fieb.), Mirabella 49 alpinus (Zett.), Macrosteles 169
albifrons (L.), Anoscopus 106 alta Walk., Ceresa 79
albiger (Germ.), Anoscopus 106 alticola Rib., Eupteryx 146
albingensis W.Wg., Cicadula 189 ampliata (W.Wg.), Edwardsiana 130
albipennis (F.), Neophilaenus 74 Anaceratagallia 92 f.
albocarinata (Stål), Nothodelphax 53 Anakelisia 36
albomarginatus Curt., Criomorphus 58 anceps (Germ.), Conomelus 38
 Index of Auchenorrhyncha genera and species 431

angulatus (Then), Psammotettix 211 Balclutha 167 f.

angulosa (Rib.), Ribautodelphax 63 barbata (Rib.), Edwardsiana 134
angusta Leth., Zygina 156 basilinea (Germ.), Euides 43
Anoscopus 106-108 Batracomorphus 101 f.
antennalis (Hpt.), Errastunus 217 bavaricus Rib., Jassargus 220
antoniae (Mel.), Dryodurgades 94 beieri (W.Wg.), Pentastiridius 27
Aphrodes 103 f. beieri W.Wg., Cixius 25
aphrodoides Hpt., Thamnotettix 186 bellevoyei (Put.), Adarrus 217
Aphrophora 76-78 bensoni (China), Verdanus 222
apicalis (Fl.), Empoasca 122 bergmani (Tull.), Edwardsiana 127
appendiculata W.Wg., Oncopsis 82 betulicola (W.Wg.), Kybos 119
Arboridia 160-162 bicarinata (H.-S.), Stiroma 41
arcuata Fieb., Cercopis 72 bicincta (Schrk.), Aphrodes 104
areatus (Stål), Pinumius 221 bifasciata (Boh.), Zonocyba 140
arenarius Rem., Arthaldeus 223 bifasciata (L.), Planaphrodes 105
argentarius Metc., Athysanus 196 bilobum (Fieb.), Agalmatium 70
argus (Marsh.), Sardius 186 binotatus (J. Shlb.), Sonronius 174
Arocephalus 208 f. bipunctata (Osh.), Kyboasca 124
artemisiae (Kbm.), Eupteryx 145 bisignata (M. & R.), Fruticidia 163
Arthaldeus 223 bisonia Kopp & Yonke, Stictocephala 79
Artianus 206 bohemani (Zett.), Diplocolenus 221
Asiraca 29 boica W.Wg., Balclutha 167
assimilis (Fall.), Sorhoanus 224 boldi (Scott), Gravesteiniella 51
assimilis (Sign.), Anoscopus 107 borealis (J. Shlb.), Criomorphus 58
aterrimus (J. Shlb.), Neophilaenus lineatus 75 borussicus W.Wg., Cixius 25
Athysanus 196 f. brabantica W.Wg., Macropsis 89
atomarius (F.), Allygidius 183 Brachyceps 66
atra Hag., Tettigometra 66 brachyptera (Boh.), Agallia 91
atrata Fieb., Tettigometra 66 brachypterus Fieb., Errhomenus 109
atricapillus (Boh.), Limotettix 199 brevipennis (Boh.), Muellerianella 51
atropunctata (Goeze), Eupteryx 142 brevis (H.-S.), Goniagnathus 164
attenuata (Germ.), Mocydiopsis 192 brunnea Mel., Eurysella 40
aubei (Perr.), Muirodelphax 52 bubalus (F.), Ceresa 79
aurantiacus (Forel), Cosmotettix 226 butleri (Edw.), Kybos 117
aurata (L.), Eupteryx 142 calamagrostidis Rem., Lebradea 224
aureola (Fall.), Erythria 113 calamagrostis Oss., Balclutha
aurita (L.), Ledra 82 Calamotettix 226
aurovittata (Dgl.), Lindbergina 137 calcarata Oss., Eupteryx 145
austriaca (Metc.), Eupteryx 142 Calligypona 49
austriaca W.Wg., Anaceratagallia 92 callosa (Then), Ossiannilssonola 126
austriacus W.Wg., Cixius 25 calyculus (Cer.), Kybos 120
Austroasca 124 cambricus China, Cixius 25
avellanae (Edw.), Edwardsiana 127 campestris (Fall.), Neophilaenus 74
avellanae Edw., Oncopsis 83 candidula (Kbm.), Edwardsiana 127
Balcanocerus 101 capnodes (Scott), Delphacodes 50
432 Appendix

carneae W.Wg., Ulopa 80 contraria (Rib.), Emelyanoviana 114

carniolicus W.Wg., Cixius 24 cornicula (Marsh.), Ophiola 198
carpini (J. Shlb.), Oncopsis 83 cornicula (Meusn.), Zyginidia 154
carri (Edw.), Fagocyba 125 cornutus (H.-S.), Enantiocephalus 227
caudatus (Fl.), Cosmotettix 225 cornutus (L.), Centrotus 79
centrorossica Zachv., Aphrodes 104 coronifer (Marsh.), Recilia 176
Centrotus 79 corticea Germ., Aphrophora 77
cephalotes (H.-S.), Psammotettix 213 coryli Le Q., Alebra 112
Ceresa 79 coryli (Toll.), Alnetoidia alneti 152
Cercopis 72 Coryphaelus 166
cerea (Germ.), Macropsis 86 Cosmotettix 225 f.
Chloriona 44-46 costalis Mats., Aphrophora 77
Chlorionidea 52 costalis (Fall.), Cosmotettix 225
Chlorita 124 f., 228 crassicornis (Panz.), Delphax 43
Cicada 70 crataegi (Dgl.), Edwardsiana 128
Cicadella 110 Criomorphus 58 f.
Cicadetta 71 cristatus (Rib.), Macrosteles 169
Cicadula 189-191 crocea (H.-S.), Mocydia 192
Circulifer 166 cruenta (H.-S.), Fagocyba 126
citrinella (Zett.), Forcipata 116 cruentatus (Panz.), Idiodonus 180
Cixidia 64 f. cunicularius (L.), Cixius 23
Cixius 22-26 curtisii (Fl.), Eupteryx 147
clavicornis (F.), Asiraca 29 cuspidata (F.), Eupelix 103
clypealis (J. Shlb.), Paraliburnia 48 cyane (Boh.), Erotettix 174
cognatus (Fieb.), Ebarrius 216 cyclops Mats., Eupteryx 145
coleoptrata (L.), Lepyronia 74 dahlbomi (Zett.), Sonronius 175
coleoptratus (F.), Issus 69 dalei (Scott), Scottianella 56
Colladonus 180 dealbata Cer., Empoasca 123
collina (Boh.), Ribautodelphax 63 debilis (Dgl.), Ribautiana 138
collina (Fl.), Eupteryx 146 decemnotata R., Eupteryx 148
collinus (Boh.), Mocuellus 227 decempunctata (Fall.), Linnavuoriana 136
Colobotettix 196 decimusquartus (Schrk.), Rhytidodus 94
communis (Ferr.), Allygus 181 decipiens Paoli, Empoasca 122
commutatus (Fieb.), Allygidius 182 decumana (Kontk.), Ophiola 198
concinna (Germ.), Eurhadina 140 dehneli Nast, Conomelus lorifer 38
concolor Fieb., Tettigometra 228 Delphacinus 39
confinis (Dhlb.), Psammotettix 215 Delphacodes 50
confinis (Reut.), Streptanus 204 Delphax 43
confinis (Zett.), Cixidia 64 Deltocephalus 175
confinis (Zett.), Thamnotettix 194 denticauda (Boh.), Acanthodelphax 53
confusa Lnv., Kelisia 34 depressa Fieb., Tettigometra 66
confusus (Fl.), Populicerus 99 Dicranotropis 54 f.
Conomelus 38 Dictyophara 65
Conosanus 201 digitatus (Rib.), Kybos 119
consobrina Curt., Agallia 92 Dikraneura 115
 Index of Auchenorrhyncha genera and species 433

dilutior (Kbm.), Thamnotettix 195 Errhomenellus 109

diminuta Rib., Aphrodes 104 Erythria 113
Diplocolenus 221 Erzaleus 227
discolor (Boh.), Javesella 60 Euconomelus 43
discolor (J. Shlb.), Ederranus 204 Euides 43
dissimilis (Fall.), Ommatidiotus 69 Eupelix 103
distincta (Fl.), Nothodelphax 54 Eupterycyba 136
distinguenda (Kbm.), Hauptidia 152 Eupteryx 142-150
distinguendus (Fl.), Jassargus 218 Eurhadina 140 f.
distinguendus (Kbm.), Euscelis 202 europaea (L.), Dictyophara 65
distinguendus (Kbm.), Tremulicerus 98 Eurybregma 41
distinguendus Kbm., Cixius 23 Eurysa 40
Ditropis 39 Eurysella 40
Ditropsis 47 Eurysula 40
divergens Kbm., Dicranotropis 54 Euscelidius 201
diversa (Edw.), Edwardsiana 128 Euscelis 202 f.
Doliotettix 196 Evacanthus 109
domino (Reut.), Hesium 194 excisa (Mel.), Unkanodes 46
douglasi (Edw.), Fagocyba cruenta var. 126 excisus (Mats.), Psammotettix 214
Doratura 177-179 exclamationis (Thnbg.), Neophilaenus 75
dorsata Edw., Chloriona 44 exigua (Boh.), Kosswigianella 55
dorsatum (Ahr.), Haematoloma 73 exilis Horv., Doratura 177
Dryodurgades 94 extrusa (Scott), Muellerianella 52
dubia (Kbm.), Javesella 61 Fagocyba 125 f.
dubius Oss., Psammotettix 214 fairmairei (Perr.), Muellerianella 51
dubius W.Wg., Cixius 26 fallax Rib., Kelisia 33
duffieldi (Le. Q.), Anoscopus 107 fasciata (Kbm.), Anakelisia 36
dumosa (Rib.), Chlorita 124 fenestratus Em., Delphacodes 59
Ebarrius 216 fenestratus (H.-S.), Neoaliturus 165
Ederranus 204 fennahi Young, Graphocephala 111
Edwardsiana 127-136, 229 fieberi (Edw.), Macrosteles 169
elaeagni Em., Macropsis 89 fieberi (Scott), Megamelodes 49
elegans (Fl.), Metidiocerus 96 Fieberiella 163 f.
elegantula (Boh.), Hyledelphax 48 filicum (Newm.), Eupteryx 150
elongata W.Wg., Rhopalopyx 187 flammigera (Geoffr.), Zygina 156
Elymana 188 flava P. Löw, Chlorionidea 52
Emelyanoviana 114 flaveola (Boh.), Paluda 186
emmeae Fieb., Trigonocranus 27 flaveola (Fl.), Xanthodelphax 56
Empoasca 121-123 flavescens (F.), Edwardsiana 129
Enantiocephalus 227 flavescens (F.), Empoasca 123
Endria 176 flaviceps (Fieb.), Pseudodelphacodes 59
erecta (Rib.), Arboridia 160 flavicollis (L.), Oncopsis 83
Erotettix 174 flavipennis (Zett.), Notus 117
Errastunus 217 flavipes (Sign.), Ditropsis 47
Errhomenus 109 flavopunctatus Mel., Errhomenellus 109
434 Appendix

flavostriatus (Don.), Anoscopus 107 grisescens (Zett.), Macustus 196

flavovarius (H.-S.), Handianus 229 Grypotes 164
flavovirens (Gill. & Bak.), Lebradea 224 guentharti Dlab., Edwardsiana 135
flori (Fieb.), Jassargus 219 guttatus Fieb., Platymetopius 180
flori (J. Shlb.), Cicadula 190 guttula (Germ.), Kelisia 30
florida Rib., Eupteryx 147 guttulifera (Kbm.), Kelisia 33
florii (Stål), Fieberiella 163 gyllenhalii (Fall.), Coryphaelus 166
Florodelphax 55 haagii Kbm., Jassus 166
Forcipata 116 f. haematoceps (M. & R.), Circulifer 166
forcipata (Boh.), Javesella 62 haematodes (Scop.), Tibicina 70
forcipata (Fl.), Forcipata 117 Haematoloma 73
formosus (Boh.), Metalimnus 208 hagemini Rem. & Jung, Kelisia 31
forneri Hpt., Aphrophora salicis 77 haglundi (J. Shlb.), Megadelphax 46
franzi (W.Wg.), Zyginidia halpina Rem. & Jung, Kelisia 32
fratercula (Edw.), Edwardsiana 129 hamata (Boh.), Dicranotropis 54
frisia (W.Wg.), Anaceratagallia 92 Handianus 197, 229
froggatti (Bak.), Edwardsiana crataegi var. 128 Hardya 185
frontalis (H.-S.), Cicadula 191 hassicus Nikusch, Cixius haupti 25
frontalis (Scott), Macrosteles 170 haupti Dlab., Cixius 25
frustrator (Edw.), Edwardsiana 130 haupti W.Wg., Kelisia 31
Fruticidia 163 haupti W.Wg., Macropsis 87
fuchsii (Kbm.), Tremulicerus 98 Hauptidia 152 f.
fulgidus (F.), Tremulicerus 98 helvolus (Kbm.), Psammotettix 213
fusca Fieb., Tettigometra 66 henribauti Dlab., Platymetopius 179
fuscinervis (Boh.), Macropsis 88 Hephathus 91
fuscovittatus (Stål), Stenocranus 37 herrichii (Kbm.), Idiocerus 96
fuscula (Zett.), Macropsis 89 Hesium 194
Gargara 78 heydenii (Kbm.), Acericerus 100
genistae (F.), Gargara 78 heydenii (Kbm.), Eupteryx 143
geometrica (Schrk.), Edwardsiana 130 heydenii Kbm., Cixius 24
germari (Zett.), Wagneripteryx 150 hippocastani (Edw.), Edwardsiana 131
glandacea (Fieb.), Macropsis 89 histrionicus (F.), Anoscopus 107
glaucescens Fieb., Chloriona 45 hofferi (Dlab.), Praganus 226
Goniagnathus 164 homophyla (Fl.), Doratura 178
graminea (F.), Macropsis 88 horvathi (Then), Recilia 176
Graphocephala 111 horvathi (W.Wg.), Macrosteles 170
Graphocraerus 183 horvathi W.Wg., Doratura 178
gratiosa Dlab., Dicranotropis 55 Hyalesthes 29
gratiosa (Boh.), Edwardsiana 131 hyalinus (Osb.), Japananus 165
gravesteini Dlab., Delphacodes 56 Hyledelphax 48
gravesteini W.Wg., Macropsis 85 hyperici (H.-S.), Zygina 159
Gravesteiniella 51 hypermaculata Rem. & Hlzg., Zygina 160
grisea Hpt., Aphrophora 77 Iassus 102
griseola Fieb., Tettigometra 67 Idiocerus 95 f.
griseombra Rem., Zygina 159 Idiodonus 180
 Index of Auchenorrhyncha genera and species 435

ignoscus (Mel.), Handianus 197 Laburrus 200

imitans (Rib.), Ribautodelphax 63 lacteinervis (Kbm.), Mimallygus 184
immaculatifrons (Kbm.), Eupteryx 145 laeta H.-S., Tettigometra 66, 67
impictifrons (Boh.), Laburrus 200 laetifica Metc., Tettigometra atra 66, 67
impressifrons (Kbm.), Metidiocerus 96 laeva (R.), Planaphrodes 106
impressopunctata Duf., Tettigometra 65-69 laevis (Rib.), Macrosteles 170
impudica Horv., Doratura 178 lamellaris (Rib.), Edwardsiana 131
impura (Boh.), Macropsis 88 laminatus (Fl.), Populicerus 99
incisa (Then), Wagneriala 116 Lamprotettix 181
incisus (Kbm.), Euscelis 202 languidus (Fl.), Arocephalus 208
inconstans (Rib.), Zygina 157 lanio (L.), Iassus 102
inermis W.Wg., Metropis 42 lanternae (W.Wg.), Edwardsiana 132
inexpectatus Rem., Psammotettix 211 Laodelphax 47
infumatus (Hpt.), Neophilaenus 75 larvatus (H.-S.), Balcanocerus 101
infuscata (J. Shlb.), Macropsis 86 lasiocarpae Oss., Cicadella 110
inquinata (Rib.), Fagocyba cruenta var. 126 latifrons (Kbm.), Metropis 42
intercedens (Lnv.), Linnavuoriana 136 Lebradea 224
intermedia (Boh.), Cicadula 192 Ledra 82
intermedia Rem., Mocydiopsis 192 lelievrei (Leth.), Eupteryx 143
intermedius Fieb., Cixius 23 lepidus (Boh.), Euconomelus 43
interruptus (L.), Evacanthus 109 leporinus (L.), Pentastiridius 28
interstinctus (Fieb.), Ebarrius 216 leptosoma (Fl.), Florodelphax 55
interstitialis (Germ.), Artianus 206 Lepyronia 74
intractabilis Kontk., Limotettix 198 lequesnei W.Wg., Megamelodes 48
intricatus (H.-S.), Phlepsius 184 lethierryi (Edw.), Edwardsiana 131
irregulata Hpt., Kelisia 31 leucophaea (Preyssl.), Tettigometra 68
irroratus Lew., Batracomorphus 102 leucophaeus (Kbm.), Errastunus 217
ishidai (Mats.), Edwardsiana 132 Liguropia 114
Issus 69 limicola (Edw.), Anoscopus 106
Japananus 165 Limotettix 198, 199 f.
Jassargus 218-220 limpidus (W.Wg.), Kybos 118
Jassidaeus 38 lindbergi (Lnv.), Kybos 119
Javesella 60 Lindbergina 137
jucunda (H.-S.), Eupterycyba 136 lineata (Perr.), Eurysa 40
juniperi (Leth.), Liguropia 114 lineatus (L.), Neophilaenus 75
Kelisia 30-35 lineolata (Horv.), Balclutha 167
kemneri (Oss.), Edwardsiana 229 lineolatus Br., Euscelis 202
kirschbaumi W.Wg., Eurhadina 140 Linnavuoriana 136 f.
koeleriae Zachv., Psammotettix 211 Litemixia 52
kolosvarensis (Mats.), Psammotettix 210 littoralis Kuntze, Doratura 179
Kosswigianella 55 lituratus (Fall.), Idiocerus 95
kozhevnikovi (Zachv.), Elymana 188 livens (Zett.), Stroggylocephalus 108
kratochvili (Lang), Arboridia 160 lividus (Edw.), Macrosteles 171
Kyboasca 124 loewii (Then), Eurhadina 141
Kybos 117-121 loginovae (Em.), Arboridia 161
436 Appendix

longicauda Rem., Mocydiopsis 193 mesomelas (Boh.), Delphacinus 39

longiceps (Kbm.), Arocephalus 208 mesopyrrhus (Kbm.), Tremulicerus 98
longicornis Sign., Micrometrina 228 Metalimnus 207 f.
longipennis (Curt.), Stenocranus 36 Metcalfa 235
longispinus W.Wg., Cixius nervosus 22 Metidiocerus 96
longiventris J. Shlb., Cicadula 189 metrius (Fl.), Erzaleus 227
lorifer Rib., Conomelus 38 Metropis 42
ludus (Dav. & De L.), Kybos 120 micantula (Zett.), Micantulina 115
lugens Germ., Ulopa 81 Micantulina 115
lugubrina (Boh.), Struebingianella 56 microcera Vilb., Macropsis 85
lugubris (Sign.), Jassidaeus 38 Micrometrina 228
lunaris (M. & R.), Zygina 155 Mimallygus 184
lunulatus (Zett.), Doliotettix 196 minima (J. Shlb.), Wagneriala 116
lurida (Fieb.), Eurysula 40 minima Rib., Kelisia 34
macrocephala Fieb., Tettigometra 68 minor (Kbm.), Neophilaenus 76
Macropsidius 91 minutus (F.), Stenocranus 37
Macropsis 84-91 Mirabella 49
Macrosteles 169-174 Mitricephalus 66
maculatus Rib., Allygus 182 mixtus (F.), Allygus 181
maculiceps Boh., Deltocephalus 175 mocsaryi (Horv.), Zyginidia 153
maculosus (Then), Macrosteles 171 Mocuellus 227
Macustus 196 Mocydia 192
major (Kbm.), Platymetopius 179 Mocydiopsis 192 f.
major (Kbm.), Stenocranus 37 modestus Scott, Allygus 182
major Hpt., Paramesus 206 moestus (Boh.), Criomorphus 58
major Uhl., Aphrophora 76 mollicula (Boh.), Emelyanoviana 114
makarovi Zachv., Aphrodes 104 monoceros Rib., Kelisia 35
mali (Edw.), Alnetoidia alneti 152 montana (Horv.), Dicranotropis 55
maligna (Walsh), Kyboasca 235 montana (Scop.), Cicadetta 71
manderstjernii (Kbm.), Erythria 113 montanus Geb. & Szwed., Speudotettix 194
marginata (H.-S.), Macropsis 86 monticola Rem., Mocydiopsis 193
marginatus (Kbm.), Streptanus 205 monticola W.Wg., Neophilaenus exclamationis 76
marginicollis (Spin.), Cixidia 65 morbillosus (Mel.), Colobotettix 196
maritimus (Perr.), Psammotettix 209 mucronatus (Rib.), Kybos 120
martigniaca Cer., Edwardsiana candidula 127 Muellerianella 51 f.
Megadelphax 46 Muirodelphax 52
Megamelodes 48 f. mulsanti (Fieb.), Macropsis 91
Megamelus 38 multinotatus (Boh.), Adarrus 216
megerlei (Fieb.), Macropsis 90 muscaeformis (Schrk.), Issus 69
Megophthalmus 81 musivus (Germ.), Myndus 27
melanopsis (Hardy), Hardya 185 Myndus 27
melissae Curt., Eupteryx 148 najas Nast, Macropsis 87
mendax (Fieb.), Macropsis 89 nanus (H.-S.), Hephathus 91
Mendrausus 220 nardeti Rem., Psammotettix 211
meridionalis Bonf., Japananus 165 nebulicola Em., Pinumius 221
 Index of Auchenorrhyncha genera and species 437

nebulosa (Ball), Endria 176 ordinaria (Rib.), Zygina 156

neglecta W.Wg., Alebra 111 orichalceus Thoms., Jassus 198
Neoaliturus 165, 166 origani Zachv., Eupteryx 144
Neophilaenus 74-76 ornata (Mel.), Cicadula 191
nervosa Vilb., Kelisia 34 ornatus (Perr.), Phlepsius 184
nervosus (L.), Cixius 22 orni L., Cicada 70
nigra (Goeze), Penthimia 103 oshanini Razv., Macrosteles 171
nigriloba (Edw.), Edwardsiana 133 ossiannilssoni Ldb., Macrosteles 172
nigrita (Kbm.), Planaphrodes 105 ossiannilssoni Nuort., Empoasca 123
nigritarsis Rem., Zygina 159 Ossiannilssonola 126
nigrolineata Scott, Eurybregma 41 pallens (Stål), Ribautodelphax 228
nitidissimus (H.-S.), Populicerus 99 pallidus (Hpt.), Neophilaenus lineatus 75
nitidulus (F.), Lamprotettix 181 pallidinervis (Dhlb.), Psammotettix 212
nivea (M. & R.), Zygina 155 pallidula (Boh.), Kelisia 33
nodosus (Rib.), Psammotettix 215 Paluda 186
notata (Proh.), Macropsis 86 paludosa (Fl.), Paradelphacodes 57
notata Curt., Eupteryx 150 pandellei (Leth.), Lindbergina 137
notatus (Mel.), Psammotettix 215 panzeri (Fl.), Cosmotettix 226
Nothodelphax 53 f. panzeri (P. Löw), Reptalus 28
notula (Germ.), Megamelus 38 paolii (Oss.), Chlorita 125
Notus 117 Paradelphacodes 57
nubilus Oss., Macrosteles 170 Paradorydium 228
obenbergeri (Dlab.), Verdanus 222 paradoxum (H.-S.), Paradorydium 228
obliqua (W.Wg.), Stiromella 39 Paraliburnia 47 f.
obliqua (Panz.), Tettigometra 68 Paralimnus 207
obscurella (Boh.), Javesella 61 paraltaicus (Orosz), Kybos 120
obsoletus (Germ.), Selenocephalus 229 Paramesus 206
obsoletus (Kbm.), Conosanus 201 Parapotes 206
obsoletus Sign., Hyalesthes 29 parvicauda Rib., Mocydiopsis 193
obtusiceps (Kbm.), Psammotettix 213 parvula (Boh.), Arboridia 161
obtusifrons (Stål), Paramesus 206 paryphasma (Fl.), Florodelphax 55
obtusivalvis (Kbm.), Jassargus 218 pascuellus (Fall.), Arthaldeus 223
ocellaris (Fall.), Errastunus 217 pauxillus (Fieb.), Mendrausus 220
ocellaris (Leth.), Ophiola 198 pectoralis Mats., Aphrophora 77
ocellata Prov., Macropsis 84 Pediopsis 84
ognevi (Zachv.), Ribautiana 139 pellax (Horv.), Laburrus 200
ogumae (Mats.), Streptanus 205 pellucida (F.), Javesella 60
ohausi W.Wg., Euscelis 203 Pentastiridius 27 f.
okaensis Zachv., Streptanus 205 Penthimia 103
Ommatidiotus 69 penthopitta (Walk.), Diplocolenus 221
Oncodelphax 57 Perotettix 195
Oncopsis 82-84 perplexus (Rib.), Kybos 120
opacipennis (Leth.), Circulifer 166 persimilis (Edw.), Cicadula 190
Ophiola 198 f. perspicillata (Boh.), Anakelisia 36
Opsius 165 petryi (Schum.), Cicadetta 71
438 Appendix

Philaenus 78 puncticollis (H.-S.), Grypotes 164

Phlepsius 184 punctifrons (Fall.), Sagatus 175
phragmitis (Boh.), Paralimnus 207 punctulum (Kbm.), Kelisia 32
pictilis (Stål), Aguriahana 151 punctum (Fl.), Arocephalus 209
picturatus (C. Shlb.), Stictocoris 197 pungens (Rib.), Ribautodelphax 64
pictus (Leth.), Perotettix 195 pusilla (Mats.), Chlorita 125
pilatoi D’Urso & Gugl., Cixidia 65 pusilla (Rib.), Arboridia 162
pilosus (Ol.), Tachycixius 26 putoni (Then), Psammotettix 214
Pinumius 221 quadrimaculatus (Sign.), Megamelodes 48
Pithyotettix 195 quadrinotata (F.), Cicadula 191
Planaphrodes 105 quadripunctulatus (Kbm.), Macrosteles 172
planicollis (Thoms.), Macropsis 87 quadrum Boh., Athysanus 197
Platymetopius 179 f. quercus (F.), Typhlocyba 139
plebeja (Edw.), Edwardsiana 133 quinquecostatus (Duf.), Reptalus 28
plurispinosa (W.Wg.), Edwardsiana 131 quinquenotata (Boh.), Cicadula 190, 192
poecilus (Fl.), Psammotettix 209 Recilia 176
poecilus (H.-S.), Stenidiocerus 97 remanei Nick., Macropsis 87
populi (Edw.), Kybos 118 remanei Orosz, Psammotettix 215
populi (L.), Populicerus 100 repletus (Fieb.), Jassargus 219
Populicerus 99 f. Reptalus 28
potentillae (Mor.), Arboridia 161 reticulata (F.), Ulopa 80
praecox Hpt., Kelisia 33 reticulatus (H.-S.), Dryodurgades 94
Praganus 226 reticulatus (Horv.), Parapotes 206
prasina (Boh.), Macropsis 85 reyi (Fieb.), Calligypona 49
preyssleri (H.-S.), Rhopalopyx 187 rhamnicola Horv., Zygina 157
proceps (Kbm.), Rhytistylus 184 rhenana W.Wg., Balclutha 167, 168
procerus (H.-S.), Handianus 229 rhodophila (Cer.), Edwardsiana 129
propinqua (Fieb.), Toya 59 rhombifer (Fieb.), Psammotettix 213
provincialis (Rib.), Hauptidia 153 Rhopalopyx 187 f.
pruinosa (Say), Metcalfa 235 Rhytidodus 94
pruni (Rib.), Balcanocerus 101 Rhytistylus 184
prunicola (Edw.), Edwardsiana 134 ribauti (Oss.), Anaceratagallia 93
Psammotettix 209-215 ribauti (Oss.), Arboridia 161
pseudocellaris (Fl.), Jassargus 218 ribauti Nick. & Rem., Acericerus 100
Pseudodelphacodes 59 ribauti W.Wg., Eurhadina 141
pteridis (Dhlb.), Empoasca 123 ribauti W.Wg., Kelisia 34
pteridis (Spin.), Ditropis 39 Ribautiana 138 f.
pulchellus (Curt.), Delphax 43 Ribautodelphax 62-64, 228
pulchella (Fall.), Eurhadina 141 rosae (L.), Edwardsiana 134
pulchra P. Löw, Zyginella 151 rosaesugans (Cer.), Edwardsiana 134
pulchripennis Asche, Litemixia 52 rosea (Fl.), Zygina 157
pulicaris (Fall.), Deltocephalus 175 roseipennis (Toll.), Zygina 157
pullula (Boh.), Oncodelphax 57 rosincola (Cer.), Zygina 158
pullula (Boh.), Zyginidia 154 rossica Zachv., Edwardsiana solearis 130
punctata (F.), Balclutha 167, 168 rostrata Rib., Eupteryx 234
 Index of Auchenorrhyncha genera and species 439

rotundiceps (Leth.), Paralimnus 207 sima Rib., Kelisia 30

rotundifrons Kbm., Idiocerus 100 similis Kbm., Cixius 26
rubroflava Lnv., Cicadula 189 similis Kbm., Idiocerus 95
rubrovittata (Leth.), Zygina 157 simillima (Lnv.), Javesella 60
rufescens Mel., Kybos 118 simillima (W.Wg.), Arboridia 161
russeola (Fall.), Ophiola 199 simplex (H.-S.), Cixius 23
rutilans (Kbm.), Metidiocerus 96 sinuata (Then), Wagneriala 116
saageri W.Wg., Eurhadina 141 smaragdula (Fall.), Kybos 119
sabulicola (Curt.), Psammotettix 212 smaragdula (Stål), Chloriona 45
sabulicola W.Wg., Kelisia 35 smreczynskii Dwor., Edwardsiana 135
Sagatus 175 sociabilis (Oss.), Edwardsiana 129
sagittarius Rib., Arocephalus 209 socialis (Fl.), Turrutus 218
sahlbergi (Fl.), Macropsidius 91 Sonronius 174 f.
salicicola (Edw.), Edwardsiana 134 sordidipennis (Stål), Macrosteles 173
salicina (Goeze), Aphrophora 78 sordidula (Stål), Megadelphax 46
salina (Hpt.), Javesella 62 sordidus (Zett.), Streptanus 205
saltuella (Kbm.), Balclutha 168 Sorhoanus 224
salviae Arz. & Vid., Eupteryx 148 soror (Lnv.), Edwardsiana 133
sanguinolenta (Scop.), Cercopis 72 sororcula (Oss.), Edwardsiana 129
sanguinosa (R.), Fruticidia 163 Sotanus 204
Sardius 186 spathulata (Rib.), Arboridia 162
sardus Rib., Macrosteles 172 speciosa (Boh.), Euides 43
saturata (Edw.), Cicadula 190 Speudotettix 194
scabripennis Edw., Megophthalmus 81 spinigera (Edw.), Edwardsiana 135
scalaris (Rib.), Ribautiana 138 spinosa (Fieb.), Acanthodelphax 53
scanicus (Fall.), Megophthalmus 81 spoliata (Horv.), Lindbergina 137
schenckii (Kbm.), Euscelidius 201 spumarius (L.), Philaenus 78
schmidtgeni (W.Wg.), Recilia 176 stachydearum (Hardy), Eupteryx 146
schmidti (W.Wg.), Sorhoanus 224 stactogalus Fieb., Opsius 165
schneideri (Günth.), Zygina 158 stali (Metc.), Javesella 62
Scleroracus 198 staminata (Rib.), Edwardsiana 127
scotti Edw., Macropsis 90 stehliki Laut., Edwardsiana 135
Scottianella 56 steini (Fieb.), Metalimnus 208
scutellaris (Fieb.), Iassus 102 stellulata (Burm.), Aguriahana 151
scutellaris (H.-S.), Zyginidia 154 Stenidiocerus 97
scutellata (Boh.), Macropsis 90 Stenocranus 36 f.
Selenocephalus 229 stenoptera (Fl.), Chloriona 45
septemnotatus (Fall.), Macrosteles 173 sticticus R., Cixius 24
septentrionalis W.Wg., Fieberiella 164 Stictocephala 79
serratulae (F.), Anoscopus 108 Stictocoris 197
sexmaculata (Hardy), Linnavuoriana 137 stigmaticalis Lew., Idiocerus 95
sexnotatus (Fall.), Macrosteles 173 stigmaticus (Germ.), Cixius 25
sicula Mats., Chloriona 44 stigmatipennis (M. & R.), Micantulina 115
signatipennis (Boh.), Eupteryx 144 Stiroma 41
signifer (Then), Hardya 185 Stiromella 39
440 Appendix

straminea (Stål), Xanthodelphax 57 tristis (Zett.), Oncopsis 82

Streptanus 204 f. trivia (Germ.), Utecha 81
striatella (Fall.), Laodelphax 47 Turrutus 218
striatula Fall., Cicada 198 Typhlocyba 139
strigilifer (Oss.), Kybos 120 ulmi (Scott), Macropsis 89
striifrons (Kbm.), Arthaldeus 223 ulmi (L.), Ribautiana 139
striola (Fall.), Limotettix 200 ulmiphagus Wils. & Clar., Edwardsiana 132
strobli (W.Wg.), Kybos 121 Ulopa 80
Stroggylocephalus 108 unciger Rib., Psammotettix 210
Struebingianella 56 undatus (De G.), Platymetopius 179
stylata (Boh.), Doratura 177 unicolor Hpt., Aphrophora salicis 77
suavis R., Zygina 157 unicolor (H.-S.), Chloriona 44
subangulata (J. Shlb.), Oncopsis 84 Unkanodes 46
subfusculus (Fall.), Speudotettix 194 untica Dlab., Eurhadina 141
substriatus (Then), Psammotettix 213 urticae (F.), Eupteryx 146
sudeticus (Kol.), Diplocolenus 221 ustulatus (M. & R.), Viridicerus 98
sulphurella (Zett.), Elymana 188 Utecha 81
sursumflexus (Then), Jassargus 220 variata Hardy, Dikraneura 115
Tachycixius 26 variatus (Fall.), Macrosteles 173
taeniatus (Horv.), Calamotettix 226 variegatus (Kbm.), Euscelidius 201
taimyrica Vilb., Hardya 185 vasconica Rib., Chloriona 46
tatraensis (Hell.), Errastunus ocellaris 217 velata (Rib.), Arboridia 162
tenella (Fall.), Eupteryx 149 venosa (Geoffr.), Anaceratagallia 93
tenerrima (H.-S.), Ribautiana 139 venosus (Germ.), Delphacodes 50
tenuis (Germ.), Hardya 185 venosus (Kbm.), Euscelis 203
tersa (Edw.), Edwardsiana 136 ventralis (Fall.), Graphocraerus 183
Tettigometra 65-68, 228 verbae Zachv., Kybos mucronatus 120 f.
Thamnotettix 194 f. Verdanus 222
thenii (P. Löw), Sotanus 204 versuta (Mel.), Arboridia 162
thoulessi Edw., Eupteryx 149 viaduensis (W.Wg.), Zyginidia 154
Tibicina 70 vicina (Horv.), Macropsis 88
tiliae (Fall.), Zygina 158 vicinus Mel., Idiocerus 95
tiliae (Germ.), Pediopsis 84 vinealis Biem., Ribautodelphax 64
tithide Ferr., Zygina 155 virescens (Panz.), Tettigometra 68
torneellus (Zett.), Colladonus 180 virgator (Rib.), Kybos 121
Toya 59 Viridicerus 98
transversa (Fall.), Ophiola 199 viridigriseus (Edw.), Macrosteles 174
tredecimpunctatus (Ldb.), Metalimnus 208 viridinervis W.Wg., Macropsis 85
tremulae (Estl.), Tremulicerus 97 viridis (L.), Cicadella 110
Tremulicerus 97 f. viridis R., Alebra 113
tricolor (Gmel.), Balclutha 167 viridula (Fall.), Chlorita 228
tricolorata Dlab., Calligypona 59 vitis (Göthe), Empoasca 123
tridentata Edw., Anomia 128 vitreus (F.), Tremulicerus 97
trifasciata (Geoffr.), Planaphrodes 105 vitripennis (Fl.), Rhopalopyx 187, 188
Trigonocranus 27 vittata (L.), Eupteryx 149
 Index of Auchenorrhyncha genera and species 441

vittata (Leth.), Austroasca 124 williamsi China, Criomorphus 59

vittifrons (Kbm.), Acericerus 100 xantha Vilb., Xanthodelphax 57
vittipennis (J. Shlb.), Kelisia 32 Xanthodelphax 56 f.
volgensis Vilb., Kybos 121 xanthoneurus (Fieb.), Sorhoanus 224
vulnerata Rossi, Cercopis 72 Xerochlorita Zachv. 124
wagneri China, Cixius 23 Zonocyba 140
Wagneriala 116 Zygina 155-160
Wagneripteryx 150 Zyginella 151
wahlbergi (Boh.), Alebra 112 Zyginidia 153 f.
442 Appendix

Index and vernacular names of plants

Taxonomy and German vernacular names after Wisskirchen & Haeupler (1998),
English vernacular names mainly after Clapham et al. (1987). Page numbers are given
only for chapter 5. Bold numbers indicate tables.

Abies alba Mill. – Silver Fir, Weiß-Tanne 237, 244, 245

Acer L. – Maple, Ahorn 275 f.
Aceraceae 275 f., 332 ff., 342, 349
Acer campestre L. – Field Maple, Feld-Ahorn 275, 276, 323
Acer monspessulanum L. – Montpelier Maple, Französischer Ahorn 275, 276
Acer negundo L. – Ash-leaved Maple, Eschen-Ahorn 275
Acer opalus Mill. – Italian Maple, Frühlings-Ahorn 275
Acer palmatum Thunb. – Japanese Maple, Fächer-Ahorn 275, 276
Acer platanoides L. – Norway Maple, Spitz-Ahorn 275, 276
Acer pseudoplatanus L. – Sycamore, Berg-Ahorn 275, 276
Acer saccharum Marsh. – Sugar Maple, Zucker-Ahorn 275
Acer saccharinum L. – Silver Maple, Silber-Ahorn 275
Achillea millefolium L. s.l. – Yarrow (group), Artengruppe Wiesen-Schafgarbe 279 f., 282 f.
Achnatherum P. Beauv. – Needlegrass, Rauhgras 321
Acinos alpinus (L.) Moench [= Calamintha alpina (L.) Lam.] – Alpine Calamint, Alpen-Steinquendel
Acinos arvensis (Lam.) Dandy [= Calamintha acinos (L.) Clairv.] – Basil Thyme, Feld-Steinquendel
Aegopodium podagraria L. – Goutweed, Giersch 277
Aesculus hippocastanum L. – Horse-chestnut, Rosskastanie 323
Agrostis L. – Bent-grass, Straußgras 309 f.
Agrostis agrostiflora (Beck) Rauschert – Slender Bent-grass, Zartes Straußgras 310
Agrostis alpina Scop. – Alpine Bent-grass, Alpen-Straußgras 310
Agrostis canina L. – Velvet Bent-grass, Hunds-Straußgras 296 ff., 310 f.
Agrostis capillaris L. – Common Bent-grass, Rotes Straußgras 296 ff., 310 f.
Agrostis castellana Boiss. & Reut. – Castilian Bent-grass, Kastilisches Straußgras 310
Agrostis rupestris All. – Rock Bent-grass, Felsen-Straußgras 310
Agrostis scabra Willd. – Rough Bent-grass, Rauhes Straußgras 310
Agrostis stolonifera L. s.l. [incl. A. gigantea Roth] – Creeping Bent-grass (group), Artengruppe
Weißes Straußgras 240, 296 ff., 310
Agrostis vinealis Schreb. [= A. coarctata Ehrh. ex Hoffm..] – Brown Bent-grass, Sand-Straußgras
296 ff., 310, 319
Aira L. – Hair-grass (partim), Haferschmiele 322, 342
Ajuga reptans L. – Bugle, Kriechender Günsel 280 f.
Alnus Mill. – Alder, Erle 253 ff., 342
Alnus alnobetula (Ehrh.) K. Koch [= A. viridis (Chaix) DC.] – Green Alder, Grün-Erle 253 f., 254 f.
Alnus glutinosa (L.) P. Gaertn. – Black Alder, Schwarz-Erle 253 ff., 254 f., 258, 351
Alnus incana (L.) Moench – Grey Alder, Grau-Erle 253 f., 254 f.
Alopecurus L. – Foxtail, Fuchsschwanz 296 ff., 322
Alopecurus aequalis Sobol. – Orange Foxtail, Rotgelber Fuchsschwanz 322
Alopecurus geniculatus L. – Marsh Foxtail, Knick-Fuchsschwanz 322
 Index and vernacular names of plants 443

Alopecurus myosuroides L. – Black Grass, Acker-Fuchsschwanz 322

Alopecurus pratensis L. – Meadow Foxtail, Wiesen-Fuchsschwanz 294, 321
Althaea officinalis L. – Marsh Mallow, Echter Eibisch 325 f., 327, 350
Althaea rosea (L.) Cav. [= Alcea rosea L.] – Hollyhock, Stockrose 325 f., 327
Amaranthaceae 333
Amelanchier lamarckii F.G. Schroed. – Juneberry, Kupfer-Felsenbirne 266 ff., 270
Amelanchier ovalis Medik. – European Serviceberry, Mitteleuropäische Felsenbirne 266 ff., 270
Amelanchier spicata (Lam.) K. Koch – Low Serviceberry, Besen-Felsenbirne 266 ff., 270
Ammophila arenaria (L.) Link – Marram Grass, Gewöhnlicher Strandhafer 296 ff., 319
Andromeda polifolia L. – Bog Rosemary, Rosmarinheide 263
Angelica L. – Angelica, Engelwurz 277
Anthoxanthum alpinum A. Löve & D. Löve – Alpine Vernal-grass, Alpen-Ruchgras 318
Anthoxanthum aristatum Boiss. – Annual Vernal-grass, Grannen-Ruchgras 318
Anthoxanthum odoratum L. – Sweet Vernal-grass, Gewöhnliches Ruchgras 296 ff., 318
Anthriscus Pers. – Chervil, Kerbel 277
Apera Adans. – Silky-bent, Windhalm 322
Apiaceae 276 f., 332 ff., 349
Araliaceae 336
Arctium L. – Burdock, Klette 282 f.
Arctostaphylos Adans. – Bearberry, Bärentraube 263
Aristolochiaceae 329
Arrhenatherum elatius (L.) P. Beauv. ex J. Presl & C. Presl – False Oatgrass, Glatthafer 294, 296 ff.,
315 f., 341
Artemisia L. – Wormwood, Beifuß 279 f.
Artemisia abrotanum L. – Lad’s Love, Eberraute 239, 279, 282 f.
Artemisia absinthium L. – Wormwood, Wermut 282 f.
Artemisia campestris L. – Field Southernwood, Feld-Beifuß 279, 282 f.
Artemisia maritima L. – Sea Wormwood, Strand-Beifuß 239, 279, 282 f.
Artemisia pontica L. – Roman Wormwood, Pontischer Beifuß
Artemisia vulgaris L. – Mugwort, Gewöhnlicher Beifuß 282 f.
Arundo donax L. – Giant Reed, Pfahlrohr 239
Asparagales 322
Asphodelus L. – Asphodel, Affodil 347
Asplenium L. – Spleenwort, Streifenfarn 243
Asplenium scolopendrium L. [= Phyllitis scolopendrium (L.) Newman] – Hart’s-tongue Fern,
Hirschzunge 243
Asteraceae 279 ff., 332 ff., 346, 348 f.
Aster linosyris (L.) Bernh. – Goldilocks, Gold-Aster 279, 282 f.
Athyrium filix-femina (L.) Roth – Lady-fern, Wald-Frauenfarn 243
Avena L. – Oat, Hafer 296 ff., 321, 351
Ballota nigra L. – Black Horehound, Schwarznessel 280 f.
Berberis vulgaris L. – Barberry, Berberitze 324
Beta vulgaris L. – Beet, Runkelrübe 328
Betonica officinalis L. – Betony, Heil-Ziest 278, 280 f.
Betula L. – Birch, Birke 253 ff., 342
444 Appendix

Betulaceae 253 ff., 331 ff., 342, 343, 349

Betula humilis Schrank – Shrub Birch, Strauch-Birke 253
Betula pendula Roth – Silver Birch, Hänge-Birke 253 f., 254 f.
Betula pubescens Ehrh. – Downy Birch, Moor-Birke 237, 253 f., 254 f.
Betula nana L. – Dwarf Birch, Zwerg-Birke 253
Blysmus Panz. ex Schult. – Flat Sedge, Quellbinse 287
Bolboschoenus maritimus (L.) Palla – Sea Club-rush, Gewöhnliche Strandsimse 287, 288 ff.
Boraginaceae 322, 333
Botriochloa ischaemum (L.) Keng – Smooth Finger-grass, Bartgras
Brachypodium P. Beauv. – False-brome, Zwenke 315
Brachypodium pinnatum (L.) P. Beauv. – Tor-grass, Fieder-Zwenke 239, 296 ff., 315
Brachypodium rupestris (Host) Roem. & Schult. – Rock False-brome, Felsen-Zwenke 315
Brachypodium sylvaticum (Huds.) P. Beauv. – Slender False-brome, Wald-Zwenke 315
Brassicaceae 332, 333 ff., 345 f.
Briza media L. – Quaking Grass, Mittleres Zittergras 296 ff., 320
Bromus L. – Brome, Trespe 316, 342
Bromus erectus Huds. – Upright Brome, Aufrechte Trespe 296 ff., 316 f., 341
Bromus inermis Leyss. – Hungarian Brome, Wehrlose Trespe 296 ff., 316 f.
Bromus ramosus Huds. – Hairy Brome, Wald-Trespe 316 f., 342
Bryophyta 329, 339
Buxus sempervirens L. – Box, Buchsbaum 324
Calamagrostis Adans. – Small-reed, Reitgras 308
Calamagrostis arundinacea (L.) Roth – Rough Small-reed, Wald-Reitgras 239, 296 ff., 308
Calamagrostis canescens (Weber) Roth – Purple Small-reed, Sumpf-Reitgras 296 ff., 308
Calamagrostis epigejos (L.) Roth – Wood Small-reed, Land-Reitgras 296 ff., 308, 319, 342
Calamagrostis pseudophragmites (Haller F.) Koel. – Bank Small-reed, Ufer-Reitgras 296 ff., 308
Calamagrostis stricta (Timm) Koeler – Bog Small-reed, Moor-Reitgras 308
Calamagrostis varia (Schrad.) Host – Mountain Small-reed, Buntes Reitgras 296 ff., 308
Calamagrostis villosa (Chaix ex Vill.) J.F. Gmel. – Shaggy Small-reed, Wolliges Reitgras 296 ff., 308
x Calammophila baltica (Flüggé ex Schrad.) Brand – Hybrid Marram, Baltischer Strandhafer 319
Calluna vulgaris (L.) Hull – Heather, Heidekraut 262 ff., 263
Caltha palustris L. – Marsh Marigold, Sumpfdotterblume 247
Calystegia sepium L. s.l. – Hedge Bindweed (group), Artengruppe Gewöhnliche Zaunwinde 328
Campanulaceae 322, 333
Cannabis sativa L. – Hemp, Hanf 248
Caprifoliaceae 337
Capsicum annuum L. – Sweet Pepper, Paprika 350
Caragana arborescens Lam. – Siberian Pea Shrub, Gewöhnlicher Erbsenstrauch
Cardamine pratensis L. – Cuckooflower, Wiesen-Schaumkraut 328
Cardaria draba (L.) Desv. – Hoary Cress, Pfeilkresse 328
Carduus L. – Thistle, Distel (partim) 279, 282 f.
Carex acuta L. [= Carex gracilis Curtis] – Slender Tufted Sedge, Schlank-Segge 288 ff., 292 f., 343
Carex acutiformis Ehrh. – Lesser Pond Sedge, Sumpf-Segge 288 ff., 292 f., 343
Carex alba Scop. – White-flowered Sedge, Weiße Segge 287, 288 ff., 293
Carex arenaria L. – Sand Sedge, Sand-Segge 287, 288 ff.
 Index and vernacular names of plants 445

Carex brizoides L. – Quaking Sedge, Zittergras-Segge 287, 288 ff.

Carex caryophyllea Latourr. [= Carex verna Chaix] – Spring Sedge, Frühlings-Segge 288 ff., 293
Carex distans L. – Distant Sedge, Entferntährige Segge 237, 288 ff., 292, 293
Carex disticha Huds. – Brown Sedge, Zweizeilige Segge 287, 288 ff.
Carex elata All. – Tufted Sedge, Steife Segge 288 ff., 292
Carex elongata L. – Elongated Sedge, Walzen-Segge 288 ff., 293
Carex ferruginea Scop. – Rusty Sedge, Rost-Segge 288 ff., 293
Carex flacca Schreb. – Glaucous Sedge, Blaugrüne Segge 287, 288 ff., 293
Carex flava L. s.l. – Yellow Sedge (group), Artengruppe Gelb-Segge 287, 288 ff.
Carex hirta L. – Hairy Sedge, Behaarte Segge 288 ff., 292
Carex humilis Leyss. – Dwarf Sedge, Erd-Segge 287, 288 ff.
Carex limosa L. – Mud Sedge, Schlamm-Segge 237, 288 ff., 292
Carex montana L. – Soft-leaved Sedge, Berg-Segge 288 ff., 292
Carex muricata L. s.l. – Prickly Sedge (group), Artengruppe Sparrige Segge 288 ff., 293
Carex nigra (L.) Reichard – Common Sedge, Braune Segge 288 ff., 292, 343
Carex panicea L. – Carnation Sedge, Hirsen-Segge 288 ff., 292
Carex paniculata L. – Greater Tussock Sedge, Rispen-Segge 287, 288 ff., 293
Carex pilosa Scop. – Ciliated Sedge, Wimper-Segge 288 ff., 293
Carex pilulifera L. – Pill Sedge, Pillen-Segge 287, 288 ff., 293
Carex pseudocyperus L. – Cyperus Sedge, Scheinzypergras-Segge 288 ff., 293
Carex remota L. – Remote Sedge, Winkel-Segge 288 ff., 293
Carex riparia Curtis – Great Pond Sedge, Ufer-Segge 288 ff., 292 f.
Carex rostrata Stokes – Beaked Sedge, Schnabel-Segge 286, 288 ff., 292
Carex sempervirens Vill. – Evergreen Sedge, Horst-Segge 288 ff., 293
Carex sylvatica Huds. – Wood Sedge, Wald-Segge 288 ff., 293
Carex vesicaria L. – Bladder Sedge, Blasen-Segge 288 ff., 293
Carex vulpina L. s.l. [incl. C. otrubae Podp.] – Fox Sedge (group), Artengruppe Fuchs-Segge 288
ff., 293
Carlina acaulis L. – Greater Carline Thistle, Silberdistel 282 f.
Carlina vulgaris L. – Lesser Carline Thistle, Golddistel 282 f.
Carpinus betulus L. – Hornbeam, Hainbuche 253 f., 254 f.
Caryophyllaceae 327, 332 ff., 345 f., 349
Castanea sativa Mill. – Sweet Chestnut, Esskastanie 239, 250, 251 f.
Catabrosa aquatica (L.) P. Beauv. – Whorl Grass, Quellgras 322
Catapodium rigidum (L.) C.E. Hubb. ex Dony – Fern Grass, Steifgras 322
Ceratophyllaceae 329
Chaerophyllum aromaticum L. – Carrot Chervil, Gewürz-Kälberkropf 277
Chaerophyllum aureum L. – Golden Chervil, Gold-Kälberkropf 277
Chaerophyllum hirsutum L. – Hairy Chervil, Rauhhaariger Kälberkropf 277
Chamaecyparis lawsoniana (A. Murr.) Parl. – Lawson’s False Cypress, Lawson-Scheinzypresse
240, 244, 245, 349
Chamaespartium sagittale (L.) Gibbs [= Genista sagittalis L.; = Genistella sagittalis (L.) Gams] – Winged
Broom, Gewöhnlicher Flügelginster 273, 274
Chenopodiaceae 322, 333, 337
Chrysanthemum vulgare L. – Tansy, Rainfarn 282 f.
446 Appendix

Cirsium acaule Scop. – Dwarf Thistle, Stengellose Kratzdistel 279, 282 f.

Cirsium arvense (L.) Scop. – Creeping Thistle, Acker-Kratzdistel 282 f.
Cirsium oleraceum (L.) Scop. – Cabbage Thistle, Kohldistel 282 f.
Cistaceae 327
Cistus L. – Cistus, Zistrose
Cladium mariscus (L.) Pohl – Great Fen Sedge, Schneidried 287
Clematis vitalba L. – Traveller’s Joy, Gewöhnliche Waldrebe 247
Clinopodium vulgare L. – Wild Basil, Wirbeldost 280 f.
Clusiaceae 327, 349
Coleanthus subtilis (Tratt.) Seidl – Sheath-grass, Scheidenblütgras 322
Colutea arborescens L. – Bladder Senna, Gewöhnlicher Blasenstrauch 273, 274
Convolvulaceae 327
Convolvulus arvensis L. – Field Bindweed, Acker-Winde 326, 327
Corispermum leptopterum (Aschers.) Iljin – Narrow-winged Bugseed, Schmalflügeliger
Wanzensame
Cornaceae 336 f.
Cornus mas L. – Cornelian Cherry, Kornelkirsche 322, 323
Cornus sanguinea L. – Dogwood, Blutroter Hartriegel 322, 323
Coronilla L. – Crownvetch, Kronwicke 273, 274
Corylus avellana L. – Hazel, Gewöhnliche Hasel 253 f., 254 f.
Corylus colurna L. – Tree Hazel, Baum-Hasel 253, 254 f.
Corylus maxima Mill. – Filbert, Lambertsnuss 253, 254 f.
Corynephorus canescens (L.) P. Beauv. [= Weingaertneria canescens (L.) Bernh.] – Grey Hair-grass,
Silbergras 296 ff., 319
Cotoneaster Medik. – Cotoneaster, Zwergmispel 264 f., 266 ff.
Crataegus L. – Hawthorn, Weißdorn 266 ff., 269 f., 272
Crepis L. – Hawk’s-beard, Pippau 279, 282 f.
Crepis paludosa (L.) Moench – Marsh Hawk’s-beard, Sumpf-Pippau 284
Cucumis sativus L. – Cucumber, Gurke 350
Cucurbitaceae 349
Cupressaceae 244 ff., 336 f., 349
Cupressus sempervirens L. – Italian Cypress, Italienische Zypresse
Cydonia oblonga Mill. – Quince, Quitte 270
Cynodon dactylon (L.) Pers. – Bermuda-grass, Hundszahngras 294, 296 ff., 320 f.
Cynosurus cristatus L. – Crested Dogstail, Wiesen-Kammgras 294, 322
Cyperaceae 284 ff., 329, 331 ff., 342, 345 f.
Cyperus L. – Galingale, Zypergras 287
Cyperus longus L. – Tall Galingale, Hohes Zypergras
Cytisus nigricans L. [= Lembotropis nigricans (L.) Griseb.] – Black Broom, Schwarzwerdender Ginster
273, 274
Cytisus scoparius (L.) Link [= Sarothamnus scoparius (L.) W.D. J. Koch] – Broom, Besenginster 273, 274
Dactylis glomerata L. – Cock’s-foot, Wiesen-Knäuelgras 296 ff., 313
Dactylis polygama Horv. – Wood Cock’s-foot, Wald-Knäuelgras 313
Dahlia pinnata Cav. – Dahlia, Dahlie 282 f.
Danthonia decumbens (L.) DC. [= Sieglingia decumbens (L.) Bernh.] – Heath Grass, Dreizahn 321
 Index and vernacular names of plants 447

Daucus carota L. – Wild Carrot, Wilde Möhre 277

Dennstaedtiaceae 336 f.
Deschampsia P. Beauv. – Hair-grass (partim), Schmiele 308 f.
Deschampsia cespitosa (L.) P. Beauv. – Tufted Hair-grass, Rasen-Schmiele 266 ff., 309, 342, 351
Deschampsia flexuosa (L.) Trin. [= Avenella flexuosa (L.) Drejer] – Wavy Hair-grass, Schlängel-
Schmiele 239, 240, 266 ff., 309
Deschampsia littoralis (Gaudin) Reut. – Lake Constance Hair-grass, Bodensee-Schmiele 309
Deschampsia media (Gouan) Roem. & Schult. – Intermediate Hair-grass, Binsen-Schmiele 309
Deschampsia setacea (Huds.) Hack. – Bog hair-grass, Borst-Schmiele 309
Deschampsia wibeliana (Sond.) Parl. – Mud Hair-grass, Elbe-Schmiele 309
Digitalis L. – Foxglove, Fingerhut 349
Digitaria sanguinalis (L.) Scop. – Hairy Finger-grass, Blutrote Fingerhirse 294, 320, 321
Dipsacaceae 327
Dryopteris Adans. – Male-fern, Wurmfarn 243
Ecballium (L.) A. Rich. – Squirting Cucumber, Spritzgurke 349
Echinochloa crus-galli (L.) P. Beauv. – Cockspur, Gewöhnliche Hühnerhirse 321
Echinops L. – Globe Thistle, Kugeldistel
Echium vulgare L. – Viper’s Bugloss, Gewöhnlicher Natternkopf 328
Elaeagnaceae 336 f.
Elaeagnus angustifolia L. – Russian-olive, Schmalblättrige Ölweide 322, 323
Eleocharis palustris (L.) Roem. & Schult. – Common Spike-rush, Gewöhnliche Sumpfbinse 287,
288 ff.
Eleocharis uniglumis (Link) Schult. – Slender Spike-rush, Einspelzige Sumpfbinse 287, 288 ff.
Elymus L. – Couch-grass, Quecke 312 f., 319
Elymus arenosus (Spenn.) Conert – Mainz Couch, Sand-Quecke 313
Elymus athericus (Link) Kerguélen [= Agropyron pungens auct. non (Pers.) Roem. & Schult.] – Sea
Couch, Dünen-Quecke 313
Elymus caninus (L.) L. – Bearded Couch, Hunds-Quecke 313
Elymus farctus (Viv.) Runemark ex Melderis – Sand Couch, Binsen-Quecke 313
Elymus hispidus (Opiz) Melderis – Hairy Couch, Graugrüne Quecke 313
Elymus obtusiflorus (DC.) Conert – Blunt-flowered Couch, Stumpfblütige Quecke 313
Elymus repens (L.) Gould [= Agropyron repens (L.) P. Beauv.] – Couch-grass, Kriech-Quecke 296
ff., 312 f., 342, 351
Epilobium angustifolium L. [= Chamaenerion angustifolium (L.) Scop.] – Fireweed, Schmalblättriges
Weidenröschen 325 f., 327
Equisetaceae 336
Equisetum arvense L. – Field Horsetail, Acker-Schachtelhalm 243
Equisetum palustre L. – Marsh Horsetail, Sumpf-Schachtelhalm 243
Equisetum sylvaticum L. – Wood Horsetail, Wald-Schachtelhalm 243
Eragrostis Wolf – Love-grass, Liebesgras 322, 342
Ericaceae 262 ff., 332 ff., 345 f.
Erica carnea L. – Spring Heath, Schnee-Heide 263, 264
Erica tetralix L. – Cross-leaved Heath, Glocken-Heide 263, 264
Eriophorum angustifolium Honck. – Common Cottongrass, Schmalblättriges Wollgras 286, 288 ff.
Eriophorum gracile W.D.J. Koch ex Roth – Slender Cottongrass, Schlankes Wollgras 286
448 Appendix

Eriophorum latifolium Hoppe – Broad-leaved Cottongrass, Breitblättriges Wollgras 286, 288 ff.
Eriophorum scheuchzeri Hoppe – White Cottongrass, Scheuchzers Wollgras 286
Eriophorum vaginatum L. – Hare’s-tail Cottongrass, Scheiden-Wollgras 286, 288 ff.
Eryngium L. – Eryngo, Mannstreu 347
Euonymus europaeus L. – Spindle Tree, Europäisches Pfaffenhütchen 324
Eupatorium cannabinum L. – Hemp Agrimony, Gewöhnlicher Wasserdost 282 f.
Euphorbiaceae 322, 333
Fabaceae 273 f., 332 ff., 345 f.
Fagaceae 250 ff., 331 ff., 342, 343, 349
Fagus sylvatica L. – Beech, Rotbuche 250, 251 f.
Falcaria vulgaris Bernh. – Longleaf, Sichelmöhre 277
Festuca L. – Fescue, Schwingel 295, 308
Festuca altissima All. – Wood Fescue, Wald-Schwingel 296 ff., 308, 319
Festuca arundinacea Schreb. – Tall Fescue, Rohr-Schwingel 295, 296 ff.
Festuca gigantea (L.) Vill. – Giant Fescue, Riesen-Schwingel 308, 342
Festuca heterophylla Lam. – Various-leaved Fescue, Verschiedenblättriger Schwingel 296 ff., 308
Festuca ovina L. s.l. [incl. F. pallens Host, F. rupicola Heuff., F. valesiaca Schleich. ex Gaudin, etc.] –
Sheep’s Fescue (group), Artengruppe Schaf-Schwingel 295, 296 ff., 308, 319, 320, 342
Festuca pratensis Huds. – Meadow Fescue, Wiesen-Schwingel 296 ff., 308
Festuca puccinellii Parl. – Dark-violet Fescue, Dunkelvioletter Schwingel 295
Festuca pulchella Schrad. – Beautiful Fescue, Schöner Schwingel 295
Festuca quadriflora Honck. – Four-flowered Fescue, Niedriger Schwingel 295
Festuca rubra L. s.l. – Red Fescue (group), Artengruppe Rot-Schwingel 295, 296 ff., 308, 319, 320
Festuca rupicaprina (Hack.) A. Kern. – Chamois Fescue, Gemsen-Schwingel 295
Festuca vivipara (L.) Sm. – Viviparous Fescue, Knollen-Schwingel
Ficus carica L. – Common Fig, Feigenbaum 324
Filipendula ulmaria (L.) Maxim. – Meadowsweet, Echtes Mädesüß 239, 266 ff., 270 f., 349
Forsythia Vahl – Forsythia, Goldflieder 324
Fragaria L. – Strawberry, Erdbeere 266 ff., 270
Frangula alnus Mill. [= Rhamnus frangula L.] – Alder Buckthorn, Faulbaum 241, 322, 323
Fraxinus excelsior L. – Ash, Gewöhnliche Esche 323, 342
Fungi 329 ff.
Galeopsis angustifolia Hoffm. – Red Hemp-nettle, Schmalblättriger Hohlzahn 278, 280 f.
Galium L. – Bedstraw, Labkraut 328, 349
Galium saxatile L. [= Galium harcynicum Weigel] – Heath Bedstraw, Harzer Labkraut
Galium verum L. s.l. – Lady’s Bedstraw (group), Artengruppe Echtes Labkraut
Genista anglica L. – Petty Whin, Englischer Ginster 273
Genista pilosa L. – Hairy Whin, Behaarter Ginster 273, 274
Genista tinctoria L. – Dyer’s Greenweed, Färber-Ginster 273, 274
Gentianaceae 333
Geraniaceae 322, 327, 349
Geranium pusillum Burm. F. – Small-flowered Crane’s-bill, Kleiner Storchschnabel 328
Geranium robertianum L. – Herb-Robert, Stink-Storchschnabel 328
Geranium sanguineum L. – Bloody Crane’s-bill, Blutroter Storchschnabel 325, 327
Glechoma hederacea L. – Ground-ivy, Gundermann 278, 280 f.
 Index and vernacular names of plants 449

Globularia cordifolia L. – Heart-leaf Globularia, Herzblättrige Kugelblume

Glyceria R. Br. – Sweet-grass, Schwaden 320
Glyceria declinata Bréb. – Small Sweet-grass, Blaugrüner Schwaden 320
Glyceria fluitans L. s.l. – Floating Sweet-grass (group), Artengruppe Flutender Schwaden 296 ff., 320
Glyceria maxima (Hartm.) Holmb. [= Poa aquatica L.; = Glyceria spectabilis Mert. & W.D.J. Koch] –
Reed Sweet-grass, Wasser-Schwaden 296 ff., 320
Glyceria nemoralis (Uechtr.) Uechtr. & Körn. – Wood Sweet-grass, Hain-Schwaden 320
Glyceria notata Chevall. – Plicate Sweet-grass, Falt-Schwaden 320
Glyceria striata (Lam.) Hitchc. – Striated Sweet-grass, Streifen-Schwaden 320
Glycyrrhiza glabra L. – Licorice-root, Süßholz 274
Graminoid monocots 329 f.
Gymnospermae 244 ff., 329 ff., 349
Hedera helix L. – Ivy, Efeu 246, 324, 340
Helianthemum alpestre (Jacq.) DC – Mountain Rock-rose, Alpen-Sonnenröschen
Helianthemum nummularium (L.) Mill. – Common Rock-rose, Gewöhnliches Sonnenröschen 325
f., 327
Helianthus annuus L. – Common Sunflower, Sonnenblume 282 f.
Helichrysum arenarium (L.) Moench – Strawflower, Sand-Strohblume 282 f.
Helictotrichon s.l. Besser – Oat-grass (partim), Wiesen-/Flaumhafer 318
Helictotrichon parlatorei (Woods) Pilg. – Alpine Oat-grass, Parlatores Wiesenhafer 318
Helictotrichon pratense (L.) Besser [= Avena pratensis L.; = Avenochloa pratensis (L.) Holub; = Avenula
pratensis (L.) Dum.] – Meadow Oat-grass, Gewöhnlicher Wiesenhafer 296 ff., 318
Helictotrichon pubescens (Huds.) Pilg. – Hairy Oat-grass, Flaumhafer 296 ff., 318, 342
Helictotrichon versicolor (Vill.) Pilg. – Variegated Oat-grass, Bunter Wiesenhafer 318
Heracleum mantegazzianum Sommier & Levier – Giant Hogweed, Riesen-Bärenklau 277
Heracleum sphondylium L. – Cow Parsnip, Wiesen-Bärenklau 277
Hieracium L. – Hawkweed, Habichtskraut 279, 282 f.
Hieracium pilosella L. – Mouse-ear Hawkweed, Mausohr-Habichtskraut 282 f., 284
Hierochloe R. Br. – Holy grass, Mariengras 322
Hippocastanaceae 336 f.
Hippocrepis comosa L. – Horseshoe Vetch, Gewöhnlicher Hufeisenklee 274
Hippophae rhamnoides L. – Sea-buckthorn, Sanddorn 322, 323
Holcus L. – Velvetgrass, Honiggras 284, 311
Holcus lanatus L. – Yorkshire Fog, Wolliges Honiggras 296 ff., 311
Holcus mollis L. – Creeping Soft-grass, Weiches Honiggras 239, 296 ff., 311
Hordelymus europaeus (L.) Jessen ex Harz – Wood Barley, Waldgerste 322, 342
Hordeum L. – Barley, Gerste 296 ff., 321, 342, 350
Horminum L. – Drachenmaul, Dragon’s-mouth 278
Humulus lupulus L. – Hop, Gewöhnlicher Hopfen 328
Hypericum L. – St John’s-wort, Joahnniskraut 349
Hypericum maculatum Crantz s.l. – Imperforate St John’s-wort (group), Artengruppe Geflecktes
Johanniskraut 325 f., 327
Hypericum perforatum L. – Perforate St John’s-wort, Tüpfel-Johanniskraut 325 f., 327
Hypochaeris L. – Cat’s Ear, Ferkelkraut 282 f.
Isolepis R. Br. – Club-rush (partim), Moorbinse 287
450 Appendix

Juglans regia L. – Walnut, Echte Walnuss 324

Juncaceae 284 f., 329, 332 ff., 345
Juncus articulatus L. – Jointed Rush, Glieder-Binse 284, 285
Juncus bufonius L. – Toad Rush, Kröten-Binse 285
Juncus conglomeratus L. – Compact Rush, Knäuel-Binse 285
Juncus effusus L. – Soft Rush, Flatter-Binse 284, 285
Juncus filiformis L. – Thread Rush, Faden-Binse
Juncus gerardii Loisel. – Saltmarsh Rush, Salz-Binse 285
Juncus inflexus L. – Hard Rush, Blaugrüne Binse 284, 285
Juncus subnodulosus Schrank – Blunt-flowered Rush, Stumpfblütige Binse 285
Juniperus L. – Juniper, Wacholder 239
Juniperus communis L. – Common Juniper, Gewöhnlicher Wacholder 244, 245
Juniperus sabina L. – Savin Juniper, Sadebaum 244
Kalmia L. – Laurel, Lorbeerrose 263
Knautia dipsacifolia Kreutzer – Teasel-leaved Knautia, Wald-Witwenblume 325, 327
Kobresia myosuroides (Vill.) Fiori – Bellard’s Kobresia, Europäisches Nacktried 287
Koeleria Pers. – Hair-grass (partim), Schillergras 318
Koeleria arenaria (Dumort.) Conert – Dune Hair-grass, Sand-Schillergras 318
Koeleria glauca (Spreng.) DC. – Glaucous Hair-grass, Blaugrünes Schillergras 296 ff., 318 f.
Koeleria macrantha (Ledeb.) Schult. – Crested Hair-grass, Zierliches Schillergras 318
Koeleria pyramidata (Lam.) P. Beauv. – Pyramidal Hair-grass, Pyramiden-Schillergras 318
Koeleria vallesiana (Honck.) Gaudin – Rock Hair-grass, Walliser Schillergras 318
Lagenaria siceraria (Mol.) Standl. – Calabash, Kalebasse 328
Lamiaceae 277 ff., 332 ff., 345 f., 348 f.
Lamium album L. – White Dead-nettle, Weiße Taubnessel 280 f.
Lamium galeobdolon L. s.l. – Yellow Archangel (group), Artengruppe Goldnessel 280 f.
Lamium maculatum L. – Spotted Dead-nettle, Gefleckte Taubnessel 278, 280 f.
Lapsana communis L. – Nipplewort, Rainkohl 282 f.
Lathyrus L. – Sweet-pea, Platterbse 274
Larix decidua Mill. – European Larch, Europäische Lärche 244, 245
Ledum palustre L. – Labrador Tea, Sumpf-Porst 263
Leersia oryzoides (L.) Sw. – Cut Grass, Reisquecke 324
Leontodon hispidus L. – Rough Hawkbit, Rauher Löwenzahn 279, 282 f.
Leonurus cardiaca L. – Motherwort, Herzgespann 280 f.
Leymus arenarius (L.) Hochst. [= Elymus arenarius L.] – Lyme-grass, Strandroggen 296 ff., 320
Ligustrum vulgare L. – Common Privet, Liguster 246, 323
Liliaceae 333, 337
Liliales 322
Loiseleuria procumbens Desv. – Alpine Azalea, Alpenazalee 263
Lolium perenne L. – Rye-grass, Ausdauerndes Weidelgras 296 ff., 321
Lonicera L. – Honeysuckle, Heckenkirsche 324
Lonicera nigra L. – Black Honeysuckle, Schwarze Heckenkirsche
Lonicera xylosteum L. – Fly Honeysuckle, Rote Heckenkirsche
Lotus corniculatus L. s.l. – Common Birdsfoot-trefoil (group), Artengruppe Gewöhnlicher Hornklee
273, 274
 Index and vernacular names of plants 451

Lotus pedunculatus Cav. [= Lotus uliginosus Schkuhr] – Greater Birdsfoot-trefoil, Sumpf-Hornklee

Lupinus polyphyllus Lindl. – Common Lupin, Vielblättrige Lupine 274
Luzula DC. – Woodrush, Hainsimse 284, 285
Luzula luzuloides (Dam.) Dandy & Wilm. [= Luzula albida (Hoffm.) DC.] – White Woodrush,
Weißliche Hainsimse
Luzula sylvestris (Huds.) Gaudin s.l. – Great Woodrush (group), Artengruppe Wald-Hainsimse
Lycopus europaeus L. – Gipsywort, Ufer-Wolfstrapp 278, 280 f.
Lysimachia vulgaris L. – Yellow Loosestrife, Gewöhnlicher Gilbweiderich 328
magnoliid complex 329
Malus Mill. – Apple, Apfel 265, 266 ff.
Malvaceae 327, 337
Marrubium vulgare L. – White Horehound, Gewöhnlicher Andorn 280 f.
Marsileaceae 324
Marsilea quadrifolia L. – Water-clover, Vierblättriger Kleefarn 324
Medicago lupulina L. – Black Medick, Hopfenklee 273, 274
Medicago sativa L. s.l. – Alfalfa (group), Artengruppe Saat-Luzerne 273, 274
Melica L. – Melick, Perlgras 321
Melica ciliata L. – Hairy Melick, Wimper-Perlgras 321
Melica nutans L. – Mountain Melick, Nickendes Perlgras 321
Melica picta K. Koch – Variegated Melick, Buntes Perlgras 321
Melica transsilvanica Schur – Transsylvanian Melick, Siebenbürger Perlgras 321
Melica uniflora Retz. – Wood Melick, Einblütiges Perlgras 296 ff., 321
Melilotus Mill. – Melilot, Steinklee
Melissa officinalis L. – Balm, Zitronen-Melisse 278, 280 f., 350
Melittis melissophyllum L. – Bastard Balm, Immenblatt 278
Mentha aquatica L. – Water Mint, Wasser-Minze 278, 280 f.
Mentha longifolia (L.) Huds. – Horse Mint, Ross-Minze 278, 280 f.
Mentha x piperita L. – Peppermint, Pfeffer-Minze 280 f.
Mespilus L. – Medlar, Mispel 270
Mibora minima (L.) Desv. – Early Sand-grass, Zwerggras 322
Micropyrum tenellum (L.) Link – Gravel Fescue, Kies-Dünnschwingel 322
Milium effusum L. – Wood Millet, Flattergras 322, 342
Molinia caerulea (L.) Moench s.l. – Purple Moor-grass (group), Artengruppe Pfeifengras 239,
240, 296 ff., 316
Myricaceae 336
Myricaria germanica (L.) Desv. – German False Tamarisk, Deutsche Tamariske 239, 322, 323
Myrica gale L. – Bog Myrtle, Gagelstrauch 324
Nardus stricta L. – Mat-grass, Borstgras 296 ff., 320
Nasturtium W.T. Aiton – Water-cress, Brunnenkresse
Nepeta cataria L. – Cat-mint, Gewöhnliche Katzenminze 280 f.
non-monocot paleoherbs 329
Nicotiana tabacum L. – Tobacco, Tabak 351
Nothofagus Blume – Southern Beech, Südbuche
Nuphar lutea (L.) Sibth. & Sm. – Yellow Water-lily, Gelbe Teichrose 324, 325
Nymphaea alba L. – White Water-lily, Weiße Seerose 324, 325
452 Appendix

Nymphaeaceae 324, 329

Ocimum basilicum L. – Basil, Basilikum 278, 280 f., 351
Oleaceae 323, 336 f.
Onagraceae 322, 327, 337
Onobrychis Mill. – Sainfoin, Esparsette 273, 274
Ononis spinosa L. – Spiny Restharrow, Dornige Hauhechel 273, 274
Onopordum acanthium L. – Scotch Thistle, Eselsdistel 282 f.
Orchidaceae 322, 333, 337
Origanum vulgare L. – Marjoram, Gewöhnlicher Dost 278, 280 f.
Orobanche L. – Broomrape, Sommerwurz 346
Paeonia L. – Peony, Pfingstrose
Panicum L. – Millet (partim), Rispenhirse 322
Papaveraceae 333
Parapholis strigosa (Dumort.) C.E. Hubb. – Hard Grass, Gekrümmter Dünnschwanz 322
Parietaria judaica L. – Common Pellitory-of-the-Wall, Mauer-Glaskraut 248 f., 328
Parietaria officinalis L. – Upright Pellitory-of-the-Wall, Aufrechtes Glaskraut 248
Pastinaca sativa L. – Wild Parsnip, Pastinak 277
Petasites Mill. – Butterbur, Pestwurz 279, 282 f.
Phalaris arundinacea L. [= Typhoides arundinacea (L.) Moench; = Baldingera arundinacea (L.) Dumort.]
– Reed-grass, Rohr-Glanzgras 296 ff., 317
Phleum L. – Cat’s-tail, Lieschgras 294, 296 ff., 321, 351
Phleum pratense L. – Timothy, Wiesen-Lieschgras 321
Phleum phleoides (L.) H. Karst. – Purple-stem Cat’s-tail, Steppen-Lieschgras 322
Phleum alpinum L. – Alpine Cat’s-tail, Alpen-Lieschgras 322
Phleum hirsutum Honck. – Rough Cat’s-tail, Matten-Lieschgras 322
Phlomis L. – Jerusalem Sage, Brandkraut 278
Phragmites australis (Cav.) Trin. ex Steud. [= Phragmites communis Trin.] – Common Reed,
Gewöhnliches Schilf 239, 260, 296 ff., 311 f., 333, 342
Picea abies (L.) H. Karst. – Norway Spruce, Gewöhnliche Fichte 244, 245, 246, 259
Picea omorika (Pancic) Purkyne – Serbian Spruce, Serbische Fichte
Picris hieracioides L. – Hawkweed Oxtongue, Gewöhnliches Bitterkraut 282 f.
Pinaceae 332 ff., 342
Pinus L. – Pine, Kiefer 244, 330, 343, 349, 351
Pinus cembra L. – Stone Pine, Zirbel-Kiefer 244, 245
Pinus mugo Turra – Mountain Pine, Latschen-Kiefer 244, 245
Pinus nigra J.F. Arnold – Black Pine, Schwarz-Kiefer 239, 244, 245
Pinus sylvestris L. – Scots Pine, Wald-Kiefer 239, 244, 245, 246
Plantaginaceae 327
Plantago lanceolata L. – Ribwort Plantain, Spitz-Wegerich 325 f., 327
Plantago major L. – Common Plantain, Breit-Wegerich 325 f., 327
Plantago media L. – Hoary Plantain, Mittlerer Wegerich 325 f., 327
Platanaceae 336
Platanus L. – Plane, Platane 323
Poa L. – Meadow-grass, Rispengras 313 f., 351
Poa alpina L. – Alpine Meadow-grass, Alpen-Rispengras 314
 Index and vernacular names of plants 453

Poa annua L. – Annual Meadow-grass, Einjähriges Rispengras 296 ff., 314

Poa badensis Haenke ex Willd. – Baden Meadow-grass, Badener Rispengras 314
Poaceae 294 ff., 329, 331 ff., 342, 343, 345 f.
Poa chaixii Vill. – Broad-leaved Meadow-grass, Wald-Rispengras 314
Poa compressa L. – Flattened Meadow-grass, Platthalm-Rispengras 314
Poa hybrida Gaudin – Hybrid Meadow-grass, Großes Rispengras 314
Poa nemoralis L. – Wood Meadow-grass, Hain-Rispengras 296 ff., 314
Poa palustris L. – Swamp Meadow-grass, Sumpf-Rispengras 315
Poa pratensis L. s.l. [incl. P. angustifolia L.] – Smooth Meadow-grass (group), Artengruppe Wiesen-
Rispengras 296 ff., 314
Poa supina Schrad. – Prostrate Meadow-grass, Läger-Rispengras 314
Poa trivialis L. – Rough Meadow-grass, Gewöhnliches Rispengras 296 ff., 314
Polygonaceae 327, 337, 346
Polygonum aviculare L. s.l. – Knotgrass (group), Artengruppe Gewöhnlicher Vogelknöterich 325, 327
Polypodium interjectum Shivas – Western Polypody, Gesägter Tüpfelfarn 243, 349
Polypodium vulgare L. – Common Polypody, Gewöhnlicher Tüpfelfarn 243, 349
Polystichum Roth. – Shield-fern, Schildfarn 243
Populus L. – Poplar, Pappel 258 ff.
Populus alba L. – White Poplar, Silber-Pappel 258, 260, 261 f.
Populus nigra L. [incl. Populus nigra “italica” (Duroi) Moench] – Black Poplar [Lombard Poplar],
Schwarz-Pappel [Pyramiden-Pappel] 258 ff, 261 f..
Populus tremula L. – Aspen, Zitter-Pappel 258, 261 f., 340
Potamogetonaceae 324
Potamogeton natans L. – Broad-leaved Pondweed, Schwimmendes Laichkraut 324, 325
Potentilla L. – Cinquefoil, Fingerkraut 349
Potentilla fruticosa L. – Shrubby Cinquefoil, Strauch-Fingerkraut 266 ff., 270
Potentilla incana P. Gaertn., B. Mey. & Scherb. [= Potentilla arenaria P. Gaertn., B. Mey. & Scherb.] –
Sand Cinquefoil, Sand-Fingerkraut 266 ff., 271
Potentilla tabernaemontani Asch. [= Potentilla neumanniana Rchb.] – Spring Cinquefoil,
Gewöhnliches Frühlings-Fingerkraut 266 ff., 271
Primulaceae 322, 333, 349
Prunella vulgaris L. – Selfheal, Kleine Braunelle 280 f.
Prunus L. – Cherry, Plum, Kirsche, Pflaume 271 ff.
Prunus avium L. [= Cerasus avium Moench] – Wild Cherry (incl. Sweet Cherry), Vogel-Kirsche
(incl. Süß-Kirsche) 266 ff., 271 f.
Prunus cerasus L. s.l. – Sour Cherry (group), Artengruppe Sauerkirsche 266 ff., 271
Prunus domestica L. – Plum, Zwetschge, Pflaume 266 ff., 271
Prunus dulcis (Mill.) D.A. Webb – Almond, Mandel 271
Prunus fruticosa Pall. – Dwarf Cherry, Zwerg-Kirsche 271
Prunus laurocerasus L. – Laurel Cherry, Lorbeer-Kirsche 271
Prunus mahaleb L. – Mahaleb Cherry, Felsen-Kirsche 271
Prunus padus L. – Bird Cherry, Trauben-Kirsche 266 ff., 271 ff.
Prunus persica (L.) Batsch – Peach, Pfirsich 271
Prunus serotina Ehrh. – Black Cherry, Späte Traubenkirsche 271
Prunus spinosa L. – Blackthorn, Schlehe 266 ff., 271 ff.
454 Appendix

Pseudotsuga menziesii (Mirb.) Franco – Douglas Fir, Douglasie 244, 245

Pteridium aquilinum (L.) Kuhn – Bracken, Gewöhnlicher Adlerfarn 243
Pteridophyta 242 f., 324, 329 ff.
Puccinellia Parl. – Saltmarsh-grass, Salzschwaden 317
Puccinellia capillaris (Lilj.) Jansen – Northern Saltmarsh-grass, Haar-Salzschwaden 317
Puccinellia distans (Jacq.) Parl. – Reflexed Saltmarsh-grass, Gewöhnlicher Salzschwaden 296 ff., 317
Puccinellia limosa (Schur) E. Holmb. – Mud Saltmarsh-grass, Sumpf-Salzschwaden 317
Puccinellia maritima (Huds.) Parl. – Common Saltmarsh-grass, Andel 296 ff., 317
Pyracantha coccinea Roem. – Fire Thorn, Feuerdorn 266 ff., 270
Pyrus communis L. s.l. - Pear (group), Artengruppe Birnbaum 265, 266 ff., 351
Quercus L. – Oak, Eiche 250 ff., 342
Quercus cerris L. – Turkey Oak, Zerr-Eiche 250
Quercus petraea Liebl. – Durmast Oak, Trauben-Eiche 239, 250 ff., 251 f.
Quercus pubescens Willd. – Downy Oak, Flaum-Eiche 250, 251 f.
Quercus robur L. – Common Oak, Stiel-Eiche 250 ff., 251 f.
Quercus rubra L. – Northern Red Oak, Rot-Eiche 250
Ranunculaceae 246, 332 ff., 345 f., 349
Ranunculus bulbosus L. – Bulbous Buttercup, Knolliger Hahnenfuß 247
Ranunculus montanus Willd. s.l. – Mountain Buttercup, Berg-Hahnenfuß 247
Ranunculus repens L. – Creeping Buttercup, Kriechender Hahnenfuß 246, 247
Rhamnaceae 336 f.
Rhamnus cathartica L. – Buckthorn, Purgier-Kreuzdorn 322, 323
Rhododendron L. – Rhododendron, Alpenrose 239, 263, 264, 350
Rhodothamnus Rchb. – Dwarf-rhododendron 263
Rhynchospora alba (L.) Vahl – White Beak-sedge, Weißes Schnabelried 287, 288 ff.
Ribes L. – Currant, Johannisbeere 324
Robinia pseudoacacia L. – Acacia, Robinie 324
Rosaceae 264 ff., 331 ff., 349
Rosa L. – Rose, Rose 271, 350
Rosa canina L. – Dog Rose, Hunds-Rose 266 ff., 271
Rosa corymbifera Borkh. – Corymb Rose, Hecken-Rose 266 ff., 271
Rosa glauca Pourr. – Glaucous Rose, Bereifte Rose 266 ff.
Rosa montana Chaix – Mountain Rose, Berg-Rose
Rosa pendulina L. – Alpine Rose, Alpen-Rose 266 ff., 271
Rosa rubiginosa L. – Sweet Briar, Wein-Rose 266 ff., 271
Rosa rugosa Thunb. – Japanese Rose, Kartoffel-Rose 239, 266 ff., 271, 349
Rosa spinosissima L. [= Rosa pimpinellifolia L.] – Burnet Rose, Bibernell-Rose 266 ff., 271
Rubiaceae 337, 346, 349
Rubus L. – Bramble, Raspberry, Brombeere, Himbeere 340
Rubus caesius L. – Dewberry, Kratzbeere 266 ff., 270
Rubus chamaemorus L. – Cloudberry, Moltebeere 270
Rubus fruticosus L. s.l. – Bramble (group), Artengruppe Echte Brombeere 241, 266 ff., 270
Rubus idaeus L. – Raspberry, Himbeere 239, 266 ff., 270
Rubus saxatilis L. – Stone Bramble, Steinbeere 270
Rumex acetosella L. – Sheep’s Sorrel, Kleiner Sauerampfer 325 f., 327
 Index and vernacular names of plants 455

Salicaceae 258 ff., 331 ff., 342, 345 f.

Salix L. – Willow, Weide 258 ff.
Salix alba L. – White Willow, Silber-Weide 258 ff., 261 f., 340
Salix appendiculata Vill. – Large-leaved Willow, Großblättrige Weide 258, 260, 261 ff.
Salix aurita L. – Eared Willow, Ohr-Weide 258 ff., 261 f.
Salix bicolor Willd. – Tea Willow, Zweifarbige Weide 258
Salix caprea L. – Goat Willow, Sal-Weide 258 ff., 261 f.
Salix cinerea L. – Grey Willow, Grau-Weide 258 ff., 261 f.
Salix daphnoides Vill. – Violet Willow, Reif-Weide 258
Salix eleagnos Scop. [= Salix incana Schrank] – Olive Willow, Lavendel-Weide 258 ff., 261 f.
Salix fragilis L. – Crack Willow, Bruch-Weide 258 ff., 261 f., 340
Salix hastata L. – Halberd Willow, Spieß-Weide 258
Salix herbacea L. – Forb Willow, Kraut-Weide 258
Salix myrsinifolia Salisb. [= Salix nigricans Sm.] – Dark-leaved Willow, Schwarzwerdende Weide
258 ff., 261 f.
Salix pentandra L. – Bay Willow, Lorbeer-Weide 258 ff., 261 f.
Salix purpurea L. – Purple Willow, Purpur–Weide 258 ff., 261 f.
Salix repens L. s.l. – Creeping Willow (group), Artengruppe Kriech-Weide 258 ff., 261 f.
Salix reticulata L. – Reticulate Willow, Netz-Weide 258
Salix retusa L. – Blunt-leaved Willow, Stumpfblättrige Weide 258
Salix triandra L. – Almond Willow, Mandel-Weide 258 ff., 261 f., 340
Salix viminalis L. – Osier, Korb-Weide 258 ff., 261 f.
Salvia officinalis L. – Sage, Echter Salbei 278, 280 f., 350
Salvia nemorosa L. – Violet Sage, Steppen-Salbei 280 f.
Salvia pratensis L. – Meadow Clary, Wiesen-Salbei 280 f.
Salvia verticillata L. – Whorled Clary, Quirlblütiger Salbei 280 f.
Sambucus ebulus L. – Danewort, Zwerg-Holunder 328
Sambucus nigra L. – Elder, Schwarzer Holunder 323
Sanguisorba minor Scop. – Salad Burnet, Kleiner Wiesenknopf
Satureja L. – Savory, Bohnenkraut 278
Saxifragaceae 333
Schoenoplectus lacustris (L.) Palla – Bulrush, Gewöhnliche Teichsimse 286, 288 ff.
Schoenoplectus tabernaemontani (C.C. Gmel.) Palla – Grey Club-rush, Salz-Teichsimse 286, 288 ff.
Schoenus ferrugineus L. – Brown Bog-rush, Rostrotes Kopfried 287, 288 ff.
Schoenus nigricans L. – Black Bog-rush, Schwarzes Kopfried 287, 288 ff.
Scirpoides holoschoenus (L.) Soják – Round-headed Club-rush, Kugelbinse 287
Scirpus radicans Schkuhr – Creeping Club-rush, Wurzelnde Simse 287, 288 ff.
Scirpus sylvaticus L. – Wood Club-rush, Wald-Simse 287, 288 ff.
Sclerochloa dura (L.) P. Beauv. – Tough Grass, Hartgras 322
Scolochloa festucacea (Willd.) Link – Swamp Grass, Schwingelschilf 322
Scrophulariaceae 327, 332 ff., 345 f., 349
Scutellaria L. – Scullcap, Helmkraut 278
Secale cereale L. – Rye, Roggen 296 ff., 321, 351
Senecio L. – Ragwort, Greiskraut 279, 282 f.
Senecio erucifolius L. – Hoary Ragwort, Rautenblättriges Greiskraut
456 Appendix

Sesleria Scop. – Moor-grass, Blaugras 319

Sesleria albicans Kit. ex Schult. [= Sesleria caerulea auct. non (L.) Arduino; = Sesleria varia auct.] –
Blue Moor-grass, Kalk-Blaugras 296 ff., 319
Sesleria disticha (Wulf.) Pers. – Two-rowed Moor-grass, Zweizeiliges Blaugras 319
Sesleria ovata (Hoppe) Kern.– Moraine Moor-grass, Eiköpfiges Blaugras 319
Setaria P. Beauv. – Bristle-grass, Borstenhirse 294, 296 ff., 321
Silene L. – Campion, Leimkraut 349
Silene flos-cuculi (L.) Clairv. [= Lychnis flos-cuculi L.] – Ragged Robin, Kuckucks-Lichtnelke 328
Solanaceae 327, 337
Solanum melangena L. – Eggplant, Aubergine
Solanum nigrum L. – Black Nightshade, Schwarzer Nachtschatten 328
Solanum tuberosum L. – Potato, Kartoffel 326, 327, 351
Solidago L. – Goldenrod, Goldrute 279, 282 f., 347
Sorbus L. – Whitebeam, Rowan, Mehlbeere, Eberesche 285 f.
Sorbus aria L. (Crantz) s.l. – Common Whitebeam (group), Artengruppe Gemeine Mehlbeere
265 f., 266 ff.
Sorbus aucuparia L. – Rowan, Eberesche 265 f., 266 ff.
Sorbus chamaemespilus (L.) Crantz – Dwarf Whitebeam, Zwerg-Mehlbeere 265
Sorbus domestica L. – True Service-tree, Speierling 265
Sorbus intermedia (Ehrh.) Pers. – Swedish Whitebeam, Schwedische Mehlbeere 265, 266 ff.
Sorbus torminalis (L.) Crantz – Wild Service-tree, Elsbeere 265, 266 ff.
Sorghum Moench – Millet (partim), Mohrenhirse
Spartina Schreb. – Cord-grass, Schlickgras 322
Spiraea L. – Bridewort, Spierstrauch 266 ff., 270
Stachys palustris L. – Marsh Woundwort, Sumpf-Ziest 280 f.
Stachys sylvatica L. – Hedge Woundwort, Wald-Ziest 278, 280 f.
Stipa L. – Feather-grass, Federgras 296 ff., 321
Stipa capillata L. – Hairy Feathergrass, Haar-Pfriemengras 319, 321
Succisa pratensis Moench – Devil’s-bit Scabious, Gewöhnlicher Teufelsabbiss 328
Syringa vulgaris L. – Lilac, Gewöhnlicher Flieder 324
Tamaricaceae 336
Tamarix L. – Tamarisk, Tamariske 239, 322, 323
Taraxacum F.H. Wigg. – Dandelion, Löwenzahn 279, 282 f.
Taxaceae 244, 336 f.
Taxus baccata L. – Yew, Europäische Eibe 244, 245
Tetraclinis articulata (Vahl) Mast. – Arartree, Gliederzypresse
Teucrium montanum L. – Mountain Germander, Berg-Gamander 280 f.
Teucrium scorodonia L. – Wood Sage, Salbei-Gamander 278, 280 f.
Thalictrum flavum L. – Common Meadow Rue, Gelbe Wiesenraute 247
Thalictrum minus L. – Lesser Meadow Rue, Kleine Wiesenraute 246, 247
Thuja L. – Arborvitae, Lebensbaum
Thymus L. – Thyme, Thymian 278
Thymus praecox Opiz – Wild Thyme, Frühblühender Thymian 280 f.
Thymus pulegioides L. – Large Thyme, Feld-Thymian 280 f.
Thymus serpyllum L. – Breckland Thyme, Sand-Thymian 278, 280 f.
 Index and vernacular names of plants 457

Thymus vulgaris L. – Garden Thyme, Echter Thymian 280 f.

Tiliaceae 257 f., 332 ff., 342, 349
Tilia L. – Lime, Linde 257, 323
Tilia cordata Mill. – Small-leaved Lime, Winter-Linde 257
Tilia petiolaris DC. – Weeping Silver Lime, Hänge-Silber-Linde 257
Tilia platyphyllos Scop. – Large-leaved Lime, Sommer-Linde 257
Tragus racemosus (L.) All. – Burdock-grass, Traubiges Klettengras 322
Trapaceae 324
Trapa natans L. – Water-nut, Wassernuss 324
Trichophorum alpinum (L.) Pers. – Alpine Deergrass, Alpen-Rasenbinse 286
Trichophorum cespitosum (L.) Hartm. – Deergrass, Gewöhnliche Rasenbinse 286, 288 ff.
“Tricolpates” 329
Trifolium campestre Schreb. – Larger Hop Clover, Feld-Klee 273, 274
Trifolium dubium Sibth. – Suckling Clover, Kleiner Klee 273, 274
Trifolium repens L. – White Clover, Weiß-Klee 273, 274
Trisetum flavescens (L.) P. Beauv. – Yellow Oat-grass, Wiesen-Goldhafer 296 ff., 321
Triticum L. – Wheat, Weizen 296 ff., 321
Typhaceae 329
Ulmaceae 246 ff., 332 ff., 342, 349
Ulmus L. – Elm, Ulme 246 ff.
Ulmus glabra Huds. [= Ulmus scabra Mill.; = Ulmus montana With.] – Wych Elm, Berg-Ulme 247 f., 248
Ulmus laevis Pall. [= Ulmus effusa Willd.] – European White Elm, Flatter-Ulme 237 247, 248
Ulmus minor Mill. [= Ulmus glabra Mill.; = Ulmus carpinifolia Gled.; = Ulmus campestris auct. non
L.] – Smooth-leaved Elm, Feld-Ulme 247 f., 248
Ulmus x hollandica Mill. [= U. minor Mill. x U. glabra Huds.] – Dutch Elm, Bastard-Ulme 247 f., 248
Ulmus pumila L. – Siberian Elm, Sibirische Ulme
Urticaceae 248 ff., 332 ff., 342, 345
Urtica dioica L. – Stinging Nettle, Große Brennessel 248 ff., 325 f., 333
Urtica kiovensis Rogow. – Kievian Nettle, Röhricht-Brennessel 248
Urtica urens L. – Small Nettle, Kleine Brennessel 248 f.
Vaccinium myrtillus L. – Bilberry, Heidelbeere 263, 264
Vaccinium oxycoccos L. – Cranberry, Gewöhnliche Moosbeere 263, 264
Vaccinium vitis-idaea L. – Foxberry, Preiselbeere 263, 264
Vaccinium uliginosum L. – Bog Bilberry, Rauschbeere 263, 264
Valeriana dioica L. – Marsh Valerian, Kleiner Baldrian 328
Valerianella locusta (L.) Laterr. – Corn Salad, Echter Feldsalat 328, 351
Ventenata dubia (Leers) Coss. – Soft-bearded Oat-grass, Zweifelhafter Grannenhafer 322
Verbascum lychnitis L. – White Mullein, Mehlige Königskerze 325 f., 327
Verbascum thapsus L. – Great Mullein, Kleinblütige Königskerze 328
Veronica L. – Speedwell, Ehrenpreis 328
Veronica chamaedrys L. – Germander Speedwell, Gamander-Ehrenpreis
Veronica montana L. – Wood Speedwell, Berg-Ehrenpreis
Viburnum L. – Arrow-wood, Schneeball 323
Viburnum lantana L. – Wayfaring-tree, Wolliger Schneeball
Vicia tenuifolia Roth – Fine-leaved Vetch, Feinblättrige Vogel-Wicke 273, 274
458 Appendix

Vicia faba L. – Horse Bean, Saubohne

Vincetoxicum hirundinaria Medik. – Angle Pod, Schwalbenwurz 326
Violaceae 333
Viola L. – Violet, Veilchen 328
Viscum L. – Mistle, Mistel 346
Vitis vinifera L. – Grape Vine, Weinrebe 323, 350 f.
Vulpia C. C. Gmel. – Vulpia Fescue, Federschwingel 322, 342
Xanthium albinum (Widder) H. Scholz – Cocklebur, Ufer-Spitzklette
Zea mays L. – Maize, Mais 296 ff., 320, 321
 Acknowledgements 459

Acknowledgements

This work would not have been possible without the help of a great many friends and
colleagues. I am particularly indebted to my academic teacher Prof. Dr. Matthias Schaefer
(Göttingen), who offered a laboratory, logistic and financial support, as well as manifold
discussion. Prof. Dr. Reinhard Remane (Marburg) and Dr. Pavel Lauterer (Brno) were
open for my visits and confirmed many difficult species. They were also open for endless
discussions on taxonomic problems and insect – plant relations. Dr. Michael Sayer, Dr.
Michael Judas, and Dr. Jürgen Schauermann (all Göttingen) provided substantial help in
logistics and software problems. Dr. Alan Stewart (Brighton), Dr. Stephen Wilson (War-
rensburg) and Dr. Sonja Migge (Göttingen) did the great job of proof-reading.
I am also particularly grateful to Heike Albrecht, Thomas Fechtler (both Göttingen)
and Monika Körner (Bayreuth), for good company in the field and help in collecting,
and to Dr. Irena Dworakowska (previously Warsaw), Prof. Dr. Alexander Emelyanov
(St. Petersburg), Dr. William della Giustina (Versailles), Heidi Günthart (Dielsdorf), Dr.
Werner Holzinger (Graz), Dr. Dimitri Tishechkin (Moscow), Dr. Sabine Walter (Freital)
and Dr. Werner Witsack (Halle/Saale), who were always open for endless discussions
on numerous aspects of taxonomy and biology. Prof. Dr. Ekkehard Wachmann (Berlin)
generously provided the cover photographs.
Axel Rothländer (Göttingen) helped in problems concerning hard- and software, and
compiled the map. Gerd Apostel, Claus and Elwira Döring (all Göttingen), Christel
Fischer (Bovenden), Renate Grüneberg, Ingrid Kleinhans (both Göttingen), Dieter
Nünchert (Friedland) and Toby Spribille (Göttingen) provided technical or logistic
support.
Material, data and other valuable information were provided by Dr. Roland Acht-
ziger (Freiberg), Christian Andres (Gamburg), Prof. Dr. Alessandra Arzone (Torino), Dr.
John S. Badmin (Canterbury), Wolfgang Billen (Lörrach), Meike Deutschmann
(Pforzheim), Dr. Sakis Drosopoulos (Athens), Peter Dynort (Öhringen), Dirk Felzmann
(Lüneburg), Dr. Frank Fritzlar (Jena), Dr. Wolfgang Fröhlich (Marburg), Thomas Funke
(Halle/Saale), Herbert Gruber (Bad Lippspringe), Kai Heller (Kiel), Rainer Hess
(Würzburg), Christian Kehlmaier (Bremen), Dr. Christian Köppel (Karlsruhe), Norbert
Maczey (Ascot), Dr. Michael Maixner (Bernkastel-Kues), Igor Malenovsky (Brno), Dr.
Peter J. Mazzoglio (Torino), Dr. Albert Melber (Hanover), Dr. Rolf Niedringhaus (Old-
enburg), Stuart McKamey (Washington, D.C.), Ute Oesterling (Mammendorf), Ing. An-
drás Orosz (Budapest), Katharina Raupach (Hanover), Gerd Reder (Flörsheim-
Dalsheim), Dr. Ralf Rombach (Rech), Judith Rothenbücher (Göttingen), Prof. Dr. Wolf-
gang Schedl (Innsbruck), Klaus Schrameyer (Heilbronn), Ute Schröder (Kiel), Gabrijel
Seljak (Brje), Dr. Peter Sprick (Hanover), Prof. Dr. Hildegard Strübing (Berlin), Prof. Dr.
Vinton Thompson (Chicago), Klaus Voigt (Ettlingen), and Alexander Weis (Munich).
Dr. Wilko Ahlrichs (Oldenburg), Prof. Dr. G.A. Anufriev (Gorky), Dr. Manfred Asche
(Berlin), Dr. Robert Biedermann (Oldenburg), Prof. Dr. Vera D’Urso (Catania), Prof. Dr.
Matija Gogala (Ljubljana), Dr. Adalgisa Guglielmino (Viterbo), Prof. Dr. Hannelore Hoch
(Berlin), Dr. Klaus Hövemeyer (Adelebsen), Prof. Dr. Stefan Scheu (Darmstadt), Dr. Tomi
Trilar (Ljubljana) and Dr. Michael R. Wilson (Cardiff) contributed by stimulating dis-
cussion on various topics.
460 Appendix

Dr. Elisabeth Bauchhenss (Schweinfurt), † Eva Groh and † Günter Groh (Neustadt/
Weinstraße), Ingrid Kammerlander (Graz), Vera Lauterer (Brno), Dorit Lichter (Jena),
Christine Mohr (Forchheim), Susanne Wurst (Darmstadt), Anke Hanxleden (Kiel), Olaf
Schindler (Albersweiler) and Dr. Günther Scholl (Schweinfurt) offered hospitality dur-
ing field trips. Dr. Kathrin Baumann (Bad Harzburg), Dr. Christian Damm (Göttingen),
Dr. Jochen Gottwald (Göttingen), Dr. Gabriele Ritschel-Kandel (Würzburg) and Dr.
Michael Sauer (Reutlingen) gave valuable information on collecting localities. Klaus
Lewejohann and Dr. Volker Wissemann (both Göttingen) helped in host plant identifi-
cation. Rudolf May (Bonn) provided data on host plant distribution. Ronald Bellstedt
(Gotha), Dr. Kees den Bieman (Ulvenhout), Dr. Thierry Bourgoin (Paris), Dr. Jürgen
Deckert (Berlin), Dr. Rainer Emmrich (Dresden), Dr. Fitz Geller-Grimm (Wiesbaden),
Matthias Hartmann (Erfurt), † Frank Hattwig (Stuttgart), † Friedrich Heller (Stuttgart),
Dr. Larry Huldén (Helsinki), Dr. Karl-Heinz Lampe (Bonn), Dr. Rauno E. Linnavuori
(Raisio), Prof. Dr. Hans-Joachim Müller (Hamburg), Dr. Wolfgang Schawaller (Stuttgart),
Prof. Dr. Klaus Schönitzer (Munich), Prof. Dr. Hans Strümpel (Hamburg) and Dr. Her-
bert Zettel (Vienna) generously offered access to museum or private collections or pro-
vided help in revision. Wolfgang Frommer (Cologne), Dr. Wilhelm Kolbe (Burscheid)
and Holger Thüs (Frankfurt) provided unpublished manuscripts. Elena Nickel (Göttin-
gen) and Olga Seeckts (Adelebsen), facilitated exchange of material and literature with
Russian colleagues and translated manuscripts. Dr. Friedrich Sander (Jena), spent much
energy in revising material and compiling data from Thuringia.
Thomas Dunz (Goslar), Achim Gagalik (Erfurt), Dr. Jörn Hildebrandt (Bremen), † Dr.
Helmut Kriegbaum (Erlangen), Johannes Mohr (Forchheim) and Johannes Voith (Augs-
burg) stimulated and promoted research and interest in Auchenorrhyncha.
I am also much indebted to my friend Anja Ritzel (Göttingen) as well as to my neigh-
bours Johannes von Grafenstein, Cornelia Lohrberg, Claudia Schmalenbach and Anna
Mira Fellechner (all Waake) for their patience, logistic help, delicious food and excellent
wine.
Last but not least, my thanks go to the staff of Pensoft Publishers (Sofia), and partic-
ularly Dr. Lyubomir Penev and Teodor Georgiev, who did much effort to get this book
into shape.