Professional Documents
Culture Documents
Herbert Nickel
Errastunus ocellaris (Fall.), Conomelus anceps (Germ.),
by courtesy of Ekkehard Wachmann. Background by courtesy of Thomas Fechtler.
and Planthoppers
of Germany
Co-published by
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
Herbert Nickel
Co-published
by
Pensoft Publishers, Sofia-Moscow
Goecke & Evers, Keltern
iv Contents
© Herbert Nickel
Co-published
by
Pensoft Publishers, Sofia-Moscow (ISBN 954-642-169-3)
Goecke & Evers, Keltern (ISBN 3-931374-09-2)
© PENSOFT Publishers
Dedicated to
Günter and Eva Groh
(† Feb 2nd 1996, and Feb 5th 1996)
vi Contents
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
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
2 Study area
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).
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
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.
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
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
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).
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
(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).
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.
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.
area size (Basset et al. 1997, see this paper also for further methods and discussion of
canopy sampling).
(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).
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
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:
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
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)
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)
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)
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)
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)
(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)
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
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)
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
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
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)
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)
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)
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
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)
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
Fischer (1972), Fröhlich (1996a), Nikusch (1976), Schiemenz (1987), Walter (1998), Wagner (1939a), HN; see
also della Giustina & Remane (1999)
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)
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)
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)
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
SMNS (leg. Dynort), Nickel (1999b); see also Anufriev & Kirillova (1998), Linnavuori (1951) (see this paper
also for identification of nymphs), Ossiannilsson (1978)
(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.
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)
(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)
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)
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)
(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)
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)
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)
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)
Frommer (1996), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998a),
Wagner (1935, 1939a), HN
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)
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)
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)
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)
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.
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
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
Fischer (1972), Reimer (1992), Schönitzer & Oesterling (1998b), Wagner (1950, 1951a), HN
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)
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)
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
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)
Fischer (1972), Frommer (pers. comm.), Remane & Fröhlich (1994b), Remane & Wachmann (1993), Wagner
(1939a, 1951a), Weis (pers. comm.), Nickel (1994), HN
Frommer (1996), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a,
1951a), HN
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)
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
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)
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
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)
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
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)
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)
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)
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)
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)
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)
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)
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
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)
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
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
Frommer (pers. comm.), Lehmann (1973a, 1973b), Müller (1978), Nicolaus (1957), Reimer (1992), SMNS,
Nickel & Remane (1996), Nickel (1994), HN
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)
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)
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)
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)
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)
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.)
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
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)
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)
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)
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)
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)
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)
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)
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
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)
Günthart (1988, and pers. comm.), Heller (1996), Remane (pers. comm.), HN; see also Günthart (1987b),
Ribaut (1936), Vidano & Arzone (1987b)
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)
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
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)
Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (pers. comm.), Schiemenz et al. (1996), Schönitzer
& Oesterling (1998b), Wagner (1939, 1951), Nickel (1994), HN
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)
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
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)
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)
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
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)
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
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)
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)
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
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)
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)
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
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)
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)
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)
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)
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)
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)
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
Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer (1992), Remane (1961a.),
Schiemenz et al. (1996), HN
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)
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
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).
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)
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)
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)
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
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)
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)
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),
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)
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.
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)
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)
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)
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
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
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
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.)
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)
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).
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.
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.
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
200 0000000000000000000000000
0 00000000000000000000000
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158 00000000000000000000
000000000000000000000000000000
Species number
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00 00000000000000000 0000000000000000000 00 00000000000000000000
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150 0 00000000000000000
00 0000000000000000000 00
00000000000000000000
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0000000000000000000 00 00000000000000000000000000000000000
00 00000000000000000
000000000000000000 0 00000000000000000000
0000000000000000000000000
00 00000000000000000
00000000000000000 00 00000000000000000000 00000
00000000000000000000 000
00 00000000000000000
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109 0 000000000000000000 0
00 00000000000000000 00
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00000000000000000000000
0000000000000000000000000000 00000000000000000
00000000000000000000 000 00000000000000000000
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000000000000000000 00 00000000000000000 00000000000000000000
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100 000000000000000000
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00 00000000000000000
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00000000000000000000 000
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000000000000000000 00 00000000000000000 00000000000000000000
00000000000000000000000 0 000000000000000000 0 0000000000000000000000000
000000000000000000
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00 00000000000000000
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00 000000000000000000 0000000000 0000000000000000000000000000
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0000000000000000 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00000 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00000000000000000000000 0000000000000000000000000 00000000000000000000000
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br ae
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Fig. 2: Species numbers and systematic affiliations of the Auchenorrhyncha fauna of Germany
General remarks and problems 237
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
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
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
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
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
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
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):
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.
Table 10. Auchenorrhyncha species utilizing Betulaceae in Germany. Explanations see Table 5.
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
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.
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
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
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.
Py racant ha coccinea
Amelanchier spp.
Sorbus aucuparia
Malus domest ica
Py rus communis
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
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
Py racant ha coccinea
Amelanchier spp.
Sorbus aucuparia
Malus domest ica
Py rus communis
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
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
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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.
Hippocrepis comosa
Colut ea arborescens
Lot us corniculat us
Cy t isus scoparius
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.
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.
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.
Glechoma hederacea
Marrubium v ulgare
O cimum basilicum
Leonurus cardiaca
Prunella v ulgaris
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
Clinopodium v ulgare
Ly copus europaeus
Melissa officinalis
O riganum v ulgare
Th. pulegioides
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.
Dahlia pinnat a
Ast er linosy ris
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
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.
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.
Schoenus ferrugineus
Eleocharis palust ris
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
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
Schoenus ferrugineus
Eleocharis palust ris
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
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
Schoenus ferrugineus
Eleocharis palust ris
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
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)
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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298 Utilization of plant resources
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
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X X x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. x x X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . x X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . x X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . ? X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . X x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . X x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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. . . . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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. . . . . . . . X X . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . X . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . X . . . . x . . . . . . . . . . . . . . . . . .
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. . . . . . . . . . . . ? X . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . x ? . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . ? . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . X . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . X . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . X . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . ? X . ? . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . ? . . ? . . . . . . . . . . . . . .
. . . . . . . ? . . . . . . . . . . . . X . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . X x . . . . . . . . . . . . .
300 Utilization of plant resources
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
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 . . . .
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. 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
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
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
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
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
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
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:
Table 23. Auchenorrhyncha species utilizing further woody host plants in Germany. Explanations see Table 5.
Hippophae rhamnoides
Elaeagnus angust ifolia
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
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
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
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
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
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
Substrate
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
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
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
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.
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
(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
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
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.
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.
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
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
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
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.
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.
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
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
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.
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
(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
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
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Fig. 12: Species numbers of 1st and 2nd degree monophagous Auchenorrhyncha on the plant families of
Germany
50
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0
Be s
Ph u s
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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
40
33
35
28 27
30
S Auchenorrhyncha
25
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0
ur
a
a
a
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is
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in
in
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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.
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
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.
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
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
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).
6.2 Dispersal
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
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)
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.
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.
6.3.1 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)
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)
Fig. 15: Dormancy phenomena in arthropods, after Müller (1970, 1992) and Witsack (1981, 1985, 2002)
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
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
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.
8 References
Abdul-Nour, H., Lahoud, L. (1996): Revision du genre Philaenus Stal, 1864 au Libanon, avec la
description d’une nouvelle espèce: P. arslani, n.sp. (Homoptera Auchenorrhyncha, Cercopi-
dae). – Nouv. Rev. d’Ent. (N.S.) 12: 297-303.
Achtziger, R. (1991): Zur Wanzen- und Zikadenfauna von Saumbiotopen. – Ber. ANL 15: 37-68.
Achtziger, R. (1995): Die Struktur von Insektengemeinschaften an Gehölzen: Die Hemipteren-
Fauna als Beispiel für die Biodiversität von Hecken- und Waldrandökosystemen. – Bayreu-
ther Forum Ökologie 20, 216 pp.
Achtziger, R., Nickel, H., Schreiber, R. (1999): Auswirkungen von Extensivierungsmaßnahmen
auf Zikaden, Wanzen, Heuschrecken und Tagfalter im Feuchtgrünland. – Schr.-R. Bayer. Landes-
amt f. Umwelt 150: 109-131.
Achtziger, R., Nickel, H. (1997): Zikaden als Bioindikatoren für naturschutzfachliche Erfolgs-
kontrollen im Feuchtgrünland. – Beitr. Zikadenkunde. 1: 3-16.
Adis, J., Basset, Y., Floren, A., Hammond, P.M., Linsenmair, K.E. (1998): Canopy fogging of an
overstorey tree – recommendations for standardization. – Ecotropica 4: 93-97.
Afscharpour, F. (1960): Ökologische Untersuchungen über Wanzen und Zikaden auf Kulturfel-
dern in Schleswig-Holstein. – Z. angew. Zool. 47: 257-301.
Al Hussein, I.A., Bergmann, S., Funke, T., Huth, J., Oelerich, H.-M., Reuter, M., Tietze, F., Wit-
sack, W. (1999): Die Tierwelt der Bergbaufolgelandschaften. – Naturschutz im Land Sachsen-
Anhalt 36: 23-40.
Alla, S., Moreau, J.P., Frerot, B. (2001a): Effects of the aphid Rhopalosiphum padi on the leafhopper
Psammotettix alienus under laboratory conditions. – Entomologia exp. appl. 98(2): 203-209.
Alla, S., Moreau, J.P., Frerot, B. (2001b): Evidence of an extractable semiochemical produced by
Rhopalosiphum padi acting on Psammotettix alienus. – Entomologia exp. appl. 99(1): 107-111.
Ammar, E.-D. (1985): Internal morphology and ultrastucture of leafhoppers and planthoppers.
– In: Nault, L.R., Rodriguez, J.G. (eds.): The leafhoppers and planthoppers. Wiley, New York,
pp. 127-162.
Andersen, N.M. (1997): Phylogenetic tests of evolutionary scenarios: The evolution of flightless-
ness and wing polymorphism in insects. – Mem. Mus. natn. Hist. nat. 173: 91-108.
Andrzejewska, L. (1962): Macrosteles laevis Rib. as an unsettlement index of natural meadow
associations of Homoptera. – Bull. Acad. pol. Sci. Cl. II. Ser. Sci. biol. 10: 221-226.
Andrzejewska, L. (1965): Stratification and its dynamics in meadow communities of Auchenor-
rhyncha (Homoptera). – Ekologia Polska (Ser. A) 13: 685-715.
386 References
Backus, E. (1985): Anatomical and sensory mechanisms of planthopper and leafhopper feeding
behavior. – In: Nault, L.R., Rodriguez, J.G. (eds.): The leafhoppers and planthoppers. Wiley,
New York, pp. 163-194.
Backus. E. (1988): Observations on the feeding behaviour of Empoasca fabae (Harris) (Cicadel-
lidae: Typhlocybinae) and the cause of hopperburn. – Proc. 6th Auchenorrhyncha Meeting,
Turin, Italy, September 7-11, 1987, pp. 493-500.
Badmin, J. (1991): Additional host plants of Eupteryx filicum (Newman) (Hem., Cicadellidae). –
Entomologist’s. mon. Mag. 127: 196.
Badmin, J. (1992): Notes on two Homoptera new to the Isles of Scilly. – Entomologist’s. mon.
Mag. 128: 62.
Badmin, J. (1997): The biology of the grassland froghopper Neophilaenus campestris at Dunge-
ness. – Dungeness Bird Observatory Report for 1996: 84-87.
Bartels, R., Fritzlar, F., Schneider, K., Schöpke, H., Wallaschek, M. (1992): Ergebnisse einer Insek-
tenaufsammlung (Saltatoria; Heteroptera; Homoptera: Auchenorrhyncha; Coleoptera) in Bit-
terfeld und Umgebung (Sachsen-Anhalt). – Kaleidoskop, Z. Päd. Hochsch. Halle 1: 28-33.
Basset, Y., Springate, N.D., Aberlenc, H.P., Delaware, G. (1997): A review of methods for sam-
pling arthropods in tree canopies. – In: Stork, N.E., Adis, J., Didham, R.K. (eds.): Canopy
Arthropods. Chapman & Hall, London. pp. 27-52.
Baugnée, J.-Y. (1995): Homoptères Auchénorhynches nouveaux pour la faune belge, récoltés
dans le sud de l’Entre-Sambre-et Meuse (Homoptera: Delphacidae, Cicadellidae). – Bull. Annls.
Soc. R. Belge Ent. 131: 161-167.
Begon, M., Harper, J.L., Townsend, C.R. (1996): Ecology. Individuals, populations and communi-
ties. 3rd ed. – Blackwell, Oxford. 1068 pp.
Bengtsson, J., Nilsson, S.G., Franc, A., Menozzi, P. (2000): Biodiversity, disturbances, ecosystem func-
tion and management of European forests. – Forest Ecology and Management 132(1): 39-50.
Benkert, D., Fukarek, F., Korsch, H. (eds.) (1996): Verbreitungsatlas der Farn- und Blütenpflan-
zen Ostdeutschlands. – Fischer, Jena. 615 pp.
Berenbaum, M.R. (1990): Evolution of specialization in insect-umbellifer associations. – Annu.
Rev. Entomol. 35: 319-343.
Bernays, E. (1996): Selective attention and host-plant specialization. – Ent. exp. appl. 80: 125-131.
Bernays, E. (1998): The value of being a resource specialist: Behavioral support for a neural
hypothesis. – Am. Nat. 15: 451-464.
Bernays, E. (1999a): Plasticity and the problem of choice in food selection. – Ann. Entomol. Soc.
Am. 92(6): 944-951.
Bernays, E. (1999b): When host choice is a problem for a generalist herbivore: Experiments with
the whitefly, Bemisia tabaci. – Ecol. Entomol. 24(3): 260-267.
Bernays, E. Funk , D. (1999): Specialists make faster decisions than generalists: Experiments
with aphids. – Proc. R. Soc. London, Biol. Sci., Ser. B 266: 151-156.
Bernays, E., Graham, M. (1988): On the evolution of host specificity in phytophagous arthro-
pods. – Ecology 69: 886-892.
Bernhardt, K.-G. (1991): Zum Auftreten von Cicadetta montana Scopoli, 1772 (Homoptera – Auchen-
orrhyncha) bei Tecklenburg und Lengerich. – Natur u. Heimat 51: 77-78.
Bernhardt, K.-G., Arnold, K. (1991): Zum Auftreten von Haematoloma dorsatum (AHRENS) und
Graphocephala fennahi YOUNG in den Räumen Münster und Osnabrück. – Natur u. Heimat
51: 75-77.
388 References
Carl, M., Schönitzer, K. (1995): Beitrag zur Kenntnis der bayerischen Zikadenfauna (Homoptera,
Auchenorrhyncha). – Nachr.-Bl. bayer. Ent. 44: 13-18.
Cerutti, N. (1937): Captures intéressantes d’Hémiptères du Valais. – Mitt. Schweiz. ent. Ges. 17:
168-172.
Cerutti, N. (1938): Trois nouvelles espèces de Cicadines du Valais. – Mitt. Schweiz. ent. Ges. 17:
189-194.
Cerutti, N. (1939a): Les Typhlocybidae du Valais. – Bull. Murithienne 56: 81-95.
Cerutti, N. (1939b): Captures intéressantes d’Hémiptères du Valais (3e liste) et description
d’espèces nouvelles. – Mitt. Schweiz. Ent. Ges. 17: 443-449.
China, W.E. (1942): A revision of the British species of Cixius Latr. (Homoptera), including the
description of a new species from Scotland. – Trans. Soc. Brit. Ent. 8: 79-110.
China, W.E. (1943): New and little-known species of British Typhlocybidae (Homoptera) with
keys to the genera Typhlocyba, Erythroneura, Dikraneura, Notus, Empoasca and Alebra. – Trans.
Soc. Brit. Ent. 8: 111-153.
Christandl-Peskoller, H., Janetschek, H. (1976): Zur Faunistik und Zoozönotik der südlichen
Zillertaler Hochalpen. – Veröff. Univ. Innsbruck 7 (101): 1-134.
Chudzicka, E. (1982): Auchenorrhyncha (Homoptera) of Warsaw and Mazovia. – Memorabilia
Zool. 36: 143-164.
Chudzicka, E. (1986a): Structure of leafhopper (Homoptera, Auchenorrhyncha) communities in
the urban green of Warzaw. – Memorabilia Zool. 42: 67-99.
Chudzicka, E. (1986b): Effects of anthropogenic pressure on leafhopper (Homoptera, Auchenor-
rhyncha) guilds. – Memorabilia Zool. 42: 100-123.
Claridge, D.W. (1986): The distribution of a typhlocybine leafhopper, Ribautiana ulmi (Homoptera:
Cicadellidae) on a specimen wych elm tree. – Ecol. Ent. 11: 31-39.
Claridge, D.W., Derry, N.J., Whittaker, J.B. (1981): The distribution and feeding of some Typhlo-
cybinae in response to sun and shade. – Acta ent. Fenn. 38: 8-12.
Claridge M.F., Nixon, G.A. (1986): Oncopsis flavicollis (L.) associated with tree birches (Betula): a
complex of biological species or a host plant utilization polymorphism? – Biol. J. Linn. Soc.
27: 381-397.
Claridge, M.F., Reynolds, W.J. (1972): Hostplant specifity, oviposition behavior and egg parasit-
ism in some woodland leafhoppers of the genus Oncopsis (Hemiptera Homoptera: Cicadel-
lidae). – Trans. R. Ent. Soc. London 124: 149-166.
Claridge, M.F., Reynolds, W.J. (1973): Male courtship songs and sibling species in the Oncopsis
flavicollis species group (Hemiptera: Cicadellidae). – J. Ent. (B) 42 (1): 29-39.
Claridge, M.F., Reynolds,W.J., Wilson, M.R. (1977): Oviposition behavior and food plant dis-
crimination in leafhoppers of the genus Oncopsis. – Ecol. Ent. 2: 19-25.
Claridge, M.F., Wilson, M.R. (1976): Diversity and distribution pattern of some mesophyll-feed-
ing leafhoppers of temperate woodland canpopy. – Ecol. Ent. 1: 231-250.
Claridge, M.F., Wilson, M.R. (1978a): Seasonal changes and alternation of food plant preference
in some mesophyll-feeding leafhoppers. – Oecologia 37: 247-255.
Claridge, M.F., Wilson, M.R. (1978b): Observations on new and little known species of Typhlo-
cybine leafhoppers (Hemiptera: Cicadellidae) in Britain. – Ent. Gaz. 29: 247-251.
Claridge, M.F., Wilson, M.R. (1981): Host-plant associations, diversity and species-area relation-
ships of mesophyll-feeding leafhoppers of trees and shrubs in Britain. – Ecol. Ent. 6: 217 -238.
Clay, K. (1990): Fungal endophytes of grasses. – Annu. Rev. Ecol. Syst. 21: 275-297.
References 391
Clay, K. (1996): Interactions among fungal endophytes, grasses and herbivores. – Res. Popul.
Ecol. 38: 191-201.
Clay, K. (1997): Fungal endophytes, herbivores and the structure of grassland communities. –
In: Gange, A.C., Brown, V.K. (eds.): Multitrophic interactions in terrestrial systems. Black-
well, Oxford. pp. 151-169.
Cobben, R.H. (1956): Bionomie der Jasside Fieberiella florii STAL (Hom. Auchenorrhyncha). –
Publ. natuurhist. Genootsch. Limburg 9: 57-82.
Cobben, R.H. (1988): What do we really know about host selection in Auchenorrhyncha? – Proc.
6th Auchenorrhyncha Meeting, Turin, Italy, September 7-11, 1987, pp. 81-92.
Cobben, R.H., Gravestein, W.H. (1958): 95 Cicaden, nieuw voor de Nederlandse fauna (Hom.
Auchenorrhyncha). – Ent. Ber. Amsterdam 18: 122-124.
Cobben, R.H., Rozeboom, G.J. (1978): Notes on Auchenorrhyncha (Homoptera) from pitfall traps
in the Gerendal Reserve (southern part of Limburg Province). – Publ. natuurhist. Genootsch.
Limburg 28: 3-15.
Cobben, R.H., Rozeboom, G.J. (1983): De invertebratenfauna van de Zuidlimburgse kalk-
graslanden. De Cicaden in bodemvallen (Hemiptera, Homoptera Auchenorrhyncha). – Na-
tuurhist. Maandbl. 72(6/7): 102-110.
Conert, H.J. (1998): Poaceae (Echte Gräser oder Süßgräser). 3. Aufl. – In: Conert, H.J., Jäger, E.J.,
Kadereit, J.W., Schutze-Motel, W., Wagenitz, G., Weber, H.E. (eds.): Gustav Hegi. Illustrierte
Flora von Mitteleuropa. Band I, Teil 3. 898 pp.
Connell, J.H. (1980): Diversity and the coevolution of competitors, or the ghost of competition
past. – Oikos 35: 131-138.
Conti, M. (1988): Recent research on leafhopper vectors of plant viruses and mycoplasmas at the
plant virus institute – Turin. – Proc. 6th Auchenorrhyncha Meeting, Turin, Italy, 7-11 Sept.
1987, pp. 447-457.
Cook, A.A. (1996): The host plants of calcareous grassland Auchenorrhyncha (Hemiptera). –
Entomol. mon. Mag. 132: 151-175.
Cook, A.G., Denno, R.F. (1994): Planthopper/plant interactions: feeding behavior, plant nutri-
tion, plant defense, and host plant specialization. – In: Denno, R.F., Perfect, T.J. (eds.): Plant-
hoppers: their ecology and management. Chapman & Hall, New York. pp. 114-139.
Courcy Williams, M. de (1989): A preliminary list of the Irish Auchenorrhyncha (Homoptera). –
Bull. Ir. Biogeog. Soc. 12: 135-148.
Crawley, M.J. (1983): Herbivory – the dynamics of animal-plant interactions. – Blackwell, Ox-
ford. 437 pp.
Crawley, M.J. (1997): Plant-herbivore dynamics. – In: Crawley, M.J. (ed.): Plant ecology. 2nd ed.
Blackwell, Oxford, pp. 401-474.
Cronquist, A. (1968): The evolution and classification of flowering plants. – Nelson, London. 396 pp.
Curry, J.P. (1994): Grassland invertebrates. – Chapman & Hall, London. 437 pp.
Darwin, C. (1859): On the origin of species. (Facsimile of the first edition). – Harvard University
Press, Cambridge/ Massachusetts. 515 pp.
Davidson, R., DeLong, D. (1938): Studies of the genus Empoasca (Homoptera, Cicadellidae). Part
V. Twelve new species of Empoasca from the United States. – Ohio J. Sci. 38: 90-96.
Davis, B.N.K. (1973): The Hemiptera and Coleoptera of Stinging Nettle (Urtica dioica L.) in East
Anglia. – J. appl. Ecol. 10: 213-237.
Davis, B.N.K. (1983): Insects on nettles. – Cambridge University Press, Cambridge. 64 pp.
392 References
Davis, B.N.K., Lawrence, C.E. (1973): Insects collected from Parietaria diffusa MERT. & KOCH
and Urtica urens L. in Huntingdonshire. – Entomologist‘s monthly Mag. 109: 252-254.
Demichelis, S., Bosco, D. (1995): Host-plant relationships and life history of some Alebra species
in Italy (Auchenorrhyncha: Cicadellidae). – Eur. J. Entomol. 92: 683-690.
Denno, R.F. (1976): Ecological significance of wing polymorphism in Fulgoroidea which inhabit
tidal salt marshes. – Ecol. Ent. 1: 257-266.
Denno, R.F., McClure, M.S., Ott, J.R. (1995): Interspecific interactions in phytophagous insects:
competition reexamined and resurrected. – Ann. Rev. Ent. 40: 297-331.
Denno, R.F., Perfect, J.T. (1994): Planthoppers: Their ecology and management. – Chapman &
Hall, New York. 799 pp.
Denno, R.F., Raupp, M.J., Tallamy, D.W., Reichelderfer, C.F. (1990): Migration in heterogenous
environments: differences in habitat selection between the wing forms of the dimorphic plan-
thopper, Prokelisia marginata (Homptera: Delphacidae). – Ecology 61(4): 859-867.
Denno, R.F., Roderick, G.F. (1990): Population biology of planthoppers. – Annu. Rev. Entomol.
35: 489-520.
Denno, R.F., Roderick, G.F. (1992): Density-related dispersal in planthoppers: Effects of inter-
specific crowding. – Ecology 73: 1323-1334.
Denno, R.F., Roderick, G.F., Olmstead, K.L., Dobel, H.G. (1991) Density-related migration in plant-
hoppers (Homoptera: Delphacidae): the role of habitat persistence. – Am. Nat. 138: 1513-1541.
Dietrich, C.H., Rakitov, R.A., Holmes, J.L., Black, W.C., IV (2001): Phylogeny of the major lineag-
es of Membracoidea (Insecta: Hemiptera: Cicadomorpha) based on 28S rRNA sequences. –
Molecular Phylogenetics and Evolution 18(2): 293-305.
Dietrich, C.H., Rothschild, M.J., Deitz, L.L. (1999): Checklist and host plants of the treehoppers
(Hemiptera: Membracidae) of North Carolina. – Proc. Entomol. Soc. Wash. 101(2): 242-262.
Dixon, A.F.G. (1966): The effect of population density and nutritive status of the host on the
summer reproductive activity of the sycamore aphid, Drepanosiphum platanoides (Schr.). – J.
Anim. Ecol. 35: 105-112.
Dixon, A.F.G. (1976): Timing of egg hatch and viability of the sycamore aphid, Drepanosiphum
platanoides (Schr.), at bud burst of sycamore, Acer pseudoplatanus L. – J. Anim. Ecol. 45: 593-603.
Dixon, A.F.G. (1984): Plant architectural complexity and alary polymorphism in tree-dwelling
aphids. – Ecol. Entomol. 9: 117-118.
Dixon, A.F.G. (1998): Aphid ecology. 2nd ed. – Chapman & Hall, London. 300 pp.
Dixon, A.F.G., Kindlmann, P., Leps, J., Holman, J. (1987): Why are there so few species of aphids,
especially in the tropics? – Am. Nat. 129(4): 580-592.
Dlabola, J. (1952): Einige neue paläarktische Zikaden und andere faunistische Bemerkungen. –
Acta ent. Mus. Nat. Pragae 28: 27-37. (In Czech.).
Dlabola, J. (1954): Křísi – Homoptera. – Fauna CSR 1, Praha. 339 pp.
Dlabola, J. (1958): Records of leafhoppers from Czechoslovakia and South European countries
(Homoptera: Auchenorrhyncha). – Acta faun. ent. Mus. Nat. Pragae 3: 7-15.
Dlabola, J. (1961a): Die Zikaden von Zentralasien, Dagestan und Transkaukasien (Homopt.
Auchenorrhyncha). – Acta ent. Mus. Nat. Pragae 34 (587): 241-358.
Dlabola, J. (1961b): Eine neue Platymetopiusart aus Mitteleuropa und weitere tschecho-
slowakische Faunistik. – Acta faun. ent. Mus. Nat. Pragae 7 (52): 5-9.
Dlabola, J. (1965): Neue Zikadenarten aus Südeuropa (Homoptera, Auchenorrhyncha). – Acta
ent. Mus. nat. Pragae 36: 657-669.
References 393
Dworakowska, I. (1973): Baguoidea rufa (Mel.) and some other Empoascini (Auchenorrhyncha,
Cicadellidae). – Bull. Acad. Pol. Sci. (Cl. II) 21: 49-58.
Dworakowska, I. (1976): Kybos FIEB., subgenus of Empoasca WALSH (Auchenorrhyncha, Ci-
cadellidae, Typhlocybinae) in Palaearctic. – Acta Zool. Cracov. 21: 387-463.
Dworakowska, I. (1977a): On some species of the subgenus Kybos Fieb. with remarks on some
other Empoascini (Auchenorrhyncha, Cicadellidae, Typhlocybinae). – Bull. Acad. Pol. Sci.
(Cl. II) 25: 609-617.
Dworakowska, I. (1977b): On several of Dlabola’s Typhlocybinae species (Auchenorrhyncha:
Cicadellidae). – Acta zool. Acad. Sci. hung., Budapest 23: 29-36.
Dworakowska, I. (1982): Typhlocybini of Asia (Homoptera, Auchenorrhyncha, Cicadellidae). –
Ent. Abh. Staatl. Mus. Tierk. Dresden 45(6): 99-181.
Dworakowska, I. (1993): Remarks on Alebra FIEB. and Eastern Hemisphere Alebrini (Auchenor-
rhyncha: Cicadellidae: Typhlocybinae). – Entomotaxonomia 15(2): 91-121.
Eastop, V.F. (1973): Deductions from the present day host plants of aphids and related insects. –
In: Emden, H.F. van (ed.): Insect/plant relationships. Blackwell, Oxford. pp. 157-178.
Eastop, V.F. (1978): Diversity of the Sternorrhyncha within major climatic zones. – In: Mound,
L.A., Waloff, N. (eds.): Diversity of insect faunas. Blackwell, Oxford. pp. 71-88.
Edwards, J. (1896): The Hemiptera-Homoptera of the British Islands. – L. Reeve, London. 271 pp.
Ehrlich, P.R., Raven, P.H. (1964): Butterflies and plants: A study in coevolution. – Evolution 18:
586-608.
Eisner, T., Eisner, M., Hoebeke, E.R. (1998): When defense backfires: detrimental effect of a plant’s
protective trichomes on an insect beneficial to the plant. – Proc. Natl. Acad. Sci. USA 95(8):
4410-4414.
Eitschberger, U. (1972): Zur Verbreitung von Cicadetta montana und Tibicen haematodes in Main-
franken (Hom., Cicadidae). – Entomol. Zs. Stuttgart 82: 210-213.
Ellenberg, H. (1996): Vegetation Mitteleuropas mit den Alpen in ökologischer Sicht. 5. Aufl. –
Ulmer, Stuttgart. 1095 pp.
Ellenberg, H., Mayer, R., Schauermann, J. (eds.) (1986): Ökosystemforschung: Ergebnisse des
Sollingprojektes. – Ulmer, Stuttgart. 507 pp.
Emeljanov A.F. (1964a): Suborder Cicadinea (Auchenorrhyncha). – Keys to the Insects of the
European USSR. 1: 337-437. (In Russian).
Emelyanov A F. (1980): An attempt of subgeneric division of the genus Tettigometra Latr. – Vestn.
Zool. 2: 51-55.
Emelyanov, A.F. (1972): New leaf-hoppers from the Mongolian People‘s Republic (Homoptera,
Auchenorrhyncha). – Insects of Mongolia 1: 199-260. (In Russian).
Emelyanov, A.F. (1982): Fulgoroidea (Homoptera) collected in the Mongolian People‘s Republic
by the entomofaunistical group of the Soviet-Mongolian complex biological expedition in
1970-1975. – Insects of Mongolia 8: 69-122. (In Russian).
Emelyanov, A.F. (1987): The phylogeny of Cicadina (Homoptera), based on comparative mor-
phological data. – Proc. Allunion Ent. Sci. 69: 19-109. (In Russian).
Emelyanov, A.F. (1993): New records of buffalo treehopper, Stictocephala bisonia Kopp & Yonke,
from temperate Asia (Homoptera: Membracidae). – Zoosystematica Rossica 2(2): 246.
Emelyanov, A.F. (2000): New records of planthoppers and leafhoppers from Russia and Kazakh-
stan (Homoptera, Cicadina). – Zoosystematica Rossica 8: 256.
396 References
Emeljanov, A.F., Gnezdilov, V.M., Gjonov, I.V. (2002): Addenda to the fauna of Cicadina (Hemi-
ptera) of Bulgaria. – Acta entomol. Slovenica 10: 83-90.
Emmrich, R. (1966): Faunistisch-ökologische Untersuchungen über die Zikadenfauna (Ho-
moptera, Auchenorrhyncha) von Grünland-Flächen und landwirtschaftlichen Kulturen des
Greifswalder Gebietes. – Mitt. Zool. Mus. Berlin 42: 61-126.
Emmrich, R. (1969): Bodenfallenfänge von Zikaden aus nordostdeutschen Laub- und Kiefern-
Mischwäldern. – Faun. Abh. Staatl. Mus. Tierk. Dresden 2: 279-294.
Emmrich, R. (1973): Zur Zikadenfauna der Insel Hiddensee (Homoptera, Auchenorrhyncha). –
Faun. Abh. Staatl. Mus. Tierk. Dresden 4: 171-176.
Emmrich, R. (1975): Faunistische Daten von Zikaden aus dem Gebiet der DDR (Homoptera,
Auchenorrhyncha). – Faun. Abh. Staatl. Mus. Tierk. Dresden 5: 241-254.
Emmrich, R. (1980): Zur taxonomischen Gliederung sowie Verbreitung von Aphrodes bicinctus
(SCHRK.) sensu RIB. (Homoptera, Auchenorrhyncha, Cicadellidae). – Faun. Abh. Staatl. Mus.
Tierk. Dresden 7: 279-284.
Emmrich, R. (1983): Ein Fund von Cercopis sanguinolenta (Scop.) im oberen Elbtal bei Dresden
(Homoptera, Auchenorrhyncha). – Ent. Nachr. Ber. 27: 273-275.
Emmrich, R. (1984): Vorkommen und Verbreitung von Cicadetta montana (Scop.) im Gebiet der
DDR, unter besonderer Berücksichtigung der Sächsischen Schweiz (Insecta, Homoptera,
Auchenorrhyncha, Cicadidae). – Faun. Abh. Staatl. Mus. Tierk. Dresden 11: 109-117.
Emmrich, R. (1986): Zur Verbreitung von Elymana kozhevnikovi (Zachv.) auf dem Gebiet der DDR
(Insecta, Homoptera, Auchenorrhyncha, Cicadellidae). – Faun. Abh. Staatl. Mus. Tierk. Dres-
den 14: 99-101.
Emmrich, R. (1989): Kommt Cicadula longiventris (J. Sahlb.) in Mitteleuropa vor? – Verh. IX. SIEEC,
Gotha 1986: 378-383.
Emmrich, R. (2001): Zur Zikadenfauna der sächsischen Braunkohlenbergbau-Folgelandschaften.
– DGaaE-Nachrichten 15(4): 134.
Farrell, B.D. (1998): “Inordinate fondness” explained: Why are there so many beetles? – Science
281: 555-559.
Feeny, P. (1976): Plant apparency and chemical defense. – Rec. Adv. Phytochem. 10: 1-40.
Feldmann, R., Bussmann, M. (1993): Die Ausbreitung der Rhododendron-Zikade (Graphocephala
fennahi Young) im Sauerland und im Hellwegraum. – Natur u. Heimat 53: 93-98.
Feldtmann, E. (1937): Nachtrag zu W. Wagner: Zikaden der Nordmark. – Bombus 2: 6-7.
Feldtmann, E. (1938): 54. (Hom. Jass.). – Bombus 2: 6-7.
Fennah, R.G. (1950): A generic revision of the Achilidae (Homoptera: Fulgoroidea) with descrip-
tions of new species. – Bull. Brit. Mus. Nat. Hist. Entomol. 1(1): 1-170.
Fieber, F.X. (1865): Synopse der europ. Arten Tettigometra. – Verh. zool.-bot. Ges. Wien. 15: 561-572.
Fieber, F.X. (1866): Grundzüge zur generischen Theilung der Delphacini. – Verh. zool.-bot. Ges.
Wien 16: 517-534.
Fieber, F.X. (1869): Synopse der europäischen Deltocephali. – Verh. zool.-bot. Ges. Wien 19: 201-222.
Fieber, F.X. (1872a): Katalog der europäischen Cicadinen, nach Originalien und Benützung der
neuesten Literatur. – Carl Gerold, Wien. I-IV, 1-19.
Fieber, F.X. (1872b): Berichtigungen zu Dr. Kirschbaum’s Cicadinen der Gegend von Wiesbaden,
Frankfurt a.M. und anderer Gegenden. – Verh. zool.-bot. Ges. Wien 12: 27-34.
Fieber, F.X. (1875): Les Cicadines d’Europe d’après les originaux et les publications les plus
récentes. Première partie. – Revue mag. Zool. (3) 3: 288-416.
References 397
Fieber, F.X. (1876): Les Cicadines d’Europe d’après les originaux et les publications les plus
récentes. Deuxième partie. – Revue mag. Zool. (3) 4: 11-268.
Fieber, F.X. (1877): Les Cicadines d’Europe d’après les originaux et les publications les plus
récentes. Deuxième partie (suite). – Revue mag. Zool. (3) 5: 1-45.
Fieber, F.X. (1878): Les Cicadines d’Europe d’après les originaux et les publications les plus
récentes. Troisième partie. – Revue mag. Zool. (3) 6: 270-308.
Fieber, F.X. (1879): Les Cicadines d’Europe d’après les originaux et les publications les plus
récentes. Quatrième partie. – Revue mag. Zool. (3) 7: 65-160.
Fischer, H. (1972): Die Tierwelt Schwabens. 21. Teil: Die Zikaden. – Ber. Naturf. Ges. Augsburg
27: 103-143.
Flint, J.H. (1990): Cicadula ornata (Melichar) (Hemiptera: Cicadellidae) new to Britain. – Ent.
Gaz. 40: 345-346.
Flor, G. (1861): Die Rhynchoten Livlands in systematischer Ordnung beschrieben. Zweiter Theil.
– Arch. Naturk. Liv-, Est- u. Kurlands 4: 1-567.
Floren, A., Schmidl, J. (1999): Faunistisch-ökologische Ergebnisse eines Baumkronen-Benebelung-
sprojektes in einem Eichenhochwald des Steigerwaldes. – Beitr. bayer. Entomofaun. 3: 179-195.
Förster, H. (1960): Zikaden-Lichtfänge. – Nachr.-bl. Oberlausitzer Insektenfreunde 4: 71-75.
Förster, H. (1961): Zur Zikadenfauna der Oberlausitz (Hemiptera – Homoptera). I. Typhlocy-
bidae. – Natura lusatica, Beitr. Erforsch. Lausitz 5: 61-72.
Franz, H. (1943): Die Landtierwelt der mittleren Hohen Tauern. – Denkschr. Akad. Wiss. Wien,
Math.-nat. Kl. 107: 1-552.
Fröhlich, W. (1996a): Beiträge zur Zoogeographie und Ökologie von Zikaden (Auchenorrhyn-
cha) auf Salzstellen in Mitteleuropa. – Thesis, University of Marburg. 205 pp.
Fröhlich, W. (1996b): Zikaden-Nachweise aus dem Gebiet des Neusiedler Sees (Österreich, Bur-
genland) und aus angrenzenden Gebieten (Insecta: Auchenorrhyncha). – Linzer biol. Beitr.
28(1): 335-347.
Fröhlich, W. (1997): Zur Salzverträglichkeit einiger Zikadenarten mitteleuropäischer Salzwie-
sen. – Beitr. Zikadenkde. 1: 17-33.
Frohne, D., Jensen, U. (1998): Systematik des Pflanzenreichs unter besonderer Berücksichtigung
chemischer Merkmale und pflanzlicher Drogen. 5. Aufl. – Wissenschaftliche Verlagsgesell-
schaft, Stuttgart. 371 pp.
Frommer, W. (1996): Untersuchungen zur Zikadenfauna (Hemiptera: Homoptera, Auchenor-
rhyncha) ausgewählter Standorte in Köln. – Decheniana (Suppl.) 35: 163-174.
Funke, T., Witsack, W. (1998): Zur Zikadenfauna der Bergbaufolgelandschaft ehemaliger Braun-
kohletagebaue in Sachsen-Anhalt (Homoptera, Auchenorrhyncha) – vorläufige Bestandsliste
– Beitr. Zikadenkde. 2: 39-51.
Futuyma, D.J. (1983): Selective factors in the evolution of host choice by phytophagous insects.
– In: Ahmad, S. (ed.): Herbivorous insects. Academic Press, New York. pp. 227-244.
Futuyma, D.J., Moreno, G. (1988): The evolution of ecological specialization. – Annu. Rev. Ecol.
Syst. 19: 207-233.
Garbarczyk, M. (1987): Leafhoppers (Homoptera, Auchenorrhyncha) and their parasitoids on
rye crops with different surroundings. – Pol. ecol. Stud. 13: 195-213.
Gatineau, F., Larrue, J., Clair, D., Lorton, F., Richard-Molard, M., Boudon-Padieu, E. (2001): A
new natural planthopper vector of stolbur phytoplasma in the genus Pentastiridius (Hemi-
ptera: Cixiidae). – Eur. J. Plant Path. 107: 263-271.
398 References
Gause, G.F. (1936): The principles of biocoenology. – Quart. Rev. Biol. 11: 320-336.
Gause, G.F. (1937): Experimental populations of microscopic organisms. – Ecology 18: 173-179.
Gauss, R. (1987): Integrierte Immigranten (Homoptera und Orthoptera) in unserer Entomofau-
na sowie zwei Zikaden-Abnormitäten. – Mitt. bad. Landesver. Naturkunde u. Naturschutz,
N.F. 14: 325-328.
Gȩbicki, C., Szwedo, J. (1998): New localities of species of the genus Cicadella Latreille, 1817 in
eastern Asia (Homoptera: Auchenorrhyncha: Cicadellidae). – Annals Upp. Sil. Mus. 8-9:
147-151.
Gȩbicki, C., Szwedo, J. (1991): Speudotettix montanus sp. nov. (Homoptera, Cicadellidae) from
Bieszczady. – Acta Biol. Silesiana 18 (35): 17-21.
Geiler, H. (1963): Artenlisten der Wanzen und Zikaden von Feldern sowie deren Abundanz und
Aktivitätsdichte während einzelner Jahre mit unterschiedlichem Witterungsverlauf. – Wiss.
Z. TU Dresden 12(2): 543-549.
Gillerfors, G. (2002): Nya arter för Sverige och nya landskapsfynd av stritar. – Entomologisk
Tidskrift 123(3): 109-116.
Gillham, M.C. (1991): Polymorphism, taxonomy and host plant association in Alebra leafhop-
pers (Homoptera: Cicadellidae: Typhlocybinae). – J. Nat. Hist. 25: 233-255.
Gillham, M.C., Claridge, M.F. (1994): A multivariate approach to host plant associated morpho-
logical variation in the polyphagous leafhopper Alnetoidia alneti (Dahlbom). – Biol. J. Linn.
Soc. 53: 127-151.
Gillham, M.C., Vrijer, P.W.F. de (1995): Patterns of variation in the acoustic calling signals of
Chloriona planthoppers (Homoptera: Delphacidae) coexisting on the common reed Phragmites
australis. – Biol. J. Linn. Soc. 54: 245-269.
Giustina, W. della (1983): La fauna de France des Cercopinae (Hom. Cicadomorpha). – Bull. Soc.
ent. Fr. 88: 192-196.
Giustina, W. della (1986): Metcalfa pruinosa (Say), nouveauté pour la fauna de France (Hom.:
Flatidae). – Bull. Soc. ent. Fr. 91(3-4): 89-92.
Giustina, W. della (1989): Homoptères Cicadellidae. Vol. 3. Compléments aux ouvrages d‘Henri
Ribaut. – Faune de France 73, Paris. 350 pp.
Giustina, W. della (1999): A review of the genus Zyginidia in France (Hemiptera: Auchenorrhyn-
cha: Cicadomorpha: Cicadellidae: Typhlocybinae). – Reichenbachia 33: 141-146.
Giustina, W. della, Balasse, H. (1999): Gone with the wind: Homoptera Auchenorrhyncha col-
lected by the French network of suction traps in 1994. – Marburger ent. Publ. 3(1): 7-42.
Giustina, W. della, Meusnier, S. (1982): Notes de chasse concernant les Cicadelles (Homoptera,
Cicadellidae). – Bull. Soc. ent. Fr. 87: 332-334.
Giustina, W. della, Navarro, E. (1993): Metcalfa pruinosa, un nouvel envahisseur? – Phytoma – La
défense des végétaux 451: 30-32.
Giustina, W. della, Remane, R. (1991): La faune de France des Delphacidae (Homoptera, Auchen-
orrhyncha). III – Récoltes et identifications de 1991. – Cahiers Natur. 47: 49-60.
Giustina, W. della, Remane, R. (1992): La faune de France des Delphacidae. II. Note de chasse
faites, pour l‘essentiel, en 1990 (Homoptera, Auchenorrhyncha). – Bull. Soc. ent. Fr. 96 (4):
313-330.
Giustina, W. della, Remane, R. (1999): La faune de France des delphacides, depuis 1985 (Ho-
moptera, Delphacidae). – Ann. Soc. entomol. Fr. (N.S.) 35 (suppl.): 274-280.
References 399
Hamilton, K.G.A. (1983b): Revision of the Macropsini and Neopsini of the New World (Rhyn-
chota: Homoptera: Cicadellidae), with notes on intersex morphology. – Mem. entomol. Soc.
Canada 123: 1-223.
Hamilton, K. (1994): Evolution of Limotettix Sahlberg (Homoptera: Cicadellidae) in peatlands,
with descriptions of new taxa. – Mem. Entomol Soc. Canada 169: 111-133.
Harborne, J.B. (ed.) (1972): Phytochemical ecology. – Academic Press, London. 272 pp.
Hartley, S.E., Jones, C.G. (1997): Plant chemistry and herbivory, or why the world is green. – In:
Crawley, M.J. (ed.): Plant ecology. 2nd ed., Blackwell, Oxford. pp. 284-324.
Harz, K. (1965): Zur Land-Fauna von Wangerooge. – Veröff. Inst. Meeresforsch. Bremerhaven 9:
210-231.
Harz, K. (1988): Zur Landfauna von Wangerooge. – Articulata 3 (5): 215-217.
Hassan, A.I. (1939): The biology of some British Delphacidae (Homoptera) and their parasites
with special reference to the Strepsiptera. – Trans. R. Ent. Soc. London 89: 345-384.
Haupt, H. (1924): Alte und neue Homoptera Mitteleuropas. – Konowia 3: 285-300.
Haupt, H. (1925): Über eine Homopteren-Ausbeute von Mittenwald und “Revision der Gattung
Cicadula Zett.”. – Mitt. Münchner Ent. Ges. 15: 9-40.
Haupt, H. (1935): Unterordnung: Gleichflügler, Homoptera. – In: Brohmer, P., Ehrmann P. Ulm-
er, G. (eds.): Die Tierwelt Mitteleuropas IV (X): 115-262.
Haupt, H., Hedicke, H. (1934): Die Fauna der Binnendüne bei Bellinchen (Oder). – Märk. Tierw.,
Berlin 1: 41-48.
Hegab, A.M., Orosz, A., Jenser, G. (1980): Observations on the larvae and imagoes of some Ally-
gus species (Homoptera). – Fol. ent. Hung. 33(1): 61-66.
Hegnauer, R. (1963): Chemotaxonomie der Pflanzen: Eine Übersicht über die Verbreitung und die
systematische Bedeutung der Pflanzenstoffe. Bd. 2. Monocotyledonae. – Birkhäuser, Basel. 540 pp.
Heller, F. (1960): Die Zikade Haematoloma dorsatum nun auch in Württemberg festgestellt. – Jh.
Ver. vaterl. Naturkde. Württemberg 115: 356-357.
Heller, F. (1975): Adarrus ocellaris (Fall.) ssp. tatraensis ssp. nova (Homoptera, Cicadellidae). –
Stuttgarter Beitr. Naturk. A, Nr. 288: 1-3.
Heller, F. (1985): Cercopis vulnerata (ROSSI) var. confluens var. nova (Homoptera, Cercopidae). –
Mitt. ent. Ver. Stuttgart 20: 16-18.
Heller, F. (1987a): Faunistische Untersuchungen im Feuchtgebiet “Unterer See” bei Horrheim
(Kreis Ludwigsburg). 5. Auchenorrhyncha, Zikaden. – Mitt. ent. Ver. Stuttgart 22: 76-92.
Heller, F. (1987b): Zwei für Deutschland neue Zikaden Japananus hyalinus und Calamotettix tae-
niatus (Homoptera: Cicadellidae). – Stuttgarter Beitr. Naturk. A, Nr. 401.
Heller, F. (1987c): Eine große Singzikade im Rosensteinpark in Stuttgart. – Mitt. ent. Ver. Stut-
tgart 22: 93-94.
Heller, F. (1994): Ribautiana debilis (Douglas), eine für Deutschland neue Kleinzikade (Homoptera:
Auchenorrhyncha). – Mitt. ent. Ver. Stuttgart 29: 119-120.
Heller, F. (1996): Vier für Deutschland neue Zikaden. Megadelphax haglundi (J. Sahlbg.), Arboridia
erecta (Rib. 1931), Fruticidia sanguinosa (Rey 1891) und Recilia horvathi (Then 1896) (Homoptera:
Auchenorrhyncha). – Mitt. ent. Ver. Stuttgart 31: 112-116.
Hempel, W., Hiebsch, H., Schiemenz, H. (1971): Zum Einfluß der Weidewirtschaft auf die Ar-
thropoden-Fauna im Mittelgebirge. – Faun. Abh. Staatl. Mus. Tierk. Dresden 3: 235-281.
Hennig, W. (1969): Die Stammesgeschichte der Insekten. – Waldemar Kramer, Frankfurt/Main.
436 pp.
402 References
Hoch, H., Asche, M. (1983): Wer ist Conomelus calabricus DLABOLA, 1979 (Homoptera Cicadina
Delphacidae). – Marburger ent. Publ. 1 (8): 1-6.
Hoch, H., Remane, R. (1983): Zur Artbildung und Artabgrenzung bei der binsenbesiedelnden
Zikadengattung Conomelus Fieber, 1866 (Homoptera Auchenorrhyncha Fulgoromorpha Del-
phacidae). – Marburger ent. Publ. 1(9): 1-115.
Hoch, H., Remane, R. (1985): Evolution und Speziation der Zikadengattung Hyalesthes Signo-
ret, 1865 (Homoptera: Auchenorrhyncha: Fulgoroidea: Cixiidae). – Marburger ent. Publ.
2(2): 1-427.
Hoffmann, B. (1980): Vergleichend ökologische Untersuchungen über die Einflüsse des kontrol-
lierten Brennens auf die Arthropodenfauna einer Riedwiese im Federseegebiet (Südwürt-
temberg). – Veröff. Naturschutz Landschaftspflege Bad.-Württ. 51/52(2): 691-714.
Hoffmann, H.-J. (1990): Zur Ausbreitung der Rhododendronzikade Graphocephala fennahi YOUNG
(Homoptera, Cicadellidae) in Deutschland, nebst Anmerkungen zu anderen Neueinwander-
ern bei Wanzen und Zikaden. – Verh. Westdt. Entom.–Tag. 1989: 285-301.
Hoffmann, H.-J. (1991): Zum Vorkommen der Bergzikade (Cicadetta montana). – Verh. Westdt.
Entom.–Tag. 1990: 201-210.
Hoffrichter, O., Tröger, E.J. (1973): Ceresa bubalus F. (Homoptera: Membracidae) – Beginn der
Einwanderung in Deutschland. – Mitt. bad. Landesverb. Naturk. Natursch., N.F. 11: 33-43.
Höhne, W. (1920): Coryphaelus gyllenhalii (Fall.), eine für die Mark Brandenburg und Deutschland
neue Zikade. – Deutsch. Ent. Z. 1920: 159-160.
Höhne, W. (1921): Paramesus reticulatus (Horváth), eine für Deutschland und die Mark Branden-
burg neue Zikade. – Deutsch. Ent. Z. 1921: 94-96.
Hollier, J.A., Brown, V.K., Edwards-Jones, G. (1994): Successional leafhopper assemblages: pat-
terns and process. – Ecological Research 9: 185-191.
Hölzel, E. (1965): Neues über Hemiptera – Homoptera (Zikaden) aus Kärnten. – Carinthia II
155/75: 115-126.
Holzinger, W. (1995a): Bemerkenswerte Zikadenfunde aus Österreich, 2. Teil (Ins.: Homoptera,
Auchenorrhyncha). – Linzer biol. Beitr. 27/2: 1123-1127.
Holzinger, W. (1995b): Bemerkenswerte Zikadenfunde aus Österreich (Homoptera, Auchenor-
rhyncha: Cicadellidae). – Linzer biol. Beitr. 27/1: 279-283.
Holzinger, W. (1995c): Zikaden (Auchenorrhyncha). – In: Wieser, C., Kofler, A., Mildner, P. (eds.):
Naturführer Sablatnigmoor. Naturwiss. Ver. Kärnten, Klagenfurt. pp. 121-128.
Holzinger, W. (1996a): Kritisches Verzeichnis der Zikaden Österreichs (Ins.: Homoptera, Auchen-
orrhyncha). – Carinthia II 186/106: 501-517.
Holzinger, W. (1996b): Die Zikadenfauna wärmeliebender Eichenwälder Ostösterreichs (Insec-
ta: Homoptera, Auchenorrhyncha). – Mitt. naturwiss. Ver. Steiermark 126: 169-187.
Holzinger, H. (1999a): Taxonomie und Verbreitung ausgewählter Zikadenarten Österreichs (In-
secta: Hemiptera: Auchenorrhyncha). – Faun. Abh. Staatl. Mus. Tierkde. Dresden 21: 259-264.
Holzinger, W.E. (1999b): Rote Liste der Zikaden Kärntens (Insecta: Auchenorrhyncha). – In: Rot-
tenburg, T., Wieser, C., Mildner, P., Holzinger, W.E. (Hrsg.): Rote Liste gefährdeter Tiere Kärn-
tens. Naturschutz in Kärnten 15: 425-450.
Holzinger, W., Fröhlich, W., Günthart, H., Lauterer, P., Nickel, H., Orosz, A., Schedl, W., Remane,
R. (1997): Vorläufiges Verzeichnis der Zikaden Mitteleuropas (Insecta: Auchenorrhyncha). –
Beitr. Zikadenkunde 1: 43-62.
404 References
Holzinger, W., Günthart, H., Lauterer, P., Nickel, H., Remane, R. (1997): Zum Herbstaspekt der
Zikadenfauna der Rabenhofteiche und Weinburger Teiche (Steiermark, Österreich; Insecta:
Auchenorrhyncha). – Beitr. Zikadenkunde 1: 37-42.
Holzinger, W., Jantscher,E., Remane, R. (1996): Erstnachweise von Zikaden aus Österreich, mit
Bemerkungen zu weiteren Arten (Ins.: Homoptera, Auchenorrhyncha). – Linzer biol. Beitr.
28/2: 1149-1152.
Holzinger, W.E., Kammerlander, I., Nickel, H. (2003, in print): The Auchenorrhyncha of Central
Europe – Die Zikaden Mitteleuropas. Volume 1: Fulgoromorpha, Cicadomorpha excl. Cicadel-
lidae. – Brill, Leiden. c. 590 pp.
Holzinger, W., Remane, R. (1994): Zikaden-Erstnachweise aus Österreich (Ins.: Homoptera
Auchenorrhyncha). – Mitt. naturwiss. Ver. Steiermark 124: 237-240.
Holzinger, W.E., Seljak, G. (2001): New records of planthoppers and leafhoppers from Slovenia,
with a checklist of hitherto recorded species (Hemiptera: Auchenorrhyncha). – Acta entomol.
Slovenica 9(1): 39-66.
Howe, H.F., Westley, L.C. (1988): Ecological relationships of plants and animals. – Oxford Uni-
versity Press, New York. 273 pp.
Hüeber, T. (1904): Catalogus insectorum faunae germanicae: Hemiptera homoptera. Systematis-
ches Verzeichnis der Zikadinen Deutschlands (und der nächst angrenzenden Landesteile). –
Jh. Ver. vaterl. Naturkde. Württemberg 60: 253-277.
Hunter, C.E., Yeargan, K.V. (1989): Development, reproduction, and competitive interactions
between two sympatric leafhopper species (Homoptera: Cicadellidae) on redbud trees. –
Environ. Entomol. 18(1): 127-132.
Hutchinson, G.E. (1959): Homage to Santa Rosalia or why are there so many kinds of animals?
– Am. Nat. 93: 145-159.
Jacobs, W., Renner, M. (1998): Biologie und Ökologie der Insekten. – 3rd ed., Spektrum, Heidel-
berg. 700 pp.
Jacobson, J., Crosby, D.G. (eds.) (1971): Naturally occurring insecticides. – Marcel Dekker Inc.,
New York. 585 pp.
Jansky, V., Okali, I. (1993): Check-list of the Slovak Auchenorrhyncha. – Zbor. Slov. nar. Muz.,
Prir. Vedy 39: 37-59.
Janzen, D. (1966): Coevolution of mutualism between ants and acacias in Central America. –
Evolution 20: 249-275.
Jensen-Haarup, A.C. (1920): Cikader. – Danmarks Fauna 24: 1-189.
Jermy, T. (1976): Insect – host-plant relationship – co-evolution or sequential evolution? – Symp.
Biol. Hung. 16: 109-113.
Johnson (1969): Migration and dispersal of insects by flight. – Methuen, London. 763 pp.
Jones, C.G., Lawton, J.H. (1991): Plant chemistry and insect species richness of British umbelli-
fers. – J. Anim. Ecol. 60: 767-777.
Jöst, H. (1962): Funde seltener Zikaden in der Pfalz. – Pfälzer Heimat 13: 24.
Jöst, H. (1966): Ein Beitrag zur Kenntnis der Zikadenfauna der Pfalz (Homoptera-Cicadina). –
Pfälzer Heimat 17: 144-146.
Judd, W.S., Campbell, C.S., Kellog, E.A., Stevens, P.F. (1999): Plant systematics. A phylogenetic
approach. – Sinauer, Sunderland/Massachusetts. 464 pp.
References 405
Kontkanen, P. (1948): Beiträge zur Kenntnis der Zikadenfauna Finnlands. III. – Ann. ent. Fenn.
14: 85-97.
Kontkanen, P. (1952): On the sibling species in the leafhopper fauna of Finland (Homoptera,
Auchenorrhyncha). – Soc. Vanamo Arch. 7: 100-106.
Kontkanen, P. (1953): Studies in insect populations I. The number of generations of some leaf-
hopper species in Finland and Germany. – Soc. Vanamo Arch. 8: 150-156.
Kopp, D.D., Yonke, T.R. (1977): Taxonomic status of the buffalo treehopper and the name Ceresa
bubalus. – Ann. ent. Soc. Am. 70: 901-905.
Körber-Grohne, U. (1990): Gramineen und Grünlandvegetationen vom Neolithikum bis zum
Mittelalter in Mitteleuropa. – Bibiotheca botanica 139: 1-105.
Körner, M., Nickel, H., Sayer, M. (2001): Zikaden an Gräsern eines norddeutschen Kiefernfor-
stes – Abundanz, Phänologie und ein Vergleich der Fangmethoden. – Beitr. Zikadenkde. 4:
33-43.
Krause, R., Emmrich, R. (1996): Zur Käfer- und Zikadenfauna einer kleinen, verwilderten Wein-
bergparzelle im Dresdner Elbtal (Coleoptera, Auchenorrhyncha). – Verh. 14. Int. Symp. Ento-
mofaunistik Mitteleuropa, SIEEC, München (4.-9.9.1994): 165-173.
Kristensen, N.P. (1975): The phylogeny of hexapod ‘orders’. A critical review of recent accounts.
– Z. zool. Syst. Evol.-forschg. 13: 1-44.
Kristensen, N.P. (1995): Forty years’ insect phylogenetic systematics. – Zool. Beitr. (N.F.) 36: 83-124.
Kudryásheva, I.V. (1975): Analysis of changes in insect weight indices in connection with the
instar structure of the population using Cicadetta montana (Scop.) (Homoptera: Cicadidae) as
an example. – Zh. Obshch. Biol. 36: 373-381. (In Russian).
Kuntze, A. (1937): Die Zikaden Mecklenburgs, eine faunistisch-ökologische Untersuchung. –
Arch. Naturgesch., N.F. 6: 299-388.
Kupka, P. (1925): Fränkische Cicadinen. – Mitt. Münchner ent. Ges. 15: 102-113.
Kupka, P. (1941): Bemerkungen zur mitteleuropäischen Cicadinenfauna. – Mitt. Dtsch. ent. Ges.
10: 57-62.
Kurir, A. (1967): Auswertung der Ergebnisse nach einem Massenbefall der Weidenschaumzikade
(Aphrophora salicina Goeze) auf der Amerikaner-Weide (Salix americana Hort.) in einer Flecht-
weidenkultur. – Z. angew. Entomol. 59: 152-186.
Lafont, R., Bouthier, A., Wilson, I.D. (1991): Phytoecdysteroids, structures, occurrence, biosyn-
thesis and possible ecological significance. – In: Hrdý, I. (ed.): Insect chemical ecology – Pro-
ceedings of a conference held in Tábor, Czechoslowakia, 12 -18 August 1990. pp. 197-214.
Lang, G. (1994): Quartäre Vegetationsgeschichte Europas. – Gustav Fischer, Jena. 462 pp.
Lang, V. (1945): Zlomky z cikadologického vyzkumu Mohelnské Reservace. – Folia ent. 8: 95-99.
Lauterborn, R. (1920): Cicadula cyanea Boheman. – Mitt. Bad. Landesver. Naturkd. 1(4): 200.
Lauterborn, R. (1923): Jassus stactogala Amyot. – Mitt. Bad. Landesver. Naturkd. 1(11): 288.
Lauterer, P. (1958): Prispewek k poznani krisu CSR. (Hom., Auchenorrhyncha). II. – Acta Mus.
Morav. 43: 125-136.
Lauterer, P. (1978): New records of leafhoppers from Czechoslovakia (Homoptera, Auchenor-
rhyncha). – Acta Mus. Morav. 63: 111-116.
Lauterer, P. (1980): New and interesting records of leafhoppers from Czechoslovakia (Homoptera,
Auchenorrhyncha). – Acta Mus. Morav. 65: 117-140.
Lauterer, P. (1983): Fagocyba cerricola sp. n. and new and interesting records of leafhoppers from
Czechoslovakia (Homoptera, Auchenorrhyncha). – Acta Mus. Morav. 68: 139-152.
References 407
Lauterer, P. (1984): New and interesting records of leafhoppers from Czechoslovakia (Homoptera,
Auchenorrhyncha) II. – Acta Mus. Morav. 69: 143-162.
Lauterer, P. (1986): New and interesting records of leafhoppers from Czechoslovakia (Homoptera,
Auchenorrhyncha) III. – Acta Mus. Morav. 71: 179-187.
Lauterer, P. (1989): New synonymy of Japananus hyalinus (Osborn, 1900), and new findings of the spe-
cies in Hungary and Bulgaria (Homoptera: Cicadellidae). – Folia entomol. Hungarica 50: 37-40.
Lauterer, P. (2000): First reports of some leafhoppers (Auchenorrhyncha: Cicadellidae) from the
Czech Republic and notes from Slovakia. – Acta Mus. Morav., Sci. biol. 85: 327-332.
Lauterer, P., Holzinger, W.E. (1996): New findings of Homoptera (Auchenorrhyncha and Psyl-
loidea) in Austria. – Acta Mus. Morav. 80: 251-253.
Lauterer, P., Malenovsky, I. (1995): Eine neue Art der Kleinzikaden (Homoptera Cicadellidae Typhlo-
cybinae) auf dem Gebiet der Bundesrepublik Deutschland. – Marburger ent. Publ. 2 (9): 76-77.
Lauterer, P., Novotný, V. (1991): New findings of leafhoppers (Homoptera, Auchenorrhyncha)
in Czechoslovakia. – Acta Mus. Morav. 76: 265-268.
Lawton, J.H. (1976): The structure of arthropod communities on bracken. – Bot. J. Linn. Soc. 73:
187-216.
Lawton, J.H. (1978): Host plant influences on insect diversity: the effects of space and time. – In:
Mound. L.A., Waloff, N. (eds.): Diversity of insect faunas. Blackwell, Oxford. pp. 105-125.
Lawton, J.H. (1982): Vacant niches and unsaturated communities: a comparison of bracken her-
bivores at sites on two continents. – J. Anim. Ecol. 51: 573-595.
Lawton, J.H. (1983): Plant architecture and the diversity of phytophagous insects. – Ann. Rev.
Entomol. 28: 23-39.
Lawton, J.H., McNeill, S. (1979): Between the devil and the deep blue sea: on the problem of
being a herbivore. – In: Anderson, R.M., Turner, B.D., Taylor, L.R. (eds.): Population dynam-
ics. Blackwell, Oxford. pp. 223-244.
Lawton, J.H., Schroder, D. (1977): Effects of plant type, size of geographical range and taxonom-
ic isolation on number of insect species associated with British plants. – Nature 265: 137-140.
Le Quesne, W. (1960): Hemiptera (Fulgoromorpha). – Handb. Ident. Br. Insects II (3). 68 pp.
Le Quesne, W. (1964): Trigonocranus emmeae Fieber (Hem., Cixiidae), new to Britain. – Entomol-
ogist‘s month. Mag. 100: 117.
Le Quesne, W. (1965): Hemiptera (Cicadomorpha) (excluding Deltocephalinae and Typhlocybi-
nae). – Handb. Ident. Br. Insects II (2a): 1-64.
Le Quesne, W. (1969): Hemiptera (Cicadomorpha) Deltocephalinae. – Handb. Ident. Br. Insects
II (2b): 65-148.
Le Quesne, W. (1972): Studies on the coexistence of three species of Eupteryx on nettle. – J. Ent.
47(1): 37-44.
Le Quesne, W. (1977): A new species of Lindbergina (Hemiptera: Cicadellidae) from Jersey. –
Annual Bull. Soc. Jersiaise 22(1): 87-90.
Le Quesne, W. (1987): Cicadella lasiocarpae OSSIANNILSSON (Hem.: Cicadellidae) new to Brit-
ain. – Ent. Gazette 38: 87-89.
Le Quesne, W., Payne, K.R. (1981): Cicadellidae (Typhlocybinae) with a checklist of the British
Auchenorrhyncha (Hemiptera, Homoptera). – Handb. Ident. Br. Insects II (2 c). 95 pp.
Leather, S.R. (1985): Does the bird cherry have its “fair share” of insect pests? An appraisal of the
species – area relationship of the phytophagous insects associated with British Prunus spe-
cies. – Ecol. Entomol. 10: 43-56.
408 References
Lehmann, W. (1973a): Untersuchungen der Zikadenfauna von Obstgehölzen. – Biol. Zbl. 92: 75-95.
Lehmann, W. (1973b): Untersuchungen der Zikadenfauna von Obstanlagen mit Hilfe von Licht-
fallen. – Biol. Zbl. 92: 625-635.
Leising, S. (1977): Über Zikaden des zentralalpinen Hochgebirges (Obergurgl, Tirol). – Alpin-
biol. Stud. IX: 1-69.
Leuchtmann, A., Schmidt, D., Bush, L.P. (2000): Different levels of protective alkaloids in grasses
with stroma-forming and seed-transmitted Epichloe/Neotyphodium endophytes. – J. Chem.
Entomol. 26: 1025-1036.
Lewis, T., Taylor, L.R. (1965): Diurnal periodicity of flight by insects. – Trans. R. ent. Soc. London
116: 393-476.
Lindberg, H. (1947): Verzeichnis der ostfennoskandischen Homoptera Cicadina. – Fauna Fenni-
ca 1: 1-81.
Lindberg, H. (1948): Materialien zu einer Monographie der Gattung Tettigometra (Homoptera,
Cicadina). – Notul. ent. 28: 1-40.
Lindblad, M., Arenö, P. (2002): Temporal and spatial population dynamics of Psammotettix alie-
nus, a vector of wheat dwarf virus. – Int. J. Pest Management 48(3): 233-238.
Lindsten, K. (1979): Planthopper vectors and plant disease agents in Fennoscandia. – In: Mar-
amorosch, K., Harris, K.F. (eds.): Leafhopper vectors and plant disease agents. Academic
Press, New York. pp. 155-178.
Linnavuori, R. (1951): Hemipterological observations. – Ann. ent. Fenn. 17: 51-65.
Linnavuori, R. (1952): Studies on the ecology and phenology of the leafhoppers (Homoptera) of
Raisio (SW-Finland). – Ann. Soc. zool.-bot. Fenn. Vanamo 14(6): 1-32.
Linnavuori, R. (1957): Remarks on some italian Delphacidae. – Boll. Soc. ent. Ital. 87: 49-52.
Linnavuori, R. (1969a): Nivelkärsäiset III. Hemiptera III. Kaskaat 1. – Animalia Fennica 12: 1-244.
Linnavuori, R. (1969b): Nivelkärsäiset IV. Hemiptera IV. Kaskaat 2. – Animalia Fennica 13: 1-312.
Löcker, H. (2003, submitted): Arborikole Zikadengilden Sloweniens. – Beitr. Zikadenkde. 6.
Logvinenko, V.N. (1975): Fulgoroidny cikadovy Fulgoroidea. – Fauna Ukrainy, Kyiv, 20(2). 287
pp. (In Ukrainian).
Louis, F., Schirra, K.-J. (1997): Grüne Rebzikade – Ein Problem? – Das deutsche Weinmagazin 14:
28-30.
Lovinger, A., Liewehr, D., Lamp, W. (2000): Glandular trichomes on alfalfa impede searching
behavior of the potato leafhopper parasitoid. – Biol. Contr. 18: 187-192.
MacArthur, R.H. (1962): Some generalized theorems of natural selection. – Proc. Natl. Acad. Sci.
USA 48: 1893-1897.
Mai, D.H. (1995): Tertiäre Vegetationsgeschichte Europas. – Gustav Fischer, Jena. 691 pp.
Maixner, M., Ahrens, U., Seemüller, E. (1995): Detection of the German grapevine yellows (Vergil-
bungskrankheit) MLO in grapevine, alternative hosts and a vector by a specific PCR proce-
dure. – Eur. J. Plant Path. 101: 241-250.
Maixner, M., Reinert, W. (1999): Oncopsis alni (Schrank) (Auchenorrhyncha: Cicadellidae) as a vector
of the alder yellows phytoplasma of Alnus glutinosa (L.) Gaertn. – Eur. J. Plant Path. 105: 87-94.
Manurung, B., Witsack, W., Fuchs, E., Mehner, S. (2001): Zur Embryonal- und Larvalentwick-
lung der Zikade Psammotettix alienus (Dahlbom, 1851) (Hemiptera, Auchenorrhyncha). – Bei-
tr. Zikadenkde. 4: 49-58.
Maramorosch, K., Harris, K.F. (1979) (eds.): Leafhopper vectors and plant disease agents. – Ac-
ademic Press, New York. 654 pp.
References 409
Marchand, H. (1952): Ein weiterer Fund von Stenocranus longipennis CURT. (Hem., Hom., Del-
phacidae) in Nordwestdeutschlands. – Faun. Mitt. Norddeutschl. 2: 4-5.
Marchand, H. (1953): Die Bedeutung der Heuschrecken und Schnabelkerfe als Indikatoren ver-
schiedener Graslandtypen. – Beitr. Ent. 3: 116-162.
Marion-Poll, F., Giustina, W. della, Mauchamp, B. (1987): Changes of electric patterns related to
feeding in a mesophyll feeding leafhopper Zyginidia scutellaris. – Entomol. exp. appl. 43: 115-
124.
Masters, G.J., Brown, V.K. (1997): Host-plant mediated interactions between spatially separated
herbivores: effects on community structure. – In: Gange, A.C., Brown, V.K. (eds.): Multitrophic
interactions in terrestrial systems. Blackwell, Oxford. pp. 217-237.
Matsumura, S. (1906): Die Cicadinen der Provinz Westpreußen und des östlichen Nachbarge-
biets. – Schr. naturforsch. Ges. Danzig, N.F. 11: 64-82.
Mattson, W.J.Jr. (1980): Herbivory in relation to plant nitrogen content. – Annu. Rev. Ecol. Syst.
11: 119-161.
Mauri, G. (1982): Nore sulla biologia della cicaletta nerorossa (Cercopis sanguinea Geoffr.) (Auchen.,
Cercopidae) e possibilità di lotta. – Mitt. Schweiz. ent. Ges. 55: 87-92.
May, Y.Y. (1978): A population study of Stenocranus minutus (Fab.) (Hemiptera: Delphacidae). –
Res. Popul. Ecol. 20: 61-78.
McClure, M.S., Price, P.W. (1975): Competition among sympatric Erythroneura leafhoppers (Ho-
moptera: Cicadellidae) on American sycomore. – Ecology 56: 1388-1397.
McClure, M.S., Price, P.W. (1976): Ecotype characteristics of coexisting Erythroneura leafhoppers
(Homoptera: Cicadellidae) on sycomore. – Ecology 57: 928-940.
McEvoy, P.B. (1986): Niche partitioning in spittlebugs (Homoptera: Cercopidae) sharing shelters
on host plants. – Ecology 67(2): 465-478.
McNeill, S. (1973): The dynamics of a population of Leptopterna dolobrata (Heteroptera: Miridae)
in relation to its food resources. – J. Anim. Ecol. 42: 495-507.
McNeill, S., Southwood, T.R.E. (1978): The role of nitrogen in the development of insect/plant
relationships. – In: Harborne, J.B. (Ed.): Biochemical Aspects of Plant and Animal Coevolu-
tion. Academic Press, London. pp. 77-98.
Melber, A. (1989): Entwicklung und Populationsdynamik der Heidezikade Ulopa reticulata (Hom.,
Auchenorrhyncha, Cicadellidae) in nordwestdeutschen Calluna-Heiden. – Zool. Jb. Syst. 116:
21-30.
Melber, A., Prüter, J., Assing, V., Sprick, P. (1996): Erste Ergebnisse der Erfassung ausgewählter
Wirbellosen-Gruppen in einer kleinen Vegetationsinsel auf den Panzerübungsflächen des NSG
Lüneburger Heide (Heteroptera; Homoptera; Auchenorrhyncha; Coleoptera, Carabidae, Sta-
phylinidae, Curculionidae). – NNA-Ber. 9(1): 93-102.
Melichar, L. (1896): Cicadinen (Hemiptera – Homoptera) von Mittel-Europa. – Felix L. Dames,
Berlin. 364 pp.
Melichar, L. (1899): Eine entomologische Reise nach dem Süden. – Ent. Jb. 1899: 138-143.
Metcalf, Z.P. (1932). General Catalogue of the Hemiptera. Fasc. IV, Fulgoroidea. Pt 1, Tettigo-
metridae. – Smith College, Northampton, Mass. 69 pp.
Metcalf, Z.P. (1967). General Catalogue of the Homoptera. Fasc. VI, Cicadelloidea. Pt 10, Euscel-
idae. – U.S.D.A., Washington, D.C. 2695 pp.
Metcalf, Z.P. (1968). General Catalogue of the Homoptera. Fasc. VI, Cicadelloidea. Pt 17, Ci-
cadellidae. – U.S.D.A., Washington, D.C. 1513 pp.
410 References
Meusel, H., Jäger E., Weinert, E. (1964): Vergleichende Chorologie der zentraleuropäischen Flo-
ra. Band 1, Kartenteil. – Fischer, Jena. 258 pp.
Meyer, G.A. (1993): A comparison of the impact of leaf- and sap-feeding insects on growth and
allocation of goldenrod. – Ecology 74: 1101-1116.
Meyer-Arndt, S, Remane, R. (1992): Phylogenie und Speziation der Fieberiellini WAGNER, 1951
(Homoptera: Auchenorrhyncha: Cicadellidae). – Marburger ent. Publ. 2 (7): 1-756.
Mityaev, I.D. (1971): Leafhoppers of Kazakhstan (Homoptera – Cicadinea). – Nauka, Alma-Ata.
212 pp. (In Russian).
Mölleken, H., Topp, W. (1997): Die Insektenfauna auf Silberweiden (Salix alba L.): Einfluß des
Geschlechts und der Pflegemaßnahmen. – Z. Ökol. Natursch. 6: 193-206.
Moosbrugger, J. (1946): Die Zikadenfauna von Vorarlberg. – Zentralbl. Gesamtgeb. Ent. 1: 65-75.
Moraal, L.G. (1996): Bionomics of Haematoloma dorsatum (Hom., Cercopidae) in relation to needle
damage in pine forests. – Anz. Schädlingskde., Pflanzenschutz, Umweltschutz 69: 114-118.
Moravskaja, A.S. (1948): To the knowledge of the genus Zyginidia (Homoptera – Cicadina). –
Nauchno-metodiches-kye Zapiski 11: 198-207. (In Russian).
Morcos, G. (1953): The biology of some Hemiptera-Homoptera. – Bull. Soc. Faoud. Ent. 37: 405-439.
Moreby, S.J., Stoate, C. (2001): Relative abundance of invertebrate taxa in the nestling diet of
three farmland passerine species, Dunnock Prunella modularis, Whitethroat Sylvia communis
and Yellowammer Emberiza citrinella in Leicestershire, England. – Agiculture, Ecosystems and
Environment 86: 125-134.
Morris, M.G. (1972): Distributional and ecological notes on Ulopa trivia GERMAR (Hem., Ci-
cadellidae). – Entomologist‘s mon. Mag. 107: 174-181.
Morris, M.G. (1990a): The Hemiptera of two sown calcareous grasslands. I. Colonization and
early succession. – J. appl. Ecol. 27: 367-378.
Morris, M.G. (1990b): The Hemiptera of two sown calcareous grasslands. III. Comparisons with
the Auchenorrhyncha faunas of other grasslands. – J. appl. Ecol. 27: 394-409.
Mühlethaler, R. (2001): Untersuchungen zur Zikadenfauna der Lebensraumtypen von Basel. –
Beitr. Zikadenkde. 4: 11-32.
Müller, H.J. (1942): Über Bau und Funktion des Legeapparates der Zikaden (Homoptera, Cica-
dina). – Z. Morph. Ökol. Tiere 38: 534-629.
Müller, H.-J. (1956): Homoptera – In: Sorauer, P. (Hrsg.): Handbuch der Pflanzenkrankheiten. 5.
Aufl., Bd. 5, 3. Lfg. pp. 150-359.
Müller, H.-J. (1957): Über die Diapause von Stenocranus minutus Fabr. (Homoptera: Auchenor-
rhyncha). – Beitr. Entomol. 7: 203-226.
Müller, H.J. (1961): Erster Nachweis einer Eidiapause bei den Jassiden Euscelis plebejus Fall. und
lineolatus Brullé (Homoptera Auchenorrhyncha). – Z. angew. Ent. 48: 233-241.
Müller, H.J. (1970): Formen der Dormanz bei Insekten. – Nova Acta Leopoldina (N.F.) 35: 7-27.
Müller, H.J. (1972): Auchenorrhyncha. Unterordnung Cicadaria, Zikaden. In: Schwenke, W.
(Hrsg.): Die Forstschädlinge Europas. 1. Band. Würmer, Schnecken, Spinnentiere, Tausend-
füssler und hemimetabole Insekten. – Paul Parey, Hamburg and Berlin, pp. 127-150.
Müller, H.J. (1976): Über die Parapause als Dormanzform am Beispiel der Imaginaldiapause
von Mocydia crocea H.-S. (Homoptera Auchenorrhyncha). – Zool. Jb. Allg. Zool. Phys. 80(3):
231-258.
Müller, H.J. (1981): Die Bedeutung der Dormanzform für die Populationsdynamik der Zwerg-
zikade Euscelis incisus (Kbm.) (Homoptera, Cicadellidae). – Zool. Jb. Syst. 108: 314-334.
References 411
Müller, H.J. (1984a): Über den Voltinismus der Dornzikade Centrotus cornutus (L.) (Homoptera,
Auchenorrhyncha: Membracidae) und die Einnischung mehrjähriger Insekten. – Zool. Jb.
Syst. 111: 321-337.
Müller, H.J. (1984b): Zur Entwicklung und Lebensweise der Larven der Dornzikade Centrotus
cornutus (L.) (Homoptera Auchenorrhyncha: Membracidae) unter besonderer Berücksichti-
gung der Kotschleuder. – Zool. Jb. Anat. 111: 385-399.
Müller, H.J. (1992): Dormanz bei Arthropoden. – Gustav Fischer, Jena. 289 pp.
Müller, J. (1969): Untersuchungen über die intrazelluläre Symbiose einiger Aetalionidae, Eu-
rymelidae und Cicadellidae (Homoptera – Auchenorrhyncha). – Zool. Jb. Syst. 96: 558-608.
Murdoch, W., Evans. F.C., Peterson, C.H. (1972): Diversity and pattern in plants and insects. –
Ecology 53: 819-829.
Musil, M. (1956): Beitrag zur Kenntnis der Entwicklungsstufen von Hyalesthes obsoletus Sign. –
Zool. List. 5(19): 17-22.
Musil, M. (1963): O vyskytu nekterych Krisu na Slovensku (Hom. Auchenorrhyncha). – Biolo-
gia, Bratislava 18 (9): 693-697.
Nast, J. (1958): Homopterological notes X – XII. – Acta zool. Crac. 2(35): 887-899.
Nast, J. (1966): Two new Palaearctic Delphacidae (Homoptera). – Bull. Acad. Pol. Sci (Cl. II) 13:
643-646.
Nast, J. (1972): Palaearctic Auchenorrhyncha (Homoptera). An annotated check list. – Polish
Scientific Publ., Warszawa. 550 pp.
Nast, J. (1975): On some Mediterranean Delphacidae (Homoptera, Auchenorrhyncha) described
by S. MATSUMURA. – Ann. zool. Warsz. 33(1): 1-15.
Nast, J. (1976a): Piewiki. Auchenorrhyncha (Cicadodea). – Katalog Fauny Polski 21. 257 pp.
Nast, J. (1976b): Piewiki (Homoptera, Auchenorrhyncha) Pienin. – Fragm. faun. 21(6): 145-183.
Nast, J. (1977): Homopterological notes XIII – XX. – Ann. zool. Warszawa 34: 27-37.
Nast, J. (1979): Palaearctic Auchenorrhyncha (Homoptera). Part 2, bibliography; addenda and
corrigenda. – Ann. zool. Warszawa 34(18): 481-499.
Nast, J. (1981): Homopterological notes XXI – XXV. – Ann. zool. Warszawa 36(14): 255-263.
Nast, J. (1982): Palaearctic Auchenorrhyncha (Homoptera). Part 3. New taxa and replacement
names introduced till 1980. – Ann. zool. Warszawa 36 (17): 289-362.
Nast, J. (1984): Notes on some Auchenorrhyncha (Homoptera), 1-5. – Ann. zool. Warszawa 37:
391-398.
Nast, J. (1986): Notes on some Auchenorrhyncha (Homoptera), 6-10. – Ann. zool. Warszawa 40:
297-307.
Nast, J. (1987): The Auchenorrhyncha (Homoptera) of Europe. – Ann. zool. Warszawa 40: 535-662.
Nault, L.R., Rodriguez, J.G., (Hrsg.) (1985): The leafhoppers and planthoppers. – Wiley, New
York. 500 pp.
Nickel, B. (1979): Untersuchungen über die Ernährung und die Ausscheidungen einheimischer
Cercopidenlarven. – Thesis, Technical University of Munich. 118 pp.
Nickel, H. (1994): Wärmeliebende Zikaden (Homoptera, Auchenorrhyncha) im südlichen Nie-
dersachsen. – Braunschweiger naturkdl. Schr. 4(3): 533-551.
Nickel, H. (1997): Zur Verbreitung und Lebensweise einiger Zikadenarten in Niedersachsen
und angrenzenden Gebieten (Homoptera, Auchenorrhyncha). – Göttinger naturkdl. Schr. 4:
151-172.
412 References
Nickel, H. (1998): Zum Vorkommen der Zwergzikade Edwardsiana rhodophila (Cerutti, 1937) in
den Wärmegebieten Ost- und Süddeutschlands (Hemiptera, Auchenorrhyncha, Cicadellidae).
– Hercynia (N.F.) 31: 277-281.
Nickel, H. (1999a): Life strategies of Auchenorrhyncha species on river floodplains in the north-
ern Alps, with description of a new species: Macropsis remanei sp. n. (Hemiptera). – Reichen-
bachia 33: 157-169.
Nickel, H. (1999b): Zum Vorkommen einiger Zikadenarten in Bayern. – Nachr.-Bl. bayer. Ent.
48(1/2): 2-19.
Nickel, H. (2001): Zikaden. – In: Der Steinberg bei Wesseln. Natur und Landschaft im Landkreis
Hildesheim 2: 105-109.
Nickel, H. (2002): Die Zikadenfauna der Hochmoore im Thüringer Wald vor 25 Jahren und heu-
te. – Naturschutzreport 19: 116-138.
Nickel, H., Achtziger, R. (1999): Wiesen bewohnende Zikaden (Auchenorrhyncha) im Gradient-
en von Nutzungsintensität und Feuchte. – Beitr. Zikadenkde. 3: 65-80.
Nickel, H., Achtziger, R., Lauterer, P., Malenovský, I., Weis, A., Witsack, W. (2001): Zur Fauna der
Zikaden, Wanzen, Blattflöhe und Augenfliegen des Kyffhäusergebirges (Hemiptera: Auchen-
orrhyncha, Heteroptera et Psylloidea; Diptera: Pipunculidae). – Beitr. Zikadenkde. 4: 75.-80.
Nickel, H., Hildebrandt, J. (2003, accepted): Auchenorrhyncha communities as indicators of dis-
turbance in grasslands (Insecta, Hemiptera) – A case study from the middle Elbe valley (North
Germany). – Agriculture, Ecosystems and Environment.
Nickel, H., Holzinger, W.E., Wachmann, E. (2002a): Mitteleuropäische Lebensräume und ihre
Zikaden (Insecta: Hemiptera: Auchenorrhyncha). – Denisia 4 (N.F.) 176: 279-328.
Nickel, H., Holzinger, W.E., Lauterer, P., Remane, R., Witsack, W. (2002b, submitted): Die Sporn-
zikadengattung Metropis Fieber, 1866 in Mitteleuropa (Hemiptera, Fulgoromorpha: Delpha-
cidae). – Beitr. Zikadenkde. 6.
Nickel, H., Remane, R. (1996): Erfassungsstand der Zikadenfauna Bayerns, mit Anmerkungen
zum Nährpflanzenspektrum und Habitat (Homoptera, Auchenorrhyncha). – Verh. 14. Int.
Symp. Entomofaunistik Mitteleuropa, SIEEC, München (4.-9.9.1994): 407-420.
Nickel, H., Remane, R. (2002): Artenliste der Zikaden Deutschlands, mit Angabe von Nähr-
pflanzen, Nahrungsbreite, Lebenszyklus, Areal und Gefährdung (Hemiptera, Fulgoromor-
pha et Cicadomorpha). – Beitr. Zikadenkde. 5: 27-64.
Nickel, H., Sander, F.W. (1996): Kommentiertes Verzeichnis der bisher in Thüringen nachgewie-
senen Zikadenarten (Homoptera, Auchenorrhyncha). – Veröff. Naturkundemus. Erfurt 15:
152-170.
Nickel, H., Witsack, W., Remane, R. (1999): Rote Liste der Zikaden Deutschlands (Hemiptera,
Auchenorrhyncha) – Habitate, Gefährdungsfaktoren und Anmerkungen zum Areal. – Beitr.
Zikadenkde. 3: 13-32.
Nicolaus, M. (1957): Zikaden und Blattflöhe aus Ost-Thüringen. – Ent. Mitt. Zool. Staatsinst.
Zool. Mus. Hamburg 1(11): 1-36.
Niedringhaus, R. (1989): Die von F. und R. Struve von 1932 bis 1938 auf Borkum gesammelten
Zikaden (Hemiptera: Auchenorrhyncha). – Natur u. Heimat 49 (3): 81-90.
Niedringhaus, R. (1991): Analyse isolierter Artengemeinschaften am Beispiel der Zikadenfauna
der ostfriesischen Düneninseln (Hemiptera: Auchenorrhyncha). – Thesis, University of Old-
enburg. 153 pp.
References 413
Orosz, A. (1977): Beiträge zur Kenntnis der Gattung Ulopa Fallén, 1814 (Homoptera: Ulopidae).
– Fol. ent. Hungarica, Ser. nov. 30(2): 95-103.
Orosz, A. (1983): The Homoptera of the Hortobagy National Park. – Natural History of the
National Parks of Hungary 2: 133-137.
Orosz, A. (1996): Data to the knowledge of the Cicadomorpha species (Homoptera: Auchenorrhyn-
cha) in the Bükk National Park. – Natural History of the National Parks of Hungary 8(2): 151-157.
Orosz, A. (1999): New Auchenorrhyncha species from Hungary (Homoptera). – Fol. ent. hung.
60: 153-164.
Ossiannilsson, F. (1944): Contributions to the knowledge of Swedish Cicadina. II., with descrip-
tion of a new species. – Opusc. ent. 9: 14-16.
Ossiannilsson, F. (1946): Halvvingar. Hemiptera. Stritar Homoptera Auchenorrhyncha. – Svensk
insektfauna 7: 1-150.
Ossiannilsson, F. (1947): Halvvingar. Hemiptera. Stritar Homoptera Auchenorrhyncha. – Svensk
insektfauna 7: 151-270.
Ossiannilsson, F. (1949): Insect drummers. – Opusc. entomol., Suppl. 10: 1-146.
Ossiannilsson, F. (1950): On the identity of Cicada spumaria Linnaeus (1758) (Hem. Hom.) with
notes on the breeding-plants of three Swedish Cercopids. – Opusc. ent. 15: 145-156.
Ossiannilsson, F. (1951): On the shape of the apodemes of the second abdominal sternum of the
males as a specific character in the genus Macrosteles Fieb. (Hom. Auchenorrhyncha). – Opusc.
ent. 16: 109-111.
Ossiannilsson, F. (1978): The Auchenorrhyncha (Homoptera) of Fennoscandia and Denmark.
Part 1: Introduction, infraorder Fulgoromorpha. – Scandinavian Science Press, Kopenhagen.
pp. 1-222.
Ossiannilsson, F. (1981): The Auchenorrhyncha (Homoptera) of Fennoscandia and Denmark.
Part 2: The Families Cicadidae, Cercopidae, Membracidae, and Cicadellidae (excl. Deltoceph-
alinae. – Scandinavian Science Press, Kopenhagen. pp. 223-593.
Ossiannilsson, F. (1983): The Auchenorrhyncha (Homoptera) of Fennoscandia and Denmark.
Part 3: The Family Cicadellidae: Deltocephalinae, Catalogue, Literature and Index. – Scandi-
navian Science Press, Kopenhagen. pp. 594-979.
Ottosson, J.G., Anderson, J.M. (1983): Number, seasonality and feeding habits of insects attack-
ing ferns in Britain: an ecological consideration. – J. Anim. Ecol. 52: 385-406.
Payne, K. (1981): The life history and host-plant relationships of Eupteryx notata Curtis (Ho-
moptera: Cicadellidae). – Entomol. mon. Mag. 117: 167-173.
Pedgley, D.E., Reynolds, D.R., Tatchell, G.M. (1995): Long-range insect migration in relation to
climate and weather: Africa and Europe. – In: Drake, V.A., Gatehouse, A.G. (eds.): Insect migra-
tion: tracking resources through space and time. University Press, Cambridge, pp. 3-29.
Perner, J. (1997): Zur Arthropodenfauna der Kalktrockenrasen im Mittleren Saaletal (Ostthürin-
gen). Teil 1: Coleoptera, Diptera, Auchenorrhyncha, Saltatoria, Araneae (Insecta et Arachni-
da). – Faun. Abh. Staatl. Mus. Tierkde. Dresden 21(3): 53-90.
Peter, H.-U. (1981): Weitere Untersuchungen zur Einnischung der Zikaden in den Halbtrocken-
rasen des Leutratals bei Jena. – Zool. Jb. Syst. 108: 563-588.
Peter, H.-U., Roth, S. (1996): Beitrag zur Kenntnis der Arthropodenfauna der Kiefernforste im
NSG “Uhlstädter Heide” bei Rudolstadt/Thüringen. 1. Das Arteninventar der Untersuchungs-
flächen. – Beitr. Ökol. 1: 83-110.
References 415
Petersen, A. (1992): Die Gräser als Kulturpflanzen und Unkräuter auf Wiese, Weide und Acker.
7. Aufl. – Akademie Verlag, Berlin. 275 pp.
Pianka, E.R. (1970): On r- and K-selection. – Am. Nat. 104: 592-597.
Pollard, D.G. (1968): Stylet penetration and feeding damage of Eupteryx melissae Curtis (Hemi-
ptera, Cicadellidae) on sage. – Bull. ent. Res. 58: 55-71.
Pollard, D.G. (1969): Directional control of the stylets in phytophagous Hemiptera. – Proc. R.
ent. Soc. London (A) 44: 173-185.
Post-Plangg, N., Hoffmann, H.-J. (1982): Ökologische Untersuchungen an der Zikadenfauna
des Bausenbergs in der Eifel – mit besonderer Berücksichtigung der Trockenrasen. – Deche-
niana, Beih. 27: 184-240.
Prestidge, R.A. (1982): Instar duration, adult consumption, oviposition and nitrogen utilization
of leafhoppers feeding on different quality food (Auchenorrhyncha: Homoptera). – Ecol.
Entomol. 7: 91-101.
Prestidge, R.A., McNeill, S. (1983a): Auchenorrhyncha – host plant interactions: leafhoppers
and grasses. – Ecol. Ent. 8: 331-339.
Prestidge, R.A., McNeill, S. (1983b): The role of nitrogen in the ecology of grassland Auchenor-
rhyncha. – In: Lee, J.A., McNeill, S., Rorison, I.H. (eds.): Nitrogen as an ecological factor.
Blackwell, Oxford. pp. 257-281.
Price, P.W. (1983): Hypothesis on organization and evolution in herbivorous insect communi-
ties. – In: Denno, R.F., McClure, M.S. (eds.): Variable plants and herbivores in natural and
managed systems. Academic Press, New York. pp. 559-596.
Price, P.W., Bouton, C.E. Gross, P., McPheron, B.A., Thompson, J.N., Weis, A.E. (1980): Interac-
tions among three trophic levels: Influence of plants on interactions between insect herbi-
vores and natural enemies. – Annu. Rev. Ecol. Syst. 11: 41-65.
Prohaska, K. (1923): Beitrag zur Kenntnis der Hemipteren Kärntens. – Carinthia II, 113/33: 32-101.
Puton, D.A. (1886): Catalogue des Hémiptères (Héteroptères, Cicadines et Psyllides) de la faune
Paléarctique. 3e Ed. – Caen. 100 pp.
Puton, D.A. (1899): Catalogue des Hémiptères (Héteroptères, Cicadines et Psyllides) de la faune
Paléarctique. 4e Ed. – Rev. Ent. 18(3/4): 3-121.
Raatikainen, M. (1960): The biology of Calligypona sordidula (Stål) (Hom., Auchenorrhyncha) –
Ann. ent. Fenn. 26: 229-242.
Raatikainen, M. (1967): Bionomics, enemies and population dynamics of Javesella pellucida (F.)
(Homoptera, Delphacidae). – Ann. Agr. Fenn. 6, Suppl. 2: 1-147.
Raatikainen, M. (1970a): Ecology and fluctuations in abundance of Megadelphax sordidulus (Stål)
(Hom., Delphacidae). – Ann. Agr. Fenn. 9: 315-324.
Raatikainen, M. (1970b): Mesopolobus graminum (Hårdh) (Hym., Pteromalidae), its population
dynamics and influence on Javesella pellucida (F.). – Ann. Agr. Fenn. 9: 99-106.
Raatikainen, M. (1971): Seasonal aspects of leafhopper (Hom., Auchenorrhyncha) fauna in oats.
– Ann. Agr. Fenn. 10: 1-8.
Raatikainen, M. (1972): Dispersal of leafhoppers and their enemies to oat fields. – Ann. Agr.
Fenn. 11: 146-153.
Raatikainen, M., Vasarainen, A. (1970): Frog-Hoppers (Hom., Cercopidae) in strawberry planta-
tions. – Ann. Agr. Fenn. 9: 290-292.
Raatikainen, M., Vasarainen, A. (1971): Comparison of leafhopper faunae in cereals. – Ann. Agr.
Fenn. 10: 119-124.
416 References
Raatikainen, M., Vasarainen, A. (1973): Early- and high-summer flight periods of leafhoppers. –
Ann. Agr. Fenn. 12: 77-94.
Rabeler, W. (1951): Biozönotische Untersuchungen im hannoverschen Kiefernforst. – Z. angew.
Ent. 32: 591-598.
Rabeler, W. (1952): Zur Kenntnis der montanen Tierwelt des Harzes. – Beitr. Naturk. Nieders. 5:
47-51.
Rabeler, W. (1957): Die Tiergesellschaft eines Eichen-Birkenwaldes im nordwestdeutschen Alt-
moränengebiet. – Mitt. flor.-soz. Arbeitsgem. (N.F.) 6/7: 297-319.
Rabeler, W. (1962): Die Tiergesellschaften von Laubwäldern (Querco-Fagetea) im oberen und
mittleren Wesergebiet. – Mitt. flor.-soz. Arbeitsgem., N.F. 9: 200-229.
Raine, J. (1960): Life history and behavior of the Bramble Leafhopper, Ribautiana tenerrima (H.-
S.) (Homoptera: Cicadellidae). – Can. Ent. 92: 10-20.
Raunkiaer, C. (1907): Planterigets Livsformer og deres Betydning for Geografien. – Kjöbenhavn,
Kristiania. English translation by Gilbert-Carter, H. (1937): Plant life forms. Claredon Press,
Oxford. 104 pp.
Raven, J.A. (1983): Phytophages of xylem and phloem: a comparison of animal and plant sap-
feeders. – Adv. Ecol. Res. 13: 135-234.
Raw, A. (1998): The third trophic level of plant defence: Neotropical social wasps’ use of odours
of freshly damaged leaves when hunting. – Rev. Bras. Zool. 15: 1075-1092.
Reclaire, A. (1944): Naamlijst der in Nederland en het aangrenzende gebied waargenomen Ci-
caden (Hemiptera-homoptera). – Ent. Ber. 11: 221-256.
Reimer, H. (1992): Beiträge zur Zoogeographie und Ökologie von Zikaden (Homoptera: Auchen-
orrhyncha) in Mittelgebirgen am Beispiel der Rhön. – Thesis, University of Marburg. 252 pp.
Remane, R. (1958): Die Besiedlung von Grünlandflächen verschiedener Herkunft durch Wan-
zen und Zikaden im Weser-Ems-Gebiet. – Z. ang. Ent. 42: 353-400.
Remane, R. (1959): Lebradea calamagrostidis gen. et spec. nov., eine neue Zikade aus Norddeutsch-
land (Homoptera Cicadina, Cicadellidae). – Zool. Anz. 163: 385-391.
Remane, R. (1960): Zur Kenntnis der Gattung Arthaldeus RIBAUT (Homoptera Cicadina, Ci-
cadellidae). – Mitt. Münchner Ent. Ges. 50: 72-82.
Remane, R. (1961a): Revision der Gattung Mocydiopsis RIBAUT (Homoptera Cicadellidae). –
Abh. math.-naturw. Kl. Akad. Wiss. Mainz 4: 3-51.
Remane, R. (1961b): Zur Kenntnis der Verbreitung einiger Zikadenarten (Homoptera Cicadina).
– Nachr.-Bl. bayer. Ent. 10: 111-114.
Remane, R. (1961c): Endria nebulosa (BALL), comb. nov., eine neue nearktische Zikade in Deutsch-
land (Homoptera Cicadina, Jassidae). – Nachr.-bl. Bayr. Ent. 10: 73-98.
Remane, R. (1961d): Die systematische Position von Deltocephalus aurantiacus FOREL (Homoptera
Cicadina, Cicadellidae). – Nachr.-bl. Bayr. Ent. 10: 1-6.
Remane, R. (1962): Einige bemerkenswerte Zikaden-Funde in Nordwest-Deutschland. – Faun.
Mitt. Norddeutschl. 2(2): 23-26.
Remane, R. (1965): Beiträge zur Kenntnis der Gattung Psammotettix HPT. – Zool. Beitr., N.F. 11:
221-245.
Remane, R. (1972): Funde der nordamerikanischen Buckelzirpe Stictocephala (früher Ceresa) bubalus
(F.) (Homoptera Auchenorrhyncha Membracidae) in Südwestdeutschland. – Faun.-ökol. Mitt.
4: 109-111.
References 417
Remane, R. (1987): Zum Artenbestand der Zikaden (Homoptera: Auchenorrhyncha) auf dem
Mainzer Sand. – Mainzer naturw. Arch. 25: 273-349.
Remane, R. (1994): Anmerkungen zum Bestand an Morphospezies der Zygina-flammigera-Gruppe
in Mitteleuropa (Homoptera Auchenorrhyncha Cicadellidae Typhlocybinae). – Marburger
ent. Publ. 2(8): 109-130.
Remane, R. (1995): Zur Verbreitung einiger Zikadenarten in Mitteleuropa, insbesondere in der
BRD (Homoptera Auchenorrhyncha). – Marburger ent. Publ. 2 (9): 71-75.
Remane, R., Achtziger, R., Fröhlich, W., Nickel, H., Witsack, W. (1998): Rote Liste der Zikaden. (Ho-
moptera, Auchenorrhyncha). – In: Binot, M., Bless, R., Boye, P., Gruttke, H., Pretscher, P. (eds.):
Rote Liste gefährdeter Tiere Deutschlands. Schr.-R. Landschaftspfl. Natursch. 55: 243-249.
Remane, R., Asche, M. (1979): Bemerkungen zur Taxonomie, Phylogenie und Verbreitung der
Gattung Conomelus FIEBER 1866 (Homoptera Cicadina Delphacidae), mit einer ergänzenden
Beschreibung von Delphacellus putoni (SCOTT 1874). – Marburger ent. Publ. 1(1): 1-132.
Remane, R., Fröhlich, W. (1991): Über Vorkommen der Zikade Cicadella lasiocarpae OSS., 1981
(Homoptera: Auchenorrhyncha, Cicadellidae) in der BRD. – Hess. faun. Briefe 11 (2): 25-31.
Remane, R., Fröhlich, W. (1994a): Vorläufige, kritische Artenliste der im Gebiet der Bundesre-
publik Deutschland nachgewiesenen Taxa der Insekten-Gruppe der Zikaden (Homoptera
Auchenorrhyncha). – Marburger ent. Publ. 2(8): 189-232.
Remane, R., Fröhlich, W. (1994b): Beiträge zur Chorologie einiger Zikaden-Arten (Homoptera
Auchenorrhyncha) in der Westpaläarktis. – Marburger ent. Publ. 2(8): 131-188.
Remane, R., Fründ, E. (1986): Beiträge zur Determinierbarkeit der Weibchen mittel- und west-
europäischer Arten der Zikaden-Gattung Cicadula ZETT. s.str. (Homoptera Auchenorrhyn-
cha Cicadellidae). – Marburger ent. Publ. 2(3): 155-178.
Remane, R., Giustina, W. della (1991): La faune de France des Delphacidae (Homoptera, Auchen-
orrhyncha). I – Récoltes d‘aout 1989. – Cahiers Natur. 47: 33-43.
Remane, R., Giustina, W. della (1993): La faune de France des Delphacidae (Homoptera, Auchen-
orrhyncha). IV. – Récoltes de 1992. – Cahiers Natur. 48: 11-24.
Remane, R., Holzinger, W. (1995): Zygina hypermaculata nov. spec., eine neue Zwergzikade aus
dem Ostalpenraum (Homoptera, Auchenorrhyncha: Cicadellidae). – Carinthia II 185/105:
713-721.
Remane, R., Jung, R. (1995): Beiträge zum Artenbestand der europäischen Kelisiinen (Auchen-
orrhyncha, Fulgoromorpha, Delphacidae). – Marburger ent. Publ. 2(9): 1-70.
Remane, R., Reimer, H. (1989): Im NSG “Rotes Moor” durch Wanzen (Heteroptera) und Zikaden
(Homoptera, Auchenorrhyncha) genutzte und ungenutzte “ökologische Lizenzen” im Ver-
gleich zu anderen Mooren und der übrigen Rhön. – Telma, Beih. 2: 149-172.
Remane, R., Wachmann, E. (1993): Zikaden: kennenlernen – beobachten. – Naturbuch Verlag,
Augsburg. 288 pp.
Renvoize, S.A., Clayton, W.D. (1992): Classification and evolution of the grasses. – In: Chap-
man, G.P. (ed.): Grass evolution and domestication. Cambridge University Press, Cambridge.
pp. 3-37.
Reuter, O.M. (1909): Charakteristik und Entwicklungsgeschichte der Hemipteren-Fauna (Het-
eroptera, Auchenorrhyncha und Psyllidae) der palaearktischen Coniferen. – Acta Soc. Sci.
Fenn. 36(1): 1-129.
Rey, C. (1891): Observations sur quelques Hémiptères-homoptères et descriptions d‘espèces
nouvelles ou peu connues. – Revue Ent. 10: 240-256.
418 References
Rhoades, D.F., Cates, R.G. (1976): Towards a general theory of plant antiherbivore chemistry. –
Rec. Adv. Phytochem. 10: 168-213.
Ribaut, H. (1931): Les espèces Francaises des groupes Erythroneura parvula (BOH.) et Erythroneura
fasciaticollis (REY) (Homoptera-Typhlocybidae). – Bull. Soc. Hist. Nat. Toulouse 62: 399-416.
Ribaut, H. (1934): Nouveaux Delphacides (Homoptera – Fulgoroidea). – Bull. Soc. Hist. Nat.
Toulouse 66: 281-301.
Ribaut, H. (1936): Homoptères Auchenorhynques (I. Typhlocybidae). – Faune de France 31, Par-
is. 231 pp.
Ribaut, H. (1939): Nouveaux genres et nouvelles espèces de la famille des Jassidae (Homoptera).
– Bull. Soc. Hist. Nat. Toulouse 73: 267-279.
Ribaut, H. (1952): Homoptères Auchenorhynques. II (Jassidae). – Faune de France 57, Paris. 474 pp.
Ribaut, H. (1959): Homoptères nouveaux pour la France. – Bull. Soc. Hist. Nat. Toulouse 94:
393-399.
Risebrow, A., Dixon, A.F.G. (1987): Nutritional ecology of phloem-feeding insects. – In: Slansky,
F., Rodriguez, J.G. (eds.): Nutritional ecology of insects, mites, spiders, and related inverte-
brates. Wiley, New York. pp. 421-448.
Roff, D.A. (1990): The evolution of flightlessness in insects. – Ecol. Monogr. 60(4): 389-421.
Röhrig, E. (1996): Die Ulmen in Europa. Ökologie und epidemische Erkrankung. – Forstarchiv
67: 179-198.
Rombach, R. (1999a): Zoogeographisch bemerkenswerte Nachweise einiger Zikadenarten (Ho-
moptera, Auchenorrhyncha) für die Nordeifel. – Marburger Ent. Publ. 3(1): 1-6.
Rombach, R. (1999b): Auswirkungen verschiedener Formen der Bewirtschaftung von Halbtrock-
enrasen auf die Zikaden (Homoptera, Auchenorrhyncha) am Beispiel der Enzian-Schiller-
gras-Rasen (Gentiano-Koelerietum) der Nordeifel (Nordrhein-Westfalen). – Thesis, Universi-
ty of Bonn. 346 pp.
Root, R.B. (1967): The niche exploitation pattern of the blue-gray gnatcatcher. – Ecol. Monogr.
37: 317-350.
Rosenthal, G.A., Janzen, D.H. (eds.) (1979): Herbivores. Their interaction with secondary plant
metabolites. – Academic Press, New York. 718 pp.
Ross, H. (1957): Principles of natural coexistance indicated by leafhopper populations. – Evolu-
tion 11: 113-129.
Ross, H.H. (1958): Further comments on niches and natural coexistence. – Evolution 12: 112-113.
Rothschild, G.H.L. (1964): The biology of Conomelus anceps (Germar) (Homoptera: Delphaci-
dae). – Trans. Soc. Br. Ent. 16: 135-148.
Rothschild, G.H.L. (1966): A study of a natural population of Conomelus anceps (Germar) (Ho-
moptera: Delphacidae) including observations on predation using the precipitin test. – J. Anim.
Ecol. 35: 413-434.
Sabelis, M.W., Baalen, M. van, Bakker, F.M., Bruin, J., Drukker, B., Egas, M., Janssen, A.R.M.,
Lesna, I.K., Pels, B., Rijn, P.C.J. van, Scutareanu, P. (1999): The evolution of direct and indirect
plant defence against herbivorous arthropods. – In: Olff, H., Brown, V.K., Drent, R.H. (eds.):
Herbivores: Between plants and predators. Blackwell, Oxford. pp. 109-166.
Sahlberg, J. (1871): Öfversigt af Finlands och den Skandinaviska halföns Cicadariae. – Notis.
Sällsk. Faun. Fl. fenn. Förh., n.s. 9(12): 1-506.
References 419
Saikkonen, K., Helander, M., Faeth, S.H., Schulthess, F., Wilson, D. (1999): Endophyte-grass her-
bivore-interactions: the case of Neotyphodium endophytes in Arizona fescue populations. –
Oecologia 212: 411-420.
Sander, F.W. (1985): Zikadenfeinde in Rasengesellschaften der DDR: Augenfliegen (Diptera,
Pipunculidae) – Bemerkungen zu Entwicklung, Verhalten und Wirtsbeziehungen. – Wiss. Z.
Friedrich-Schiller-Univ. Jena. Naturwiss. R. 34: 609-624.
Sander, F.W., Achtziger, R., Lauterer, P., Funke, T., Holzinger, W.E., Kammerlander, I., Nickel, H.,
Orosz, A., Witsack, W. (1999): Beitrag zur Kenntnis der Zikaden- und Blattflohfauna (Auchen-
orrhyncha und Psylloidea) der Oberlausitz unter besonderer Berücksichtigung des Natur-
schutzgebietes “Niederspree”. – Beitr. Zikadenkde. 3: 81- 89.
Schaefer, M. (1973): Untersuchungen über Habitatbindung und ökologische Isolation der Zikaden
einer Küstenlandschaft (Homoptera: Auchenorrhyncha). – Arch. Natursch. Landschaftsforsch.
13: 329-352.
Schaefer, M. (1981): The arthropod fauna of high salt marshes. – In: Smit, C.S., den Hollander, J.,
van Wingerden, W.K.R.E., Wolff, W.J. (eds.): Terrestrial and freshwater fauna of the Wadden
Sea area. pp. 57-67. Stichting Veth tot Steun aan Waddenonderzoek, Leiden.
Schaefer, M. (1992): Wörterbuch der Ökologie. 3. Aufl. – Gustav Fischer, Jena. 433 pp.
Schedl, W. (1995): Einwanderung der Amerikanischen Büffelzikade (Stictocephala bisonia Kopp
& Yonke, 1977) nach Österreich. 2. Beitrag. (Homoptera: Auchenorrhyncha, Membracidae). –
Stapfia 37: 149-152.
Schedl, W. (1998): Die Verbreitung und Biologie von Gargara genistae (Fabricius 1775) in Öster-
reich (Homoptera: Auchenorrhyncha: Membracidae). – Stapfia 55: 607-612.
Schedl, W. (2000): Taxonomie, Biologie und Verbreitung der Singzikaden Mitteleuropas (Insec-
ta: Homoptera: Cicadidae et Tibicinidae). – Ber. nat.-med. Ver. Innsbruck 87: 257-271.
Schiemenz, H. (1964): Zikaden (Homoptera Auchenorrhyncha) von einer Rohbodenkippe des
Braunkohlentagebaues Böhlen. – Abh. Ber. Naturk.-Mus. Görlitz 39 (16): 1-8.
Schiemenz, H. (1969): Die Zikadenfauna mitteleuropäischer Trockenrasen (Homoptera, Auchen-
orrhyncha). Untersuchungen zu ihrer Phänologie, Ökologie, Bionomie und Chorologie. –
Ent. Abh. staatl. Mus. Tierk. Dresden 36: 201-280.
Schiemenz, H. (1970): Beiträge zur Insektenfauna der DDR: Verzeichnis (check list) der im Ge-
biet der Deutschen Demokratischen Republik bisher festgestellten Zikaden (Homoptera:
Auchenorrhyncha). – Beitr. Ent. 20: 481-502.
Schiemenz, H. (1971a): Die Zikadenfauna (Homoptera Auchenorrhyncha) der Erzgebirgshoch-
moore. – Zool. Jb. Syst. 98: 397-417.
Schiemenz, H. (1971b): Zur Zikaden- und Heuschreckenfauna des Naturschutzgebietes “Stein-
holz – Harsleber Berge”. – Naturk. Jber. Mus. Heineanum 5/6: 47-52.
Schiemenz, H. (1973): Zur Zikadenfauna (Homoptera Auchenorrhyncha) der Naturschutzge-
biete “Steppenheide am Großen Seeberg”, “Schwellenburg”, “Alperstedter Ried” und “Ves-
sertal”. – Abh. Ber. Mus. Nat. Gotha 1973: 70-79.
Schiemenz, H. (1975): Die Zikadenfauna der Hochmoore im Thüringer Wald und im Harz (Ho-
moptera, Auchenorrhyncha). – Faun. Abh. staatl. Mus. Tierk. Dresden 5: 215-233.
Schiemenz, H. (1976): Die Zikadenfauna von Heide- und Hochmooren des Flachlandes der DDR
(Homoptera, Auchenorrhyncha). – Faun. Abh. Staatl. Mus. Tierk. Dresden 6: 39-54.
420 References
Sequeira, A.S., Farrell, B.D. (2001): Evolutionary origins of Gondwanan interactions: How old
are Araucaria beetle herbivores? – Biol. J. Linn. Soc. 74: 459-474.
Sergel, R. (1984a): Ein weiterer Nachweis der Cixiide Hyalesthes obsoletus SIGNORET in Fran-
ken. – Abh. Naturw. Ver. Würzburg 25: 81-82.
Sergel, R. (1984b): Der Europäische Laternenträger in Unterfranken. – Abh. Naturw. Ver.
Würzburg 25: 72-80.
Sergel, R. (1986): Hyalesthes obsoletus SIGNORET in Germany (Homoptera: Fulogoroidea: Cixi-
idae). – Biol. Z. 1(1): 90-93.
Sergel, R. (1987): On the occurence of the Rhododendron-leafhopper, Graphocephala fennahi
YOUNG 1977, in the Western Palaearctic region (Homoptera, Cicadellidae). – Anz. Schädling-
skde., Pflanzensch., Umweltsch. 60: 134-136.
Settle, H.W., Wilson, L.T. (1990): Invasion by the variegated leafhopper and biotic interactions:
parasitism, competition, and apparent competition. – Ecology 71: 1461-1470.
Sforza, R., Bourgoin, T., Wilson, S.W., Boudon, P.E. (1999): Field observations, laboratory rearing
and descriptions of immatures of the planthopper Hyalesthes obsoletus (Hemiptera: Cixiidae).
– Eur. J. Ent. 96(4): 409-418.
Shear, W.A. (1991): The early development of terrestrial ecosystems. – Nature 351: 283-289.
Simberloff, D., Dayan, T. (1991): The guild concept and the structure of ecological communities.
– Annu. Rev. Ecol. Syst. 22: 115-143.
Simoes de Andrade, G. (1997): Stictocephala alta (Walker, 1851) sp. rev., comb. n., the correct name
for the “buffalo treehopper”, with S. bisonia Kopp & Yonke, 1977 as a new synonym, and
notes on Hadrophallus bubalus (Fabricius, 1794) comb. n. (Homoptera: Membracidae). – Trans.
Am. Entomol. Soc. 123: 289-295.
Smreczynski. S. (1954): Materialy do fauny pluskwiakow (Hemiptera) Polski. – Fragm. faun. 7:
1-146.
Snodgrass, R.E. (1958): Evolution of arthropod mechanisms. – Smithonian misc. Coll. 138(2): 1-77.
Sorensen, J.T., Campbell, B.C., Gill, R.J., Steffen-Campbell, J.D. (1995): Non-homophyly of Auchen-
orrhyncha (“Homoptera”), based upon 18S rDNA phylogeny: eco-evolutionary and cladistic
implications within Pre-Heteropterodea Hemiptera (s.l.) and a proposal for new monophyl-
etic suborders. – Pan-Pacific Entomologist 71(1): 31-60.
Southwood, T.R.E., Moran, V.C., Kennedy, C.E.J. (1982): The assessment of arboreal insect fauna:
comparisons of knockdown sampling and faunal lists. – Ecol. Entomol. 7: 331-340.
Southwood, T.R.E., Moran, V.C., Kennedy, C.E.J. (1982): The richness, abundance and biomass
of the arthropod communities on trees. – J. Anim. Ecol. 51: 635-649.
Stadler, H. (1922): Tettigonia haematodes in Unterfranken. – Arch. Naturgesch. 88 (10): 160.
Stein, B., Bogon, K. (1990): Zum Vorkommen der Bergzikade, Cicadetta montana (Insecta: Homoptera)
in Nordhessen/Südniedersachsen und Westthüringen. – Göttinger naturk. Schr. 2: 65-72.
Stewart, A. (1986a): Descriptions and key to the nymphs of Eupteryx (Curtis) leafhoppers (Ho-
moptera: Cicadellidae) occurring in Britain. – Syst. Entomol. 11: 365-376.
Stewart, A. (1986b): Nymphal colour/pattern polymorphism in the leafhoppers Eupteryx urticae
(F.) and Eupteryx cyclops Matsumura (Hemiptera: Auchenorrhyncha): spatial and temporal
variation in morph frequencies. – Biol. J. Linn. Soc. 27: 79-101.
Stewart, A. (1988): Patterns of host-plant utilization by leafhoppers in the genus Eupteryx (Hemi-
ptera: Cicadellidae) in Britain. – J. Nat. Hist. 22: 357-379.
422 References
Then, F (1896): Neue Arten der Cicadinen-Gattungen Deltocephalus und Thamnotettix. – Mitt.
Naturwiss. Ver. Steiermark 32: 165-197.
Then, F (1897): Fünf Cicadinen-Species aus Österreich. – Mitt. Naturwiss. Ver. Steiermark 33:
102-116.
Thüs, H. (1996): Zikaden der Weiden um Balingen-Zillhausen (Schwäbische Alb). – Naturkdl.
Beitr. DJN 31: 36-51.
Tingey, W.M. (1985): Plant defensive mechanisms against leafhoppers. – In: Nault, L.R., Rod-
riguez, J.G. (eds.): The leafhoppers and planthoppers. Wiley, New York. pp. 217-234.
Tishechkin, D.Y. (1993): Review of Macropsis species (Homoptera, Cicadellidae, Macropsinae)
feeding on Hippophae rhamnoides with description of new species. – Zool. Zh. 72(2): 54-60. (In
Russian).
Tishechkin, D.Y. (1998): Acoustic signals and morphological characters of leafhoppers from Aph-
rodes bicinctus group from central European Russia. – Zool. Zh. 77(6): 669-676 (In Russian).
Tishechkin, D.Yu. (2000): On taxonomic status of Cicadella lasiocarpae (Homoptera, Cicadellidae).
– Zool. Zh. 79(7): 863-867. (In Russian).
Tonkyn, D.W., Whitcomb, R.F. (1987): Feeding strategies and the guild concept among vascular
feeding insects and microorganisms. – Curr. top. vector res. 4: 179-199.
Torka, V. (1905): Tettigometra obliqua Panz. – Zeitschrift für wissenschaftliche Insektenbiologie 1:
451-455.
Törmälä, T. (1982): Evaluation of five methods of sampling field layer arthropods, particularly
the leafhopper community, in grassland. – Ann ent. Fenn. 48: 1-16.
Tromellini, C., Alma, A., Vidano, C. (1988): Biotaxonomy and ecology of Rhytidodus decimusquar-
tus in Piedmont, Italy (Rhynchota Auchenorrhyncha). – Proc. 6th Auchenorrhyncha Meeting,
Turin, Italy, September 7-11, 1987, pp. 259-266.
Truman, J.W., Riddiford, L.M. (1999): The origins of insect metamorphosis. – Nature 401: 447-452.
Trümbach, H. (1959): Die Zikaden und Psylliden der Umgebung Erlangens, eine systematisch-
ökologische Untersuchung. – Sitz.-ber. phys.-med. Soz. Erlangen 79: 102-151.
Tscharntke, T. (1999): Insects on common reed (Phragmites australis): Community structure and
the impact of herbivory on shoot growth. – Aquatic Botany 64: 399-410.
Tscharntke, T., Greiler, H.-J. (1995): Insect communities, grasses, and grasslands. – Annu. Rev.
Entomol. 40: 535-558.
Tulowitzki, I. (1990): Einfluß der Beweidung auf die Populationsstruktur der Kleinzikade Psam-
motettix putoni (Hom. Auch.) in der Salzwiese der schleswig-holsteinischen Westküste. – Verh.
Ges. Ökol. 19(2): 152-162.
Valle, R.R., Kuno, E., Nakasuji, F. (1989): Competition between laboratory populations of green
leafhoppers, Nephotettix spp. (Homoptera: Cicadellidae). – Res. Popul. Ecol. 31: 53-72.
Vera, F.W.M. (2000): Grazing ecology and forest history. – CABI, Oxon. 506 pp.
Vidano, C. (1959b): Sulla identificazione specifica di alcuni Erythroneurini europei (Hemiptera, Typhlo-
cybidae). – Estratto dagli Annali del Museo Civico di Storia Naturale di Genova 71: 328-348.
Vidano, C. (1961): L’influenza microclimatica sui caratteri tassonomici in Tiflocibidi sperimen-
talmente saggiati – Memorie dalle Societa Entomologica Italiana 40: 144-167.
Vidano, C. (1962): La Empoasca lybica Bergevin nuovo nemico della Vite in Italia – L’Italia agrico-
la 4: 329-346.
Vidano, C. (1963a): Deviazione trofica ampelofila della Ceresa bubalus Fabricius e rispondenza
reattiva del vegetale. – Atti Acc. Sci. Torino 98: 193-212.
424 References
Vidano, C. (1963b): Eccezionali strozzature anulari caulinari provocate da Ceresa bubalus Fabri-
cius in Vitis. – Annali Fac. Sci. Agr. Univ. Stud. Torino 2: 57-108.
Vidano, C. (1963c): Alterazioni provocate da insetti in Vitis osservate, sperimentate e comparate.
– Annali Fac. Sci. Agr. Univ. Stud. Torino 1: 513-644.
Vidano, C. (1965): A contribution to the chorological and oecological knowledge of the Europe-
an Dikraneurini (Homoptera Auchenorrhyncha). – Zool. Beitr. (N.F.) 11: 343-367.
Vidano, C. (1967): Sintomatologia esterna ed interna da insetti fitomizi su Vitis – Annali della
Accademia di Agricoltura di Torino 109: 117-136.
Vidano, C., Arzone, A. (1978): Typhlocybinae on officinal plants. – Auchenorrhyncha newsletter
1: 27-28.
Vidano, C., Arzone, A. (1985): Zyginidia pullula: distribuzione nel territorio e ciclo biologico. –
REDIA 68: 135-150.
Vidano, C., Arzone, A. (1987a): Typhlocybinae of broad-leaved trees and shrubs in Italy. 2. Betu-
laceae. – Estratt. Bollet. Inst. Ent. “Guido Grandi” 16: 257-267.
Vidano, C., Arzone, A. (1987b): Typhlocybinae of broad-leaved trees and shrubs in Italy. 4. Fa-
gaceae. – REDIA 70: 171-189.
Vidano, C., Arzone, A., Meotto, F. (1987): Dati morfologici, biologici e fitopatologici su Grapho-
cephala fennahi (Homoptera: Auchenorrhyncha) nuovo fitomizo di Rhododendron spp. in Italia.
– La difesa delle piante 10(1): 101-112.
Vidano, C., Bosco, D., Arzone, A. (1990): Dioecia obligata in Lindbergina (Homoptera Auchenor-
rhyncha Cicadellidae). – REDIA 73: 293-306.
Vilbaste, J. (1960): Eine Revision der Sammlung von G. Flor. 1. Homoptera: Cicadina: Fulgoroi-
dea. – Eest. NSV Tead. Akad. Toim. Biol. 9 (2): 135-143. (In Russian).
Vilbaste, J. (1961): Neue Zikaden (Homoptera: Cicadina) aus der Umgebung von Astrachan. –
Eesti NSV Tead. Akad. Toimet. 10: 315-331. (In Russian).
Vilbaste, J. (1962): Über die Arten Rhopalopyx preyssleri (H.-S.) und Rh. adumbrata (C.R. Sahlb.)
(Homoptera, Iassidae). – Notul. Ent. 42: 62-66.
Vilbaste, J. (1965): Über die Zikadenfauna Altais. – Akademia Nauk. Tartu. 144 pp. (In Russian).
Vilbaste, J. (1968a): Preliminary key for the identification of the nymphs of North European
Homoptera Cicadina. I. Delphacidae. – Ann. ent. Fenn. 34: 65-74.
Vilbaste, J. (1968b): K faune tsikadovykh Primorskogo kraia. – Valgus, Tallinn. 179 pp.
Vilbaste, J. (1971): Eesti tirdid I. – Valgus, Tallinn. 283 pp.
Vilbaste, J. (1972): Kelisia nervosa n.sp. – a new Delphacid species from Lithuania. – Eesti NSV
Tead. Akad. Toimet. 21: 75-77. (In Russian).
Vilbaste, J. (1973): Revision of the collection of G. Flor. – Eesti NSV Tead. Akad. Toimet. 22: 15-28.
Vilbaste, J. (1974): Preliminary list of Homoptera – Cicadinea of Latvia and Lithuania. – Eesti
NSV Tead. Akad. Toimet. 23: 131-163.
Vilbaste, J. (1975): Zur Larvalsystematik der Zikaden. – Verh. Vl. Int. Symp. Entomofaun. Mit-
teleuropa 343-346.
Vilbaste, J. (1976): A Revision of Homoptera – Cicadinea described by S. Matsumura from Eu-
rope and the Mediterranean area. – Eesti NSV Tead. Akad. Toimet. 25: 25-36.
Vilbaste, J. (1980): On the Homoptera – Cicadinea of Kamchatka. – Ann. zool. Warsz. 35: 367-
418.
Vilbaste, J. (1982): Preliminary key for the identification of the nymphs of North European Ho-
moptera Cicadinea. II. Cicadelloidea. – Annls. zool. Fenn. 19: 1-20.
References 425
Wagner, W. (1947b): Stenocranus major KB. in der Umgebung Hamburgs. – Bombus 1(42): 183.
Wagner, W. (1948a): Neue deutsche Homopteren und Bemerkungen über schon bekannte Arten.
– Verh. naturw. Heimatforsch. Hamburg 29: 72-89.
Wagner, W. (1948b): Zikaden und Blattflöhe. Der Stand ihrer faunistischen Erforschung in Nied-
ersachsen. – Beitr. Naturkde. Niedersachsen 1(4): 18-25.
Wagner, W. (1948c): Aphrophora salicina Goeze und forneri Haupt. – Bombus 48: 208-210.
Wagner, W. (1949a): Die deutschen Arten der Gattung Oncopsis Burmeister (Hemipt. Homopt.).
– Verh. Ver. naturw. Heimatforsch. Hamburg 30: 1-25.
Wagner, W. (1949b): Drei neue Typhlocybiden aus der Steiermark. – Zentbl. Ges. Ent. 3: 43-45.
Wagner, W. (1950a): Balclutha boica n. sp., eine neue Jasside aus Bayern. – Ber. naturforsch. Ges.
Augsburg 3: 97-100.
Wagner, W. (1950b): Die salicicolen Macropsis-Arten Nord- und Mitteleuropas. – Notul. Ent. 30: 81-114.
Wagner, W. (1951a): Verzeichnis der bisher in Unterfranken gefundenen Zikaden (Homoptera
Auchenorrhyncha). – Nachr. naturw. Mus. Stadt Aschaffenburg 33: 1-54.
Wagner, W. (1951b): Beitrag zur Phylogenie und Systematik der Cicadellidae (Jassidae) Nord-
und Mitteleuropas. – Comment. Biol. Helsinki 12(2): 1-44.
Wagner, W. (1952a): Bemerkungen zur Zikadenfauna des nördlichen Westdeutschlands. – Faun.
Mitt. Norddeutschl. 2: 2-4.
Wagner, W. (1952b): Eine neue Rhopalopyx-Art aus Spanien (Homoptera – Auchenorrhyncha). –
Eos 28: 83.84.
Wagner, W. (1953): Eine neue Macropsis-Art (Hemiptera Homoptera) aus den Niederlanden. –
Ent. Ber. 14: 232-234.
Wagner, W. – (1954): Die Fulgoroidea der Omer-Cooper-Expedition in die Lybische Wüste. –
Bull. Soc. Fouad Ier Ent. 38: 211-224.
Wagner, W. (1955): Neue mitteleuropäische Zikaden und Blattflöhe (Homoptera). – Ent. Mitt.
zool. Staatsinst. zool. Mus. Hamburg 1(6): 163-194.
Wagner, W. (1958): Über eine Zikaden-Ausbeute vom Großen Belchen im Schwarzwald (Ho-
moptera Auchenorrhyncha). – Ent. Mitt. Zool. St.-Inst. Zool. Mus. Hamburg 40: 433-443.
Wagner, W. (1961): Berichtigungen zum Kapitel “Homoptera Auchenorrhyncha (Zikaden)”. –
Die Nordostalpen im Spiegel ihrer Landtierwelt 2: 790. Innsbruck.
Wagner, W. (1963a): Dynamische Taxionomie, angewandt auf die Delphaciden Mitteleuropas. –
Mitt. Hamburg. zool. Mus. Inst. 60: 111-180.
Wagner, W. (1963b): Revision der europäischen Arten dreier Gattungen der Homoptera-Cicadi-
na Dryodurgades Zakhvatkin, Fieberiella Signoret und Phlepsius Fieber. – Ent. Mitt. zool. St.-
Inst. zool. Mus. Hamburg 45: 423-435.
Wagner, W. (1964): Die auf Rosaceen lebenden Macropsis-Arten der Niederlande. – Ent. Ber.
Amsterdam 24: 123-136.
Wagner, W. (1967a): Die Singzikaden (Homoptera, Cicadidae) aus dem Pliozän von Willershausen.
– Ber. Naturhist. Ges. Hannover 111: 91-94.
Wagner, W. (1967b): Taxonomie der Gattung Paluda De Long 1937 (Homoptera Euscelidae). –
Zool. Beitr., N.F. 13: 479-499.
Wagner, W. (1970): Oliarus beieri, eine neue Zikade aus den Alpen. – Ann. Naturhist. Mus. Wien
74: 281-284.
Wagner, W., Franz, H. (1961): Unterordnung Homoptera. Überfamilie Auchenorrhyncha
(Zikaden). – Die Nordostalpen im Spiegel ihrer Landtierwelt 2: 74-158. Innsbruck.
References 427
Wagner, W., Wagner, E. (1938): Hemipterenfunde aus Baden. – Mitt. Bad. Landesver. Naturk.
(N.F.) 3: 402-405.
Wais, A. (1990): Biologie und Ernährung von Zwergzikaden (Homoptera: Auchenorrhyncha) an
Kartoffeln und deren Fähigkeit zur Übertragung des Kartoffel-Y-Virus (PVY). – Thesis, Uni-
versity of Göttingen. 82 pp.
Waloff, N. (1973): Dispersal by flight of leafhoppers. – J. appl. Ecol. 10: 705-730.
Waloff, N. (1975): The parasitoids of the nymphal and adult stages of leafhoppers (Auchenor-
rhyncha: Homoptera) of acidic grassland. – Trans. R. ent. Soc. London 126(4): 637-686.
Waloff, N. (1979): Partitioning of resources by grassland leafhoppers (Auchenorrhyncha, Ho-
moptera). – Ecol. Ent. 4: 379-385.
Waloff, N. (1980): Studies on grassland leafhoppers and their natural enemies. – Adv. Ecol. Res.
11: 81-215.
Waloff, N. (1983): Absence of wing polymorphism in the arboreal, phytophagous species of
some taxa of temperate Hemiptera: an hypothesis. – Ecol. Entomol. 8: 229-232.
Waloff, N., Jervis, M.A. (1987): Communities of parasitoids associated with leafhoppers and
planthoppers in Europe. – Adv. Ecol. Res. 17: 281-402.
Waloff, N., Solomon, M.G. (1973): Leafhoppers of acidic grassland. – J. appl. Ecol. 10: 189-212.
Waloff, N., Thompson, P. (1980): Census data and analysis of populations of some leafhoppers
(Auchenorrhyncha, Homoptera) of acidic grassland. – J. Anim. Ecol. 49: 395-416.
Walter, D.E., Proctor, H.C. (1999): Mites: ecology, evolution and behaviour. – CABI, Wallingford,
Oxon. 322 pp.
Walter, S. (1975): Larvenformen mitteleuropäischer Euscelinen (Homoptera, Auchenorrhyncha).
– Zool. Jb. Syst. 102: 241-302.
Walter, S. (1978): Larvenformen mitteleuropäischer Euscelinen (Homoptera, Auchenorrhyncha).
Teil II. – Zool. Jb. Syst. 105: 102-130.
Walter, S. (1996): Zikaden als Indikatoren für die Bewertung von Landschaftseinheiten. Ein
Beispiel zur Charakterisierung der Drömlingsniederung (Sachsen-Anhalt). – Ber. 2. Auchen-
orrhyncha-Tagung 15.9.-17.9.1995 in Marburg: 15-24.
Walter, S. (1998): Grünlandbewertung mit Hilfe von Zikaden (Homoptera, Auchenorrhyncha):
ein Beispiel aus dem Osterzgebirge. – Beitr. Zikadenkde. 2: 13-38.
Walter, S., Emmrich, R. (1995): Kommentiertes vorläufiges Verzeichnis der Zikaden (Homoptera,
Auchenorrhyncha) im Freistaat Sachsen. – Mitt. Sächs. Ent. 28: 11-23.
Ward, L.K. (1988): The validity and interpretation of insect foodplant records. – Br. J. Ent. Nat.
Hist. 1: 153-162.
Webb, M.D., Vilbaste, J. (1994): Review of the leafhopper genus Balclutha KIRKALDY in the
Oriental Region (Insecta: Homoptera: Cicadellidae). – Ent. Abh. Staatl. Mus. Tierkde. Dres-
den 56(3): 55-87.
Weber, A., Maixner, M. (1998): Survey of populations of the planthopper Hyalesthes obsoletus
Sign. (Auchenorrhyncha, Cixiidae) for infection with the phytoplasma causing grapevine
yellows in Germany. – J. Appl. Ent. 122: 375-381.
Weis, A., Schönitzer, K. (2001): Der Wandel der Zikadenfauna am “Lochhauser Sandberg”, einem
kleinflächigen Reliktbiotop bei München. – Beitr. Zikadenkde. 4: 59-68.
Wheeler, W.C., Whiting, M., Wheeler, Q.D., Carpenter, J.M. (2001): The phylogeny of the extant
hexapod orders. – Cladistics 17: 113-169.
428 References
Whitcomb, R.F., Hicks, A.L., Lynn, D.E. (1987): Geographic variation in host relationships of
leafhoppers (Homoptera: Cicadellidae) in North American grasslands. – In: Wilson, M.R.,
Nault, L.R. (eds.): Proc. 2nd Int. Workshop Leafhoppers Planthoppers Economic Importance,
Provo, Utah, USA, 28th July – 1st Aug. 1986, CIE, London, pp. 293-325.
Whitcomb, R.F., Hicks, A.L., Lynn, D.E. (1988): Host specifity: a major mechanism enhancing
insect diversity in grasslands. – In: Davis, A., Stanford, G. (eds.): Proc. 10th N. Am. Prairie
Conf., Denton, Texas, June 22nd – 26th, 1986. Native Prairies Ass. Texas, Dallas. 6 pp.
Whitcomb, R.F., Kramer, J., Coan M.E., Hicks, A.L. (1987): Ecology and evolution of leafhopper-
grass host relationships in North American grasslands. – Curr. Top. Vector Res. 4: 121-178.
White, T.C.R. (1993): The inadequate environment. Nitrogen and the abundance of animals. –
Springer, Berlin. 425 pp.
Whittaker, J.B. (1965a): The biology of Neophilaenus lineatus (L.) and N. exclamationis (Thunberg)
(Homoptera: Cerco-pidae) on Pennine moorland. – Proc. R. ent. Soc. London (A) 40: 51-60.
Whittaker, J.B. (1965b): The distribution and population dynamics of Neophilaenus lineatus (L.)
and N. exclamationis (Thun.) (Homoptera: Cercopidae) on Pennine moorland. – J. Anim. Ecol.
34: 277-297.
Whittaker, J.B. (1968): Polymorphism of Philaenus spumarius (L.) (Homoptera, Cercopidae) in
England. – J. Anim. Ecol. 37: 99-111.
Whittaker, J.B. (1971): Population changes in Neophilaenus lineatus (L.) (Homoptera: Cercopidae)
in different parts of its range. – J. Anim. Ecol. 40: 425-443.
Whittaker, J.B. (1973): Density regulation in a population of Philaenus spumarius (L.) (Homoptera:
Cercopidae). – J. Anim. Ecol. 42: 163-172.
Whittaker, J.B. (1984): Responses of sycamore (Acer pseudoplatanus) leaves to damage by a typhlo-
cybine leaf hopper, Ossiannilssonola callosa. – J. Ecol. 72: 455-462.
Wiegert, R.G. (1964): Population energetics of meadow spittlebugs (Philaenus spumarius L.) as
affected by migration and habitat. – Ecol. Monogr. 34: 217-241.
Wilson, M.R. (1978): Descriptions and key to the genera of the nymphs of British woodland
Typhlocybinae (Homoptera). – Syst. Entomol. 3: 75-90.
Wilson, M.R. (1981): Identification of European Iassus species (Homoptera: Cicadellidae) with
one species new to Britain. – Syst. Entomol. 6: 115-118.
Wilson, M.R., Asche, M. (1998): Cicada clavicornis Fabricius, 1794 (currently Asiraca clavicornis;
Insecta, Homoptera): proposed conservation of the specific name. Case 3040. – Bull. Zool.
Nomenclature 55(2): 93-95.
Wilson, M.R., Claridge, M.F. (1991): Handbook for the identification of leafhoppers and plant-
hoppers of rice. – CABl, Wallingford. 142 pp.
Wilson, M.R., Claridge, M.F. (1999): Species differentiation in the Edwardsiana lethierryi (Edwards)
species group (Hemiptera: Auchenorrhyncha: Cicadomorpha: Cicadellidae: Typhlocybinae).
– Reichenbachia 33(16): 123-130.
Wilson, S.W. (1988): Delphacidae of Alaska (Homoptera: Fulgoroidea). – Great Basin Naturalist
Memoirs 2: 335-343.
Wilson, S.W. (1992): The Delphacidae of Yukon Territory, Canada (Homoptera: Fulgoroidea). –
Insecta mundi 6: 79-100.
Wilson, S.W., Mitter, C, Denno, R.F., Wilson, M.R. (1994): Evolutionary patterns of host plant use
by delphacid planthoppers and their relatives. – In: Denno, R.F., Perfect, T.J. (eds.): Planthop-
pers: their ecology and management. Chapman & Hall, New York. pp. 7-113.
References 429
Wisskirchen R., Haeupler, H. (1998): Standardliste der Farn- und Blütenpflanzen Deutschlands.
– Ulmer, Stuttgart. 765 pp.
Witsack, W. (1971): Experimentell-ökologische Untersuchungen über Dormanzformen von
Zikaden (Homoptera – Auchenorrhyncha). I. Zur Form und Induktion der Embryonaldor-
manz von Muellerianella brevipennis (Boheman) (Delphacidae). – Zool. Jb. Syst. 98: 316-340.
Witsack, W. (1973): Experimentell-ökologische Untersuchungen über Dormanzformen von
Zikaden (Homoptera – Auchenorrhyncha). II. Zur Ovarial-Parapause und obligatorischen
Embryonal-Diapause von Philaenus spumarius (L.) (Aphrophoridae). – Zool. Jb. Syst. 100:
517-562.
Witsack, W. (1981): Zum weiteren Ausbau des ökologischen Systems der Dormanzformen. –
Zool. Jb. Syst. 108: 502-518.
Witsack, W. (1985): Dormanzformen bei Zikaden (Homoptera Auchenorrhyncha) und ihre öko-
logische Bedeutung. – Zool. Jb. Syst. 112: 71-139.
Witsack, W. (1991): Simultane Embryonaldormanzen bei Euscelis incisus (Kbm.) (Homoptera
Auchenorrhyncha) als populationsökologische Mehrfachabsicherung für das Überleben im
Winter. – Zool. Jb. Syst. 118: 287-307.
Witsack, W. (2002): Dormanzformen mitteleuropäischer Zikaden. – Denisia 4: 471-482.
Wolf, H. (1976): Die Berg-Singzikade in Hessen und in Mitteleuropa. – Jb. nass. Ver. Naturk. 103:
18-23.
Wonn, L. (1956): Ökologische Studien über die Zikadenfauna der Mainzer Sande. – Jb. Nassau.
Ver. Naturk. 92: 81-122.
Wood, T.K. (1987): Host plant shifts and speciation in the Enchenopa binotata Say complex. – Proc.
2nd Int. Workshop on leafhoppers and planthoppers of economic importance, Provo, Utah,
USA, 28th July–1st Aug. 1986, pp. 361-368.
Wood, T.K. (1988): Consequences of membracid life histories mediated by plant phenology. –
Proc. 6th Auchenorrhyncha Meeting, Turin, Italy, September 7-11, 1987, pp. 247-253.
Wood, T.K., Guttman, S.I. (1981): The role of host plants in the speciation of treehoppers: an
example from the Enchenopa binotata complex. – In: Denno, R.F., Dingle, H. (eds.): Insect life
history patterns. Springer, New York. pp. 39-54.
Wood, T.K., Guttman, S.I. (1982): Ecological and behavioral basis for reproductive isolation in
the sympatric Echenopa binotata Say complex (Homoptera: Membracidae). – Evolution 26:
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-
ptera). – Syst. Entomol. 26(1): 1-13.
Young, D.A., Frazier, N.W. (1954): A study of the leafhopper genus Circulifer Zachvatkin (Ho-
moptera, Cicadellidae). – Hilgardia 23: 25-52.
Zabel, J., Tscharntke, T. (1998): Does fragmentation of Urtica habitats affect phytophagous and
predatory insects differentially? – Oecologia 116: 419-425.
Zachvatkin, A.A. (1948): Novye cikady (Homoptera – Cicadina) Srednerusskoy fauny. – Nauch-
no-metodicheskie Zapiski Glavnogo Upravlenie po Zapovednikam 11: 177-185.
430 Appendix
Appendix
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.
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
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.