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The Leafhoppers and Planthoppers of Germany
Auchenorrhyncha, the planthoppers and leafhoppers, occur in high density and species richness in almost all terrestrial ecosystems, forming an important component of food webs. They may transmit plant pathogens such as viruses and mycoplasmas, and a few species are among the worlds’ worst pests of cereals and other crops. Due to the lack of a comprehensive key, the central European fauna has long been neglected by entomologists, and as a consequence, the knowledge of their life history was rather small. Host relations, in particular, were insufficiently known. This work summarises an extensive data collection combined with a widely scattered literature on a large geographical scale. For the first time, a systematic overview of phenology, food plants, habitat requirements, geographical and altitudinal distribution and economic importance of all Auchenorrhyncha species known from Germany is provided, based on up to date taxonomy. The guild composition of all plant groups on family and species level is analysed and discussed in detail. Further analyses deal with life strategies such as host specificity, dispersal and life cycles. The book is a highly welcome synthesis and reference for all students of biodiversity as well as for ecologists, entomologists, botanists and phytopathologists.
Herbert Nickel
Cover illustration (from top to bottom): Zygina flammigera (Geoffr.), Errastunus ocellaris (Fall.), Conomelus anceps (Germ.), by courtesy of Ekkehard Wachmann. Background by courtesy of Thomas Fechtler.
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The Leafhoppers and Planthoppers of Germany Patterns and strategies in a highly diverse group of phytophagous insects Herbert Nickel
Co-published by Pensoft Publishers, Sofia–Moscow Goecke & Evers, Keltern
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Contents THE LEAFHOPPERS AND PLANTHOPPERS OF GERMANY (HEMIPTERA, AUCHENORRHYNCHA): Patterns and strategies in a highly diverse group of phytophagous insects Herbert Nickel
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Contents
The Leafhoppers and Planthoppers of Germany (Hemiptera, Auchenorrhyncha): Patterns and strategies in a highly diverse group of phytophagous insects Herbert Nickel
Co-published by Pensoft Publishers, Sofia-Moscow Goecke & Evers, Keltern
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THE LEAFHOPPERS AND PLANTHOPPERS OF GERMANY (HEMIPTERA, AUCHENORRHYNCHA): Patterns and strategies in a highly diverse group of phytophagous insects
© Herbert Nickel
Pensoft Series Faunistica No 28 ISSN 1312-0174 Co-published by Pensoft Publishers, Sofia-Moscow (ISBN 954-642-169-3) Goecke & Evers, Keltern (ISBN 3-931374-09-2)
Pensoft Publishers, Acad. G. Bonchev Str., Bl.6, 1113 Sofia, Bulgaria Fax: +359-2-870-45-08, e-mail:
[email protected], www.pensoft.net
Vertrieb in Deutschland, den anderen Mitgliedsstaaten der EU und der Schweiz Goecke & Evers, Keltern (in Antiquariat Goecke & Evers) Inh. Erich Bauer Sportplatzweg 5, D-75210 Keltern, Germany Fax: 07236-7325; International Fax (++49) (0)7236-7325; E-mail:
[email protected]
First published 2003
© PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner.
Printed in Bulgaria, May 2003
Contents
Dedicated to
Günter and Eva Groh († Feb 2nd 1996, and Feb 5th 1996)
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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
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4.1.5 Tettigometridae Germar, 1821 .......................................................................... 65 4.1.6 Issidae Spinola, 1839 .......................................................................................... 69 4.1.6.1 Caliscelinae Amyot & Serville, 1843 .................................................. 69 4.1.6.2 Issinae Spinola, 1839 ............................................................................. 69 4.2 Cicadomorpha Evans, 1946 – Leafhoppers (s.l.) ..................................................... 70 4.2.1 Cicadidae Leach, 1815 (s.l.) – Cicadas ............................................................ 70 4.2.2 Cercopidae Leach, 1815 – Froghoppers .......................................................... 72 4.2.2.1 Cercopinae Leach, 1815 ........................................................................ 72 4.2.2.2 Aphrophorinae Amyot & Serville, 1843 ............................................ 74 4.2.3 Membracidae Rafinesque, 1815 – Treehoppers ............................................. 78 4.2.4 Cicadellidae Latreille, 1825 - Leafhoppers (s.str.) ......................................... 80 4.2.4.1 Ulopinae Le Peletier & Serville, 1825 ................................................. 80 4.2.4.2 Megophthalminae Kirkaldy, 1906 ....................................................... 81 4.2.4.3 Ledrinae Kirschbaum, 1868 ................................................................. 82 4.2.4.4 Macropsinae Evans, 1935 ..................................................................... 82 4.2.4.5 Agalliinae Kirkaldy, 1901 ..................................................................... 91 4.2.4.6 Idiocerinae Baker, 1915 ......................................................................... 94 4.2.4.7 Iassinae Amyot & Serville, 1843 ........................................................ 101 4.2.4.8 Penthimiinae Kirschbaum, 1868 ....................................................... 103 4.2.4.9 Dorycephalinae Oman, 1943 ............................................................. 103 4.2.4.10 Aphrodinae Haupt, 1927 .................................................................. 103 4.2.4.11 Cicadellinae Latreille, 1825 .............................................................. 109 4.2.4.12 Typhlocybinae Kirschbaum, 1868 .................................................... 111 4.2.4.13 Deltocephalinae Fieber, 1869 ........................................................... 163 4.3 Dubious records ......................................................................................................... 228 4.4 Short-term changes .................................................................................................... 230 4.4.1 Declines ............................................................................................................. 230 4.4.2 Range expansions ............................................................................................. 233 4.4.3 Introductions ..................................................................................................... 234 4.4.4 Occasional influxes .......................................................................................... 235 4.5 General remarks on the German Auchenorrhyncha fauna ................................. 235 5 Utilization of plant resources ........................................................................................ 237 5.1 Plant taxa ..................................................................................................................... 237 5.1.1 General remarks and problems ..................................................................... 237 5.1.2 Plant groups and their associated Auchenorrhyncha guilds .................... 242
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5.1.2.1 Pteridophyta ........................................................................................ 242 5.1.2.2 Gymnospermae ................................................................................... 244 5.1.2.3 Ranunculaceae ..................................................................................... 246 5.1.2.4 Ulmaceae .............................................................................................. 246 5.1.2.5 Urticaceae ............................................................................................. 248 5.1.2.6 Fagaceae ................................................................................................ 250 5.1.2.7 Betulaceae (incl. Corylaceae) ............................................................. 253 5.1.2.8 Tiliaceae ................................................................................................ 257 5.1.2.9 Salicaceae .............................................................................................. 258 5.1.2.10 Ericaceae ............................................................................................. 262 5.1.2.11 Rosaceae .............................................................................................. 264 5.1.2.12 Fabaceae .............................................................................................. 273 5.1.2.13 Aceraceae ............................................................................................ 275 5.1.2.14 Apiaceae .............................................................................................. 276 5.1.2.15 Lamiaceae ........................................................................................... 277 5.1.2.16 Asteraceae ........................................................................................... 279 5.1.2.17 Juncaceae ............................................................................................ 284 5.1.2.18 Cyperaceae ......................................................................................... 284 5.1.2.19 Poaceae ............................................................................................... 294 5.1.2.20 Further plant groups ......................................................................... 322 5.1.3 Differences between plant taxa ...................................................................... 329 5.1.3.1 Major plant clades ............................................................................... 329 5.1.3.2 Plant families ....................................................................................... 331 5.2 General conclusions on food plant choice ............................................................. 338 5.3 Stratification and plant architecture ....................................................................... 340 5.4 Utilized substrates ..................................................................................................... 344 5.4.1 Phloem ............................................................................................................... 345 5.4.2 Xylem ................................................................................................................. 345 5.4.3 Mesophyll .......................................................................................................... 348 5.4.4 Fungi .................................................................................................................. 349 5.5 Pests on cultivated plants ......................................................................................... 350 6 Life strategies .................................................................................................................... 352 6.1 Host specificity and its possible causes .................................................................. 352 6.1.1 Diet width in Auchenorrhyncha .................................................................... 352 6.1.2 Plant apparency ................................................................................................ 355
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6.1.3 Plant diversity .................................................................................................. 357 6.1.4 Plant defence ..................................................................................................... 358 6.1.5 Nitrogen ............................................................................................................. 361 6.1.6 Interspecific competition ................................................................................ 361 6.1.7 Resource predictability ................................................................................... 365 6.1.8 Neural constraints ............................................................................................ 365 6.1.9 Regional monophagy ...................................................................................... 365 6.1.10 Endophytic fungi ........................................................................................... 366 6.2 Dispersal ...................................................................................................................... 366 6.2.1 Wing length ....................................................................................................... 366 6.2.2 Migration ........................................................................................................... 369 6.2.3 Colonization ...................................................................................................... 372 6.3 Life cycle ...................................................................................................................... 374 6.3.1 Voltinism ............................................................................................................ 374 6.3.2 Dormancy .......................................................................................................... 377 7 Summary ............................................................................................................................ 381 8 References .......................................................................................................................... 385 Appendix ............................................................................................................................... 430 Index of Auchenorrhyncha genera and species .......................................................... 430 Index and vernacular names of plants ......................................................................... 442 Acknowledgements ......................................................................................................... 459
1 Introduction And God said, Let the waters under the heaven be gathered together unto one place, and let the dry land appear: and it was so. And the earth brought forth grass, and herb yielding seed after his kind, and the tree yielding fruit, whose seed was in itself, after his kind. And God made the beast of the earth after his kind, and cattle after their kind, and every thing that creepeth upon the earth after his kind: and God saw that it was good. Genesis 1: 9, 12, 25
Auchenorrhyncha, the planthoppers and leafhoppers, feed on plant sap and occur in high density and species richness in most terrestrial ecosystems. Their functional significance 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; Andrzejewska 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 Pipunculidae (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 & Rodriguez 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. However, their importance may have been underestimated, because they must ingest large amounts of an unbalanced substrate to receive adequate quantities of nutrients. Honeydew of Auchenorrhyncha is rather thinly distributed, but may be a significant and predictable 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.
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Introduction
Auchenorrhyncha identification often requires dissection and examination of the genitalia. There is, however, no modern, comprehensive identification book covering the whole of the central European fauna, although Holzinger et al. (2003) have started to work on such a compilation. Students have to use various keys of adjacent countries (Haupt 1935; Ribaut 1936, 1952; Dlabola 1954; Ossiannilsson 1978, 1981, 1983), monographs and original descriptions, which may diverge in species interpretations and differ in accuracy. Hence, the interest in this group has been rather limited for a long time. Most sampling methods are rather unspecific and do not allow an assignment to certain host species and microhabitats. This can only be done by labourious suction sampling of single plants or by direct observation and collecting of single specimens. Hence, the knowledge of distribution, life histories and insect – plant relationships in Auchenorrhyncha is rather incomplete. The available literature includes a number of ecological and faunistic contributions, although most species statements are brief, unprecise, unsystematic, of only regional validity or even contradictory. On the one hand, many papers have been written by ecologists who had only little time to become acquainted with the complicated taxonomy and who often changed the subject afterwards, on the other hand, most experienced taxonomists deal with the fauna of more southern countries. Therefore, a critical review and summary of information on a current taxonomic state with special reference to food plant relationships does not yet exist. The same is true for an analysis of a complete Auchenorrhyncha fauna and the utilized flora. This work is intended to be the foundation of such a summary, based on all the available data on life cycle, food plants, habitats, life strategies, as well as geographical and altitudinal distribution of all species found in Germany. Many sections of this work will be purely descriptive, and discussions on a number of points will be far from exhaustive. Some aspects will be analysed elsewhere in more detail. Special emphasis is laid on correct taxonomy, which is regarded here as a major basis for research on all aspects of biological diversity. During the process of writing this book, morphological and biomolecular evidence has grown that the Auchenorrhyncha may not be a monophyletic taxon (Bourgoin 1986, 1993; Campbell et al. 1995; Sorensen et al. 1995). However, very recently, Yoshizawa & Saigusa (2001) insisted on the classical opinion of a sister group relationship between the Fulgoromorpha and Cicadomorpha proposed by Hennig (1969) and Kristensen (1975), although based only on the reduction of a sclerite in the fore wing base, which was interpreted as a synapomorphy. Hence, the term Auchenorrhyncha may become obsolete in the future, but is certainly practical from an ecological and historical point of view. Further ecological studies of plant-sucking insects should treat Fulgoromorpha and Cicadomorpha separately and include other groups, notably Psylloidea, Aphidoidea and most Heteroptera.
Study area
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2 Study area
2.1 General remarks The study area is the Federal Republic of Germany (Fig. 1). Species of neighbouring countries were not taken into consideration, even if they have been found in adjacent regions, e.g. the Vosges, the central Alps and the Bohemian Forest. All information on life history is taken exclusively from Germany, if not otherwise stated. Thus, certain ecological traits of species may be different in other parts of the range, particularly life cycle, food plant utilization and altitudinal distribution. This introduction to the study area includes only a brief overview of the most important geographical units and their position, geology, morphology, climate, soils, vegetation and human impact. Much of this information is taken from Liedtke & Marcinek (1994) and Ellenberg (1996).
2.2 The north German plain The north German plain generally comprises the glacially affected lowlands to the north of the hilly regions which occupy much of the central part of Germany. It also includes the basins of Westphalia, the lower Rhine, Cologne and Leipzig. The major part is below 100 m a.s.l., although very small portions near the southern edge are above 200 m a.s.l. The area is covered by vast Pleistocene and Holocene deposits. The soils are usually sandy and poor except in the more loamy marshes near the coast and the river valleys (including the periglacial drainage systems called Urstromtäler), as well as the loess belt stretching along the southern fringe, which is among the most productive farming grounds all over Europe. The climate is suboceanic, but with more atlantic traits in the western parts, the winters usually being rather mild (with mean January temperatures above freezing point) and – compared with other regions – somewhat cooler summers (mean July temperatures around 17 oC, although slightly lower near the coast). Continental features increase 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 exceed 18 oC and are thus comparable with those in southwest German viticultural regions. Likewise, the annual rainfall decreases from west to east, with usually more than
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Study area
Fig. 1: Map of Germany
The region of the Mittelgebirge
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700 mm to the west of a line from Schwerin to Braunschweig (roughly the 11th meridian), 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 ombrotrophic 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 inland in the vicinity of geologic faults and salt works. The formerly vast forests have been strongly reduced by human activities, their original 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 nutrient immissions.
2.3 The region of the Mittelgebirge The area between the foreland of the Alps and the north German plain is treated here as a unit, for reasons of simplicity and due to the lack of zoogeographical borders. The northern limit is formed by the drop-off towards the northern plains, which is locally obscured by loess deposits of the Börde belt. The Danube valley and the slopes of the Swabian Jura down to the Lake Constance basin (Hegau) are the southern limit. The morphological diversity of the region is conspicuous. Major parts consist of various Paleozoic and Mesozoic rocks, except the Pleistocene and Holocene sediments of the larger river valleys (notably the upper Rhine plain) and the volcanoes of the Eifel, Westerwald, 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 reaching 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.
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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 Erzgebirge 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 summer 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 SaaleUnstrut 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 Hessen, 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 Bavarian 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 altitudes 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 water. 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 species. Nowadays this type of grassland has often been abandoned due to its low productivity. In some places, however, it is sustained for reasons of conservation and landscape management.
The Alps, their foothills and foreland
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2.4 The Alps, their foothills and foreland These regions represent a single unit in terms of physical geography, but not from a biogeographical point of view. The northern border is the valley of the Danube, which flows along the southern slopes of the Swabian and Franconian Jura and the Bavarian Forest. Most of the southern border is formed by the rugged limestone crests of the northern Alps. The altitudes range from 300 to 400 m a.s.l. in the Danube valley, the Lake Constance basin and along the lower courses of the Inn and Isar, to 500 to 900 m a.s.l. in the moraine regions and the alpine valley bottoms, and to almost 3000 m a.s.l. on the highest mountains. The foreland of the Alps is essentially a depression filled with debris of diverse origin 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 rolling, 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 Donaumoos and Dungau and at the edge of the Alps. The annual precipitation is largely determined 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, poplar 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 vegetation 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 limestone may be covered with monospecific stands of scots pine. Most deciduous tree species 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-
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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 vegetation 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 prevail, 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
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3 Material and methods
3.1 Literature data The most important literature on distribution and ecology is listed in Table 1, although 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 material, 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 misidentified. Due to the fact that the most important progress in Auchenorrhyncha taxonomy was made after focusing on genitalic morphology, which was applied in Germany for the first time only by Wagner (1935 and thereafter), most publications before that were excluded from analysis, unless the material had been revised. In general, it appears that northern and eastern parts of Germany are better studied than southern and western parts. Moreover, the coverage is very uneven, with regions sampled for almost 150 years by a number of collectors, e.g. the vicinity of Mainz, as well as conspicuous gaps almost without any data, e.g. the Saarland (see chapter 3.4).
3.2 Museum collections Compared to some other insect groups, e.g. Lepidoptera and Coleoptera, Auchenorrhyncha are not well represented in most museum collections. Moreover, collectors have focused on large and conspicuous species, like Cicadoidea, Cercopidae and Membracidae, whereas the highly diverse groups of Delphacidae, Typhlocybinae and Deltocephalinae 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
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Material and methods
Table 1. References of important data on ecology and distribution of Auchenorrhyncha in Germany Reference
Study area
Period
Achtziger (1991, 1995) Bieman (1987a) Bittner & Remane (1977) Bornholdt (1991) Emmrich (1966) Fischer (1972) Fröhlich (1996a, 1997) Frommer (1996) Funke & Witsack (1998) Haupt (1925) Heller (1987a) Hildebrandt (1990a, 1995b) Kirschbaum (1868) Kuntze (1937) Marchand (1953) Müller (1978) Nickel (1994) Nickel (1997) Nickel (1999a, 1999b) Nickel & Achtziger (1999) Nickel & Remane (1996) Nickel & Voith (in prep.) Nicolaus (1957) Niedringhaus (1991) Niedringhaus (1997) Niedringhaus & Olthoff (1993) Nikusch (1976) Post-Plangg & Hoffmann (1982) Remane (1958) Remane (1961a) Remane (1987) Remane (1994) Remane & Fröhlich (1994b) Schaefer (1973) Schiemenz (1969) Schiemenz (1971a, 1975, 1976) Schiemenz (1987, 1988, 1990) Schiemenz et al. (1996) Schwörbel (1957) Trümbach (1959) Wagner (1935 ff.) Wagner (1939a) Wagner (1950)
Franconia Europe Rhön Hessen, Franken Vicinity of Greifswald Bavarian Swabia Central Europe Urban areas of Cologne Vicinity of Halle Bavarian Alps Vicinity of Ludwigsburg Weser estuary Vicinity of Mainz Mecklenburg Weser-Ems region Leutratal near Jena Southern Niedersachsen Niedersachsen Bavaria Central Europe Bavaria Bavarian and Allgäu Alps Eastern Thuringia East Frisian Islands Emsland North-western Germany Vogelsberg Eifel Weser-Ems region Western Palearctic Mainzer Sand Central Europe Western Palearctic Bottsand near Kiel Eastern Germany Eastern Germany Eastern Germany Eastern Germany Spitzberg near Tübingen Vicinity of Erlangen North-western Germany Mainzer Becken Central Europe
1989-94
Focus
Hedges, forest edges Review of Ribautodelphax 1973-75 Bog fauna 1986-88 Effects of management 1963 Faunistics, ecology since 1813 Faunistic review 1991-94 Salt marshes 1994 Faunistics, ecology 1996-97 Former mining areas 1924 Faunistics, taxonomy 1979-86 Faunistics, ecology 1985-88 Effects of inundation mostly 1860s Taxonomy, faunistics mostly 1936 Faunistics, ecology 1951 Grassland habitats 1971-74 Hillside gradient 1990-94 Faunistics, ecology 1990-96 Faunistics, ecology 1993-98 Faunistics, ecology 1990-1999 Meadow fauna mainly since 1950 Species list, faunistics 1987-2001 Altitudinal distribution mainly since 1925 Faunistics, ecology 1977-88 Island colonization 1989-94 Faunistics, ecology 1982-92 Faunistics, ecology 1972-74 Faunistics, ecology 1974-75 Dry hillsides 1954-55 Land use effects, ecology Review of Mocydiopsis 1955-85 Ecology, faunistics Review of Zygina 1956-93 Faunistics, ecology 1971-72 Coastal habitats 1963-66 Dry grassland 1969-74 Bog fauna Faunal review Faunal review 1952-55 Faunistics, ecology 1954-57 Faunistics, ecology mostly after 1920 Faunistics, taxonomy mostly 1930s Faunistics, taxonomy Review of Macropsis
Unterfranken Oberfranken Mainly vicinity of Berlin Middle Rhine Mainly eastern Germany Mainly Brandenburg Thuringia Baden-Württemberg Eastern Thuringia Thuringia Mainly eastern Germany Mainly north Germany Germany Mainly Baden Mainly Mainz basin Mainly former GDR Southern Bavaria Borkum Mainly Swabia Germany
Naturwissenschaftliches Museum der Stadt Aschaffenburg
Naturkundemuseum Bamberg
Zoologisches Museum Berlin (ZMB)
Zoologisches Forschungsinstitut und Museum Alexander König (ZFMK) Staatliches Museum für Tierkunde Dresden (MTD)
Deutsches Entomologisches Institut, Eberswalde (DEI)
Naturkundemuseum Erfurt (NME) F. Heller (now partim SMNS) Museum für Naturkunde Gera Museum der Natur Gotha (NMPG) Zoologisches Institut der Universität Halle (ZIH)
Zoologisches Institut und Museum Hamburg (ZIMH) H. Haupt (merged in H.J. Müller’s, now MTD)
Staatliches Museum für Naturkunde Karlsruhe (SMNK) C.L. Kirschbaum (now Museum Wiesbaden, MWNH) H.J. Müller (now MTD)
Zoologische Staatssammlung München (ZSM) R. Struve (now Westfälisches Landesmuseum für Naturkunde, Münster)
Staatliches Museum für Naturkunde Stuttgart (SMNS) W. Wagner (now partim ZIMH)
Study area
Collection
Nickel (unpublished) Nickel (unpublished)
(1996), Nickel (unpublished) Schönitzer & Oesterling (1998a, 1998b) Niedringhaus (1989)
Voigt (pers. comm.) Wagner (1939a) Schiemenz (1987, 1988, 1990), Schiemenz et al.
Nickel (unpublished) Schiemenz (1987, 1988, 1990), Schiemenz et al. (1996), Wagner (1939a), Nickel (unpublished)
(1996), Emmrich (pers. comm.) Schiemenz (1987, 1988, 1990), Schiemenz et al. (1996) Nickel & Sander (1996) Nickel (unpublished) Nickel & Sander (1996) Nickel & Sander (1996) Schiemenz (1987, 1988, 1990), Schiemenz et al. (1996)
Schiemenz (1987, 1988, 1990), Schiemenz et al. (1996), Nickel (unpublished) Nickel (unpublished) Schiemenz (1987, 1988, 1990), Schiemenz et al.
Nickel (in prep.)
Wagner (1951a)
Revision
Table 2. Overview of the collections included in this study (study areas specified for larger collections only refer to Germany)
Museum collections
11
12
Material and methods
(ZIMH). Furthermore, Wagner used to exchange material freely with numerous colleagues, e.g. F. Heller, H.-J. Müller and M. Nicolaus, which is now found in their collections. 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 collection 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 revisions (Fieber 1866; Ribaut 1936; Vilbaste 1962; Emelyanov 1964a).
3.3 Own data Gathering of material and data started in 1990, focusing on food plant relationships, habitat requirements and distribution, with emphasis on the southern and eastern half of Niedersachsen, Franconia and northern and central parts of Thuringia, furthermore on the Bavarian and Allgäu Alps, Palatinate and southern Baden. Smaller collections have been taken in the following regions: eastern Holstein, East Friesia, Lüneburger Heide, western Mecklenburg, Drömling, Oderbruch, northern Hessen, Oberlausitz, Nahe valley, Swabian Jura and southeastern Bavarian Forest. Altogether, more than 500 sites have been sampled, the number of specimens taken exeeds 250.000.
3.4 Sampling deficiencies Sampling deficiencies can either exist on a specific level (meaning that certain species 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-represented in many samples, particularly certain species of Delphacidae (e.g. among the genera 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 European Auchenorrhyncha species spending its whole life cycle in the ground, although single macropterous specimens may be caught on dispersal flights (section 4.1.1). Furthermore, 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 nitidulus (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 identified. 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, Stuttgart, 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, Uckermark, middle and southern parts of Brandenburg, lower Rhine, Münsterland, Moselle basin, Saarland, north Baden, eastern Bavaria and certain parts of Württemberg.
3.5 Sampling methods 3.5.1 General remarks Auchenorrhyncha occur in almost all types of terrestrial habitats in humid to semiarid climates. Their abundance in the herb and canopy layer often exceeds 1000 ind./ m2, and even the soil and litter layer are utilized by some taxa or developmental stages. Thus, due to high densities, non-quantitative sampling can be done easily. However, it must be taken into consideration that the small-scale distribution of host specialists is often clumped. Gathering of quantitative data requires more effort and is mainly applied to the herb layer, but not to the shrub and canopy layer. Frequent blurring of sampling results is caused by individuals which migrate or passively drift, or have simply fallen down from trees. Such specimens do not belong to the autochthonous fauna, but may affect results considerably, particularly species number and diversity indices. In general, records of single and macropterous individuals as well as of monophages away from their host plants should be treated with care, especially during windy wheather, on hot summer days with strong thermals and on mountain ridges. In the following sections, the commonly-used sampling methods will be enumerated and discussed. Much of the author’s material was gathered by non-quantitative sweep-netting with special reference to potential food plants. Additional material originated from suction catches, pitfall and Malaise traps as well as ground photo-eclectors.
14
Material and methods
3.5.2 Quantitative methods At present, suction sampling is the only practicable way of gaining data on the abundance of herb layer Auchenorrhyncha (Haas 1980; Törmälä 1982). For this purpose, a conventional, petrol-driven leaf-blower can be converted by fixing a flexible or stiff tube at the suction opening. The tube is supplied with a fine-mesh collection bag, fixed around the inlet either by a cuff or by rubber bands (Stewart & Wright 1995). In addition, the apparatus may be mounted on a backpack frame, but can also be carried with one hand. Finally, the sample is put into a plastic bag or a glass jar, either in alcohol or a cold bag. Depending on the vegetation, there can be a large amount of litter and soil, which requires a considerable effort for sorting out the insects. For preliminary studies and, if only adults and older nymphs shall be sampled, it may be sufficient to put the whole catch into an open box. Auchenorrhyncha climbing and hopping up the sides can be collected with an aspirator, thus reducing the sorting effort, although some macropterous individuals may escape. After long experience they may even be identified and released on the spot. However, a number of species, notably those belonging to Macrosteles, Psammotettix and Jassargus, cannot be identified in the field even by experienced taxonomists, as well as most smaller instars of nymphs. Hence, such individuals should always be checked in the laboratory. Unknown nymphs can be reared until emergence. For experiments requiring statistical analysis, this method has shown to be very efficient in most habitats with a complete cover of grasses and herbs, particularly in various types of grassland and in open forests. In heterogenous habitats it yields much better results than sweep-netting, which almost entirely neglects epigeic species and is much affected by surface qualities and vegetation structure. Ground photo-eclectors can be used for estimating densities of species spending the nymphal stage in the soil and litter and move up to higher layers after molting into the adult stage, namely among Cixiidae, Cicadidae and Cercopinae. Distribution may be clumped around patches of oviposition, however. Hence, larger numbers of samples may be needed for obtaining statistically significant results. This method is only practicable within more extensive programs sampling all groups of arthropods emerging from the soil. Although cixiid planthoppers may be common in some forest habitats, they have rarely been treated on a specific level. On the other hand a tent-shaped photo eclector put up only for a few hours for sampling the herb layer Auchenorrhyncha was shown to be of little efficiency (Törmälä 1982). The soil and litter fauna can be investigated by taking cores plus vegetation and succeeding extraction. The recommended diameter of the corer should be at least around 20 cm. Extraction according to Kempson et al. (1963) requires a special apparatus, but is very efficient, although estimates for the herb layer are usually too low due to escaping individuals. Canopy sampling for quantitative results is critical. Chemical knockdown, which is now commonly applied in tropical forests, has occasionally been done in temperate latitudes (e.g. Southwood et al. 1982; Floren & Schmidl 1999). Auchenorrhyncha have not been treated on a specific level, but there may be potential for gaining accurate estimates for canopy abundances. However, it is difficult to relate these numbers to
Semi-quantitative methods
15
area size (Basset et al. 1997, see this paper also for further methods and discussion of canopy sampling). 3.5.3 Semi-quantitative methods The term „semi-quantitative“ is here referred to methods, which can be standardized, but do not yield estimates, which can be related to the ground area size. Comparability of the results is more or less limited. In sites with similar height and structure of the vegetation and even surface, for instance in most anthropogenic grasslands, standardized sweep-netting provides good results, which can be compared within the same study and on similar plots (Witsack 1975). The sweep net mouth should be D-shaped or square in order to cover much of the soil surface. Numbers of individuals and species caught are often high, although epigeic species are usually much under-represented. However, comparison can be critical in tall and inhomogeneous vegetation, with changing time of day and weather conditions and between different collectors. Furthermore, comparisons between different species of the same plot are critical because of differing vertical distribution (Andrzejewska 1965; Novotný 1992; Peter 1981). In fenland habitats with tussock sedges and on forest floors with only scattered vegetation this method is not appropriate. For special questions, it may be used along homogeneous fringes of forests and hedges (Achtziger 1995). Sweep-netting was done for a large proportion of the existing studies on central European grassland Auchenorrhyncha (e.g. Emmrich 1966; Marchand 1953; Niedringhaus 1991; Nikusch 1976; Remane 1958; Schiemenz 1969; Achtziger & Nickel 1997; Nickel & Achtziger 1999). Pitfall traps also yield high numbers of individuals, but fewer species. This may be caused by the clumped distribution of many species, particularly those living monophagously on tussock grasses and sedges. The traps usually catch all the epigeic species not sampled by sweep-netting (Agalliinae, Anoscopus spp., Streptanus spp.). Thus, using pitfall traps for Auchenorrhyncha is only efficient as a supplement of sweep-netting. However, both methods combined sample almost the complete range of herb-layer species, although a direct search may be necessary in large tussocks. An important shortcoming is the fact, that relation to area size is not given, and comparability may be critical even within the same study. Individual and species numbers caught in tree eclectors may show high variability, depending on tree species, site and exposure. Nymphs and adults of obligate canopy species, which have fallen down and climb up afterwards, are most numerous. Also, obligate vertical migrants move up the tree trunks after emergence and are caught. All these species are capable of flight, so the proportion of sampled individuals in relation to the total number is unknown. Also, it is difficult to relate catch numbers and area size. Altogether, tree eclectors offer information, which may be valuable, although difficult to interpret, particularly in closed forests. Some work has been done by Büchs (1988) and Nickel (unpublished data). Other catches have been counted on supra-specific level only (e.g. Ellenberg et al. 1986). At present, the Malaise trap is one of the most efficient methods for sampling species living in taller and woody vegetation, in particular those migrating either vertically
16
Material and methods
(from one layer to another) or horizontally (from one plant to another), notably among Cixiidae, Empoasca, Zyginidia, Zygina, Arboridia, Allygus, Allygidius and Thamnotettix. Numbers of individuals and species are often very high and may exceed 3000 specimens 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 qualities in relation to sun-exposure and individual plant properties. On shrubs and along forest margins with deep-hanging branches, a standardized beating was carried out. A few good flyers and momentarily sucking individuals may escape, but, depending on the study question, samples within the same study and on similar sites may be comparable (Achtziger 1995). 3.5.4 Qualitative methods If only species lists and food plant records are needed, non-quantitative sweep-netting and direct search on the plants are very efficient. Specimens are taken selectively with an aspirator, thus avoiding large catches of other arthropod groups and common species, which have to be sorted out in the laboratory. When sampling large and heterogeneous plots with quantitative or semi-quantitative methods, search on single plant species may offer useful supplementary data on communities and regional host preferences. Therefore, two to four samples per season can produce almost the complete range of species, provided the student has a good knowledge of food plants. Moreover, residence of most species can be proved by laying emphasis on nymphal records. In forests, however, recording of arboricolous species is critical, except along glades and rides with deep-hanging branches. Thus, some arboricolous species are frequently underrecorded because they spend most of their life in tree canopies.
3.6 Literature on identification There is no single concise identification key for central European Auchenorrhyncha yet. Earliest major works, mostly dating from the second half of the 19th century, notably by Herrich-Schäffer (1829 ff.), Flor (1861), Kirschbaum (1868), J. Sahlberg (1871), Fieber (1875, 1876, 1877, 1878, 1879), Edwards (1896) and Melichar (1896) form an indispensable foundation for taxonomic research, but can hardly be used for identification today. However, Fieber, and particularly, Edwards had a surprisingly well-developed knowledge of genital morphology, despite insufficient optical gear at that time.
Literature on identification
17
This knowledge has long been ignored, notably in the series „Danmarks Fauna“ (Jensen-Haarup 1920), „Fauna van Nederland“ (Blöte 1927, 1943) and „Tierwelt Mitteleuropas“ (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 landmark for future work worldwide. Most species are described in detail, together with excellent drawings of the genital apparatus. It encompasses most of the German leafhopper fauna and shows deficiencies only in groups, the taxonomy of which is critical still today (notably Macropsinae, Zygina). Della Giustina (1989) completed both volumes with new information on taxonomy, nomenclature and faunistics, including numerous 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 standard work on the Auchenorrhyncha fauna of all Fennoscandia and Denmark, which was published in the series „Fauna Entomologica Scandinavica“. It contains numerous 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 ˇ in the series of „Fauna CSR“. 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 research on Auchenorrhyncha bioacoustics, demonstrating that apodemes of male abdominal 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 Auchenorrhyncha by collecting numerous figures from the original literature and encompassing two volumes. Written in Finnish and published in the series “Suomen Eläimet” (Animalia Fennica), this work received little attention, despite its high standard. Similarly, the Fulgoromorpha, Cicadidae, Cercopidae and Membracidae of Estonia were comprehensively described by Vilbaste (1971). However, treating only a small portion of the central European fauna and written in Estonian, it is of limited value for many users. The series of „Fauna Ukraini“ includes a volume on the Ukrainian Fulgoromorpha, written by Logvinenko (1975), which allows the identification of numerous central Eu-
18
Material and methods
ropean species, particularly of delphacids, thus forming an important supplement to the remaining works. In some cases, however, species interpretations should be checked and compared with those of western European authors. Remane & Wachmann (1993) offered an introduction provided with numerous excellent colour photographs, with a broad and concise overview of biology and diversity of the central European Auchenorrhyncha fauna. Although it is not intended to be an identification guide, identification of conspicuous species can easily be done. Furthermore, it can be used for visual confirmation of dichotomous keys. Work on the compilation of a new key to the central European Auchenorrhyncha is in progress and has just resulted in the completion of the first volume covering the Fulgoromorpha and Cicadomorpha excl. Cicadellidae (Holzinger et al. 2003, in print). Useful supplementary information can also be taken from Mitjaev (1971) on the Auchenorrhyncha of Kazakhstan and from Anufriev & Emelyanov (1988) for those of the Far East of the former Soviet Union. Apart from these, there are numerous taxonomic papers and monographs on certain groups or geographical areas, which are indispensable for identification of some species. Above all, W. Wagner has to be mentioned, who did an enormous amount of work on the Auchenorrhyncha fauna of large parts of central Europe as well as the Mediterranean region. By describing and revising taxa and clarifying uncertain names, he paved the way for modern research on this group (Wagner 1937a, 1939a, 1939b, 1941a, 1944, 1948a, 1949a, 1949b, 1950, 1952, 1953, 1955, 1963b, 1964 and others). More recent descriptions and revisions, which have not or only partially been integrated into the above mentioned major works, have been presented by Asche (1980, 1982a), den Bieman (1987a), Booij (1981), Dworakowska (1971, 1972a, 1976, 1993), Emelyanov (1964b), Gillham (1991), Hoch & Remane (1983, 1985), Lang (1945), Lauterer (1958), Lindberg (1948), Linnavuori (1957), Nast (1981), Remane (1961a, 1965, 1994), Remane & Holzinger (1995), Remane & Jung (1995), Ribaut (1934), Tishechkin (1998, 2002), Vilbaste (1972), Wilson & Claridge (1999) and Nickel (1999a). Further important help for identification of certain taxa is offered by Asche (1994), della Giustina (1983), Heller (1987a, 1987b), Holzinger (1999a), Nast (1966) and Remane & Fröhlich (1991). For Cicadula and Jassargus it has been demonstrated that females within diverse genera, which were by then considered indistinguishable, can be identified after dissection of the oviposition apparatus (Remane & Fründ 1986; Schulz 1976). Thus, further research on female genitalic morphology of difficult taxa may have the potential to solve urgent taxonomic problems (see Bourgoin 1993). Usually identification of nymphs is possible at least in later instars and down to generic level. Study of nymphs can be easily done by sampling separate food plants, thus collecting both adults and nymphs together, and by rearing unknown nymphs in the laboratory, if possible on their field host plant. However, literature on nymphal morphology is scanty. Wagner (1950) was the first to realize that for Macropsis, which was one of the most difficult Auchenorrhyncha groups by then, the nymphs are often easier to identify than adults. It was Vilbaste (1968a, 1975, 1982), who achieved great progress in nymphal morphology, who stressed its importance for systematics and who compiled a key for northern European taxa. A little later, Kathirithamby (1971, 1973) studied and described the nymphs of 22 selected British species of Cicadellidae as was done
Literature on identification
19
by Walter (1975, 1978) for a number of central European Euscelinae. Wilson (1978) produced 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 political 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 mainly 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 Ossiannilsson (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 generation 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 Germany it is confined to much lower altitudes and rarely occurs above 400 m a.s.l. The encounter frequency in host plant stands is roughly divided into three classes (high, intermediate, low). However, this parameter does not refer to individual numbers and is much biased by sampling methods and conditions. Distributional borders are described only roughly due to shortage of faunistic data. Normally, larger cities are mentioned (at distances of no more than 30 km from the locality) as well as more important geographical units. Maximum altitudinal records are only stated, if residence is proved or likely. If mountain data from Germany are insufficient, literature from alpine areas of Austria, Switzerland, Italy and France are quoted. It has to be stressed here that mountain slopes and particularly ridges often attract migrating and drifted Auchenorrhyncha, including many species originating from much lower altitudes and different habitats. Some unusually high altitude data in the literature may refer to such records. Another important aspect is the downward shift of vegetation zones towards the north, which amounts to almost 500 m from the Alps to the Harz Mountains (Ellenberg 1996). After each species chapter references are listed. References for identification are only given in critical cases as well as for species which are not or not sufficiently treated in the main identification works (della Giustina 1989; Holzinger et al. 2003; Ossiannilsson 1978, 1981, 1983; Ribaut 1936, 1952). The following abbreviations were used: I – XII B M E egg nymph
Months of occurrence of adults Beginning (1st to 10th day of month) Mid (11th to 20th day) End (21st to 31st day) Overwintering in the egg stage Overwintering in the nymphal stage
22
Review of species
adult gen. m a.s.l. MNHN MTD SMNS ZFMK ZIMH ZMB ZMH ZSM HN
Overwintering in the adult stage Generation Meters above sea level Muséum National d’Histoire Naturelle, Paris Museum für Tierkunde Dresden Staatliches Museum für Naturkunde Stuttgart Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn Zoologisches Institut und Museum Hamburg Zoologisches Museum Berlin Zoological Museum Helsinki Zoologische Staatssammlung Munich Collection of Herbert Nickel, Göttingen
4.1 Fulgoromorpha Evans, 1946 – Planthoppers 4.1.1 Cixiidae Spinola, 1839 Nymphal ecology of the Cixiidae is poorly known; most species live in the soil and litter layer feeding on roots and perhaps fungi (Wilson et al. 1994). Adults are usually found above ground on various dicotyledonous, often woody plants and are usually considered to be polyphagous. However, the host range of nymphs may in fact be narrower, simply because roots of other plant species are out of reach at least under trees. Numbers of sampled adults are generally low, except in Malaise traps and around patches of emergence shortly after it. In some groups there is an evolutionary trend towards retaining cryptic life habits during adult stage. Much of the taxonomy of species treated here was clarified by Wagner (1939a), who studied material mainly from central and southern parts of Europe, and China (1942), who focused on western European material. However, the Cixiidae still comprise some of the most critical taxa among European planthoppers (see below). Very recently, Holzinger et al. (2003, in press.) compiled an up to date key to central European species.
Cixius nervosus (Linnaeus, 1758) Two morphs occur in central Europe: the nominate morph and the var. longispinus W. Wagner, 1955. The latter’s aedeagal theca has a much longer lateral spine. Apparently, in the study area they differ neither in ecology nor distribution. They are of equal abundance and often occur together; intermediate specimens have been found. However, it is uncertain which morph corresponds to the Linnean type material.
M V – E IX; nymph, 1 gen. Adults are found on various deciduous shrubs and trees, often Salix, also Alnus, Betula, Corylus, Tilia and others; in wet to damp habitats of various types (along forest roads and margins, hedges, in open forests, on solitary trees, even among subalpine alder scrub and alpine dwarf shrubs), often near water. In Germany the most common and widespread species among cixiid planthoppers, frequently up to at least 1850 m a.s.l. in the Bavarian Alps and also occurring at highest altitudes of the Mittelgebirge, as well as on the Frisian Islands. Achtziger (1991), Fischer (1972), Niedringhaus (1991, 1997), Remane (1987), Schiemenz (1987), Schwoerbel (1957), Voigt (1978), Wagner (1935, 1939a, 1951a, 1955), HN and others
Cixiidae Spinola, 1839
23
Cixius cunicularius (Linnaeus, 1767) B V – E IX; nymph, 1 gen. Usually along sunny margins of forests and hedges; adults are found on various deciduous shrubs and trees (Corylus, Betula, Alnus, Ulmus and others). The species is widespread throughout Germany (although numbers of collected specimens are usually low), up to at least 1200 m a.s.l. in the Mittelgebirge and the Bavarian Alps. Fischer (1972), Schiemenz (1987), Schönitzer & Oesterling (1998a), Wagner (1935, 1939a), HN and others
Cixius simplex (Herrich-Schäffer, 1835) Mainly E VI – M X, according to Schiemenz (1987) also M V; nymph, 1 gen. Habitat requirements are poorly known; the species is found in dry to moist habitats, adults mainly along woody margins (published food plants are Alnus, Betula, Tilia, Prunus spinosa, Pinus), occasionally also among herbaceous vegetation of wet meadows and fens. Widespread in the lowlands of Germany, but seldom recorded. Northernmost localities are near Bremen, Lauenburg, Rostock, Greifswald and on the island of Rügen, up to at least 600 m a.s.l. in the Mittelgebirge; not found in Denmark and Scandinavia. Achtziger (1991), Fischer (1972), Nikusch (1976), Post-Plangg & Hoffmann (1982), Remane (1987), Schiemenz (1987), Schönitzer & Oesterling (1998a), Schwoerbel (1957), SMNS, Wagner (1939a, 1940a, 1951a), Weis & Schönitzer (2001), HN; see also Nast (1987), Ossiannilsson (1978)
Cixius wagneri China, 1942 After Wagner (1939a) revised Cixius species of central and southern Europe, China (1942) pointed out that Wagner’s interpretation of C. intermedius Fieber, 1870 referred to a new species and created a name for it.
E V – E IX; nymph, 1 gen. In semi-open stands of deciduous woody plants in damp to dry habitats, often in vineyard situations, but also in floodplains. Food plants of adults are probably various deciduous trees, shrubs and tall-growing herbs. In Germany restricted to the western parts, recorded from the Rhine valley between the Kaiserstuhl and Bacharach, the valley of the Nahe (Schloßböckelheim) and the northern Eifel Mountains (near Mechernich), between 80 and 400 m a.s.l. Remane & Fröhlich (1994b), Rombach (1999a), HN
Cixius distinguendus Kirschbaum, 1868 Most records E VI – E IX, according to Schiemenz (1987) also E V; nymph, 1 gen. In closed forests and along their margins; most records of adults in stands of Fagus, Quercus, Picea and Pinus. Widespread in Germany, but rather localized, mostly collected in low numbers only. The highest known localities are at 650 m a.s.l. in the Bavarian foothills of the Alps. Fischer (1972), Mölleken & Topp (1997), Niedringhaus & Olthoff (1993), Rombach (1999b), Schiemenz (1987), Wagner (1939a, 1940a), Nickel (1999b), HN
24
Review of species
Cixius sticticus Rey, 1891 According to Remane & Fröhlich (1994b) the relationship between this species and C. carniolicus Wagner, 1939, described from Austria, should be investigated. The description of the latter is mainly based upon slight differences in wing venation. Furthermore, there is a strong similarity to C. alpestris W.Wg. (see below). In fact, all three taxa may be conspecific (see also Holzinger et al. 2003).
M VI – E VIII; nymph, 1 gen. In warm and rather dry sites, mainly on dry grassland with scattered shrubs and along sunny forest margins, occasionally also in open oak and pine forests. Adults are usually swept from various shrubs (Rosa and others). In Germany rather localized and at the northeastern edge of the range (but see taxonomic remark above), at present known from southern Niedersachsen (Bodenwerder, Bad Sachsa), the Ahr valley (Altenahr), the Rhön Mountains (near Hünfeld), Thuringia (near Arnstadt, Jena and Gera), along the middle Rhine (Bacharach) and the Nahe valley (e.g. Schloßböckelheim, Bad Münster), up to 500 m a.s.l. Reimer (1992), Remane & Fröhlich (1994b), Schiemenz (1987), Nickel (1997), HN
Cixius alpestris W. Wagner, 1939 See taxonomic remarks above.
So far E VI – B IX; nymph, 1 gen. Reported from high mountains of Poland, Slovakia, Austria, northern Italy and France. This species has recently been recorded in the Berchtesgaden and Chiemgau Alps (Königsbach- and Gotzentalalm, St. Bartolomä, Geigelstein) and the Fricken above Garmisch-Partenkirchen, between 700 and 1700 m a.s.l. Furthermore, ‡‡ which cannot be indentified with certainty have also been found in other sites (e.g. Rotspitz above Hindelang). Thus, it is probably more widespread in the Bavarian and Allgäu Alps. It has been reported to favour “besonnte Schutthänge und Heidewälder” in Austria; specimens from the Engadine (Switzerland) have been taken among a group of Pinus mugo in subalpine scree. Nickel & Voith (unpublished data), HN; see also Günthart (1987a), Wagner & Franz (1961)
Cixius heydenii Kirschbaum, 1868 B VI – B IX; nymph, 1 gen. Largely confined to the Alps and the Carpathians; in Germany known only from the Bavarian and Allgäu Alps and the Bavarian Forest. Adults are usually swept in the shrub and herb layers of damp to moist habitats, mainly in open stands of trees and shrubs of the upper montane and subalpine belt, as well as in alpine dwarf shrub heaths between 1200 and 2000 m a.s.l. (in North Tyrol, Austria, up to 2200 m a.s.l.), although single individuals have been collected down to 600 m a.s.l. Biedermann (pers. comm.), Fischer (1972), Remane & Fröhlich (1994b), HN; see also Christandl-Peskoller & Janetschek (1976), Leising (1977)
Cixiidae Spinola, 1839
25
Cixius beieri W. Wagner, 1939 Cixius haupti Dlabola, 1949 C. beieri Wagner, 1939 has been described from the South Tyrolese Alps (Italy). Specimens from Mittenwald in the Bavarian Alps were named C. montanus Haupt, 1924, which turned out to be a primary homonym, later replaced by C. haupti Dlabola, 1949. According to Remane & Fröhlich (1994a) C. haupti Dlab. and C. beieri W.Wg. are likely to be conspecific. The only character separating them is the length of a lateral spine on the aedeagal theca, which, as in C. nervosus (L.), is variable. Specimens with intermediate length of this spine from the Vogelsberg were described as C. haupti hassicus Nikusch, 1976, but are also found in other regions of the Mittelgebirge. Hence, all three morphs are treated here as a single species.
B VI – E VIII; nymph, 1 gen. In Germany the more common and widespread form with short aedeagal spine (haupti morph) occurs in the Bavarian Alps, the Rhön Mountains, Black Forest, Vogelsberg, Meißner, Solling, Harz Mountains, Thuringian Forest and the Erzgebirge, between 400 and 1200 m a.s.l. German specimens with an elongate aedeagal spine (beieri morph) have been found only in the Harz Mountains and Black Forest. This is mainly a species of open, montane coniferous forests also found along the edges of bogs, meadows and clearings. In central Europe it is confined to the Alps as well as rather cool parts of the Mittelgebirge. Adults are often swept from spruce but also from pine and deciduous shrubs. Bittner & Remane (1977), Haas (1975), Nikusch (1976), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1975, 1987), HN
Cixius cambricus China, 1935 C. borussicus Wagner, 1939 was described from East Prussia and C. austriacus Wagner, 1939 from South Tyrol (Italy). Later, however, Wagner considered them both as conspecific with C. cambricus China (see Christandl-Peskoller & Janetschek 1976; Nast 1987; Ossiannilsson 1978). On the other hand slight differences in the morphology of the aedeagus and the anal tube as well as diverging habitat preferences may indicate the existence of distinct species.
M V – M IX; nymph, 1 gen. Described from the Scottish Highlands, but found in Germany mainly in xerothermic sites with scattered shrubs in the Saale-Unstrut region (between Halberstadt, Sondershausen, Jena, Naumburg and Eisleben), in the Elbe valley near Meißen and in the Kaiserstuhl. All these localities are below 400 m a.s.l. On the other hand there are several sites in the subalpine belt of the Bavarian and Allgäu Alps between 1800 and 2200 m a.s.l. Reported from alpine locations between 1750 and 2430 m a.s.l. in North Tyrol (Austria) and from xerothermic hillsides on Quercus, Acer, Carpinus and Rhododendron in Ukraine. MTD, Schiemenz (1987), ZIMH, HN; see also Christandl-Peskoller & Janetschek (1976), Logvinenko (1975)
Cixius stigmaticus (Germar, 1818) M V – M VII, according to Schiemenz (1987) until B IX; nymph, 1 gen. In the interior and along the margins of deciduous forests in moderately moist to moderately dry sites. Adults were swept from Betula, Alnus, Salix, Quercus, Rubus and other woody plants. Widespread in the lowlands of Germany, although scattered and usually found in
26
Review of species
low numbers only; highest localities are at 550 m a.s.l. in the Mittelgebirge. It is likely that the species has declined in abundance since most records date back before 1970. Fischer (1972), Post-Plangg & Hoffmann (1982), Schiemenz (1987), Wagner (1935, 1939a), HN
Cixius dubius W. Wagner, 1939 B V – E VI, occasionally until E VII; nymph, 1 gen. Adults usually in the shrub and canopy layer of dense to open forests of beech, oak and pine in damp to moderately dry sites (on Corylus, Quercus and others). There are only scattered records from the middle parts of Germany, notably between the Danube and the northern edge of the Mittelgebirge, notably Thuringia (locus typicus: Kleinfurra near Nordhausen), southern Niedersachsen, middle Hessen and Unterfranken, but also from near Bitterfeld, Quedlinburg, Cologne, Bonn and Tübingen, up to at least 500 m a.s.l. The species is not reported from the Netherlands, France and Switzerland, but from Belgium, the Czech Republic, Austria and northern Italy. The Polish records are uncertain. Reimer (1992), Remane & Fröhlich (1994b), Schiemenz (1987), Schwoerbel (1957), Wagner (1939a, 1951a), Nickel (1994), HN
Cixius similis Kirschbaum, 1868 M V – B VIII; nymph, 1 gen. In raised and intermediate bogs, occasionally also on peaty and acidic substrates of other sites (e.g. moist heathlands, open forests and clearings). Adults are usually swept from low-growing woody plants, preferentially Betula pubescens, Pinus, Vaccinium myrtillus and V. uliginosum. Occurs rather localized in Germany, mainly in bog areas of the north German plain, in rather cool parts of the Mittelgebirge (notably Harz Mountains, Thuringian Forest, Erzgebirge, Rhön Mountains, Spessart), as well as in the foothills of the Alps, up to at least 1000 m a.s.l. Bittner & Remane (1977), Fischer (1972), Krause et al. (1992), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1987), Strübing (1955), Wagner (1939a, 1951a), Nickel (2002), HN
Tachycixius pilosus (Olivier, 1791) E IV – M VII; nymph, 1 gen. Adults on various deciduous woody plants (Prunus spinosa, Quercus, Betula, Populus and others) in damp to dry sites, in grassland with scattered shrubs, as well as in open forests. Widespread and rather common at lower elevations in Germany (up to at least 500 m a.s.l.), but apparently absent from large parts of the Mittelgebirge and the Alps. Fischer (1972), Niedringhaus (1991, 1997), Remane (1987), Rombach (1999b), Schiemenz (1987), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Wagner (1935, 1939a), HN
Trigonocranus emmeae Fieber, 1876 E V – E VII; nymph, 1 gen.
Cixiidae Spinola, 1839
27
Life habits poorly known. According to very few existing data, nymphs and brachypterous 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 European deciduous forest zone (northeastern Spain, France, England, northern Italy, Switzerland, Austria, Croatia, Slovakia, Bulgaria, southern Russia). Emmrich (2001), Funke & Witsack (1998), Coll. R. Köhler, Remane & Fröhlich (1994b), Coll. M. Sayer, Nickel & Voith, unpublished data), Nickel (2001), HN; also Emeljanov et al. (2002), Gnezdilov (2000)
Myndus musivus (Germar, 1825) B VI – M VIII; nymph, 1 gen. Adults are found on temporarily flooded sand and gravel banks along unchanneled rivers on willows, notably Salix viminalis and S. triandra. In Germany, this species occurs at the western and northern edge of the range, notably in the valleys of the Elbe and Rhine and their larger tributaries (near Dresden, Meißen, Leipzig, Merseburg, Halle, Stendal, Aschaffenburg, Wiesbaden and Bonn), as well as near Rinteln an der Weser. All localities are below 150 m a.s.l. It is likely to have suffered a dramatic decline due to canalization of rivers, since most records are from before 1950. Haupt (1924, 1935), Kirschbaum (1868), Schiemenz (1987), Wagner (1951a, and unpublished data), HN
Pentastiridius beieri (W. Wagner, 1970) So far only B VI – E VI, according to Nast (1977) in Poland and Ukraine E V – E VII; nymph, 1 gen. On willows on almost bare gravel banks of unchanneled mountain rivers. Has also been taken from Myricaria germanica in Ukraine and from Alnus in the Engadine (Switzerland). This species is only known from the Alps, Beskids and Carpathians. In Germany there are only two records from the upper course of the Isar near Vorderriß, 800 m a.s.l.: 23.VI.1997, 1 †, 1 ‡, as well as 04.VI.1998, 11 ††, 7 ‡‡, on tumbled shrubs of Salix purpurea and S. eleagnos. Very recently populations in eastern France which have been suspected to transmit stolbur phytoplasma in sugar beet fields have been provisionally
28
Review of species
assigned to this species. But regarding its specific habitat requirements, it is likely that another species is involved. Nickel (1999a); see also Gatineau et al. (2001), Günthart (1987a), Nast (1977), Remane & Fröhlich (1994b)
Pentastiridius leporinus (Linnaeus, 1761) E V – B VIII, occasionally until B IX; nymph, 1 gen. In moderately wet to wet sites which are at most rarely flooded, mainly in fens, along rivers, as well as in ruderal and brackish habitats. Adults usually on Phragmites australis, according to published data also on Scirpus, Carex and Eriophorum. In Germany this species occurs only locally, with strongholds in the foothills of the Alps and in the upper Rhine plain; further scattered records are from the Wetterau, the upper Main valley, the island of Amrum, Holstein, the rain shadow area to the east of the Harz Mountains and the Oberlausitz, up to at least 750 m a.s.l. Some of the older literature records from southern Germany may refer to P. beieri (W.Wg.). Fischer (1972), Fröhlich (1996a), Schiemenz (1987), Wagner (1935, 1939a, 1970), Wagner & Wagner (1938), Walter (pers. comm.), HN
Reptalus panzeri (P. Löw, 1883) M VI – B VIII; nymph, 1 gen. In xerothermic sites with scattered shrubs, usually on sunny hillsides or on plateaus. Adults are often found on Rosa and Prunus spinosa, but also on other woody plants (Clematis, Salix, Crataegus, Pinus and others). In Germany restricted to rather warm regions between the Danube and a line running from Koblenz to Frankfurt am Main, Fulda, Gotha, Magdeburg, Leipzig and Dresden, notably in viticultural areas (SaaleUnstrut region, Elbe valley around Meißen, Unterfranken, upper and middle Rhine valley, valleys of the Moselle, Nahe and Neckar, and southern Baden). The highest localities are at 500 m a.s.l. only. Maixner (pers. comm.), Reimer (1992), Schiemenz (1987), Schönitzer & Oesterling (1998a), Schwoerbel (1957), SMNS, Wagner (1939a, 1951a), HN
Reptalus quinquecostatus (Dufour, 1833) Specimens published under this name show considerable morphological variation and may belong to several distinct species; see, for instance, Dlabola (1954), Logvinenko (1975), Musil (1956), Mityaev (1971), Holzinger et al. (2003).
So far only B VII – B VIII; nymph, 1 gen. Reported mainly from southern Europe and southwestern parts of Asia. In Germany there are only 6 records from the upper Rhine plain and Franconia: Speyer, Bruchgraben, 100 m a.s.l., July 1989, in Malaise trap catches; Staffelbach, 240 m a.s.l., 11.VII.1949, 1 †; Coburg, Krebsgrund, 300 m a.s.l., 15.VII.1950, 1 †, 1 ‡; Kitzingen, Klosterforst, 09.VII.1994, 1 ‡, on Salix cinerea; Gerolzhofen, Eichelmannsee, 08.VIII.1994, 1 ‡, on Salix cinerea; Erlangen, Exerzierplatz, 280 m a.s.l., 23.VII.2001, 1 †, 1 ‡. Apparently the species prefers moist sites
Asiracinae Motschulsky, 1863
29
with scattered shrubs and tall herbs on sandy or loamy substrates in warm situations. In Ukraine and Slovakia it is considered to be mesophilous and was collected on tall herbs and low-growing shrubs, notably Melilotus, Carduus, Cirsium, Caragana, Cerasus and Spiraea. Has also been recorded in salt marshes in Hungary. Remane (1995), Nickel (1999b), HN; see also Logvinenko (1975)
Hyalesthes obsoletus Signoret, 1865 E V – M VIII; nymph, 1 gen. This species is restricted in central Europe to rather warm sites mainly in viticultural areas. It lives in sun-exposed vineyard situations on abandoned land, along walls and sunny embankments, also in old gardens and in disturbed patches in dry grassland, usually on stony or crumby soils. In Germany it is only known from the valleys of the Rhine (between Lörrach and Koblenz), Neckar, Main, Nahe and Moselle, as well as the Saale-Unstrut region near Naumburg and Halle, up to 350 m a.s.l. From the Moselle valley Convolvulus arvensis and, less frequently, Ranunculus bulbosus, Urtica dioica, Senecio erucifolius and Artemisia vulgaris have been reported as the main food plants of nymphs and adults. Adults were also recorded on Solanum nigrum, Galeopsis angustifolia, Daucus carota, Chrysanthemum vulgare and Calystegia sepium. Woody plants such as Vitis and Salix were only found to hold single specimens. The species is known as a potential vector of grapevine yellows (also in southwestern Germany) and is reported from southern and eastern Europe as a vector of Stolbur disease of tomatoes and potatoes. Hoch & Remane (1985), Maixner (1994), Remane (1987), Schiemenz (1987), Schwoerbel (1957), Sergel (1984a, 1986), Wagner (1939a), Weber (pers. comm.), Weber & Maixner (1998), HN; see also Brˇcák (1979), Sforza et al. (1999) (see this paper also for nymphal morphology)
4.1.2 Delphacidae Leach, 1815 4.1.2.1 Asiracinae Motschulsky, 1863 Asiraca clavicornis (Fabricius, 1794) Recently Dolling (1996) suggested that Asiraca clavicornis (F.) might be a junior synonym of Cimex aequinoctialis Scopoli, 1763 and Cicada quadristriata Gmelin, 1790, but this was dismissed by Wilson & Asche (1998). On the one hand it is not quite clear to which species these names refer, on the other hand synonymisation would cause far-reaching nomenclatural changes concerning several supraspecific taxa.
Adults I – XII, mainly B VIII – M VI; adult, 1 gen. This species is found in sunny, moderately dry to dry, usually disturbed sites. It is usually taken in low numbers only, but is widespread in the warmer regions in the south and east of Germany (mainly in the valleys of Rhine, Main and Danube, the rain shadow area to the east of the Harz Mountains and Thuringian Basin), where it is found in abandoned vineyards, ruderal sites, along waysides, etc., preferentially near patches of both bare ground and tall herbs or shrubs. The food plants are poorly known; adults are usually found in the herb layer as well as on shrubs. Hence, the species is probably
30
Review of species
polyphagous. The northern edge of its range runs through the Mittelgebirge, approximately 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 species is strikingly coloured and easy to identify, thus, it is likely to have declined in central Europe, as has been presumed for England. Frommer (1996), Krause & Emmrich (1996), Nikusch (1976), Post-Plangg & Hoffmann (1982), Schiemenz (1987), Schwoerbel (1957), Wagner (1939a, 1941a, 1951a), HN; see also Kirby (1992), Nast (1976a)
4.1.2.2 Kelisiinae W. Wagner, 1963 Kelisia guttula (Germar, 1818) Most published records of this taxon are in need of revision due to possible confusion with K. sima Rib. and K. sabulicola W.Wg. (see Remane & Jung 1995); this is particularly true for all coastal sites of north Germany as well as for most other countries outside central Europe.
B VI – E X; egg, 1 gen. On Carex flacca, perhaps also on other low-growing sedges, in temporarily dry to moderately wet sites usually on nutrient-poor soils, mainly calcareous grassland, montane meadows and fens. Widespread but local in Germany with strongholds in the foothills of the Alps (up to at least 1200 m a.s.l.) and the limestone areas of the Mittelgebirge region; there are also a few confirmed localities in the north German plain. Fischer (1972), Müller (1978), Remane (1962, and pers. comm.), Remane & Jung (1995), Rombach (1999b), Schiemenz (1987), Nickel & Achtziger (1999), HN; see also Bieman & Booij (1984)
Kelisia sima Ribaut, 1934 The identity of this species, which was originally described as Kelisia pascuorum var. sima Ribaut, 1934, was clarified by Remane & Jung (1995). Without any further comments it was treated as a distinct species by Wagner & Franz (1961). Moreover, recent field studies showed that the habitats and host plants are different from those of K. guttula (Germ.), although both species may occur syntopically.
B VII – B X; egg, 1 gen. In wet to temporarily wet, sunny to moderately shady sites on basic to acidic substrates, mainly in spring mires and fens, as well as on fairly overgrown gravel banks of alpine rivers, sometimes also in temporarily wet forest glades and clearings. Sedges of the Carex flava group are the host plants. Most German locations are in the Alps and their foothills (valleys of the Iller, Lech and Isar, as well as the Chiemgau and Berchtesgaden Alps); there are also scattered records from the Mittelgebirge (southern Black Forest, 1100 m a.s.l., Rhön Mountains, Main valley, middle Hessen, Thuringian Forest, Erzgebirge) and on the island of Borkum. The distribution is poorly known because many older records of K. guttula (Germ.) are in need of revision. Outside Germany the species is known from the French Pyrenees, Trentino (Italy), Styria (Austria) and the Baltic Sea island of Gotland. Remane & Jung (1995) (see this paper also for identification), Walter (1998), ZIMH, Nickel & Achtziger (1999), Nickel & Sander (1996), HN; see also Wagner & Franz (1961)
Kelisiinae W. Wagner, 1963
31
Kelisia irregulata Haupt, 1935 E VII – E X; egg, 1 gen. On Carex flacca in wet to temporarily dry, sunny to slightly shady, usually basic and nutrient-poor sites, mainly calcareous grassland, montane meadows, calcareous small-sedge fens, occasionally also in open forests (coppice-with-standards, waysides, clearings, etc.). Widespread in southern and middle parts of Germany and fairly common in areas of Jurassic and Triassic limestone, notably in the Alps and their foothills, on the Swabian and Franconian Jura (locus typicus: Blaubeuren near Ulm), in Mainfranken and Thuringia; usually found in intermediate frequency on the host. At present, localities at the edge of the range include Bad Münstereifel, Osnabrück, Warburg, Göttingen, Nordhausen and Jena. Occurs up to at least 1200 m a.s.l. Otherwise reported from Slovakia, Austria and France only. Nikusch (1976), Reimer (1992), Rombach (1999a), Schiemenz (1987), Schwoerbel (1957), Trümbach (1959), Nickel & Achtziger (1999), Nickel (1994), HN; see also Jansky & Okali (1993), Remane & della Giustina (1991), Remane & Guglielmino (2002), Wagner & Franz (1961)
Kelisia haupti W. Wagner, 1939 This delphacid was first collected by Haupt (1935) near Freyburg an der Unstrut (locus typicus) and Karlstadt am Main, but erroneously identified as K. melanops Fieber, 1878. Later Wagner (1939a) revised this material and described it as a new species. Older literature records have to be considered doubtful due to the rather complicated situation within the genus in general and due to the recent splitting of the Kelisia haupti group by Remane & Jung (1995).
B VII – E X; egg, 1 gen. Monophagous on Carex humilis on sparsely vegetated dry grassland, as well as in open pine and oak forests, usually on xerothermic hillsides and plateaus on limestone or gypsum, occasionally also on more acidic substrates (recorded up to at most 500 m a.s.l.). In Germany only scattered, although locally dominant in suitable sites. Known from warmer regions of Thuringia (Kyffhäuser, Thuringian Basin and surrounding Triassic limestone plates, notably in the basins of Saale, Gera, Werra and Wipper), of SaxonyAnhalt (northeastern edge of the Harz Mountains, Unstrut valley) and the Main-SaaleTauber area (between Tauberbischofsheim, Würzburg, Karlstadt, Bad Kissingen and Bad Neustadt), also found in southern Niedersachsen (southern edge of the Harz Mountains near Bad Sachsa). Müller (1978), Remane & Jung (1995), Schiemenz (1969, 1987), SMNS, Nickel (1994), HN
Kelisia hagemini Remane & Jung, 1995 This species and the following have only recently been split from K. haupti W.Wg. after morphometric differences of the male genitalia (Remane & Jung 1995). In Germany all three species apparently occur parapatrically.
M VIII – E VIII, in northern Italy E IX; egg(?), 1 gen. Also on Carex humilis and until very recently known only from high mountains of southern Europe (Pyrenees, southern Alps and Mt. Olymp/Greece) as well as Carinthia (Austria). In Germany it was recorded from the Swabian Jura and the Lake Constance Basin:
32
Review of species
Fridingen, 800 m a.s.l., 21.VIII.1998, 2 ††, 4 ‡‡, and Sipplingen, 450 m a.s.l., 20.VIII.1998, 1 †, in both sites on xerothermic calcareous cliff-tops with scattered trees. Nickel (1999b), HN; see also Remane & Jung (1995), Holzinger (1999b)
Kelisia halpina Remane & Jung, 1995 See taxonomic remark above.
In Austria and the Czech Republic B VIII – E IX (Remane & Jung 1995); egg(?), 1 gen. This species is only known from a few localities in Niederösterreich, Carinthia, North Tyrol (Austria), Südtirol (Italy), Graubünden (Switzerland) and Moravia (Czech Republic). It lives in xerothermic, although temporarily seepy, usually basic sites, on Carex humilis. There is a recent record from the Bavarian Alps: Vorderriß, 850-1000 m a.s.l., 22.VIII.1998, 12 ††, 12 ‡‡, on steep, south-facing calcareous hillsides with scattered stands of pine. Nickel (1999b), HN; see also Remane & Jung (1995), Holzinger (1999b)
Kelisia vittipennis (J. Sahlberg, 1868) M VII – B XI, occasionally until V; egg, 1 gen. Mainly in raised and intermediate bogs where it is often among the dominant species during mid and late summer; less commonly found in peaty meadows and calcareous sedge fens. Host plants are cottongrasses (Eriophorum vaginatum, E. angustifolium, E. latifolium). Records published from dry calcareous grassland in Britain are likely to refer to other species, perhaps K. irregulata Hpt. Rather widespread in Germany but local, mainly in the remnant bog areas of the north German plain, as well as in the Mittelgebirge, the Alps and their foothills. Occurs up to at least 1900 m a.s.l. in North Tyrol (Austria) and Switzerland, although highest known localities in Germany are only at 1000 m a.s.l. Nikusch (1976), Remane (1958), Schiemenz (1987), Wagner (1935, 1939a), Nickel & Achtziger (1999), Nickel (2002), HN; see also Günthart (1987a), Leising (1977)
Kelisia punctulum (Kirschbaum, 1868) M VII – M XI; egg, 1 gen. In tall-sedge swamps in moderately wet to temporarily flooded, moderately eutrophic, sunny to shady sites (fens, fen woods, also wet meadows and along ditches), mainly (perhaps exclusively) on Carex acutiformis. Widespread in southern and middle parts of Germany and locally dominant in suitable sites. There are also scattered records from the north German plain (in Holstein extending northward at least to Itzehoe, Bad Oldesloe and Bad Schwartau), from Ireland, England and eastern Poland, but not from Denmark and Scandinavia. Highest localities are at 550 m a.s.l. only; single individuals collected at higher altitudes are probably mostly vagrants. Heller (1987a), Marchand (1953), Nikusch (1976), Reimer (1992), Remane (1958), Schiemenz (1987), Wagner (1935), Nickel & Achtziger (1999), Nickel & Remane (1996), HN; see also Le Quesne (1960), Nast (1976a, 1987)
Kelisiinae W. Wagner, 1963
33
Kelisia guttulifera (Kirschbaum, 1868) According to Wagner (1939a) K. fallax Ribaut, 1934, described from southern France, is conspecific with K. guttulifera (Kbm.), although Wagner did not see Ribaut’s type material. However, in central Europe there are two ecologically distinct forms (see below), which may turn out to show also morphological differences after a more thorough study, as was recently demonstrated for Kelisia guttula (Germ.) and K. sima Rib. by Remane & Jung (1995) (see above).
M VII – M X; at least ‡‡ frequently until IV/V; ad.(?), 1 gen. Usually in beech and oak forests (often along roads and ditches) in moist or seepy sites on Carex sylvatica and C. remota, occasionally in alder fen woods on C. elongata and in open wetlands (spring mires, wet meadows, sand pits, etc.) on Carex vulpina, C. distans, C. paniculata and perhaps other species. In Germany at least scattered and presumably often overlooked; mainly in middle and southern parts, although apparently absent or rare to the south of the Danube and in the northwest German plain. The northern edge of the range runs from Wales through England, the Netherlands, northern Germany (extending at least to Höxter, Braunschweig, Uelzen, Rendsburg, Rostock and Neustrelitz) and Poland to Lithuania. Highest localities in the Mittelgebirge are at 500 m a.s.l. The locus typicus is near Wiesbaden. Schiemenz (1987), Trümbach (1959), Wagner (1935, 1939a), ZIMH, Nickel (1994), HN
Kelisia praecox Haupt, 1935 M VIII – B VI; adult, 1 gen. In temporarily wet sites on Carex brizoides, usually in open forests, clearings and forest meadows, also on other sedge species in straw meadows of the foothills of the Alps. Widespread in the eastern half of Germany up to 700 m a.s.l., but rather scattered, although locally abundant and probably under-recorded. Localities at the western edge of the range are near Biberach, Tübingen, Aschaffenburg, Gießen, Halle and Neubrandenburg. Not reported from Fennoscandia, Denmark, the Netherlands, Belgium, Switzerland and Italy, but from two isolated localities in the French Rhone Alps. Fischer (1972), Remane & Fröhlich (1994b), Schiemenz (1987), Schwoerbel (1957), SMNS, Trümbach (1959), Wagner (1951a), HN; see also della Giustina & Remane (1999), Nast (1987)
Kelisia pallidula (Boheman, 1847) M VII – E X; egg, 1 gen. Monophagous on Carex panicea in temporarily wet to peaty, basic to acidic, usually sunny sites, preferentially spring mires, small-sedge fens, peaty meadows and intermediate bogs. Widespread in Germany, though sporadic, fairly common in the foothills of the Alps, but in rather small and scattered populations in most regions of the Mittelgebirge and the north German plain. The highest localities are at 1100 m a.s.l. Niedringhaus & Olthoff (1993), Nikusch (1976a), Reimer (1992), Schiemenz (1987), SMNS, Wagner (1935, 1939a), Nickel & Achtziger (1999), Nickel & Remane (1996), HN
34
Review of species
Kelisia nervosa Vilbaste, 1972 Described from Lithuania, but strongly resembling K. confusa Lnv. (see below), which occurs mainly in southeastern Europe.Vilbaste (1972) did not discuss the relationship towards the latter species and was apparently not aware of it. Thus, the relationship between K. nervosa Vilb. and K. confusa Lnv. is in need of revision; they are probably conspecific.
In Lithuania M VIII – E VIII (Vilbaste 1974); egg(?), 1 gen. This taxon is altogether known only from 6 localities in Lithuania, eastern Poland and eastern Germany. The two German records are both from the margins of intermediate bogs in the vicinity of Lübben and comprised fairly large populations: Halbe, end of VIII.1995, and Krausnick, 29.VIII.2001. The latter population lived in tussocks of Carex elata. Remane (1995), HN; see also Nast (1976a), Vilbaste (1972)
Kelisia confusa Linnavuori, 1957 See taxonomic remarks above.
This appears to be an essentially southeast European species living in swampy and moderately eutrophic sites among yet undetermined tall sedges (Carex sp.). Recently it was recorded for the first time in Styria (Austria) and in the south of the upper Rhine plain: Rust, 160 m a.s.l., 03.VIII.1997, 1 † (brach.), and Endingen, 200 m a.s.l., 18.VIII.1998, 1 † (macr.), all in tall-sedge swamps, perhaps on Carex acutiformis. HN; see also Holzinger et al. (1997), Nast (1987)
Kelisia minima Ribaut, 1934 So far M VII – B IX, in France according to Ribaut (1934) until XI; egg(?), 1 gen. Apparently this is a very rare species, which is only known from altogether 8 localities: three older records from northern and southern France, „sur Cyperus longus“, another one from Switzerland (Valais, Dugny, in a spring mire on a hillside, 600 m a.s.l.) and 4 from Germany: island of Hiddensee, Vitte, 03.IX.1965, 1 †, in brackish grassland; Wetterau, Münzenberg, July and August 1966 and 1967; eastern Niedersachsen, Schreyahn near Lüchow, 21.VII.1994, 5 ††, 1 ‡, and Thuringian Basin near Erfurt, Stotternheim, 13.VII.1997, 1 †, 3 nymphs, the latter two in inland saltmarshes in tussocks of Carex distans. The Hiddensee record was already published as K. guttulifera (Kbm.). Emmrich (pers. comm.), Remane & Fröhlich (1994b), Nickel (1997), HN; see also Schiemenz (1987)
Kelisia ribauti W. Wagner, 1938 Belongs to a species group which is subject to diverging interpretations of species distinction and nomenclature among various authors (Asche & Remane 1982). In particular there are populations living in rather montane or boreal habitats, whereas others are widespread in the Mediterranean basin. However, morphological distinction is difficult.
B VII – E X; egg, 1 gen. In peaty meadows, intermediate bogs and spring mires, also along the edge of raised bogs (in the lagg zone, along ditches, in peat diggings, etc.). Probably lives on sedges
Kelisiinae W. Wagner, 1963
35
(notably Carex nigra), although often found in patches with Eriophorum angustifolium or E. latifolium. Is also reported from Carex elongata. Status and distribution in Germany are very similar as in K. vittipennis (J. Shlb.), with which it often co-occurs, although usually in lower abundance. Found up to 1900 m a.s.l. in North Tyrol (Austria) and the Engadine (Switzerland), up to at least 1200 m a.s.l. in the Bavarian and Allgäu Alps and the southern Black Forest. Marchand (1953), Nikusch (1976a), Reimer (1992), Schiemenz (1987), Strübing (1955), Wagner (1939a, 1951a), Wagner & Franz (1961), Nickel & Achtziger (1999), Nickel (1997, 2002), HN; see also Günthart (1987a), Leising (1977)
Kelisia sabulicola W. Wagner, 1952 Most older records of this species were published as K. guttula (Germ.) until Wagner (1938a, 1939a, 1952) pointed out that there are morphologically and ecologically different populations. Some authors (e.g. Nast 1987) consider K. sabulicola W.Wg. a junior synonym of K. pannonica Matsumura, 1910, which was described only after 2 ‡‡ from Hungary and Germany. This book follows the interpretation of Ossiannilsson (1978) and Remane & Fröhlich (1994a), who suggest to retain Wagner’s unambiguous name.
Adults I – XII, mainly E VI – V; adult, 1 gen. Monophagous on Carex arenaria on sand dunes near the coasts of the North and Baltic Sea, as well as inland in open sandy grassland of the north German plain. Most records are from the islands (Heligoland, East and North Frisian Islands, Hiddensee, Rügen); locally also found inland, southward at least to Oldenburg, Dömitz, Magdeburg, Dessau, Potsdam and Dresden; all sites are below 150 m a.s.l. Has been reported from the region of the Mittelgebirge, but these specimens should be revised. The locus typicus is Horst an der Seeve near Hamburg. Niedringhaus (1991, 1994), Remane (1958), Schaefer (1973), Schiemenz (1987), Wagner (1952a), HN
Kelisia monoceros Ribaut, 1934 M VI – E X; egg, 1 gen. Largely confined in central Europe to the Alps and their foothills, preferentially living in various kinds of low-productivity grassland, often with Carex sempervirens, notably on sunny slopes, in open pine forests and on riverine gravel banks. Much more scattered to the north of it where usually found on Carex vulpina ssp. otrubae and C. muricata, in inland salt marshes and disturbed, dry or temporarily dry sites on loamy or clayey substrates, mainly in mining areas and pastures. Most German records are from southern Bavaria (up to at least 1600 m a.s.l., but frequently found above 2000 m a.s.l. in the southern Alps), in the lowlands extending northward to Mainz, Gießen, Halle, Leipzig, Dresden and Görlitz. Apart from a single specimen from the botanical garden in Hanover, there are no records from the north German plain, although the species is reported from Scandinavia. Fischer (1972), Marchand (1953), Remane (1987), Remane & Fröhlich (1994b), Schiemenz (1987), Nickel (1999b), HN; see also Ossiannilsson (1978)
36
Review of species
Anakelisia fasciata (Kirschbaum, 1868) Mainly B VII – X, ‡‡ occasionally until V; mainly egg, 1 gen., according to Witsack (1985) overwintering takes place mainly in thermic quiescence (also a few ‡‡); see also Lauterer (1984). In tall-sedge swamps in wet to temporarily flooded, moderately eutrophic, sunny to shady sites (fens, lake shores and fen woods; absent from mown meadows), on Carex riparia, perhaps also on other species of tall sedges (C. acutiformis and C. acuta). Widespread in Germany, although locally absent or overlooked (for instance, not recorded from Schleswig-Holstein, Brandenburg, Northrhine-Westphalia, Bavaria to the south of the Franconian Jura). Appears to be restricted to lowlands and river valleys; highest localities are at 350 m a.s.l. only. The locus typicus is near Frankfurt am Main. Heller (1987a), Hildebrandt et al. (1998), Remane & Fröhlich (1994b), Schiemenz (1987), Witsack (1985), Nickel & Achtziger (1999), Nickel & Remane (1996), Nickel (1994), HN; see also Lauterer (1984)
Anakelisia perspicillata (Boheman, 1845) E VII – E X; egg, 1 gen. Rather eurytopic (although more xerophilous towards the north) in sunny and oligotrophic sites with short vegetation, ranging from dry to temporarily wet and from basic to acidic conditions, mainly pastures, submontane meadows, dry forest margins and calcareous small-sedge swamps. The main host plant on basic substrates is Carex flacca, and C. pilulifera has been recorded in more acidic sites. Widespread in Germany with strongholds in the Alps and their foothills (frequently up to at least 1050 m a.s.l. in Bavaria, but extending to 1700 m a.s.l. in eastern Austria) and the limestone regions in middle parts, but rather scarce in the north German plain. Nikusch (1976), Reimer (1992), Schiemenz (1987), Wagner (1935), Nickel & Achtziger (1999), Nickel & Remane (1996), Nickel (1994), HN; see also Wagner & Franz (1961)
4.1.2.3 Stenocraninae W. Wagner, 1963 Stenocranus longipennis (Curtis, 1837) E VII – E X; mainly egg(?), 1 gen. (see also Strübing 1963). This is a western European species occurring in rather scattered populations in large tussocks of Carex paniculata in wet fenlands of western and northern parts of Germany. From there it is known only from 5 localities: Friedrichsruh near Hamburg, Stolzenau an der Weser, Denkershausen near Northeim, as well as Erlenbach and Fischbach (220 m a.s.l.), both near Pirmasens. Otherwise it is only reported from Ireland, England, the Netherlands, France and Spain. Marchand (1952), Strübing (1963), Wagner (1947a), ZIMH, Nickel (1994), HN; see also Bieman (1993), Nast (1987), Remane & Fröhlich (1994b)
Stenocraninae W. Wagner, 1963
37
Stenocranus major (Kirschbaum, 1868) Adults I – XII, mainly E VIII – M VI; adult, 1 gen. (see also Strübing 1963). In moderately wet to temporarily flooded, sunny to slightly shaded, rather eutrophic sites, often near standing or running water, also along ditches, in wet ruderal sites (sand pits, mining areas, etc.) and in fen woods, rarely on mown meadows. Phalaris arundinacea is the host plant. Widespread and common in most parts of Germany; in autumn the species is locally extremely abundant and may also be found in stands of Calamagrostis epigejos (but apparently without breeding). Found in Schleswig-Holstein northward only to Pinneberg, Kiel and Lübeck, although there are few records from the Baltic Sea islands of Zealand (Denmark), Bornholm and Öland, and from Skane (all Sweden). Highest localities are at 800 m a.s.l. in the Mittelgebirge and Bavarian Alps. Fischer (1972), Niedringhaus (1997), Remane (1987), Schiemenz (1987), Schwoerbel (1957), Wagner (1939a, 1947b), Nickel & Achtziger (1999), Nickel (1997), HN
Stenocranus minutus (Fabricius, 1787) Adults I – XII, mainly E VIII – E VI; adults overwinter in primary parapause before oviposition, 1 gen. (Müller 1957, 1958; Witsack 1981). On Dactylis glomerata and probably D. polygama in sunny to slightly shaded, moderately dry to moist and moderately eutrophic sites, mainly pastures, meadows, field margins, waysides and open forests; usually absent or only as a temporary stray in most conventionally managed meadows. Widespread and common in Germany, particularly in middle and southern parts, although largely confined to locations below 600 m a.s.l.; single specimens were recorded up to 1000 m a.s.l. Achtziger & Nickel (1997), Fischer (1972), Müller (1978), Post-Plangg & Hoffmann (1982), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz (1987), Schöpke (1996), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN; see also May (1978)
Stenocranus fuscovittatus (Stål, 1858) B VIII – B VII; adult, 1 gen. In moderately wet to temporarily flooded, oligotrophic to moderately eutrophic sites (mainly fens, spring mires, peaty meadows, etc.); host plants are various species of sedges (Carex elata, C. paniculata, C. panicea, presumably also C. acutiformis, C. acuta and others). The species is widespread and fairly common in the middle and southern parts of Germany, but there are only scattered localities in the north German plain where it lives at the edge of the range. It frequently occurs up to 600 m a.s.l. in the foothills of the Alps (possible vagrants found up to 950 m a.s.l.), although only below 350 m a.s.l. in southern Niedersachsen. Northernmost localities are near Stolzenau an der Weser, Mölln, Rostock and Neustrelitz. Also reported from England and northern Poland; records from the Netherlands, however, have turned out to be erroneous. Heller (1987a), Kuntze (1937), Marchand (1953), Nikusch (1976), Schiemenz (1977, 1987), Wagner (1939a, 1951a), Nickel & Achtziger (1999), Nickel (1997), HN; see also Bieman (1993), Le Quesne (1960), Nast (1976a)
38
Review of species
4.1.2.4 Delphacinae Leach, 1815 Jassidaeus lugubris (Signoret, 1865) Adults I – XII, mainly E VIII – E V; adult, 1 gen. In sparsely vegetated dry grassland in sunny, basic to acidic sites, preferentially near the base of fine-leaved species of fescue (Festuca ovina group), perhaps also on Stipa capillata. Rather localized in Germany and on the northern edge of the range, mainly confined to the warmer regions of southern and eastern parts, but perhaps overlooked. Northernmost records are from near Bad Neuenahr, Warburg, Wolfenbüttel, Sondershausen, Halberstadt, Parchim and Waren an der Müritz; the highest localities are at 550 m a.s.l. Kuntze (1937), Nikusch (1976), Reimer (1992), Remane (pers. comm.), Schiemenz (1987), SMNS, Wagner (1939a), Walter (pers. comm.), Nickel & Remane (1996), Nickel (1997), HN
Megamelus notula (Germar, 1830) B VI – E X, according to Schiemenz (1987) single ‡‡ until V; mainly egg, 1(?) gen. Eurytopic in stands of tall sedges of various species in moderately wet to temporarily flooded sites (from oligotrophic intermediate bogs to eutrophic meadow depressions, even ditches within fields). Host plants are various species of Carex (C. acutiformis, C. acuta, C. rostrata, C. nigra, C. disticha and others). Widespread in Germany and often dominant in suitable sites, extending up to at least 1350 m a.s.l. in the Mittelgebirge and the Bavarian Alps, up to 1500 m a.s.l. in North Tyrol (Austria). Heller (1987a), Niedringhaus (1991), Nikusch (1976), Remane (1958), Schiemenz (1971a, 1975, 1987), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN and others
Conomelus anceps (Germar, 1821) E VI – E X; egg, 1 gen. (see also Witsack 1985). In moist to temporarily flooded, sunny to moderately shady sites (notably meadows and pastures, fens, intermediate bogs, forest clearings, ruderal sites). Host plants are various species of rushes (Juncus effusus, J. inflexus, J. articulatus and others). Widespread in Germany and common in many places, found up to at least 1200 m a.s.l. in the Mittelgebirge and the Bavarian Alps. Maximum densities within tussocks of Juncus effusus in England were over 8000 nymphs (1st and 2nd instar) or 1000 adults per sqm. Bittner & Remane (1977), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Schiemenz (1971a, 1987), Schwoerbel (1957), Wagner (1935), Witsack (1985), Nickel & Achtziger (1999), HN and others; see also Rothschild (1964, 1966)
Conomelus lorifer Ribaut, 1948 This species occurs from Ukraine, the Balkans and Italy, to Poland, Germany and France. Two endemic subspecies, C. l. calabricus Dlabola, 1979 and C. l. lorifer Ribaut, 1948, inhabit southern and middle Italy. C. l. dehneli Nast, 1966 occurs in the remaining parts of the range. These sub-
Delphacinae Leach, 1815
39
species show clinal morphological intergradation into each other (Remane & Asche 1979; Hoch & Asche 1983; Hoch & Remane 1983).
B VI – M X; egg, 1 gen. On Juncus effusus and perhaps other species of rushes, usually in seepy or temporarily wet sites. Appears to favour disturbed habitats and sandy substrates such as sand and gravel pits, mining areas and forest tracks, but has also been found in spring mires and floodplain depressions. In Germany at the northern edge of the range and apparently rather scarce; only found near Gardelegen, Nauen, Nordhausen, Halle, Bad Salzungen, Bad Kissingen and Speyer, but probably under-recorded. All German localities are situated below 300 m a.s.l., but the species was found up to 1400 m a.s.l. in northern Italy. Funke & Witsack (1998), Remane & Fröhlich (1994b), Sander (pers. comm.), Wagner (unpublished data), HN
Delphacinus mesomelas (Boheman, 1850) M V – M IX; nymph, 1(?) gen. In sunny, low-vegetated, dry to wet sites of low productivity. In the north German plain on heaths and other dry and sandy habitats, to the south of it mainly on moderately dry to wet, even peaty meadows usually at submontane altitudes, as well as on various types of dry grassland. Host plants are fine-leaved species of fescue (mainly Festuca ovina group and F. rubra) and perhaps additional grasses. Widespread in northern and middle parts of Germany, but rather local; not recorded to the south of the Danube, but occurring up to at least 1100 m a.s.l. in the southern Black Forest. Bornholdt (1996), Kuntze (1937), Niedringhaus (1991), Reimer (1992), Rombach (1999b), Schaefer (1973), Schiemenz (1969, 1971, 1987), Wagner (1935, 1951a), Nickel & Achtziger (1999), HN
Stiromella obliqua (W. Wagner, 1948) So far B V – M VI, in Austria according to Moosbrugger (1946) until B VII; nymph, 1 gen. The range of this species extends from Yakutia through Mongolia and Kazakhstan to Europe, where it is only known from a few records from Estonia, Latvia, Ukraine, Austria (Niederösterreich and Vorarlberg, reported from Schoenus ferrugineus, up to at least 1000 m a.s.l.) and a single locality in Unterfranken near Schweinfurt (Grettstadt, 210 m a.s.l., 15.V.1993, 17.VI.1994 and 06.V.2001, a large population in a temporarily dry calcareous sedge fen, perhaps on Schoenus nigricans). Nickel & Remane (1996), HN; see also Anufriev & Emelyanov (1988), Moosbrugger (1946, and unpublished data), Nast (1987), Vilbaste (1974), Wagner & Franz (1961)
Ditropis pteridis (Spinola, 1839) M V – B VIII; nymph, 1 gen. Monophagous on Pteridium aquilinum in moderately moist to moderately dry, usually acidic and moderately shady sites, mainly in open forests and along their margins, occasionally in clearings and pastures. Widespread in Germany and in intermediate to
40
Review of species
high frequency on the host, but not recorded to the south of the Danube, where it is probably overlooked. Highest localities are at 1100 m a.s.l. in the southern Black Forest. Reimer (1992), Schiemenz (1987), Wagner (1935, 1939a, 1951a), HN; see also Morcos (1953)
Eurysa lineata (Perris, 1857) B V – E VII; nymph, 1 gen. On various substrates in the grass layer of open, usually thermophilous deciduous and coniferous forests (mainly oak and pine) and along their margins, as well as around hedges, solitary trees and shrubs on rather dry grassland. Host plants are various grasses (Poa nemoralis, Deschampsia flexuosa, Melica uniflora and others). In Germany mainly confined to lower altitudes of middle and southern parts; the northernmost localities are near Bonn, Warburg, Hildesheim and Malchin, the highest ones at 500 m a.s.l. To the north of these there are only single records from southern Sweden, England and Ireland. Achtziger (1991), Post-Plangg & Hoffmann (1982), Rombach (1999b), Schiemenz (1987), Wagner (1939a), Nickel (1994), HN; see also Le Quesne (1960), Ossiannilsson (1978)
Eurysella brunnea (Melichar, 1896) M V – E VII, occasionally until B IX; nymph, 1 gen. Like E. lineata (Perr.) in open, grassy forests and near woody margins in dry grassland, vineyards, etc., but usually in higher-vegetated sites and rarely syntopic. The host plants are grasses, often Brachypodium pinnatum in basic sites, but the species is also found on siliceous substrates, where Poa nemoralis is likely to be the host. Lives mainly near the base of its hosts and is probably much under-recorded, but is usually collected in pitfall traps and with suction samplers. In Germany living at the northern edge of the range and known mainly from rather warm regions in the valleys of the Rhine and Main, and along their tributaries (Ahr, Nahe, Lahn, Fränkische Saale), as well as in the rain shadow area to the east of the Harz Mountains; highest localities are at 550 m a.s.l.; the northernmost records (which appear to be more or less isolated) are from the vicinity of Bonn, Biedenkopf, Warburg, Göttingen, Quedlinburg and Jena. Biedermann (pers. comm.), Kupka (1941), Reimer (1992), Remane & Fröhlich (1994b), Schiemenz (1987), Wagner (1951a), HN; see also Asche (1994), Stewart (1997)
Eurysula lurida (Fieber, 1866) B V – B IX; nymph, 2 gen. In moderately dry to wet sites on small-reed, mainly in forest clearings, sand and coal pits, along forest roads, etc., on Calamagrostis epigejos, locally also on C. canescens in fen woods, fens and intermediate bogs. Single macropterous individuals, however, are frequently found in other habitats. The species is widespread in Germany, but is found only at lower altitudes (to at least 600 m a.s.l.). It is often overlooked due to its secretive habits, although easily seen after direct search on the base of the host. Fischer (1972), Niedringhaus & Olthoff (1993), Schiemenz (1987), Strübing (1955), Walter (pers. comm.), Nickel (1999b), HN
Delphacinae Leach, 1815
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Eurybregma nigrolineata Scott, 1875 E IV – M VII; nymph, 1 gen. In sunny, moderately dry to damp, occasionally moderately moist, usually disturbed sites, which can be quickly colonized by macropterous individuals (along waysides, railway lines and ditches, in abandoned fields and vineyards, low-input meadows, etc., also inland salt marshes). Host plants are tall grasses (mainly Elymus repens, at least locally also Holcus lanatus and Dactylis glomerata). In central Europe this species is suspected to expand its range north- and westward since the 1950s and 60s, although it is known from England since the 19th century (including brachypterous specimens). From eastern Germany (Saale-Unstrut region) it is recorded since 1942, in the west only since 1960. At present it is widespread and locally common in most parts of the country, although not yet recorded to the south of the Danube. Localities on the Baltic Sea island of Rügen, near Schnackenburg, Elmshorn and Bonn may be near the northern and western edge of the range, since there are no records from the Netherlands and Denmark. Largely confined to lowlands below 500 m a.s.l.; macropterous and also brachypterous individuals from higher altitudes and the lower Ems area are perhaps vagrants. In France it is widespread at least in middle and northern parts, in southern Sweden, Belgium, Switzerland and northern Italy it was recently found for the first time. Achtziger (1991), Bornholdt & Remane (1993), Niedringhaus (1997), Reimer (1992), Remane (pers. comm.), Remane & Fröhlich (1994b), Rombach (1999a), Schiemenz (1987), SMNS, Nickel & Achtziger (1999), Nickel (1994), HN; see also Baugnée (1995), della Giustina & Remane (1999), Gillerfors (2002), Günthart (2000), Le Quesne (1960), Nast (1972, 1987), Webb (pers. comm.)
Stiroma affinis Fieber, 1866 M V – E VIII; nymph, 1 gen. In deciduous and coniferous forests, usually in moderately dry to moist sites on various grasses (Brachypodium pinnatum, Calamagrostis arundinacea, Poa nemoralis, Deschampsia flexuosa, Festuca altissima and others). Widespread in the lowlands of Germany and fairly common, locally extending up to 1000 m a.s.l. in the Mittelgebirge and the foothills of the Alps, up to 1500 m a.s.l. in the central Alps of Switzerland. Fischer (1972), Niedringhaus (1997), Reimer (1992), Schiemenz (1987), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN; see also Günthart (1987a)
Stiroma bicarinata (Herrich-Schäffer, 1835) M V – E VIII; nymph, 1 gen. Among rather tall grasses in moderately moist to wet sites, usually in low-input meadows, pastures and abandoned fields (in the lowlands only in cool valley bottoms or in the vicinity of shrubs and trees), in deciduous and coniferous forests (in more moist and shady sites than St. affinis Fieb., but locally syntopic) and in subalpine pastures and tall herb stands. Often in tussocks of Deschampsia cespitosa, but also on other grasses (e.g. Holcus mollis, Glyceria maxima, Dactylis glomerata, Elymus repens, Arrhenatherum elatius).
42
Review of species
Widespread in Germany and fairly common, also in the higher parts of the Mittelgebirge and the Alps, where frequently recorded up to at least 1800 m a.s.l. Bittner & Remane (1977), Fischer (1972), Reimer (1992), Schiemenz (1987), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN and others
Metropis latifrons (Kirschbaum, 1868) M V – B VII; nymph, 1 gen. On fine-leaved species of fescue (Festuca ovina group) in xerothermic, basic as well as acidic, sunny to moderately shady sites on hillsides and plateaus. In Germany at the northeastern edge of the range and only known from a few localities in the valleys and basins of the Rhine, Nahe, Main and Fränkische Saale, mostly on Triassic limestone, aeolian sand and porphyry; the highest localities are at 550 m a.s.l. Furthermore, there is a confirmed record from inland dunes of the lower Oder valley (Bielinek, Poland). Asche et al. (1983), Remane (1987), Remane & Fröhlich (1994b), Wagner (1939a, 1951a), Nickel et al. (2002b), HN; see also Haupt (1931)
Metropis inermis W. Wagner, 1939 So far B V – E VI, nymph, 1 gen. This is apparently an eastern steppe species with isolated populations in eastern parts of Germany, from where it is known only from 6 sites in eastern Bavaria (Naab valley near Kallmünz and Krachenhausen, Laaber valley near Eilsbrunn and Undorf, Altmühl valley near Meihern, all between 350 and 450 m a.s.l.) and eastern Thuringia: Steudnitz near Jena. The latter record comprised a single macropterous † only, probably a stray individual. The Bavarian records were from dry and sun-exposed hillsides and plateaus of Jurassic limestone on a green subspecies of Festuca ovina. The Thuringian specimen was collected in a dry grassland strongly influenced by phosphate immissions. Achtziger (pers. comm.), Remane (1961b), Schiemenz (1987), Nickel et al. (2002b)
Achorotile albosignata (Dahlbom, 1850) In Scandinavia VI – VIII (Ossiannilsson 1978), nymph(?), 1(?) gen. The range of this species extends from the Altai through western Siberia to central and western Europe, where it is known only from rather isolated localities. From Germany there are only two records: a single † from the vicinity of Erlangen, which has been taken in high summer in the mid 1950s „ … in einem trockenen Kiefernwaldstück auf Waldgräsern …“. There is also a published record from Berlin dating from 1901: “In der Umgebung von Berlin ist dieses Insekt häufig auf Festuca- und Agrostis-Arten, besonders auf sterilen, sandigen Hängen.” Was also collected on species of Festuca in open pine forests in the mountains of eastern and central Spain and southern France. Due to the widespread occurrence of sensory pits in the nymphal stage in delphacids, some older reports of this species from central Europe are likely to be based on misidentifications. Matsumura (1906), Nast (1976a), Trümbach (1959) (Remane vid.); see also Remane & Fröhlich (1994b)
Delphacinae Leach, 1815
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Euconomelus lepidus (Boheman, 1847) E VI – M X; egg, 1 gen. Usually in sunny to moderately shady, temporarily or permanently wet, basic to acidic sites with fresh to brackish water (meadow depressions, spring mires, inland salt marshes, sand and coal pits). The main host plant is Eleocharis palustris s.l.; various reports from Juncus must be considered as unreliable regarding host identification. Widespread in Germany, but often occurring in small pockets only, although locally abundant; highest localities are at c. 700 m a.s.l. in Upper Bavaria, but found at 1000 m a.s.l. in the Tyrolian Alps (Austria). Niedringhaus (1991), Schiemenz (1987), Wagner (1935, 1939a), Nickel & Achtziger (1999), Nickel & Remane (1996), HN
Delphax crassicornis (Panzer, 1796) B VII – B IX; egg, 1 gen. Monophagous on Phragmites australis, usually near the base of the stems. Apparently prefers reeds which are not mown and flooded at most during winter; locally also in open fen and floodplain forests and salt marshes. In Germany widespread though sporadic; most records are from the northeastern half (mainly to the east of a line from Sylt to Hamburg, Braunschweig, Gotha and Würzburg), but also found in the Weser-Ems area, the upper Rhine plain and the Allgäu (c. 800 m a.s.l.). Fischer (1972), Remane (pers. comm.), Schiemenz (1987), Strübing (1960), HN
Delphax pulchellus (Curtis, 1833) E VI – M IX; egg, 1 gen. Monophagous on Phragmites australis, like the preceding species mainly near the ground and in sites which are flooded not at all or only in winter. In exceptional cases, both species are found together. The strongholds are in saltmarshes inland and along the coast, as well as in low-input meadows and pastures with only scattered reed; apparently absent from forests. Widespread in Germany but only localized, mainly near the coast and in the north German plain, but also recorded from middle parts (although restricted to valley bottoms) and the foothills of the Alps, where extending up to at least 700 m a.s.l. Niedringhaus (1991), Schaefer (1973), Schiemenz (1987), Wagner (1939a), Nickel & Achtziger (1999), Nickel & Remane (1996), Nickel (1997), HN
Euides basilinea (Germar, 1821) Delphax speciosa Boheman, 1845
B V – E VIII; nymph, 1 - 2 gen., depending on spring weather conditions (Strübing 1960). Monophagous on Phragmites australis. This is the most widespread among central European reed-dwelling Auchenorrhyncha species (both in geographical and ecological
44
Review of species
terms); along shores of lakes and ponds, in fen woods, fens and intermediate bogs, near fresh and brackish water, along the coast as well as inland. Is often overlooked due to secretive habits; thus, there are only few published data. However, it is widespread in Germany, extending to at least 800 m a.s.l. in the Bavarian Alps. Frommer (1996), Heller (1987a), Jöst (1966), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1987), Strübing (1960), Voigt (1978), Wagner (1935), Nickel & Remane (1996), HN
Chloriona unicolor (Herrich-Schäffer, 1836) Is often cited to be originally described from western Germany (e.g. Nast 1987). In fact Herrich-Schäffer (1836) only states: „Ein Weib von Wien. Mann und Weib von Zürich von Hrn. Bremy.“ The type material is lost anyway, so the identity cannot be clarified anymore. However, Fieber (1866), who was the first revisor, published illustrations of the male genital segment of several Chloriona species. His interpretation was also adopted by Asche (1982a).
So far M VI – E VIII; nymph, 2(?) gen. Monophagous on Phragmites australis. In Germany only known from three localities in inland salt marshes and calcareous spring mires in Hessen (Wetterau near Münzenberg, 150 m a.s.l., several records, between 1966 and 1993) and Thuringia (Heilingen near Orlamünde, 30.VI.1965, and Röttelmisch near Kahla, 16.VII.1968, both at c. 200 m a.s.l.). Reported from 1400 m a.s.l. in the Engadine (Switzerland). Asche (1982a), Fröhlich (1996a), Schiemenz (1987); see also Günthart (1987a), Nast (1984)
Chloriona sicula Matsumura, 1910 Chloriona flaveola Lindberg, 1948 sec. Asche (1982a)
So far E V – M VIII; nymph, 2 gen. Monophagous on Phragmites australis. In Germany at the northeastern edge of the range and only known from 5 sites along the upper course of the Rhine between Waldshut and Speyer, but probably under-recorded. The habitats in these localities included various freshwater sites (along streams, in sand pits, a quarry and a garden pond), between 100 and 400 m a.s.l. Remane & Fröhlich (1994b), HN; see also Nast (1984)
Chloriona dorsata Edwards, 1898 So far E V – M VII; nymph, 1(?) gen. Monophagous on Phragmites australis in various habitats (along rivers, oxbow lakes, ponds, fens, ditches, inland salt marshes and others). The first records for Germany have been published only recently, but the species is rather widespread at least in middle parts. Furthermore, it is recorded from the foothills of the Alps (Murnau, 620 m a.s.l.), the upper Rhine plain (Speyer), the lower part of the Elbe valley (Magdeburg, Bleckede) and eastern Holstein (Kiel). Apparently, the German localities are at the south-
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ern and eastern edge of the range; however, the taxonomic relationship with taxa described from central Asia should be checked. Remane & Fröhlich (1994b), Nickel (1994, 1999b), HN; see also Nast (1976a, 1987)
Chloriona glaucescens Fieber, 1866 M V – M IX; nymph, 2 gen. Monophagous on Phragmites australis, but largely confined to saline sites, rarely in freshwater sites with strongly fluctuating water tables and temporary salt accumulation in the upper soil layer. In Germany mainly near the coasts of the North and Baltic Sea, also in inland saltmarshes, sand pits and coal mines, mainly in Saxony-Anhalt and Thuringia, locally in southern Niedersachsen, Hessen and Saxony (Werra valley, Wetterau, near Bautzen and Boxberg); all localities are below 150 m a.s.l. Fröhlich (1997), Funke & Witsack (1998), Schaefer (1973), Schiemenz (1987), Strübing (1960), Wagner (1935, 1939a), HN
Chloriona smaragdula (Stål, 1853) E V – B VIII, according to Schiemenz (1987) until B IX; nymph, 1 - 2 gen. Monophagous on Phragmites australis, almost exclusively inland. Eurytopic in freshwater sites (even along ditches within fields), also at the edge of inland salt marshes, but largely absent from coastal salt marshes near the North and Baltic Sea. Single specimens from straw meadows are presumably vagrants. Apart from that, this species is widespread in low-lying regions of Germany; it is usually the most common among reed-dwelling Auchenorrhyncha species; in the foothills of the Alps it frequently occurs up to 800 m a.s.l. Fischer (1972), Heller (1987a), Niedringhaus (1997), Reimer (1992), Schiemenz (1987), Strübing (1960), Wagner (1935), HN
Chloriona stenoptera (Flor, 1861) So far B VI – B VII, in Poland until E VII; nymph, 1(?) gen. Monophagous on Phragmites australis; recorded only a few times in central Europe: Bialowieza (eastern Poland), 09.VI.1949, 1 †, and 21.VII.1949, 4 ††, 15 ‡‡, in an open, peaty forest of spruce, pine and birch; Baden-Württemberg: Horrheim near Stuttgart, 200 m a.s.l., 07.VI.1983, 1 †, 3 ‡‡, in a “non-flooded stand of reed“; Schopfloch near Esslingen (Swabian Jura), 750 m a.s.l., 07.VII.1975, 2 ††. There are three more recent records near the Austrian and Bavarian edge of the Alps: Pinswang am Lech (North Tyrol, Austria), 830 m a.s.l., 30.VI.1997, 1 †, in reeds along a small brooklet in a meadow; Isar floodplain near Vorderriß, 800 m a.s.l., 04.VI.1998, 3 ††, in reeds growing among willow scrub; Gröbenzell near Munich, 500 m a.s.l., June 1999. Otherwise this species is only known from Estonia, Latvia and Kazakhstan. Heller (1987a), SMNS, Weis & Schönitzer (2001), Nickel (1999a); see also Mityaev (1971), Nast (1958)
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Review of species
Chloriona vasconica Ribaut, 1934 So far mainly E V – M VII, according to Schiemenz (1987) until B IX; nymph, 1 - 2 gen. Monophagous on Phragmites australis, usually along rivers and streams, lakes, ponds, oxbow lakes, in wet ruderal sites and inland salt marshes. Widespread in the lowlands of Germany, but only scattered, recorded up to 600 m a.s.l. only. Heller (1987a), Schiemenz (1987), Wagner (1935), Nickel & Remane (1996), Nickel (1994), HN
Unkanodes excisa (Melichar, 1898) E V – B IX; nymph, 2 gen. In Europe largely confined to the coasts of the Baltic Sea; otherwise reported only from Yakutia, the Kurile Islands and Alaska. Lives on Leymus arenarius, usually on primary and white dunes. Found in Germany only locally along the Baltic Sea and in one inland site near Königs Wusterhausen. The latter record may refer to vagrant individuals or originate from a secondary plantation. Schaefer (1973), Schiemenz (1987), Wagner (1935), HN; see also Anufriev & Emelyanov (1988), Wilson (1988)
Megadelphax haglundi (J. Sahlberg, 1871) In Sweden B VI – E VII, in Austria E VIII (Ossiannilsson 1978; Nast 1986); nymph(?), 1(?) gen. There is only a single German record from the Swabian Jura: Warmberg near Oberstetten, c. 800 m a.s.l., 23.VI.1968, 1 † on a xerothermic, stony, calcareous upland with dwarf blackthorn, juniper and groups of various trees. In Europe otherwise recorded from southern Sweden, Bohemia, Moravia (both Czech Republic), Niederösterreich (Austria) and the French Massif Central; also reported from Mongolia. The host plants are unknown, but are likely to include species of Poaceae. Heller (1996); see also Anufriev & Emelyanov (1988), Nast (1987), Remane & Fröhlich (1994b)
Megadelphax sordidula (Stål, 1853) B V – E IX; nymph, 2 gen. On Arrhenatherum elatius in moderately dry to damp, occasionally also moist meadows, although in most agricultural areas nowadays largely confined to narrow strips along waysides and ditches. The species is widespread in middle and southern parts of Germany, but has only rarely been recorded in the north German plain. To the north of the Danube it occurs only at lower altitudes (usually below 500 m a.s.l.), in the Bavarian Alps it is found up to at least 800 m a.s.l. Is not known from Denmark and Britain, but common in southern and middle parts of Sweden and Finland. The species is reported from there to invade wheat and oat fields and to be a potential vector of Phleum green stripe virus (PGSV). Achtziger (1991), Fischer (1972), Günthart (1988), Marchand (1953), Müller (1978), Reimer (1992), Schiemenz (1987), Schwoerbel (1957), Wagner (1935, 1951a), Nickel & Achtziger (1999), HN; see also Ossiannilsson (1978), Raatikainen (1960, 1970a)
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Laodelphax striatella (Fallén, 1826) E IV – B X; nymph, 2 gen. This species lives in unstable populations in dry to moist, usually anthropogenically affected sites (mainly fallow fields, fertilized meadows, waysides) on various grasses, including cereal crops, perhaps also on Cyperaceae and Juncaceae. The bulk of individuals is macropterous and highly vagile. In Germany continuously reproducing populations with a higher proportion of brachypters are largely confined to rather warm lowlands in the southern half (mainly to the south of the Main, perhaps also in Thuringia and Saxony). Macropterous individuals may occur in larger numbers, but with much varying reproductive success, more or less in the whole country, with single individuals also in the north German plain and the higher parts of the Mittelgebirge and the Alps, were they may be found up to 1800 m a.s.l. The species is reported to be noxious to cereal crops (including rice) by transmission of various diseases, notably Maize rough dwarf virus (MRDV) and Barley yellow striate mosaic virus (BYSMV) in warmer countries. In most parts of central Europe, however, the species is probably too scarce to have an economic impact. Fischer (1972), Günthart (1988), Marchand (1953), Niedringhaus & Olthoff (1993), Remane (1987), Schiemenz (1971a, 1975), Schwoerbel (1957), Wagner (1935), Nickel & Achtziger (1999), Nickel (1997), HN; see also Brˇcák (1979), Conti (1988)
Ditropsis flavipes (Signoret, 1865) B V – B X; nymph, 2 gen. Rather widespread and locally common in the Bavarian Alps, the Allgäu and the Swabian and Franconian Jura; in sunny, moderately or temporarily dry pastures and meadows, scattered and less abundant in base-rich and increasingly dry grassland habitats of lower altitudes towards the north. The host plant is Bromus erectus. The northernmost localities, some of which appear to be rather isolated, are on a line from Mechernich (northern Eifel Mountains) to Lorch (Rheingau), Bad Wildungen, Bad Frankenhausen and Eberswalde. Found up to 1000 m a.s.l. on the Swabian Jura and in the Bavarian Alps, up to 1400 m a.s.l. in Vorarlberg (Austria). Bornholdt (1991), Fischer (1972), Remane & Fröhlich (1994b), Rombach (1999a), Schiemenz (1987), Wagner (1939a), Nickel & Achtziger (1999), HN; see also Moosbrugger (1946)
Paraliburnia adela (Flor, 1861) M V – B IX; nymph, (1-?) 2 gen. Monophagous on Phalaris arundinacea, usually in temporarily flooded, rather eutrophic, sunny to shady sites, notably along rivers, streams and ditches, in willow-poplar floodplain woods and in low-input meadows. Lives near the base of the host plant and is rarely caught with the sweep-net. In the older literature Glyceria has been reported as host plant, which is caused by confusion of ‡‡ with those of Struebingianella lugubrina (Boh.). There are only few published records; however, P. adela (Fl.) is widespread in
48
Review of species
Germany, fairly common in most parts and can be easily found when searching directly on the ground. Extends up to at least 800 m a.s.l. in the foothills of the Alps. Fröhlich (1996a), Kirschbaum (1868), Remane (1962), Schiemenz (1987), Nickel & Achtziger (1999), Nickel & Remane (1996), HN; see also Lauterer (1980)
Paraliburnia clypealis (J. Sahlberg, 1871) E V – B IX; nymph, (1-?) 2 gen. This is an essentially north European species, which is known in central Europe only from scattered records in the Netherlands, the Czech Republic and the northern half of Germany. There are more or less isolated localities in the north German plain, the Harz Mountains (Clausthal-Zellerfeld, 550 m a.s.l.), as well as middle Hessen (Schweinsberg and Großenmoor, up to 300 m a.s.l.). Lives in wet, usually acidic and peaty, sunny to shady sites (mainly fen woods, intermediate bogs and fens) in tall stands of Calamagrostis canescens; but there are also host reports of Rhynchospora and Eriophorum from the Netherlands. Is perhaps under-recorded due to secretive habits and clumped occurrence. Niedringhaus & Olthoff (1993), Remane (1962), Remane & Fröhlich (1994b), Schiemenz (1987), Nickel & Sander (1996), HN; see also Bieman & Booy (1984), Dlabola (1970)
Hyledelphax elegantula (Boheman, 1847) E IV – E IX; nymph, 2 gen., perhaps 1 at higher altitudes. In the grass layer of deciduous and coniferous forests in moderately dry to moist sites, also under alleys and even solitary trees and shrubs. Rarely and in low numbers in treeless heaths and bogs, but presumably only as vagrant. Lives on various grasses, notably Calamagrostis arundinacea, Brachypodium pinnatum, Holcus mollis, Molinia caerulea, Deschampsia flexuosa, in the lowlands of southern and middle parts of Germany, but at higher altitudes and in the north German plain, it is largely confined to the latter species. Widespread in Germany, found up to at least 1300 m a.s.l. in the Mittelgebirge and the Bavarian Alps, up to the subalpine belt in Austria and Switzerland. Bittner & Remane (1977), Fischer (1972), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1987), Wagner (1935, 1939a), HN and others; see also Günthart (1987a), Wagner & Franz (1961)
Megamelodes lequesnei W. Wagner, 1963 Was misinterpreted by Scott (1870) and since then erroneously published as M. quadrimaculatus (Sign.) – see below. Much later Wagner (1963), who had revised Signoret’s types, discovered this error and created a nomen novum (in a footnote!). Hence, older records of M. quadrimaculatus (Sign.) are in need of revision.
In Britain VIII – III (Le Quesne 1960); adult(?), 1 gen. This is a species with insufficiently known biology and distribution, which was recorded from Germany only from a single site in Holstein: Vollstedter See near Rendsburg, November 1970, 1 † in a pitfall trap in a bog. In Britain it is reported to live mainly in
Delphacinae Leach, 1815
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base-rich fens and has been suspected to be associated with Juncus subnodulosus; in France it is widespread and lives in fens with dominating tall sedges. Remane (pers. comm.), Remane & Fröhlich (1994b); see also Kirby (1992)
Megamelodes quadrimaculatus (Signoret, 1865) Liburnia fieberi Scott, 1870
Adults I – XII, mainly VIII – VI; adult, 1 gen. In wet to temporarily wet, mesotrophic to moderately eutrophic, neutral to basic sites, mainly spring mires, fens and wet meadows, but also inland salt marshes and overgrown sand pits. The host relationships are not well known. Adults are often found in the vicinity of rushes (Juncus inflexus and J. articulatus) and sedges (Carex spp.), and the species has also been published from grasses (notably Poa spp.) and dicotyledons. However, the few definite host records (of large numbers of adults and nymphs) are from mixed stands of non-flowering and thus inconspicuous Festuca pratensis and F. rubra. In Germany this species is found only sporadically, at the edge of the range, and often in very small pockets only, at least up to 500 m a.s.l. Northernmost localities are near Bonn, Herne, Hanover, Hamburg, Neumünster, Rostock and Frankfurt an der Oder. It is not found in northern Europe and Poland; from Denmark there is only a single record from the 19th century. It is probably often overlooked due to its specific habitat requirements, the late appearance and epigeic habits. Köhler (pers. comm.), Maczey (pers. comm.), Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Schiemenz (1987), Wagner (1935, and unpublished data), Nickel & Achtziger (1999), Nickel (1997, 1999b), HN; see also Bieman & Booy (1984), Nast (1987), Ossiannilsson (1978)
Calligypona reyi (Fieber, 1866) Mainly E VI – M IX, at least in favourable years M V; nymph(?), 1 - 2 gen. Usually in permanently flooded sites near fresh or moderately saline, usually standing water, rarely also in temporarily dry sites or near running water; mostly along shores of lakes and ponds, occasionally in sand and gravel pits, saltmarshes and garden ponds. Host plants are Schoenoplectus lacustris and Sch. tabernaemontani. Until recently there were only a few published records from Germany (mainly near the coasts of the North and Baltic Sea, as well as near Waren an der Müritz, Leipzig, Gera and Frankfurt am Main). However, the species is widespread all over the country and extends at least up to 730 m a.s.l. in the Allgäu and Upper Bavaria. May be fairly abundant in larger stands of host plants. Harz (1988), Remane & Fröhlich (1994b), Schiemenz (1987), Wagner (1939a), Nickel & Remane (1996), Nickel (1997), HN
Mirabella albifrons (Fieber, 1879) B IV – E IX; nymph, 2 gen.
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Review of species
On Calamagrostis epigejos in sunny to slightly shady, moderately dry to moderately moist sites, mainly in forest clearings, ruderal sites, sand and gravel pits, mining areas and on railway embankments; rarely in rather moist sites on C. canescens. Widespread in southern and eastern parts of Germany and common in favourable sites, but at the edge of the range. Northern- and westernmost localities are on a line from Marburg to Göttingen, Braunschweig, Uelzen and Greifswald. However, the species is also reported from the Netherlands and the Paris basin. The highest localities are at c. 650 m a.s.l. in Upper Bavaria. Fischer (1972), Remane & Fröhlich (1994b), Schiemenz (1987), Walter (pers. comm.), Nickel (1994), HN; for identification see Dlabola (1954)
Delphacodes capnodes (Scott, 1870) Adults I – XII, mainly E VIII – E VI; adult, 1 gen. In tall-sedge swamps, reeds (also in moderately saline sites) and intermediate bogs, the latter usually with open stands of birch or pine. It lives close to the ground, often deep in hummocks of Sphagnum and is rarely collected; most specimens are macropterous individuals on dispersal flight. Host plants are sedges and cottongrasses. In fens it is found on tall sedges (Carex acutiformis, C. acuta and C. riparia are reported from the Czech Republic and Slovakia); in peaty sites of Germany and the Netherlands it is apparently associated with Eriophorum angustifolium. Populations in fenland habitats consist mainly or even exclusively of ‡‡, the biology of which is poorly known. Pseudogamous, triploid ‡‡ from a Greek population have been found to respond to acoustic signals of Dutch †† of D. capnodes and were also inseminated, but without passing the male genome to their offspring. In Germany this species appears to be rare and rather localized although it is widespread; the highest localities are at 600 m a.s.l. only. Fröhlich (1996a), Funke & Witsack (1998), Niedringhaus & Olthoff (1993), Remane (1958), Schiemenz (1987), Strübing (1955), Wagner (1938b, 1940a), Nickel (1999b), HN; see also den Bieman & de Vrijer (1987), Lauterer (1980)
Delphacodes venosus (Germar, 1830) Adults I – XII, mainly B IX – E VII; adult, 1 gen. In tall grass stands in moderately shady to sunny, peaty to moist, sometimes also damp or moderately dry, usually cool sites, mainly low-input meadows and pastures, fens, intermediate bogs and fen woods. The host plants include Deschampsia cespitosa, Festuca arundinacea, Agrostis spp., Molinia caerulea and other grasses, and probably sedges (Carex spp.). Has repeatedly been considered as tyrphophilous or even tyrphobiotic. However, in central Europe most populations are clearly found in moderately eutrophic, moist to temporarily wet sites. The species is widespread in Germany and fairly common, but often overlooked. Most specimens are caught in pitfall traps. Frequently found up to at least 1000 m a.s.l. in the Mittelgebirge and the Bavarian Alps. Fischer (1972), Schiemenz (1987), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN; see also Lauterer (1980), Ossiannilsson (1978)
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Gravesteiniella boldi (Scott, 1870) B V – E X; nymph, 2 gen. On Ammophila arenaria on white and grey dunes near the coast, occasionally also on open inland dunes of the north German plain. In Germany locally common on the East and North Frisian Islands, Heligoland and along the Baltic Sea coast, also found inland near Hamburg and Dömitz. Is also reported from eastern parts of Austria, where A. arenaria is absent. Niedringhaus (1991, 1994), Schaefer (1973), Schiemenz (1987), Wagner (1935), HN; see also Holzinger & Remane (1994), Ossiannilsson (1978)
Muellerianella brevipennis (Boheman, 1847) E VI – E X; egg, 1 - 2 gen. (see Drosopoulos 1977, Witsack 1971). On Deschampsia cespitosa in sunny to shady, moderately moist to wet, usually temporarily waterlogged sites, mainly meadows, pastures, abandoned grassland, fens, clearings, open forests, also forest roads. In Germany widespread and in intermediate to high frequency on the host, also at higher altitudes of the Mittelgebirge, found up to at least 1500 m a.s.l. in the Allgäu Alps. Fischer (1972), Nikusch (1976), Reimer (1992), Schiemenz (1987), Wagner (1939a), Nickel & Achtziger (1999), HN
Muellerianella fairmairei (Perris, 1857) Was formerly considered to be conspecific with the following species, until Booij (1981) demonstrated that they are clearly distinct. Thus, most older records are in need of revision. The complicated biology was studied by Drosopoulos (1977), Booij (1981, 1982) and Booij & Guldemond (1984).
E VI – E X; egg, 2 gen., probably 1 at higher altitudes. In wet to damp, usually cool and moderately eutrophic sites, mainly meadows, pastures and forest glades. Food plants are Holcus lanatus and H. mollis. At least the former is also the oviposition plant under long-day conditions. Under short-day conditions, overwintering eggs are laid exclusively on Juncus effusus, which is left, however, immediately after hatching. The species reaches the eastern edge of its range in Germany where it is restricted to oceanic climates of the western and northwestern parts. In the Mittelgebirge reproducing populations were found up to 1000 m a.s.l. Easternmost localities are near Lörrach, Todtnau (Black Forest), Emmendingen, Bad Nauheim, Laasphe, Brilon, Hannoversch Münden, Lüneburg, Boizenburg and Rendsburg, as well as in Jutland (Denmark) and southern Sweden. At present, the species is expanding its range and has apparently colonized most parts of eastern Germany. Recently it has been recorded in numerous sites where it was absent some years ago, e.g. near Gardelegen, Northeim, Göttingen, Brandenburg, Schwedt (Oder) and Pirna. Populations near the edge of the range may include up to 100% of triploid, pseudogamous ‡‡. Booij (1982), Booij & Guldemond (1984), Drosopoulos (pers. comm.), Fröhlich (1996a), K. Heller (pers. comm.), Niedringhaus & Olthoff (1993), Rothenbücher (pers. comm.), Walter (pers. comm.), Nickel & Achtziger (1999), Nickel (1997), HN
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Review of species
Muellerianella extrusa (Scott, 1871) Was formerly considered to be conspecific with the preceding species (e.g. by Ossiannilsson 1978) or confused with it (see above). Older records of M. fairmairei (Perr.), mainly those from the central and eastern parts of central Europe, usually refer to M. extrusa (Scott).
E VI – E X; egg, according to Schiemenz (1987) occasionally adult, 1 gen. On Molinia caerulea in peaty to moist, also temporarily moist sites, mainly straw meadows, margins and degenerated stages of bogs and open forests. Recorded from most parts of Germany, rather wide and densely spread in the Alps and their foothills and the Mittelgebirge, but more scattered in the lowlands, where the species has probably much decreased due to drainage, fertilizing and ploughing of former habitats. Occurs up to at least 1000 m a.s.l. in the Mittelgebirge, to at least 1200 m a.s.l. in the Allgäu Alps. Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1987), Nickel & Achtziger (1999), Nickel & Remane (1996), HN; for identification see Booij (1981), Ossianilsson (1983)
Chlorionidea flava P. Löw, 1885 B V – B IX; nymph, 1 - 2 gen. On Sesleria albicans, usually in open pine forests in dry to temporarily dry and basic sites on sun-exposed calcareous and gypsum hillsides; in the foothills of the Alps also on rather dry riverine gravel banks, at higher altitudes locally in open pastures. In Germany at the northwestern edge of the range and known from Thuringia (basins of Saale, Unstrut, Ilm and Werra), Mainfranken, the Franconian and Swabian Jura and the Alps and their foothills, also found near Eschwege and Göttingen. Highest localities are at c. 1000 m a.s.l. in the Swabian Jura and at c. 1300 m a.s.l. in the Bavarian and Allgäu Alps, but the species is reported from the alpine belt in Austria. Haupt (1925), Remane & Fröhlich (1994b), Schiemenz (1987), Nickel (1994), HN; see also Wagner & Franz (1961)
Litemixia pulchripennis Asche, 1980 B VIII – B X; egg, 1 gen. This striking species was described only 20 years ago from western France. It lives deep in tussocks of Molinia caerulea in open pine forests on temporarily moist, acidic and sandy substrates. More recently it was found in three sites in the northern upper Rhine plain (near Speyer, Germersheim and Neustadt an der Weinstraße, all at c. 100 m a.s.l.). Remane & Fröhlich (1994b), HN; see also Asche (1980), della Giustina & Remane (1992), Remane & della Giustina (1991)
Muirodelphax aubei (Perris, 1857) E IV – B X; nymph, 2 gen. In grass stands in rather dry, sunny, often disturbed sites usually with some bare ground, for instance on open sand of inland and tertiary coastal dunes, in military training areas, dry abandoned fields, in mining areas, etc., also on strongly grazed calcareous hill-
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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 below 400 m a.s.l., one locality in the foothills of the Alps is at c. 800 m a.s.l. Fischer (1972), Niedringhaus (1991), Nikusch (1976), Remane (1987), Schiemenz (1987), Schwoerbel (1957), Trümbach (1959), Wagner (1935, 1939a, 1951a), HN
Acanthodelphax denticauda (Boheman, 1845) M IV – B X; nymph, 2 gen. On Deschampsia cespitosa in moist to wet meadows, pastures and abandoned grassland, locally also in forests. Widespread in Germany and in intermediate frequency on the host, although often underrepresented in sweep-net samples. Found up to at least 850 m a.s.l. in the Mittelgebirge and the foothills of the Alps. Bittner & Remane (1977), Fischer (1972), Niedringhaus & Olthoff (1993), Schiemenz (1987), Schwoerbel (1957), Wagner (1935, 1939a), Wagner & Franz (1961), Nickel & Achtziger (1999), Nickel (1997), HN
Acanthodelphax spinosa (Fieber, 1866) M IV – M X; nymph, 2 gen., probably 1 at higher altitudes. On Festuca rubra and, less frequently, F. ovina in sunny, damp to moist, occasionally also peaty to dry sites (mainly oligotrophic or wet meadows, fens, dry grassland, ruderal sites, waysides, etc.). The species is largely absent from intensively managed meadows, which may be invaded by single macropterous individuals on dispersal flight, but without long-termed reproductive success. It is widespread in Germany and common in many places, although more scattered in northern parts, reaching the edge of the range in Holstein (Heide and Lübeck). Occurs up to at least 1500 m a.s.l. in the Mittelgebirge and the Bavarian Alps, up to c. 2000 m a.s.l. in North Tyrol and Styria (both Austria). Bittner & Remane (1977), Fischer (1972), Marchand (1953), Nikusch (1976), Reimer (1992), Schiemenz (1987), Wagner (1935), Nickel & Achtziger (1999), HN; see also Leising (1977), Wagner & Franz (1961)
Nothodelphax albocarinata (Stål, 1858) Was not distinguished from N. distincta (Fl.) until Kontkanen (1952) clarified the identity of both species. The majority of the older central European records, for instance Haupt (1935) and Wagner (1948b), refer to N. distincta (Fl.).
B V – B IX; nymph, 2 gen. This is a rare and tyrphobiotic species, occurring in central Europe only in very isolated sites near the south-western edge of the range. Found along hollows in raised and intermediate bogs; the host plant is probably Carex limosa. In Germany there are only 12 confirmed localities between Halle, Eilenburg, Bautzen, Weißwasser and Luckau, as
54
Review of species
well as in Upper Bavaria (Sachsenkam and Seeshaupt), up to at most 700 m a.s.l. To the west of these, there is only a single specimen from the French Jura and a more recent record from near Apeldoorn (Netherlands). Remane & Fröhlich (1994b), Schiemenz (1976, 1987), Walter (pers. comm.), HN; see also Bieman & Rozeboom (1993), Remane & della Giustina (1993)
Nothodelphax distincta (Flor, 1861) M IV – B IX, at higher altitudes mainly V – VII; nymph, 1 - 2 gen. Tyrphobiotic in open raised bogs on Eriophorum vaginatum. In Germany occurring rather scattered in the remnant bog areas of the north German plain, the highest parts of the Mittelgebirge (mainly Solling, Harz Mountains, Thuringian Forest, Erzgebirge, Rhön Mountains, Black Forest) and the foothills of the Alps. The highest localities are at 900 m a.s.l. Bittner & Remane (1977), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1958), Schiemenz (1971a, 1975, 1976, 1987), Wagner (1948b), Nickel & Remane (1996), Nickel (1997, 2002), HN
Dicranotropis hamata (Boheman, 1847) M IV – E X; nymph, 2 gen., according to Schiemenz (1987) only 1 at higher altitudes. Eurytopic on various grasses in moist to moderately dry, sunny to moderately shady, often disturbed sites (mainly waysides, ruderal sites, forest clearings, abandoned fields, occasionally also fertilized meadows and pastures). Host plants are Elymus repens, Dactylis glomerata, Holcus lanatus, H. mollis, Festuca arundinacea and other grasses. Widespread and common in Germany, extending to at least c. 1100 m a.s.l. in the Mittelgebirge and the Alps. Is reported from Scandinavia and former Czechoslovakia as a potential vector of cereal virus diseases, but is unlikely to cause major damage at least in central Europe. Achtziger (1991), Fischer (1972), Niedringhaus (1991), Remane (1987), Schiemenz (1987), Wagner (1935), Nickel & Achtziger (1999), HN and others; see also Brˇcák (1979), Lindsten (1979)
Dicranotropis divergens Kirschbaum, 1868 M V – B X; nymph, 1 gen. In oligotrophic to moderately eutrophic meadows and pastures of higher altitudes in moderately dry to moist, occasionally also wet sites. Lives on Festuca rubra, perhaps also on additional grasses. In Germany particularly widespread and common in the Bavarian and Allgäu Alps and their foothills up to at least 1800 m a.s.l. (but found up to 2200 m a.s.l. in Austria and Switzerland). Also occurring in most of the higher Mittelgebirge: Bavarian Forest, Oberpfälzer Wald, Fichtelgebirge, Frankenwald, Rhön Mountains, Thuringian Forest, Erzgebirge, as well as Black Forest, Swabian Jura and Harz Mountains, usually above c. 600 m a.s.l., locally down to 400 m a.s.l. Repeated records of single brachypters in lowland areas are probably caused by passive drift or shortterm reproduction of vagrants. Achtziger (1991), Bittner & Remane (1977), Kupka (1941), Reimer (1992), Schiemenz (1987), Wagner (1958), Nickel & Achtziger (1999), HN; see also Günthart (1987a), Leising (1977)
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Dicranotropis montana (Horváth, 1897) Dicranotropis gratiosa Dlabola, 1997 (see Holzinger 1999a)
In Austria B VI – E VIII (Dlabola 1997; Moosbrugger 1946); nymph, 1 gen. In Europe only known from a few records from the higher parts of the Carpathians (Romania) and Alps (Austria, northern Italy, Bavaria). In Germany the only locality is in the Allgäu: Rotspitze above Hindelang, 1800 m a.s.l., 04.VIII.1995, 7 ††, 6 ‡‡ on Deschampsia cespitosa in a subalpine pasture close to a brooklet. Is probably more widespread in the German Alps and has been found near Scharnitz (North Tyrol, close to the Bavarian border). Was recorded in the Dolomites (Italy) from grasses in an open coniferous forest. Nickel (1999b); see also Asche (1982b), Remane & Fröhlich (1994b); for identification see Holzinger (1999a)
Florodelphax leptosoma (Flor, 1861) E IV – M X; nymph, 2 gen., probably 1 at higher altitudes. On rushes (Juncus articulatus, J. effusus, J. subnodulosus, J. conglomeratus and others) in moderately wet to temporarily flooded, sunny to slightly shady sites, mainly low-input meadows, spring mires and intermediate bogs, locally also mining areas and sand pits. Wide and densely spread in Germany, but often only in small populations, extending up to at least 1100 m a.s.l. in the Mittelgebirge and the Alps, but found at 1500 m a.s.l. in North Tyrol (Austria). Fischer (1972), Nikusch (1976), Niedringhaus (1991), Reimer (1992), Schiemenz (1987), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN and others
Florodelphax paryphasma (Flor, 1861) M V – E IX; nymph, 2 gen. On Carex disticha in moderately eutrophic and temporarily waterlogged or flooded sites (mainly wet meadows, fens, spring mires, floodplain depressions, shores of ponds and lakes, etc.). In Germany the species has often been overlooked and was until recently only known from very few localities mainly in the northern plain. However, according to more recent data, it is also widespread in the middle parts where it often occurs in high frequency and individual numbers on the host. At present, southernmost localities are known from the vicinity of Speyer, Ludwigsburg, Ansbach, Ilmenau and Dresden, only up to 450 m a.s.l. Has been recorded in central France, the Pyrenees and the Sierra de Cuenca of central Spain, but not yet in Austria, Italy and Switzerland. Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Schiemenz (1987), SMNS, Wagner (1952a), Walter (pers. comm.), Nickel & Achtziger (1999), HN; see also della Giustina & Remane (2001)
Kosswigianella exigua (Boheman, 1847) M III – B X; nymph, 2 gen. In sunny, oligotrophic and low-vegetated, often grazed grasslands in moderately dry to dry, occasionally also damp or very dry sites on various substrates (both sandy to solid,
56
Review of species
basic to acidic). Host plants are fine-leaved species of fescue (Festuca ovina group), although additional grasses (Corynephorus canescens, Deschampsia flexuosa) have been reported. Widespread in Germany and locally abundant, extending up to at least 1100 m a.s.l. in the Mittelgebirge and the Alps. Fischer (1972), Niedringhaus (1991), Remane (1987), Rombach (1999b), Schiemenz (1987), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN and others
Scottianella dalei (Scott, 1870) Delphacodes gravesteini Dlabola, 1971 sec. Anufriev (1980)
In England according to Waloff & Solomon (1973) IV – VIII; nymph, 2 gen. Occurs from Morocco through western Europe northward to England, known from Germany only from two records of single, brachypterous †† in the extreme west: Wahner Heide near Cologne, c. 50 m a.s.l., 08.VIII.1982, and Dudenhofen near Speyer, c. 110 m a.s.l., 25.IV.1989. Was found among grasses in moist sites in France („around ponds, lakes, marshes, etc.“), on Agrostis capillaris in England. Bieman & Booy (1984), Remane (1995); see also della Giustina (pers. comm.), Waloff & Solomon (1973)
Struebingianella lugubrina (Boheman, 1847) E IV – M IX; nymph, 2 gen. On species of sweet-grass (Glyceria maxima, G. fluitans group) in wet, usually temporarily flooded, rather eutrophic, sunny to slightly shady sites, mainly near standing and running water (along rivers, streams, ditches, in floodplain depressions), in wet meadows, alder fens, as well as on temporarily dry bottoms of ponds. Widespread and fairly common in the lowlands of Germany, in intermediate to high frequency on both host species; rather scattered at higher altitudes and extending up to at least 900 m a.s.l. Fischer (1972), Niedringhaus (1997), Reimer (1992), Schiemenz (1987), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), Nickel (1997), HN
Xanthodelphax flaveola (Flor, 1861) B VI – B IX; nymph, 1(?) gen. In sunny to slightly shady grass stands in moderately acidic, moderately dry to moist sites, mainly low-productivity pastures and meadows, usually on loamy or sandy substrates. The host plant is Poa pratensis. In Germany rather localized and uncommon, mainly in the eastern parts of the north German plain and the Mittelgebirge (Harz Mountains, Eichsfeld, Eifel Mountains, Rhön Mountains, Thuringian Forest, Erzgebirge, Oberpfälzer Wald, Donnersberg, Swabian Jura, Black Forest), as well as in the foothills of the Alps, extending up to at least 1200 m a.s.l. Has apparently declined due to intensification of grassland management. It is not reported from Switzerland; from France it is only known from a few localities in the southern half.
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Fischer (1972), Fröhlich (1996a), Nikusch (1976), Schiemenz (1987), Walter (1998), Wagner (1939a), HN; see also della Giustina & Remane (1999)
Xanthodelphax straminea (Stål, 1858) B V – M IX; nymph, 2 gen. In sunny to moderately shady, wet to moderately dry, acidic to neutral, oligotrophic to moderately eutrophic sites, mainly in spring mires, intermediate bogs and wet meadows, usually on sandy or peaty soils, locally also in moderately saline sites; on bentgrass (Agrostis canina, A. stolonifera, A. capillaris.). Widespread in Germany, although scattered; more common in the sand and bog areas in the northeastern half, but rather localized in the southwest, where recorded from Upper Bavaria, Oberpfalz, Mainfranken and the upper Rhine between Lörrach and Wiesbaden. Highest localities are at 850 m a.s.l. in the Thuringian Forest and the foothills of the Alps. Bittner & Remane (1977), Günthart (1988), Schiemenz (1971a, 1987), Wagner (1939a), Nickel & Achtziger (1999), Nickel (1999b), HN
Xanthodelphax xantha Vilbaste, 1965 So far E VI – B VIII; nymph, 1(?) gen. Described from the Altai and Poland, in Germany found only in two localities: Gauting near Munich, c. 580 m a.s.l., 1959 and 1961, in a clearing within a spruce forest, and Serrahn near Neustrelitz, c. 70 m a.s.l., 1973 and 1974, along the edge of an intermediate bog. Possible host plants in both sites are species of Calamagrostis (C. canescens or C. epigejos) and Molinia caerulea. Otherwise known in Europe only to occur in Latvia, Belorussia and north-eastern Poland. Remane & Fröhlich (1994b), Schiemenz (1976); see also Nast (1966), Vilbaste (1965)
Paradelphacodes paludosa (Flor, 1861) E IV – B IX; nymph, 1 - 2 gen. In wet, usually peaty, acidic as well as basic sites, mainly intermediate bogs, spring mires and fens. Host plants are sedges, probably Carex rostrata on acidic, C. panicea on more basic substrates. Widespread all over Germany, but scattered, with strongholds in the bog areas of the north German plain and along the edge of the Alps, rather uncommon elsewhere. The highest sites are at 1200 m a.s.l. in the Allgäu Alps. Fröhlich (1996a), Marchand (1953), Remane (1958), Schiemenz (1987), Wagner (1946), Nickel & Achtziger (1999), Nickel & Remane (1996), HN
Oncodelphax pullula (Boheman, 1852) M V – M VIII; nymph, 1 gen. In sedge swamps of wet to peaty, acidic as well as basic sites, mainly intermediate bogs, calcareous small-sedge fens and birch swamps. The species lives on sedges, mainly –
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Review of species
perhaps exclusively – on Carex nigra. It is widespread in Germany, but only very scattered, highest localities are at 800 m a.s.l. in the Bavarian Alps. Fischer (1972), Hoffmann (1980), Niedringhaus & Olthoff (1993), Remane (1962), Remane & Fröhlich (1994b), Walter (1998), Nickel & Achtziger (1999), HN; see also Lauterer (1978)
Criomorphus albomarginatus Curtis, 1833 B V – E VII; nymph, 1 gen. In wet to damp, sunny to shady sites, mainly low-input meadows, abandoned grassland, open forests, fens, waysides, ditches, forest margins, etc., avoiding high-intensity grassland. Host plants are various species of grasses (Festuca rubra, Deschampsia cespitosa, D. flexuosa, Calamagrostis arundinacea and others), a few records are also from Juncaceae (Luzula spp.). The species is widespread in the lowlands of Germany and locally common; extending to at least 1100 m a.s.l. in the Mittelgebirge, up to 1500 m a.s.l. in the Bavarian Alps. Bornholdt (1996), Haas (1975), Niedringhaus (1991), Nikusch (1976), Remane (pers. comm.), Schiemenz (1987), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN and others
Criomorphus borealis (J. Sahlberg, 1871) B V – E VII; nymph, 1 gen. This is a Siberian species, its range extending through northern Germany westward to the Netherlands and through the Erzgebirge, the Thuringian and Bohemian Forest to the Harz Mountains and eastern Bavaria. In the north German plain it lives only locally on Calamagrostis canescens in intermediate bogs, fens, and fen woods (found near Pinneberg, Hamburg, Sassnitz/Rügen, Neustrelitz, Berlin and Niesky). In the Harz Mountains, Thuringian Forest and Erzgebirge, however, it is common in many places and often dominant in stands of Calamagrostis villosa in spruce forests and in clearings mainly of the submontane belt (between 350 and 1000 m a.s.l.). Niedringhaus & Olthoff (1993), Schiemenz (1976, 1987), Strübing (1960), Remane & Fröhlich (1994b), HN; see also Bieman & Booy (1984), Novotný (1995)
Criomorphus moestus (Boheman, 1847) So far E V – B VII; nymph, 1 gen. Largely confined to peatlands of northern Europe. From Germany there are several published records most of which are based on misinterpretations. The only confirmed records are from two intermediate bogs near Berlin, both at 50 m a.s.l.: Teufelsmoor near Müggelheim, a population among Calamagrostis canescens in 1951 and 52, and Maggelhansluch near Zernsdorf, 30.V.2002, 2 ††, 2 ‡‡). C. canescens is also reported as host plant from Sweden, but the second German site was a mixed stand of Carex, Phragmites and Eriophorum vaginatum without Calamagrostis. Moreover, most recently studied stands of C. canescens in northern Germany were infested by C. borealis (J. Shlb.) only. Strübing (1955), Coll. Maczey (vid. HN); see also Ossiannilsson (1944), Vilbaste (1960), Walter & Emmrich (1995), Nickel & Sander (1996)
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Criomorphus williamsi China, 1939 So far E V – B VII, in England according to Hassan (1939) B V – M VII; nymph, 1 gen. Described from England, elsewhere in Europe only recorded from Moravia (Czech Republic), Hungary, middle Russia and northern Germany, where known from only three localities (all below 50 m a.s.l., all specimens brachypterous): Bremen, Stadtwald, 1982, 1 †, in an open oak-birch forest; Hamburg, Vier- und Marschlande, 01.VII.1982 and 20.VI.1983, 1 †, respectively, in abandoned fields; Amt Neuhaus near Lüneburg: Zeetze, 30.V.2000 and 07.VI.2002, a large population, in lush grass stands along a ditch, perhaps on Poa palustris or Poa trivialis. The only record from Moravia (1 brachypterous †) is from an open forest. The species is reported to live on grasses usually in moist sites in England, in meadows with Rumex in middle Russia, in floodplain meadows in Kazakhstan. Is probably more widespread at least in lowland central Europe. Hildebrandt (1986), Tharsen (1987), ZIMH, HN; see also Anufriev & Kirillova (1998), Emelyanov (pers. comm.), Hassan (1939), Lauterer (1983), Le Quesne (1960), Nast (1987), Orosz (1983)
Pseudodelphacodes flaviceps (Fieber, 1866) Specimens of Calligypona tricolorata Dlabola, 1961a and Delphacodes fenestratus Emelyanov, 1972 from central Asia have been kindly sent by A.F. Emelyanov (St. Petersburg) and exchanged for Ps. flaviceps (Fieb.) from the Bavarian Alps. All turned out to be at least congeneric. Nomenclatural changes will be published elsewhere.
So far B VI – E VIII, according to Fischer (1972) also M IV; nymph, 2 gen. On sparsely vegetated, often almost bare sand and gravel banks of unchanneled rivers in the Alps and their foothills. In Germany very rare and only found along the Lech (near Augsburg, 1936, i.e. before drainage) and the upper Isar: 1961 and 1994 single macropterous †† near Ascholding and Wallgau, from 1996 to 1998 several records (in pure stands of Calamagrostis pseudophragmites and including brachypterous individuals) in altogether 4 sites between Krün and Fall as well as along the Rißbach near Vorderriß. All localities are between 500 and 900 m a.s.l. Otherwise, in Europe only known from old records from the Rhine floodplains in Vorarlberg (Austria) and Switzerland (probably Rhine valley, too). Another record from the vicinity of Trieste (now Italy, but erroneously cited from former Yugoslavia) is in need of verification. Fischer (1972), Fröhlich (1996b), Remane (pers. comm.), Nickel (1999a); see also Melichar (1899), Moosbrugger (1946), Nast (1987)
Toya propinqua (Fieber, 1866) So far VIII – B X; nymph, 2(?) gen. This is a pantropical species, in Europe largely confined to southern parts. Its occurrence to the north of the Alps is perhaps only erratical. It lives on Cynodon dactylon and perhaps other grasses usually in disturbed sites. Macropterous individuals on dispersal flight can be drifted over long distances. Found in Germany only in the northern upper Rhine plain (Mainz, Darmstadt and Speyer) and in the Lech floodplains near Landsberg (the latter probably stray individuals). A published record from the surroundings of Berlin is based
60
Review of species
on misidentified specimens of Paraliburnia clypealis (J. Shlb.). In urban areas of Mainz the species was found in 1993 through 1995, but not in 1996 and 1997, with brachypterous individuals at least in 1994. Was recently reported from the French and Swiss side of the southern upper Rhine plain near Colmar and in urban areas of Basel. Fischer (1972), Remane (1987), Strübing (1955, and pers. comm.), Thüs (pers. comm), Wonn (1956); see also della Giustina & Balasse (1999); Drosopoulos (1982), Mühlethaler (2001), Nast (1975), Wagner (1954)
Javesella discolor (Boheman, 1847) In the lowlands mainly B V – M VII, at higher altitudes VI – IX; nymph, 1 gen. Lives in the undergrowth of deciduous and coniferous forests, in shady forest glades, at the edges of bogs (invading into the interior along ditches) and in subalpine pastures and scrub. Host plants are mainly grasses (Deschampsia cespitosa, D. flexuosa, Calamagrostis canescens and others), probably also various species of Cyperaceae and Juncaceae (Carex, Scirpus, Juncus, Luzula). Widespread in Germany and particularly common in the submontane and montane belt (in the Bavarian Alps and the Allgäu frequently extending up to at least 1800 m a.s.l.); less common in the lowlands, locally absent from warm regions. Fischer (1972), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1971a, 1975, 1987), Schwoerbel (1957), Wagner (1935, 1939a), HN; see also Novotný (1995), de Vrijer (1986)
Javesella simillima (Linnavuori, 1948) According to Remane & Fröhlich (1994a) the status of German specimens require further investigation, particularly their relationship with J. discolor (Boh.). However, J. simillima (Lnv.) is much smaller and paler, and the aedeagus is triangular. German specimens seen by the author were morphologically and ecologically clearly distinct from the preceding species. Furthermore, both taxa are reported as distinct species from large parts of arctic Europe, Asia and North America (Anufriev & Emelyanov 1988; Ossiannilsson 1978; Wilson 1988, 1992). Thus, there is no reason to doubt their specific separation.
E V – E VII; nymph, 1 gen. According to existing data this taxon is largely confined to arctic regions of Asia and North America and occurs very locally in bogs of northern Europe, the Polish Tatra and the mountains of northern Bohemia and Moravia (Czech Republic). Furthermore, there are 6 localities in intermediate and raised bogs in the Erzgebirge, the Thuringian Forest, the Dresdener Heide and the Niederlausitz, between 180 and 960 m a.s.l. Most specimens in these sites were found on Eriophorum angustifolium, but Carex rostrata is reported as a further host. Emmrich (1975), Schiemenz (1971a, 1975), Nickel (2002), HN; see also Anufriev & Emelyanov (1988), Lauterer & Novotný (1991), Ossiannilsson (1978), Wilson (1988, 1992)
Javesella pellucida (Fabricius, 1794) B IV – M X; nymph, 2 gen. This is a eurytopic species living in various wet to dry, sunny to moderately shady sites. Food plants are predominantly grasses (Festuca, Elymus, Dactylis, Poa, Deschampsia, Agrostis, Lolium, Calamagrostis, Phleum and others), less commonly Cyperaceae, Juncaceae
Delphacinae Leach, 1815
61
and perhaps also Equisetaceae. Populations with brachypterous individuals are usually 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. Highest 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 grassland, 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), Remane (1987), Remane & Wachmann (1993), Schiemenz (1971a, 1975, 1987), Wagner (1935), Nickel & Achtziger (1999), HN and others; see also Brˇcák (1979), Nuorteva (1965), Raatikainen (1967, 1970b)
Javesella dubia (Kirschbaum, 1868) E IV – M X; nymph, 2 gen. On grasses, usually Agrostis stolonifera and A. capillaris, probably also on other species, in damp to moderately wet, sunny to shady sites (mainly meadows and pastures, forest roads, ruderal sites, abandoned fields, moist forests and low-input lawns of gardens and parks), under natural conditions presumably in salt marshes, along shores of lakes and rivers and in fen woods. Widespread and common in most parts of Germany, in the Mittelgebirge and the Bavarian Alps up to at least 900 m a.s.l., probably only temporary populations at higher altitudes; in the Swiss Alps nymphs were found at 1200 m a.s.l. Has been reported as a potential vector of several plant viruses, but is probably only of minor economic importance. Fischer (1972), Kuntze (1937), Reimer (1992), Remane (1987), Schiemenz (1987), Wagner (1939a), Nickel & Achtziger (1999), HN and others; see also Brˇcák (1979), Günthart (1987a)
Javesella obscurella (Boheman, 1847) M IV – M IX; nymph, 2 gen. In temporarily wet, sunny to moderately shady sites on loamy and clayey substrates, locally also on slightly saline soils. Primary habitats are scarcely vegetated mud banks along lake and pond shores and in floodplain depressions, secondary ones are pastures and meadows, waysides and mining areas. Host plants are mainly grasses (adults mostly on Alopecurus geniculatus and Glyceria fluitans), perhaps also species of Cyperaceae and Juncaceae. Widespread and common in Germany, also in higher parts of the Mittelgebirge and the Alps, brachypterous individuals usually below 1200 m a.s.l., macropters frequently found up to 2000 m a.s.l., where breeding seems unlikely, however. Is reported as a vector of Oats sterile dwarf virus (OSDV) from Finland, but probably without economic importance. Fischer (1972), Heller (1987a), Leising (1977), Niedringhaus (1991, 1997), Schiemenz (1987), Schwoerbel (1957), Wagner (1935), Nickel & Achtziger (1999), HN; see also Brˇcák (1979), de Vrijer (1986), Lindsten (1979)
62
Review of species
Javesella salina (Haupt, 1924) E V – E VIII; nymph, 1(?) gen. In saline sites along the coast and inland, rarely in fens and peaty meadows. The host plants are unknown, perhaps Juncus gerardii, Puccinellia distans or Briza media. Found in Germany only very locally, mainly along the coast of the Baltic Sea (westward at least to Heiligenhafen) and in the rain shadow area to the east of the Harz Mountains (between Magdeburg, Halberstadt, Bad Frankenhausen and Halle). Also recorded in rather isolated sites near Hamburg, Bad Salzungen, Bad Nauheim and Ansbach (400 m a.s.l.). Fröhlich (1996a), Kuntze (1937), Schiemenz (1987), Wagner (1935, 1940a), ZIMH, Nickel (1999b), HN; see also Lauterer (1980)
Javesella forcipata (Boheman, 1847) E V – E VIII, according to Schiemenz (1987) B V – B IX; nymph, 1 gen. In cool and moderately eutrophic, temporarily wet or flooded, shady to moderately shady sites, often with lush grassy vegetation (mountain and fen forests, forest meadows, tall herb stands, spring mires, etc.). Host plants are various grasses (Deschampsia cespitosa, Holcus mollis, probably also Poa trivialis, Calamagrostis canescens, Agrostis stolonifera; recorded on Poa annua in the Netherlands). The species is widespread in Germany, with strongholds in the submontane and montane belt, more scattered in the lowlands, found to at least 1000 m a.s.l. in the Mittelgebirge, frequently up to 1500 m a.s.l. in the Bavarian and Allgäu Alps. Nikusch (1976), Reimer (1992), Schiemenz (1975, 1987), Wagner (1935), Nickel & Achtziger (1999), Nickel & Remane (1996), Nickel (1997), HN; see also de Vrijer (1986)
Javesella stali (Metcalf, 1943) So far B V – E VIII; nymph, 2 gen. This is a pioneer species in stands of horsetail in early successional stages on sandy to loamy substrates, mainly sand pits and river banks, in southwestern Germany also found along waysides, in fallow fields and ruderal sites; the host plant is Equisetum arvense. In Germany only rarely recorded (though probably overlooked) and at the northern and western edge of the range ; at present there are 16 known localities near Berlin, in the Spreewald, along the middle course of the Main and the upper Isar (750 m a.s.l.) and along the southern and middle Rhine (between Freiburg and Bonn). It is also reported from near Stettin (northwestern Poland) and the eastern half of France (Cahors/Lot, Savoie, Paris). Fechtler (pers. comm.), Maczey (pers. comm.), Schiemenz (1987), Nickel & Remane (1996), Nickel (1999a), HN; also della Giustina (pers. comm.), della Giustina & Remane (1991), Remane & della Giustina (1993), Wagner (1941a)
Ribautodelphax albostriata (Fieber, 1866) B IV – B X; nymph, 2 gen.
Delphacinae Leach, 1815
63
In grass stands usually in sunny, oligotrophic to moderately eutrophic, moderately dry to damp, occasionally also moist sites (e.g. various types of dry grassland, meadows, ruderal sites, waysides, forest clearings); single macropterous individuals on dispersal flight, however, may be found in most terrestrial habitats. Lives monophagously on Poa pratensis (including ssp. angustifolia). Widespread in Germany and common in many places, although apparently absent from the northwest German plain; border localities are near Aachen, Cologne, Hildesheim, Hamburg and Lübeck. Highest records of brachypterous individuals are from 950 m a.s.l. on the Swabian Jura, from 1200 m a.s.l. in the Allgäu Alps. Fischer (1972), Frommer (pers. comm.), Günthart (1988), Reimer (1992), Remane (1987), Schiemenz (1987), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN and others
Ribautodelphax angulosa (Ribaut, 1953) So far B V – E VIII; nymph, 2 gen. In sunny, moderately dry, occasionally also moderately moist, usually acidic and oligotrophic sites (mainly dry grassland on sandy or siliceous substrates as well as lowproductivity pastures and meadows). The host plant is Anthoxanthum odoratum. Despite wide host distribution this species is apparently rare and very localized, but has often been misidentified. At present there are only 10 verified records from Germany, most of which have been gathered in or near the northern upper Rhine plain: near Frankfurt am Main, Wiesbaden, Ebernburg an der Nahe, Speyer and Bruchsal, as well as Donnersberg near Rockenhausen. Other localities are widely scattered: Oberstdorf/ Allgäu, c. 800 m a.s.l., Höhbeck near Lüchow, Bornhagen near Bad Heiligenstadt, Schkeuditz near Halle, Sprey near Bautzen and Saupsdorf (Elbsandsteingebirge). Records published until 1987 may refer to other species and are in need of revision. Bieman (1987a, and pers. comm.), Wagner (1939a), Walter (pers. comm.), Nickel (1997), HN
Ribautodelphax collina (Boheman, 1847) E IV – B X; nymph, 2 gen. Monophagous on Agrostis capillaris in damp to dry, sunny, oligotrophic and usually acidic sites (dry grassland on sand and other siliceous substrates, abandoned grassland, occasionally low-productivity meadows). The species is widespread and locally common in the north German plain, but in middle and southern parts of Germany, localities are rather scattered and largely confined to areas of sand, sandstone and other acidic substrates (mainly valleys of Rhine and Main, Mittelfranken, Thuringia); highest localities are at 600 m a.s.l. Some of the older published records may refer to R. vinealis Bieman. Bieman (1987a, 1987b), Remane (1987), Schiemenz (1987), Walter (1999), Nickel & Achtziger (1999), HN
Ribautodelphax imitans (Ribaut, 1953) M V – E VIII; nymph, 2 gen. Monophagous on Festuca arundinacea, in temporarily moist to temporarily wet, rather eutrophic, usually sunny sites, notably meadows, pastures and abandoned grassland.
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Review of species
The range includes southern and western parts of Europe; in Germany the species is apparently restricted to the southwest (found in the Kaiserstuhl, Lake Constance Basin, upper valley of the Neckar near Tübingen, Main and Regnitz valley near Schweinfurt and Forchheim, as well as middle Hessen near Selters and Marburg), up to at least 400 m a.s.l. Bieman (1987a, 1987b), Remane & Fröhlich (1994b), Schwoerbel (1957), Nickel & Achtziger (1999), Nickel & Remane (1996), HN
Ribautodelphax pungens (Ribaut, 1953) Older records, particularly some from northeastern Germany, are in need of revision and may refer to other species of the genus.
E III – B X; nymph, 2 gen. Monophagous on Brachypodium pinnatum in damp to dry, sunny to moderately shady and usually basic sites; particularly common and often among the dominant species in calcareous grassland and open pine forests, but some populations are reported to comprise varying proportions of triploid, pseudogamous ‡‡. Widespread in middle and southern parts of Germany, less common and at the edge of the range in the north. Northernmost localities are on a line from Aachen to Bonn, Bielefeld, Hildesheim, Parchim, Rostock and Rügen; extending up to 1200 m a.s.l. in the Allgäu Alps. Bieman (1987a, 1987b, 1988), Reimer (1992), Remane (1987), Rombach (1999b), Schiemenz (1987), Nickel (1994), HN
Ribautodelphax vinealis Bieman, 1987 In the Netherlands M V – M IX; nymph, 2 gen. (Bieman 1987a). Monophagous on Agrostis vinealis in sandy dry grassland. Was until recently only reported from the Netherlands. From Germany there are three known localities not higher than 150 m a.s.l.: Stolzenau an der Weser, 1951; Quedlinburg, Harslebener Berge, 31.VII.1957; Pevestorf near Schnackenburg an der Elbe, 21. and 29.V.1959. A revision of older records up to 1987 may reveal more localities. Bieman (1987a, 1987b, and pers. comm.), Marchand (1953), Remane & Fröhlich (1994b), Nickel (1997)
4.1.3 Achilidae Stål, 1866 Cixidia confinis (Zetterstedt, 1828) In Sweden according to Ossiannilsson (1978) M V – B VIII; nymph, 1 gen. Nymphs as well as adults in fissures and under the bark of dead trunks of pine, feeding on mycelium of fungi. From central Europe there are only two records, both from the Bavarian Alps in the vicinity of Bad Reichenhall: Jochberg near Weißbach an der Alpenstraße, 800 m a.s.l., 22.VII.1974, 1 †, and Winkl near Berchtesgaden, 700 m a.s.l., 22.VIII.1996, 1 †, on a tree trunk in an open pine forest on a southwest-facing slope. Otherwise known only from Sweden, Finland, Estonia and middle Russia.
Dictyopharidae Spinola, 1839
65
SMNS (leg. Dynort), Nickel (1999b); see also Anufriev & Kirillova (1998), Linnavuori (1951) (see this paper also for identification of nymphs), Ossiannilsson (1978)
Cixidia pilatoi D’Urso & Guglielmino, 1995 Populations of this species living in south-central Europe were published as Cixidia marginicollis (Spinola, 1839) by most authors. The true C. marginicollis (Spin.), however, is so far only known from Sicily (D’Urso & Guglielmino, 1995), and differences between German specimens and the newly described C. pilatoi D’Urso & Gugl. from Italy appear to be insignificant.
So far B VI – M VI, in Austria and the Czech Republic until M IX (Holzinger 1995; Lauterer 1978; Wagner & Franz 1961); nymph, 1 gen. Known from Germany only from the Kaiserstuhl (Badberg, c. 350 m a.s.l., 10.VI.1952, 2 ‡‡, and 20.VI.1967, 1 †, 1 ‡). Probably lives – like most other species of Cixidia – on and under the bark of dead trees feeding on hyphae of fungi. Is considered to be associated with deciduous trees, notably Quercus. Coll. H.J. Müller, Remane (pers. comm.), Wagner & Franz (1961); see also Holzinger (1995a, 1996), Lauterer (1978)
4.1.4 Dictyopharidae Spinola, 1839 Dictyophara europaea (Linnaeus, 1767) Mainly M VII – M X, according to Schiemenz (1987) also E VI; egg, 1 gen. In sunny, xerothermic sites on various substrates, preferentially in disturbed patches with some bare ground (oviposition is done into the soil), locally even in dry ruderal sites. Probably polyphagous, most individuals found on dicotyledonous herbs, but the species is also reported from dwarf shrubs and even trees. In Germany it lives at the northern edge of the range, with strongholds in warmer regions in southern and eastern parts (rain shadow area to the east of the Harz Mountains, upper Rhine plain, valleys of Neckar, Main, Nahe, Danube). Northernmost localities (which appear to be more or less isolated) are on a line from Bonn to Gießen, Gotha, Sondershausen, Stendal, Nauen and Stralsund, the highest ones are only at 400 m a.s.l. in the Lake Constance basin. Haupt (1924), Müller (1942), Nikusch (1976), Remane (1987), Schiemenz (1987), Schönitzer and Oesterling (1998a), Schwoerbel (1957), Sergel (1984b), SMNS, Voigt (1978), Wagner (1939a, 1951a), HN
4.1.5 Tettigometridae Germar, 1821 Among central European Auchenorrhyncha the taxon Tettigometra Latr. is probably the most critical group concerning identity and distinction of species. This is caused by only poor differences in genitalic morphology and a considerable intraspecific variation of head shape and body colouration. Life history is another enigma: presumably all – at least central European – species are associated with ants; the character of these relationships, however, is largely unknown. Müller (1942) has observed oviposition on the surface of stones, which may, to some extent, explain the extreme reduction of the female genitalia and the preference of habitats with bare patches. It has been suggested that the nymphs live in ants’ nests feeding on plant roots
66
Review of species
(Emelyanov 1987), but also nymphal development near the plant base has been observed (see Müller 1956). Rearing over more generations has failed so far. Hence, species interpretations in this book may not be identical with those of the original describers, particularly since the whereabouts 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 Mitricephalus Sign. are only subgenera. The situation among black taxa in central Europe is particularly problematic. Apparently, there are at least two species in Germany. For this book, they were preliminarily distinguished by the following characters: larger specimens (5 - 5.5 mm), which are rather broad-built, usually shiny black, but also blackish- to orange-brownish above (particularly on the vertex), are interpreted as T. fusca Fieb. The front is usually concave, the vertex showing two lateral bulbosities. In contrast, smaller specimens (usually 3.3 – 4.3 mm), most of which are blackish, but occasionally with dark or even pale brown fore wings (particularly after degreasing with benzine) and with a short, but much variable head, are treated as T. atra Hag. This opinion is based on the study of museum material from many parts of Europe determined by various authors, but is perhaps not in accordance with the interpretation of Fieber (1865). It should be stressed that brown-winged specimens have been labelled and also published as T. atra laetifica Metcalf, 1932, but also as T. laeta H.-S. It should also be noted that the identity of T. depressa Fieber, 1865 and T. atrata Fieber, 1872 is considered here as dubious.
Tettigometra atra Hagenbach, 1825 T. atra laetifica Metcalf, 1932 (see taxonomic remarks above!)
B VIII – M VI; adult, 1 gen. In xerothermic, sunny to moderately shady sites on basic to acidic substrates (mainly dry grassland and margins of pine and oak forests, occasionally also managed meadows). Is reported to live in ant nests (Lasius, Tetramorium). Most adults are swept in the low vegetation, a few also from woody plants. In Germany at the northern edge of the range, rather scattered and with strongholds in warmer parts (Saale-Unstrut region, Rhine-Main region, Mainfranken, Neckar valley, Franconian Jura, slopes of the upper Rhine valley). Northernmost localities, some of which are not verified, are near Bacharach (middle Rhine), Schlüchtern, Jena, Quedlinburg, Neustrelitz and Frankfurt an der Oder, up to at least 600 m a.s.l. in the foothills of the Alps. Nikusch (1976), Schiemenz (1987), Schönitzer & Oesterling (1998a), SMNS, Schwoerbel (1957), Wagner (1939a, 1951a), HN
Tettigometra fusca Fieber, 1865 See taxonomic remarks above! Due to a misunderstanding between the authors, this species was published for Thuringia as T. atrata Fieb. by Nickel & Sander (1996).
B VIII – M VI; adult, 1 gen. In spatially diverse dry grassland with open stands of trees or shrubs, usually on basic substrates on xerothermic plateaus or on upper slopes. Was found in nests of Formica pratensis Retzius several times. In Germany only very locally and mainly in limestone areas. Most records (including recent ones) are from the Triassic limestone regions of eastern Thuringia (near Jena and Arnstadt), Franconia (between Würzburg, Karlstadt and Bad Kissingen, also near Coburg), the valleys of Tauber and Kocher (Schweinberg
Tettigometridae Germar, 1821
67
near Buchen, Crispenhofen near Künzelsau). Also published from near Erlangen, München and Starnberg. The highest sites are situated at 600 m a.s.l. Apparently, many populations have vanished during recent decades. Biedermann (pers. comm.), Heller (pers. comm.), Schiemenz (1987), Schönitzer & Oesterling (1998a), SMNS, Trümbach (1959), Wagner (1951a), ZIMH, ZMB, Nickel & Sander (1996), Nickel & Remane (1996)
Tettigometra griseola Fieber, 1865 So far B VIII – M VI; adult, 1gen. Usually in xerothermic sites with at least scattered shrubs or trees, mainly on limestone, loess and schist; another record is from a seepy, temporarily wet slope. In Germany known only from 5 localities in the Kaiserstuhl area (Schelingen, Vogtsburg, Istein), along the middle Rhine (Lorch) and in the foothills of the Alps (Fischen am Ammersee), up to at most 600 m a.s.l. Remane (pers. comm.), Wagner & Franz (1961), HN
Tettigometra impressopunctata Dufour, 1846 Populations living in Germany may belong to at least two different species. Moreover, there are older reports of T. depressa Fieber, 1865, T. atrata Fieber, 1872, and others, the identity of which is unclear and which are perhaps conspecific with T. impressopunctata Duf.
Mainly B VIII – B VI, occasionally also VII; adult, 1 gen. Along sunny woodland margins in dry to temporarily dry, basic to acidic, usually rather warm sites, mainly in grazed dry grasslands, along spatially diverse forest margins and on gravel banks of alpine rivers with scattered shrubs. There are also older records from fenland habitats in the foothills of the Alps. Adults are found among herbaceous vegetation as well as on shrubs and trees (Quercus, Taxus, Myricaria and others). In Germany at the northern edge of the range, presumably declined in recent decades. Most localities are in the foothills of the Alps and in warmer regions of southern and eastern parts (valleys of Rhine, Main and Neckar, as well as in Thuringia), northernmost sites, which appear to be more or less isolated, are near Siegen, Göttingen, Bad Frankenhausen and Magdeburg. The highest known sites are at 900 m a.s.l. in the Bavarian Alps, at 1200 m a.s.l. in Switzerland. Schiemenz (1987), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Wagner (1939a, 1951a), Nickel (1994), HN; see also Günthart (1987a)
Tettigometra laeta Herrich-Schäffer, 1835 Has been confused with brown-winged specimens of T. atra Hag. (= T. atra var. laetifica Metc.; see above).
This is apparently a Mediterranean species, which is reported to live in ants’ nests, but no further details are known on the life history. The only record from Germany is the original description: “Bei Regensburg selten”, but the type material is unfortunately lost. Herrich-Schäffer (1835a); see also Kirschbaum (1868), Metcalf (1932), Wagner (1939a)
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Review of species
Tettigometra leucophaea (Preyssler, 1792) Tettigometra obliqua (Panzer, 1799)
Mainly B VIII – M VI, occasionally also VII; adult, 1 gen. This species is published from various, usually spatially diverse xerothermic sites, often with scattered trees or shrubs, bare sand or rocks. In older papers, notably from the beginning of the 20th century, mass occurrences have been reported with subsequent damage to rye, wheat and oats, but the species has declined dramatically in Germany, as in most parts of central Europe. Formerly it was widespread in middle and southern parts and has occurred northward at least to Frankfurt am Main, Lauenburg, Ludwigslust, Waren an der Müritz and Angermünde, extending to at least 500 m a.s.l. in the foothills of the Alps. The current status is uncertain; the latest records are from the 1960s and 1970s (Saale valley near Jena, lower Oder valley near Seelow and Bad Freienwalde). Many of the older localities have been checked since without positive records. Thus, the species is probably extinct in Germany. Fischer (1972), Kirschbaum (1868), Kuntze (1937), Lindberg (1948), Remane (1987), Schönitzer & Oesterling (1998a), Schiemenz (1987), Schwoerbel (1957), SMNS, Torka (1905), Wagner (1935, 1939a, 1951a, and unpublished data)
Tettigometra macrocephala (Fieber, 1865) E VII – E VI; adult, 1 gen. Along xerothermic and spatially diverse woodland margins in basic as well as acidic sites, mostly in vine growing regions. In Germany known only from a few isolated localities at the northern edge of the range and apparently declining, altogether known from 17 sites, mainly in Triassic limestone areas of eastern Thuringia, Franconia and northern Baden (near Jena, Bad Kissingen, Hammelburg, Karlstadt, Tauberbischofsheim), but in the Rhine-Nahe region (Schloßböckelheim, Bad Münster, Wöllstein, Lorch) also on porphyry, Keuper and other substrates; also found along the upper courses of the Main, Neckar and Danube (Bamberg, Tübingen, Ulm), between 150 and 500 m a.s.l. Biedermann (pers. comm.), Dynort (pers. comm.), Lindberg (1948), Schiemenz (1987), Schwoerbel (1957), SMNS, Wagner (1939a, 1951a), HN
Tettigometra virescens (Panzer, 1799) Most vivid green individuals of Tettigometra smaller than 5 mm in central Europe have been labelled as T. virescens (Panz.). However, they have also been confused with T. laeta H.-S. and perhaps other species. Furthermore, the identity of numerous varieties mostly described from southern Europe (see Metcalf 1932) is not clear. In contrast, the morphological variation of specimens originating from Germany and adjacent areas is surprisingly low.
E VIII – M VI; adult, 1 gen. Along woody margins in xerothermic sites on basic to acidic substrates (mainly Triassic limestone, gypsum, schist). Most adults are swept from low vegetation, in Poland also from Pinus and Juniperus. Rare in Germany and at the northern edge of the range; only known from a few localities, usually on upper slope and plateau situations: Saale valley
Caliscelinae Amyot & Serville, 1843
69
(near Jena), Thuringian Basin (Gotha), Main and Tauber region (near Frankfurt am Main, Tauberbischofsheim, Karlstadt, Hammelburg, Bad Kissingen, Bad Königshofen) and the valley of the middle Rhine (near Bacharach), between 150 and 350 m a.s.l. Biedermann (pers. comm.), Haupt (1924), Müller (1978), Reimer (1992), SMNS, Wagner (1939a, and unpublished data), Nickel (1999b), HN; see also Smreczynski (1954)
4.1.6 Issidae Spinola, 1839 4.1.6.1 Caliscelinae Amyot & Serville, 1843 Ommatidiotus dissimilis (Fallén, 1806) Older German reports of O. concinnus Horváth, 1905, a species otherwise published only from Hungary (see Nast 1987), are dubious and probably refer to O. dissimilis (Fall.).
B VII – M X; egg, 1 gen. In raised and intermediate bogs on Eriophorum vaginatum. In Germany only scattered, but locally abundant, mainly in the north German plain and along the edge of the Alps, also in the Black Forest; apparently absent from most regions of the Mittelgebirge. The highest localities are at 900 m a.s.l. Fischer (1972), Remane (1958), Schiemenz (1976, 1987), Wagner (1935), HN
4.1.6.2 Issinae Spinola, 1839 Issus coleoptratus (Fabricius, 1781) Mainly E VI – M X, in warmer regions from E V onwards; nymph, 1 gen. On woody plants in damp to dry sites, in closed deciduous and mixed forests as well as in semi-open tree stands (e.g. along forest margins, in parks, cemeteries, etc., also on solitary trees on grassland). Adults on various deciduous trees (mainly Quercus and Acer, also Corylus, Betula, Ulmus, Tilia, Sorbus, Clematis and others), hibernating nymphs found on Hedera, Ligustrum, Juniperus and Taxus, as well as in the litter layer. In Germany the species occurs at the northern limit of the range, which runs approximately along the northern edge of the Mittelgebirge (northernmost records near Cologne, Northeim, Quedlinburg, Leipzig and Kamenz); densely spread to the south of this line, although sweep net catches usually contain only low numbers. Frequently found up to 1000 m a.s.l. in the Bavarian Alps. Achtziger (1991), Büchs (1988), Fischer (1972), Frommer (1996), Post-Plangg & Hoffmann (1982), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Wagner (1939a, 1951a), Nickel (1997), HN
Issus muscaeformis (Schrank, 1781) So far E V – E VIII; nymph, 1 gen. Apparently not ecologically distinct from I. coleoptratus (F.), although extending further northward in Europe. Found on Quercus and Corylus along sunny forest margins and on
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Review of species
solitary trees and shrubs. Is reported from almost the whole of the European deciduous forest zone, but recorded from Germany from 9 localities only: upper and lower course of the Elbe (Pirna, Harburg, and Winsen an der Luhe), Bergisches Land (Hückeswagen), Fulda valley (Kassel), Saale-Unstrut region (Freyburg), Bavarian Forest (Hochoberndorf and Ulrichsberg, both near Deggendorf) and Berchtesgaden Alps (St. Bartolomä), extending to at least 700 m a.s.l. Emmrich (pers. comm.), Remane (pers. comm.), Coll. Sander, SMNS, Wagner (1935, and unpublished data), HN; see also Holzinger (1996b), Ossiannilsson (1978), Wagner & Franz (1961)
Agalmatium bilobum (Fieber, 1877) This is a Mediterranean species, known from Germany only from a single record on the East Frisian Islands: Wangerooge, 17.VI.1960, in white dunes „eine gute Population“, which was no more found in the 1980s. Hence, this was apparently a temporary occurrence. Harz (1965, 1988), Niedringhaus (1991), Remane & Fröhlich (1994a)
4.2 Cicadomorpha Evans, 1946 – Leafhoppers (s.l.) 4.2.1 Cicadidae Leach, 1815 (s.l.) – Cicadas
Cicada orni Linnaeus, 1758 VI – B IX; nymph, development takes about 4 years (Schedl 2000). This species is widespread and common in Mediterranean regions. Obviously, there are occasional influxes into viticultural regions of southwest Germany. Two more recent records are based on observations only, but the species can be easily identified: Stuttgart, 30.VII. -12.VIII.1986, 1 † singing; Öhringen, 05.VII.1997, 1 ‡, 3 weeks later song heard. Furthermore, there is a record from the Pliocene of Willershausen (southern Niedersachsen). Nymphs and adults are reported to be polyphagous, adult feeding has been recorded on Pinus, Pistacia, Olea, Fraxinus and others. Dynort (pers. comm.), Heller (1987c), Wagner (1967); see also Boulard & Mondon (1995), Schedl (2000)
Tibicina haematodes (Scopoli, 1763) M IV – B VIII; nymph, development takes several years, three according to Schedl (2000). In dry grassland and abandoned vineyards with scattered shrubs and trees, usually in xerothermic upper slope situations on limestone and Triassic sandstone. According to older reports, the nymphs feed on roots of Vitis vinifera and Prunus spinosa, oviposition takes place on the latter. Adults are usually found in the tree and shrub layer and are reported from Pinus sylvestris, Quercus pubescens, Fraxinus excelsior, Acer spp., Prunus spinosa and others. Single individuals may perform long dispersal flights and often land
Cicadidae Leach, 1815 (s.l.) – Cicadas
71
in exposed sites such as hilltops, solitary trees and even vineyard workers. In Germany the species is restricted to isolated localities in viticultural regions in southwestern parts between 100 and 350 m a.s.l., but many records date back before 1940. The strongholds are in Rheinhessen (between Bad Kreuznach, Alzey and Nierstein – also recently), the Main valley near Volkach and between Karlstadt and Ochsenfurt (all recently), as well as in the valleys of Neckar and Enz and in the adjacent Stromberg and Kraichgau (between 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 & Mondon (1995)
Cicadetta montana (Scopoli, 1772) According to Gogala & Trilar (1998, 1999) this is a complex of several species, which are distinct mainly bioacoustically. Central European populations probably belong to at least two species. The song of the more widespread form is long-lasting and continuous, with only short and irregular interruptions. Another form is apparently rare and known from Germany only from a few localities at the northern edge of the Thuringian Forest (Tieftal near Arnstadt) and the Tauber valley near Tauberbischofsheim and Bad Mergentheim. Its song consists of two different and alternating echemes, the first of which lasts several seconds, the second is very short. However, it should be noted, that in many localities the type of song has not been recorded. Thus, the short-singing form is perhaps more widespread. For French populations with a short song, Boulard & Mondon (1995) use the name C. petryi (Schuhmacher, 1924), which has been created for specimens from the Kyffhäuser in Thuringia, based on a slightly different colouration. However, it is uncertain whether this name is correct. All Kyffhäuser populations recorded during recent years clearly performed the long-lasting and continuous type of song, and morphological characters for distinction between the two groups have not yet been found (Gogala & Trilar, pers. comm.). See also Schedl (2000).
M V – M VII; nymph, development takes several years. Among open stands of shrubs or trees in moderately dry to dry, sun-exposed sites, usually on hillsides or plateaus, mainly in dry pastures and meadows and along sunny forest margins. Widespread but scattered in the southern and middle parts of Germany, with strongholds in viticultural regions in the southwest and in the rain shadow area to the east of the Harz Mountains, usually below 500 m a.s.l., but frequently at 750 m a.s.l., along the upper course of the Isar and on the Swabian Jura, locally even up to 1000 m a.s.l. in the Black Forest. The northernmost localities are near Ahrweiler, Beckum/Westphalia, Ibbenbüren, Holzminden, Göttingen, Nordhausen, Quedlinburg, Naumburg an der Saale, Finsterwalde and Fürstenwalde. The species is absent from Denmark, but occurs very locally in southern Scandinavia. Andres (pers. comm.), Ant (1991), Back (1976), Bernhardt (1991), Bußmann & Feldmann (1995), Eitschberger (1972), Emmrich (1984), Felzmann (pers. comm.), Hoffmann (1991), Kless (1971), Niehuis & Weitzel
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Review of species
(1996), Reimer (1992), Schedl (2000), Schiemenz (1988), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Stein & Bogon (1990), Vogel (1935), Wolf (1976), Nickel (1994), HN and others; see also Ossiannilsson (1981)
4.2.2 Cercopidae Leach, 1815 (s.l.) – Froghoppers 4.2.2.1 Cercopinae Leach, 1815 Cercopis arcuata Fieber, 1844 In eastern Austria B V – B VII (Holzinger 1996b); nymph, 1 gen. In Germany known only from 4 records from the Alps and their foothills, all probably dating before 1950: Schellenberg, 03.VI.1938, and Almbachklamm, 05.VI.1938, both in the vicinity of Berchtesgaden (ssp. haupti Priesner, 1923), also „Bayerisches Allgäu“ and Geiselgasteig to the south of Munich, c. 550 m a.s.l. In warmer regions of the Czech Republic, Austria and France this species lives in pine or oak forests; most adults were found on dicotyledonous herbs. Wagner & Franz (1961), ZIMH, ZSM; see also Holzinger (1996b), Lauterer (pers. comm.)
Cercopis sanguinolenta (Scopoli, 1763) B V – E VII; nymph, 1 gen. In sunny, moderately dry to dry sites on basic to acidic substrates in warm situations, notably in dry grassland and abandoned vineyards, preferably in disturbed patches with tall, but scattered vegetation. Food plants are various grasses and herbs; most feeding adults are reported from Arrhenatherum elatius, also on Salvia pratensis, Brachypodium pinnatum, Falcaria vulgaris, Cardaria draba, Galium verum, Thalictrum minus and others. Germany is at the northern edge of the species’ range, with only isolated populations, mainly in the rain shadow area to the east of the Harz Mountains (between Sondershausen, Halle, Leipzig, Eisenberg, Jena and Kölleda, as well as up along the Saale to Rudolstadt) and in the Rhine-Nahe region (Bacharach, Schloßböckelheim, Bad Münster). A few isolated localities are situated in the Elbe valley near Meißen and the foothills of the Alps near Starnberg, c. 600 m a.s.l. In the northeastern Alps of Styria (Austria) the species is less xerophilous and frequently occurs in the upper montane belt, locally up to 1600 m a.s.l. Biedermann (1998), Emmrich (1983), Jöst (1966), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1988), Schönitzer & Oesterling (1998a), HN; see also Wagner & Franz (1961)
Cercopis vulnerata Rossi, 1807 In the lowlands B V – M VII, at higher altitudes VI – E VIII; nymph, 1 gen. In moderately dry to moderately wet, sunny to moderately shady sites, mainly abandoned grassland, forest glades, open forests, along waysides, streams and ditches, absent from mown meadows. Adults mainly on various tall herbs and grasses (Urtica, Aegopodium, Lupinus, Chrysanthemum, Filipendula, Arrhenatherum and others); the only
Cercopinae Leach, 1815
73
published records of nymphal feeding (from Switzerland) are from Arrhenatherum elatius and Dactylis glomerata. Widespread in Germany and common in southern and central 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 northern Germany and at the edge of the range, extending at least to the East Frisian Islands, Bremerhaven, Itzehoe, Ratzeburg, Rostock and Strasburg. Fischer (1972), Kehlmaier (2000), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Wagner (1935, 1939a), HN and others; see also Mauri (1982)
Haematoloma dorsatum (Ahrens, 1812) E IV – E VII, occasionally until E VIII; nymph, 1 gen. Adults on pine (Pinus sylvestris, additional species, notably Pinus nigra, in more southern and western parts of Europe) in open forests in damp to dry sites, usually on limestone or sandy soils. Nymphs live in the litter layer and feed on fine roots and basal stems of grasses (preferentially Deschampsia flexuosa on acidic substrates, probably Brachypodium pinnatum and others on basic ones). Numbers of swept adults may be large in grassy forest floor vegetation after emergence in May, but rather small in the lower part of the tree layer from June onwards. Here they feed on pine needles and mate; oviposition takes place in the grass layer. In the Netherlands and the Mediterranean region there may be local damage in pine plantations. In Germany at present the species occurs only in the western half (mainly below 400 m a.s.l., but to at least 700 m a.s.l. on the Swabian Jura) and apparently is extending its range: collected for the first time in 1935 near Bonn, from 1936 onwards in the Rhine-Nahe region, from where it was published as new to Germany, recorded in 1952 near Karlsruhe, in 1955 in the Kaiserstuhl, since 1956 along the northeastern edge of the Black Forest near Calw, later in large parts of Swabia (eastward at least to Schwäbisch Gmünd and Bad Urach), since 1969 in middle Hessen and in the Weser-Ems area, since the mid 1970s in the middle Main region between Karlstadt, Würzburg, Kitzingen and Schweinfurt, as well as along the Fränkische Saale, at least since 1984 in the foothills of the Alps of Upper Swabia north of Ravensburg, found in 1987 on the East Frisian island of Borkum, later also on Norderney, since 1989 in the Senne (Westphalia), in 1993 along the upper Main near Staffelstein, in 1996 near Lake Constance (Sipplingen), in 1997 in the Lüneburger Heide to the south of Fallingbostel, 2001 in the Grabfeld of southern Thuringia, and 2002 in the Altmark (Salzwedel, Klötze). Apparently, pine forest areas in central Thuringia, the Wendland of Niedersachsen and in Mecklenburg, which have been investigated during recent years, were not yet colonized. Achtziger (pers. comm.), Bernhardt & Arnold (1991), Bußmann & Feldmann (1995), Fritzlar (pers. comm.), Gauss (1987), Haeseler & Niedringhaus (1988), Heller (1960), Hess (pers. comm.), Jöst (1966), Kupka (1941), Niedringhaus & Olthoff (1993), Nikusch (1976), Remane (1987), Remane & Wachmann (1993), Schulze (1992), SMNK, SMNS, Voigt (pers. comm.), Wagner (1939a), Witsack (pers. comm.), ZFMK, Nickel & Remane (1996), HN; see also Holzinger (pers. comm.), Moraal (1996)
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Review of species
4.2.2.2 Aphrophorinae Amyot & Serville, 1843 Lepyronia coleoptrata (Linnaeus, 1758) E V – M X; egg, 1 gen. Usually in sunny, dry to moist, occasionally also temporarily moist or even wet sites, mainly ruderal habitats, waysides, dunes, dry grassland (often in disturbed patches) and low-input meadows; also reported from saline sites near the coast of the Baltic Sea. Adults often on grasses; in the foothills of the Alps, nymphs were found on Equisetum, Pedicularis, Eupatorium, Solidago, Artemisia, Carex, Phragmites, Menyanthes and various umbelliferous herbs, in Sweden on Vaccinium, Polygonum, Anthriscus, Chamaenerion, Potentilla, Hypericum, Ranunculus, Plantago, Galium, as well as shrubs and trees (Salix, Populus, Betula, Corylus). Widespread in Germany, locally common in southern and eastern parts, but otherwise rather scattered or rare, extending to at least 1250 m a.s.l. on sun-exposed hillsides in the Mittelgebirge and along the edge of the Alps. Apparently absent from northwestern parts and extending at least to Bonn, Marburg, Eschwege, Nordhausen, Magdeburg, Lüchow and Bad Oldesloe. Fischer (1972), Kuntze (1937), Heller (1987a), Nickel (1979), Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (pers. comm.), Schiemenz (1988), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Wagner (1939a, 1951a), Nickel & Achtziger (1999), Nickel (1997), HN; see also Ossiannilsson (1981)
Neophilaenus albipennis (Fabricius, 1798) Mainly B VI – B IX, occasionally E V – B X; egg, 1 gen. Monophagous in stands of Brachypodium pinnatum in moderately dry to damp, also temporarily moist, usually slightly shady and basic sites, notably in dry grassland and open stands of oak and pine. Widespread and locally common in limestone areas in middle and southern parts of Germany, but with distribution border approximately along the northern edge of the Mittelgebirge. Northernmost localities are near Bonn, Höxter, Wolfenbüttel, Halberstadt, Halle, Gera and Görlitz. In the Bavarian and Allgäu Alps it frequently occurs up to at least 1200 m a.s.l., single individiuals were even recorded at 1600 m a.s.l. This is one of the very few Auchenorrhyncha species of which metapopulation dynamics have been studied in detail. Biedermann (1997, 2000), Deutschmann (pers. comm.), Rabeler (1962), Reimer (1992), Rombach (1999a), Schiemenz (1988), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Nickel (1994), HN
Neophilaenus campestris (Fallén, 1805) E V – B X; egg, 1 gen. In ruderal or grazed, usually moderately dry and sunny sites on various substrates (sandy to clayey, basic to acidic), mainly grassy ruderal sites, mining areas, waysides and military training areas, also grazed or trampled patches in dry grassland and inland dunes. Host plants are various grasses (Agrostis capillaris, probably Elymus repens, Arrhenatherum elatius, Poa compressa and others), some adults ascend onto Pinus and other woody plants during hot summer days. Widespread in the lowlands of Germany (below 600 m a.s.l.), but usually at low densities, with strongholds in warmer regions.
Aphrophorinae Amyot & Serville, 1843
75
Neither recorded to the northwest of a line running from Düsseldorf to Osnabrück, Hamburg and Lübeck, nor to the south of the Danube. Reimer (1992), Remane (1987), Schiemenz (1988), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), Nickel (1997), HN; see also Badmin (1997)
Neophilaenus exclamationis (Thunberg, 1784) According to Wagner (1955) three subspecies occur in Germany: ssp. exclamationis (Thunberg, 1784) in the north German plain, ssp. monticola W. Wagner, 1955 in the middle parts (up to at least 1000 m a.s.l.) southward to the edge of the Alps and ssp. alpicola W. Wagner, 1955 in the Bavarian and Allgäu Alps above 600 m a.s.l. The exact borders between their ranges are unclear, however.
E V – B X; egg, 1 gen. In open forests and along their margins (mostly under pine or oak), usually in oligotrophic, damp to moderately dry, basic as well as acidic sites, also in grassland of the subalpine and alpine belt of the Alps. In most lowland regions the main host plant is clearly Festuca ovina, but the species may also live on Deschampsia flexuosa and additional grasses. It is widespread in Germany, but uncommon, frequently recorded in the Bavarian and Allgäu Alps at 2000 m a.s.l., in Austria and Switzerland up to at least 2500 m a.s.l. Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998a), Wagner (1935, 1939a, 1955), HN; see also Günthart (1987a), Leising (1977), Whittaker (1965)
Neophilaenus infumatus (Haupt, 1917) E V – B X; egg, 1 gen. In sun-exposed, usually low-growing grass stands with incomplete cover of vegetation in dry to moderately dry sites. Host plants are usually (perhaps exclusively) greyish-green subspecies of the Festuca ovina group. In Germany this species is restricted to steppe sites on gypsum and porphyry of Thuringia and Saxony-Anhalt. These localities are at the western edge of the range, although the species is locally dominant. Recorded between Halberstadt, Bernburg, Halle, Jena, Erfurt, Gotha and Sondershausen, up to 350 m a.s.l. Single specimens (perhaps vagrants) were also found in Brandenburg and Saxony. It is reported up to 1600 m a.s.l. in dry valleys of the central Alps of Austria and Switzerland. Schiemenz (1969, 1988), HN; see also Günthart (1987a, 1997), Wagner & Franz (1961)
Neophilaenus lineatus (Linnaeus, 1758) According to Remane & Wachmann (1993) this may be a complex of several species. In Germany, besides ssp. lineatus (Linnaeus, 1758), there are populations, which have been described as Philaenus lineatus pallida Haupt, 1917 (on dunes and in other types of dry sandy grassland of northwestern Germany) and Philaenus aterrimus J. Sahlberg, 1871 (in raised bogs of the Mittelgebirge and the Alps). Ossiannilsson (1981) and most other authors consider them both as ecological forms of N. lineatus (L.).
M VI – B XI; egg, 1 gen. Eurytopic usually in tall stands of Poaceae and Cyperaceae in sunny to shaded, wet to moderately dry sites, mainly in bogs of various types, low-input meadows and pas-
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Review of species
tures, ruderal sites, open forests, salt marshes along the coast and inland, and coastal dunes. Widespread and common all over Germany, frequently found up to 1600 m a.s.l., occasionally at 1800 m a.s.l. in the Bavarian Alps. Host plants are species of Poaceae (Calamagrostis spp., Deschampsia spp., Festuca spp., Agrostis spp., Holcus spp., Ammophila arenaria and others), Cyperaceae (Carex spp., Trichophorum cespitosum and others), probably also Juncaceae (Juncus spp.) and other families. Fischer (1972), Nickel (1979), Niedringhaus (1991), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1988), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN and others; see also Halkka et al. (1977), Novotný (1995), Whittaker (1965a, 1965b, 1971)
Neophilaenus minor (Kirschbaum, 1868) E V – E X; egg, 1 gen. In sunny, oligotrophic, moderately dry to dry sites on various substrates (acidic to basic, preferably sand, but also on solid rock), with sparse cover of vegetation. Host plants are mainly Corynephorus canescens, Festuca ovina, Koeleria glauca, and probably additional species of fine-leaved grasses. Widespread in the sand areas of the north German plain (northward at least to Pinneberg and Eutin) and often among the dominant species in heaths and Corynephorus swards, but rather scattered to the south, where largely restricted to inland dune areas and dry hillsides; highest localities are at 500 m a.s.l. in the foreland of the Alps, at 600 m a.s.l. in the Mittelgebirge. Not reported from Denmark, Norway and Sweden, but from southeastern Finland, Latvia and Lithuania. Frommer (pers. comm.), Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987, and pers. comm.), Schiemenz (1988), Schönitzer & Oesterling (1998a), Wagner (1935, 1951a), HN; see also Ossiannilsson (1981)
Aphrophora alni (Fallén, 1805) B VI – E X; egg, 1 gen. Eurytopic among various stands of woody plants in wet to dry sites, usually along shores of lakes, ponds, rivers, streams and hedges, as well as in parks, forests (mainly along edges and roads) and xerothermic sites with scattered shrubs. Adults are usually swept from deciduous shrubs and trees (Salix, Alnus, Populus, Betula, Prunus spinosa, Corylus, Rosa, Cytisus and many others), nymphs in Scandinavia were found close to the ground on various herbaceous plants (Polygonum, Filipendula, Trifolium, Angelica, Galium, Ranunculus, Hieracium, Viola and others), also on adventitious shoots of Salix, Betula and Alnus. Widespread and common in most parts of Germany, frequently occurring up to at least 1500 m a.s.l. in the Mittelgebirge and the Alps. Achtziger (1991), Fischer (1972), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz (1988), Wagner (1935, 1939a), HN and others; see also Halkka et al. (1977), Ossiannilsson (1981)
Aphrophora major Uhler, 1896 Aphrophora alpina Melichar, 1900 sec. Anufriev & Emelyanov (1988), Komatsu (1997b)
M VII – E IX; egg, 1 gen.
Aphrophorinae Amyot & Serville, 1843
77
In moist to wet sites with scattered trees or shrubs and along forest margins, mainly in fens and intermediate bogs, along the edge of raised bogs and in cool meadow valleys. Adults on Betula and Salix, but also in stands of tall sedges and reeds. The nymphal host plants are not properly known; Phragmites and Pedicularis have been recorded in Bavaria, Salix and Artemisia in Japan. Thus, it is likely that adults after emergence ascend from the herbaceous layer to shrubs and trees, as in A. alni (Fall.). The species is rather localized in Germany, it is usually collected in low numbers only. Most localities are in the Bavarian and Swabian foothills of the Alps (between 500 and 1000 m a.s.l.) and in the northwest German plain (near Lingen, Meppen, Vechta, Cloppenburg, Hanover, Bremen, Pinneberg). Furthermore, there are a few localities along the upper and middle Rhine (Germersheim, Düsseldorf, Cologne), as well as an unverified record from near Berlin. In Poland it is only known from northeastern parts, in France only from the north; it is not reported from Fennoscandia, Denmark and Belgium. Frommer (1996), Haupt (1924), Jöst (1966), Mölleken & Topp (1997), Nickel (1979), Niedringhaus & Olthoff (1993), Schiemenz (1988), Schönitzer & Oesterling (1998a), SMNS, ZMB, Nickel (1999b), HN; see also della Giustina & Remane (2001), Komatsu (1997b), Nast (1976a, 1987)
Aphrophora corticea Germar, 1821 M VI – M X; egg, 1 gen. In forests of Pinus sylvestris and along their margins, in temporarily moist to dry, basic as well as acidic sites. Nymphal development takes place among low vegetation on pine saplings, on Vaccinium myrtillus, Calluna vulgaris and other woody plants, probably also on herbs. After emergence, adults migrate up into the canopy layer, but oviposition presumably takes place near the ground. Widespread in Germany, but locally absent, with strongholds in the pine forest areas of Jurassic and Triassic limestone, sandstone and aeolian sand, as well as in the north German plain. Occurs up to at least 1400 m a.s.l. in the Bavarian Alps. Fischer (1972), Schiemenz (1988), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Wagner (1935, 1939a), HN and others
Aphrophora pectoralis Matsumura, 1903 Aphrophora salicis forneri Haupt, 1919; Aphrophora costalis Matsumura, 1903 sec. Komatsu (1997b) Recently some material of the extensive Matsumura collection was revised by Komatsu (1997a, 1997b, 1997c). A. costalis Mats., which was in use in the European literature since Vilbaste (1968b), was declared a junior synonym of A. pectoralis Mats.
B VI – E IX; egg, 1 gen. Along shores of running and standing water, in meadow valleys, forest glades and other cool sites; adults and nymphs live on various species of willows (mainly Salix caprea, S. purpurea, S. cinerea, S. aurita, locally S. viminalis, S. alba, S. myrsinifolia and others). Widespread in Germany, but usually in low to intermediate frequency, with strongholds in cool situations in the Mittelgebirge and along the edge of the Alps, where extending up to at least 1200 m a.s.l. Is not recorded from Mecklenburg-Vorpommern, the northern half of Poland and Denmark, but from large parts of Fennoscandia and the Baltic countries.
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Review of species
Frommer (1996), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998a), Wagner (1935, 1939a), HN
Aphrophora salicina (Goeze, 1778) Aphrophora grisea Haupt, 1919; Aphrophora salicis unicolor Haupt, 1919 Many published reports, including recent ones, are in need of revision due to former taxonomic uncertainties, which were only resolved by Wagner (1948c) in a paper, which has received little attention.
B VI – B X; egg, 1 gen. Adults and nymphs on Salix alba and S. purpurea, less frequently on S. viminalis, S. cinerea, S. fragilis, a handful of records also from S. aurita, S. caprea, S. triandra and others. A few nymphs (probably fallen individuals) have been found in the herb layer. Widespread in Germany and usually in intermediate frequency, but more confined to lowlands and the larger river valleys than A. pectoralis Mats. Usually found below 600 m a.s.l., locally up to 850 m a.s.l. Fischer (1972), Heller (1987a), Lauterer & Malenovsky (1995), Niedringhaus (1991), Reimer (1992), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Wagner (1935, 1939a), HN
Philaenus spumarius (Linnaeus, 1758) E V – B XI; egg, 1 gen. (see also Witsack 1973) Extremely eurytopic and polyphagous. Adults and spittle masses usually on dicotyledonous herbs (but also on grasses, ferns, horsetails, dwarf shrubs, and even adventitious shoots of shrubs and trees) in peaty to dry, sunny to moderately shady sites such as meadows, pastures, fens, open forests, shores of running and standing water, waysides, ruderal sites, dry grassland (where usually confined to margins and disturbed patches), primary dunes, salt marshes, subalpine stands of herbs, etc. Often abundant in low-input meadows and abandoned fields with dominating dicotyledonous herbs, where spittle masses may be extremely abundant on Silene flos-cuculi, Cirsium arvense, Urtica dioica, Ranunculus repens, Filipendula ulmaria and others. Also found in low numbers on plant taxa largely avoided by other Auchenorrhyncha, e.g. Rubiaceae, Boraginaceae, Primulaceae, Brassicaceae and Orchidaceae, but usually rare or absent in intensively managed grassland, where reproductive success is uncertain. Widespread and common in Germany, highest localities at 1800 m a.s.l. in the Bavarian and Allgäu Alps. Achtziger (1991), Fischer (1972), Nickel (1979), Niedringhaus (1991), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1988), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN and others; see also Halkka (1978), Halkka et al. (1967, 1978), Stewart & Lees 1996, Wiegert (1964), Whittaker (1968, 1973)
4.2.3 Membracidae Rafinesque, 1815 – Treehoppers Gargara genistae (Fabricius, 1775) E VI – M X; egg, 1 gen. In sunny to moderately shady, damp to dry sites on acidic to basic substrates. Adults and nymphs usually stay in small aggregations on woody species of Fabaceae and are
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79
attended by ants. The northern edge of the range runs through Jutland, but in Germany the species is widespread, though scattered, with strongholds both in areas of sand and sandstone (where usually occurring in intermediate to high frequency on Cytisus scoparius in heaths, on roadside embankments and along forest margins) and on sun-exposed, 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 arborescens, rarely on herbaceous species such as Coronilla, perhaps also Onobrychis). Food plant records from other countries include Cytisus nigricans, Trifolium campestre, Coronilla sp., Melilotus sp., Medicago sp., Glycyrrhiza sp. and Caragana arborescens. Highest localities in Swabia and the Bavarian foothills of the Alps are at 650 m a.s.l. only. Frommer (pers. comm.), Niedringhaus (1997), Post-Plangg & Hoffmann (1982), Schiemenz (1988), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN and others; see also Müller (1956), Ossiannilsson (1978), Schedl (1998)
Centrotus cornutus (Linnaeus, 1758) B V – M X, mainly M V – B VIII; nymph (usually 3rd and 5th instar), semivoltine (Müller 1984a). In moderately moist to dry sites with tall herbs or shrubs, usually along woodland margins and among herb stands in dry grassland, along hedges, forest roads and margins, along shores as well as in the interior of open forests with luxuriant undergrowth. Nymphs usually stay near the base of the host (mainly Vincetoxicum, Cirsium, Carduus and Urtica), adults ascend to the upper herb and shrub layer for feeding and oviposition and are often swept from low-growing woody plants (Populus, Quercus, Rubus, Prunus and others). Widespread in Germany, but often collected in low numbers only. Found up to at least 1200 m a.s.l. in the Bavarian Alps, up to 1500 m a.s.l. in Switzerland. Bußmann & Feldmann (1995), Fischer (1972), Heller (1987a), Müller (1984b), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998a), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN and others; see also Günthart (1987a)
Stictocephala bisonia Kopp & Yonke, 1977 Several nomenclatural changes have been proposed for this species in recent years. In most older papers the name Ceresa bubalus (Fabricius, 1794) was used. Caldwell (1949) placed it into the genus Stictocephala Stål, 1869. Kopp & Yonke (1977) studied Fabricius‘ type and found that it is not conspecific with the species introduced into the Old World, and created the new name bisonia. More recently De Andrade (1997), after a revision of Walker’s material, suggested that the species has already been described as Ceresa alta Walker, 1851. However, according to Dietrich et al. (1999) this synonymy is doubtful, and the identity of Walker’s type, which is a female, is uncertain.
M VII – E X; egg, 1 gen. Along woody margins and in tall herb stands, usually near rivers, streams and ditches; oviposition is reported to take place on low-growing, woody species of Rosaceae (apple, pear, plum, cherry, peach, blackthorn and others, thus locally becoming noxious), willows (e.g. Salix purpurea, S. aurita) and others, nymphs are usually found on herbaceous plants (Solidago gigantea, Medicago sativa, also invading cultivations). Additional
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food plants reported in the literature include Rosa, Cornus, Crataegus, Vitis, Populus, Ulmus, Coronilla, Melilotus, Chrysanthemum and Artemisia. The species was introduced from North America into Europe (presumably with twigs of fruit trees) where it was reported for the first time in 1912 from former Hungary. Later it spread over most parts of southern Europe (extending northward to Belgium in oceanic climates) and has reached parts of North Africa, Kirghizia and Kazakhstan. In Germany recorded for the first time in 1966 along the southern upper Rhine (Isteiner Klotz) and in 1972 in numerous sites between Lörrach and Kehl, although already found in 1952 in Alsace near Strasbourg (France). Meanwhile it occurs in the valleys of the Black Forest, down the Rhine valley at least to Speyer (since 1996) and Darmstadt (since 2000), up the Rhine to Radolfzell near Lake Constance, 400 m a.s.l. (1998) and up the Neckar valley to Nürtingen (since 1995). Has been reported to cause damage on grapevine and fruit trees by oviposition into young twigs in Italy. Dynort (pers. comm.), Gauss (1987), Hoffmann (1990), Hoffrichter & Tröger (1973), Müller (1956), Reder (pers. comm.), Remane (1972), Schuler (1952), SMNS, Voigt (pers. comm.), HN; see also Arzone et al. (1987), Emelyanov (1993), Günthart (1980), Nast (1987), Schedl (1995), Vidano (1963a, 1963b)
4.2.4 Cicadellidae Latreille, 1825 - Leafhoppers (s.str.) 4.2.4.1 Ulopinae Le Peletier & Serville, 1825 Ulopa carneae W. Wagner, 1955 In Austria adults and nymphs recorded IV – VIII (Wagner & Franz 1961); probably semivoltine, like the following species (see below). This species is known only from a few records in the eastern parts of the Alps between 700 and 2200 m a.s.l. on Erica carnea in open pine and spruce forests and in subalpine scrub. So far there are only two German records from the Bavarian Alps: Winkl near Berchtesgaden, 700 m a.s.l., 22.VIII.1996, 2 nymphs; Schwarzeck near Ramsau, 1000 m a.s.l., 24.VIII.1996, 1 †, 2 ‡‡, 2 nymphs. Nickel (1999b); see also Leising (1977), Wagner & Franz (1961)
Ulopa reticulata (Fabricius, 1794) Adults and nymphs I – XII; nymph and adult, semivoltine (Melber 1989). In peaty to moderately dry, sunny to moderately shady sites on acidic substrates, mainly heaths, bogs and forest margins, occasionally also in open pine and oak forests; the host plant is Calluna vulgaris. Widespread in Germany and particularly common in the north German plain and along the edge of the Alps, more scattered in middle parts, where locally absent. Frequently found at 1300 m a.s.l. in the Mittelgebirge and the Bavarian and Allgäu Alps, up to 1600 m a.s.l. in Styria (Austria). Fischer (1972), Melber (1989), Niedringhaus (1991), Reimer (1992), Schiemenz (1971a, 1975, 1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN; see also Wagner & Franz (1961)
Megophthalminae Kirkaldy, 1906
81
Utecha trivia (Germar, 1821) According to Orosz (1977) specimens which probably originate from the same locus typicus, but named Ulopa lugens Germar, 1821, belong to a distinct species. The majority of European taxonomists, however, consider it only as a macropterous form of U. trivia (Germ.). The latter view is adopted here.
Adults I – XII, mainly E VII – V; adult (apparently only ‡‡), 1 gen. In sunny and oligotrophic, moderately dry to dry and low-vegetated (preferentially grazed) sites usually on basic substrates. The host relationships are poorly known; the few published reports include Hippocrepis comosa, Echium vulgare, in England Plantago lanceolata. In Germany rather scattered and mainly in southern and eastern parts; northernmost localities are on a line from Bad Münstereifel to Mainz, Aschaffenburg, Bad Kissingen, Gotha, Sondershausen, Wittenberg and Dresden, frequently occurring at 700 m a.s.l. on the Swabian Jura, but recorded up to 1300 m a.s.l. in the Chiemgau Alps. Reimer (1992), Rombach (1999a), Schiemenz (1988), Schönitzer & Oesterling (1998b), SMNS, Wagner (1939a, 1951a), HN; see also Morris (1972), Szwedo (2001)
4.2.4.2 Megophthalminae Kirkaldy, 1906 Megophthalmus scabripennis Edwards, 1915 So far only E VI; egg, 1 gen. This species‘ range includes the Mediterranean region and parts of western Europe, where it is normally reported from sunny margins and open, xerothermic forests. From Germany there are only two records from sun-exposed rocky slopes in the valleys of the southern upper Rhine and the Nahe: Isteiner Klotz near Lörrach, c. 300 m a.s.l. and Bad Münster, c. 200 m a.s.l., 21.VI.1965, 2 ††, 1 ‡. Remane & Fröhlich (1994b), Coll. Kobel-Lamparski (Remane det.); see also Guglielmino (1993), Holzinger (1995b)
Megophthalmus scanicus (Fallén, 1806) E VI – E X; egg, 1 gen. In sunny, wet to moderately dry sites, mainly low-input meadows and pastures, heaths, fens, abandoned fields and waysides. Single macropterous individuals may be swept among tall herbs, shrubs and trees. Host plants are various species of Fabaceae (Trifolium dubium, Medicago lupulina, Lotus pedunculatus and others). Widespread and fairly common in most parts of Germany, frequently found in the Allgäu at 1200 m a.s.l., in the Bavarian Alps occasionally at 1500 m a.s.l. Achtziger (1991), Fischer (1972), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN and others; see also Wagner & Franz (1961)
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4.2.4.3 Ledrinae Kirschbaum, 1868 Ledra aurita (Linnaeus, 1758) M VI – B X, rarely E III; nymph (hibernating twice), semivoltine (Schiemenz 1988). In moist to moderately dry sites, in closed forests as well as in semi-open tree stands. Lives on the bark of various deciduous trees, preferentially Quercus, but frequently also Betula, Populus, occasionally Tilia, Fagus, Malus and Acer, also reported from Alnus and Corylus. Widespread in the lowlands of Germany, but collected in low numbers only, usually in light traps and tree eclectors, rarely in the sweep-net. Highest localities in the Mittelgebirge and the foreland of the Alps at 550 m a.s.l. only. Achtziger (1991), Büchs (1988), Fischer (1972), Frommer (1996), Niedringhaus (1997), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN
4.2.4.4 Macropsinae Evans, 1935 Oncopsis alni (Schrank, 1801) M V – E VIII, at higher altitudes until IX; egg, 1 gen. Monophagous on alders (Alnus glutinosa and A. incana) near standing or running water, in fen woods and on seepy slopes. Widespread in Germany, common and in high frequency on both alder species (also on planted A. incana in lowlands), extending up to at least 1000 m a.s.l. in the Mittelgebirge and the Bavarian Alps, found up to the subalpine belt in the northeastern Alps of Austria. Was recently found to be a potential vector of the alder yellows phytoplasma. Fischer (1972), Günthart (1988), Maixner & Reinert (1999), Niedringhaus (1991, 1997), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1949a, 1951a), HN; see also Wagner & Franz (1961)
Oncopsis appendiculata W. Wagner, 1944 E V – B VIII; egg, 1 gen. On Betula pendula, rarely on B. pubescens, usually on solitary trees or along forest margins and alleys. Often occurs together with other species of Oncopsis, but apparently confined to damp or dry sites of lower altitudes. Widespread in Germany, but uncommon, locally absent and only in low to intermediate frequency. The highest localities are in the foreland of the Alps at c. 600 m a.s.l. Freese (pers. comm.), Frommer (pers. comm.), Schiemenz (1988), SMNS, Wagner (1949a), Nickel & Remane (1996), HN
Oncopsis tristis (Zetterstedt, 1840) B VI – B VIII, occasionally until B IX; egg, 1 gen. Monophagous on birches (Betula pendula and B. pubescens), but highly eurytopic, in raised bogs as well as dry grassland with scattered trees, on solitary trees as well as in open
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forests, often also in urban areas. Widespread and common in Germany, in high frequency on the hosts, although in smaller numbers than O. flavicollis (L.). Highest localities are at 1200 m a.s.l. in the Alps and the Black Forest; in North Tyrol (Austria) breeding is reported from 1900 m a.s.l. Fischer (1972), Niedringhaus (1991, 1997), Reimer (1992), Remane & Reimer (1989), Schiemenz (1988), Wagner (1951a), HN; see also Leising (1977)
Oncopsis avellanae Edwards, 1920 So far E V – E VI; egg, 1 gen. On Corylus avellana (in Britain also on Alnus glutinosa) in damp to dry woodland sites, usually along forest roads and margins and on solitary shrubs in meadows and pastures. Widespread at least in the western half of Germany where occurring in low to intermediate frequency on the host. Recorded from the Kaiserstuhl, Allgäu, Rhön Mountains, Mainfranken, middle Hessen, southern Niedersachsen and adjacent parts of Thuringia, the Lüneburger Heide, the island of Borkum, eastern Holstein and Angeln, up to 800 m a.s.l. According to current knowledge these localities are at the eastern edge of the range, but the species is likely to be under-recorded or misidentified and may be more widespread in Europe. Otherwise it is only known from France, the Netherlands and Great Britain. Fischer (1972), Remane & Fröhlich (1994b), HN; see also Claridge et al. (1977), della Giustina & Remane (2001), Nast (1987)
Oncopsis carpini (J. Sahlberg, 1871) M V – E VIII; egg, 1 gen. On Carpinus betulus in various sites, usually along forest margins and on solitary trees, but also in the interior of forests (where difficult to sample with the sweep-net), in parks and even in hedges in urban areas. Widespread in Germany and usually in high frequency on the host, although only scarcely recorded from the northwest German plain; highest localities are at 800 m a.s.l. in the Mittelgebirge and the Alps. Achtziger (1991), Fischer (1972), Frommer (pers. comm.), Günthart (1988), Post-Plangg & Hoffmann (1982), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1949a), HN
Oncopsis flavicollis (Linnaeus, 1761) group This taxon is morphologically extremely variable. In Wales, according to bioacoustic and morphometric studies of Claridge & Reynolds (1973) and Claridge & Nixon (1986), it is likely to consist of three distinct species. This may also be true for central European populations.
M V – VII, occasionally until B IX; egg, 1 gen. Eurytopic on birches (Betula pendula and B. pubescens) in much variable sites, in forests as well as on solitary trees, in bogs, parks, gardens, in dry grassland and others. Breeding has been documented at 1900 m a.s.l. near the tree line of the Tyrolian Alps (Austria). Widespread in Germany, extremely abundant and in very high frequency on the
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hosts. Frequently recorded at 1200 m a.s.l. in the Black Forest and the Allgäu, but probably extending higher up. Fischer (1972), Niedringhaus (1991, 1997), Reimer (1992), Remane & Reimer (1989), Schiemenz (1988), Wagner (1935, 1939a, 1951a), HN and others; see also Leising (1977)
Oncopsis subangulata (J. Sahlberg, 1871) E V – E VIII; egg, 1 gen. On birches, preferentially on Betula pendula, in Wales also reported from B. pubescens. Widespread in Germany, but less common than the preceding species, only in low to intermediate frequency and apparently confined to lower altitudes (found only up to 700 m a.s.l.), with strongholds in river floodplains. Fischer (1972), Freese (pers. comm.), Günthart (1988), Niedringhaus (1997), Schiemenz (1988), Wagner (1949a), HN; see also Claridge & Reynolds (1972)
Pediopsis tiliae (Germar, 1831) B VI – B VIII, occasionally M IX; egg, 1 gen. On lime (Tilia cordata, T. platyphyllos, perhaps also ornamental species), mainly in forests, along alleys and on solitary trees (also in urban areas). Widespread in the lowlands of Germany and in intermediate to high frequency, but usually low abundance; more scattered in north Germany and the foreland of the Alps. Highest localities at 600 m a.s.l. only. Fischer (1972), Günthart (1988), Reimer (1992), Schönitzer & Oesterling (1998b), Remane & Wachmann (1993), Schwoerbel (1957), Wagner (1935, 1939a), HN
Most members of the genus Macropsis are very difficult to identify, due to only slight differences in genitalic structures and considerable intraspecific variation of colouration. Hence, distribution patterns shown in this book are often incomplete. Wagner (1941a, 1950, 1953, 1964) contributed fundamentally 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), Tishechkin (1999) and Nickel (1999a) added further information. Just before this book was printed, Tishechkin (2002) presented a detailed revision of east and central European species, including a key with all relevant characters. In many cases, a reliable identification is only possible after these papers.
Macropsis albae W. Wagner, 1950 Hamilton (1980) suggested that this name was a junior synonym of M. ocellata Provancher, 1872, which is suspected to be introduced from Europe to North America. However, Provancher’s type is lost; the author did not even give information on sex and type locality and thus, its identity is highly uncertain. Later, Hamilton (1983b) designated a neotype from Canada, choosing an individual of the introduced species dwelling on Salix alba. Although Nast (1987), Schiemenz (1988), and others followed him, this opinion is rejected here, due to the extremely critical taxonomic situation among green species of Macropsis, and the unambiguous description of M. albae W.Wg.
E V – E VIII; egg, 1 gen. On Salix alba, probably also on its hybrids, usually near running or standing water. Widespread particularly in middle and southern parts of Germany and usually in high
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frequency on the host; less common in the north German plain, where reaching the edge of the range. Northernmost localities are on the island of Langeoog, near Hamburg, Kiel and Neubrandenburg, recorded up to at least 750 m a.s.l. near the edge of the Alps. The locus typicus is Campow near Ratzeburg. Lauterer & Malenovsky (1995), Mölleken & Topp (1997), Niedringhaus (1991), Reimer (1992), Remane (pers. comm.), Schiemenz (1988), Wagner (1950, 1951a), HN
Macropsis gravesteini W. Wagner, 1953 So far M VI – M VIII; egg, 1 gen. Like the preceding species on Salix alba, but rather uncommon and in low frequency. The distribution in Germany and Europe is imperfectly known, but the species has probably often been overlooked or misidentified. German records are from near Lübeck, Dannenberg, Düsseldorf, Halle, Marburg and Bad Nauheim, mostly in larger river valleys below 200 m a.s.l. Mölleken & Topp (1997), Remane (pers. comm.), Remane & Fröhlich (1994b), HN
Macropsis prasina (Boheman, 1852) Recently Tishechkin (1994) clarified the status of two further pale green Macropsis species, both described from central Asia and living on Salix viminalis. At least one of them – M. microcera Vilbaste, 1982 – is also found in European Russia. Hence, much of the central European specimens – including most of those cited in this book – are in need of revision. However, a number of German specimens which have been revised by Tishechkin (pers. comm.) belonged to M. prasina (Boh.).
B VI – B IX; egg, 1 gen. On hairy-leaved willows in damp to wet sites, in the study area Salix cinerea, S. viminalis, S. aurita and S. caprea. Apparently widespread and common in the lowlands of Germany, but imperfectly documented; in many places it occurs in intermediate to high frequency on the hosts. Frequently recorded to at least 600 m a.s.l. in the Mittelgebirge and the Alps, occasionally up to 1200 m a.s.l. Mölleken & Topp (1997), Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992), Wagner (1950), Nickel & Remane (1996), HN; also Tishechkin (pers. comm.)
Macropsis viridinervis W. Wagner, 1950 B VI – M VIII; egg, 1 gen. On Salix triandra, presumably also S. pentandra, in river floodplains, along streams and lake shores. Often overlooked or misidentified, but occurring at least scattered in the lowlands of southern and middle parts of Germany, with strongholds in the larger river valleys, usually in intermediate frequency on the host, although reaching the edge of the range. Northernmost records from near Rinteln an der Weser, Hamburg (locus typicus), Dannenberg, Halle and Dresden. Also recorded to the south of Stettin (northwestern Poland) and up to 700 m a.s.l. in the foothills of the Alps. Nicolaus (1957), Wagner (1950, and unpublished data), HN; see also Holzinger (1999a)
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Macropsis notata (Prohaska, 1923) Macropsis vestita Ribaut, 1952 sec. Tishechkin (2002)
B VI – E VIII; egg, 1 gen. In intermediate frequency on Salix triandra, less commonly on S. fragilis, usually in river floodplains. Imperfectly documented from Germany, but probably widespread in the lowlands – at least between the Danube and the coast. Northernmost localities are near Höxter, on the island of Amrum, near Plön, Heiligenhafen, Bad Frankenhausen and Meißen, only up to at most 400 m a.s.l. Not recorded from Denmark and Scandinavia, in Poland apparently confined to southern parts. Nicolaus (1957), Remane (pers. comm.), Schwoerbel (1957), SMNS, Wagner (1950, 1951a), HN; see also Nast (1976a), Ossiannilsson (1981)
Macropsis marginata (Herrich-Schäffer, 1836) M V – E VIII; egg, 1 gen. Monophagous on Salix purpurea, eurytopic near running or standing water. Distribution in Germany imperfectly documented, but common and in high frequency on the host, at least in middle and southern parts. The northern edge of the range apparently runs through northern Germany; northernmost localities are near Cologne, Rinteln an der Weser, Hamburg and Greifswald, up to 1300 m a.s.l. in the Bavarian Alps. The locus typicus is near Regensburg. Fischer (1972), Frommer (pers. comm.), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1950), HN
Macropsis infuscata (J. Sahlberg, 1871) B VI – E VIII; egg, 1 gen. At lower altitudes (including the Mittelgebirge and the foothills of the Alps, up to at least 900 m a.s.l.) this species lives on Salix caprea in forest glades and clearings, along forest margins and roads, in hedges and ruderal sites; in the Alps and their foothills (recorded between 700 and 1200 m a.s.l.) it is also found on S. myrsinifolia along running water and on seepy slopes. It is widespread and common in Germany and occurs in intermediate to high frequency on S. caprea, less frequently on S. myrsinifolia. Fischer (1972), Frommer (pers. comm.), Niedringhaus (1997), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1950, 1951a), HN
Macropsis cerea (Germar, 1837) Central European populations currently interpreted as M. cerea (Germ.) are likely to comprise several distinct species.
B VI – B IX; egg, 1 gen. On various species of willows, in the lowlands mainly Salix viminalis, S. triandra, S. fragilis, S. cinerea and S. caprea, in the Mittelgebirge and Alps mainly S. aurita, S. purpurea and S. myrsinifolia. Widespread in Germany, occurring in intermediate to high frequen-
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cy, but rather low individual numbers on the hosts. Highest localities are at 1200 m a.s.l. in the Mittelgebirge and the Allgäu Alps. Fischer (1972), Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1950, 1951a), HN
Macropsis remanei Nickel, 1999 E VI – M IX; egg, 1 gen. Monophagous on Salix eleagnos along streams and rivers in mountains and their foreland. This species was described only recently from the Bavarian Alps. At present, there are 10 German localities, near Berchtesgaden (locus typicus: Ramsau), along the upper Isar between Mittenwald and Bad Tölz, and along the Loisach near Garmisch-Partenkirchen, at altitudes between 650 and 1400 m a.s.l. Furthermore, the species has been found in various localities in northern Italy (Friuli Venezia Giulia: Tagliamento) and Slowenia. Probably it is more widespread in the Alps and perhaps other European mountains. Nickel (1999a), HN; see also Löcker (2003), Tishechkin (2002)
Macropsis najas Nast, 1981 M VI – E VIII; egg, 1 gen. Until recently this species was only known from the original description from Poland, but later it was found in large parts of central Europe. Meanwhile there are more than 25 records mainly from middle parts of Germany (approximately between a line running from Cologne to Göttingen, Stendal and Görlitz in the north and another one from Speyer to Würzburg and Bayreuth in the south, also near Lüneburg). It lives monophagously on Salix alba along rivers and streams, recorded only below 400 m a.s.l. Otherwise it is also known from northern Italy, eastern France and Austria. Frommer (pers. comm.), Remane (1995), Nickel (1997), HN; see also Holzinger et al. (1996), Nast (1981)
Macropsis haupti W. Wagner, 1941 This species has been mistaken for M. cerea (Germ.) for a long time, until Wagner (1941a), after studying Germar’s types, recognized its distinctiveness and created a new name. However, Ossiannilsson (1948) studied the type of Pediopsis planicollis Thomson, 1870, originally described from Sweden, and erroneously declared them to be conspecific, thus rendering M. haupti W.Wg. a junior synonym. Subsequently, Wagner (1950), in his exhaustive monograph on central European Macropsis species, accepted this opinion. After studying this paper, Ossiannilsson (1954) realized his error and declared M. planicollis (Thoms.) a junior synonym of M. cerea (Germ.).
B VI – M IX; egg, 1 gen. Monophagous on Salix purpurea, mainly along rivers and streams. Often syntopic with M. marginata (H.-S.), although usually in lower numbers and frequency, recorded only up to 1000 m a.s.l. Widespread in middle and southern parts of Germany, but only scattered, reaching the edge of the range in the north German plain. Northernmost localities are near Rinteln an der Weser, Dannenberg and Frankfurt an der Oder; also reported from northwestern parts of Poland (to the south of Stettin).
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Review of species
Fischer (1972), Reimer (1992), Schönitzer & Oesterling (1998b), Wagner (1950, 1951a), HN
Macropsis impura (Boheman, 1847) B VI – M VIII; egg, 1 gen. On Salix repens, occasionally on low-growing individuals of Salix aurita. Strongholds in Germany on coastal dunes near the North and Baltic Sea, where locally common; very sporadic elsewhere, usually in peaty wetlands of the northern plains, the Mittelgebirge and along the edge of the Alps, where found to at least 800 m a.s.l. (but up to 1050 m a.s.l. in Styria, Austria); not recorded from southwestern parts (Baden-Württemberg, Rheinland-Pfalz, Nordrhein-Westfalen). Niedringhaus (1991), Remane & Reimer (1989), Schiemenz (1988), Schönitzer & Oesterling (1998b), Trümbach (1959), Wagner (1950), HN
Macropsis fuscinervis (Boheman, 1845) M VI- M VIII, according to Wagner (1951a) from M V onwards; egg, 1 gen. Monophagous on Populus tremula, usually along forest margins and roads, in clearings and ruderal sites. Widespread in Germany, but only in intermediate frequency and low individual numbers on the host, recorded up to 700 m a.s.l. in the Mittelgebirge and the Bavarian Alps. Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1941a, 1941c), HN
Macropsis graminea (Fabricius, 1798) B VI – M VIII; egg, 1 gen. Monophagous on Populus nigra, on the nominate form as well as on var. italica, occasionally also on hybrids, where reproductive success may be uncertain; mainly in river floodplains, parks and urban settlements, usually in intermediate to high frequency, but low individual numbers. In Germany apparently confined to southern and middle parts (to at least 600 m a.s.l.), localities near Cologne, Göttingen, Hamburg, Lübeck and Berlin are probably near the northern edge of the range, although there are a few older records from Norway and Sweden. Frommer (pers. comm.), Günthart (1988), Nikusch (1976), Reimer (1992), Schiemenz (1988), Wagner (1941a, 1941c, 1951a), HN; see also Ossiannilsson (1981)
Macropsis vicina (Horváth, 1897) So far M VI – B VIII; egg, 1 gen. In Europe this species occurs mainly in southern parts, but extends locally northward to Champagne (France), Germany and eastern Poland. The host plant is Populus alba. From Germany it is only known from 4 records in the Kaiserstuhl of southwestern Baden, the Kyffhäuser of Thuringia and the vicinity of Berlin: Achkarren, c. 200 m a.s.l., 18.VI.1971, 3 ††, 12 ‡‡ and nymphs; Oberbergen, c. 400 m a.s.l., 15.VI.2002, 1 †, 1 ‡; Günserode, 160 m a.s.l., 08.VII.1995, 2 ††; Berlin, Tegel, c. 40 m a.s.l., 10.VIII. 2002. Remane & Fröhlich (1994b), Stewart (pers. comm.), HN; see also Nast (1976a)
Macropsinae Evans, 1935
89
Macropsis elaeagni Emelyanov, 1964 In the Czechia Republic, Slovakia and Austria B VI – B IX (Holzinger & Remane 1994; Lauterer 1984); egg, 1 gen. Originally this species was confined to Kazakhstan and southern Russia and lives on Elaeagnus. In recent decades it has expanded its range and appeared in the Balkan region and southeastern parts of central Europe, where it has colonized secondary plantations along waysides and in public parks. Dispersal is suspected to take place mainly through cuttings. In Germany it was found only in two localities in the northern upper Rhine plain: Weiterstadt, 120 m a.s.l., 03.VIII.1996, 29 ††, 15 ‡‡, and Germersheim, 17.VIII.1998, 3 ‡‡, both along waysides on E. angustifolia, c. 120 m a.s.l., but it is probably more widespread at least in southern parts. HN; see also Emelyanov (1964b), Lauterer (1984)
Macropsis glandacea (Fieber, 1868) Pediopsis mendax Fieber, 1868; Pediopsis ulmi Scott, 1873
B VI – B IX; egg, 1 gen. On elm, usually Ulmus minor and U. x hollandica, at least single specimens found on U. laevis, usually in river floodplains, parks, along alleys and hedges. In Germany at the northern edge of the range, largely restricted to the valleys of larger rivers and their basins (so far found along the Rhine, Main, Neckar, Lahn, Danube, Altmühl, Leine, Unstrut, Saale, Elbe and Neiße). The northernmost localities are near Cologne, Göttingen, Hamburg, Plön, Stendal and Berlin, the highest ones at 400 m a.s.l. Frommer (pers. comm.), Nikusch (1976), Remane (1987), Remane & Wachmann (1993), Schiemenz (1988), SMNS, Wagner (1939a, 1941a, 1941c), Nickel (1994, 1999b), HN
Macropsis brabantica W. Wagner, 1964 The specific identity of this taxon has yet to be demonstrated. According to Wagner (1964) it lives monophagously on Rubus caesius and could not be reared on R. idaeus in the laboratory. However, German populations of Macropsis on R. caesius, most of which clearly belong to M. fuscula (Zett), often include some individuals showing intermediate colouration of M. fuscula (Zett.) and M. brabantica W.Wg.
So far only E VIII; probably egg, 1 gen. In Germany only reported from two sites in Brandenburg to the south of Berlin: Klein Köris and Groß Köris, 22. and 24.VIII.1995, „an einer niedrigwüchsigen Rubus-Art (Rubus caesius?) ( ... ) sowohl im Schatten eines Mischwaldes als auch in einem Erlenbruchwald“. The species was described from several localities in the Netherlands and has since been reported only from Germany and middle Russia (see below). Remane (1995); see also Tishechkin (2002), Wagner (1964)
Macropsis fuscula (Zetterstedt, 1828) B VI – M IX, occasionally until B X; egg, 1 gen.
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Review of species
On Rubus idaeus, R. fruticosus and R. caesius in various types of forests, in clearings, along hedges, in tall herb stands and gardens. Widespread in Germany, frequently occurring to at least 1300 m a.s.l. in the Mittelgebirge and the Alps, but single specimens were found up to 1600 m a.s.l. Has been identified as a potential vector of Rubus stunt disease (RSD) in various parts of Europe. Fischer (1972), Frommer (1996), Niedringhaus (1991, 1997), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1939a, 1964), HN; see also Brˇcák (1979)
Macropsis scotti Edwards, 1920 So far E VII – M VIII, in Britain VII – X (Le Quesne 1965); egg, 1 gen. This species occurs from Morocco through western Europe to England and Ireland. From Germany it is recorded only in two localities in the extreme west: Eifel Mountains, Niederzissen, c. 300 m a.s.l., mid August 1974, 1 ‡, along a hedge, and Stuttgart, c. 250 m a.s.l., 21.VII.1937, 1 specimen. Is reported to live monophagously on Rubus fruticosus. Post-Plangg & Hoffmann (1982) (Remane det.), Tishechkin (2002, and pers. comm.); see also Wagner (1964)
Macropsis megerlei (Fieber, 1868) B VII – M VIII; egg, 1 gen. Until recently this species was known to occur in central Europe from a few records only. It lives on shrubs of roses in sunny, xerothermic sites (mainly on limestone, gypsum, schist, etc.). Host plants are mainly Rosa rubiginosa, less frequently R. spinosissima and R. canina; occasionally also ornamental R. glauca and R. rugosa. In Germany it is widespread at least in the middle parts and difficult to collect, but probably fairly common on favourable sites. There are records from Mainfranken, northern Baden, middle Hessen, Thuringia, Palatinate, as well as along the northern and southern edge of the Harz Mountains, up to 400 m a.s.l. To the north of Germany it is known only from two single specimens from southern Norway. Kupka (1925), Wagner (1939a), Schiemenz (1988), SMNS, Nickel (1997), HN; see also Lauterer (2000), Ossiannilsson (1981)
Macropsis scutellata (Boheman, 1845) B VI – M IX; egg, 1 gen. Lives monophagously on Urtica dioica in damp to moderately wet, sunny to moderately shady sites, usually in river floodplains, along shores of lakes and ponds and in ruderal sites with tall herbaceous vegetation. Widespread in Germany, although seldom recorded due to secretive life habits, and apparently largely confined to lowland areas (up to at most c. 600 m a.s.l.); however, there is a single specimen from the Allgäu Alps at 1200 m a.s.l. Fischer (1972), Frommer (pers. comm.), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a, 1952, 1964), HN
Agalliinae Kirkaldy, 1901
91
Macropsis mulsanti (Fieber, 1868) In Switzerland and Austria so far M VII – B IX; egg, 1 gen. Monophagous on Hippophae rhamnoides, reported to live on sunny and rocky slopes in Switzerland. In Europe this species is mainly confined to southern parts; to the north of the Alps it has only been found in a few localities near Lake Constance and along the upper course of the Rhine: Bludenz and Röthis (both Vorarlberg, Austria), Strasbourg (Alsace, France) and southern Baden: Heitersheim, 06.VIII.1997, 3 ‡‡ on a roadside embankment. HN; see also Moosbrugger (1946), Remane & Fröhlich (1994b), Tishechkin (1993)
Hephathus nanus (Herrich-Schäffer, 1835) Palearctic species of this genus were recently reviewed by Tishechkin (1999), proving that the Asian H. achilleae Mityaev, 1967 also occurs in the eastern half of Europe, notably in large parts of European Russia, southern Moravia (Czech Republic), Hungary and probably Finland. Thus, it may be more widespread in central Europe as well. The author also pointed out that the drawings of Ossiannilsson (1981) do not refer to H. nanus (H.-S.), but to H. achilleae Mity.
So far E VI – E VIII; egg, 1 gen. In sunny, oligotrophic and xerothermic sites with short vegetation, usually in heavily grazed pastures on basic substrates, perhaps associated with Cirsium acaule. In Germany only locally and at the northern edge of its range, mainly in warmer regions of eastern and southern parts (Saale-Unstrut region, Thuringian Basin, Mainfranken, valleys of the Neckar, Nahe and upper Rhine). Northernmost localities are near Mainz, Northeim, Magdeburg and Berlin; found up to at least 500 m a.s.l. near Arnstadt and Munich, but there is also an unconfirmed record from at least 1000 m a.s.l. from the Bavarian Alps. The locus typicus is near Regensburg. Haupt (1925), Remane (1987), Schönitzer & Oesterling (1998b), Schiemenz (1969, 1988), SMNS, Wagner (1939a), ZIMH, Nickel (1994), HN; for identification see Tishechkin (1999)
Macropsidius sahlbergi (Flor, 1861) M VI – B VIII; egg, 1 gen. (Schiemenz 1988). This species occurs mainly in steppe regions of central Asia and southeastern Europe, with rather isolated populations in xerothermic grassland sites of central Europe, from where it is reported to live on Artemisia campestris. In Germany it is confined to northeastern parts and known only from 5 localities near Pasewalk, Angermünde, Eberswalde, Stendal and Meißen, all below 150 m a.s.l. The last record dates from 1965; thus the current status is uncertain. Schiemenz (1969, 1988), Wagner (1941a)
4.2.4.5 Agalliinae Kirkaldy, 1901 Agallia brachyptera (Boheman, 1847) B VII – E X; eggs overwinter in secondary parapause terminated by cold, 1 gen. (Witsack 1985)
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Review of species
In sunny to shady, damp to wet, occasionally moderately dry or peaty sites, notably low-input meadows and pastures, abandoned fields and forests. Presumably this species lives polyphagously on various species of Fabaceae and Asteraceae (Taraxacum, also reported from Trifolium, Onobrychis, Achillea, as well as Rumex). Widespread in Germany and fairly common, found up to at least 900 m a.s.l. in the Mittelgebirge and the Alps; frequently recorded at 2000 m a.s.l. in Trentino (northern Italy). Niedringhaus (1991), Nikusch (1976), Reimer (1992), Remane (1958), Schiemenz (1988), Wagner (1935), Nickel & Achtziger (1999), HN; see also Ossiannilsson (1981)
Agallia consobrina Curtis, 1833 M VII – M V, from M VI onwards along the upper Rhine; adult (only ‡‡?), 1 gen. In moderately shady, moderately dry to moist sites on acidic to basic substrates, usually in rather warm situations, mainly open deciduous forests with luxuriant growth of herbs (coppices, coppice-with-standards, occasionally also floodplain forests and alder fen woods), dry grassland and abandoned vineyards with scattered shrubs and trees. Lives on various species of Lamiaceae (Glechoma, Teucrium, probably also Lamium, Stachys and others), perhaps also grasses and Urtica dioica. In Germany widespread between the Danube and the northern edge of the Mittelgebirge, and particularly common in viticultural regions, rather scattered in northern Germany and not recorded to the south of the Danube. Most localities are below 500 m a.s.l., although found up to at least 900 m a.s.l. in the southern Black Forest. Kuntze (1937), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schiemenz (1988), Schwoerbel (1957), Trümbach (1959), Wagner (1935, 1939a, 1951a), HN
Anaceratagallia austriaca W. Wagner, 1955 So far B III – M VI, in the Valais (Switzerland) according to Dlabola (1971) E IX; adult, 1(?) gen. According to existing data, this species is a rare endemic of the Alps and south German limestone regions. In Germany it was only found in the Franconian Jura, Kipfenberg, c. 350 m a.s.l., 06. and 08.III.1961, „eine individuenreiche Population ( ... ) auf einem Steppenheide-Südhang ( ... ) unter Thymus-Polstern“, as well as in the middle Main valley, Gambach, 300 m a.s.l., 04.VI.1993, 3 ‡‡, and 19.VI.1994, 2 ‡‡, along the upper edge of a sun-exposed, steep and rocky hillside. Otherwise only reported from a few xerothermic sites in Styria (Austria), the Vintschgau (Italy) and the Valais (Switzerland), between 800 and 2300 m a.s.l. Remane (1961b), HN; see also Wagner (1955), Dlabola (1971), Günthart (1987a)
Anaceratagallia frisia (W. Wagner, 1939) This taxon appears to be closely related to A. laevis Ribaut, 1935, which is widespread from central Asia to western Europe, but it is geographically isolated and confined to Poland (inland dunes along the lower Oder), northern and middle parts of Germany, the Netherlands (West Friesian Islands), and perhaps Great Britain (see Remane 1995).
Agalliinae Kirkaldy, 1901
93
E VI – E V; adult (only ‡‡), 1(?) gen. Very scattered, but locally abundant in various xerothermic sites with incomplete cover of vegetation, in dune areas with dominating Corynephorus and Festuca on the East Frisian Islands (Norderney, Baltrum, Langeoog, Spiekeroog, Wangerooge; locus typicus: Borkum), as well as in steppe-like dry grassland in the Kyffhäuser and along the lower Oder, one single record is from a mining area to the south of Berlin. Highest localities are at 250 m a.s.l. The host plants are not known, but may include various species of Fabaceae or Thymus. Bröring & Niedringhaus (1989a, 1989b), Niedringhaus & Olthoff (1993), Remane (1995), Schiemenz (1969, 1988), Wagner (1939b), HN
Anaceratagallia ribauti (Ossiannilsson, 1938) I – XII, mainly VIII – VI; adult (at least a few ††), oocytes overwinter in photoperiodically controlled oligopause, ‡‡ in thermic quiescence, 1 gen. (Witsack 1985, and pers. comm.). In sunny, dry to damp, occasionally also moist sites on acidic to basic substrates, mainly disturbed, grazed or trampled patches in dry grassland, in low-and medium-input meadows, abandoned fields, mining areas, along waysides and on inland dunes. Often syntopic with A. venosa (Geoffr.), but more dominant in acidic and sandy sites and confined to the lowlands. Most populations apparently live on Plantago lanceolata and P. major, but various dicotyledonous herbs have been reported as additional hosts from laboratory rearings (particularly Medicago, Trifolium and Onobrychis, as well as some species of Lamiaceae and Scrophulariaceae). Widespread in the lowlands of Germany, with strongholds in warmer regions and the north German plain, where locally common and dominant in favourable sites, but rare or absent in most parts of the Mittelgebirge, the Alps and their foreland. The highest localities are only at 500 m a.s.l., but at 1500 m a.s.l. in the Engadine (Switzerland). Achtziger (1991), Fischer (1972), Niedringhaus & Olthoff (1993), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schwoerbel (1957), Wagner (1939a, 1951a), Witsack (1985), Nickel & Achtziger (1999), HN; see also Günthart (1987a)
Anaceratagallia venosa (Geoffroy, 1785) Mainly M VII – M X, in southwestern Germany from E V onwards; according to Witsack (1985) eggs overwinter in secondary parapause terminated by cold and followed by thermic quiescence, 1 gen. In similar habitats as A. ribauti (Oss.) and occasionally syntopic, but generally preferring dryer, more basic and less vegetated sites (including quarries and old farm roads) and extending up to higher altitudes in the Mittelgebirge and Alps. Lives on low-growing dicotyledonous plants (apparently mainly Lotus corniculatus, but also reported from Hippocrepis comosa and Thymus). Widespread in Germany and locally common, usually among the dominant species in sparsely vegetated calcareous grasslands of southern and eastern parts of Germany; less common in other habitats, but found to at least
94
Review of species
1000 m a.s.l. in the Erzgebirge, the Bavarian and Black Forest, frequently up to 2000 m a.s.l. in the Bavarian and Allgäu Alps. In North Tyrol (Austria) breeding has been recorded in the alpine belt at 2450 m a.s.l. Fischer (1972), Müller (1978), Niedringhaus (1991), Reimer (1992), Remane (1987), Rombach (1999b), Wagner (1937b, 1939a, 1951a), Witsack (1985), HN; see also Leising (1977)
Dryodurgades antoniae (Melichar, 1907) B VIII – B XI; egg, 1 gen. This is a western European species known only from Portugal, Spain, France, the Netherlands and western Germany. Records from Bulgaria are in need of revision. In Germany there are 8 known sites. Localities at the edge of the range include Cologne, Braunschweig, Marburg and Landau/Pfalz. Lives on Cytisus scoparius in clearings and abandoned fields, along margins, embankments, etc.; highest records are from c. 350 m a.s.l. only. Frommer (pers. comm.), Jöst (1962), Remane & Fröhlich (1994b), Nickel (1994), HN
Dryodurgades reticulatus (Herrich-Schäffer, 1834) B VIII – E V; adult, 1 gen. Along forest margins and in the vicinity of shrubs in xerothermic limestone sites. Vicia tenuifolia is the host plant, although adults were swept from Pinus sylvestris, where hibernation may take place. In Germany confined to the southern parts, only known from a few records from near Regensburg (locus typicus), Stuttgart and the Triassic limestone belt of Mainfranken and adjacent areas of Baden and Thuringia (near Tauberbischofsheim, Würzburg, Karlstadt, Bad Neustadt and Meiningen). Highest localities in these regions are at c. 550 m a.s.l., but at 1300 m a.s.l. in the Engadine (Switzerland). Remane & Fröhlich (1994b), Wagner (1963), HN; see also della Giustina (1989), Günthart (1987a), Wagner (1963)
4.2.4.6 Idiocerinae Baker, 1915 Rhytidodus decimusquartus (Schrank, 1776) B VII – X, occasionally ‡‡ until E IV; mainly egg, 1 gen., published statements of 2 gen. (Müller 1956) refer to North America. On Populus nigra nigra and P. n. italica, probably also on hybrids, in river floodplains, urban settlements and open cultivated areas. Single individuals are swept from other woody species (e.g. Salix) and often fly into houses during autumn and winter nights. Widespread in the lowlands of Germany and usually in intermediate frequency, but low individual numbers; highest localities are at 550 m a.s.l. near the edge of the Alps. Feldtmann (1938), Fischer (1972), Müller (1956), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1939a, 1951a), HN; see also Tromellini et al. (1988)
Idiocerinae Baker, 1915
95
Idiocerus lituratus (Fallén, 1806) B VII – B X; egg, 1 gen. Usually in rather cool sites (notably fens, intermediate bogs, meadow valleys, forest glades, coastal dunes), mainly on hairy-leaved species of willows (Salix cinerea, S. aurita, S. caprea), near the coast preferentially on Salix repens. Single vagrants may be swept from S. viminalis, S. purpurea, S. alba, S. fragilis and other woody plant species. Widespread in Germany, occurring in intermediate to high frequency on the host plants, frequently found up to at least 900 m a.s.l. in the Mittelgebirge and the foothills of the Alps. Fischer (1972), Niedringhaus (1991, 1997), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN
Idiocerus vicinus Melichar, 1898 B VIII – B VI; adult (only ‡‡?), 1 gen. Along rivers and streams and in gravel pits, occasionally on roadside embankments, mainly on Salix purpurea, at least occasionally also on S. eleagnos. In Germany found only along the edge of the Alps and in the upper Rhine plain between Freiburg and Speyer, between 100 and 1150 m a.s.l., usually in low to intermediate frequency, but locally in large numbers. Nickel (1999a), HN
Idiocerus similis Kirschbaum, 1868 M VII – B X; egg, 1 gen. Monophagous on Salix purpurea along rivers and streams, in sand and gravel pits, occasionally also on roadside embankments. Widespread in middle and southern parts of Germany, found in low to intermediate frequency and rather low numbers on the host. The northern edge of the range runs roughly along the northern edge of the Mittelgebirge. Northernmost localities are near Cologne, Göttingen, Hamburg, Lüchow, Nauen and Potsdam, frequently found up to at least 900 m a.s.l. in the Bavarian Alps. Frommer (pers. comm.), Günthart (1988), Reimer (1992), Schiemenz (1988), Wagner (1935, 1939a), Nickel & Remane (1996), Nickel (1994), HN
Idiocerus stigmaticalis Lewis, 1834 M VI – E IX, occasionally until B XI; egg, 1 gen. On narrow-leaved species of willows near running and standing water, preferentially on Salix alba, but found in lower numbers on many others, notably S. fragilis, S. triandra, S. eleagnos, also S. viminalis, S. purpurea, S. caprea, S. cinerea and others; single individuals on dispersal flight may be swept from other deciduous trees such as Populus, Betula, Alnus and Acer. Widespread in Germany and common in most regions, in high frequency at least on S. alba; the highest records are at 1000 m a.s.l. in the Bavarian Alps on S. eleagnos.
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Review of species
Fischer (1972), Lauterer & Malenovsky (1995), Mölleken & Topp (1997), Niedringhaus (1991, 1997), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Trümbach (1959), Wagner (1935, 1939a), HN
Idiocerus herrichii (Kirschbaum, 1868) B VII – E V; adult, 1 gen. On Salix alba and S. fragilis near running or standing water. Widespread in the lowlands of Germany, but rather scattered and usually in low frequency and individual numbers, with strongholds in the north German plain and in larger river valleys (recorded along the Rhine, Neckar, Main, Lahn, Danube, Isar, Saale and their main tributaries, Elbe, Oder). The highest localities in these regions are at 600 m a.s.l., but at 1200 m a.s.l. in the Engadine (Switzerland). Achtziger (1991), Fischer (1972), Mölleken & Topp (1997), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), SMNS, Wagner (1939a, 1951a), HN; see also Günthart (1987a)
Metidiocerus elegans (Flor, 1861) Mainly E VII – X, occasionally III – M VI; egg (also adult?), 1 gen. On grey-leaved willows (Salix caprea, S. cinerea, S. aurita) in cool and moist situations, notably meadow valleys, river floodplains, fens and forest glades. Widespread in Germany, but rather scattered, in low frequency and individual numbers, found to at least 1100 m a.s.l. in the Mittelgebirge and the Bavarian Alps, at 1730 m a.s.l. in Switzerland. Fischer (1972), Niedringhaus & Olthoff (1993), Reimer (1992), Remane & Reimer (1989), Schiemenz (1988), Wagner (1935), Nickel (1999b), HN; see also Günthart (1987b)
Metidiocerus rutilans (Kirschbaum, 1868) Adults I – XII, mainly M VII – B VI; adult, 1 gen. Usually on narrow-leaved species of willows (Salix viminalis, S. purpurea, S. triandra, S. alba, less frequently S. eleagnos, S. fragilis), occasionally on S. caprea; recorded on Picea abies in autumn and winter. Widespread in middle and southern parts of Germany and in intermediate to high frequency on the 4 first-mentioned willow species, but often in low individual numbers. The edge of the range runs through northern Germany; the northernmost localities are near Lingen (Ems), Goslar, Dannenberg, Stendal and Neubrandenburg. Found up to at least 900 m a.s.l. in the Mittelgebirge, up to 1000 m a.s.l. in the Bavarian Alps. Fischer (1972), Mölleken & Topp (1997), Niedringhaus (1997), Schiemenz (1988), Schwoerbel (1957), Wagner (1951a), Nickel (1994), HN
Metidiocerus impressifrons (Kirschbaum, 1868) Adults I – XII, mainly M VII – M VI; Ad, 1 gen.
Idiocerinae Baker, 1915
97
On narrow-leaved species of willows, preferentially Salix viminalis, occasionally also S. purpurea and S. triandra, usually in river floodplains near water. In Germany recorded only from scattered sites and living at the northern edge of the range, although presumably under-recorded or misidentified. Most localities are along the Rhine, Main, Danube, Elbe, Saale, Oder and Neiße, single records also from the valleys of the Leine, Ahr, Lahn, Dill, Lech (near Füssen, 800 m a.s.l.) and Isar (near Mittenwald, 900 m a.s.l.); extending northward at least to Düsseldorf, Northeim, Lauenburg, Magdeburg, Berlin and Angermünde. At least single individuals have been found at 1200 m a.s.l. in the Engadine (Switzerland). Fischer (1972), Mölleken & Topp (1997), Remane (pers. comm.), Remane & Wachmann (1993), Schiemenz (1988), Wagner (1939a), Nickel (1994, 1999b), ZIMH, HN; see also Günthart (1987a)
Stenidiocerus poecilus (Herrich-Schäffer, 1835) M VII – M VI; adult, 1 gen. On Populus nigra nigra and P. n. italica, usually in river floodplains, although often away from water. Hibernating individuals were taken from coniferous trees in Piemonte (Italy). In Germany rather localized and mainly along the larger rivers (Elbe between Dresden and Boizenburg, Saale near Jena and Rudolstadt, Main near Aschaffenburg, Lahn near Marburg, Rhine between Waldshut and Cologne, Nahe near Bad Münster); a few records also from the surroundings of Berlin and Leipzig, the Oberlausitz, as well as near Böblingen and Balingen (up to 600 m a.s.l.). The locus typicus is near Regensburg. Frommer (pers. comm.), Günthart (1988), Remane (pers. comm.), Schiemenz (1988), SMNS, Wagner (1939a, 1951a), HN; see also Giordano et al. (2001)
Tremulicerus tremulae (Estlund, 1796) M VI – B XI; egg(?), 1 gen. Monophagous on Populus tremula, mainly in forest glades and along margins, but also in closed stands. Widespread in middle and northern parts of Germany and in low to intermediate frequency on the host, but not recorded to the south of the Main, where perhaps overlooked. Reported from the French side of the upper Rhine plain and most parts of Austria (although some of these records are in need of revision). The highest localities are at 700 m a.s.l. in the Mittelgebirge. Niedringhaus (1997), Reimer (1992), Schiemenz (1988), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN; see also Ribaut (1952), Wagner & Franz (1961)
Tremulicerus vitreus (Fabricius, 1803) B VI – B X; egg, 1 gen. On Populus nigra nigra, P. n. italica and their hybrids, in river floodplains and open cultivated land (also in urban settlements). Widespread in lowland areas of middle and southern parts of Germany and in intermediate to high frequency on the hosts; less common in northern Germany, where apparently at the edge of the range. Northern-
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Review of species
most localities are on the island of Norderney, near Lübeck, Halle and Berlin, found only up to 550 m a.s.l. in the Mittelgebirge and the foreland of the Alps. To the north of it, there is only one record from the 19th century from Denmark; another record is from near Stettin (northwestern Poland). Feldtmann (1938), Günthart (1988), Lauterer & Malenovsky (1995), Schiemenz (1988), Schwoerbel (1957), Wagner (1935, 1939a, 1941a), ZMB, Nickel & Remane (1996), Nickel (1997), HN; see also Ossiannilsson (1981)
Tremulicerus distinguendus (Kirschbaum, 1868) E VI – E X; egg, 1 gen. On Populus alba in river floodplains, parks and open cultivated land, vagrants also on other poplar species. Widespread in the lowlands of Germany, occurring northward to the East Frisian Islands, Holstein and Mecklenburg, usually in intermediate frequency and rather low individual numbers on the host, to the north of it only known from a few records from Denmark and southern Sweden. Highest localities are at 500 m a.s.l. in the Mittelgebirge and the foreland of the Alps. Frommer (pers. comm.), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Wagner (1935), HN; see also Gillerfors (2002), Ossiannilsson (1981)
Tremulicerus fulgidus (Fabricius, 1775) Idiocerus mesopyrrhus Kirschbaum, 1868 Idiocerus fuchsii Kirschbaum, 1868 (see Nickel & Remane 2002)
Adults I – XII, mainly E VII – B VII; adult (also ††), 1 gen. On Populus nigra italica and P. n. nigra along rivers and streams and in open cultivated land, occasionally also on poplar hybrids (but breeding has not been recorded on these); in winter on Picea, reported from France also on Calluna, Hedera, Pinus and Cupressus. Widespread in middle and southern parts of Germany and usually in intermediate frequency on the host, but confined to lowland areas (up to at most 600 m a.s.l.) and apparently absent from the north, the range extending at least to a line running from Cologne to Marburg, Göttingen, Stendal and Leipzig. Frommer (1996), Remane & Fröhlich (1994b), Schiemenz (1988), Nickel (1994), HN; see also Ribaut (1952)
Viridicerus ustulatus (Mulsant et Rey, 1855)
Adults I – XII, mainly M VII – M VI; adult, 1 gen. On Populus alba in river floodplains, parks and along waysides. Widespread in middle and southern parts of Germany, occurring in high frequency, although rather low individual numbers, but more scattered to the north of it, where presumably only for some decades. Northernmost records – all dating from the 1990s and many from urban settlements – are from the island of Norderney, near Göttingen, Dannenberg, Berlin and Frankfurt an der Oder; found only up to 500 m a.s.l. in the foreland of the Alps.
Idiocerinae Baker, 1915
99
Fischer (1972), Frommer (pers. comm.), Remane & Fröhlich (1994b), Remane & Wachmann (1993), Wagner (1939a, 1951a), Weis (pers. comm.), Nickel (1994), HN
Populicerus albicans (Kirschbaum, 1868) M VI – B X; egg, 1 gen. Monophagous on Populus alba in river floodplains, parks and along waysides, also in urban settlements. Widespread in the lowlands of Germany (up to at least 600 m a.s.l.) and usually in high frequency and individual numbers on the host, even on much isolated tree individuals. Fischer (1972), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a, 1951a), HN
Populicerus confusus (Flor, 1861) M VI – B X; egg, 1 gen. In river floodplains, along streams, in forest glades and among subalpine scrub, mainly on grey-leaved species of willows (Salix cinerea, S. aurita, S. caprea, S. viminalis, in the Alps and their foothills also S. myrsinifolia and S. appendiculata), occasionally found in rather low numbers on additional species (S. alba, S. fragilis, S. triandra). Widespread in Germany and fairly common, occurring up to at least 1200 m a.s.l. in the Mittelgebirge, up to 1500 m a.s.l. in the Bavarian Alps. Fischer (1972), Mölleken & Topp (1997), Niedringhaus (1991, 1997), Reimer (1992), Remane (1987), Remane & Reimer (1989), Schiemenz (1988), Wagner (1935), 1939a, 1951a), HN and others
Populicerus nitidissimus (Herrich-Schäffer, 1835) B VI- M X; egg, 1 gen. On Populus nigra italica, P. n. nigra and their hybrids, in cultivated land, urban settlements and river floodplains (also in closed forests). Widespread in the lowlands of Germany, in high frequency and individual numbers on the hosts, although less common in the north German plain, extending only up to 600 m a.s.l. in the Mittelgebirge and the foreland of the Alps. Büchs (1988), Günthart (1988), Mölleken & Topp (1997), Niedringhaus (pers. comm.), Reimer (1992), Remane (1987), Schiemenz (1988), ZIMH, HN
Populicerus laminatus (Flor, 1861) M VI – M X; egg, 1 gen. Monophagous on Populus tremula, mainly in forests and along their margins. Widespread in Germany, apparently only in low to intermediate frequency, but locally high individual numbers. A single tree stem eclector catch in northern Thuringia revealed 33.000 individuals within one season. The highest localities are at 700 m a.s.l. in the Mittelgebirge and the foothills of the Alps.
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Frommer (1996), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a, 1951a), HN
Populicerus populi (Linnaeus, 1761) M VI – M X; egg, 1 gen. Like the preceding species monophagous on Populus tremula, but more eurytopic, in higher frequency and usually high individual numbers, often also found on solitary trees and shrubs. Widespread in Germany, extending up to at least 1000 m a.s.l. in the Mittelgebirge and the Bavarian Alps, found at 1480 m a.s.l. in Switzerland. Achtziger (1991), Fischer (1972), Niedringhaus (1991, 1997), Reimer (1992), Remane & Reimer (1989), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1951a), HN; see also Günthart (1987b)
Acericerus heydenii (Kirschbaum, 1868) E VII – E VI; adult, 1 gen. Usually on Acer pseudoplatanus in forests, river floodplains and urban settlements, in lower numbers also on A. platanoides, A. campestre and A. monspessulanum. Often flies into houses during autumn and winter nights. Widespread in middle and southern parts of Germany and – at least on A. pseudoplatanus – in intermediate frequency, but low individual numbers. The northernmost localities are near Cologne, Göttingen, Kiel, Stendal, Berlin and Fürstenwalde. Found up to at least 1000 m a.s.l. in the Bavarian Forest, up to at least 1250 m a.s.l. in the Bavarian and Allgäu Alps. Fischer (1972), Frommer (pers. comm.), Reimer (1992), Remane (pers. comm.), Remane & Wachmann (1993), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1939a), Weis & Schönitzer (2001), Nickel (1994), HN
Acericerus ribauti Nickel & Remane, 2002 Idiocerus rotundifrons Kirschbaum, 1868 sensu Ribaut (1952) (see Nickel & Remane 2002)
Adults I – XII, mainly B VIII – V; adult, 1 gen. Preferentially on Acer campestre, in lower numbers also on A. pseudoplatanus and A. platanoides, usually along sun-exposed forest margins, in parks and urban settlements, flying into houses during autumn and winter nights. Widespread in middle and southern parts of Germany and in intermediate to high frequency on A. campestre, but at the edge of the range; northernmost localities are near Cologne, Göttingen, Halle and Berlin, mainly below 600 m a.s.l., but recorded at 800 m a.s.l. on the Swabian Jura. Fischer (1972), Frommer (1996), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1988), Schönitzer & Oesterling (1998b), Weis & Schönitzer (2001), Nickel (1994), HN
Acericerus vittifrons (Kirschbaum, 1868) Idiocerus rotundifrons Kirschbaum, 1868 (see Nickel & Remane 2002)
Adults I – XII, mainly B VIII – V; adult, 1 gen.
Iassinae Amyot & Serville, 1843
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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, another site near Neubrandenburg may be rather isolated, although there is a very recent record from southern Sweden (Skåne). Büchs (1988), Frommer (1996), Kuntze (1937), Reimer (1992), Schiemenz (1988), Schwoerbel (1957), Wagner (1951a), Weis & Schönitzer (2001), Nickel (1994), HN; see also Gillerfors (2002)
Balcanocerus larvatus (Herrich-Schäffer, 1835) E VI – M X; egg, 1 gen. On Prunus spinosa in sun-exposed, damp to dry sites, mainly pastures and meadows with scattered shrubs, embankments, forest margins and hedges. In Germany widespread between the Danube and the northern edge of the Mittelgebirge, extending to at least 750 m a.s.l. and occurring in high frequency at least in warmer regions, although seldom recorded. Northernmost localities are near Bonn, Brilon, Göttingen, Wolfenbüttel, Halle, Leipzig and Dresden; there is also an old record from near Neubrandenburg. Not reported from Denmark and Scandinavia. Achtziger (1991, 1994), Heller (1987a), Kuntze (1937), Post-Plangg & Hoffmann (1982), Remane & Fröhlich (1994b), Rombach (1999a), Schiemenz (1988), Schwoerbel (1957), Trümbach (1959), Wagner (1939a, 1951a), Walter (1998), Nickel (1994), HN; see also Ossiannilsson (1981)
Balcanocerus pruni (Ribaut, 1952) M VIII – B VI; adult, 1 gen. Like the preceding species on Prunus spinosa, but more thermophilous, in Germany largely confined to viticultural regions of southwestern parts. Recorded from the Ahr valley near Altenahr, the middle Rhine near Cologne and Bacharach, the Nahe valley near Schloßböckelheim and Mainfranken (Karlstadt and Hammelburg), between 50 and 350 m a.s.l. Otherwise only reported from France, Italy, the former Yugoslawia, Bulgaria, Greece and southern Russia (Dagestan). Frommer (pers. comm.), Remane & Fröhlich (1994b), Nickel & Remane (1996), HN; see also Dlabola (1961a), Drosopoulos et al. (1986), Nast (1987)
4.2.4.7 Iassinae Amyot & Serville, 1843 Batracomorphus allionii (Turton, 1802) E VI – B IX; egg, 1 gen. In forest glades, clearings, heaths, etc., usually in oligotrophic, more or less acidic, damp to moderately dry sites. The host plant is Cytisus scoparius, occasionally also Genista tinctoria and perhaps additional species of woody Fabaceae. The distribution in Germany is imperfectly known; strongholds are apparently in sand and sandstone regions in
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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), Wagner (1935, 1939a), Nickel (1999b), HN
Batracomorphus irroratus Lewis, 1834 E VI – M IX; egg, 1 gen. In xerothermic grassland and along sunny margins of pine and oak forests, usually on limestone and other basic substrates. The main host plant is Helianthemum nummularium, although other species of Helianthemum have rarely been swept. In Germany occurring rather localized and only in middle parts, mainly in limestone regions (Saale-Unstrut region, southern Niedersachsen, Thuringian Basin, Eifel Mountains, Rhine-Main region, Mainfranken, Tauber valley, as well as the Franconian and Swabian Jura, usually found below 500 m a.s.l., but up to at least 800 m a.s.l. on the Swabian Jura. The northern edge of the range roughly follows a line from Bonn to Marburg, Göttingen, Wernigerode, Eisleben and Leipzig. Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Remane (pers. comm.), Rombach (1999a), Schiemenz (1988), SMNS, Wagner (1939a, 1951a), Nickel (1994), HN
Iassus lanio (Linnaeus, 1761) E VI – B XI; egg, 1 gen. Eurytopic on oaks (Quercus robur, Qu. petraea), in the interior of forests as well as along their margins and on solitary trees, also in urban settlements. Widespread in the lowlands of Germany, in high frequency on both oak species, also on the East Frisian Islands, extending to at least 600 m a.s.l. in the foothills of the Alps. Achtziger (1991), Fischer (1972), Gotsmy & Schopf (1992), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz (1988), Wagner (1935, 1939a), HN and others
Iassus scutellaris (Fieber, 1868) E VI – B X; egg, 1 gen. Monophagous on elms, usually on Ulmus minor and U. x hollandica, perhaps also on U. laevis, but not recorded from U. glabra. Usually along alleys and hedges and on solitary trees, mainly in river floodplains, parks and along waysides, presumably also in closed forests, where difficult to sample with the sweep-net. Widespread in lowland middle parts of Germany (to at least 500 m a.s.l.), in high frequency but low individual numbers at least on U. minor and U. x hollandica. Furthermore, there are single records from the area to the south of the Danube and the north German plain. Northernmost localities are near Cologne, Bielefeld, Hanover, Hamburg, Parchim and Rostock. Apparently absent from Denmark and Scandinavia. Fischer (1972), Feldtmann (1937), Frommer (1996), Remane (1987), SMNS, Wagner (1935), ZIMH, Nickel (1994), HN; see also Ossiannilsson (1981)
Penthimiinae Kirschbaum, 1868
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4.2.4.8 Penthimiinae Kirschbaum, 1868 Penthimia nigra (Goeze, 1778) B V – M VIII; nymph, 1 gen. On shrubs and low-growing trees in dry to damp sites in warm situations (notably abandoned vineyards, dry grassland, ruderal sites, forest margins, coppice-with-standards, etc.). Food plants are various deciduous shrubs and trees (adults found on Quercus, Populus and others). Occurs in Germany only locally and at the northern edge of the range (usually below 400 m a.s.l.), with strongholds in viticultural regions, notably along the upper and middle Rhine and its tributaries (Ahr, Moselle, Nahe, Main), the Elbe near Dresden and Meißen, as well as in the Saale-Unstrut region between Naumburg, Halle and Leipzig; further localities, some of which appear to be rather isolated, are near Augsburg, Bonn, Berlin and in the Oberlausitz. Fischer (1972), Maixner (pers. comm.), Post-Plangg & Hoffmann (1982), Remane (1987, and pers. comm.), Remane & Wachmann (1993), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1939a, 1951a), HN
4.2.4.9 Dorycephalinae Oman, 1943 Eupelix cuspidata (Fabricius, 1775) I – XII, probably semivoltine, hibernation as nymph and adult. In sunny to moderately shady, dry to damp, occasionally also moist or even temporarily wet sites (various types of dry grassland, dunes, low-productivity meadows and pastures, ruderal sites, waysides, sunny forest margins, moderately saline sites, etc.), mainly on Festuca ovina, perhaps also on F. rubra and additional grasses. Widespread in Germany and fairly common in favourable sites, recorded up to 1300 m a.s.l. in the Black Forest, up to 1500 m a.s.l. in the Engadine (Switzerland). Bornholdt (1996), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1939a, 1951a), Nickel & Achtziger (1999), HN; see also Günthart (1987a)
4.2.4.10 Aphrodinae Haupt, 1927 For a long time, there have been diverging opinions on systematics of the genus Aphrodes Curt. (s.str.). Until recently, most authors distinguished only two species in central Europe (Emmrich 1980; Nast 1976b). The interpretation followed here is adopted from Tishechkin (1998), who essentially clarified biosystematic, although not nomenclatural problems. Accordingly, the Aphrodes bicincta group includes 4 distinct species with differences in bioacoustics, morphology and life history (see below). Hence, most published records from central Europe are in need of revision. Furthermore, there is a number of older names, which might have priority for some taxa, but this would require investigation of a vast material. The grammatical gender of Aphrodes is female (see Metcalf 1963).
Aphrodes aestuarina (Edwards, 1908) Perhaps only a subspecies or ecological form of A. makarovi Zachv.
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In England VII – IX; egg(?), 1 gen. (Kirby 1992). At present this taxon is only known from coastal regions of the North and Baltic Sea (England, Poland, Niedersachsen). Recorded in Germany only from the East Frisian Islands and the vicinity of Cuxhaven, where it is probably restricted to coastal salt marshes and moderately saline meadows and pastures. The host plants are unknown. Niedringhaus (1991, and pers. comm.), Wagner (1937c); see also Kirby (1992); for identification see Tishechkin (1998)
Aphrodes bicincta (Schrank, 1776) sensu Tishechkin (1998) Aphrodes makarovi Zachv. sensu Ossiannilsson (1981)
B VI – X, single specimens until I; egg, 1(?) gen. Due to very recent taxonomic changes this species’ distributional status in central Europe is rather unclear. Apparently mainly in sun-exposed, damp to dry, basic to acidic sites, notably calcareous hillsides, inland dunes, heaths, low-productivity meadows, dry waysides and abandoned fields. Probably widespread in Germany, but with strongholds in the lowlands, most verified records are from below 500 m a.s.l. Host plants are species of Fabaceae and perhaps other plant families. Was identified as a vector of Clover phyllody (CP), Clover dwarf (CD) and perhaps other plant diseases in various parts of Europe, but the species identity of these populations is uncertain (see above). The latter is also true for populations found in inland salt marshes and on dry hillsides at higher altitudes of the Alps and the Mittelgebirge. Emmrich (1980), HN; see also Brˇcák (1979); for identification see Tishechkin (1998)
Aphrodes diminuta Ribaut, 1952 Aphrodes centrorossica Zachvatkin, 1953; A. bicincta (Schrank, 1776) sensu Ossiannilsson (1981)
E VI – E X; egg, 1 gen. In oligotrophic, wet to temporarily dry sites, mainly in fens dominated by Carex, in lowinput meadows and pastures. Is reported to live on species of Fabaceae in Russia. The distribution in Germany is imperfectly known, but the species is probably widespread, although uncommon, with strongholds in the submontane and montane belt in middle and southern parts, but also in the north German plain. It is frequently found at 1200 m a.s.l. in the Mittelgebirge, up to at least 1500 m a.s.l. in the Bavarian Alps. Emmrich (1980), Remane (pers. comm.), Remane & Wachmann (1993), HN; for identification see Tishechkin (1998)
Aphrodes makarovi Zachvatkin, 1948 Mainly E VI – E X, single individuals occasionally until winter; egg, 1 gen. Eurytopic in moderately wet to damp, sunny to shady sites, often in disturbed or eutrophic patches, notably fertilized meadows and pastures, tall herb and grass stands, abandoned fields, ruderal sites, shores of running and standing water, ditches, forest roads, alder fen woods, etc., also slightly saline sites inland and near the coast. Host
Aphrodinae Haupt, 1927
105
plants are Taraxacum, Cirsium, Urtica dioica, Rumex, and probably additional dicotyledonous herbs. Widespread and common in the lowlands of Germany, but usually below c. 800 m a.s.l., although imperfectly documented. Remane (pers. comm.), HN; for identification see Tishechkin (1998)
Planaphrodes bifasciata (Linnaeus, 1758) B VI – E IX; egg, 1 gen. In sunny to shady, moderately wet to damp sites, mainly meadows and pastures of the submontane belt dominated by Nardus or Trisetum and dicotyledonous herbs, also in lowland areas in moist deciduous forests with a luxuriant herb layer (e.g. floodplain and fen woods), among stands of tall herbs as well as along margins of bogs. No host plants are known from central Europe, but in field choice experiments conducted in Britain, adults and nymphs have been found in numbers on several grasses, notably on Helictotrichon pubescens, Brachypodium pinnatum, Briza media, Bromus erectus, Festuca ovina, F. rubra, Trisetum flavescens, as well as on the sedge Carex flacca. The distribution in Germany is imperfectly documented due to secretive habits. Apparently, most localities are in the Alps and the Mittelgebirge (mainly between 400 and 1200 m a.s.l., up to 1500 m a.s.l. in Switzerland), e.g. Solling, Harz and Rhön Mountains, Frankenwald, Thuringian Forest, Erzgebirge and Vogelsberg, but there are also records from valley situations and the north German plain. Bornholdt (1996), Fischer (1972), Nikusch (1976), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1937c), Nickel & Achtziger (1999), HN; see also Cook (1996), Günthart (1987a)
Planaphrodes nigrita (Kirschbaum, 1868) M VI – E X; egg, 1 gen. Like the preceding species, mainly in mountainous areas, often syntopic and difficult to distinguish ecologically. Less frequently found, in the lowlands apparently preferring forests in damp sites, but rather wet habitats at higher altitudes (e.g. spring mires, boggy spruce forests, occasionally grassland). The host plants are unknown, a few individuals were taken from Calamagrostis canescens and Juncus articulatus, but the species occurs in many sites where these plants are absent. Widespread in Germany, but imperfectly documented, from lowland areas up to at least the upper montane belt, found up to 1500 m a.s.l. in the Bavarian Alps, at 2250 m a.s.l. in North Tyrol (Austria). Emmrich (1969), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1988), Schönitzer & Oesterling (1998b), Wagner (1937c), Nickel & Achtziger (1999), Nickel & Remane (1996), Nickel (1997), HN; see also Christandl-Peskoller & Janetschek (1976)
Planaphrodes trifasciata (Geoffroy, 1785) sensu Ribaut (1952) The name given in the original description is not valid, since it is a primary homonym of Cicada trifasciata De Geer, 1773, which is in turn a junior synonym of P. bifasciata (Linnaeus, 1758) (see Metcalf 1963). Therefore, Hamilton (1975) has, apparently without studying type material, pro-
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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 strongholds in bog and heath areas of the north German plain, the Mittelgebirge and the Alps, as well as in warmer regions of southern and middle parts. Highest localities are at 1000 m a.s.l. in the Thuringian Forest and at 1300 m a.s.l. in the Chiemgau Alps. Achtziger (1991), Fischer (1972), Remane (1987), Schiemenz (1969, 1975, 1988), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN
Anoscopus albifrons (Linnaeus, 1758) M VI – E X, single individuals until winter; egg, 1 gen. In sunny to moderately shady, moderately dry to moist sites, mainly heaths, open forests, low-productivity meadows and pastures. Host plants are various grasses (Deschampsia flexuosa, Holcus mollis, Brachypodium pinnatum and others). Widespread in Germany and common in favourable habitats, also in higher parts of the Mittelgebirge, up to at least 1500 m a.s.l. in the Bavarian Alps. Achtziger (1991), Fischer (1972), Haas (1975), Niedringhaus (1991, 1997), Reimer (1992), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN
Anoscopus limicola (Edwards, 1908) M VII – E IX; egg, 1 gen. This species is apparently an endemic of western European coasts, living among grasses in salt marshes. The host plant is probably Puccinellia maritima. In Germany it is known only from the islands of Sylt and Amrum, all East Frisian Islands, where locally dominant, as well as from a few mainland sites (near Varel, Bremerhaven and Cuxhaven). Further verified records exist only from Ireland, England, southern Sweden, the Netherlands and the Atlantic and Mediterranean coasts of France. Hildebrandt (1995), Niedringhaus (1991), Niedringhaus & Olthoff (1993), Wagner (1937c), ZIMH; see also della Giustina & Remane (2001), Gravestein (1965), Kirby (1992), Nast (1987), Ossiannilsson (1981)
Anoscopus albiger (Germar, 1821) E VI – M XI; egg, 1 gen. Usually in moderately saline sites inland and near the coast, also in ruderal habitats (preferentially coal mining areas) and peaty sedge swamps. Host plants are not known, but are likely to include grasses. Rather scattered in Germany, reaching the northern edge of the range; mainly along the coast of the North Sea and in inland salt marsh and
Aphrodinae Haupt, 1927
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coal mining areas of eastern parts, notably Saxony-Anhalt, Thuringia and Saxony, single records are from near Oldenburg, Wolfenbüttel, Göttingen, Cologne, Gießen, Frankfurt am Main, as well as the French part of the upper Rhine plain, usually below 250 m a.s.l. To the north of Germany it is known only from the Baltic Sea islands of Bornholm and Gotland. Fröhlich (1996a), Frommer (1996), Funke & Witsack (1998), Niedringhaus & Olthoff (1993), Schiemenz (1988), Wagner (1937c, 1939a), Nickel (1997), HN; see also Gillerfors (2002), Ribaut (1952)
Anoscopus alpinus (W. Wagner, 1955) Described from northeastern parts of the Alps, but is perhaps conspecific with A. assimilis (Signoret, 1879), which is reported mainly from southwestern parts of Europe. The description of A. duffieldi (Le Quesne, 1964), known only from a single coastal site in England (see also Remane & Fröhlich 1994b) is based merely upon slight differences of the aedeagus tip, body colouration and vertex length. However, in most Anoscopus species, these characters are subject to considerable intraspecific variation. Hence, morphological and zoogeographic evidence suggests that at least the two latter taxa are conspecific. In France A. assimilis (Sign.) is reported from coastal lowlands of western and northwestern parts (Ribaut 1952; della Giustina 1989).
M VII – E IX; egg, 1 gen. In heaths and bogs dominated by dwarf shrubs, extending from the submontane to the alpine belt. In Germany largely confined to the Bavarian and Allgäu Alps (between 1100 and 2100 m a.s.l., but up to 2970 m a.s.l. in North Tyrol, Austria) and to the higher Mittelgebirge, notably the Erzgebirge, Thuringian Forest and southern Black Forest (between 650 and 1450 m a.s.l.) The host plants are presumably grasses associated with Vaccinium myrtillus, Calluna vulgaris and other dwarf shrubs. Remane & Fröhlich (1994b), Schiemenz (1971a, 1975, 1988), Walter (pers. comm.), Nickel & Voith (unpublished data), Nickel (2002); see also Christandl-Peskoller & Janetschek (1976), Wagner (1955)
Anoscopus flavostriatus (Donovan, 1799) B VII – B XI, according to Schiemenz (1988) single individuals until IV; egg, 1 gen. In grass stands of various types, usually in rather cool, wet to damp sites (mainly lowinput meadows and pastures, forest glades, fens, abandoned fields and open forests). The host plants are grasses (Deschampsia flexuosa, Dactylis glomerata, Elymus repens and others). The species is widespread in Germany and fairly common in most regions (up to at least 1000 m a.s.l.), more scattered or locally absent only in warm and dry areas (notably the rain shadow area to the east of the Harz Mountains, Mainfranken, upper Rhine plain). Achtziger (1991), Fischer (1972), Niedringhaus (1991), Reimer (1992), Remane (1958), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN
Anoscopus histrionicus (Fabricius, 1794) Despite distinctive colouration of ††, this species has been misidentified or misinterpreted. Thus, a number of records compiled by Schiemenz (1988) are in need of revision and refer to other species, including most, if not all Thuringian records.
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Review of species
M VI – M IX; egg, 1 gen. Is reported to live in moist to moderately dry, usually sunny and oligotrophic sites, e.g. heaths, low-input meadows and open forests. Host plants are probably grasses. In Germany mainly in northern and eastern parts and perhaps at the edge of the range. The westernmost localities are on the East Frisian Islands from Borkum to Wangerooge, near Hamburg, Hanover, Dessau, Leipzig, Fürth, Regensburg and Munich, up to 500 m a.s.l. The majority of records dates back at least 40 years, more recent ones are only known from the East Frisian Islands. In France the species is apparently confined to the Alps and the Pyrenees, but it is reported to occur in most European countries including the Netherlands, Belgium and the British Islands. Herrich-Schäffer (1834), Niedringhaus (1991), Schönitzer & Oesterling (1998b), Wagner (1935); see also della Giustina (pers. comm.), Holzinger (pers. comm.), MNHN (Bourgoin pers. comm.), Nast (1987)
Anoscopus serratulae (Fabricius, 1775) B VII – M X; egg, 1 gen. In grass stands in moderately dry to moist, occasionally also wet, usually sunny sites, mainly more or less strongly fertilized meadows and pastures, ruderal sites, lawns in parks and gardens, also moderately saline sites. Host plants are grasses (Elymus repens, Dactylis glomerata, Holcus spp.). The species is widespread in the lowlands of Germany and fairly common, the highest localities are at c. 700 m a.s.l. only, but it is recorded from 900 m a.s.l. in North Tyrol (Austria). Achtziger (1991), Niedringhaus (1991), Nikusch (1976), Schiemenz (1988), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN; see also Günthart & Thaler (1981)
Stroggylocephalus agrestis (Fallén, 1806) M VII – E X, at least occasionally until VI; egg(?), 1 gen. In sedge stands in various wet sites (mainly fens, intermediate bogs, wet meadows, fen woods, also slightly saline sites); host plants are usually tall species of Carex (C. acutiformis, C. acuta, C. nigra and others), perhaps also Bolboschoenus maritimus. Widespread in the lowlands of Germany and fairly common in favourable sites; the highest localities are at c. 800 m a.s.l. Fischer (1972), Hoffmann (1980), Kuntze (1937), Reimer (1992), Schiemenz (1976, 1988), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN
Stroggylocephalus livens (Zetterstedt, 1840) E VII – E VI; adult (and nymph?), 1 (perhaps only half) gen. This is a tyrphophilous species usually living in intermediate bogs and fens, but it is also reported from raised bogs (where probably restricted to margins) and birch fens. Presumably it lives on Carex, perhaps on Eriophorum. Uncommon in Germany and mainly in remnant bog areas of the north German plain, small populations are also found in the Bavarian and Swabian foothills of the Alps and the Mittelgebirge (Harz Mountains, Thu-
Cicadellinae Latreille, 1825
109
ringian Forest, Erzgebirge, Rothaargebirge, Rhön Mountains, Fichtelgebirge), as well as in the upper Rhine plain, up to at least 900 m a.s.l. Achtziger (pers. comm.), Bittner & Remane (1977), Hoffmann (1980), Kuntze (1937), Schönitzer & Oesterling (1998b), Remane (1958), Schiemenz (1971a, 1973, 1975, 1976, 1988), Wagner (1935), Walter (pers. comm.), Nickel (2002), HN
4.2.4.11 Cicadellinae Latreille, 1825 Evacanthus acuminatus (Fabricius, 1794) B VI – B IX; egg, 1 gen. In moderately shady to shady, moist to moderately dry, more or less eutrophic sites, usually with a luxuriant herb layer, mainly in deciduous and mixed forests of various types and along woody margins (in dry grassland, abandoned vineyards, etc.); absent from mown meadows. Host plants are various dicotyledonous herbs; adults were taken from Glechoma, Lamium, Stachys, Valeriana, Verbascum and others. Widespread in Germany, particularly in middle and southern parts, although usually collected in low individual numbers only, more scattered in the north German plain. Frequently found up to 1200 m a.s.l. in the Bavarian and Allgäu Alps, up to at least 1500 m a.s.l. in Carinthia (Austria) and the Engadine (Switzerland). Büchs (1988), Fischer (1972), Niedringhaus (1991), Post-Plangg & Hoffmann (1982), Reimer (1992), Schiemenz (1988), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN and others; see also Günthart (1987a), Prohaska (1923)
Evacanthus interruptus (Linnaeus, 1758) M VI – M X; egg, 1 gen. In sunny to shady, wet to damp, usually rather eutrophic sites, mainly in low-input meadows, fens, along various herbaceous and woody margins (shores of running or standing water, forest roads and edges, ditches, etc.), in forests and subalpine tall herb stands. Adults on Arctium, Epilobium angustifolium, Cirsium, Eupatorium, Lamium, Petasites, Senecio, Urtica and many others, occasionally also Rubus idaeus. Widespread and common in Germany, particularly in rather cool situations (but apparently absent from the East Frisian Islands). Large populations have been found up to at least 1850 m a.s.l. in the Bavarian and Allgäu Alps; the highest localities in North Tyrol (Austria) and the Engadine (Switzerland) are at 2200 m a.s.l. Achtziger (1991), Fischer (1972), Niedringhaus (1991), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1988), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN and others; see also Günthart (1987a)
Errhomenus brachypterus Fieber, 1866 Has been subject to taxonomic confusion, which was finally cleared by Lauterer (1983). Accordingly, there is a considerable intraspecific variability of colouration, head shape and body sculpturation, resulting in the description of Errhomenellus flavopunctatus Melichar, 1914, which is, however, a younger synonym.
110
Review of species
I – XII, probably semivoltine, hibernation as nymph and adult. In the litter layer of deciduous and coniferous forests (mainly under Fagus, Quercus, Picea, Abies and Pinus) in wet to damp, basic as well as acidic sites, also found up to at least 1850 m a.s.l. in subalpine scrub in the Bavarian and Allgäu Alps (reported to live under Alnus alnobetula, Pinus mugo and Rhododendron, up to 2000 m a.s.l. in Austria). Usually recorded in pitfall traps and sieved litter, rarely in sweep-net samples and under stones. The host plants are unknown. Apparently, there is not a single plant species or even a plant family common to all sites. Moreover, there is often a complete lack of any herbaceous undergrowth. Thus, the species may live polyphagously on fine roots of trees and shrubs, which are within easy reach of its long rostrum. The distribution in Germany is imperfectly documented, but it seems to be widespread and locally even common in southern and middle parts (mainly above 300 m a.s.l.). According to present data, the border of the range runs along des northern edge of the Mittelgebirge (northernmost localities are on a line from Aachen to Dortmund, Minden, Goslar, Quedlinburg, Leipzig and Dresden); a singe record from near Hamburg dates from 1895. Large numbers have been sampled in acidic and boggy spruce forests in higher parts of the Harz Mountains, Thuringian Forest and Erzgebirge, also in limestone beech forests of southern Niedersachsen and northern Hessen and in mixed forests on Cretaceous chalk and dolomite in the Veldensteiner Forst near Nuremberg. More or less scattered records are known from other regions (Solling, Eifel Mountains, Franconian Jura, Breisgau, Swabian and Bavarian Allgäu, also in the Vosges (eastern France) and the southern Dutch province of Limburg. Achtziger (1991), Coll. Bauchhenss, Drees (2000), Haupt (1935), Schiemenz (1971a, 1975, 1988), Coll. Sprick, SMNS, Wagner (1938c), Nickel (1994), HN; see also Cobben & Rozeboom (1978), Hölzel (1965), Wagner & Franz (1961)
Cicadella lasiocarpae Ossiannilsson, 1981 So far E VI – B IX; egg, 1 gen. Was described not long ago from Sweden, but has been found in large parts of the Palearctic since (from eastern Siberia and Korea to Ireland, Wales and the northern half of Germany). At present there are about 20 known localities in Germany from near Lüchow, Haldensleben, Cologne, Dresden, Bautzen, Marburg, and Fulda, up to 300 m a.s.l., usually in wet to peaty, mesotrophic sites (peaty meadows and pastures, fens and intermediate bogs), often with dominating sedges, particularly Carex nigra. In most cases it occurs at high densities, syntopically with C. viridis (L.). Frommer (pers. comm.), Remane (pers. comm.), Remane & Fröhlich (1991), Nickel & Achtziger (1999), Nickel (1997), HN; see also Dmitriev (1998), Le Quesne (1987), Ossiannilsson (1981), G¸ebicki & Szwedo (1998), Tishechkin (2000), Walter (pers. comm.)
Cicadella viridis (Linnaeus, 1758) Mainly B VII – M X; 1 gen.; in warmer regions E V – B XI, 2 gen., according to Witsack (1985), hibernation takes place in the embryonic stage, in secondary parapause terminated by cold.
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111
In various sunny to moderately shaded, temporarily flooded to moist, occasionally also rather dry sites. Polyphagous on rushes, sedges and grasses, probably also on dicotyledons, although most commonly on Juncus. Oviposition into saplings and low branches of trees has been reported to cause damage in forest and fruit-tree plantations. Highest abundances usually occur in fens, intermediate bogs, wet meadows and pastures, but the species is also found along bog margins, in open forests, clearings, ruderal sites, mining areas, along forest roads and ditches, occasionally even in dry grassland (including nymphs). It is widespread in Germany and common in most regions, in the Bavarian Alps frequently found up to at least 1200 m a.s.l., but also in the subalpine belt in Styria (Austria). Fischer (1972), Müller (1956, 1972), Niedringhaus (1991), Reimer (1992), Schiemenz (1988), Schmutterer (1953), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN and others; see also Wagner & Franz (1961)
Graphocephala fennahi Young, 1977 M VII – M X, in southwestern Germany B VII – XI; egg, 1 gen. This is a Nearctic species which was introduced probably with its host plants and which occurs in Europe at least since the 1930s, initially in England, since the 1970s also in Switzerland, France, the Netherlands and Germany (at that time generally reported as G. coccinea Forster, 1771). Discovered in Germany in 1978 in Mönchengladbach, afterwards found in Oldenburg in 1982, in Düsseldorf, Frankfurt am Main and Stuttgart in 1983, in Bremen and Hamburg in 1984, in Berlin in 1988. At present, it is widespread in much of the western half below 550 m a.s.l., and particularly common in the basins of the Rhine and Ems. In recent years there is a number of records from the eastern half, indicating a further eastward range expansion. Hamburg, Dannenberg, Hanover, Gardelegen, Berlin, Göttingen, Freiberg, Dresden, Bayreuth, Schweinfurt, Munich, Stuttgart and Freiburg are localities near the spreading front, although it is also reported from Vienna (Austria). The species lives in parks, cemeteries and gardens (in England also in open forests) on ornamental species of Rhododendron, which is necessary for nymphal development, although some individuals may be swept from other woody plants such as Acer, Platanus, Tilia and others. In England and France it is often mentioned in connection with Pycnostysanus azaleae (Peck.), a phytopathogenic fungus causing bud blast on Rhododendron. However, it is uncertain whether pathogen transmission is conducted or facilitated by the leafhopper, since detailed studies are lacking. Achtziger (pers. comm.), Bußmann (1992), Feldmann & Bußmann (1993), Hoffmann (1990), Körner (pers. comm.), Niedringhaus & Olthoff (1986, 1993), Olthoff (1986), Remane (pers. comm.), Remane & Wachmann (1993), Sergel (1987), SMNS, Walter (pers. comm.), Weis (pers. comm.), Nickel & Remane (1996), Nickel (1994), HN; see also Arzone et al. (1987), della Giustina (1989), Kaiser-Mauer (1977), Morcos (1953)
4.2.4.12 Typhlocybinae Kirschbaum, 1868 Alebra neglecta W. Wagner, 1940 E VI – B IX; egg, 1 gen. Oligophagous on Carpinus betulus, Prunus padus and Crataegus, occasionally also on Prunus avium; most individuals are swept in open stands of trees and shrubs and along forest
112
Review of species
margins. Widespread in Germany, but not common and imperfectly documented. The edge of the range runs through north Germany; northernmost localities are near Bremen, Hamburg, Plön and Neubrandenburg, extending to at least 700 m a.s.l. in the Allgäu. Frommer (1996), Günthart (1988), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1990), SMNS, Wagner (1940b), Nickel & Remane (1996), HN
Alebra coryli Le Quesne, 1977 M VI – B IX; egg, 1 gen. At present only known from the western half of Europe (British Isles, Netherlands, France, Switzerland, Poland, Czech Republic and Italy) and Canada (where presumably introduced), but probably more widespread and formerly misidentified. Lives on Corylus avellana in forests, along their margins, as well as on solitary shrubs, also on C. maxima and C. colurna in urban settlements. Widespread in Germany, apparently common and in rather high frequency in most better studied areas, e.g. large parts of Bavaria, middle Hessen and southern Niedersachsen; also recorded along the upper and middle Rhine, the Wendland of Niedersachsen, the Oberlausitz and the Oderbruch, up to at least 700 m a.s.l., found at 860 m a.s.l. in Switzerland. Frommer (pers. comm.), Remane & Fröhlich (1994b), Schönitzer & Oesterling (1998b), Nickel (1994), HN; see also della Giustina (1989), Dworakowska (1993), Gillham (1991) (see this paper also for identification), D’Urso (1995), Günthart (1987b), Lauterer & Novotný (1991)
Alebra wahlbergi (Boheman, 1845) E V – M IX; egg, probably 1 gen. (see also Demichelis & Bosco 1995). Polyphagous and eurytopic on various deciduous trees, the majority of individuals found on Acer campestre, Tilia cordata, Ulmus minor and Carpinus betulus, in lower numbers also on Acer platanoides, A. pseudoplatanus, Ulmus glabra, Sorbus intermedia, S. aria and Betula pendula; in Britain also on Aesculus hippocastanum, Prunus avium, Alnus glutinosa and Castanea sativa. Widespread in Germany and usually among the commonest arboricolous leafhopper species of lower altitudes, found up to at least 800 m a.s.l. in the Mittelgebirge and the foothills of the Bavarian Alps, at 1300 m a.s.l. in the Valais (Switzerland). Achtziger (1991), Frommer (1996), Niedringhaus (1997), Olthoff (1986), Remane (1987), Schiemenz (1990), Wagner (1935, 1939a), HN, and others; see also Cerutti (1939a), Claridge & Wilson (1981)
Alebra albostriella (Fallén, 1826) This taxon and the following have long been treated as conspecific, but are considered to be distinct species since Gillham (1991). Accordingly, there are differences in colouration and in the morphology of sternal apodemes of ††, as well as in host preferences. The latter, however appear to be variable in different parts of Europe: A. albostriella (Fall.) lives monophagously on Quercus robur in Wales, but on various Fagaceae species in Piemont (see below). The taxonomic situation (also within remaining Alebra species) is further complicated by the occurrence of various colour morphs.
E VI – M X; egg, 1 gen., in northern Italy according to Demichelis & Bosco (1995) occasionally 2.
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113
Usually on Quercus robur, at least adults occasionally on Qu. petraea, Alnus, Betula, and additional deciduous species, in closed forests as well as in open stands and on solitary trees. From Italy Quercus petraea, Qu. cerris and Castanea sativa have been reported as additional host plants. Apparently widespread in the lowlands of Germany, although published records partially refer to the following taxon. Highest localities in the Mittelgebirge and the foothills of the Alps are at 750 m a.s.l. Achtziger (1991), Frommer (1996), Niedringhaus (1997), Remane (1987), Schiemenz (1990), Wagner (1935, 1939a), HN, and others; see also Demichelis & Bosco (1995); for identification see Gillham (1991)
Alebra viridis (Rey, 1894) M VII – M X; egg, 1 gen., 2 according to Demichelis & Bosco (1995) in northern Italy. Usually on Quercus petraea in forests and along their margins, at least single individuals also on Qu. robur; in Wales breeding has also been recorded on Qu. cerris, in Italy additionally on Castanea sativa. Apparently widespread in the lowlands of Germany (extending northward at least to Cologne, Lüneburg and Berlin), but less common than the preceding taxon and – particularly in the eastern half – only poorly documented. The highest localities are at 550 m a.s.l. in the Mittelgebirge, at 1300 m a.s.l. in the Valais (Switzerland). Frommer (1996), Nickel & Remane (1996), Nickel (1994), HN; see also Cerutti (1939a), Demichelis & Bosco (1995), Gillham (1991); for identification see Gillham (1991)
Erythria aureola (Fallén, 1806) M V – E X; egg, 2 gen., perhaps 1 at higher altitudes, perhaps 3 in warmer regions. On dwarf shrubs in sunny, sparsely vegetated, moderately dry to dry sites; in the north German plain mainly on heathland on Calluna vulgaris, in middle and southern parts on various substrates and usually on Thymus. Widespread in Germany, scattered and locally absent in the Mittelgebirge, but often among dominant species in favourable places; highest localities are at 1000 m a.s.l. in the Mittelgebirge, at 1800 m a.s.l. in the Bavarian and Allgäu Alps. Is reported to live in on Globularia cordifolia and Teucrium montanum in the alpine belt of Switzerland (up to 2280 m a.s.l.) and eastern parts of Austria. Fischer (1972), Reimer (1992), Remane (1987), Schiemenz (1969, 1990), Trümbach (1959), Wagner (1935, 1939a, 1951a), HN; see also Günthart (1987a), Wagner & Franz (1961)
Erythria manderstjernii (Kirschbaum, 1868) Adults I – XII, mainly B VIII – B VI; adult, 1 gen. On dicotyledonous herbs and dwarf shrubs in sunny to shady, usually damp to moderately wet sites of the Mittelgebirge and the Alps; mainly in open coniferous and mixed forests, subalpine scrub and alpine grassland, also near streamlets. Adults are usually found on various dicotyledons, e.g. Ranunculus (at least R. montanus, R. repens), Thymus, Senecio, Eupatorium, Vaccinium, in southwestern parts of the Alps also Chrysanthemum, Crepis, Leontodon, Trifolium, Prunella, Plantago and Chaerophyllum. Widespread and com-
114
Review of species
mon in the Bavarian and Allgäu Alps, between 800 and 2000 m a.s.l., recorded at 2400 m a.s.l. in North Tyrol (Austria) and the Valais (Switzerland). Also in higher parts of the Mittelgebirge (usually above c. 600 m a.s.l., but extending down to c. 450 m a.s.l. in cool and shady situations), known from the Black Forest (syntype from near Freudenstadt), the Swabian Jura, Rhön Mountains, Fichtelgebirge, Vogelsberg, Thuringian Forest, Erzgebirge, Eifel Mountains, Rothaargebirge and Harz Mountains. Fischer (1972), Kirschbaum (1868), Nikusch (1976), Rabeler (1952), Reimer (1992), Remane (pers. comm.), Schiemenz (1990), HN; see also Cerutti (1939a), Christandl-Peskoller & Janetschek (1976), Dobler (1985), Leising (1977), Vidano (1959a)
Liguropia juniperi (Lethierry, 1876) So far B VIII – IX; adult, 1(?) gen. This species is known to live on scaly species of Cupressaceae (Cupressus sempervirens, Juniperus phoenicea, Tetraclinis articulata) in the Mediterranean region. In Germany it was found only along the upper and middle Rhine and in the Nahe valley (urban areas of Mainz, Frankfurt, Darmstadt and Cologne, as well as near Schloßböckelheim), below 200 m a.s.l. In Mainz the species was reported from 1993 through 1998 in parks and gardens on ornamental Chamaecyparis. Specimens in Darmstadt were swept from Ch. lawsoniana. Recently it was also published from the French side of the southern upper Rhine plain near Colmar. This is probably one of the rare cases of an invading host specialist exclusively attacking a plant which was introduced from other countries than the insect. Frommer (1996), Thüs (pers. comm.), Fröhlich & Nickel (unpublished data), HN; see also della Giustina & Balasse (1999)
Emelyanoviana contraria (Ribaut, 1936) So far M VIII – B IX; egg(?), 1(?) gen. Only known from the Pyrenees (France, Andorra), the Alps (North Tyrol, Piemonte, Hautes-Alpes) and mountainous regions of England and Scotland. Most records are from sun-exposed, rocky, rather dry calcareous hillsides from the upper montane to the alpine belt (found between 1500 and 2500 m a.s.l. in the Alps). Helianthemum sp. is reported as host plant in Britain. Recently the species has been recorded in Malaise trap samples from the Allgäu Alps: Ponten above Hindelang, c. 1850 m a.s.l., mid August until mid September 1998, altogether 6 ††, 4 ‡‡. Nickel & Voith (unpublished data); see also della Giustina & Meusnier (1982), Kirby (1992), Leising (1977), Remane (1961b), Remane & Fröhlich (1994b), Ribaut (1952), Woodroffe (1971)
Emelyanoviana mollicula (Boheman, 1845) M V – E X; egg (occasionally adult), 2 gen., perhaps 1 at higher altitudes. In sunny, dry to damp, occasionally also moist sites, mainly dry grassland, low-input meadows and pastures, and ruderal sites, occasionally also gardens. Host plants are
Typhlocybinae Kirschbaum, 1868
115
various herbs, mainly Salvia pratensis, Teucrium, Origanum, Thymus, and other species of Lamiaceae, as well as Verbascum spp.; is also reported from Heracleum, Mentha, Satureja, Cannabis, Parietaria, Artemisia, Fragaria, and additional dicotyledons. Widespread in Germany and common particularly in warmer regions, but apparently absent from most parts of the northwest German plain, frequently found up to at least 1350 m a.s.l. in the Mittelgebirge and the Bavarian Alps. Is also reported from the alpine belt in Switzerland (up to 2200 m a.s.l.), but breeding is uncertain in these sites. Achtziger (1991), Fischer (1972), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Rombach (1999b), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), Nickel (1997), HN; see also Günthart (1987a), Vidano (1965)
Dikraneura variata Hardy, 1850 Adults I – XII; but hibernation probably mainly in the egg stage, 2 gen. Among grasses, usually in acidic, damp to moderately dry, shady to moderately shady sites, locally and in lower abundances also on basic substrates; mainly in open forests and along woody margins, occasionally in parks and gardens. Often occurs in high abundance on Deschampsia flexuosa, preferentially under pine, beech, oak or spruce, in lower numbers on fescue (Festuca ovina, F. rubra, F. heterophylla). Widespread and common in northern and middle parts of Germany (including the Bavarian Forest, Swabian and Franconian Jura, and southern Black Forest, where found up to at least 1200 m a.s.l.), but apparently absent to the south of the Danube. Achtziger (1991), Reimer (1992), Remane (1987), Schiemenz (1990), Trümbach (1959), Wagner (1935, 1939a), HN, and others
Micantulina stigmatipennis (Mulsant et Rey, 1855) M V – B X; egg, 2 gen. On mullein (Verbascum lychnitis, perhaps also additional species) in sunny, moderately dry to dry, often disturbed sites on basic as well as acidic substrates. In Germany the species is rather scattered in lowland areas, at the western edge of the range; westernmost localities are near Frankfurt an der Oder, Potsdam, Dessau, Sondershausen, Jena and Meißen, three localities in the Franconian Jura near Eichstätt and Regensburg between 400 and 450 m a.s.l. appear to be rather isolated. Schiemenz (1990), Schumacher (1923), ZIMH, HN; see also Vidano (1965)
Micantulina micantula (Zetterstedt, 1840) In Italy B VII – E V; adult, probably 1 gen. (Vidano 1965). In Germany only known from two localities in Bavaria: Mertingen near Donauwörth, c. 400 m a.s.l., 05.V.1935, and Altmühl valley near Arnsberg, 450 m a.s.l., 27.IX.1996, 4 ††, 4 ‡‡, on a xerothermic hillside on Jurassic limestone with scattered shrubs and trees on Thalictrum minus. From Italy the species is reported from various species of Thalictrum in meadows, near shrubs and in forests, up to 1500 m a.s.l. Otherwise known from Scan-
116
Review of species
dinavia, northern Italy, former Yugoslavia, Greece, and large parts of boreal Asia; records from other countries may be dubious. Fischer (1972), HN; see also Drosopoulos et al. (1986), Nast (1972, 1987), Vidano (1965)
Wagneriala incisa (Then, 1897) So far M VII – E VIII; 1(?) gen.; in northern Italy according to Vidano (1965) E V – B XI; egg, at least 2 gen. In Germany only known from three records from Westphalia and Bavaria: Weser valley near Höxter, c. 200 m a.s.l., 1955 or 56, 1 specimen in an open beech forest on a steep, south-facing slope of Triassic limestone; Mainfranken: Sulzthal, 250 m a.s.l., 11.VII.1994, 1 ‡, and Berchtesgaden Alps: Winkl near Berchtesgaden, 700 m a.s.l., 22.VIII.1996, 2 ‡‡, both in open pine forests on xerothermic limestone plateaus and slopes. From northern Italy the species is reported to live in the undergrowth of chestnut and oak coppices with sedges, notably Carex caryophyllea, C. muricata and C. montana. In the German sites, the latter species is likely to be the host plant. Otherwise it is only known from Romania, former Yugoslavia, Austria, Italy, and the Baltic Sea islands of Gotland and Wollin. Rabeler (1962), Nickel (1999b); see also Holzinger (1999b), Nast (1987), Ossiannilsson (1981), Vidano (1965), Wagner (1955)
Wagneriala minima (J. Sahlberg, 1871) Mainly B VII – E VIII, occasionally also M VI and M X, nymphs so far B VII; egg(?), 1(?) gen. In sun-exposed, xerothermic pine forests on limestone, occasionally also on basic blown sand; mainly on Carex humilis, perhaps also on C. alba and additional sedge species. In Germany only known from the basin of the Saale (near Freyburg an der Unstrut, Bad Frankenhausen, Jena, Eisenberg and Arnstadt), the valley of the Altmühl between Eichstätt and Kelheim, the upper Isar between Vorderriß and Wolfratshausen, and the vicinity of Mainz, up to at least 1000 m a.s.l., but found at 1500 m a.s.l. in South Tyrol (Italy). Fischer (1972), Remane (1987, and pers. comm.), Schiemenz (1990), HN; see also Günthart (1987a)
Wagneriala sinuata (Then, 1897) E VI – B X; egg(?), 1(?) gen. On Carex flacca usually in moderately shady, rather dry and basic sites, mainly open pine forests and thermophilous margins of forests and shrubs on calcareous hillsides. In Germany at the northern edge of the range, known at present only from 12 localities in the northern Eifel Mountains, the Thuringian Eichsfeld, the Vogelsberg, Mainfranken and the Franconian Jura, between 200 and 500 m a.s.l. Remane & Fröhlich (1994b), Rombach (1999b, and pers. comm.), Wagner (1955), Nickel (1999b), HN
Forcipata citrinella (Zetterstedt, 1828) E V – E X; egg, 2 gen., probably 1 at higher altitudes.
Typhlocybinae Kirschbaum, 1868
117
In sedge stands in oligotrophic, usually peaty to temporarily moist (locally temporarily dry), basic to acidic sites, mainly straw meadows, fens and intermediate bogs; also in raised bogs, where confined to marginal slopes, ditches and peat diggings, rarely and in low numbers also in dry (often temporarily dry) grassland. Host plants are low-growing species of sedges, perhaps C. flacca, C. nigra and others. Widespread in Germany and locally common, particularly to the south of the Danube and in the Mittelgebirge, but rather scattered or even locally absent at lower altitudes where there was probably a dramatic decline due to intensification of agricultural management in recent decades. Found up to at least 1700 m a.s.l. in the Bavarian and Allgäu Alps, up to 1870 m a.s.l. in the French Alps. Fischer (1972), Marchand (1953), Reimer (1992), Schiemenz (1971a, 1976, 1990), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN; see also della Giustina (1989)
Forcipata forcipata (Flor, 1861) This species has often been confused and misinterpreted. Ribaut (1936) and Wagner (1939a) treated it as Dicraneura citrinella (Zett.).
B VI – E X, according to Schiemenz (1990) also M V; egg, 2 gen., probably 1 at higher altitudes. In open deciduous and coniferous forests and along their margins (mainly under pine and spruce, but also beech, oak and alder), in moist to dry sites from the lowlands up to the upper montane belt, at higher altitudes (above c. 1000 m a.s.l.) more in sunny and also wet to peaty sites (tall- and small-sedge swamps, alpine grassland). Host plants are various sedges (Carex flacca, C. humilis, probably also C. sylvatica, C. brizoides and others), woodrushes (Luzula spp.) and perhaps grasses. Widespread in Germany and common particularly in the limestone regions of middle and southern parts, found in high abundances up to 2000 m a.s.l. in the Bavarian and Allgäu Alps, in the French Alps even at 2500 m a.s.l. Fischer (1972), Niedringhaus (1997), Reimer (1992), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), HN; see also della Giustina (1989)
Notus flavipennis (Zetterstedt, 1828) M V – E X; egg, 2 gen., probably 1 at higher altitudes. In moderately wet to temporarily flooded, usually sunny sites, mainly in tall-sedge swamps, meadows, along ditches and margins of bogs, in wet ruderal sites, etc., usually on various species of tall sedges (notably Carex acutiformis, C. acuta, C. nigra, C. rostrata, C. vesicaria, C. paniculata, C. disticha), also found in slightly saline sites on Bolboschoenus maritimus. Widespread and common in Germany and one of the dominant species of many wetland sites, extending up to at least 1600 m a.s.l. in the Alps. Bittner & Remane (1977), Fischer (1972), Niedringhaus (1991), Reimer (1992), Schaefer (1973), Schiemenz (1988), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others
Kybos butleri (Edwards, 1908) B VI – B X; egg, 2 gen. On Salix triandra, less frequently on S. cinerea and S. aurita, along shores of rivers, streams, lakes and ponds, also on S. repens on dunes along the coasts of the North and Baltic Sea.
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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 reported from all adjacent countries and is probably much under-recorded. Frommer (pers. comm.), Niedringhaus (1991, 1997), Remane (pers. comm.), Wagner (1935, 1939a, 1955), Nickel & Remane (1996), HN; see also Dworakowska (1976), Nast (1987)
Kybos rufescens Melichar, 1896 B VI – B X; egg, 2 gen. On Salix purpurea along rivers and streams. Widespread in Germany, in intermediate to high frequency on the host, although less common in the north German plain. Frequently found up to at least 1000 m a.s.l. along alpine rivers in Bavaria, up to 1500 m a.s.l. in the Engadine (Switzerland). The locus typicus is near Munich. Fischer (1972), Frommer (pers. comm.), Nikusch (1976), Reimer (1992), Remane (1987), Schwoerbel (1957), Voigt (1978), Wagner (1951a, 1955), HN; see also Günthart (1987a)
Kybos limpidus (W. Wagner, 1955) B VI – E IX; egg, 2 gen. In floodplains of rivers and streams and along lake shores, mainly on Salix viminalis, less frequently on S. triandra, presumably also on S. pentandra. Widespread in the lowlands of Germany, in intermediate to high frequency and often large numbers on the main host, recorded up to 600 m a.s.l. in the foreland of the Alps. The locus typicus is Niedermarschacht near Hamburg. Dworakowska (1976), Emmrich (1975), Reimer (1992), Wagner (1955), Nickel & Remane (1996), HN
Kybos abstrusus (Linnavuori, 1949) M VI – M X; egg, 2 gen. On Populus nigra italica and P. n. nigra, preferentially in river floodplains, also in urban settlements (parks, sports grounds, etc.). Rather scattered in Germany, at the northwestern edge of the range (although perhaps under-recorded), most localities are known from larger river valleys: upper Rhine (from Speyer to Mainz, also on the Swiss and French side), Main (from Frankfurt to Haßfurt), Isar (Munich), Lahn (Marburg), Weser (Hannoversch Münden), Leine (Göttingen), Elbe (near Lüchow), also near Arnstadt and Balingen, up to at least 550 m a.s.l. Remane (1987, and pers. comm.), Wagner (1955), Nickel & Remane (1996), Nickel (1997), HN; see also Lauterer (1996)
Kybos populi (Edwards, 1908) B VI – E X; egg, 2 gen. On poplars (Populus tremula, P. nigra, P. alba, also hybrids) in floodplains of rivers and streams, in forests (usually along margins and roads, as well as in clearings), also in
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urban settlements. Widespread in Germany, in intermediate to high frequency on the hosts, found up to at least 800 m a.s.l. in the Mittelgebirge and the foothills of the Alps. Niedringhaus (1991, 1997), Reimer (1992), Remane (1987), Remane & Reimer (1989), Schiemenz (1990), Wagner (1935, 1955), Nickel & Remane (1996), HN
Kybos lindbergi (Linnavuori, 1951) and K. betulicola (W. Wagner, 1955) Empoasca betulicola W. Wagner, 1955 Dworakowska (1976), Nast (1972, 1987) and Ossiannilsson (1981) treat Kybos lindbergi (Lnv.) and K. betulicola (W.Wg.) as two distinct species. According to the first author, both hybridise in central Europe (although laboratory rearing has not been carried out). German specimens can only rarely be clearly assigned to either of the two. Transitional specimens have recently also been recorded from the Czech Republic (Lauterer 1996). The drawings of Dworakowska (1976) show considerable intraspecific variation in genitalic morphology and even appear contradictory in comparison with those of Ossiannilsson (1981) regarding the shape of the pygofer appendages. Furthermore, there are no differences in host preferences at least in central Europe. Hence, both forms are likely to belong to a single species with pronounced variability of male genitalic structures. In this case Linnavuori’s name has priority. In addition, there is a nomenclatural problem: according to Hamilton (1983a), K. betulicola (W.Wg.) is a junior synonym of Empoasca luda Davidson & De Long, 1938 described from North America. However, De Long’s drawing shows a species without lateral appendages of the aedeagus, which may belong to the butleri or limpidus group, but is clearly distinct from European birch-dwelling specimens. Hence, this opinion is refused, and Linnavuori’s name will be used here until more detailed biosystematic research and further investigations of type material have been carried out. Nevertheless, the European birch-feeding species occurs on introduced Betula pendula in Canada and the United States (I. Dworakowska, pers. comm.).
B VI – E X; egg, 2 gen. On birches (Betula pendula and B. pubescens) in wet to moderately dry sites, on solitary trees and along alleys, as well as in open forests. In the western half of Germany, including Thuringia, specimens with more or less intermediate genitalic morphology are widespread, although not yet recorded to the south of the Danube. Specimens from Berlin and Saxony increasingly show characters of the lindbergi type. Usually occurs in intermediate to high frequency and intermediate individual numbers on both species of birch. Highest localities are at 800 m a.s.l. in the German Mittelgebirge, at 1485m a.s.l. in the Swiss Alps. Niedringhaus (1991, 1997), Reimer (1992), Remane & Reimer (1989), Schiemenz (1990), Wagner (1955), Nickel & Remane (1996), HN; see also Günthart (1987a)
Kybos digitatus (Ribaut, 1936) In France VIII (della Giustina 1989); egg, 2 gen. Recorded in 2000 in the extreme southwest of Germany: Weil am Rhein, in a sand pit, probably on Salix purpurea or S. eleagnos. Otherwise only known from a few localities in southern and southwestern France; most of these records are from undetermined species of Salix. Mühlethaler (2001, and pers. comm.); see also Dworakowska (1977), della Giustina (1989)
Kybos smaragdula (Fallén, 1806) E V – E X; egg, 2 gen.
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On Alnus glutinosa and A. incana (also reported from A. alnobetula from Italian parts of the Alps), mainly along rivers, streams and lake shores, as well as on seepy slopes. Widespread and common in Germany and in high frequency on the host plants at least in lowland areas (up to c. 600 m a.s.l.), frequently, but less common in the Mittelgebirge and the Alps up to 1000 m a.s.l., recorded at 1600 m a.s.l. in the Engadine (Switzerland). Achtziger (1991), Fischer (1972), Heller (1987a), Niedringhaus (1991, 1997), Reimer (1992), Remane & Reimer (1989), Schiemenz (1990), Wagner (1935, 1939a, 1951a), HN; see also Günthart (1987a), Vidano & Arzone (1987a)
Kybos calyculus (Cerutti, 1939) So far E VI – E VIII (Schiemenz 1990); egg, 2(?) gen. At present this species is only known from Poland, England, Switzerland and eastern parts of Germany, where it was recorded in two sites in the 1980s: Dubringer Moor (near Hoyerswerda), E VIII 1987, and Berlin (Biesdorf), E VI – E VIII 1983. It is reported to live on Betula pubescens and is apparently confined to lower altitudes. The highest locality is the locus typicus in Switzerland (Chippis, Valais, c. 600 m a.s.l.). Schiemenz (1990); see also Cerutti (1939a), Dworakowska (1976)
Kybos strigilifer (Ossiannilsson, 1941) K. perplexus (Ribaut, 1952), showing very similar genitalic structures, has been described from the French side of the upper Rhine plain and has been reported from various mountainous regions of eastern and central Europe, as well as from central Asia. Like K. strigilifer (Oss.), it lives on Salix caprea and can be distinguished only by irregular spines on the aedeagus, which are more numerous and more strongly developed. However, German specimens of K. strigilifer (Oss.) show great variability of this character, even within the same populations and on the same host individual. It should also be mentioned that the few published notes on this problem are controversial. Günthart (1974), who reared the species under laboratory conditions, reported variability in the shape of the aedeagus and the anal tube appendages. However, if one follows Dworakowska (1976), Günthart’s drawings in fact indicate K. perplexus (Rib.). Hence, there appears to be no character clearly distinguishing these forms, which may in fact be conspecific. Furthermore, K. paraltaicus (Orosz, 1996) described after a single vagrant † from Hungary is likely to belong to this group.
M VI – E IX; egg, 2 gen. On Salix caprea, S. cinerea and S. myrsinifolia, mainly in forest glades, clearings, mining areas, fens, intermediate bogs, and along streams and ditches. Widespread in Germany, usually in intermediate frequency on the host plants, recorded up to 1200 m a.s.l. in the Allgäu Alps. Individuals clearly bearing characters of K. perplexus (Rib.) have been found in southeastern parts, notably Lower Bavaria. Niedringhaus & Olthoff (1993), Post-Plangg & Hoffmann (1982), Reimer (1992), Schiemenz (1990), Wagner (1955), Weis & Schönitzer (2001), Nickel & Remane (1996), HN; see also the drawings of della Giustina (1989), Dworakowska (1976, 1977a), Günthart (1974)
Kybos mucronatus (Ribaut, 1933) Has been treated as specifically distinct from Kybos mucronatus verbae Zachvatkin, 1953 by Dworakowska (1976) and later, Nast (1987) and della Giustina (1989). However, all figured specimens show an enormous intraspecific variation of the shape of aedeagus, anal tube appendages, ster-
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nal and tergal apodemes. The Bavarian specimen (see below) showed sternite apodemes as described for verbae, but tergite apodemes as shown for mucronatus (cf. della Giustina 1989). Thus, at least central European populations of both taxa are considered here as conspecific. Doubts on the distinctiveness have also been expressed by Günthart (1997).
In Poland and Switzerland VII – IX; egg, 1(?) gen. (Dworakowska 1976, Günthart 1987b). Largely confined to the region of the Alps and the Carpathians. Reported to live on Alnus glutinosa, but also on Salix. In Germany only known from three trap catches in the Rhineland and the Bavarian Alps: Krefeld, c. 40 m a.s.l., c. 1970, Burscheid, 200 m a.s.l., 3 individuals, July 1986, and Garmisch-Partenkirchen, Friedergries, c. 900 m a.s.l., 1 †, September 2000. Found between 450 and 1475m a.s.l. in the Swiss Alps. Kolbe & Bruns (1988, and pers. comm.), Coll. Munk. (Remane det.), Nickel & Voith (unpublished data); see also Dworakowska (1976), Günthart (1987b)
Kybos strobli (W. Wagner, 1949) In Switzerland VI – IX (Günthart 1987b); egg, 2(?) gen. This is apparently an endemic species of the Alps and the Carpathians, which is reported to live on Alnus incana. At present there are only two records from Germany, both comprising single †† only: Ulm (Swabian foreland of the Alps), c. 500 m a.s.l., 03.VIII.1908, and Hindelang (Allgäu Alps), 1200 m a.s.l., 02.VIII.1995. Recorded from altitudes between 460 and 1560 m a.s.l. in Switzerland. Dworakowska (1976) (see this paper also for identification), Nickel (1999b); see also Günthart (1987b)
Kybos virgator (Ribaut, 1933) Kybos volgensis Vilbaste, 1961, described from the Astrakhan region (southern Russia) on the basis of differences in body size and adeagus shape, has been reported from Sweden, Switzerland and eastern Germany (Ossiannilsson 1981; Günthart 1987a; Sander et al. 1999). However, due to the high variability and incongruent descriptions of Russian and Swedish specimens the specific identity is in need of further clarification.
B VI – E X; egg, 2 gen. On Salix alba and S. fragilis in floodplains of rivers and streams, locally also on S. viminalis, S. aurita, S. cinerea and S. babylonica (also far from water), few specimens found on S. pentandra. Widespread and common in the lowlands of Germany, usually in high frequency and large numbers particularly on Salix alba and S. fragilis, but recorded only up to 600 m a.s.l. in the Mittelgebirge and the foreland of the Alps. Fischer (1972), Lauterer & Malenovsky (1995), Mölleken & Topp (1997), Niedringhaus (1991, 1997), Reimer (1992), Wagner (1935, 1939a, 1951a), HN, and others
Empoasca affinis Nast, 1937 †† E VII – B XI, ‡‡ probably until V/VI, but these cannot be identified with certainty; ‡‡(?), 1(?) gen. Along margins of shrubby and herbaceous vegetation in sunny, damp to dry sites, mainly dry grassland, ruderal sites, abandoned vineyards, forest margins, hedges, etc.; most
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individuals are swept from deciduous shrubs and trees (Rubus, Ulmus, Carpinus, Corylus, Populus, Salix and others), and in the herb layer, often in low numbers only. In Germany 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, Sondershausen, Berlin and Frankfurt an der Oder; single individuals have been found near Cologne, Helmstedt, Lauenburg and Kaufbeuren, c. 650 m a.s.l., but usually below 400 m a.s.l. Fischer (1972), Frommer (1996), Reimer (1992), Remane (1987), Remane (pers. comm.), Schiemenz (1990), Schönitzer & Oesterling (1998b), Wagner (1951a, 1952), HN
Empoasca apicalis (Flor, 1861) E VIII – M V; adult, 1 gen. (see Günthart 1987a; Lauterer 1980). This species‘ range essentially includes Siberia and Scandinavia, with isolated populations in central Europe. From Germany there are only three records from southern parts: Illasberg near Füssen, c. 800 m a.s.l., 13.V.1951; upper Neckar valley near Rottweil, 23.IX.1956, and Breitbrunn am Ammersee, 23.VIII.1959, in both sites on Sambucus ebulus. In the Engadine (Switzerland), nymphs and adults were found on Lonicera xylosteum at c. 1100 m a.s.l. in spruce forests with luxuriant undergrowth. In Moravia (Czech Republic), single individuals were recorded on Lonicera nigra between 290 and 620 m a.s.l. in shady forests. Finnish populations have been reported to migrate from L. xylosteum to Alnus incana, Juniperus and Picea in autumn. Fischer (1972), Remane & Fröhlich (1994b); see also Günthart (1987a), Lauterer (1980), Nuorteva (1948, 1952a)
Empoasca decipiens Paoli, 1930 Adults I – XII, mainly M VII – E V; adult (including ††!), 2(?) gen. In various, damp to dry, sunny to moderately shady sites, usually along margins of herbaceous and shrubby vegetation, in ruderal sites, gardens and fields. Mainly on weeds in the latter, but locally causing feeding damage on cucumbers and sweet pepper in south German greenhouses. Also reported to be noxious in various cultures of vegetables and fruits in more southern European countries, although species identification may not be accurate in some cases. Host plants are mainly dicotyledonous herbs belonging to many families (Rosaceae, Fabaceae, Solanaceae, Caprifoliaceae, Asteraceae and others) and – particularly in autumn – also on shrubs (Rosa, Rubus, Cornus, Corylus, Sambucus and others), during hibernation presumably again on herbs. Widespread and common in the lowlands of middle and southern parts of Germany, more scattered and increasingly preferring synanthropic habitats towards the north. Usually confined to altitudes below 500 m a.s.l., but locally found up to 800 m a.s.l. along the edge of the Bavarian Alps. Reported from 1300 m a.s.l. in the Valais (Switzerland). From Scandinavia there are only two known records from southern Norway and southern Sweden; likewise, there is only a single record from Denmark. Achtziger (1991), Frommer (1996), Niedringhaus & Olthoff (1993), Post-Plangg & Hoffmann (1982), Raupach (pers. comm.), Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1951a), HN, and others; see also Cerutti (1939a), della Giustina (1989), Gillerfors (2002), Günthart (1971a, 1987a), KobletGünthardt (1975), Ossiannilsson (1981), Wagner & Franz (1961)
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Empoasca pteridis (Dahlbom, 1850) Adults I – XII, mainly M VIII – E X, according to laboratory and field studies of Günthart (pers. comm.) †† and ‡‡ hibernate (the former only in very low numbers), according to Remane (1987) hibernation takes place mainly in the egg stage, 2(?) gen. In moist to damp, occasionally also wet or dry, sunny to moderately shady, usually disturbed sites, mainly fertilized meadows and pastures, herbaceous margins, fields, ruderal sites, waysides, etc., but found in low numbers and perhaps only as vagrants in most other types of non-forested terrestrial habitats (e.g. dry grassland, wet meadows, also on various deciduous shrubs and trees). Host plants are various herbs including cultivated potato. Widespread and common at lower altitudes of Germany, locally found up to c. 1000 m a.s.l. in the Mittelgebirge and along the edge of the Alps. Achtziger (1991), Reimer (1992), Remane (1987), Schiemenz (1990), Wagner (1935, 1939a), Wais (1990), Nickel & Achtziger (1999), HN, and others; see also Günthart (1971b)
Empoasca ossiannilssoni Nuorteva, 1948 This taxon is likely to be conspecific with E. dealbata Cerutti, 1939, which was described after a single † from the Valais (Switzerland). Only one further † from Styria (Austria, on Prunus padus) has been recorded since (Wagner & Franz 1961). A number of very similar taxa has been described also from Siberia (see Anufriev & Emelyanov 1988). However, the shape of the pygofer appendages, which has been used as the most important distinguishing character in this genus, has been found to be much variable (e.g. Günthart 1974; Nickel, unpublished data). Thus, a revision of Palearctic species would be much welcome.
In Sweden and Finland E VII – B IV; adult, 1 gen. (see Ossiannilsson 1981). Until recently, this taxon has been known only from the coniferous forest zone of Eurasia and the Ukrainean Carpathians. A more recent record is from the Bavarian foothills of the Alps: Hackensee near Dietramszell, 660 m a.s.l., 09.IX.1994, 1 †, along a forest margin near a lake shore with dominating Picea abies, Prunus padus and Corylus avellana. Data on the biology are scarce. In Sweden and Latvia, the species has been found on Prunus padus and in a birch forest; hibernation is suspected to take place on Picea. HN; see also Anufriev & Emelyanov (1988), Nast (1972), Ossiannilsson (1981)
Empoasca vitis (Göthe, 1875) The use of the name E. flavescens (F.) for this species until the second half of the 20th century is caused by misinterpretation of Fabricius‘ description, which was finally clarified by Wagner (1961) after a study of the type material.
Adults I – XII, mainly B VII – M V; adult (also ††), 1 gen. Among all central European arboricolous leafhoppers, this species shows the widest range of host plants, living extremely polyphagously on various woody plants: Fagus, Quercus, Betula, Alnus, Carpinus, Corylus, Ulmus, Rosa, Rubus, Malus, Sorbus, Prunus, Acer, Frangula, Rhamnus, Cornus, Vitis (in southern Europe and southwestern Germany causing feeding damage in some years), Populus, Salix, Tilia, Viburnum, Lonicera, Fraxinus, even on introduced species (Platanus, Aesculus, etc.), frequently also in the herb layer (at least adults on Solanum tuberosum, Althaea, Helianthus and Petasites). In late summer
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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 typhlocybine leafhopper, frequently occurring up to the subalpine tree line, found up to at least 1500 m a.s.l. in Bavaria, up to 2000 m a.s.l. in Switzerland. Achtziger (1991), Fischer (1972), Herrmann et al. (1999), Louis & Schirra (1997), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1990), Schruft & Wegner-Kiß (1999), Wagner (1935), Wais (1990), HN and others; see also Cerutti (1939a), Günthart (1987a)
Austroasca vittata (Lethierry, 1884) E V – B X; egg, 2 gen. On Artemisia absinthium in moderately dry, usually disturbed sites (ruderal sites, waysides, abandoned vineyards, etc.), occasionally also on A. maritima in inland saltmarshes. In Germany, at the northwestern edge of the range, this species has been recorded since the 1920s, occurring only locally and rather scattered in warm-summer regions, with strongholds in Saxony, the rain shadow area to the east of the Harz Mountains, and the Rhine valley between Mannheim and Cologne, single records are from the Wendland (eastern Niedersachsen), Mecklenburg, the Uckermark, northern Hessen and Mainfranken, usually below 300 m a.s.l., but reported from 1500 m a.s.l. in the Engadine (Switzerland). Fröhlich (1996a), Frommer (1996), Schiemenz (1990), Walter (pers. comm.), Nickel (1997), HN; see also Günthart (1987a)
Kyboasca bipunctata (Oshanin, 1871) B VI – E IX; egg, 2 gen. On Ulmus minor and U. x hollandica, perhaps also U. laevis, mainly on low-growing individuals (coppices, saplings), usually in sunny or moderately shady sites. Has been found on Cannabis and Glycyrrhiza in Poland and the Czech Republic. In Germany only recorded in very scattered localities, almost exclusively in urban habitats in river valleys (Monheim near Düsseldorf, Cologne, Berlin, Halle, Bitterfeld, Göttingen, Arnstadt, Jena, Görlitz and Badberg/Kaiserstuhl, also near Stettin, Poland), up to 400 m a.s.l.; is also known from a few localities to the west and north (southern France, Netherlands, England and southern Finland). Frommer (pers. comm.), Kolbe & Bruns (pers. comm.), Schiemenz (1990), MTD, Nickel (1994), HN; see also della Giustina (1989), Dworakowska (1973), Haupt & Hedicke (1934), Lauterer (1984), Nast (1987)
Chlorita dumosa (Ribaut, 1933) This species and the following belong to a subgroup, which was named Xerochlorita Zachvatkin, 1953. Most authors treat it as a subgenus of Chlorita Fieb. However, Zachvatkin did not provide a description, and consequently, the name is not valid and has to be treated as a nomen nudum.
M V – M X; egg, at least 2 gen. In cushions of thyme (Thymus praecox, Th. serpyllum, probably Th. pulegioides) in sunny, dry to moderately dry, low-vegetated, usually grazed sites, preferentially on sand and
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limestone, but also on other well-drained substrates. In Germany at the northwestern edge of the range, with strongholds in warmer regions of southern and eastern parts. Most records are from the rain shadow area to the east of the Harz Mountains, Thuringia and Mainfranken; rather scattered localities are near Cologne, Mainz, Heidelberg, Regensburg, Göttingen, Frankfurt an der Oder and Bautzen, up to 450 m a.s.l. Frommer (pers. comm.), Reimer (1992), Remane (1987), Schiemenz (1990), Wagner (1941a), Nickel (1994), HN
Chlorita pusilla (Matsumura, 1906) See taxonomic remark above!
B VI – E IX; egg, at least 2 gen. This species is altogether known only from 7 localities in dune regions of western Europe: Brodnica (previously Strasburg, to the northeast of Warsaw/Poland), “zahlreiche Exemplare (...) auf niedrigen krautartigen Pflanzen”; Vieille-St.-Girons, 26.VI.1996, VieuxBoucau-les Bains, 06.IX.1997 (both south-western France); Gartz near Angermünde, 12.VIII.1937, on Thymus cushions; Weinberg near Perleberg, 1965 and 1966, altogether 9 ††, 2 ‡‡ on a “Schafschwingel-Schillergras-Sandtrockenrasen”; Boizenburg, 15.VI. 1998, 9 ††, 50 ‡‡, as well as Gerwisch near Magdeburg, 01.VI.2000, 2 ††, 3 ‡‡, the two latter records on inland dunes on Thymus serpyllum. Schiemenz (1969), Wagner (1941a), HN; see also della Giustina & Remane (2001), Matsumura (1906)
Chlorita paolii (Ossiannilsson, 1939) B V – M X; egg, at least 2 gen. In sunny and oligotrophic, damp to dry sites on various substrates, mainly dry grassland, low-productivity meadows and pastures, ruderal sites and roadside embankments. Host plants are Achillea millefolium and Artemisia campestris, locally A. absinthium and A. vulgaris. Widespread and common in many places of eastern and southern Germany, but becoming rather scarce towards the northwest, where living at the edge of the range, and towards the edge of the Alps. Border localities are known from near Koblenz, Marburg, Göttingen, Braunschweig, Uelzen and Hamburg; frequently recorded at 1100 m a.s.l. in higher parts of the Mittelgebirge. Not reported from Denmark and southern Sweden, although widespread in Mecklenburg. Also common on the German side of the upper Rhine plain, although in France known only from the western Alps (up to 1700 m a.s.l.), Burgundy and Auvergne. Furthermore, there is a single record from the Netherlands. Büttner (1964), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), Nickel (1994), HN; also Cobben & Gravestein (1958), della Giustina (1989, and pers. comm.)
Fagocyba carri (Edwards, 1914) So far E V – B IX; egg, 2 gen. On oaks (Quercus robur and Qu. petraea) in forests and along their margins. Probably widespread in the lowlands of Germany, but only poorly documented and apparently
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in rather low frequency and individual numbers; neither recorded in northeastern parts (Mecklenburg-Vorpommern, Saxony-Anhalt) nor in the foreland of the Alps. The highest localities are at 500 m a.s.l. in the Mittelgebirge. Achtziger (1991), Günthart (1988), Niedringhaus (1991, 1997), Reimer (1992), Remane (1987), Schiemenz (1990), Wagner (1935), HN
Fagocyba cruenta (Herrich-Schäffer, 1838) Typhlocyba douglasi Edwards, 1878; Typhlocyba inquinata Ribaut, 1936 This taxon is considered here as a single species comprising three morphs differing only in characters of colouration, but not in habitat preferences, host plants, phenology and distribution, although the more strongly pigmented reddish (= cruenta) morph apparently occurs only in the summer generation. Some authors, however, treat the yellowish white F. douglasi (Edw.), the reddish F. cruenta (H.-S.), and F. inquinata (Ribaut, 1936), which shows a fumose clavus, as distinct species. Striking intraspecific differences in colouration are also found within other species of Typhlocybini, e.g. Eurhadina and Eupteryx.
M V – E X; egg, 2 gen., perhaps 1 at higher altitudes. The more widespread yellowish white douglasi morph is eurytopic and polyphagous on various deciduous woody plants, in closed forests as well as on solitary trees. Preferentially on Fagus and Carpinus (where locally extremely abundant), but also – roughly in order of decreasing densities – on Acer, Tilia, Ulmus, Prunus, Sorbus, Alnus, Betula, Corylus, Populus, Salix, Aesculus, Quercus, Rosa, Crataegus, Fraxinus and others, even Juglans. Very common and widespread all over Germany, locally with a low proportion of female individuals of var. inquinata (Rib.). Frequently found up to 1800 m a.s.l. in the Bavarian and Allgäu Alps, and perhaps reproducing on subalpine shrubs. The more scattered and uncommon reddish cruenta morph shows the same host preferences and geographic distribution, although it has not been recorded in the north German plain (but in Denmark and southern Sweden, as well as near the alpine tree line). Most individuals are found among large numbers of the douglasi morph, but the earliest central European specimens date from the first decade of July. Achtziger (1991), Bittner & Remane (1977), Frommer (1996), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1990), Schönitzer & Oesterling (1998b), SMNS, Wagner (1935, 1939a), HN, and others; see also Claridge & Wilson (1976, 1981), Nast (1987), Ossiannilsson (1981)
Ossiannilssonola callosa (Then, 1886) B VI – M IX; egg, 1(?) gen. On Acer pseudoplatanus in forests, parks and on solitary trees, usually in cool sites (valley bottoms, shady slopes, near water). Widespread in Germany and usually in intermediate frequency on the host, but only poorly documented, frequently occurring up to at least 1300 m a.s.l. in the Bavarian and Allgäu Alps, although single specimens were found at 1500 m a.s.l. Reimer (1992), Remane & Reimer (1989), Schiemenz (1990), Wagner (1935), Nickel (1999b), HN; see also Whittaker (1984)
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Edwardsiana alnicola (Edwards, 1924) So far E VI – B VIII; egg, 2(?) gen. On alders (Alnus glutinosa, A. incana) in moist to wet sites, often near water. The distribution in Germany is rather poorly documented; the species is known only from scattered records in the western half, between the North Sea coast and the Alps, where it occurs at least in low frequency and individual numbers. Highest localities are at 1000 m a.s.l. in Bavaria, but at 1660 m a.s.l. in the Engadine (Switzerland). The species is not reported from France, Belgium and the Netherlands; thus, the western edge of its range may run through Germany. Achtziger (1991), Frommer (pers. comm.), Kolbe & Bruns (1988), Niedringhaus & Olthoff (1993), Wagner (1935), Nickel (1997), HN; see also Günthart (1987a)
Edwardsiana avellanae (Edwards, 1888) Typhlocyba staminata Ribaut, 1931 Many authors consider E. staminata (Rib.) to be a distinct species (e.g. Nast 1972, 1987; Ossiannilsson 1981). Compared to E. avellanae (Edw.), two tiny appendages of much variable length on the aedeagus tip are the only distinctive characters. Moreover, most, if not all of these individuals are pipunculized or otherwise abnormal. Body size, colouration, host plant, habitat and distribution (also within Germany) are almost identical to E. avellanae (Edw.). Therefore, Lauterer (pers. comm.), della Giustina (1989) and Le Quesne & Payne (1981) consider them conspecific. This interpretation is adopted here.
B VI – M X; egg, 2 gen. Monophagous on Corylus avellana; most individuals are swept along forest margins and roads, generally in more shady and cool sites than E. spinigera (Edw.). Widespread in Germany, fairly common and in intermediate frequency, found up to at least 850 m a.s.l. in the Mittelgebirge and the Bavarian Alps, up to 1400 m a.s.l. in the Valais (Switzerland). Achtziger (1991), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1990), Trümbach (1959), Wagner (1935, 1939a), HN; see also Cerutti (1939a)
Edwardsiana bergmani (Tullgren, 1916) B VI – B IX; egg, 2 gen., probably 1 at higher altitudes. On birches (Betula pendula, B. pubescens) and alders (Alnus glutinosa, A. alnobetula), usually in moist, cool or shady habitats (e.g. bogs, forests, subalpine alder shrub, near streams). Widespread in Germany, but rather scattered and in low frequency and individual numbers, found up to at least 1500 m a.s.l. in the Bavarian Alps. Achtziger (1991), Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1990), HN
Edwardsiana candidula (Kirschbaum, 1868) E. candidula martigniaca (Cerutti, 1939), described from Switzerland is distinguished only by slight differences in shape and length of the aedeagus appendages, but apparently not in life history. Recently, it has been treated as a distinct species by various authors (e.g. Nast 1987; Lauterer, pers. comm.). Both forms occur in central Europe, although only the nominate one has
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been recorded from Germany. The drawings of Ribaut (1936) show the latter, those of Ossiannilsson (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 frequency on the host in most regions, but apparently absent from northwestern parts. Localities at the edge of the range include Cologne, Göttingen, Lauenburg and Lübeck; found only up to 500 m a.s.l. in the foreland of the Alps. Frommer (pers. comm.), Remane (1987, and pers. comm.), Schiemenz (1990), Wagner (1935, 1939a), Nickel & Remane (1996), Nickel (1997), HN
Edwardsiana crataegi (Douglas, 1876) A morph with different shape and length of the aedeagus appendages has been described as Typhlocyba froggatti Baker, 1925. However, according to laboratory rearings of Günthart (1971b) the offspring of a single ‡ showed both the aedeagus type of the nominate form and of froggatti Bak.; the latter is generally considered to be a variety. Regarding individual numbers, it dominates in most parts of the study area.
E V – M X; egg, 2 gen. On woody species of Rosaceae, mainly Malus, Crateagus, Prunus domestica, P. avium, P. spinosa; is also reported from Mespilus and Sorbus from Switzerland and Wales. Has been reported to cause damage on apple trees in southern Europe and elsewhere. Widespread and common in Germany, found up to at least 700 m a.s.l. in the Mittelgebirge, up to 1400 m a.s.l. in the Swiss Alps. Achtziger (1991), Frommer (1996), Lehmann (1973a), Müller (1956), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), HN, and others; see also Claridge & Wilson (1981), Günthart (1971b, 1987a)
Edwardsiana diversa (Edwards, 1914) According to China (1943), the name Anomia tridentata Edwards, 1928, which had been created for a parasitized specimen, is a junior synonym of E. diversa (Edw.). Wagner (1935) published records of parasitized individuals from Bremen, the aedeagus of which resembled the one of A. tridentata Edw., but he argued that its specific status may be uncertain. China’s opinion is apparently based only on the fact that A. tridentata Edw. (only 1 †!) had been taken from Cornus. Thus, its status is doubtful, since parasitized Edwardsiana males are often impossible to identify. Hence, the occurrence of E. diversa (Edw.) in northern Germany is not proven.
B VI – E X; egg, 2 gen. On dogwood (Cornus sanguinea, C. mas) in damp to dry, usually eutrophic or basic sites, mainly dry grassland, along hedges, forest margins and roads, as well as in parks. In Germany at the northern edge of the range, only known from widely-separated localities, but apparently widespread in middle parts and at least locally in intermediate to high frequency on the hosts (up to at least 450 m a.s.l.). Not found in the north German plain and to the south of the Danube; the northernmost localities are near Cologne, Göttingen and Magdeburg.
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Frommer (pers. comm.), Lehmann (1973a, 1973b), Müller (1978), Nicolaus (1957), Reimer (1992), SMNS, Nickel & Remane (1996), Nickel (1994), HN
Edwardsiana flavescens (Fabricius, 1794) For a long time, most authors distinguished E. fratercula (Edwards, 1908) and E. sororcula (Ossiannilsson, 1936) after differences in shape and length of the lateral aedeagus appendages. However, since Wagner & Franz (1961) both names are treated as junior synonyms of E. flavescens (F.).
E V – E X; egg, 2 gen. Preferentially on Carpinus betulus and Fagus sylvatica in forests, parks and along hedges, occasionally also on other species of deciduous woody plants (e.g. Salix, Quercus, Acer, Ulmus and Betula). Widespread in Germany and usually common on beech and hornbeam, frequently found up to at least 900 m a.s.l. in the Mittelgebirge and the Bavarian Alps, single specimens at 1200 m a.s.l. Bittner & Remane (1977), Förster (1961), Niedringhaus & Olthoff (1993), Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Schiemenz (1990), Wagner (1935, 1939a), Nickel & Remane (1996), HN
Edwardsiana rhodophila (Cerutti, 1937) E VI – B X; egg, 2 gen. Previously, this species was only known from the Mediterranean region, notably southern France, Switzerland (Valais), Greece, northern Italy and eastern Spain, reported from „ ... speziellen Rosa-Arten in mehr oder weniger montanen Bereichen“. Recently also recorded in the Saale-Unstrut region and Mainfranken, where it lives on Rosa rubiginosa in xerothermic sites, mainly in upper slope and plateau situations on limestone and gypsum. At present there are altogether 9 German localities from the surroundings of Nordhausen, Artern, Freyburg an der Unstrut, Ilmenau, Gotha, as well as Karlstadt am Main, some of them holding high individual numbers. Found up to 1530 m a.s.l. in Spain, but only up to 410 m a.s.l. in Germany. Nickel (1998); see also Cerutti (1937), Remane & Fröhlich (1994b)
Edwardsiana sociabilis (Ossiannilsson, 1936) M VI – B X; egg, 2 gen. This species is reported from northern and central parts of Europe, but also from Georgia. In Germany it occurs mainly in urban areas and along hedges on ornamental roses, mainly Rosa rugosa. Probably it is more widespread, but is only known from scattered localities (East Frisian Islands, Bremen, Hamburg, Göttingen, Marburg and surroundings, Hof, Görlitz, Steingaden/Upper Bavaria, up to at most 800 m a.s.l.). Furthermore, populations have been recorded in near-natural habitats on Filipendula ulmaria in tall herb stands of rather wet or cool sites (northeastern Harz Mountains, Elbe valley near Dresden, Vogelsberg, Thuringian Forest, Bad Königshofen im Grabfeld). The species is reported to be common on cultivated Rosa sp. and Rubus idaeus in gardens in southern
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Finland, and from Filipendula ulmaria in Sweden. This is one of the rare documented cases of a stenophagous native insect species infesting an introduced plant. Achtziger (1991), Emmrich (1975), Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Wagner (1937g), HN; see also Linnavuori (1952), Ossiannilsson (1981)
Edwardsiana frustrator (Edwards, 1908) E V – E X; egg, 2 gen. On various deciduous woody plants, without any apparent host preference: Corylus, Acer, Quercus, Aesculus (based on laboratory rearings or nymphal records in Wales and Switzerland), also occasionally found on Carpinus, Tilia, Ulmus, Fagus, Betula, Malus, Prunus, Rosa, Crateagus, and probably additional species. Widespread in the lowlands of Germany, but usually in rather low numbers, found only up to 600 m a.s.l. in the Mittelgebirge. Not recorded from Mecklenburg-Vorpommern, where probably overlooked. Achtziger (1991), Frommer (1996), Niedringhaus & Olthoff (1993), Post-Plangg & Hoffmann (1982), Reimer (1992), Schiemenz (1990), Wagner (1935, 1939a), HN; see also Claridge & Wilson (1976, 1981), Günthart (1974)
Edwardsiana ampliata (W. Wagner, 1948) Edwardsiana solearis rossica Zachvatkin, 1948 Resembling E. frustrator (Edw.) in genital morphology and life habits. Both are considered only as distinct subspecies by Zachvatkin (1948) and Anufriev (pers. comm.).
B VI – B X; egg, 2 gen. Described from Silesia (Poland) and since then mainly reported from the deciduous forest region of eastern Europe. Recently it was recorded in Mainfranken, where it is currently known from altogether 6 localities near Schweinfurt and Bad Königshofen, along forest margins on various deciduous trees (Acer campestre, A. pseudoplatanus, Corylus avellana, Quercus robur), between 200 and 300 m a.s.l.; all records comprised only 1 or 2 †† respectively. Apparently polyphagous, reported to live on Quercus and Tilia in Moravia (Czech Republic), mainly on Tilia cordata in Poland, on Quercus, Corylus and Cerasus in middle parts of Russia. Nickel & Remane (1996), HN; see also Anufriev & Kirillova (1998), Chudzicka (1986a), Lauterer (pers. comm.), Zachvatkin (1948)
Edwardsiana geometrica (Schrank, 1801) E VI – M X; egg, 2 gen., perhaps 1 at higher altitudes. On alders in various damp to wet sites, usually on Alnus glutinosa in lowland regions, preferentially on A. incana in the Mittelgebirge and the Alps. Widespread in Germany and, compared to other Edwardsiana species, well documented due to the striking colouration. Frequently found up to at least 1000 m a.s.l. in the Mittelgebirge and the Bavarian and Allgäu Alps, single specimens up to 1800 m a.s.l. Also found in the subalpine belt in the Engadine and Valais (both Switzerland), where presumably living on Alnus alnobetula.
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Achtziger (1991), Fischer (1972), Frommer (pers. comm.), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Remane & Reimer (1989), Schiemenz (1990), Wagner (1935, 1939a), HN; see also Cerutti (1939a), Günthart (1987a), Vidano & Arzone (1987a)
Edwardsiana gratiosa (Boheman, 1852) B VII – M X; egg, 2 gen. Monophagous on Alnus glutinosa, usually near standing or running water. Known from sporadic localities in the lowlands of Germany, but probably under-recorded (e.g. not yet found to the south of the Danube, nor in Rhineland-Palatinate and MecklenburgVorpommern, but from adjacent regions of Poland and France). Usually in intermediate frequency, but low individual numbers on the host; highest localities are in the Mittelgebirge at 600 m a.s.l. only. Frommer (1996), Heller (1987a), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer (1992), Schiemenz (1990), Wagner (1935, 1939a), Nickel & Remane (1996), HN; see also Nast (1976a), Ribaut (1936)
Edwardsiana lamellaris (Ribaut, 1931) This species is only known from a few countries in central and southern Europe and is apparently rare in most parts of its range. Specimens published from the surroundings of Halle turned out to belong to E. prunicola (Edw.) after a revision. The only definitive record from Germany is from a moderately dry grassland site with scattered oaks near Stuttgart: Eichenhain near Riedenberg, c. 400 m a.s.l., 08.VI.1994, 2 ††, 1 ‡, on Rosa canina. This plant is also likely to be the host in Moravia (Czech Republic). From Italy, however, the species is reported to live monophagously on Quercus petraea and Qu. robur. Perhaps there is a host shift between the first and the second generation. Lehmann (1973a), SMNS (Heller leg. et det.); see also Lauterer (1984), Vidano & Arzone (1987b)
Edwardsiana lethierryi (Edwards, 1881) B VI – B X; egg, 2 gen. On Acer campestre, Tilia cordata and T. platyphyllos (in England and Wales also on Aesculus hippocastanum), in lower numbers and without evidence for breeding also on other deciduous trees (Ulmus, Acer platanoides, Corylus and others). Widespread at least in the lowlands of Germany (including the East Frisian Islands, Heligoland, Holstein and Mecklenburg), but poorly documented; highest known localities are at only 550 m a.s.l. in the Mittelgebirge. Recorded in northern Europe only from the island of Gotland and southern Norway. Fischer (1972), Frommer (1996), Niedringhaus (1991, 1994), Nikusch (1976), Reimer (1992), Schiemenz (1990), Schwoerbel (1957), Wagner (1939a), HN; see also Claridge & Wilson (1981), Günthart (1974), Ossiannilsson (1981)
Edwardsiana plurispinosa (W. Wagner, 1935) The recent revision of the E. lethierryi group presented by Wilson & Claridge (1999) could only partially be taken into consideration for this book. Populations hitherto treated as E. hippocastani (Edw.) sensu Ribaut were split into two species, but the name was shown to be a junior
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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. alnobetula) in forests and in semi-open habitats; single individuals (probably without reproduction) are found on most native deciduous tree species. Widespread in the lowlands of Germany, but poorly documented; usually in intermediate to high frequency on the hosts, up to at least 700 m a.s.l., although E. hippocastani (Edw.) is mentioned from Alnus at 1600 m a.s.l. in the Swiss Alps. Niedringhaus & Olthoff (1993), Wagner (1935), HN; see also Cerutti (1939a), Claridge & Wilson (1976, 1981); for identification see Wilson & Claridge (1999)
Edwardsiana ulmiphagus Wilson & Claridge, 1999 E V – B X; egg, 2 gen. On elms (Ulmus minor, U. glabra, U. laevis) in forests, parks and on solitary trees; single vagrants also on other deciduous trees. Widespread in the lowlands of Germany, but poorly documented, usually in intermediate to high frequency and individual numbers on all three elm species. Highest localities are at 700 m a.s.l. in the Mittelgebirge, but E. hippocastani (Edw.) has been reported from Ulmus at 1230 m a.s.l. in the Engadine (Switzerland). HN; see also Claridge & Wilson (1976, 1981), Günthart (1987a); for identification see Wilson & Claridge (1999)
Edwardsiana ishidai (Matsumura, 1932) Opinions on the status of this taxon and the following are contradictory: according to Dworakowska (1982), Nast (1972, 1987) and Ossiannilsson (1981) Typhlocyba lanternae W.Wg. is a junior synonym of T. ishidai Mats.; but Lauterer (1984) and Le Quesne & Payne (1981) argue that there are differences in the shape of the aedeagus and in host choice. Evidence from German populations supports the latter view, although the total number of records of E. lanternae (W.Wg.) is extremely low. Until the problem has been resolved, both shall be treated as separate taxa.
E V – M X; egg, 2 gen. E. ishidai (Mats.) is apparently widespread in middle and southern parts of Germany, extending northward at least to Cologne, Göttingen, Nordhausen and Eberswalde. Is usually swept along forest margins and roads, in parks, and from solitary trees, recorded up to 350 m a.s.l. only. Host plants are Ulmus minor, U. glabra, U. laevis, and perhaps Corylus avellana. Not recorded from France, but from the Baden and Palatinate part of the upper Rhine plain. Frommer (pers. comm.), Nicolaus (1957), SMNS, Nickel & Sander (1996), Nickel (1994, 1999b), HN; see also Günthart (1987b)
Edwardsiana lanternae (W. Wagner, 1937) See taxonomic remarks above.
So far E VI – B IX; egg, 2 gen.
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Described after 2 †† collected in street lamp domes in Hamburg-Altona, shortly afterwards 1 † was recorded in the Duvenstedter Brook to the north of Hamburg. There are two more recent records from the Emsland near Lingen „1 † in einem Alnus-dominierten Feuchtgebüsch, 04.VII.89, 1 † in einer Mischgehölzhecke, 04.IX.89“. Reported from Alnus glutinosa and Sorbus aucuparia in England and Wales. Otherwise the distribution of this taxon is poorly known, since it is usually considered to be conspecific with the preceding one (see above). Wagner (1937d, 1941b), Niedringhaus & Olthoff (1993); see also Claridge & Wilson (1976), Le Quesne & Payne (1981)
Edwardsiana soror (Linnavuori, 1950) In the Czech Republic and Slovakia B VI – B X (Lauterer 1980); egg, 2 gen. The range of this species, which is usually reported from Alnus incana, extends from eastern Siberia to the eastern half of central Europe. There are only two German records: Oberlausitz: Quoos, c. 150 m a.s.l., 02.IX.1958, 17 †† „auf Teichdämmen an Alnus“, and Mittelfranken: Langensendelbach, 340 m a.s.l., 09.VII.2001, 1 †, on Alnus incana in the undergrowth of an open pine forest. Has been found at 930 m a.s.l. in Carinthia (Austria). Förster (1961), HN; see also Anufriev (1975), Anufriev & Emelyanov (1988), Holzinger (1999b), Lauterer (1980)
Edwardsiana nigriloba (Edwards, 1924) So far M VI – M VIII; egg, 2 gen. Monophagous on Acer pseudoplatanus in mountain forests, river floodplains and parks, generally preferring rather cool sites. Widespread in Germany (even found on Heligoland), but only in low frequency, probably overlooked, frequently found up to at least 800 m a.s.l. in the Mittelgebirge, but single specimens at 1700 m a.s.l. in the Bavarian Alps. Not recorded from Baden-Württemberg, Brandenburg and Mecklenburg-Vorpommern, but from France, Switzerland and Poland. In more northern parts of Europe, the species is only known from a few records from southern Sweden and Scotland. Niedringhaus & Olthoff (1993), Remane & Reimer (1989), Schiemenz (1990), Wagner (1935, 1939a), Walter (1998), Nickel (1999b), HN; see also Le Quesne & Payne (1981), Nast (1976a, 1987), Ossiannilsson (1981)
Edwardsiana plebeja (Edwards, 1914) M VI – M X; egg, 2 gen. Monophagous on elms (Ulmus minor, U. laevis, U. glabra); usually swept along forest margins and alleys and on solitary trees. Widespread in the lowlands of Germany, in intermediate frequency on the hosts, but only scattered and with strongholds in river valleys, the highest localities are at only c. 450 m a.s.l. in the German Mittelgebirge, but at 800 m a.s.l. in the Engadine (Switzerland). Fischer (1972), Schiemenz (1990), Wagner (1935, 1939a), Nickel (1999b), HN; see also Günthart (1987a)
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Edwardsiana prunicola (Edwards, 1914) Typhlocyba barbata Ribaut, 1931 E. prunicola (Edw.) was described from Prunus, E. barbata (Rib.) from Salix. Most authors treat them as conspecific due to the apparent lack of morphological differences (e.g. Nast 1972, 1987; Ossiannilsson 1981).
B VI – M X; egg, 2 gen. Usually on hairy-leaved willows (Salix cinerea, S. aurita, S. viminalis, occasionally also S. caprea) in various wetlands, as well as on species of Prunus (mainly P. spinosa and P. domestica), along hedges and forest margins and in fruit-tree plantations, a few records also on ornamental Viburnum lantana. Widespread in Germany, usually in intermediate frequency and individual numbers, but poorly documented (e.g. not recorded from Bavaria to the south of the Danube); highest localities are at 600 m a.s.l. in the Mittelgebirge, at 1250 m a.s.l. in the Swiss Alps. Achtziger (1991, 1994), Frommer (1996), Günthart (1988), Niedringhaus (1991), Reimer (1992), Remane (1987), Ribaut (1936), Schiemenz (1990), SMNS, Wagner (1935, 1939a), HN; see also Günthart (1987b)
Edwardsiana rosae (Linnaeus, 1758) B V – E X, occasionally XI/XII; egg, 2 gen., perhaps only 1 at higher altitudes. On various roses (Rosa canina, R. corymbifera, R. rubiginosa, R. rugosa and others) in moderately shady to sunny, damp to dry sites; in lower numbers and preferentially (perhaps exclusively) in the second generation also on other woody species of Rosaceae (Prunus spinosa, species of Rubus and Sorbus, cultivated cherry, plum, peach and apple, in Wales also recorded from cultivated strawberries). Widespread and common all over Germany, frequently found up to at least 1500 m a.s.l. in the Bavarian Alps. Locally this species is extremely abundant on ornamental roses in gardens and parks and may cause feeding damage. Achtziger (1991), Lehmann (1973a, 1973b), Müller (1956), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz (1990), Schwoerbel (1957), Trümbach (1959), Wagner (1935), HN; see also Claridge & Wilson (1978a), Günthart (1971b)
Edwardsiana rosaesugans (Cerutti, 1939) M VII – E IX; egg, 1(?) gen. On wild roses (mainly Rosa pendulina, a few individuals have been found on R. canina agg.), in open forests, shrubs and pastures from the montane to the subalpine belt. In Germany only known from 8 localities in the Bavarian and Allgäu Alps (Ponten above Hindelang, Schachen above Garmisch-Partenkirchen, Sojernspitze above Mittenwald, Brauneck above Lenggries, as well as Roßfeld, Ahornbüchsenkopf, Jenner and Wimbach valley, all above Berchtesgaden) between 900 and 1800 m a.s.l. Remane & Fröhlich (1994b), Nickel (1999b), HN; see also Cerutti (1939a), Claridge & Wilson (1978b), Lauterer (1983)
Edwardsiana salicicola (Edwards, 1885) M VI – E X; egg, 2 gen., perhaps only 1 at higher altitudes.
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On hairy-leaved willows (Salix cinerea, S. aurita, S. caprea, occasionally also S. viminalis and S. appendiculata) in rather wet or cool sites, e.g. along margins of bogs and fens, in wet meadows, forest glades, and among subalpine scrub. Probably widespread in Germany, although poorly documented. Usually in intermediate to high frequency and rather high individual numbers on the hosts, recorded at least up to 900 m a.s.l. in the Mittelgebirge, up to 1500 m a.s.l. in the Bavarian Alps. Frommer (1996), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Remane & Reimer (1989), Schiemenz (1990), Trümbach (1959), Wagner (1935, 1939a), Nickel (1999b), HN
Edwardsiana spinigera (Edwards, 1924) M VI – M IX; egg, 2 gen. On Corylus avellana; locally syntopic with E. avellanae (Edw.), but generally preferring dryer and warmer sites, reaching the edge of the range in the north German plain. At least scattered and in low frequency in southern and middle parts of Germany, apparently scarce in the north (recorded near Bremen and Lüchow). Not reported from Denmark and Scandinavia. Highest localities are at 850 m a.s.l. in the Mittelgebirge, at 1350 m a.s.l. in the Swiss Alps. Achtziger (1991), Frommer (1996), Niedringhaus & Olthoff (1993), Post-Plangg & Hoffmann (1982), Reimer (1992), Schiemenz (1990), SMNS, Trümbach (1959), Nickel (1997), HN; see also Günthart (1987b)
Edwardsiana smreczynskii Dworakowska, 1971 So far M VI – M IX, in Krakow according to Dworakowska (1971) until M X; egg, 2(?) gen. So far altogether known only from 8 localities: Crakow (locus typicus), Paris (described almost synchronously as E. guentharti Dlabola, 1971!), Caen (Normandy), Warsaw, Bajtava (Slovakia), Basel (Switzerland), Main valley near Schweinfurt (Unterfranken), 220 m a.s.l., 12.IX.1994, 8 ††, on Ulmus spec., and Oberbergen (Kaiserstuhl) near Freiburg, 14.VI.2002, 2 ††, on Ulmus minor. Nymphal development takes place on elm (perhaps Ulmus laevis), but in Crakow a few adults have also been found on other shrubs and trees (Cornus, Crataegus, Betula, Malus, Rosa, Salix). Malenovsky (pers. comm.), Nickel (1999b); see also Chudzicka (1982), Dworakowska (1971), della Giustina (1999), Günthart (1974), Lauterer (1980), Mühlethaler (2001)
Edwardsiana stehliki Lauterer, 1958 M VI – B IX; egg, 2 gen., in Slovakia until M XI (Lauterer 1984). On hazel (Corylus avellana, in Slovakia also C. maxima, but not C. colurna) in damp to dry sites. From Germany there are only two records, both from the Oberlausitz: „Umgebung Bautzen, 1957-61 (häufig)“, as well as Ebersbach near Görlitz, 01.IX.1998, 1 †, on C. avellana. Is reported from Sorbus aria and Viburnum at 600 m a.s.l. in Switzerland. Otherwise only known from Sweden, Poland and Austria. Förster (pers. comm. to Lauterer 1984), HN; see also Günthart (1987b)
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Edwardsiana tersa (Edwards, 1914) E V – E IX; egg, 2 gen. On Salix viminalis, usually in valleys of rivers and streams, perhaps also on other species of willows (S. triandra, S. pentandra). Widespread in the lowlands of Germany, up to at least 650 m a.s.l., often in high frequency and individual numbers on the host, but not recorded to the south of the Danube. In Austria the species is only reported from a single locality in Styria (Judenburg, 680 m a.s.l., on Salix daphnoides), in France southward to Paris; recently found in Basel (Switzerland). Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1990), Schwoerbel (1957), Wagner (1935), HN; see also Holzinger (1995a), della Giustina (1989)
Eupterycyba jucunda (Herrich-Schäffer, 1837) B VII – E X; egg, 1 gen. Monophagous on Alnus glutinosa, mainly near standing or running water, also in forests of moist to wet sites and mining areas. Widespread in the lowlands of Germany, in intermediate to high frequency, but usually low individual numbers, recorded only up to 600 m a.s.l. in the Mittelgebirge and the foothills of the Alps. Achtziger (1991), Fischer (1972), Frommer (pers. comm.), Heller (1987a), Niedringhaus (1991, 1997), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1990), Wagner (1935, 1939a, 1951a), HN
Linnavuoriana decempunctata (Fallén, 1806) Adults I – XII, mainly B VIII – VI; adult, 1 gen. On birches (Betula pendula and B. pubescens), usually in cool sites, e.g. river basins (also in urban parks), fens and bogs; hibernating individuals were found on Picea abies. In Germany probably much under-recorded due to late emergence, but apparently widespread and usually in low to intermediate frequency, up to at least 1000 m a.s.l. Frommer (1996), Niedringhaus & Olthoff (1993), Schiemenz (1990), Schönitzer & Oesterling (1998b), SMNS, Wagner (1935), HN
Linnavuoriana intercedens (Linnavuori, 1949) Has been treated as a subspecies of L. decempunctata (Fall.), but is likely to be specifically distinct due to constant differences in the morphology of the aedeagus and different host plant (Lauterer & Novotný 1991).
So far B VII, in Moravia at least until M X (Lauterer & Novotný 1991); ad.(?), 1 gen. So far only known from Norway, Sweden, Finland, northern Russia, Latvia and Czechia, but is perhaps more widespread, and published records of L. decempunctata (Fall.) from Alnus in Poland, Switzerland and Italy should be revised. There is a single recent record from southeastern parts of Germany: Langensendelbach near Forchheim, 340 m a.s.l., 09.VII.2001, 2 ††, 1 ‡ (freshly emerged), on Alnus incana in a forest clearing. Is apparently monophagous on A. incana. The altitudes of Bohemian localities range from
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200 to 730 m a.s.l. Specimens reported from the Engadine and the Valais (both Switzerland) at altitudes between 1500 and 1700m a.s.l. may also belong to this species. HN; see also Ossiannilsson (1981), Lauterer & Novotný (1991), Günthart (1987a)
Linnavuoriana sexmaculata (Hardy, 1950) E VII – X, single ‡‡ frequently until M VI; mainly egg(?), 1 gen. On hairy-leaved species of willows in various wetland habitats (along rivers and streams, in bog areas, near lakes and ponds, in mining areas, etc.) as well as in forest glades; usually in high frequency on Salix viminalis, less frequently on S. cinerea, S. caprea, S. aurita, as well as S. alba, in the Alps and their foothills also on S. eleagnos. Has been taken from Pinus and Picea in winter. Widespread and fairly common in Germany, although usually found in low numbers only; the highest localities are at 700 m a.s.l. in the Mittelgebirge, at 1000 m a.s.l. in the Bavarian Alps, at 1520 m a.s.l. in the Engadine (Switzerland). Frommer (1996), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1939a), Nickel (1999b), HN; see also Günthart (1987a)
Lindbergina aurovittata (Douglas, 1875) According to Le Quesne (1977) this taxon shows pronounced intraspecific variation in morphology of the aedeagus. Accordingly, he considers Lindbergina pandellei (Lethierry, 1878) to be a junior synonym of L. aurovittata (Dgl.). Several authors, however, have treated Typhlocyba pandellei spoliata Horváth, 1907, originally described only as a variety of the latter, as a distinct species (e.g. Drosopoulos et al. 1986; D’Urso 1995). Compared to L. aurovittata (Dgl.), differences in life history are apparently weak, if present at all (Vidano & Arzone 1987b; Vidano et al. 1990; Bosco & Arzone 1991).
E V – E X; egg, 2 gen. The range of this taxon includes the Mediterranean region and parts of western Europe. Since the beginning of the 1990s it has been found in 6 localities in the extreme west of Germany: middle Rhine valley near Cologne, upper Rhine valley near Bruchsal, Nahe valley near Schloßböckelheim, Neckar valley near Heilbronn (large populations on cultivated Rubus fruticosus and perhaps noxious), as well as Palatinate, Haardtrand near Wachenheim and Annweiler (on Quercus petraea), all sites between 50 and 200 m a.s.l.; also reported from the French side of the southern upper Rhine plain near Colmar. From southern and western Europe a remarkable host shift has been described: in Wales, overwintering eggs are laid in leaves of evergreen Rubus (presumably mainly R. fruticosus), adults of the spring generation migrate to other woody species for oviposition (mainly Quercus robur and Fagus sylvatica, also Alnus glutinosa, Corylus avellana, Betula pubescens, Quercus cerris, Qu. petraea); the summer generation emerging from these eggs return to Rubus for laying the winter eggs. In Piedmont (Italy), however, winter eggs are laid on evergreen oaks (Quercus ilex and Qu. suber); the spring generation migrates to deciduous species (Quercus pubescens, Qu. cerris) and other trees. Schrameyer (pers. comm.), Frommer (pers. comm.), Fröhlich & Nickel (unpublished data), HN; see also Claridge & Wilson (1978a), della Giustina & Balasse (1999), Vidano & Arzone (1987b), Vidano et al. (1990)
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Ribautiana alces (Ribaut, 1931) So far M VI – B IX, unidentified ‡‡ of either this or the following species until B X; egg, 2(?) gen. On oak (Quercus robur, perhaps also Qu. petraea) in rather warm sites, preferentially in upper slope or plateau situations and along sunny forest margins. In Germany apparently at the northeastern edge of the range, only recorded in middle parts (Donnersberg/Palatinate, Mainzer Sand, middle Rhine valley near Bacharach, Ahr valley to the south of Bonn, middle Hessen, surrounding plates of the Thuringian Basin near Artern and Arnstadt), up 450 m a.s.l., but up to 1300 m a.s.l. in the Valais (Switzerland). Not recorded from Denmark, Scandinavia and Poland. Remane (1987, and pers. comm.), Wagner (1939a), HN; see also Cerutti (1939a), Nast (1987)
Ribautiana scalaris (Ribaut, 1931) So far mainly M VII – M IX, single records B VI and B X; egg, 2(?) gen. Like the preceding species, on oak in rather warm sites, but preferring Quercus petraea instead of Qu. robur; adults being usually swept along sun-exposed forest margins, occasionally in alleys or groups of trees. Widespread at least in middle and southern parts of Germany, but sporadic, in low frequency and low individual numbers; not recorded in the area to the south of the Danube. Northernmost localities are near Herne, Hamburg, Göttingen and Eberswalde, but the species has been found in southern Scandinavia and Poland. The highest German sites are at 400 m a.s.l. only, but at 1300 m a.s.l. in the Valais (Switzerland). Köhler (pers. comm.), Niedringhaus & Olthoff (1993), Schwoerbel (1957), Trümbach (1959), Wagner (1939a), Wagner & Franz (1961), Nickel (1997), HN, see also Cerutti (1939a), Chudzicka (1982), Claridge & Wilson (1981), Ossiannilsson (1981)
Ribautiana debilis (Douglas,1876) E V – B XII; egg (presumably also single ‡‡), 2 gen. The first German report of this species dates only from 1994. Shortly afterwards it was found in numerous sites in the valleys of the upper and middle Rhine, the Nahe, Neckar and Main, as well as in middle Hessen. Most specimens were sampled on Rubus fruticosus and R. caesius, along hedges and forest margins and in the undergrowth of parks and open forests. The northeastern edge of the range follows a line from Cologne to Marburg, Würzburg, Heilbronn, Stuttgart and Freiburg; highest localities are at 300 m a.s.l. only. In France and Britain the species is reported from various woody plants, preferentially members of the Rosaceae (Malus, Pyrus, Prunus, Cydonia, Rosa, Rubus), Quercus, Alnus, also Corylus, Betula, Tilia, Ulmus and others. Frommer (1996), Heller (1994), Remane (1995), Schrameyer (pers. comm.), HN; see also Le Quesne & Payne (1981), Ribaut (1936)
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Ribautiana tenerrima (Herrich-Schäffer, 1834) E V – E X, occasionally until III; egg, 2 gen. On Rubus fruticosus, R. idaeus and R. caesius in semi-open cultivated areas (forest margins and clearings, gardens, hedges, etc.), in low numbers also on other woody plants, mainly Quercus, Salix, Acer, Prunus and Corylus. In Italy and Wales, breeding is reported to take place on Rubus and Corylus only. Widespread and common in the lowlands of Germany, but probably under-recorded; only single specimens found to the south of the Danube and above c. 600 m a.s.l. (in the Mittelgebirge), but reported from 900 m a.s.l. in the Valais (Switzerland). Achtziger (1991), Frommer (1996), Heller (1994), Niedringhaus (1991), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schiemenz (1990), Wagner (1935, 1939a), HN; see also Arzone & Vidano (1987a), Claridge & Wilson (1981), Raine (1960)
Ribautiana ognevi (Zachvatkin, 1948) So far E V – M IX; egg, 2 gen. This species’ range essentially includes the eastern part of the European deciduous forest zone, extending westward to Bohemia (Czech Republic) and eastern parts of Austria and Germany, where it is currently known from 5 localities up to 200 m a.s.l. only: Main floodplains near Schweinfurt, Oder valley near Wriezen and Angermünde, and Elbe floodplains near Gorleben and Burg. Lives on Ulmus laevis usually in river floodplains, apparently in intermediate frequency and low to intermediate individual numbers. Is perhaps much under-recorded, but has not been found in host stands in the valleys of the Rhine and Leine. Nickel (1999b), HN; see also Dlabola (1958, 1977), Emelyanov (1964a) (see this paper also for identification)
Ribautiana ulmi (Linnaeus, 1758) E V – B XI; egg, 2 gen. On elms – in the lowlands usually Ulmus minor and U. laevis, in more mountainous regions mainly U glabra – in forests and open stands as well as on solitary trees. Widespread and fairly common in Germany, in high frequency and individual numbers on the hosts, occurring up to at least 1000 m a.s.l. in the Bavarian Alps, up to at least 1350 m a.s.l. in the Engadine (Switzerland). Achtziger (1991), Bittner & Remane (1977), Fischer (1972), Reimer (1992), Remane (1987), Schiemenz (1990), Wagner (1935, 1951a), HN; see also Günthart (1987a)
Typhlocyba quercus (Fabricius, 1777) Mainly M VI – M IX, occasionally until IV; egg, 1(?) gen. In semi-open cultivated areas, mainly along forest margins, in river floodplains, fruittree plantations and along hedges, usually on species of Prunus and Quercus (P. padus, P. avium, P. spinosa, P. domestica, Qu. robur, Qu. petraea); in low numbers also on other woody species (Corylus, Carpinus, Fagus and others); in Wales breeding has been found to take
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place on these (as well as on Castanea and ornamental Nothofagus). Widespread in Germany, but often in intermediate frequency and low individual numbers only. Highest localities are at 900 m a.s.l. in the Mittelgebirge and Bavarian Alps, at 1240 m a.s.l. in the Engadine (Switzerland). Achtziger (1991), Fischer (1972), Lehmann (1973a), Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992), Remane (1987), Schiemenz (1990), Schwoerbel (1957), Trümbach (1959), Wagner (1935, 1939a), HN; see also Claridge & Wilson (1976, 1981), Günthart (1987a)
Zonocyba bifasciata (Boheman, 1851) E VI – M X; egg, 1(?) gen. On Carpinus betulus, Ulmus minor and U. glabra, in closed forests as well as in semi-open tree stands, occasionally even along hedges. Single specimens found on Acer, Tilia, Alnus, and other deciduous trees, but probably without breeding. Widespread in the lowlands of Germany, but often in low individual numbers only, usually in high frequency on Carpinus, but less frequently on Ulmus; highest localities are at 600 m a.s.l. Frommer (1996), Günthart (1988), Niedringhaus (1997), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), HN, and others; see also Claridge & Wilson (1981)
Eurhadina concinna (Germar, 1831) B VI – B X; egg, 1 gen. Preferentially on Quercus robur and Qu. petraea in forests and semi-open stands, in lower numbers on Fagus, Betula, Alnus (in Wales breeding was found to take place on these), Tilia and others. Widespread in the German lowlands, usually in intermediate to high frequency on both oak species, but recorded only up to 700 m a.s.l. in the Mittelgebirge and the foothills of the Alps. Achtziger (1991), Frommer (1996), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), HN; see also Claridge & Wilson (1976, 1981)
Eurhadina kirschbaumi W. Wagner, 1937 So far M VII – B IX; egg, 1 gen. On Quercus petraea; adults are usually swept along forests roads and margins, mainly in acidic, occasionally also basic, usually damp to moderately dry sites. In Germany rather sporadic and in low frequency, but perhaps overlooked. Recorded from the surroundings of Hamburg (locus typicus), Berlin, Göttingen, Artern, Ahrweiler, Koblenz, Mainz, Saarbrücken, Aschaffenburg, Würzburg, Schweinfurt, Arnstadt, Niesky and Ludwigsburg. Highest localities are at 450 m a.s.l. only. Is also reported to live on Quercus pubescens, Qu. cerris and Castanea sativa in the Mediterranean region; these potential host species have not yet been sampled in Germany, however. Post-Plangg & Hoffmann (1982), Remane (pers. comm.), SMNS, Wagner (1937a, 1939a, 1951a), ZMB, Nickel (1994), HN; see also Drosopoulos (1999), Vidano & Arzone (1987b)
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Eurhadina loewii (Then, 1886) Eurhadina untica Dlabola, 1967 After Dlabola (1967a) north and central European populations with unforked lateral appendages of the aedeagus are conspecific with Eurhadina untica Dlabola, 1967 described from Ulmus pumila in Mongolia. The author also suggested that individuals with forked appendages mainly reported from the southern half of Europe belong to a distinct species (E. loewii s.str.). This opinion is rejected here. According to Lauterer (pers. comm.) the type of E. loewii (Then) shows an anomalous spinulation of the aedeagus branches.
M VI – E IX; egg, 1 gen. On Acer pseudoplatanus, often in shady or cool situations (preferentially on slopes and in river floodplains, but also in parks and gardens). Widespread in Germany, in low to moderate individual numbers and in intermediate frequency on the host, found to at least 900 m a.s.l. in the Bavarian Alps, up to 1290 m a.s.l. in the Engadine (Switzerland). Bittner & Remane (1977), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer (1992), Schiemenz (1990), SMNS, Wagner (1935, 1939a), Nickel & Remane (1996), HN; see also Günthart (1987a, 1987b)
Eurhadina pulchella (Fallén, 1806) B VI – B X; egg, 1 gen. On Quercus robur and Qu. petraea, usually in forests and semi-open stands, occasionally on solitary trees; in low numbers also on Betula, Fagus, Carpinus and others, but probably without breeding. Widespread in the lowlands of Germany, usually in high frequency and intermediate individual numbers on both host species, found up to at least 750 m a.s.l. in the Allgäu, but reported from 1300 m a.s.l. in the Valais (Switzerland). Achtziger (1991), Fischer (1972), Frommer (1996), Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992), Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), HN; see also Cerutti (1939a), Claridge & Wilson (1981), Vidano & Arzone (1987b)
Eurhadina ribauti W. Wagner, 1935 E V – M X; egg, 2(?) gen. Like the preceding species, mainly on Quercus robur and Qu. petraea, occasionally Fagus, Carpinus, Tilia, Alnus and others, but breeding apparently only on Quercus. Widespread in Germany, but apparently confined to lowland regions (recorded up to 450 m a.s.l. only), usually in low to intermediate frequency and individual numbers. Not recorded to the south of the Danube, except from a single locality in the Lake Constance Basin. Niedringhaus & Olthoff (1993), Remane (1987), Schiemenz (1990), Wagner (1935, 1939a), Nickel & Remane (1996), HN; also Claridge & Wilson (1981)
Eurhadina saageri W. Wagner, 1937 E VI – B X; egg, 1 gen. On oak, mainly (perhaps exclusively) on Quercus robur. Adults are usually swept along forest margins and in semi-open host stands in lowland areas and river basins. The few
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host records from other European countries also refer exclusively to Qu. robur. In Germany only very sporadic, in low frequency and individual numbers, at the northern edge of the range. Border localities are near Wilhelmshaven, Hamburg (locus typicus), Lübeck, Wolfsburg, Gera and Kamenz, up to at most 550 m a.s.l. Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Schiemenz (1990), Wagner (1937a, 1951a), HN; also Anufriev & Kirillova (1998), Lauterer (pers. comm.)
Eupteryx atropunctata (Goeze, 1778) M V – E X; egg, at least 2 gen. Polyphagous in rather tall herb stands in wet to dry, usually sunny sites (low-input grassland, waysides, ruderal habitats, abandoned fields, fens and moderately saline sites, also synanthropic in gardens and fields). Host plants are mainly Lamiaceae (Mentha, Salvia, Clinopodium, Origanum, Ballota, Melissa and others), also Verbascum thapsus, Thalictrum flavum, Solanum tuberosum, Pastinaca, presumably also Cirsium and Urtica dioica. Furthermore, Lavatera, Althaea, Nepeta, Lamium, Beta vulgaris and Vicia faba are mentioned in the literature. Widespread and common in Germany, found up to at least 900 m a.s.l. in the Mittelgebirge and the Bavarian Alps. Is reported from 1500 m a.s.l. in Switzerland and 1700 m a.s.l. in Austria, but without information on breeding at these altitudes. Fischer (1972), Fröhlich (1996a), Niedringhaus (1991, 1997), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), Wais (1990), Nickel & Achtziger (1999), HN, and others; see also Günthart (1987a), Stewart (1988), Wagner & Franz (1961)
Eupteryx aurata (Linnaeus, 1758) M V – E X; egg, 2 gen. Like E. atropunctata (Goeze) in tall herb stands, but usually in cool or moderately shady, moist to temporarily flooded sites, often near water, along forest roads and among subalpine scrub, in lower numbers in gardens and ruderal sites. Host plants are tall herbs, in the first generation Urtica dioica, in the second also various species of Asteraceae (mainly Petasites, Arctium, Senecio, Cirsium, Dahlia), Lamiaceae (mainly Lamium, Mentha), Apiaceae (Heracleum sphondylium, also H. mantegazzianum), Malvaceae (Althaea rosea) and others. Literature records include Humulus lupulus and Solanum tuberosum, as well as additional species of the plant families mentioned above. Widespread and very common all over Germany, also in higher parts of the Mittelgebirge, frequently found up to 1700 m a.s.l. in the Bavarian and Allgäu Alps. Fischer (1972), Niedringhaus (1991, 1997), Reimer (1992), Remane (1987), Schiemenz (1990), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN; see also Stewart (1988)
Eupteryx austriaca (Metcalf, 1968) Published data on life history and distribution appear to be contradictory, perhaps due to taxonomic problems. The status of north German lowland populations, in particular, is in need of revision.
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M VI – B X; egg, 1(?) gen. Mainly on Knautia dipsacifolia in damp to moist, moderately shady sites of the montane and subalpine belt, usually along sunny forest margins and roads, as well as in subalpine scrub. In Germany largely confined to the Alps and the southern Black Forest, between 750 and 1850 m a.s.l., in intermediate frequency on the host. On the other hand, there are older records from Succisa pratensis in wetland habitats of northern Germany (Albersdorf/Holstein, 24.VII.1933, and Duvenstedter Brook near Hamburg, 22.VIII.1937). Fischer (1972), Wagner (1935, 1941b), HN
Eupteryx heydenii (Kirschbaum, 1868) According to Nast (1987), this species has been described after two syntypes, one † from Switzerland and one ‡ from Germany. However, according to Fieber (1872a, 1872b), who studied the type material, the ‡ (collected near Bad Rippoldsau, Black Forest) belongs to the preceding species.
So far E VI – E IX; egg, 1(?) gen. In tall herb stands in the submontane to the subalpine belt, usually along forest margins and roads, along streams and ditches, and in montane pastures. The host plant is Chaerophyllum hirsutum, perhaps also Ch. aureum. Furthermore, Petasites has been reported from the Austrian and Swiss Alps, but these feeding records are probably erroneous. There are only few published data from Germany, although the species is widespread and rather common at least in the Alps and their foothills; there is also a number of localities in the Mittelgebirge (Harz Mountains, Rhön Mountains, Thuringian Forest, Frankenwald, Vogtland, Erzgebirge, Franconian Jura and Oberpfälzer Wald, probably also in other hilly regions). According to existing data, these are at the northern edge of the range. Found between 400 and 1850 m a.s.l. in Germany, up to 2000 m a.s.l. in Styria (Austria). Fischer (1972), Reimer (1992), Schiemenz (1990), Schönitzer & Oesterling (1998b), Walter (pers. comm.), HN; see also Cerutti (1939a), Günthart (1974), Wagner & Franz (1961)
Eupteryx lelievrei (Lethierry, 1874) B VI – E X; egg, 2 gen. On Betonica officinalis in temporarily wet to temporarily dry, usually moderately shady sites, mainly along forest margins, near shrubs, and in low-input meadows, preferentially on clayey soils. In Germany only very sporadic, in low frequency and low densities; at present there are only 10 known localities: southern Niedersachsen (near Göttingen), Thuringian Basin (near Erfurt), middle Hessen (Wermertshausen), Taunus (Niederreifenberg), Franconia (Sulzheim and Gesees), Kaiserstuhl (Vogtsburg) and Bavarian Alps (Berchtesgaden, Ramsau, and Nußdorf am Inn, c. 1300 m a.s.l.). Otherwise only known from a few records from Belgium, France, Switzerland, Austria, the Czech Republic, Slovakia, Bulgaria and middle Russia. Fechtler (pers. comm.), Remane & Fröhlich (1994b), Nickel & Voith (unpublished data), Nickel (1994), HN; see also Lauterer (1983, 1984), Nast (1987)
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Eupteryx origani Zachvatkin, 1948 B VI – B X; egg, 2 gen., probably 1 at higher altitudes. On Origanum vulgare, in oligotrophic, moderately dry to damp, moderately shady sites; mainly along herbaceous margins in dry grassland, montane meadows and pastures, preferably near trees or shrubs. In Germany only very sporadic and known from scattered localities in the Bavarian Alps (up to at least 1600 m a.s.l.), the Franconian and Swabian Jura, the Frankenwald, the Thuringian Forest, the middle Rhine and its tributaries (basins of Ahr, Lahn and Dill) and the north German plain (near Hamburg and Berlin). To the west of this, there are records from Scotland, England, the Netherlands, Switzerland and south-eastern France (Savoie). Has been reported from Veronica chamaedrys, but these data require confirmation. Remane (1961b), Remane & Fröhlich (1994b), Post-Plangg & Hoffmann (1982), Schiemenz (1990), SMNS, HN; see also della Giustina & Remane (2001), Günthart (1997), Le Quesne & Payne (1981), Nast (1987)
Eupteryx signatipennis (Boheman, 1847) E VI – B X; egg, 2 gen. On Filipendula ulmaria, usually in wet and cool sites (in fens, along shores and ditches, in pastures and abandoned meadows, apparently absent from habitats mown more than once a year). From Germany there are only very few published records, but the species has been found in most regions except the northeast (Mecklenburg-Vorpommern, Brandenburg, Saxony-Anhalt), where it is probably overlooked. The highest localities are at 1200 m a.s.l. in the Allgäu Alps and the southern Black Forest. Bittner & Remane (1977), Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer (1992), Remane (1962), Remane & Fröhlich (1994b), Rombach (pers. comm.), Schiemenz (1990), Nickel & Achtziger (1999), Nickel & Remane (1996), HN
Eupteryx adspersa (Herrich-Schäffer, 1838) So far M VI – E IX, in Poland according to Dworakowska (1970a) V – X; egg, 2 gen. Along dry waysides and in ruderal sites on various substrates (mainly sand, limestone and gypsum), on Artemisia absinthium. The German localities are at the northwestern edge of the range; there are only c. 15 records along the middle Rhine and its tributaries, since 1890 (St. Goarshausen, Bacharach, Limburg an der Lahn), in the rain shadow area to the east of the Harz Mountains (near Eisleben, Freyburg an der Unstrut, Artern), near Lüchow, Berlin, Göttingen and Meißen. The occurrence at the locus typicus near Regensburg (Bavaria) is not confirmed due to formerly diverging interpretations of the species’ identity, and because Herrich-Schäffer’s collection has been lost during the last war, although the most closely related species, E. artemisiae (Kbm.) is not known from southern Germany to the south of the Main river (see below). All German localities of E. adspersa (H.-S.) are below 250 m a.s.l., but the species is reported from up to 1650 m a.s.l. in the Engadine and Valais (both Switzerland). Dworakowska (1970a; 1972a), Herrich-Schäffer (1838), Remane (pers. comm.), Schiemenz (1990), Walter & Emmrich (1995), Nickel (1997), HN; see also Cerutti (1939a), Günthart (1987a)
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Eupteryx artemisiae (Kirschbaum, 1868) E V – M X; egg, 2 gen. In salt marshes along the coasts of the North and Baltic Sea, also locally inland in saline sites and old gardens. Reports from dry grassland habitats probably refer to the preceding species. Usually on Artemisia maritima, but on A. abrotanum in gardens. Known in Germany from the East and North Frisian Islands (where locally abundant), the Jade Bay, the coasts of Schleswig-Holstein, and the island of Poel; further populations occur in the rain shadow area to the east of the Harz Mountains (Solgraben Artern), in Hamburg, Göttingen and Marburg (botanical gardens), as well as in Frankfurt am Main (locus typicus, no habitat given). All localities are below 200 m a.s.l. Niedringhaus & Olthoff (1993), Remane (pers. comm.), Schiemenz (1990), Wagner (1935, and unpublished data), Nickel (1997), HN; for identification of nymphs see Stewart (1986a)
Eupteryx calcarata Ossiannilsson, 1936 B V – E X; egg, 2 gen. On Urtica dioica, in sunny, damp to moist sites; generally preferring more sunny and dry habitats than the other Urtica-dwelling congeners, e.g. along ditches and streams in open fields, in ruderal sites and mining areas, occasionally also in pastures. Widespread in the lowlands of Germany, but only localized, rarely found above 400 m a.s.l., a single record is from c. 750 m a.s.l. in the Allgäu. Fischer (1972), Heller (1987a), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1962), Schiemenz (1990), Schwoerbel (1957), Wagner (1939a, 1951a), Nickel (1994), HN
Eupteryx cyclops Matsumura, 1906 E V – E X; egg, 2 gen. On Urtica dioica, mainly near standing or running water (also along ditches within fields), herbaceous and woody margins, and in open forests. In comparison to other species of Eupteryx on this host, with preference for moderately sunny, moist and higher-altitude sites. Widespread and common all over Germany, frequently found up to at least 1000 m a.s.l. in the Mittelgebirge, the Bavarian and Allgäu Alps (single individuals up to 1600 m a.s.l.), up to 2000 m a.s.l. in the Valais (Switzerland). Achtziger (1991), Fischer (1972), Heller (1987a), Mölleken & Topp (1997), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz (1990), HN, and others; see also Cerutti (1939a), Günthart (1987a), Stewart (1988), Stiling (1980a); for identification of nymphs see Stewart (1986b)
Eupteryx immaculatifrons (Kirschbaum, 1868) M VI – E X; egg, 2 gen. Formerly, this species was often overlooked or misinterpreted. It lives on Lamium maculatum in moist to damp, usually shady sites, mainly in floodplain forests along rivers and streams as well as on seeping slopes. In Germany recorded only from middle and southern parts (locus typicus: Mombach near Mainz), apparently at the northern edge of
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the range, but it may be more widespread, together with its host. Border localities are near Bonn, Marburg, Göttingen, Bad Frankenhausen, Dresden and Görlitz. Has been found at 700 m a.s.l. in the Bavarian foothills of the Alps, but to the north of the Danube it is usually confined to locations below 500 m a.s.l. Büchs (1988), Emmrich (1975), Wagner (1939a), Remane & Fröhlich (1994b), Nickel (1994, 1999b), HN
Eupteryx urticae (Fabricius, 1803) E V – E X; egg, 2 gen. On Urtica dioica in damp to moderately wet, usually moderately shady to shady sites, preferentially in various types of forests (often along roads and margins) and along shores of standing or running water (also ditches). In Wales and England individuals of the second generation may occur in numbers on Urtica urens and Parietaria judaica, although overwintering eggs are found on U. dioica only. Widespread and common in Germany, found up to at least 900 m a.s.l. in the Bavarian Alps, up to 1500 m a.s.l. in Switzerland. Achtziger (1991), Mölleken & Topp (1997), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), HN, and others; see also Günthart (1987a), Stewart (1988, and pers. comm.), Stiling (1980a); for identification of nymphs see Stewart (1986b)
Eupteryx collina (Flor, 1861) This species has often been misinterpreted; thus, many older reports refer to other species. After a revision of Flor’s collection Vilbaste (1973) concluded that Eupteryx alticola Ribaut, 1936 is a junior synonym of E. collina (Fl.), and that E. florida Rib., which had originally been described only as a variety, is a distinct species. However, there is some evidence that records published even recently as E. collina (Fl.) may refer to a complex of several species.
So far M VI – E IX; egg, 1 gen., probably 2 in lower altitudes. On Mentha longifolia along streams and ditches, as well as in seepy pastures. Is also reported from Acinos alpinus in the Valais (Switzerland) and from various species of Lamiaceae in Italian gardens. In Germany the species is largely confined to the Bavarian and Allgäu Alps and their foothills, mainly between 700 and 1700 m a.s.l. A few isolated localities are near Augsburg (c. 500 m a.s.l.) and in the vicinity of Berlin (c. 60 m a.s.l.). Most if not all other lowland reports refer to E. florida Rib. or ‡‡ which are difficult to identify. Fischer (1972), Remane (1961b), Schiemenz (1990), ZMB, HN; see also Arzone (pers. comm.), Günthart (1987a), Vidano & Arzone (1978)
Eupteryx stachydearum (Hardy, 1850) sensu Dworakowska (1972a) This taxon and the following have long been considered to be conspecific, until Dworakowska (1972a) pointed out that there are differences in head colouration and particularly in the shape of the aedeagus (see also Vilbaste 1973). This opinion has been followed by some authors, e.g. della Giustina (1989) and Remane & Fröhlich (1994a), but not by others, e.g. Nast (1987) and Ossiannilsson (1981). Until further studies have been done, both shall be treated here as different species, although differences are only slight and transitional stages may occur. Apparently,
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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 northern edge of its range, whereas E. stachydearum (Hardy) is also reported from Denmark and southern 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 Ossiannilsson (1981); for identification see Dworakowska (1972a), della Giustina (1989)
Eupteryx curtisii (Flor, 1861) See taxonomic note on the preceding species.
E V – M X; egg, 2 gen. Generally more in warmer and less shady sites than E. stachydearum (Hardy) and apparently not syntopic. Preferentially on Teucrium scorodonia in open oak and pine forests and along their margins, in acidic, usually moderately dry sites, perhaps also on Stachys sylvatica and Clinopodium vulgare as well as on other species of Lamiaceae. The distribution in Germany is poorly known. There are confirmed records from the area to the north of a line from Lörrach to Singen am Hohentwiel and Passau, and to the south of a line from Düsseldorf to Lengerich, Hannoversch Münden and Herzberg am Harz, usually below 600 m a.s.l., but also from sun-exposed slopes up to 1100 m a.s.l. in the southern Black Forest. Büchs (1988), Kolbe & Bruns (pers. comm.), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Nickel & Sander (1996), Nickel (1997), HN
Eupteryx florida Ribaut, 1952 Mainly M V – E X, occasionnally until V; egg, in low numbers also adult, at least 2 gen. In moist to moderately dry, usually moderately shady sites, often on Ballota nigra along nitrophilous herbaceous and woody margins, on Stachys sylvatica in open forests and along their margins, as well as in gardens on Salvia officinalis and Melissa officinalis; further populations have been found on Clinopodium vulgare, Lamium album, Mentha longifolia, Stachys palustris and Glechoma hederacea. In Germany widespread and fairly common between the Danube and the north edge of the Alps (up to at least 600 m a.s.l., although single specimens were found at 1300 m a.s.l.), but uncommon in the north German plain, where increasingly synanthropic and at the edge of the range. Border localities are near Lingen (Ems), Bremen, Kiel, Rostock and Eberswalde.
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Frommer (1996), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987, and pers. comm.), Schiemenz (1990), Nickel & Remane (1996), HN; see also Stewart (1988); for identification of nymphs see Stewart (1986a)
Eupteryx salviae Arzone & Vidano, 1994 So far only VIII; egg, 2(?) gen. So far only known from northeastern Italy and the former Yugoslawia (Serbia, Croatia, Montenegro), where reported from various species of Lamiaceae (Salvia officinalis, Teucrium polium, Thymus pulegioides, Th. praecox). Recently, a population was found on Salvia officinalis in a garden in Stuttgart. Remane (pers. comm.), Nickel & Remane (2002); see also Arzone & Vidano (1994)
Eupteryx decemnotata Rey, 1891 M V – E X; egg, 2 gen. Until recently this species was only known from the Mediterranean region (southern France, Italy) and Switzerland. Apparently, there is a dramatic recent range expansion. The first German record is from Malaise traps from near Speyer in 1989; since then it has been found in more than 30 sites in gardens, urban green and abandoned vineyards in the upper Rhine plain (Weil am Rhein, Annweiler, Neustadt an der Weinstraße, Wachenheim, Lorsch, Wiesbaden), in the basins of the Neckar, Nahe, Main and Regnitz (Stuttgart, Schloßböckelheim, Würzburg, Bamberg, Forchheim), in Cologne, Münster, Marburg, Oldenburg, Hildesheim, Göttingen, Erfurt and Berlin. Host plants are various species of ornamental and officinal Lamiaceae (mainly Salvia officinalis, also Melissa officinalis, Nepeta cataria and Thymus); all localities are below 300 m a.s.l. Recently it was also recorded from the French and Swiss side of the southern upper Rhine plain near Colmar and urban areas of Basel, from Vienna, Graz (both eastern Austria) and Nova Gorica (western Slowenia). Fechtler (pers. comm.), Frommer (pers. comm.), Remane (1995, and pers. comm.), Remane & Fröhlich (1994b), Nickel & Remane (1996), HN; see also della Giustina & Balasse (1999), Günthart (1987b), Mühlethaler (2001), Nast (1987), Holzinger & Seljak (2001), Vidano & Arzone (1978)
Eupteryx melissae Curtis, 1837 Mainly E V – B X, some adults and nymphs until V (see also Stewart 1988); (mainly?) egg, 2(?) gen. This species occurs mainly in southern and western parts of Europe. In Germany it appears to prefer synanthropic sites and is known only from a few locations: Elend (Harz Mountains), near Eisleben (recorded in 1910) and Quedlinburg (on Mentha and Marrubium vulgare), Göttingen (botanical gardens as well as private gardens and parks), also found in urban areas and gardens of Weil am Rhein, Stuttgart, Cologne, Münster, Oldenburg, Marburg, Hildesheim, Erfurt and Berlin. Most recent records were from ornamental Nepeta cataria and Salvia officinalis, a few also from Leonurus cardiaca and Althaea officinalis, the latter in the second generation only. All localities are situated be-
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low 250 m a.s.l. except the first (a single specimen at c. 500 m a.s.l.). There is some evidence 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 officinalis have been reported as additional hosts. Fechtler (pers. comm.), Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Remane (pers. comm.), Schiemenz (1990), Nickel (1994), HN; see also Stewart (1988); for identification of nymphs see Stewart (1986a)
Eupteryx thoulessi Edwards, 1926 So far E VI – B X, according to Wagner (1941b) rarely until B XI; egg, 2 gen. In sunny, wet to temporarily flooded sites, mainly tall-sedge swamps and reeds, locally also in sand pits and moderately saline sites. Host plants are Mentha aquatica and occasionally Lycopus europaeus; the latter probably only in the second generation. Furthermore, M. spicata and M. arvensis have been reported. In Germany the species occurs rather locally, with strongholds in the northern half. Most records are from the north German plain and southern Niedersachsen (up to at most 300 m a.s.l.); localities in the Thuringian Basin (near Erfurt) and the southern upper Rhine plain (near Freiburg) appear to be rather isolated. Kuntze (1937), Niedringhaus & Olthoff (1993), Schiemenz (1990), Wagner (1941b), Nickel (1997), HN; see also Ribaut (1936), Stewart (1988); for identification of nymphs see Stewart (1986a)
Eupteryx tenella (Fallén, 1806) B V – E X; egg, 2 gen. On Achillea millefolium along herbaceous margins in moderately shady, damp to dry, basic to acidic sites, mainly in dry grassland, along hedges and forest edges, occasionally also in old gardens; apparently absent from mown meadows. Widespread in the middle parts of Germany, but rather uncommon, most populations being confined to very small sites. Not recorded to the south of the Danube; very few localities are in the north German plain (near Hamburg, Bad Schwartau, Rathenow and Seelow). Found up to c. 700 m a.s.l. in the Mittelgebirge. Achtziger (1991), Frommer (pers. comm.), Nikusch (1976), Reimer (1992), Remane (1987), Schiemenz (1990), Wagner (1939a), HN; for identification of nymphs see Stewart (1986a)
Eupteryx vittata (Linnaeus, 1758) M V – E X; egg, 2 gen. On various dicotyledons in sunny to shady, wet to moderately moist sites, mainly lowinput meadows and deciduous forests. Favoured host plants are Ranunculus repens and Glechoma hederacea, but the species also lives on Ajuga reptans, Valeriana dioica and others. Furthermore, from Britain it is reported to live on Veronica montana, Prunella vulgaris, Plantago major and Crepis paludosa, although some host species may be utilized in the second generation only. Widespread in Germany and fairly common, extending to at least 1350 m a.s.l. in the Mittelgebirge and the Alps.
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Achtziger (1991), Bittner & Remane (1977), Fischer (1972), Haas (1975), Niedringhaus (1997), Reimer (1992), Schiemenz (1990), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others; see also Stewart (1988); for identification of nymphs see Stewart (1986a)
Eupteryx notata Curtis, 1837 M V – E X; egg, 2 gen., at higher altitudes probably 1. Among low-growing herbs in oligotrophic and sunny, basic to acidic, dry to damp, occasionally also rather wet sites, mainly low-productivity meadows and pastures, dry grassland, heaths and roadside embankments, occasionally straw meadows and calcareous sedge fens. Usually on Hieracium pilosella and Leontodon hispidus, but also reported from Prunella, Thymus, Hypochaeris and Plantago. Widespread in Germany and fairly common, with strongholds in regions with less intensive agriculture (e.g. at higher altitudes, on sandy soils or in rather dry climates). Frequently found up to 1700 m a.s.l. in the Bavarian and Allgäu Alps, up to 2100 m a.s.l. in North Tyrol (Austria), and even at 2250 m a.s.l. in Switzerland. Achtziger (1991), Fischer (1972), Kuntze (1937), Reimer (1992), Remane (1987), Schiemenz (1969, 1990), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN; see also Günthart (1987a), Leising (1977), Morcos (1953), Payne (1981), Stewart (1988); for identification of nymphs see Stewart (1986a)
Eupteryx filicum (Newman, 1853) B VI – E X; egg, 2 gen. In damp to moderately dry, often acidic, more or less shady and rocky sites, usually in open forests (often on roadside embankments), locally in gardens and on walls. Host plants are various species of ferns, mainly Polypodium; in England and Wales P. vulgare and P. interjectum in the first generation, additionally Polystichum, Dryopteris, Phyllitis, Asplenium, Athyrium and Pteridium in the second. The range essentially includes the Mediterranean region and western Europe (northward to England, Wales and Ireland). From Germany it was first published only recently and is known mainly from western parts, usually below 550 m a.s.l., where it is locally common and occurs in high frequency. Localities near Remscheid, Cologne, Göttingen, Ilmenau, Gotha, Tauberbischofsheim, Bad Dürkheim and Freiburg may be situated at the edge of the range, but the species is probably under-recorded. A recent eastward range expansion, as in other western European species, is unlikely; because it is known from the Polish Beskid and Tatra Mountains since 1911 and 1925, respectively. Fechtler (pers. comm.), Frommer (pers. comm.), Remane & Fröhlich (1994b), HN; see also Badmin (1991, 1992), Smreczynski (1954), Stewart (1988); for identification of nymphs see Stewart (1986a)
Wagneripteryx germari (Zetterstedt, 1840) M VI – M X, occasionally until IV/V; egg, 1 gen. On Pinus sylvestris and P. mugo, reported also from P. nigra and P. cembra in the Swiss Alps; eurytopic in forests as well as on solitary trees, in bogs as well as among subalpine scrub. Widespread in Germany and in intermediate to high frequency, found up to 1950 m a.s.l. in the Bavarian and Allgäu Alps, up to 2150 m a.s.l. in Switzerland.
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Fischer (1972), Frommer (1996), Niedringhaus (1997), Reimer (1992), Remane & Reimer (1989), Schwoerbel (1957), Trümbach (1959), Wagner (1935, 1939a), HN, and others; see also Cerutti (1939a), Günthart & Günthart (1983)
Aguriahana pictilis (Stål, 1853) In Latvia E VI (Vilbaste 1974), in Austria M IX (Wagner & Franz (1961), egg(?), 1 gen.(?) This species is often listed for Germany, even in more recent literature, probably based upon an old statement of Typhlocyba picta F. (= Eupteryx atropunctata Goeze) which was copied by later authors. However, the only definite record has been gathered only very recently in the southern Black Forest: Föhrenmoos near Ibach, 1000 m a.s.l., 18.VIII.2001, 10 ‡‡, in a shady spruce bog. All individuals were swept from Vaccinium myrtillus (which is also reported to be the host plant in other European countries) growing under Betula pubescens, but the densities were extremely low. Thus it is likely that these individuals had fallen down from the tree canopies and that B. pubescens is the true host plant. Moreover, most, if not all species of the genus are purely arboricolous. Otherwise the species is also reported to occur in other parts of central Europe, but most localities appear to be rather isolated. Kirschbaum (1868), Sahlberg (1871), HN; see also Dworakowska (1972b), Metcalf (1968), Nast (1987), Ossiannilsson (1981)
Aguriahana stellulata (Burmeister, 1841) E VI – B X; egg, 1(?) gen. Eurytopic in semi-open cultivated areas (river floodplains, parks, fruit-tree plantations, forest margins, etc.), usually on Tilia cordata, Prunus avium (including fruit trees), P. spinosa, Populus nigra, Betula pendula, Acer platanoides and A. campestre, also reported from Aesculus hippocastanum in Switzerland. Widespread in the lowlands of Germany, but usually in low to intermediate frequency and low individual numbers only, highest localities are at 550 m a.s.l. in the Bavarian foreland of the Alps, but at 1200 m a.s.l. in the Engadine (Switzerland). Carl & Schönitzer (1995), Frommer (1996), Lehmann (1973a), Niedringhaus (1997), Olthoff (1986), Reimer (1992), Schwoerbel (1957), Wagner (1935, 1939a), Nickel (1997), HN; see also Günthart (1987a)
Zyginella pulchra P. Löw, 1885 Adults I – XII, mainly M VIII – B VI; adult, 1 gen. Mainly on Acer pseudoplatanus, less frequently on A. campestre, A. platanoides and A. monspessulanum, also occasionally (but probably not breeding) on Quercus, Alnus and Crataegus; overwintering individuals were found on Picea. Adults are usually swept along more or less shady forest margins as well as in parks and on roadside trees in urban areas. In the 1930s the only known German localities were near Mainz, but during the past two decades it was recorded from scattered sites over most of the middle parts of Germany (roughly between the Danube and a line from Cologne to Marburg, Göttingen, Erfurt and Dresden). Since 1985 some populations, which appear to be rather isolated, have also
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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. Furthermore, 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 awareness of the species in late summer, when collecting activity on trees is generally low. Frommer (1996), Niedringhaus & Olthoff (1986, 1993), Wagner (1939a), Remane & Fröhlich (1994b), SMNS, Nickel (1994, 1999b), HN; see also Dworakowska (1969), Nast (1976a)
Alnetoidia alneti (Dahlbom, 1850) Some authors argued that this might be a complex of several species, particularly based on the smaller body size, specific host association, and reduced yellowish pigmentation both in A. alneti f. coryli (Tollin, 1851) and A. alneti f. mali (Edwards, 1915). After a detailed analysis of morphometrics and colouration, Gillham & Claridge (1994) concluded that alder and hazel populations formed the extremes of a considerable variation, but are clearly conspecific with the nominate morph. They suggested that differences in morphology may be host-induced.
E V – M X; egg, 2 gen. Polyphagous on deciduous trees and shrubs in forests, in semi-open cultivated areas, and on solitary trees, preferentially in damp to wet sites, often abundant and in high frequency on Alnus glutinosa and A. incana, common also on Acer, Carpinus, Prunus, Corylus, Tilia, Cornus, Aesculus, in lower numbers on Quercus, Ulmus, Crataegus, Betula, Salix and Fagus. Widespread and common in the lowlands of Germany, frequently found up to 900 m a.s.l. in the Mittelgebirge and the foothills of the Alps, up to 1400 m a.s.l. in the Engadine (Switzerland). Achtziger (1991), Fischer (1972), Lehmann (1973a), Niedringhaus (1991, 1997), Reimer (1992), Remane (1987), Schiemenz (1990), HN, and others; see also Claridge & Wilson (1976, 1981), Gillham & Claridge (1994), Günthart (1987a)
Hauptidia distinguenda (Kirschbaum, 1868) Adults I – XII, mainly B IX – M VI; adult, 1 gen. Habitat requirements imperfectly known; usually found in xerothermic sites among ruderal herb stands and along herbaceous margins, often in vineyard situations; Geranium robertianum and G. pusillum are reported as food plants. Presumably described from the surroundings of Mainz, although the holotype label is no longer readable. From Germany known only from 6 localities (all below 300 m a.s.l.) along the upper and middle Rhine and its tributaries: Ahr near Altenahr, Nahe near Schloßböckelheim and Bad Münster am Stein, Neckar near Heilbronn (on Lagenaria siceraria), Rhine valley near Bacharach, and Wachenheim an der Weinstraße. Fröhlich & Nickel (unpublished data), Remane (1961b, and pers. comm), Schrameyer (pers. comm.), Wagner (1939a); see also Cerutti (1939a), Günthart (1987a), Ossiannilsson (1981)
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Hauptidia provincialis (Ribaut, 1931) So far VIII – I, ad.(?), 1(?) gen. This is a Mediterranean species so far only known from southern France, Italy, Croatia, Greece and Cyprus. From summer 2000 until spring 2002, a large, synanthropic population caused feeding damage on cultivated, perennial Ocimum basilicum and Valerianella locusta, in a greenhouse in southern Germany: Ulm, c. 500 m a.s.l. From southern countries, it is reported to feed on Parietaria, Urtica, Geranium and Solanum melangena and is apparently polyphagous on a number of plants avoided by most other leafhoppers. Coll. R. Dengler (det. HN); see also Dworakowska (1970d), Drosopoulos et al. (1986), della Giustina (1989)
Zyginidia franzi (W. Wagner, 1944) According to Dworakowska (1970b) this species has already been described as Erythroneura silvicola var. alpicola Cerutti, 1939. This interpretation was adopted by Nast (1972, 1987), della Giustina (1989) and others, who used the name Z. alpicola (Cer.). However, this name was created on the basis of a single †, the description of which included only two short notes on wing colouration and shape of the anal tube appendages, which only slightly diverged from typical Z. mocsaryi (Horv.). A type revision (P. Mazzoglio, pers. comm.) confirmed the conspecifity with the latter and Cerutti’s correct identification.
So far E VII – B X, according to Wagner & Franz (1961) in Styria (Austria) also E VI; egg(?), 1 gen. Confined to the Alps and Carpathians; in Germany found only in 5 localities between 1250 and 1800 m a.s.l.: Ponten above Hindelang, Deutenhausen near Füssen, Friederspitz above Garmisch-Partenkirchen, Sojernspitze above Mittenwald, and Königsbachalm above Berchtesgaden. Lives in open pine forests and alpine grassland, usually on limestone, presumably on Sesleria spp. Has been found above 2400 m a.s.l. in the Engadine (Switzerland). Fischer (1972), Nickel & Voith (unpublished data); see also Günthart (1987a, 1997), Lauterer (1984), Wagner & Franz (1961)
Zyginidia mocsaryi (Horváth, 1910) B VI – E IX, occasionally until M XI; egg, 2(?) gen. On Sesleria albicans in open pine and spruce forests, usually in warm situations on sunexposed calcareous or gypsum slopes, locally also on Festuca altissima in acidic and rather cool habitats. Rather localized in Germany and at the northern edge of the range, but abundant in favourable sites, with strongholds in the Bavarian Alps, the Franconian Jura, and the Triassic limestone and gypsum regions of Thuringia. Localities at the edge of the range include Bad Münstereifel, Höxter, Göttingen, Nordhausen, Jena and Dresden. Found at 900 m a.s.l. in the southern Black Forest and up to at least 1200 m a.s.l. in the Allgäu Alps, although single individuals were recorded at 1600 m a.s.l. Müller (1978), Rabeler (1962), Remane & Fröhlich (1994b), Rombach (1999b), Schiemenz (1990), Nickel (1994), HN
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Zyginidia pullula (Boheman, 1845) In Italy I – XII; ad, 4 gen. (A. Arzone pers. comm.). Has been reported from Germany in older papers, but most cases are likely to refer to the following species. On the other hand Z. pullula (Boh.) is reported to live in Denmark, the Czech Republic and Italy. From Germany, however, there is only a single confirmed record from eastern Bavaria: Oberpfälzer Jura, Naab valley near Kallmünz, 09.IX.1960, 1 †, which is likely to have been a vagrant. From Italy this species is reported to live on various grasses such as Elymus, Cynodon, Echinochloa, Hordeum, Sorghum, Triticum, Zea, Bromus, Digitaria, Setaria, Molinia and others (locally causing feeding damage to cultivated maize and other cereals), usually in disturbed sites such as fallow fields and waysides. Remane & Fröhlich (1994b); also Lauterer (pers. comm.), Metcalf (1968), Ossiannilsson (1981), Vidano & Arzone (1985)
Zyginidia scutellaris (Herrich-Schäffer, 1838) sensu Ribaut (1936) It is uncertain whether Ribaut’s interpretation, which is now commonly accepted, refers to the species described by Herrich-Schäffer. The holotype (a single ‡ from near Regensburg) is lost. At present, resident populations of this species are apparently absent from this region, but it may have occurred there during the 19th century, at least as a vagrant.
Adults I – XII, mainly M VIII – M VI; adult, 1(?) gen. Mainly in ruderal sites, disturbed dry grassland, abandoned vineyards, etc., undertaking flights in summer, and often found in large numbers in abandoned fields, fallows, winter cereals, maize fields, fertilized meadows, and along waysides. Host plants are various grasses (Festuca, Poa, Dactylis and others). The range includes the western Mediterranean region as well as France, Germany, Belgium, the Netherlands and England. In Germany not mentioned in older papers, and apparently present only since the 1950s (common in the Kaiserstuhl area in 1953, in middle Hessen since the 1960s). At present, permanent populations (which may show extremely high abundances) are known mainly from the southwestern half (Lake Constance basin, upper and middle Rhine, valleys of the Neckar, Main and Regnitz, Eifel Mountains, middle Hessen, up to at least 400 m a.s.l.), and very recently also in the Saale valley between Rudolstadt and Naumburg. In the latter localities, the species was definitely not present during the 1960s and 1970s. Furthermore, there are records of single individuals, which are probably vagrants, from the Kyffhäuser, the Thuringian Forest, the Altmühl and Neckar valley, and the Bavarian Alps and their foreland. Is reported to cause feeding damage on maize in France. In southwestern Germany, damage has been observed mainly on sweet corn; the economic importance, however, has not been quantified, and is perhaps negligible. Bornholdt & Remane (1993), Frommer (pers. comm.), Günthart (1988), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (pers. comm.), Rombach (1999b), Schiemenz (1990), Schrameyer (pers. comm.), Schwoerbel (1957), Walter (pers. comm.), ZIMH, HN; see also Della Giustina (1989), Kirschbaum (1868), Marion-Poll et al. (1987), Remane (1987), Wagner (1939a, 1951a), Wonn (1956)
Zyginidia viaduensis (W. Wagner, 1941) Has recently been treated as conspecific with Z. cornicula Meusnier, 1982, which lives in the subalpine belt of the western Alps and the Pyrenees (Mazzoglio 1989). This opinion is chal-
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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 Germany there are 7 known localities: Boizenburg an der Elbe, Gerwisch near Magdeburg, Groß Machnow near Zossen, Niederlehme near Königs Wusterhausen, Oderberg near Eberswalde, Mescherin (locus typicus) and Gartz, both near Angermünde. Schiemenz (1990), Wagner (1941a, 1944), HN; for identification see Wagner (1941a)
Zygina nivea (Mulsant & Rey, 1855) So far B VIII; adult(?), 1(?) gen. On Populus alba, in France also reported from P. nigra, Salix eleagnos and S. alba. From Germany there are only 4 records from the upper and middle Rhine between 50 and 220 m a.s.l.: Burkheim, 02.VIII.1997, Buggingen, 06.VIII.1997, small populations on P. alba in river floodplains, as well as in urban areas of Cologne, two records between 1993 and 1995. Presumably, this species is more widespread in western parts of Germany. On the other hand older reports either refer to Edwardsiana candidula (Kbm.) or originate from areas outside present-day Germany. Frommer (pers. comm.), HN; see also Ribaut (1936)
Zygina tithide Ferrari, 1882 So far only M IX; in the Valais (Switzerland) according to Cerutti (1939a) hibernation as adults, 1(?) gen. After a revision a single † collected near Jena (Thuringia, Leuchtenburg near Kahla, 400 m a.s.l., 17.IX. 1967), which had formerly been published as Z. nivea (M. & R.), turned out to belong to Z. tithide Ferr. However, due to the absence of the host plant at the collecting locality, a permanent occurrence is not proven. The nearest known localities are in Poland (near Poznan), Bohemia (Czech Republic), eastern Austria and the Valais (Switzerland). From the latter region this species is reported to live on Populus alba and other deciduous trees and shrubs, migrating to coniferous trees, Juniperus, and Rubus fruticosus for hibernation. Coll. Müller, Schiemenz (1990); see also Cerutti (1939a), Dworakowska (1977b), Giordano et al. (2001), Nast (1976a), Remane & Fröhlich (1994b)
Zygina lunaris (Mulsant et Rey, 1855) In Strasbourg (France) VII – III; adult, 1 gen. (Lauterer & Malenovsky 1995). This species was recently found in parks in the upper Rhine floodplains in Kehl and Weil am Rhein and adjacent suburbs (Baden, c. 140 and 260 m a.s.l.), as well in Strasbourg (Alsace, France) and Basel (Switzerland), where it is rather common on Salix alba, S. fragilis, S. babylonica and S. purpurea, migrating to Acer and Rosa in October and to
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various coniferous trees for hibernation. Otherwise it is only known to occur in the western Mediterranean region. Lauterer & Malenovsky (1995), Mühlethaler (2001, and pers. comm.), HN
Zygina angusta Lethierry, 1874 Populations living on Rosaceae differ slightly in colouration from those on Fagaceae and may belong to a distinct species (Remane 1994). Moreover, since the description of Z. nigritarsis Rem., most published records are in need of revision.
E VII – B VI; adult, 1 gen. In semi-open stands of shrubs and trees, usually in dry to damp sites; on the one hand on hedges and shrubs such as Crataegus, Rosa and Prunus spinosa, on the other hand along forest margins on Quercus robur, in lower numbers also on Qu. petraea, Fagus, Carpinus, Prunus padus and others; found on coniferous trees (Picea, Pinus, Juniperus) during hibernation, on Ribes in spring. The species is poorly documented, but apparently widespread in Germany between the foothills of the Alps and the coast, common at least in middle and southern parts; the highest localities are at 700 m a.s.l. Achtziger (1995a), Frommer (1996), Niedringhaus (1997), Reimer (1992), Remane (1987, and pers. comm.), Nickel (1997), HN; for identification see Remane (1994)
Zygina flammigera (Geoffroy, 1785) M VII – B VI; adult, 1 gen. On various woody plants, in summer mainly on Rosaceae; the main hosts are several species of Prunus, Malus, Crataegus, Sorbus, and perhaps Rosa. Also recorded on Fagus, Quercus and others, usually on Picea in winter, occasionally on Pinus, Buxus, Juniperus, Rubus and Hedera. Widespread in the lowlands of Germany and common, found up to 950 m a.s.l. in the Mittelgebirge and the Bavarian Alps, at 1280 m a.s.l. in the Engadine (Switzerland), at 1400 m a.s.l. in the Trentino (northern Italy). Achtziger (1995a), Frommer (1996), Niedringhaus (1991, 1997), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987, 1994), Schiemenz (1990), HN; see also Günthart (1987a)
Zygina ordinaria (Ribaut, 1936) This taxon is often confused with Z. tiliae (Fall.) and other species of Zygina. According to Remane (1994) it may comprise a complex of several distinct species.
Adults I – XII, mainly M VII – E V; adult, 1 gen. On narrow-leaved species of willows, often near running water, as well as in heath and bog habitats, usually on willows, notably Salix alba, S. triandra, S. viminalis, S. fragilis, S. purpurea, S. repens and S. pentandra, occasionally also on S. eleagnos and additional species. The overwintering sites are unknown. Widespread in Germany, but poorly documented; usually in intermediate frequency, but low individual numbers on the hosts, found up to at least 950 m a.s.l. in the Bavarian Alps. Frommer (pers. comm.), Reimer (1992), Remane (pers. comm.), Nickel (1997, 1999b); for identification see Remane (1994), Günthart (1974)
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Zygina rosea (Flor, 1861) According to Remane & Fröhlich (1994b) the name Z. roseipennis (Tollin, 1851), which is used by many authors for this taxon (e.g. Nast 1972, 1987), is doubtful and should be rejected. Furthermore, literature records from central and southern Europe probably refer to other species and should be revised.
So far B VIII – M II, in Sweden according to Ossiannilsson (1981) until B VI; adult, 1 gen. In raised and intermediate bogs, and along their margins. Nymphal development and most adults on Betula pubescens, hibernation takes place on conifers, mainly Pinus sylvestris (although there is one record from Pseudotsuga douglasi). In Germany largely restricted to the remnant bog areas of the northern plains, recorded near Meppen, Lingen (Ems), Oldenburg, Hamburg, Hanover and Ratzeburg (probably overlooked in Mecklenburg and Brandenburg), also found near Fulda and Munich (550 m a.s.l.). Niedringhaus (1997), Remane (1962), Remane & Fröhlich (1994b), Wagner (1941b), Weis (pers. comm.); see also Linnavuori (1952), Nuorteva (1952a)
Zygina rubrovittata (Lethierry, 1869) Adults I – XII; adult, 2(?) gen. In sunny, usually dry and sandy sites (mainly heaths, edges of pine forests, drained raised bogs) on Calluna vulgaris, which is also the winter food plant. In Germany locally common in the northern plains, but rather scattered and restricted to regions of sand and sandstone in middle parts (middle Hessen, Thuringian Forest, Erzgebirge, Pfälzer Wald, northern upper Rhine plain, Mittelfranken, Danube valley); apparently absent from the Alps, their foothills and foreland. The highest localities are at 450 m a.s.l. Frommer (pers. comm.), Remane (pers. comm.), Schiemenz (1990), Struve (1939), Trümbach (1959), Wagner (1935), HN; see also Badmin (1995)
Zygina suavis Rey, 1891 sensu Ossiannilsson (1981) This is a rather critical taxon, the identity and distinction of which are subject to diverging opinions in the literature (see Vidano 1959b, 1961). Perhaps there is another distinct species in central Europe, showing a very similar colouration (see Remane 1994), but living on deciduous trees (probably Quercus or Tilia). However, until further studies have been done, Vidano’s opinion should be followed, treating Zygina rhamnicola Horváth, 1903 and Erythroneura inconstans Ribaut, 1936 as junior synonyms of Z. suavis R. Another problem is caused by the fact that Z. schneideri Günth., which is apparently widespread in Europe, was described only in 1974. Thus most data published before that are in need of revision.
M VII – E V; adult, 1(?) gen., according to Günthart (1974) in Switzerland 2 gen. In moist to dry, basic as well as acidic sites, mainly open forests, pastures with scattered shrubs, and various margins (notably hedges, forest and bog edges), preferentially on Frangula alnus, at least in low numbers also on Rhamnus cathartica, on Picea abies during hibernation. In Germany only poorly documented and often confused, but widespread in the lowlands, where occurring in intermediate frequency at least on Frangula alnus. Northernmost localities are near Papenburg, Kiel and Königs Wusterhausen, but the species is also reported from southern parts of Sweden and Finland and from north-
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western Poland. Found up to at least 600 m a.s.l. in the German Mittelgebirge, up to 1200 m a.s.l. in the Engadine (Switzerland). Niedringhaus (1997), Reimer (1992), Remane (1987, and pers. comm.), Schiemenz (1990), Nickel (1994), HN; see also Günthart (1974, 1987a), Nast (1976a), Ossiannilsson (1981); for identification see Remane (1994)
Zygina schneideri (Günthart, 1974) See taxonomic notes on Z. suavis R.
Adults I – XII, mainly M VII – B VI; adult, 1(?) gen. On woody species of Rosaceae in rather warm sites, mainly on sunny hillsides and along hedges on Prunus spinosa, Rosa canina and R. rubiginosa, as well as in urban settlements on Rosa rugosa, Potentilla fruticosa, Amelanchier, Cotoneaster, and other ornamental shrubs; hibernation probably takes place in the litter layer. Widespread in middle parts of Germany, between the Danube and a line from Cologne to Braunschweig, Halle and Frankfurt an der Oder, particularly common and in high frequency in warm regions, found up to 900 m a.s.l. on the Swabian Jura. Is also reported from England, southern Sweden, southern Norway. Achtziger (1995a), Bornholdt & Remane (1993), Frommer (pers. comm.), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987, 1994), Remane & Fröhlich (1994b), Rombach (1999a), Nickel (1994), HN; see also Le Quesne & Payne (1981), Ossiannilsson (1981); for identification see Günthart (1974), Remane (1994)
Zygina tiliae (Fallén, 1806) Adults I – XII, mainly E VII – B VI; adult, 1(?) gen. In open woodland and along forest margins, in wet to damp, occasionally also rather dry sites. The main host is Alnus glutinosa. Lower numbers are also swept from Alnus incana, Tilia cordata (where breeding has been found to take place at least locally) and other woody species (notably Prunus padus, Salix and Betula); at least occasionally found on Picea abies in winter. Widespread and locally common in Germany, at least to the north of the Danube, although poorly documented due to identification problems. Recorded below 450 m a.s.l. only, but reported from 1190 m a.s.l. in the Engadine (Switzerland). Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987, and pers. comm.), Nickel & Remane (1996), HN; see also Günthart (1987a); for identification see Remane (1994)
Zygina rosincola (Cerutti, 1939) Mainly E VII – V; adult, 1(?) gen. Usually in rather warm and sun-exposed sites on Rosa (from which nymphal development is reported), Prunus spinosa and Crataegus; adults occasionally also on Quercus and Betula; in winter found on coniferous trees (Pinus, Picea) and Juniperus. The distribution in Germany is poorly documented due to difficulties in determination. Apparently widespread, but uncommon at least in the western half (Bavaria, Hessen, southern Niedersachsen, Westphalia) between the Danube and the northern edge of the Mittelgebirge; not recorded from eastern parts (from Mecklenburg-Vorpommern to Saxony and Thuring-
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ia), where probably overlooked. The highest localities are at only 500 m a.s.l. From Switzerland this species is reported from Rosa glauca, R. pendulina, R. montana, Sorbus aucuparia, Prunus padus and Malus, in altitudes up to 1520 m a.s.l. Remane & Fröhlich (1994b), Nickel & Remane (1996), Nickel (1994), HN; see also Günthart (1987b, and pers. comm.); for identification see Günthart (1974), Remane (1994)
Zygina griseombra Remane, 1994 B VII – B V; adult, 1 gen. Nymphal development takes place on Carpinus betulus, usually along margins of forests and in their interior; after emergence adults migrate to other woody plants and are recorded on coniferous trees during winter, frequently on Crataegus in more open sites in spring. In Germany the species is only known from the region between the Main and the northern edge of the Mittelgebirge, where it occurs in rather low frequency and individual numbers (although probably under-recorded). Northernmost localities are near Altenahr, Marburg (locus typicus: Elnhausen), Göttingen, Bad Frankenhausen and Dresden, at least up to 500 m a.s.l. Outside Germany it has been recorded so far only from northern France, Moravia (Czech Republic), Styria (Austria), the canton of Zurich (Switzerland) and southern Italy. Remane (1994, and pers. comm.), Nickel & Remane (1996), Nickel (1994), HN; see also Günthart (2000), Holzinger & Remane (1994); see also della Giustina & Remane (2001), Lauterer (2000), for identification see Remane (1994)
Zygina nigritarsis Remane, 1994 This species has been described only recently and is perhaps widespread in the European deciduous forest zone. Thus, a revision of older material labelled as Z. angusta Leth. or Z. tiliae (Fall.) is likely to reveal more records.
M VIII – B VI; adult, 1 gen. Nymphs on Prunus padus, mainly in floodplains of rivers and streams, adults migrating to coniferous trees (Picea, Pinus) for hibernation; found on Crataegus, Ribes and other woody plants in spring (also in parks, gardens, dry grassland and other habitats). The distribution in Germany is only poorly documented, but the species is widespread at least between the foreland of the Alps and the northern edge of the Mittelgebirge, although apparently scattered and in low frequency; northernmost localities are near Oldenburg, Hamburg, Plön, Bad Frankenhausen and Bitterfeld, recorded up to 550 m a.s.l. Outside Germany only known from North Tyrol (Inn valley near Mils, Austria), but probably more widespread. Frommer (pers. comm.), Neumann (1997), Remane (1994), ZIMH, ZIMH, HN; for identification see Remane (1994)
Zygina hyperici (Herrich-Schäffer, 1836) M V – M X; egg, 2 gen. Monophagous on Hypericum perforatum in sunny, damp to dry, usually disturbed sites (ruderal habitats, waysides, railway embankments, mining areas, grazed or abandoned grass-
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land, etc.). Widespread in the lowlands of Germany, in intermediate to high frequency on the host, but rather scattered in the northwestern plain and to the south of the Danube, usually found below 500 m a.s.l., locally up to 1000 m a.s.l. in the southern Black Forest. Achtziger (1991), Frommer (pers. comm.), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schiemenz (1990), Wagner (1935, 1939a), HN
Zygina hypermaculata Remane & Holzinger, 1995 So far E VII – E IX; egg, 1 gen. This species has only recently been described from Upper Bavaria and Styria (Austria). It is known only from the montane and subalpine belt (between 1300 and 1900 m a.s.l.) of the northeastern Alps, living on Hypericum maculatum. In this region, however, it appears to be widespread and fairly common. The locus typicus is Brauneck above Bad Tölz (1500 m a.s.l., 07.VIII.1961), the paratypes are from three sites in Styria. Further Austrian records are from Carinthia and Upper Austria. In Germany the species has been found in the Berchtesgaden and Karwendel Alps (Roßfeld, Schneibstein and Reiteralm, all above Berchtesgaden, Sojernspitze near Mittenwald), in herb stands on sunny hillsides as well as in open montane coniferous forests. Remane & Holzinger (1995), HN; see also Holzinger (1999b)
Arboridia kratochvili (Lang, 1945) So far M VI – B X; egg(?), 2(?) gen. Until recently this species was only known from a few localities in southern Poland, the Czech Republic and Slovakia. Since 1993 it has been recorded from altogether 6 sites on Triassic limestone plateaus in the Main-Saale region of Franconia (Thüngersheim, Gambach, Wiesenfeld, Aschfeld, Gössenheim and Machtilshausen, all c. 300 m a.s.l.). In these sites it is locally abundant on Potentilla tabernaemontani in xerothermic grassland with low-growing and incomplete cover of vegetation (Trinio-Caricetum humilis). Nickel & Remane (1996), HN; see also Dlabola (1977), Lauterer (pers. comm.), Nast (1976a); for identification see Lang (1945)
Arboridia erecta (Ribaut, 1931) So far B VIII – B VI, adult, 1(?) gen. On deciduous trees and shrubs in warm situations; reported to live on Acer campestre and occasionally Castanea sativa in Italy, also on Prunus, Quercus, Corylus and Tilia in Switzerland and France. In Germany the species lives at the northern edge of its range and is only known from 4 records in Baden-Württemberg and Bavaria: Markgröningen near Stuttgart, 230 m a.s.l., 17.IX.1980, 1 †, at light; Schwörstadt near Lörrach, 290 m a.s.l., 05.IV.1987, in forest litter, and 02.VI. 1987, on Malus and Prunus avium, altogether 7 ††, 5 ‡‡; Freiburg, municipal cemetery, 250 m a.s.l., 04.VIII.1997, 4 ††, 2 ‡‡, on Acer platanoides; Schönhofen near Regensburg, 450 m a.s.l., 16.IX.1997. Recorded at 1530 m a.s.l. in the Engadine (Switzerland).
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Günthart (1988, and pers. comm.), Heller (1996), Remane (pers. comm.), HN; see also Günthart (1987b), Ribaut (1936), Vidano & Arzone (1987b)
Arboridia parvula (Boheman, 1845) A. potentillae (Moravskaja, 1948) was described from Potentilla arenaria (= P. incana) in southern Russia. Apparently, it can be distinguished from A. parvula (Boh.) only by the narrowing distal half of the aedeagus stem. Specimens found on Potentilla incana in several xerothermic localities in eastern parts of Germany may belong to this taxon. Provisionally, however, they shall be treated as A. parvula (Boh.), but the problem requires further studies of morphology and ecology.
Adults I – XII, mainly M VII – B VI; adult, 1(?) gen. Usually along woody margins in dry to damp (occasionally also moist) sites, often in rather warm situations, notably dry grassland with scattered shrubs and trees, along hedges and forest margins. Adults on herbaceous and shrubby species of Rosaceae (Potentilla incana, Filipendula ulmaria, Rubus idaeus and others), in lower numbers also on various deciduous trees (Quercus, Prunus padus and others); most winter records are from herbaceous vegetation. Widespread in Germany and locally common, particularly in warmer regions, but rather sporadic and uncommon in most parts of the Mittelgebirge and the north German plain, not reported from Mecklenburg-Vorpommern. The highest localities are at 1400 m a.s.l. in the Bavarian Alps, but the species is reported to occur frequently in the subalpine belt up to 1800 m a.s.l. in Switzerland. Niedringhaus & Olthoff (1993), Post-Plangg & Hoffmann (1982), Reimer (1992), Schwoerbel (1957), Schiemenz (1990), Nickel & Remane (1996), HN; see also Cerutti (1939a), Günthart (1987a)
Arboridia ribauti (Ossiannilsson, 1937) M VII – B VI; adult, 1 gen. On deciduous trees (mainly Carpinus betulus, but also Tilia cordata and Quercus), usually along forest margins and roads, sometimes also along hedges. Winter specimens have been found on Calluna vulgaris and Hedera helix. In Germany recorded only between the edge of the Alps and a line from Düsseldorf to Göttingen, Halle and Dresden, where usually occurring in intermediate to low frequency and rather low numbers. Highest localities are at 750 m a.s.l. in the Black Forest, but at 1400 m a.s.l. in the Trentino (northern Italy) and at 1300 m a.s.l. in the Valais (Switzerland). Frommer (1996), Kolbe & Bruns (pers. comm.), Reimer (1992), Schiemenz (1990), Wagner (1951a), HN; see also Cerutti (1939a); for identification see Wagner (1939a)
Arboridia simillima (W. Wagner, 1939) Recently A. loginovae (Emelyanov, 1964), known from the Altai Mountains, Siberia and Kazakhstan, has been reported from Moravia (Czech Republic) by Lauterer (2000). Morphology and ecology are very similar as in A. simillima (W.Wg.). Southwest German populations of the latter have been found to show a considerable morphological variation in the shape of the aedeagus. Thus, the distinction of these two taxa is in need of clarification.
Adults I – XII, mainly M VII – M V; adult, 1(?) gen.
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On sun-exposed shrubs and in open xerothermic woodland, mainly in slope and plateau 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 Kyffhäuser, Hainleite, and valleys of Unstrut, Saale and Ilm). Further localities are situated in the Eifel Mountains and along the upper course of the Isar (Vorderriß, c. 1000 m a.s.l.). The species is probably under-recorded. Post-Plangg & Hoffmann (1982), Schiemenz (1990), Wagner (1939a), Nickel & Remane (1996), HN; for identification see Wagner (1939a)
Arboridia velata (Ribaut, 1952) E VII – M V; adult, 1 gen. Along sunny margins of oak forests, usually on rather warm upper slopes. At present known from Germany only from the Lake Constance Basin, the valleys of the Rhine (between Waldshut and Koblenz), Nahe and Neckar, the slopes along the Danube between Kelheim and Passau, the Kyffhäuser, the upper course of the Saale near Bad Blankenburg, and the Niederlausitz. The majority of individuals has been swept from oaks (Quercus petraea and Qu. robur, perhaps also Qu. pubescens), a few – probably vagrants – also from neighbouring woody species. Highest localities are at c. 500 m a.s.l., but at 1200 m a.s.l. in the Valais (Switzerland). Günthart (1988), Schiemenz (1990), SMNS, Nickel (1999b), HN; see also Cerutti (1939a), Ribaut (1931, 1936), Vidano & Arzone (1987b)
Arboridia spathulata (Ribaut, 1931) According to Dworakowska (1970c) this is a junior synonym of A. versuta (Melichar, 1897). However, the latter was described after a ‡, and most ‡‡ in this genus cannot be identified with certainty. In contrast, Ribaut’s description contains exact and unambiguous drawings of † genitalia. Hence, the identity of A. versuta (Mel.) is considered here as dubious.
So far only E X; adult(?), 1(?) gen. In central Europe this species has only rarely been found. The only German record is from Franconia: Bad Windsheim, c. 320 m a.s.l., 20.X.1937. From Italy, it is reported to live monophagously on Quercus petraea, Qu. robur, Qu. pubescens and Qu. cerris. Has been found at 1250 m a.s.l. in the Valais (Switzerland). Wagner (1939a); see also Cerutti (1939a), Vidano & Arzone (1987b)
Arboridia pusilla (Ribaut, 1936) Adults I – XII, mainly B VII – M VI, adult, 2(?) gen. On Geranium sanguineum, along woody and herbaceous margins in rather warm situations, occasionally also in open pine and oak forests and in abandoned vineyards. Often syntopic with A. simillima (W.Wg.), but in different layers of the vegetation. In Germany
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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 Tuttlingen and Regensburg) and Kaiserstuhl. The highest localities are at 900 m a.s.l. Bornholdt & Remane (1993), Krause & Emmrich (1996), Remane (1987), Schiemenz (1990), Wagner (1939a), Nickel & Remane (1996), HN
Fruticidia bisignata (Mulsant et Rey, 1855) E VIII – B VII; adult, 1 gen. The range of this species essentially includes the Mediterranean region, but extends northeastward to the upper and middle parts of the Rhine valley and its tributaries. Border localities are near Cologne, Koblenz, Bad Nauheim, Lohr am Main, Heidelberg and Freiburg. Lives in various types of dry grassland with scattered shrubs or trees and along sunny forest margins, usually on woody species of Rosaceae (mainly Crataegus, but also Malus, Amelanchier and Quercus); winter specimens have been found on Calluna and coniferous trees. The highest localities are at c. 300 m a.s.l. only. Frommer (pers. comm.), Günthart (1988), Post-Plangg & Hoffmann (1982), Remane (pers. comm.), SMNS, Wagner (1939a), Nickel (1999b), HN
Fruticidia sanguinosa (Rey, 1891) A single †, which may have been a vagrant, was found near Heidelberg (Sandhausen), 27.IX.1982, on Crataegus, along the margin of a xerothermic inland dune. Otherwise, this species is poorly known and only reported to occur in Mediterranean regions of France and Italy, feeding on Crataegus, Rosa and Prunus, usually syntopically with the preceding one. Heller (1996); see also D’Urso (1995), Ribaut (1936)
4.2.4.13 Deltocephalinae Fieber, 1869 Fieberiella florii (Stål, 1864) M VII – M X, single individuals B VI; egg, 1 gen. A polyphagous species living on woody plants in open shrubland and along various margins, mainly in dry grassland with scattered shrubs, but also in ruderal sites, along rivers, streams and hedges, in parks and gardens. Host plants are usually Rosa, Prunus spinosa, Ligustrum vulgare and Rubus idaeus (found only synanthropic on the two latter species), at least single adults also on Quercus, Ulmus, Cytisus, Salix, Syringa, Elaeagnus and others. In Germany at the northern edge of the range, with strongholds in the warmer regions (mainly the basins of the Rhine, Main and Neckar). Northernmost records are
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from near Cologne, Bad Nauheim, Schlüchtern, Sondershausen, Berlin and Frankfurt an der Oder, up to at least 400 m a.s.l., but reported from 880 m a.s.l. in Switzerland. Cobben (1956), Frommer (pers. comm.), Meyer-Arndt & Remane (1992), Schiemenz et al. (1996), Wagner (1963), Nickel & Remane (1996), HN; see also Günthart (1987b)
Fieberiella septentrionalis W. Wagner, 1963 B VII – E X (in southwestern Germany from E V onwards); egg, 1 gen. On Prunus spinosa and Rosa species in dry grassland, ruderal sites, along forest margins and hedges, but also synanthropic in gardens and parks, where usually found on Rosa rugosa, Ligustrum vulgare, Rubus idaeus, R. fruticosus, Rhododendron, Spiraea, Cotoneaster, Solidago canadensis and S. gigantea. Generally in similar sites as the preceding species and syntopical in some places, but with a different distribution pattern. A tongue-shaped part of the range extends into central parts of Germany from the southeast. According to existing data, border localities include Regensburg, Ansbach, Landau (Pfalz), Bad Kreuznach, Koblenz, Cologne, Minden, Quedlinburg, Halle and Eberswalde. Furthermore, the species occurs locally in urban settlements near the coast and in the foreland of the Alps (Bremen, Hamburg, Rostock, Munich), where it is perhaps extending its range. The highest localities are at c. 550 m a.s.l., the locus typicus is Jena. Frommer (1996), Jöst (1966), Meyer-Arndt & Remane (1992), Niedringhaus & Olthoff (1986, 1993), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1963), HN
Grypotes puncticollis (Herrich-Schäffer, 1834) E VI – M X, according to Schiemenz et al. (1996) also M V; egg, 1 gen. On pines in forests and along their edges as well as on solitary trees, in Germany recorded on Pinus sylvestris only, in Styria (Austria) also on P. nigra. Single individuals on dispersal flight are frequently swept from herbaceous vegetation of various habitats (meadows, dry grassland, etc.). Widespread in Germany, in high frequency and at intermediate to high densities on the host, found up to at least 900 m a.s.l. in the Mittelgebirge and the Bavarian Alps; recorded at 1400 m a.s.l. in Switzerland. Fischer (1972), Niedringhaus (1991), Remane (1987), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a), HN; see also Günthart (1987b), Holzinger (pers. comm.)
Goniagnathus brevis (Herrich-Schäffer, 1835) B VIII – E VI, occasionally also VII; adult, 1 gen. In sunny, moderately dry to dry and sparsely vegetated sites on various substrates (limestone, gypsum, porphyry, diabas, sandstone, sand, etc.). In several occasions, Thymus pulegioides and Th. praecox were recorded as food plant. In Germany confined to the southern half and at the edge of the range; northernmost localities are on a line from Koblenz to Marburg, Göttingen, Quedlinburg, Halle, Leipzig and Bautzen. Found up to 800 m a.s.l on the Swabian Jura and in the foothills of the Alps.
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Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (pers. comm.), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1939, 1951), Nickel (1994), HN
Japananus hyalinus (Osborn, 1900) Japananus meridionalis Bonfils, 1981
B VIII – M X; egg, 1 gen. Described from North America, where it is probably introduced; the main range includes the deciduous forest zones of eastern Asia and Europe. Several authors have suggested that it was also introduced to Europe, based on the fact that it has been found on ornamental trees (notably Acer japonicum and A. palmatum, both originating from Japan) and on the late discovery, which happened only in 1942 in Styria (Austria). Since then it was also found in Catalonia (Spain), southern France, Switzerland, southwest Germany, northern Italy, the Czech Republic, Slovakia, Hungary, Serbia, Romania, Bulgaria and southern Russia. However, many, if not most European records are from native maples in habitats which suffered little anthropogenic influence, although this is only partially true for the German localities, most of which are in urban areas: Freiburg (Günterstal, 2001, on Acer campestre), Stuttgart (Hofen, 1985, on A. campestre, and Rosensteinpark, 1985, on A. pseudoplatanus), Neckarsulm, Heilbronn (both 2001, on A. platanoides, A. palmatum and A. campestre), Munich (2001, on ornamental Acer) and Cologne (1994), up to 550 m a.s.l. The species is monophagous on maple; Acer campestre is apparently the main host in Europe. Billen & Schrameyer (pers. comm., det. Achtziger), Frommer (1996), Heller (1987b), Weis (pers. comm.); see also Arno et al. (1988), Arzone et al. (1987), della Giustina (1989), Lauterer (1980, 1984, 1989), Mühlethaler (2001), Remane & Fröhlich (1994b)
Opsius stactogalus Fieber, 1866 E VI – B X; egg, 1 gen. In central Europe this species was originally restricted to gravel banks along rivers in the Alps and their foothills. At least since the middle of the 19th century it has been reported from Tamarix species in gardens and parks of the lowlands and is now widespread and common all over Germany (for instance in northern Germany recorded from the East Frisian Islands, Kiel and Rostock). Since the beginning of the 20th century it is also known from North America, where it was probably introduced. From natural habitats, it is at present only known from the Isar between Mittenwald and Wolfratshausen (formerly also along the Saalach, Lech and upper Rhine), where it is locally common on Myricaria germanica. Fischer (1972), Frommer (1996), Kirschbaum (1868), Kuntze (1937), Lauterborn (1923), Niedringhaus (1991), Remane (pers. comm.), Schönitzer & Oesterling (1998b), Wagner (1939a), Nickel (1999a), HN; see also Hamilton (1983a)
Neoaliturus fenestratus (Herrich-Schäffer, 1834) group The taxonomy of this group is unclear and in need of a thorough revision. In Germany alone there may be two, or perhaps even three morphologically and ecologically distinct forms.
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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 grassland, inland dunes, rocky slopes, abandoned fields and vineyards, waysides and mining 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 common in rather warm situations mainly below c. 400 m a.s.l., but uncommon in western parts of the north German plain as well as in the Bavarian and Allgäu Alps, where frequently recorded up to 1800 m a.s.l. Also reported from alpine habitats of Austria and Italy, but apparently absent from Denmark, Norway and Sweden. From Israel it is reported as a vector of Safflower phyllody in fields of Carthamus tinctorius, although it is uncertain whether these populations are conspecific with the central European ones. Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Schwoerbel (1957), Wagner (1951a), Nickel (1994), HN; see also Klein (1992), Leising (1977), Ossiannilsson (1983)
Circulifer opacipennis (Lethierry, 1876) sensu Young & Frazier (1954) Belongs to a taxonomically critical species group, which lives mainly in rather warm and dry regions of the Old and New World. Some taxa are suspected to transmit various virus and MLO diseases (e.g. Klein 1992). The identity of populations found in Germany is not quite clear. According to Nast (1972, 1987) Circulifer is a junior synonym of Neoaliturus, and Cicadula opacipennis Leth. and Jassus haagii Kirschbaum, 1868 (locus typicus near Mainz, described after 2 ‡‡) are junior synonyms of C. haematoceps (Mulsant & Rey, 1855). According to Young & Frazier (1954) and Oman (1970), however, Circulifer is a distinct genus and C. opacipennis (Leth.) a distinct species. This interpretation is followed here until further research has been done.
This taxon is reported from various open habitats in the Mediterranean region and the Near East where it probably lives polyphagously on dicotyledonous herbs and dwarf shrubs (adults found on Atriplex, Cistus, Rosmarinus, Marrubium, Salicornia, Portulaca, Thymus, Beta and others, often on succulents). Populations in central Europe may be only unstable. Presumably 2 ‡‡ collected near Mainz and described as Jassus haagii Kirschbaum, 1868 („Mombach, Sandhügel, zwischen niederen Pflanzen, Ende April“) belong to this taxon. Recently, it was also found in the Oberlausitz (Lauta, 1994, 1 † in a yellow tray), in mining areas near Halle, as well as on flat roofs of Basel (Switzerland). Funke (pers. comm.), Al Hussein et al. (1999), Kirschbaum (1868), Wagner (1939a), Walter & Emmrich (1995); see also della Giustina (1989), Mühlethaler (2001), Remane (1987), Young & Frazier (1954)
Coryphaelus gyllenhalii (Fallén, 1826) So far B VII – B IX; egg, 1 gen. Along shores of mesotrophic lakes and ponds, in dense, often slightly shaded stands of Schoenoplectus lacustris. In Germany at the western and southern edge of the range and confind to the eastern half, mainly in glacial lakes of the northeast German plain and Upper Bavaria, as well as in fish pond areas of the Oberlausitz, Thuringian Forest and southern Harz Mountains. Western- and southernmost localities, which appear to be
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strongly isolated from each other, are near Berlin, Bad Sachsa, Arnstadt, Ilmenau, Seeshaupt and Penzberg (600 m a.s.l.). Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Nickel (1997), HN
Balclutha boica W. Wagner, 1950 For this taxon the name B. lineolata (Horváth, 1904) was commonly used for several decades. However, Webb & Vilbaste (1994) recently studied Horváth’s type material collected in central Asia and concluded that it is a junior synonym of B. punctata (F.). This opinion is also supported by Emelyanov (pers. comm.).
In Latvia and Lithuania according to Vilbaste (1974) M VIII – M V; adult(?), 1(?) gen. Reported mainly from the eastern half of Europe. The only German record is from the type locality in the Bavarian foreland of the Alps: Landsberg am Lech, 575m a.s.l., 02.IX.1941, 3 †† „im hohen Gras der Auen am linken Lechufer ..., der Fundort wurde noch 1941 durch Baumaßnahmen zerstört“. Reported from forests and forest glades in Latvia and Lithuania, from Picea abies in spring in Sweden. Otherwise only published from Estonia, Poland, the Czech Republic, Slovakia, Bulgaria and Greece. Reports from other countries may refer to other species. Fischer (1972), Wagner (1950); see also Anufriev & Emalyanov (1988), Drosopoulos et al. (1986), Nast (1987), Ossiannilsson (1983), Vilbaste (1974), Webb & Vilbaste (1994)
Balclutha calamagrostis Ossiannilsson, 1961 Adults I – XII, mainly M VIII – E V; adult, 1 gen. In scattered stands of Calamagrostis epigejos, occasionally also on C. pseudophragmites, in sunny to moderately shady, moderately dry to moderately moist sites, mainly in mining areas, ruderal sites, forest glades, as well as on gravel banks of alpine rivers. Hibernation presumably takes place in the grass layer. In Germany only poorly documented and often misidentified, but apparently widespread and common in favourable sites, e.g. in southern and eastern Niedersachsen, in parts of Saxony-Anhalt, Thuringia, Saxony, northern Baden, as well as in Upper Bavaria, found up to at least c. 800 m a.s.l. in the Rhön Mountains. So far there are only few further records from northern and central parts of Europe (Sweden, Finland, Baltic states, Poland, the Czech Republic, Austria, eastern France), but the range is imperfectly known. German localities may be near its southern and western border. Fischer (1972), Funke & Witsack (1998), Remane (1987), Remane & Reimer (1989), Rombach (1999a), Schiemenz et al. (1996), Walter (pers. comm.), Nickel (1997), HN; see also della Giustina & Remane (2001), Holzinger (1996b), Nast (1987), Ossiannilsson (1983), Szwedo et al. (1996)
Balclutha punctata (Fabricius, 1775) sensu Wagner (1939) Balclutha lineolata (Horváth, 1904) sec. Webb & Vilbaste (1994) The taxonomic situation of European species of Balclutha is critical due to strong morphological similarities. Further complications arose by the installation of a neotype for the name B. punctata (F.) by Blocker (1967), who chose a specimen of the following species, which in turn had been described as B. rhenana W. Wagner 1939 (see below). Following Blocker, Nast (1986, 1987) decided that B. tricolor (Gmelin, 1790) should be the valid name for B. punctata (F.) sensu Wagner.
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However, there is no evidence at all that Blocker’s neotype is conspecific with the species described 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 habitats and are found on coniferous trees in winter (notably Picea, Pinus, Taxus). The species 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), Schiemenz et al. (1996), Schwoerbel (1957), Wagner (1935, 1939a), HN; see also Günthart (1987a), Wagner & Franz (1961)
Balclutha rhenana W. Wagner, 1939 Described from near Mainz and considered to be conspecific with B. punctata (F.) in a revision of Nearctic Balclutha species by Blocker (1967), declaring a specimen of B. rhenana (W.Wg.) from the Baltic Sea island of Rügen as neotype. This opinion is rejected here (see above).
Adults I – XII, mainly M VII – M VI; adult, 1 gen. On Phalaris arundinacea usually in wet and rather eutrophic sites, usually near standing or running water (including ditches) as well as in abandoned grassland, probably hibernating in the grass layer. Widespread in Germany, in high frequency on the host (e.g. along the edge of the Bavarian Alps, up to at least 750 m a.s.l., in the valleys of the Danube, Neckar and Main, in Thuringia, southern Niedersachsen, the lower Elbe valley), but often overlooked or confused with the preceding species. Thus, the number of published records is low. From western parts of central Europe the species is only known from the Netherlands and Switzerland, but not from Belgium and France. Hence, localities along the upper and middle Rhine between Freiburg and Cologne may be at the edge of the range. The locus typicus is Heidesheim near Mainz. Fischer (1972), Frommer (pers. comm.), Reimer (1992), Schiemenz et al. (1996), SMNS, HN; see also Nast (1987)
Balclutha saltuella (Kirschbaum, 1868) So far only M VIII – E VIII; adult(?), 1(?) gen. In most parts of central Europe this species probably occurs as a temporary stray only, living on grasses on various sunny, usually more or less disturbed sites. Although described from the surroundings of Wiesbaden 130 years ago (from a single ‡), there are only three further and very recent German records, from Upper Bavaria, southern Niedersachsen and the Lake Constance Basin: Vorderriß, Isar floodplains, 800 m a.s.l., 27.VIII.1996, 1 †, 1 ‡, on gravel banks; Ebergötzen, 170 m a.s.l., 19.VIII.1996, 1 ‡, and
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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 Republic) and the middle Rhone valley (France). Kirschbaum (1868), Nickel (1999a); see also della Giustina (pers. comm.), Lauterer (pers. comm.), Ribaut (1952)
Macrosteles alpinus (Zetterstedt, 1828) M VII – E IX; egg, 1 gen. In central Europe, this species is confined to stands of tall sedges, in usually sunny, wet to boggy sites of the upper montane and subalpine belt of the Alps and Sudetes. In Germany it has only been recorded in 4 localities in the Allgäu and Bavarian Alps (Oberstdorf, Wannenkopf, 1650 m a.s.l., 1936; Oberstdorf, Geißalpsee, c. 1600 m a.s.l., 06.VIII.1995; Berchtesgaden, Priesberger Moos, 1350 m a.s.l., 10.VIII.1997, and Berchtesgaden, Funtensee, 1600 m a.s.l., August and September 1992, in Malaise traps). Densities in these sites were usually high. Has been found between 1160 and 2380 m a.s.l. in North Tyrol (Austria) and the Engadine (Switzerland). Otherwise it is reported from large parts of the arctic and boreal zone of Europe, Asia and North America. Wagner (1939a), Nickel (1999b), HN; see also Dlabola (1970), Günthart (1987a), Leising (1977), Nast (1972), Vilbaste (1980)
Macrosteles cristatus (Ribaut, 1927) E V – M X; egg, 2 gen. A pioneer species in fallow fields, ruderal sites, mining areas, along field paths and forest roads, on dry pond bottoms, in cereal crops, etc.; low numbers of individuals (some of which may be vagrants) are also found in moderately moist to moderately dry fertilized meadows and pastures and other habitats. Host plants are grasses and herbs, perhaps also species of Cyperaceae. The original habitats presumably include gravel and mud banks along rivers. Widespread all over Germany and particularly common in southern and middle parts. Breeding takes place up to at least 800 m a.s.l., but single individuals are frequently recorded at 1800 m a.s.l. in the Bavarian and Allgäu Alps. Has been shown to transmit some yellows diseases in various parts of Europe, but probably without economic relevance. Achtziger (1991), Fischer (1972), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Tharsen (1987), Wagner (1935), Nickel & Achtziger (1999), HN; see also Brˇcák (1979)
Macrosteles fieberi (Edwards, 1889) E V – B X; egg, 2 gen. In hollows and on floating mats in raised and intermediate bogs, nowadays mainly in old and overgrown peat diggings. The host plant is Eriophorum angustifolium. Literature records from other types of habitats such as salt marshes, sand pits, etc. should be revised. In Germany apparently confined to the northern half; at present, there are only
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6 verified records from the Weser-Ems region, North Friesland, eastern Holstein, the Dübener Heide and the Solling, up to 450 m a.s.l. Hildebrandt et al. (1998), Remane (1958), Schiemenz et al. (1996), Nickel (1997), HN
Macrosteles frontalis (Scott, 1875) E V – B X; egg, 2 gen., perhaps 1 at higher altitudes. In stands of horsetail in wet to moderately dry sites, usually along field paths and margins, railway embankments, in mining areas, but also in fens, intermediate bogs and along lake shores. The main host plants are Equisetum arvense and E. palustre, but the species is also reported to live on E. sylvaticum. Widespread in Germany and fairly common in the Alps and their foothills, rather sporadic elsewhere. The highest localities are at 1350 m a.s.l. in the Berchtesgaden Alps. Fischer (1972), Remane (1958), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a, 1951a), HN
Macrosteles horvathi (W. Wagner, 1935) Among populations of northern Germany and the Bavarian Alps, single specimens have been found which resemble M. nubilus Ossiannilsson, 1936 described from Sweden (Wagner 1939a, HN). Until further evidence has been presented, these are considered to be abnormal individuals of M. horvathi (W.Wg.). Ossiannilsson (1954) himself was not convinced of the specific distinctiveness of M. nubilus Oss.
E V – M X; egg, 2 gen., according to Leising (1977) 1 in the North Tyrolian Alps (Austria). A hygrophilous pioneer species of disturbed and wet, usually waterlogged, seepy or slightly saline sites, often in gravel, sand and clay pits, clearings, along temporarily wet shores of lakes and ponds, also in spring mires with predominating rushes and moderately saline wet meadows and pastures (near the coast of the North and Baltic Sea as well as inland). Host plants are rushes (Juncus articulatus, J. gerardii, probably also J. bufonius and additional species). Widespread in Germany, but only scattered, recorded up to 1500 m a.s.l. in the Bavarian Alps. In North Tyrol (Austria) and the Swiss Alps large populations have been found in the subalpine belt, low numbers even up to 2250 m a.s.l. Fischer (1972), Niedringhaus (1991), Reimer (1992), Schiemenz et al. (1996), Wagner (1935), HN; see also Günthart (1987a), Leising (1977)
Macrosteles laevis (Ribaut, 1927) M V – M X; egg, 2 gen., perhaps 1 at higher altitudes. Another pioneer species of sparsely vegetated mud banks along rivers and lakes, secondarily, and often in high abundances, in fertilized meadows, ruderal sites, on temporarily dry ponds bottoms, in cereal crops and mining areas. Lives polyphagously on grasses, rushes, sedges, and also dicotyledonous herbs. Widespread in Germany and particularly abundant in the southern half (although with strong annual fluctuations of numbers), but with decreasing density towards the north. Breeding takes place at least up to 900 m a.s.l. in the Mittelgebirge, single individuals have been found at 1800 m a.s.l.
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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 organisms from various European countries, notably European aster yellows (EAY), stolbur, 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 Andrzejewska (1962), Brˇcák (1979), Leising (1977), Ossiannilsson (1983)
Macrosteles lividus (Edwards, 1894) B VI – E IX; egg, 2 gen. In temporarily or permanently flooded, mesotrophic to moderately eutrophic, usually sunny sites, notably shores of ponds, sand and gravel pits, as well as moderately saline marshes, inland and near the coast. The host plant is Eleocharis palustris (probably including E. uniglumis). Rather scattered in Germany, apparently at the southwestern edge of the range, although probably under-recorded. Altogether there are c. 20 sites, mainly along the coasts of the North and Baltic Sea and in the eastern half; westernmost localities, most of which appear to be rather isolated, are on the island of Borkum, near Hanover, Göttingen, Bad Brückenau (Rhön Mountains) and Vorderriß (upper Isar, 800 m a.s.l.). Is also reported from England, the Netherlands and northern France, but not from Belgium and Switzerland. Niedringhaus & Olthoff (1993), Remane (pers. comm.), Schiemenz et al. (1996), Wagner (1939a, 1941b), Walter (pers. comm.), Nickel & Remane (1996), Nickel (1997, 1999a), HN; see also della Giustina & Remane (2001)
Macrosteles maculosus (Then, 1897) E V – B X; egg, 2 gen. In sunny, damp to moderately dry, usually sandy or gravelly sites of early successional stages with sparse or low-growing vegetation cover, mainly trampled patches, ruderal sites, gravel and paved sites (even within cities). The host plant is Polygonum aviculare. In Germany apparently at the northwestern edge of the range, known from a few records in the southeastern half only. Border localities are near Balingen (Swabian Jura), Stuttgart, Wetzlar, Marburg, Göttingen, Salzgitter, Helmstedt, Quedlinburg, Halle, Leipzig and Meißen; is also reported from near Stettin (Poland). The highest localities are at 700 m a.s.l. in the Rhön Mountains, at 1000 m a.s.l. in the Engadine (Switzerland). Fröhlich (1996a), Rabeler (1952), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Wagner (1941a), Nickel (1994), HN; see also Günthart (1987a)
Macrosteles oshanini Razviazkina, 1957 So far M VIII – M IX, in England according to Le Quesne (1969) VI; egg, 2(?) gen. Most records of this species are from the eastern half of Europe. It lives in wet sites, preferentially fens with strongly fluctuating water tables or moderate salinity, also near shady forest margins. The host plants are unknown, but may include species of Cyper-
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aceae or Poaceae. In Germany found only in 6 localities in Holstein (Steinbek), Mecklenburg (Hagenow, Leussow, Neukloster), Brandenburg (Trechwitz) and Thuringia (Jena), all below c. 150 m a.s.l. Kuntze (1937), Müller (pers. comm.), ZIMH, HN; see also Ossiannilsson (1983)
Macrosteles ossiannilssoni Lindberg, 1954 B VI – B X; egg, 1 - 2 gen. In wet, usually peaty sites (intermediate bogs, spring mires, peaty meadows and pastures). The host plants are probably Carex species and Rhynchospora alba, perhaps also Juncus spp. and grasses. Vagrants may be encountered far away from typical habitats. In Germany autochthonous populations are known from scattered sites in the northwestern plains as well as in the Erzgebirge, Frankenwald, Bavarian and Black Forest, and the Bavarian Alps. Found up to at least 1500 m a.s.l. in Upper Bavaria, up to 2370 m a.s.l. in the Engadine (Switzerland). Niedringhaus & Olthoff (1993), Reimer (1992), Walter (1998), Nickel & Achtziger (1999), Nickel & Sander (1996), Nickel (1999b), HN; see also della Giustina (1989), Günthart (1987a, 1987b), Leising (1977), Nast (1976a)
Macrosteles quadripunctulatus (Kirschbaum, 1868) E V – B X; egg, 2 gen. In sunny, disturbed, usually sandy and sparsely vegetated, moderately dry to dry sites (sand pits, abandoned fields, military training areas, waysides, trampled patches in dry grassland, etc.), in warmer regions of southern Germany also on loamy substrates (notably in vineyards). Setaria and Panicum, as well as Corispermum have been reported as host plants. The species is uncommon in Germany, rather localized, largely confined to sand and viticultural regions (up to at most 300 m a.s.l.); it has been recorded in the north German plain, middle Hessen, the Thuringian and Leipzig Basin, along the upper and middle Rhine, in the Main valley and Mittelfranken. The locus typicus is Mombach near Mainz. Is reported as a potential vector of yellows diseases in other countries, but is unlikely to cause major effects in central Europe due to the scattered occurrence. Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Remane (1987), Schiemenz et al. (1996), Trümbach (1959), Wagner (1935), HN
Macrosteles sardus Ribaut, 1948 So far E V – E IX; egg, 2 gen. Usually in moist, moderately shady sites in river floodplains and along lake shores. Is reported to live on grasses in Kazakhstan, but no further details are known on habitat and host plants. In Germany known only from a few records from eastern Holstein (near Plön and Lübeck), the lower course of the Elbe (near Lüneburg) and the upper Rhine plain (near Freiburg, Speyer, Heidelberg and Darmstadt), all below 200 m a.s.l., but found at 700 m a.s.l. in Trentino (northern Italy). Remane (1962), Remane & Fröhlich (1994b), Nickel (1997), HN; see also Mityaev (1971)
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Macrosteles septemnotatus (Fallén, 1806) B VI – E X; egg, 2 gen. In wet, sunny to moderately shady sites, mainly abandoned grassland, fens, shores of running and standing water, and fen woods. The host plant is Filipendula ulmaria. Widespread in Germany, fairly common, in intermediate to high frequency on the host, recorded up to 1200 m a.s.l. in the Allgäu Alps. Fischer (1972), Heller (1987a), Nikusch (1976), Reimer (1992), Schiemenz et al. (1996), Wagner (1935), Nickel & Achtziger (1999), HN
Macrosteles sexnotatus (Fallén, 1806) M V – M X; egg, 2 gen. (see also Witsack 1985). The life history is very much like in M. cristatus (Rib.) and M. laevis (Rib.). M. sexnotatus (Fall.) frequently occurs syntopically with these as a pioneer species in fertilized meadows and pastures, as well as in cultivated crops, but in general, it is more hygrophilous and often dominant in moist, eutrophic meadows, moist ruderal sites, and on mud banks with sparse vegetation, also in moderately saline grassland. Migrating individuals are sometimes found in large numbers in other habitats. Host plants are Poaceae, Cyperaceae, and probably also dicotyledonous herbs. Widespread in Germany and very common; frequently up to at least 1200 m a.s.l. in the Bavarian and Allgäu Alps, at least occasionally up to 1600 m a.s.l. Achtziger (1991), Fischer (1972), Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992), Schiemenz et al. (1996), Wagner (1935), Walter (1996, 1998), Nickel & Achtziger (1999), HN, and others
Macrosteles sordidipennis (Stål, 1858) E V – E IX; egg, 2 gen. In salt marshes along the coast and inland. The main host plant is Puccinellia distans, but P. maritima and Juncus gerardii have also been reported. In Germany known from the East Frisian Islands, the North Sea coast between Emden and Cuxhaven, the Baltic Sea coast near Heiligenhafen, as well as from inland salt marshes in Holstein, Saxony-Anhalt, Thuringia, middle Hessen and eastern Niedersachsen, also from sites heavily affected by phosphate immissions near Jena. The highest localities are at c. 250 m a.s.l. Fröhlich (1996a, and pers. comm.), Hildebrandt (1995), Müller (1985), Niedringhaus & Olthoff (1993), Schiemenz et al. (1996), Wagner (1937a), HN; see also Linnavuori (1952)
Macrosteles variatus (Fallén, 1806) B VI – E X; egg, 2 gen. In eutrophic, moderately moist to moderately wet, usually moderately shady sites with lush herb layer, mainly along shores and margins in floodplains of rivers and streams, along forest roads, shaded ditches, etc., preferentially on Urtica dioica, perhaps also on other species of dicotyledonous herbs. Widespread in Germany, fairly common in most
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valley bottoms, but often overlooked; recorded up to at least 800 m a.s.l. in the Mittelgebirge and at the edge of the Alps. Fischer (1972), Heller (1987a), Mölleken & Topp (1997), Niedringhaus (1997), Reimer (1992), Remane (pers. comm.), Lauterborn (1920), Schiemenz et al. (1996), Trümbach (1959), Wagner (1935), HN
Macrosteles viridigriseus (Edwards, 1922) E V – M X; egg, 2 gen. Mainly in temporarily dry or moderately saline pastures and meadows (often in river floodplains or floodplain depressions, occasionally in intensively managed sites), also in rather low-vegetated spring mires and fens and, as a pioneer species, on sparsely vegetated banks of gravel, sand and mud. Records from other habitats probably refer to vagrants. Host plants are various grasses, probably also Cyperaceae. Has been found in most parts of Germany, but often in very low numbers only. Records of large and reproducing populations are more localized, but have been found up to 850 m a.s.l. in the Thuringian Forest, up to 1250 m a.s.l. in Austria. Has been reported as a vector of Clover phyllody (CP) in Britain, but apparently without economic importance. Emmrich (1966), Marchand (1953), Niedringhaus & Olthoff (1993), Schiemenz et al. (1996), Walter (1996), Nickel & Achtziger (1999), HN; see also Brˇcák (1979), Wagner & Franz (1961)
Erotettix cyane (Boheman, 1845) So far M VII – M IX; egg, 1 gen. On floating water plants, usually on small ponds and lakes in cool and moderately shady situations. Host plants are Potamogeton natans, Nuphar lutea, Nymphaea alba, and probably additional species of floating plants. Has been reported from Trapa natans and Marsilea quadrifolia in Austria. In Germany only very sporadic, known from about 20 localities, mostly to the east of a line running from Schwerin to Kassel, Würzburg and Munich, recorded up to 600 m a.s.l. Further localities, which appear to be isolated, are situated in the Weser-Ems region and the southern upper Rhine plain, although the species is also reported from the Netherlands, Belgium and France. Funke & Witsack (1998), Coll. Förster (Niedringhaus pers. comm.), Schiemenz et al. (1996), Walter (1996), Nickel & Remane (1996), HN; see also Holzinger (1995b)
Sonronius binotatus (J. Sahlberg, 1871) M VII – B IX; egg, 1 gen. This species is widespread in the Siberian and north European taiga zone, with rather isolated populations in central European mountain regions. It lives in forest clearings and along roads on Epilobium angustifolium; at least adults are also found on other species of Epilobium. In Germany only known from altogether 4 localities in the Bavarian Alps (Mittenwald), the Vogtland (near Plauen) and the Rhön Mountains (to the east of Fulda), between 500 and 1000 m a.s.l. Bittner & Remane (1977), Haupt (1925), Reimer (1992), Walter & Emmrich (1995), Nickel (1999b)
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Sonronius dahlbomi (Zetterstedt, 1840) M VII – B IX; egg, 1 gen. Known from Germany only from altogether 6 records in the Allgäu and Bavarian Alps (in the vicinity of Oberstdorf, Mittenwald and Lenggries, between 700 and 1100 m a.s.l.). Found on Epilobium angustifolium in cool and moderately shady sites, usually in or near woodland; from Sweden Rubus idaeus, Alchemilla spec. and Filipendula ulmaria have been reported as additional food plants. Haupt (1925), Remane (pers. comm.), Wagner (1939a); see also Ossiannilsson (1983)
Sagatus punctifrons (Fallén, 1826) M VI – M IX; egg, 1 gen. Along rivers and streams, on gravel banks, dunes and in mining areas, on narrow-leaved species of willows; most individuals (including nymphs) found on Salix purpurea, S. triandra and S. viminalis, low numbers also on other species (S. pentandra, S. fragilis, S. alba). Furthermore, S. repens is the main host in coastal areas, S. eleagnos in the Alps and their foothills. Widespread at lower and middle altitudes of Germany (up to at least 950 m a.s.l.), in intermediate to high frequency, locally abundant in favourable sites. Fischer (1972), Frommer (pers. comm.), Niedringhaus (1991), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a), HN
Deltocephalus maculiceps Boheman, 1847 E VI – M IX; egg, 1 gen. This is a western European species of intermediate and heathland bogs. Eriophorum vaginatum has been reported as host plant, but there is more evidence that the species lives on Molinia caerulea. It is rare in Germany and restricted to northern parts, recorded from the Weser-Ems area, the Lüneburger Heide, Holstein, Mecklenburg, the Niederlausitz and the vicinity of Halle, up to c. 100 m a.s.l. Otherwise it is only reported from the island of Gotland (Sweden), Denmark, Ireland, England, the Netherlands, northwestern Poland, France and Spain; records from other countries are doubtful. Remane (1958, and pers. comm.), Schiemenz et al. (1996), Wagner (1935), HN; see also Le Quesne (1969), Nast (1976a, 1987), Ossiannilsson (1983)
Deltocephalus pulicaris (Fallén, 1806) M V – E X; egg, 2 gen., according to Leising (1977) 1 in the North Tyrolian Alps (Austria). In sunny to moderately shady, wet to moderately dry sites, often extremely abundant in low-vegetated sheep and cattle pastures, fertilized meadows, in lower numbers also along forest roads, in fens, moderately saline habitats and lawns in parks, gardens and even sports grounds. Host plants are various grasses (Agrostis, Poa, Dactylis, Elymus, Lolium and others). Widespread in Germany, very common in most parts, also in the Mittelgebirge and the Alps, where frequently found up to at least 1500 m a.s.l. In Austria
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and Switzerland large populations occur at 2100 m a.s.l., at least single individuals were recorded at 2510 m a.s.l. Fischer (1972), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others; see also Günthart (1984, 1987a), Leising (1977), Wagner & Franz (1961)
Recilia coronifer (Marshall, 1866) E VI – M X; egg, 1 gen. In moderately shady, occasionally also sunny, wet to damp, usually acidic sites, mainly open oak and pine forests (often in clearings, along margins and roads), bog margins and straw meadows. Host plants are Holcus mollis and, less frequently, Molinia caerulea. Widespread in the lowlands of Germany (up to at least 600 m a.s.l.), but usually in low individual numbers, at the northern edge of the range in Schleswig-Holstein; northernmost localities are near Heide, Rendsburg and Eckernförde. Achtziger (1991), K. Heller (pers. comm.), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1958, and pers. comm.), Schiemenz et al. (1996), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN
Recilia horvathi (Then, 1896) So far E VIII – E IX; egg(?), 1(?) gen. This species’ range extends from Mongolia to central Europe, where it occurs in rather isolated populations in xerothermic sites. In Germany there are only two known localities in the northern upper Rhine plain near Heidelberg: Pferdstrieb and Pflege Schönau, both near Sandhausen, c. 110 m a.s.l., 26.VIII.1981 and 27.IX.1982, several ††, on inland sand dunes on Corynephorus canescens. There are further records from Moravia (Czech Republic), Styria (Austria), Slowenia, northern Italy and Ticino (Switzerland). Heller (1996); also Lauterer (pers. comm.), Then (1896), Remane & Fröhlich (1994b)
Recilia schmidtgeni (W. Wagner, 1939) So far E VIII – X; egg(?), 1(?) gen. Like the preceding species, known in Germany only from two localities in sand areas of the northern upper Rhine plain: Mainz, 24.VIII.1951, 2 ††, „in einem Pflanzgarten bei Uhlerborn“, and Dudenhofen near Speyer, Oktober 1989, „in Anzahl in Cynodon-Beständen in feuchteren Senken am Rand eines Dünengebietes“. Has been recorded in moderately saline sites in eastern parts of Austria. In the Mediterranean region the species lives in various open habitats, often in disturbed patches. Remane & Fröhlich (1994b), Wonn (1956); see also Fröhlich (1996a)
Endria nebulosa (Ball, 1900) B VII – M IX; egg, 1 gen.
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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. Moreover, it is very difficult to collect due to its epigeic life habits; thus it is probably much under-recorded in most parts of its range. Funke & Witsack (1998), Remane (1961c), Schiemenz et al. (1996), Nickel (1997), HN; see also Arzone et al. (1987), Lauterer (1980)
Doratura exilis Horváth, 1903 B VI – E IX; egg, 1 gen., perhaps 2 in southern Germany or in warm years. On usually grey-leaved subspecies of Festuca ovina in oligotrophic, sunny, low-vegetated, often strongly grazed sites on dry, basic as well as acidic substrates. Widespread in the lowlands of Germany, but only scattered, locally dominant on sandy dry grassland of the north German plain and on calcareous hillsides in the middle parts. Not recorded to the south of the Danube, but found up to at least 750 m a.s.l. on the Swabian Jura. The northernmost localities are near Hamburg and Lauenburg; the distribution border apparently runs through southern parts of Holstein. Not reported from Schleswig, Denmark and Scandinavia, except southern Sweden. Reimer (1992), Remane (1958, 1987), Schiemenz et al. (1996), SMNS, Trümbach (1959), Wagner (1935, 1951a), HN; see also Ossiannilsson (1983)
Doratura stylata (Boheman, 1847) B VI – M X, in warmer regions M V – XI; egg, 1 gen., perhaps 2 in southwestern Germany. In oligotrophic and sunny, dry to damp, occasionally also moist sites, mainly low-productivity grassland, waysides and ruderal sites, often dominant in meadows and pastures on nutrient-poor substrates of the submontane belt. The main host plants are Festuca rubra, F. ovina and Agrostis capillaris, probably also additional species of fine-leaved grasses, e.g. Poa pratensis angustifolia and Nardus stricta. Widespread in Germany and particularly common in regions with less intensive agriculture, found up to at least 1200 m a.s.l. in the Black Forest, but frequently in the alpine belt (up to 2200 m a.s.l.) of the Austrian and Swiss Alps. Achtziger (1991), Fischer (1972), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Remane (1987), Schiemenz (1969), Schiemenz et al. (1996), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others; see also Günthart (1987a), Leising (1977), Wagner & Franz (1961)
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Doratura horvathi W. Wagner, 1939 M VI – B X; egg, 1 gen. In sunny, moderately dry to dry, basic (occasionally also acidic), usually grazed sites with low-growing and sparse grass cover. The host plant is Helictotrichon pratensis. In Germany at the northern edge of the range, only scattered, but locally dominant, notably in regions of limestone, gypsum and porphyry of middle parts, between the foothills of the Alps and the northern edge of the Mittelgebirge (Eifel Mountains, southern Niedersachsen, eastern Harz Mountains, Thuringia, Rhine-Nahe region, Main and Rhön region, Tauber valley and Franconian Jura, also foreland of the Alps and Kaiserstuhl). The northernmost localities are near Mechernich, Brilon, Göttingen, Halberstadt, Halle and Jena, up to at least 700 m a.s.l. Otherwise the species is reported only from central Poland, Moravia (Czech Republic), Slovakia, Austria and Italy. Reimer (1992), Rombach (1999a, 1999b), Schiemenz et al. (1996), SMNS, Wagner (1939a, 1951a), ZIMH, Nickel (1994), HN; see also D’Urso (1995), Holzinger & Remane (1994), Lauterer (1984), Nast (1976a)
Doratura impudica Horváth, 1897 M VI – E IX; egg, 1 gen. Among sparse stands of tall grasses, usually on acidic, sandy to gravelly soils in moderately dry to dry, often moderately shady sites, mainly in clearings of pine forests, in the vicinity of inland dunes and in sandy ruderal sites. The host plant is usually Calamagrostis epigejos, perhaps also Elymus spp. (in northern Italy Elymus athericus). In Germany with strongholds in the northeastern plains, where locally common in favourable sites, but perhaps at the western edge of the range. Westernmost localities are on the island of Amrum, near Lüneburg, Halle, Leipzig, Dresden and Görlitz, up to at most 200 m a.s.l. Published records from the Mittelgebirge refer to stray or misidentified individuals. Is also reported to occur in northern and eastern parts of France and in the Netherlands. Bartels et al. (1992), Funke & Witsack (1998), Remane (pers. comm.), Schiemenz (1969), Schiemenz et al. (1996), Walter (pers. comm.), Nickel (1997), HN; see also Della Giustina (1989), Gravestein (1976)
Doratura homophyla (Flor, 1861) M V – M X; egg, 2 gen. In sunny, sparsely vegetated grass stands in damp to dry, moderately eutrophic, often disturbed sites, usually on sandy, occasionally also loamy substrates (disturbed patches in dry grassland, abandoned fields, low-productivity pastures of cattle or sheep, waysides, military training areas, etc.). Host plants are grasses (Agrostis capillaris, probably also Elymus repens, Festuca rubra, Poa compressa and others). Locally common in the lowlands of Germany (up to at most 400 m a.s.l.) and largely confined to sand and gypsum regions (north German plain, upper and middle Rhine, valleys of the Main, Danube and Regnitz, Thuringian and Leipzig Basin, Oberlausitz). Frommer (1996), Niedringhaus (1997), Remane (1987), Schiemenz et al. (1996), Trümbach (1959), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN
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Doratura littoralis Kuntze, 1937 Was originally described as a variety of D. homophyla (Fl.), after specimens from near Rostock, but treated as a distinct species by Wagner (1941b), and subsequently, Dworakowska (1968). However, the morphological differences appear to be very small; after a revision of more material they may turn out to fall within the range of intraspecific variation of D. homophyla (Fl.).
So far VI – VIII; egg, 1(?) gen. Altogether known only from very few records from Heligoland, eastern Holstein (Hohwachter Bucht), Mecklenburg (between Rostock and Ahrenshoop) and the lower Oder valley (to the south of Stettin, Poland), mainly on white and grey dunes near the coast; the Polish locality, however, is an inland dune. Ammophila arenaria has been suspected to be the host plant. Kuntze (1937), Niedringhaus & Olthoff (1993), Schiemenz et al. (1996), Wagner (1941b); for identification see Dworakowska (1968)
Platymetopius undatus (De Geer, 1773) and Platymetopius henribauti Dlabola, 1961 P. henribauti Dlab. was described after highly variable characters (shape of forehead and 7th sternite of ‡‡), with contradictory information on food plants, without revising the type material 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 conclusion, 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 vegetation (Helianthemum and other species) or from oak and birch. Apparently, there was a dramatic decline during recent decades. Kirschbaum (1868), Remane (1987), Schiemenz et al. (1996), Wagner (1935, 1939a), ZIMH, HN
Platymetopius major (Kirschbaum, 1868) B VI – M X; egg, 1 gen., 2 in southern Germany. Along sunny woodland margins in damp to dry, basic to acidic sites (dry grassland with scattered trees and shrubs, spatially diverse forest margins, etc.). Nymphs are found among low vegetation (locally in cushions of Helianthemum nummularium, but also on other plants), adults ascend various species of deciduous trees and shrubs (mainly Quercus, Betula, Acer, also Crataegus, Prunus, Fagus and others). Widespread in middle and southern parts of Germany and fairly common in warmer regions, but rather localized in the north, where recorded from Holstein, the Wendland of Niedersachsen and Mecklenburg (although apparently absent from the Weser-Ems area). To the north of it, there
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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 hillsides of the Bavarian Alps. Fischer (1972), Post-Plangg & Hoffmann (1982), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN; see also Ossiannilsson (1983)
Platymetopius guttatus Fieber, 1869 So far B VII – E VIII; egg, 1 gen. In open xerothermic forests and along their margins, mainly in sun-exposed slope or plateau situations. Adults are usually found on Betula pendula and Quercus; the food plants of nymphs are unknown, but are likely to include low-growing herbs. From Germany known only from about 15 scattered localities in the valleys of the Nahe, Main and Franconian Saale, and in warmer regions of Thuringia and Saxony-Anhalt (near Nordhausen, Bad Frankenhausen, Querfurt, Jena, Gera and Rudolstadt), up to at most 400 m a.s.l., but the species may be under-recorded. Not reported from north Germany, but from southern parts of Norway and Sweden. Perner (1997), Schiemenz et al. (1996), Wagner (1939a), Wagner & Franz (1961), Nickel & Remane (1996), HN; see also Anufriev & Kirillova (1998), Ossiannilsson (1983)
Idiodonus cruentatus (Panzer, 1799) B VII – M X; egg, 1 gen. In temporarily moist to moderately dry, basic as well as acidic sites, usually in open stands of trees or shrubs with varying proportions of evergreens. Nymphs are found among low-growing vegetation, most adults have been swept from Tilia, Betula, Fagus, Rubus, Vaccinium, Calluna, and additional species of woody plants and dwarf shrubs. Widespread and locally common in the southern half of Germany, mainly in open pine and spruce forests in areas of Jurassic and Triassic limestone, and among subalpine scrub, up to at least 1750 m a.s.l. in the Bavarian Alps, up to 2200 m a.s.l. in alpine dwarf shrub heaths of North Tyrol (Austria) and the Engadine (Switzerland); rather scattered in the northern half, where mainly confined to raised bog margins, occasionally in forest clearings and glades. Fischer (1972), Reimer (1992), Rombach (1999a), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a), HN; see also Günthart (1987a), Leising (1977)
Colladonus torneellus (Zetterstedt, 1828) B V – E VIII, in the Czech Republic and Slovakia according to Lauterer (1980) M IV – B VII; nymph, 1 gen. In mixed and coniferous forests (usually under Picea or Pinus) in cool sites with shrubby undergrowth, also reported from deciduous forests in the Czech Republic and Slovakia. Not much is known on the species’ life history; perhaps it is a vertical migrant which quickly ascends into the canopy layer after emergence. Recorded in larger numbers from Melica
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uniflora in Moravia (Czech Republic), also on various shrubs and dwarf shrubs (Salix, Betula, 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), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), HN; see also Günthart (1987a), Lauterer (pers. comm.), Ossiannilsson (1983), Ribaut (1952)
Lamprotettix nitidulus (Fabricius, 1787) B VII – E X; egg, 1 gen. In deciduous forests in damp to moist, usually eutrophic sites, mainly flood-plain and fen woods, as well as oak-hornbeam forests. Nymphs are found in low-growing herbaceous vegetation (often among Poaceae, Lamium or Urtica), adults after emergence quickly ascend various deciduous trees (Quercus, Alnus, Betula, Ulmus, Tilia and others). Widespread in the lowlands of Germany, but seldom found (although easily recorded in numbers with suitable methods such as tree eclectors, light traps, and sweeping of nymphs on the forest floor). Highest localities are at 600 m a.s.l. in the Mittelgebirge and the foothills of the Alps. Büchs (1988), Frommer (1996), Kupka (1925), Niedringhaus & Olthoff (1993), Reimer (1992), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a), HN
Allygus communis (Ferrari, 1882) Was originally described as a variety of the following species, and for a long time considered to be conspecific with it, until Ossiannilsson (1983) pointed out that there are considerable differences in genitalic structures in both sexes. Thus, a number of older records of A. mixtus (F.) may refer to A. communis (Ferr.).
M VI – M X; egg, 1 gen. Along sunny forest margins and in semi-open stands of trees; in damp to dry sites, locally syntopic with A. mixtus (F.), but generally preferring warmer and drier habitats. Nymphs are found among low-growing herbaceous vegetation, presumably on grasses and dicotyledons, adults usually on Quercus and Betula. Widespread in the lowlands of Germany, but less common than the following species, and with strongholds in rather warm regions of low altitudes, recorded only up to 600 m a.s.l. in the Mittelgebirge and the foothills of the Alps. Günthart (1988), Kolbe & Bruns (1988), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987), Schönitzer & Oesterling (1998b), SMNS, ZIMH, Nickel & Remane (1996), Nickel (1997), HN
Allygus mixtus (Fabricius, 1794) Most older records are in need of revision and may refer to A. communis (Ferr.) (see above).
E VI – M X; egg, 1 gen.
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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, Frangula, Pinus and others). Widespread in Germany and fairly common in many places, frequently found up to 1300 m a.s.l. in the Mittelgebirge and the Bavarian Alps. Achtziger (1991), Frommer (1996), Günthart (1988), Niedringhaus (1991, 1997), Reimer (1992), Remane (1987), Schönitzer & Oesterling (1998b), SMNS, Nickel (1997), HN
Allygus maculatus Ribaut, 1952 B VII – B X; egg, 1 gen. In oak forests in basic to acidic, temporarily moist to dry sites of lower altitudes (recorded up to 450 m a.s.l. only). Nymphal development takes place in the herb layer (mainly on grasses), most adults are swept from oaks (Quercus robur and Qu. petraea). In Germany widespread, but seldom found (at present c. 30 localities only), mainly between the Danube and the northern edge of the Mittelgebirge, also near Berlin and Ratzeburg. To the north of it, there are only two old reports from Jutland and southern Sweden. Most sweep net records comprise only single individuals, larger numbers have been collected in light traps. Fischer (1972), Rabeler (1962), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Schwoerbel (1957), HN; see also Ossiannilsson (1983)
Allygus modestus Scott, 1876 M VI – B X; egg, 1 gen. In open deciduous forests and semi-open cultivated habitats (along forest margins, alleys, etc.), usually in moderately wet to damp, occasionally also moderately dry sites. Nymphs feed on grasses (but perhaps also on dicotyledonous herbs), adults on Quercus, Ulmus, Alnus, Betula, Acer, Prunus and other deciduous trees. Widespread in Germany, including the northern plains, but apparently confined to the lowlands (up to at least 550 m a.s.l. in the foreland of the Alps). North German localities are near the northern border of the range, which runs through England, Jutland, southern Sweden and northwestern Poland. Frommer (1996), Günthart (1988), Niedringhaus & Bröring (1988), Remane (1987), Schiemenz al. (1996), SMNS, Trümbach (1959), Wagner (1935, 1939a, 1951a), HN; see also Gillerfors (2002), Le Quesne (1969), Nast (1976a), Ossiannilsson (1983)
Allygidius commutatus (Fieber, 1872) M VI – B X; egg, 1 gen. Along sunny forest margins and in open stands of deciduous and coniferous trees of moderately moist to dry, basic to acidic sites, preferentially in warm situations. Host plants are mainly tall grasses (Brachypodium pinnatum, Calamagrostis arundinacea, C. epigejos, Melica uniflora and others), at least some adults are found on deciduous trees (Quercus, Ulmus, Betula and others). Widespread and fairly common in middle and southern
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parts of Germany (although usually collected in small numbers only), up to at least 800 m a.s.l. in the Mittelgebirge and the foothills of the Alps, with decreasing density of localities in the north German plain; apparently absent from the Weser-Ems area. However, the species is reported to be widespread in southern Scandinavia. Achtziger (1991), Fischer (1972), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN; see also Ossiannilsson (1983)
Allygidius abbreviatus (Lethierry, 1878) M VI – B X; egg, 1 gen. In basic to acidic, xerothermic sites, usually in the vicinity of shrubs or trees. Nymphs are found in low-growing herbaceous vegetation; at least some adults ascend various deciduous trees. In Germany confined to southwestern parts. The status in Germany has long been uncertain, since there was only a single proved but inaccurate record („Rheinhessen“). Further records dating before 1940 are not verified or were corrected later. Has recently been found in 5 localities in the northern Eifel Mountains and along the Palatinate Haardt (near Bad Münstereifel, Grünstadt and Neustadt an der Weinstraße, all between 150 and 350 m a.s.l.). Otherwise only reported from Belgium, France, the Czech Republic, Slovakia, Hungary, Austria and Italy. Rombach (1999a), Wagner & Franz (1961), HN; see also Puton (1886), Wagner (1935, 1941b), Hegab et al. (1980) (see this paper also for identification of nymphs), Lauterer (pers. comm.), Nast (1987)
Allygidius atomarius (Fabricius, 1794) Mainly B VI – M IX, according to Schiemenz et al. (1996) M V – E X; egg, 1 gen. In warm, moderately moist to moderately dry sites with semi-open stands of trees, mainly in river floodplains, along forest margins, in coppice-with-standards, etc. The nymphs live among herbaceous vegetation, probably on tall grasses, most adults are swept from Ulmus, Quercus and other deciduous trees. In Germany at the northern edge of the range, found only sporadically and in low numbers, at lower altitudes (up to at least 600 m a.s.l.), mainly in the larger river valleys (Rhine, Main, Elbe, Saale, Unstrut, Havel, Oder and others). Northernmost localities are near Cologne, Göttingen, Boizenburg, Stendal, Oranienburg and Frankfurt an der Oder. Is perhaps under-recorded. Achtziger (1991), Frommer (1996), Remane (1987), Wagner (1935, 1939a), Walter (1998), Nickel (1997), HN
Graphocraerus ventralis (Fallén, 1806) M V – B IX; egg, 1 gen. In sunny, usually oligotrophic, moderately dry to damp, occasionally also moist or saline sites, mainly low-input meadows and pastures, ruderal sites, grey and inland dunes, abandoned vineyards and waysides. Feeds on various grasses (probably Anthoxanthum odoratum, Poa pratensis and others. In field choice experiments in Britain, adults and nymphs fed on Arrhenatherum elatius, Helictotrichon pubescens and Trisetum flavescens). Widespread in Germany and locally common, but rather scattered in regions of inten-
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sive agriculture. Found up to at least 900 m a.s.l. in the Mittelgebirge, up to 2000 m a.s.l. in the French Alps. Achtziger (1991), Fischer (1972), Fröhlich (1996a), Kuntze (1937), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Remane (1987), Schaefer (1973), Schiemenz (1969), Schiemenz et al. (1996), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN, and others; see also Cook (1996), della Giustina (1989)
Mimallygus lacteinervis (Kirschbaum, 1868) So far M VIII – M IX, according to Moosbrugger (1946) in Vorarlberg (Austria) E VII; egg, 1 gen. On low-growing willows (Salix purpurea, probably also S. eleagnos) on almost bare gravel banks of alpine rivers. Older reports of Hippophae rhamnoides as host plant presumably refer to Salix eleagnos. Known from Germany only from the Iller near Oberstdorf, the Isar between Mittenwald and Fall, and the Friedergries near Garmisch-Partenkirchen; all sites between 750 and 900 m a.s.l. The Iller population, which has been recorded in 1937, is probably extinct due to drainage. Franz (1943), MTD, Nickel & Voith (unpublished data), Nickel (1999a); see also Kirschbaum (1868), Moosbrugger (1946)
Phlepsius intricatus (Herrich-Schäffer, 1838) B VIII – B V; adult (only ‡‡?), 1 gen. This species is reported to occur from central Asia to the Mediterranean region, known from Germany only from several records in the Kaiserstuhl (Badberg near Oberbergen, c. 400 m a.s.l.), the latest one dating from 1977. Lives in xerothermic grassland sites with scattered shrubs; adults are at least occasionally swept from shrubs, but the food plants are unknown. Remane (pers. comm.), Wagner (1963), ZIMH
Phlepsius ornatus (Perris, 1857) So far E VII – M IV; adult, 1 gen. In Germany known only from altogether 4 localities in the Nahe valley (Lemberg, Bad Münster, Ebernburg, Schloßböckelheim, all between c. 150 and 300 m a.s.l.) in xerothermic, acidic porphyry sites with scattered shrubs. Most adults were swept in low-growing herbaceous vegetation, a few also from shrubs. The food plants are unknown. Jöst (1966), Wagner (1963), HN
Rhytistylus proceps (Kirschbaum, 1868) E VI – M X, according to Schiemenz et al. (1996) from B VI onwards; egg, 1 gen. In sparse and low-growing stands of grasses, in sunny, moderately dry to dry sites (mainly sandy dry grassland, heaths, open margins of pine forests, occasionally chalk and sandstone hillsides, etc.). The main host plant is Festuca ovina; additional species reported in the literature include Corynephorus canescens, Nardus stricta and Carex arenaria, but
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these are probably erroneous. The species is widespread and fairly common in sandy areas of the north German plain, but more localized to the south of it, being largely restricted to areas of aeolian sand in larger river valleys in southern parts (along the upper Rhine, the Main and the Danube); the highest localities are at c. 500 m a.s.l. only. Niedringhaus (1991, 1997), Post-Plangg & Hoffmann (1982), Remane (1987), Schiemenz (1969), Schiemenz et al. (1996), Wagner (1935, 1939a), Nickel & Remane (1996), HN; see also Ossiannilsson (1983)
Hardya melanopsis (Hardy, 1850) Is reported from mountainous regions of western and southern European countries (Nast 1987), also from arctic and subarctic parts of Siberia (Anufriev & Emelyanov 1988), although those population have been described as H. taimyrica Vilbaste, 1969. In general, the taxonomy of European populations may be critical.
So far only B VIII, in England according to Le Quesne (1969) VII – V; adult, 1 gen. Known in Germany only from a single site in the upper montane belt of the southern Black Forest: Schauinsland, Steinwasen, 1100 m a.s.l., 05.VIII.1997, a large population (including nymphs) in a pasture with scattered rocks and spruce trees. An older published record from Rheinhessen refers to the following species. In Britain the species has been recorded on heathland sites with dominating Festuca ovina and F. rubra. HN; see also Le Quesne & Morris (1971), Wagner (1939a, 1955)
Hardya signifer (Then, 1897) B VII – B V; adult, 1 gen. On xerothermic hillsides with open grassy vegetation and scattered trees or shrubs, usually (perhaps exclusively) on species of fescue belonging to the Festuca ovina group. Known from Germany only from a few isolated sites, on gypsum, Jurassic limestone and porphyry in warm areas of southern and eastern parts, all between 150 and 450 m a.s.l. (Kyffhäuser, valleys of the Altmühl and Nahe), although locally in high abundance. Originally described from alpine habitats of Salzburg state (Austria), and also reported from the alpine belt of Switzerland and France (Alps and Pyrenees). Otherwise published from Lithuania, Switzerland, Spain, Italy, the former Yugoslawia and Bulgaria. Some of these records, however, should be confirmed. Remane (1961b), Schiemenz (1969), Wagner (1955), HN; see also della Giustina (1989), Günthart (2000), Then (1897)
Hardya tenuis (Germar, 1821) Specimens currently assigned to this taxon in Germany belong to two different species, one of which is of unknown identity. The latter is mainly found in southwestern limestone regions. The characters of ecology and distribution given below only refer to H. tenuis (Germ.) sensu Ribaut (1952).
Mainly M VII – X, ‡‡ occasionally VI; adult(?), 1 gen. Among sparse vegetation in moderately shady, moderately dry to dry, usually acidic and sandy, rarely basic sites. Mainly found along forest margins and under solitary
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trees (notably Pinus and Quercus) in heathland, occasionally in abandoned vineyards and dry grassland, often confined to small patches. Host plants are grasses (Festuca ovina, probably also Poa nemoralis, Agrostis capillaris and others). In Germany only very scattered, mainly in large pine forest areas of northern parts, as well as in inland dune areas (northern upper Rhine plain, Danube, Main and Regnitz valley, Mittelfranken), locally also on sandstone and limestone in warm situations (valleys of the Fränkische Saale, Main and Naab), recorded up to 400 m a.s.l. Has apparently vanished from some former localities; most records date back several decades. In addition, some published records are based on misidentified specimens. Reimer (1992), Schiemenz et al. (1996), Trümbach (1959), Wagner (1935, 1939a), HN; see also Niedringhaus (1991), Wagner (1941b)
Sardius argus (Marshall, 1866) Thamnotettix aphrodoides Haupt, 1935 described from near Bonn is likely to be a junior synonym, although the type material could not be found during a revision of Haupt’s collection.
E V – B X, egg(?), 2 gen.; according to Le Quesne (1969) in England III, IV and VII – XI. This species’ range includes the Mediterranean region and parts of western Europe. It lives in oligotrophic and acidic, usually moderately dry sites, preferentially in grazed heathland. Host plants are grasses, perhaps Agrostis capillaris or Danthonia decumbens. Uncommon in Germany, at the eastern edge of the range, only recorded from Rheinland-Pfalz and Hessen, as well as near Aschaffenburg and probably Bonn (from the latter location published as Th. aphrodoides Haupt, 1935). Highest localities are at c. 350 m a.s.l. only. Has apparently vanished from a number of sites during recent decades. Haupt (1935), Kirschbaum (1868), Post-Plangg & Hoffmann (1982), Remane & Fröhlich (1994b), Wagner (1951a), HN; see also Cobben & Rozeboom (1983), Le Quesne (1969)
Paluda flaveola (Boheman, 1845) E VI – M X; egg, 1 gen. Among tall grasses in sunny to moderately shady, peaty to damp, usually acidic and rather cool sites (mainly clearings, open forests, fens, intermediate bogs, margins of raised bogs, abandoned meadows and ruderal sites; apparently absent from mown grassland). Host plants are species of small-reed (Calamagrostis epigejos, C. canescens, C. villosa, C. arundinacea), but also further grasses (perhaps Deschampsia cespitosa, Phalaris arundinacea, Molinia caerulea and others). Widespread in the northeastern half of Germany and fairly common in favourable sites (up to at least 900 m a.s.l.), but apparently at the edge of the range; southwesternmost localities are known from the Eifel and Rhön Mountains, Mainfranken, the Black and Bavarian Forest. Is not reported from Austria, Switzerland and France. Niedringhaus & Olthoff (1993), Reimer (1992), Remane (pers. comm.), Remane & Fröhlich (1994b), Schiemenz (1971a), Schiemenz et al. (1996), Wagner (1941a), Nickel & Remane (1996), HN; see also della Giustina & Remane (2001)
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Rhopalopyx adumbrata (C. Sahlberg, 1842) sensu Vilbaste (1962) Was considered to be conspecific with Rh. preyssleri (H.-S.) for a long time, until Vilbaste (1962) pointed out that there are pronounced differences in the male genitalia and in habitat preferences, although without studying the type material. Concerning life history, both species are similar in phenology, distribution and habitat requirements. However, there are differences in moisture requirements (despite some overlap) and host choice (see below). Thus they may co-occur in damp sites, but on different plants.
B VII – B X; egg, 1 gen. In usually low-growing stands of grasses in oligotrophic, moderately wet to moderately dry, sunny to moderately shady, acidic to basic sites, mainly low-productivity meadows and pastures, heathland and calcareous hillsides. Definite host plant records all refer to Festuca rubra and, less commonly, F. ovina. Widespread in Germany, but at the edge of the range; westernmost localities are on the East Frisian Islands, in the Eifel Mountains and the Black Forest. The species is also reported from England and the French Alps, where it is found up to 2000 m a.s.l., although highest localities in the German Mittelgebirge and the Bavarian Alps are only at 1500 m a.s.l. Niedringhaus & Olthoff (1993), Reimer (1992), Remane & Fröhlich (1994b), Rombach (1999b), Nickel & Achtziger (1999), Nickel (1997), HN; see also Le Quesne (1969)
Rhopalopyx preyssleri (Herrich-Schäffer, 1838) sensu Vilbaste (1962) See taxonomic notes on the preceding species.
B VI – M X; egg, 1 gen. Usually in sunny, occasionally also slightly shaded grass stands in damp to dry, basic as well as acidic sites, mainly in disturbed habitats (ruderal sites, abandoned fields and vineyards, waysides, forest clearings, dry grassland, etc.), locally syntopic with the preceding species. Is often found among Calamagrostis epigejos, Brachypodium pinnatum and Elymus repens, which have been suggested to be the host plants, but apparently, the species lives exclusively on Poa pratensis angustifolia. Widespread in Germany, but often found in low numbers only. Highest localities are at 800 m a.s.l. in the foothills of the Alps and on the Swabian Jura. Achtziger (1991), Frommer (1996), Niedringhaus (1991), Reimer (1992), Schiemenz et al. (1996), Nickel (1997), HN
Rhopalopyx elongata W. Wagner, 1952 This species, bearing a diagnostic pygophor, was already shown by Ribaut (1936) as Thamnotettix vitripennis (Fl.), after specimens collected in the French Roussillon region. The type series of Jassus vitripennis Flor, 1861, however, included specimens both from Latvia and southern France, which later turned out to belong to different species. Thus, after strict interpretation of nomenclatural rules, the species’ name would have been fixed by Ribaut. Later Wagner (1952b), who was not aware of that, and who used the name vitripennis also for the central and north European populations, found out that there were two distinct species and described Rh. elongatus W.Wg. (locus typicus in Spain). In order to avoid further complicating of the nomenclatural situation, Ribaut (1959) did not insist on the priority of his interpretation. This opinion is adopted here. The grammatical gender of Rhopalopyx is female (see Ribaut 1939).
So far E VI – M X; egg, 2(?) gen.
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Review of species
Occurs mainly in southwestern parts of Europe, but has recently been found in 4 localities 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-exposed sites on limestone and sand. The host plants are apparently grasses. No further details are known on the life history. Frommer (pers. comm.), Rombach (1999a); see also Nast (1987)
Rhopalopyx vitripennis (Flor, 1861) Is usually considered to be a much variable taxon comprising a number of varieties, which were formerly treated as distinct species. These varieties include Rh. parvispinus W. Wagner, 1948, Rh. monticola Ribaut, 1939, Thamnotettix graminis Haupt, 1935, and Th. andropogonis Haupt, 1924 (see Wagner 1967). See also taxonomic notes on the preceding species.
E V – M X; egg, 2 gen. In oligotrophic, sunny, moderately dry to dry, usually short or sparsely vegetated sites on various substrates. Host plants are fine-leaved species of fescue (Festuca ovina group, perhaps also F. rubra). Widespread in the lowlands of Germany, often among the dominant species in the sand areas of the north German plain and in warmer regions of southern and eastern parts, but rather sporadic elsewhere. Essentially a species of low altitudes, but occasionally found up to 700 m a.s.l. in the Mittelgebirge and the foreland of the Alps, up to 1050 m a.s.l. in Switzerland. However, small individual numbers are highly vagile; thus reproduction should be ascertained in submontane and montane localities. Fischer (1972), Marchand (1953), Reimer (1992), Remane (1958, 1987), Schiemenz et al. (1996), Struve (1939), Trümbach (1959), Wagner (1935, 1939a, 1951a), Walter (1998), HN; see also Günthart (1987b)
Elymana kozhevnikovi (Zachvatkin, 1938) M VII – E IX; egg, 1 gen. Among tall grassy undergrowth of open forests, mainly under beech, pine, spruce and oak; host plants are several species of small-reed (Calamagrostis arundinacea, C. varia, C. canescens, but not C. epigejos). This is an eastern species reaching its distribution limit in Germany. So far it has been found in mountain forests and pine heaths of the Bavarian Alps and their foothills, the Harz Mountains, Elbsandsteingebirge and Erzgebirge, in low-input forests and coppice-with-standards in Thuringia and Unterfranken, as well as in an alder fen near the southwestern edge of the Harz Mountains, between 250 and 1400 m a.s.l. Emmrich (1986), Fischer (1972), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Nickel (1994), HN; for identification of nymphs see Dmitriev (1999), Walter (1978)
Elymana sulphurella (Zetterstedt, 1828) E VI – M X; egg, 1 gen. (see also Witsack 1985). Eurytopic in grass stands of wet to moderately dry, sunny to moderately shady sites, mainly in low-input meadows and pastures, abandoned fields, clearings, shady patch-
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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), Schiemenz et al. (1996), Wagner (1935), Nickel & Achtziger (1999), HN, and others; see also Günthart (1987a), Morcos (1953); for identification of nymphs see Dmitriev (1999), Walter (1978)
Cicadula albingensis W. Wagner, 1940 Mainly M VII – B X, occasionally B VI – E X; egg, 1 gen. In wet to peaty, meso- to moderately eutrophic, sunny to moderately shady sites (mainly in spring mires and along headwaters of streams, also in fens and intermediate bogs, straw and peaty meadows). The preferred host plant is Scirpus sylvaticus, but there are also a few records from tall-sedge stands (Carex acuta, C. acutiformis and others). In Germany near the western and southern edge of the range; westernmost localities are near Freiburg, Pirmasens, Koblenz, Marburg, Göttingen and Hamburg (locus typicus). Has been found in France and Belgium, but is not reported from the Netherlands, Switzerland and Italy. Widespread in Germany, locally common at submontane altitudes, but rather scattered in the lowlands. Highest localities are at 1100 m a.s.l. in the Bavarian Alps and the Black Forest. Fischer (1972), Fröhlich (1996a), Heller (1987a), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Trümbach (1959), Wagner (1940b, 1951a), Nickel & Achtziger (1999), Nickel (1997), HN
Cicadula rubroflava Linnavuori, 1952 Some authors (e.g. Ossiannilsson 1983) use the name C. longiventris (J. Sahlberg, 1871) for this species. However, this name is based on the description of ‡‡, which could not be identified with certainty until Remane & Fründ (1986). Therefore, Linnavuori’s name should be kept until the type material has been revised (see Remane & Fröhlich 1994b).
B VIII – M X; egg, 1 gen. On Carex brizoides in forests and forest glades on seepy or waterlogged substrates, usually in stands of spruce and pine in cool situations. In Germany only very sporadic, at the western and southern edge of the range; at present there are only 15 known localities, mostly in Saxony (Elbsandsteingebirge, Oberlausitz, Dresdener Heide, Vogtland) and the foothills of the Alps (Ammersee area, near Kelheim and Biberach), also in the Bavarian Forest (near Freyung), Vogelsberg (Waldensberg) and Main valley (Volkach), between 200 and 750 m a.s.l. Emmrich (1989), Remane & Fröhlich (1994b), Remane & Fründ (1986), Schönitzer & Oesterling (1998b), Walter (pers. comm.), HN
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Review of species
Cicadula persimilis (Edwards, 1920) B VI – E X; egg, 2 gen., probably 1 at higher altitudes. On Dactylis glomerata in sunny to slightly shady, moist to moderately dry sites, notably moderately eutrophic meadows and pastures, abandoned fields, ruderal sites, waysides, etc. In Germany widespread and common in most regions, although less common and confined to field margins in regions of more intensive agriculture. Frequently occurs up to at least 1800 m a.s.l. in the Bavarian and Allgäu Alps. Achtziger (1991), Fischer (1972), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others; see also Morcos (1953)
Cicadula quinquenotata (Boheman, 1845) E VII – E X; egg, 1 gen. In intermediate bogs of dune valleys and moraine depressions. The host plant is not known, most authors suggest an unidentified species of Carex or Eriophorum. In Germany the species is confined to the northern plains; southernmost localities are on the East Frisian Islands, near Hamburg, Lauenburg, Waren an der Müritz, and Luckenwalde. All German sites are below 100 m a.s.l., but the species is reported from altitudes above 1500 m a.s.l. in the Austrian Alps. Kuntze (1937), Niedringhaus (1991), Remane & Fründ (1986), Schiemenz et al. (1996), Wagner (1935, 1937e); see also Holzinger (1999b), Ossiannilsson (1983), Wagner & Franz (1961)
Cicadula saturata (Edwards, 1915) B VII – M X; egg, 1 gen. In wet, usually peaty and acidic sites (intermediate bogs, fens, floating mats of Sphagnum, peaty meadows); host plants are sedges, mainly Carex nigra, perhaps also C. rostrata. In Germany at the western and southern edge of the range, with strongholds in rather cool areas of the north German plain, the Mittelgebirge and the Bavarian and Swabian foothills of the Alps, found up to 1500 m a.s.l. in Upper Bavaria. Not reported from Austria, Switzerland, Italy and Belgium, but from the Netherlands and eastern parts of France. Bittner & Remane (1977), Fröhlich (1996a), Hoffmann (1980), Nikusch (1976), Reimer (1992), Schiemenz et al. (1996), SMNS, Wagner (1935, 1946), Nickel & Achtziger (1999), Nickel (1997, 1999b, 2002); see also Cobben & Gravestein (1958), della Giustina (pers. comm.), den Bieman (pers. comm.), Nast (1987), Ribaut (1952)
Cicadula flori (J. Sahlberg, 1871) Mainly E VI – E X, in some years from E V onwards; egg, 2 gen. In stands of tall sedges in wet to temporarily flooded, usually moderately eutrophic sites, mainly fens and straw meadows, occasionally also in fen woods, often syntopic with C. quadrinotata (F.) and locally dominant, but more stenotopic and restricted to rather warm situations. Host plants are tall sedges (preferentially Carex acuta, perhaps also C. acutiformis, C. elata and additional species). Published records from central Europe are sparse,
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but the species is apparently widespread in most lowland areas of northern and middle Germany (e.g. Holstein, southern and eastern Niedersachsen, Thuringia, Hessen, Franconia). Highest localities are at 500 m a.s.l. in the Bavarian foreland of the Alps. Hildebrandt (1995), Niedringhaus (1991), Reimer (1992), Remane (1958), Schiemenz et al. (1996), Trümbach (1959), Wagner (1946), Weis & Schönitzer (2001), Nickel & Achtziger (1999), HN; see also Lauterer (1986)
Cicadula quadrinotata (Fabricius, 1794) B VI – E X; egg, 2 gen., 1 at higher altitudes. Eurytopic in peaty (also temporarily flooded) to moist, locally also moderately dry, sunny to moderately shady sites, often dominant in fens, intermediate bogs, low-input meadows, along ditches, and in peat diggings in raised bogs, also found in ruderal sites, moderately saline marshes and fen woods. In regions of more intensive agriculture, however, the species is often restricted to ditches and other margins. Host plants are Carex acutiformis, C. acuta, C. rostrata, C. nigra, C. elata, C. disticha, C. brizoides, C. hirta and others; few individuals have also been found on Bolboschoenus maritimus and Eriophorum angustifolium. Widespread in Germany and among the dominant species in many wetland habitats, although population sizes are likely to have decreased due to draining and intensification of grassland management. Occurs up to at least 1500 m a.s.l. in the Allgäu Alps, but up to 2400 m a.s.l. in North Tyrol (Austria) and the Engadine (Switzerland). Bittner & Remane (1977), Fischer (1972), Marchand (1953), Reimer (1992), Schiemenz (1971a), Schiemenz et al. (1996), Wagner (1935), Nickel & Achtziger (1999), HN; see also Günthart (1987a), Leising (1977), Novotný (1995)
Cicadula frontalis (Herrich-Schäffer, 1835) Mainly M VII – M X, in southern parts and in warm years from B VI onwards; egg, 1 (-2?) gen. In sunny to moderately shady, wet to temporarily flooded, usually moderately eutrophic sites, mainly fens and shores of ponds and streams, occasionally in fen woods; apparently absent from mown sites. Host plants are tall-growing sedges, mainly Carex acutiformis and C. riparia, perhaps also C. pseudocyperus and C. acuta. Widespread in the lowlands of Germany, but uncommon, usually in lower abundances than the remaining species of Cicadula. Occurs up to at least 700 m a.s.l. in the foothills of the Alps, but apparently only below 500 m a.s.l. in regions to the north of the Danube. Frommer (1996), Niedringhaus & Olthoff (1993), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1935, 1951a), HN
Cicadula ornata (Melichar, 1900) So far only B VII – M VII, in Scandinavia VII – IX (Ossiannilsson 1983); egg, 1(?) gen. This species’ range includes the whole boreal zone of the Palearctic and the Nearctic. In Europe it was hitherto only known from England, Sweden, Finland, Estonia, Latvia and Poland. Is reported to live on tall sedges in river floodplains, but no further details are known on habitat requirements. From Germany there is only a single record from suburban
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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 northwestern parts of Germany referred to C. quinquenotata (Boh.) and have been corrected later. Hence, the occurrence of C. intermedia (Boh.) in Germany is not proven. Emmrich & Nickel (unpublished data), Schiemenz et al. (1996), Wagner (1935, 1937e); see also Flint (1990), Nast (1987), Ossiannilsson (1983), Vilbaste (1974)
Mocydia crocea (Herrich-Schäffer, 1837) Adults I – XII, mainly B VIII – B VI; adult, 1 gen. (see also Müller 1976). In sunny to moderately shady, dry to temporarily moist sites on basic to acidic substrates, usually in dry grassland, ruderal sites, and along waysides, locally also in straw meadows, temporarily dry salt marshes and open, thermophilous forests of oak or pine. Host plants are Brachypodium pinnatum, Bromus erectus, Calamagrostis epigejos, Elymus repens and other tall grasses. In Germany at the northern edge of the range; widespread and locally common in middle and southern parts, but rather localized in the north German plain; recorded northward to Cologne, Hanover, Wolfsburg, Magdeburg and Neubrandenburg. Usually found below 500 m a.s.l., except in the higher limestone regions (Bavarian Alps, Swabian Jura) where occurring up to at least 1000 m a.s.l. Achtziger (1991), Fischer (1972), Fröhlich (1996a), Frommer (pers. comm.), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Rombach (1999a), Schiemenz et al. (1996), Schwoerbel (1957), Nickel & Achtziger (1999), Nickel (1994), HN, and others
Mocydiopsis attenuata (Germar, 1821) E VII – V; adult, 1 gen. On species of fescue (Festuca ovina group, F. rubra, F. heterophylla) usually in moderately shady, damp to moderately dry, acidic as well as basic sites; mainly along margins of forests of pine, oak and beech (occasionally also in their interior), as well as in low-productivity grassland with scattered trees or shrubs. The distribution in Germany (as well as in other parts of Europe) is poorly documented due to identification problems; however, the species is apparently widespread (although not yet recorded from MecklenburgVorpommern and Baden-Württemberg), but densities are usually low. Most localities are below 400 m a.s.l., but there is a single record from the foothills of the Alps at 750 m a.s.l. Frommer (1996), Niedringhaus (1991), Nikusch (1976), Remane (1961a, 1987), Schiemenz et al. (1996), HN
Mocydiopsis intermedia Remane, 1961 E VII – V; adult, 1 gen. In moderately dry to dry, moderately shady sites on basic to moderately acidic substrates, usually in xerothermic grassland with scattered trees or shrubs and along sunny forest margins, probably associated with Poa pratensis angustifolia. In Germany this species occurs at the northern edge of the range and is largely confined to warmer low-
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land regions between the Danube and the northern edge of the Mittelgebirge (mainly along the upper and middle Rhine, Franconian Jura, Mainfranken, Saale-Unstrut region), up to at least 550 m a.s.l. Border localities are near Bonn, Marburg, Bad Neustadt an der Saale, Bad Frankenhausen, Quedlinburg, Halle and Leipzig. It is not reported from Belgium, the Netherlands, the British Islands, northern Europe and Poland. Has apparently disappeared in a number of localities. Funke & Witsack (1998), Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1961a, 1987), Remane & Fröhlich (1994b), Schiemenz et al. (1996), HN
Mocydiopsis longicauda Remane, 1961 B VII – V; adult, 1 gen. In moderately dry to dry, basic to acidic sites, usually in the vicinity of trees or shrubs (mainly dry grassland, meadows with scattered fruit trees, forest margins, etc.). Host plants are fine-leaved species of fescue (Festuca ovina group). A tongue-shaped part of the range extends from southeastern Europe into Germany, with strongholds in Thuringia, Mainfranken and the Franconian Jura. Border localities are near Regensburg, Eichstätt, Würzburg, Bad Neustadt an der Saale, Gotha, Bad Frankenhausen, Halberstadt, Haldensleben, Dessau and Gera, at least up to 500 m a.s.l. Müller (1978), Reimer (1992), Remane (1961a), Remane & Fröhlich (1994b), Schiemenz et al. (1996), HN
Mocydiopsis monticola Remane, 1961 E VII – V; adult, 1 gen. On Holcus mollis in damp to moist, moderately shady, usually acidic sites, mainly open oak forests, clearings, coppice-with-standards, forest margins, etc. In Germany only known from the middle parts, between the Danube and the northern edge of the Mittelgebirge, up to at least 400 m a.s.l., although probably under-recorded. Northernmost localities are near Cologne, Frankenberg an der Eder, Artern and Görlitz. The species is neither reported from more northern parts of Europe, nor from Belgium, the Netherlands, Poland and the Czech Republic, but it is probably overlooked or misidentified. Frommer (pers. comm.), Remane (pers. comm.), Remane & Fröhlich (1994b), HN; for identification see Remane (1961a)
Mocydiopsis parvicauda Ribaut, 1939 E VII – IV; adult, 1 gen. On Agrostis capillaris in oligotrophic and acidic, sunny to moderately shady, moderately moist to dry sites (mainly pastures and heaths, also clearings, forest margins, coppicewith-standards, etc.). Widespread in western parts of Germany, but usually at low densities, more scattered towards the east, where apparently at the northeastern edge of the range. Border localities are on a line from Kiel to Hamburg, Stendal, Leipzig and Hoyerswerda. The species is found up to at least 600 m a.s.l. in the Mittelgebirge and the foreland of the Alps.
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Frommer (pers. comm.), Niedringhaus & Olthoff (1993), Nikusch (1976), Reimer (1992), Remane (1961a.), Schiemenz et al. (1996), HN
Speudotettix subfusculus (Fallén, 1806) This is a rather polymorphic species with much varying body size, wing length and colouration intensity. Variation is particularly pronounced in upper montane and subalpine populations of the Alps. Speudotettix montanus G¸ebicki & Szwedo, 1991, described on the basis of merely two specimens from a mountain site in southeastern Poland, is likely to be within this range of variation and should be revised.
Mainly E IV – M VIII, at higher altitudes until M IX; nymph, 1 gen. Eurytopic in deciduous and coniferous forests and along their margins, in wet to moderately dry, basic to acidic sites, also among subalpine scrub and in alpine grassland. Nymphs are found among low vegetation (often on sedges and grasses); most adults feed on various deciduous trees and shrubs (Betula, Corylus, Alnus, Populus, Prunus and others), probably on dwarf shrubs above the alpine tree line. Widespread and common in Germany, also in higher parts of the Mittelgebirge and the Bavarian and Allgäu Alps, where frequently encountered up to 2000 m a.s.l.; in Switzerland even found up to 2400 m a.s.l. Achtziger (1991), Fischer (1972), Frommer (1996), Niedringhaus (1997), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Schwoerbel (1957), Wagner (1935, 1939a), HN, and others; see also Günthart (1987a)
Hesium domino (Reuter, 1880) E VI – B X; egg, 1 gen. In wet to moderately dry, basic to acidic sites with semi-open stands of deciduous trees (fens, intermediate bogs, low-productivity meadows and pastures, dry grassland, forest margins and glades, clearings, etc.). Adults usually on Betula pendula and B. pubescens, occasionally on Alnus, Ulmus and other trees, nymphs in low-growing herbaceous vegetation, probably feeding on grasses (Festuca rubra, Holcus mollis and others). Widespread in Germany, except in northwestern parts, and in intermediate frequency, but low individual numbers. Apparently absent from the northwest German plain, the Netherlands, Denmark and the British Isles; border localities are near Bonn, Siegen, Warburg, Northeim, Goslar, Hamburg and Rostock. Recorded up to at least 1200 m a.s.l. in the Mittelgebirge and Bavarian Alps, up to 1750 m a.s.l. in the French Alps. Achtziger (1991), Fischer (1972), Jöst (1966), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Wagner (1935, 1939a), HN, and others; see also della Giustina (1989)
Thamnotettix confinis (Zetterstedt, 1828) Mainly M V – M VIII, at montane and subalpine altitudes VI – IX; nymph, 1 gen. Polyphagous in deciduous and coniferous forests of damp to moist, acidic to basic sites, as well as among subalpine scrub. Nymphal development takes place in low-growing herbaceous vegetation; at least some adults ascend various deciduous trees or shrubs (Quercus, Betula, Rubus and others). Widespread all over Germany, found up to at least
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1800 m a.s.l. in the Bavarian and Allgäu Alps, up to 2200 m a.s.l. in North Tyrol (Austria) and the Engadine (Switzerland). Fischer (1972), Frommer (1996), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Schwoerbel (1957), Wagner (1935, 1939a), HN; see also Günthart (1987a), Leising (1977)
Thamnotettix dilutior (Kirschbaum, 1868) B V – B X; nymph, 1 gen. In open oak forests and along their margins, in moderately dry to damp, acidic as well as basic sites. Nymphs are found among low-growing herbaceous vegetation (on Deschampsia flexuosa and other grasses), occasionally on shrubs, most adults on oaks (Quercus robur and Qu. petraea). Widespread in Germany, but restricted to rather low altitudes (found up to 400 m a.s.l. only, but reported to occur at 800 m a.s.l. in Ticino, Switzerland). Locally common in warmer regions in southern and eastern parts (often abundant in Malaise traps), but more localized and less common towards the north, not recorded to the south of the Danube. Post-Plangg & Hoffmann (1982), Niedringhaus (1997), Remane (1987), Schiemenz et al. (1996), Wagner (1935, 1939a, 1951a), HN; see also Günthart (1987b)
Pithyotettix abietinus (Fallén, 1806) E V – M IX, single individuals until XI; nymph, 1 gen. In coniferous and mixed forests of higher altitudes, secondarily also in lowland conifer plantations. Nymphs and adults on Picea abies, perhaps also on Abies alba. In Germany originally confined to the natural host range in the Alps and at higher altitudes of the Mittelgebirge, where locally common, but nowadays also widespread in the lowlands, even found in parks and cemeteries. However, the species is seldom recorded, since adults usually stay in the canopy layer. Highest localities are at 1700 m a.s.l. near the alpine tree limit of Upper Bavaria. Fischer (1972), Frommer (1996), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Trümbach (1959), Wagner (1935, 1939a, 1951a), HN, and others
Perotettix pictus (Lethierry, 1880) E V – E VII, occasionally until X; nymph, 1 gen. Like P. abietinus (Fall.) on Picea abies and perhaps Abies alba, mainly in the canopy layer of open forests, along forest roads and edges. Less common than the preceding species and more restricted to forest areas of higher altitudes; in Germany only found between 300 and 1700 m a.s.l., at the northern edge of the range; recorded from the Alps and their foreland, the Black Forest, the Erzgebirge, the Rhön Mountains, middle Hessen, Thuringia, the Rothaargebirge and the Harz Mountains. Reimer (1992), Remane (pers. comm.), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), SMNS, Nickel & Remane (1996), HN
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Colobotettix morbillosus (Melichar, 1896) M V – E VIII, single individuals until IX; nymph, 1 gen. Another species living on Picea abies and perhaps Abies alba, usually in higher situations; most adults are swept in open forests and along their margins. In Germany probably under-recorded, but apparently more or less restricted to the alpine foothills and the Mittelgebirge (up to at least 1200 m a.s.l.), but occasionally found in the north German plain. According to present data, the range is largely confined to central Europe and includes southern Finland, France (only Savoie), Poland, Switzerland, Austria and the Czech Republic; the species is not reported from Russia. Fischer (1972), Reimer (1992), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Trümbach (1959), Wagner (1935, 1939a), HN; see also della Giustina (1989), Emelyanov (pers. comm.), Günthart (pers. comm.), Holzinger (1996a), Lauterer (pers. comm.), Nast (1976a, 1987), Ossiannilsson (1983)
Macustus grisescens (Zetterstedt, 1828) E IV – M VIII, at higher altitudes occasionally until IX; nymph, 1 gen. In tall-growing stands of sedges and grasses in peaty to damp, sunny to moderately shady sites, mainly in fens, intermediate and raised bogs, straw meadows, fen woods, along ditches, as well as in forest clearings, abandoned fields and open forests. Host plants are sedges (Carex acutiformis, C. paniculata, probably also C. acuta, C. rostrata, C. nigra, and others), grasses (Calamagrostis arundinacea, C. epigejos, Festuca arundinacea, Dactylis glomerata, Elymus repens and others) and woodrushes (Luzula spp.). Widespread in Germany and fairly common in favourable sites, found up to at least 1900 m a.s.l. in the Bavarian and Allgäu Alps, up to 2170 m a.s.l. in the Engadine (Switzerland). Fischer (1972), Niedringhaus (1991), Reimer (1992), Schiemenz et al. (1996), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN; see also Günthart (1987a)
Doliotettix lunulatus (Zetterstedt, 1838) B V – B VIII; nymph, 1 gen. In moderately shaded stands of grasses, usually in wet to moderately moist, acidic to moderately basic sites, mainly at submontane altitudes, mainly in deciduous and coniferous forests and along their margins, probably associated with Agrostis stolonifera. In Germany only localized, but fairly common in favourable habitats, with strongholds in the Mittelgebirge (Eifel Mountains, Rothaargebirge, Vogelsberg, hilly regions of Thuringia, Harz Mountains, Swabian Jura and others) and the foothills of the Alps, occasionally also found in the lowlands (near Hamburg, Berlin, Wiesbaden and Landau/Pfalz). Highest localities are at 1000 m a.s.l. Fischer (1972), Fröhlich (1996a), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a), HN
Athysanus argentarius Metcalf, 1955 B VI – E IX, according to Schiemenz et al. (1996) B V – B XI; egg, 1 gen.
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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 moderately saline sites. Host plants are various grasses (Calamagrostis spp., Elymus repens, Dactylis glomerata, Arrhenatherum elatius, Deschampsia cespitosa, Festuca arundinacea, Holcus spp. and others). Widespread and fairly common in the lowlands of Germany, found up to 900 m a.s.l. in the Mittelgebirge and the foothills of the Alps. Achtziger (1991), Fröhlich (1996a), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a), Walter (pers. comm.), Nickel & Achtziger (1999), HN
Athysanus quadrum Boheman, 1845 B VII – M X; egg, 1 gen. In sunny, moist to wet, usually mesotrophic sites, mainly calcareous sedge fens and non-fertilized peat and straw meadows. The host plants are unknown, but may include dicotyledonous herbs (perhaps Lathyrus, Inula, or Filipendula). Has been found in large parts of Germany, but most records date before 1970. Highest sites are at 600 m a.s.l. in the Bavarian foothills of the Alps. German localities are apparently near the southern and southwestern edge of the range. The species is reported from the Netherlands, Belgium and Austria, but not from Switzerland; the French records are uncertain. However, it has been found in the Kaiserstuhl, close to the French and Swiss border. Fischer (1972), Marchand (1953), Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN; also della Giustina (pers. comm.), Nast (1987)
Handianus ignoscus (Melichar, 1896) Species identities within this genus are obscured by the loss of some type material, notably of H. flavovarius (Herrich-Schäffer, 1835) and H. procerus (Herrich-Schäffer, 1835). The species interpretation in this book is adopted from Dlabola (1954) and Emelyanov (1964a).
So far B VII – B IX; egg, 1 gen. Has been published from eastern and southeastern Europe and from several Asian countries, but the total range is unknown due to taxonomic uncertainties. Is reported to live on dry sites on woody species of Fabaceae, mainly Cytisus (s.l.), and is known from Germany only from 5 acidic and sandy localities: Wahrburg (near Stendal), 1925, Falkenberg (to the west of Torgau), 1966, 1967, and 1973, as well as Erlangen, between 1954 and 1957. Furthermore reported from „Brandenburg“, but without details. Recently, the species was found in another site near Erlangen: Schießplatz Tennenlohe, 09.VII.2001, 10 ††, 11 ‡‡, 04.IX.2001, 1 †, 2 ‡‡, on Cytisus scoparius in a former military training area on aeolian sand. The highest localities are at c. 300 m a.s.l. Schiemenz et al. (1996), Trümbach (1959), Wagner & Franz (1961), HN; see also Emelyanov (1964a), Herrich-Schäffer (1835b)
Stictocoris picturatus (C. Sahlberg, 1871) E VI – E X; egg, 1 gen.
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In sunny and low-vegetated grassland of dry to temporarily dry, usually basic, occasionally 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 Fabaceae, notably Ononis spinosa and Genista tinctoria, occasionally Chamaespartium sagittale, perhaps also herbaceous species (Trifolium?). In Germany only scattered, at the northwestern 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 Valais (Switzerland). Border localities include Koblenz, Marburg, Bad Lauterberg, Quedlinburg, Lauenburg and Neubrandenburg. Kupka (1925), Nikusch (1976), Reimer (1992), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1951a), HN; see also Prohaska (1923)
The following taxon Ophiola Edwards, 1922 has recently been placed into the genus Limotettix J. Sahlberg, 1871 s.l. as a subgenus by Hamilton (1994). Instead of Ophiola Edw., this author prefers Scleroracus Van Duzee, 1894 as name. The latter, however, is rejected by most European authors as a nomen nudum (e.g. Nast 1987; Ossiannilsson 1983; Remane & Fröhlich 1994a, and others).
Ophiola cornicula (Marshall, 1866) This species was misinterpreted for a long time, and various names have been used (notably Jassus orichalceus Thomson, 1869, and Limotettix intractabilis Kontkanen, 1949). Kontkanen (1948) showed that in central and northern Europe there are at least three species with distinct genitalia and morphometrics. Later Le Quesne (1962) studied Marshall’s type material and clarified the nomenclatural situation.
E VI – E IX; egg, 1 gen. In acidic, moderately dry to peaty, sunny to moderately shady sites, notably in heaths, raised bogs (preferentially in drained patches) and open pine forests. Feeds on Calluna vulgaris, probably also on species of Vaccinium. Widespread in Germany, but largely restricted to areas of sand, sandstone, bogs and other acidic substrates, mainly in the north German plain, the submontane and montane belt of the Mittelgebirge and along the edge of the Alps; the highest localities are at 1200 m a.s.l. Bittner & Remane (1977), Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987), Schiemenz et al. (1996), HN
Ophiola decumana (Kontkanen, 1949) Cicada striatula Fallén, 1806 (p. H.) Identity and species differentiation in this taxon are not satisfactorily studied. Perhaps in central Europe there is a complex of several distinct species. Furthermore, the identity of O. ocellaris (Lethierry, 1880) is uncertain; it has been described from France and Austria and is reported from large parts of the western Palearctic by Nast (1972, 1987), but has not been mentioned by Hamilton (1994) in his generic review.
E V – B X; egg, 2 gen. In sunny, moderately dry to dry, acidic to basic sites of early successional stages, usually on gravelly to sandy, locally also on loamy substrates, mainly mining areas, trampled patches (preferring middle strips of dirt roads), military training areas, abandoned fields, low-input vineyards, as well as gravel banks of alpine rivers. Host plants are Polygonum
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aviculare, Rumex acetosella, and perhaps further dicotyledonous herbs; from Sweden Vaccinium myrtillus and Calluna vulgaris have been reported. Widespread in Germany, but only localized; most older records are in need of revision. There are recent verified records from various parts of the north German plain, the warmer regions in eastern and southern parts (mainly Saale-Unstrut region, valleys of Main and Regnitz, upper Rhine plain), and the Alps (upper course of the Isar, Berchtesgaden); usually found below 300 m a.s.l., but up to at least 900 m a.s.l. along alpine rivers. Niedringhaus & Olthoff (1993), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Nickel (1999a), HN; see also Ossiannilsson (1983)
Ophiola russeola (Fallén, 1826) Has been treated by some authors as conspecific with O. plutonia (Uhler, 1877). According to Hamilton (1994), the latter is a distinct species confined to the New World.
E VI – B X; egg, 1 gen. On dwarf shrubs (mainly Calluna vulgaris, but also Erica carnea, Vaccinium oxycoccus and others) in raised bogs, heaths and open pine forests, usually in acidic sites, but also on limestone gravel along alpine rivers. Widespread in Germany and locally common in heath and bog areas of the north German plain and the Mittelgebirge, in sandstone and inland dune areas, as well as along the edge of the Alps. Highest localities are at 1200 m a.s.l. in the Allgäu Alps and the Black Forest; small numbers are also reported from the alpine belt of the Engadine (Switzerland) and Styria (Austria), but reproduction is not proven in these altitudes. Niedringhaus & Olthoff (1993), Reimer (1992), Schönitzer & Oesterling (1998b), Strübing (1955), HN; see also Günthart (1987a)
Ophiola transversa (Fallén, 1826) E VI – M IX, according to Schiemenz et al. (1996) from B VI onwards; egg, 1 gen. The range of this species extends from Mongolia to central Europe, apparently with isolated populations in the Alps and the Pyrenees. It inhabits oligotrophic grassland usually in moderately dry or temporarily dry sites (mainly heaths, locally also calcareous hillsides). In Germany there are only sporadic populations, mainly in sand areas of the northeastern half; border localities are on the island of Borkum, near Fallingbostel, Halle, Leipzig and Görlitz. Another locality in southern Thuringia, at 550 m a.s.l., appears to be isolated. Some populations from Germany (as well as from the Italian Alps and the Pyrenees) were taken from Achillea millefolium. Recorded at 1900 m a.s.l. in the Slovakian Tatra Mountains. Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Wagner (1935); see also Lauterer (1984)
Limotettix atricapillus (Boheman, 1845) E VI – E IX; egg, 1 gen.
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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, Dresden), a few isolated sites are in the Lake Constance area (Bodanrück) and the Bavarian foothills of the Alps (Oberstdorf, 850 m a.s.l., and Inzell, 700 m a.s.l.). Has vanished from some former localities. The host plants are poorly known; Carex has been stated in the literature. The Inzell record comprised a large population living on Rhynchospora alba in an overgrown peat digging. MTD, Schiemenz et al. (1996), Wagner (1941a, 1943), Wagner & Wagner (1938), Remane (1962), Nickel (1999b)
Limotettix striola (Fallén, 1806) B VI – E X; egg, 2 gen., probably 1 in bog habitats and at higher altitudes. In various habitats in sunny, peaty or temporarily flooded sites: in moderately saline meadows, floodplain depressions, raised and intermediate bogs, also as a pioneer species in sand and gravel pits. Host plants are various species of Cyperaceae (mainly Eleocharis, probably also Schoenoplectus and Trichophorum). Published records from rushes are probably based on host misidentification. Widespread in Germany and locally fairly common and abundant, mostly recorded along the coasts of the North and Baltic Sea, in inland salt areas (notably in Thuringia and Saxony-Anhalt), along larger river valleys, and in bog areas, rather scattered elsewhere; found up to at least 1350 m a.s.l. in the Allgäu and Bavarian Alps. Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Remane (1958), Schiemenz et al. (1996), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN
Laburrus impictifrons (Boheman, 1852) B VII – B X; egg, 1 gen. This is a steppe species living monophagously on Artemisia campestris in sunny, moderately dry to dry sites, usually on acidic and sandy substrates. In Germany largely confined to the northeastern half, at the edge of the range, with strongholds in sand areas of Mecklenburg-Vorpommern, Brandenburg, Saxony-Anhalt and Saxony; otherwise only known from isolated sites along the lower course of the Elbe, on gypsum hillsides in Thuringia, and inland sand dunes along the Rhine (Mainz, Darmstadt, Heidelberg), Main (Aschaffenburg, Karlstadt, Volkach, Bamberg) and Danube (Kelheim); the highest localities are at 350 m a.s.l. only. Furthermore, there are a few 19th century records from southwestern France (Gironde, Landes). Remane (1987), Schiemenz et al. (1996), SMNS, Wagner (1935, 1939a, 1951a), Walter (pers. comm.), Nickel (1997), HN; see also Ribaut (1952)
Laburrus pellax (Horváth, 1903) B VI – B X; egg, 1 gen. Monophagous on Aster linosyris in dry grassland in sunny, usually basic sites, mainly on limestone and gypsum, occasionally porphyry. This is another xerothermophilous steppe species, its range extends from central Asia to southern parts of Germany. From
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there it is known from scattered localities in the Nahe and upper Rhine valley (also on the French side), southern Palatinate, Mainfranken, the Tauber valley, southern parts of the Franconian Jura, and the Kyffhäuser, up to 400 m a.s.l. only. Remane & Wachmann (1993), Schönitzer & Oesterling (1998b), SMSN, Wagner (1939a, 1951a), Nickel (1999b), HN; see also della Giustina & Remane (2001)
Euscelidius schenckii (Kirschbaum, 1868) B VII – M X, according to Schiemenz et al. (1996) from M VI onwards; egg, 1 gen. In disturbed, moist to moderately dry, sunny to moderately shady sites, mainly ruderal habitats, tall herb stands (preferentially Urtica dioica), and disturbed patches in dry grassland and vineyards, apparently polyphagous on various dicotyledonous herbs. Widespread in Germany, but seldom recorded due to secretive life habits close to the ground; the highest localities are at 500 m a.s.l. only. The locus typicus is Weilburg an der Lahn. Achtziger (1991), Hildebrandt (1986), Reimer (1992), Schiemenz et al. (1996), Trümbach (1959), Wagner (1951a), HN
Euscelidius variegatus (Kirschbaum, 1858) B VII – M X; egg, 1 gen. Like the preceding species in disturbed sites, but – despite broad overlap – preferring warmer and drier situations and more sandy substrates, mainly ruderal habitats, abandoned fields, sand and gravel pits, waysides, vineyards, and even gardens. Many records comprise only single individuals, which may have been on dispersal flight; thus, some published data on habitat requirements and food plants are perhaps uncertain. The host plants are unknown; presumably, the species lives polyphagously on various dicotyledonous herbs. In Germany it is largely confined to warm regions in southern and eastern parts (mainly upper and middle Rhine valley and its larger tributaries, the rain shadow area to the east of the Harz Mountains), at the northwestern edge of the range. Border localities are on a line from Cologne to Gießen, Bad Frankenhausen, Halberstadt and Berlin; the highest sites are at 350 m a.s.l. only. The locus typicus is Mombach near Mainz. Is reported as a vector of Clover phyllody (CP) from France. Frommer (1996), Remane (1987, and pers. comm.), Rombach (1999a), Schiemenz et al. (1996), SMNS, Wagner (1939a, 1951a), HN; see also Brˇcák (1979)
Conosanus obsoletus (Kirschbaum, 1858) B VII – M X, according to Schiemenz et al. (1996) M V – M XI; egg, 1 gen. In sunny, peaty to damp or temporarily waterlogged sites, usually in fens, low-input meadows and pastures, and salt marshes, also found in intermediate bogs, forest clearings, ruderal sites, and along waysides and ditches. Host plants are rushes (Juncus effusus, J. filiformis, J. inflexus, J. gerardii and others) and tall grasses (Festuca arundinacea, perhaps Elymus repens, Arrhenatherum elatius and others). Widespread in Germany and fairly common, the highest localities are at 1200 m a.s.l. in the Black Forest.
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Bittner & Remane (1977), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Reimer (1992), Remane & Wachmann (1993), Schiemenz (1971a), Schiemenz et al. (1996), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others
Euscelis distinguendus (Kirschbaum, 1858) E VI – B X; egg, 1 gen. In sunny, oligotrophic, basic to acidic sites; in the Bavarian and Allgäu Alps and the Black Forest mainly found in temporarily moist to moderately dry low-productivity pastures and meadows up to at least 1600 m a.s.l. (up to 2100 m a.s.l. in North Tyrol, Austria), in the lowlands preferring moderately dry grassland on limestone or siliceous substrates (also on sand), particularly on abandoned or disturbed pastures and fields. Lives on species of Asteraceae, probably Taraxacum or Picris hieracioides, perhaps also Hieracium and others. The German sites are at the northwestern edge of the range; the species is widespread and fairly common in the Alps and their foothills, but rather sporadic elsewhere, mainly in regions of limestone and sand. Border localities are near Mainz (locus typicus), Göttingen and Schleswig. Achtziger (1991), Remane (1987), Remane (pers. comm.), Schiemenz et al. (1996), Strübing (1966, and pers. comm.), Wagner (1939a, 1951a), Nickel (1997), HN; see also Leising (1977)
Euscelis incisus (Kirschbaum, 1858) E III – XI; nymph, usually 2 gen., at higher altitudes egg, 1 gen. (Müller 1954); 3rd and 4th instar nymphs and eggs, at higher altitudes only the latter, overwinter in thermic quiescence; all other stages and adults usually die in autumn (Müller 1981; Witsack 1981). In usually sunny, moist to moderately dry, eutrophic to oligotrophic sites, mainly in meadows, pastures, abandoned fields, along waysides and ditches, and even on lawns in gardens and parks; often abundant in low-vegetated, fertilized pastures and meadows in sun-exposed situations. Host plants are various species of Fabaceae (mainly Trifolium spp.) and grasses (in England Trisetum flavescens, Briza media, Helictotrichon pubescens, Arrhenatherum elatius, Bromus erectus and others). Widespread in Germany and very common, frequently found up to 1000 m a.s.l. in the Alps and on the Swabian Jura, but usually below 700 m a.s.l. elsewhere. Records from higher altitudes probably refer to stray individuals. Has been identified as a vector of various mycoplasma-like organisms, such as Clover phyllody (CP), Clover dwarf (CD), Stolbur and Parastolbur in various European countries. Fischer (1972), Niedringhaus (1991), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN; see also Brˇcák (1979), Cook (1996)
Euscelis lineolatus Brullé, 1832 E V – B XI; egg, 1 gen. This species is widespread in the Mediterranean region and western Europe. In Germany it is restricted to the extreme western parts. Border localities are near Prüm and
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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 Bremen. 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 England 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 Brˇcák (1979), Prestidge & McNeill (1983)
Euscelis ohausi W. Wagner, 1939 E VI – M X; egg, 1 gen. This species is confined to western parts of Europe; it is only reported from Portugal, France, Switzerland, Germany, Belgium, the Netherlands, Britain (northward to Scotland), Jutland and northwestern Poland. It lives on woody species of Fabaceae in sunny, occasionally also in moderately shady, damp to moderately dry sites, on acidic substrates (mainly sand and sandstone). There are two morphs in Germany, which are commonly regarded as subspecies: the ssp. ohausi W. Wagner, 1939 is found on Genista anglica, in heath and dune areas of northern Germany (Lüneburger Heide, in the vicinity of Hamburg and Ludwigslust, and on the north Frisian islands of Amrum and Sylt); the ssp. singeri W. Wagner, 1951 lives on Cytisus scoparius along margins of low-productivity grassland and forest glades, mainly in middle and southern parts. Border localities, some of which appear to be rather isolated, are near Cologne, Marburg, Nordhausen, Torgau an der Elbe, Niesky, Aschaffenburg, Erlangen and Oberstdorf im Allgäu (800 m a.s.l.). Is apparently absent from the Czech Republic and Austria. Frommer (1996), Remane (1967, and pers. comm.), Schiemenz et al. (1996), Strübing & Schwarz-Mittelstaedt (1988), Trümbach (1959), Wagner (1951a), Walter (pers. comm.), HN
Euscelis venosus (Kirschbaum, 1868) M VII – E IX; egg, 1 gen. In oligotrophic, basic, damp to moderately dry (also temporarily dry), usually shortvegetated pastures (occasionally meadows), in the Alps mainly on sun-exposed hillsides, in lower altitudes also on north-facing slopes. The host plants are unknown, but are likely to include species of Asteraceae (perhaps Carlina acaulis and C. vulgaris). In Germany the species is restricted to the southern half and occurs at the northern edge of the range, with strongholds in the Bavarian and Allgäu Alps and their foothills, where it is locally fairly common up to at least 1500 m a.s.l., but very sporadic in middle parts. Border localities are near Mainz (locus typicus), in the Vogelsberg and Meißner, near Herzberg am Harz, Stendal and Potsdam. Fischer (1972), Nikusch (1976), Remane (pers. comm.), Schiemenz et al. (1996), Wagner (1939a, 1951a), Nickel (1994), HN
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Review of species
Ederranus discolor (J. Sahlberg, 1871) So far B VI – B IX; probably egg, 1(?) gen. At present, this species is only known from northern Russia (Karelia), southern Finland and Brandenburg; its habitat requirements are poorly known. Phragmites australis and Glyceria maxima have been reported as host plants; thus, the species is likely to live in rather wet sites. The two German record are from inland salt marshes with Phragmites australis and Juncus gerardii near the cities of Brandenburg and Potsdam: Trechwitz, Netzener See, c. 30m a.s.l., 05.VI.1993, 1 †, 09.IX.1993, 4 ††, 1 ‡, and Gröben, Gröbener See, c. 30m a.s.l., 10.IX.1993, 6 ††, 4 ‡‡. Fröhlich (1996a), Remane & Fröhlich (1994b); see also Emelyanov (1964a), Lindberg (1947), Ossiannilsson (1983), Sahlberg (1871)
Sotanus thenii (P. Löw, 1885) So far B VII – B VIII, in Tyrol (Austria) E VI – E IX; nymph, 1 gen. (Dobler 1985; Leising 1977) This species is confined to grasslands of damp, well-drained sites of the subalpine to the upper alpine belt of the Alps; probably it lives on an unidentified species of Poaceae. From Germany it is only known from 5 localities in the Allgäu near Oberstdorf, between 1700 and 2050 m a.s.l.: Hochvogel, Rotspitz, Nebelhorn, Rotkopf and Mindelheimer Hütte (below Liechelkopf), but it is probably more widespread in Bavaria. Otherwise only reported from Austria, Switzerland and northern Italy, where found between 1500 and 3050 m a.s.l. Fischer (1972), Schönitzer & Oesterling (1998b), HN; see also Dobler (1985), Günthart (1987a), Leising (1977), Nast (1987), Thaler (pers. comm.), Wagner & Franz (1961)
Streptanus aemulans (Kirschbaum, 1868) M VI – E X; egg, 2(?) gen. In grass stands in sunny to moderately shady, usually damp to moist, occasionally also moderately dry or wet sites (mainly meadows, pastures, forest glades, ruderal habitats, waysides, also white and grey dunes near the coast and moderately saline sites). Host plants are various grasses, often Holcus lanatus, H. mollis and Elymus repens. Widespread in Germany and common in most regions, but usually below 900 m a.s.l. Achtziger (1991), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Wagner (1935, 1951a), Walter (pers. comm.), Nickel & Achtziger (1999), HN, and others
Streptanus confinis (Reuter, 1880) E VI – B X; egg, 1 gen. Among tall grasses in sunny to shady, peaty to moist (preferentially waterlogged) sites, mainly abandoned or low-input meadows and pastures, fens and fen woods, often in rather cool situations, locally also in moderately saline sites. Usually (perhaps exclusively) deep in tussocks of Deschampsia cespitosa; seldom recorded due to secretive life habits. In Germa-
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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 Netherlands, Poland and Austria; in France only known from a single locality in the Jura, but probably under-recorded. The highest localities are at 800 m a.s.l. in the Rhön Mountains, but single specimens have been found at 1800 m a.s.l. in the Bavarian Alps. Bittner & Remane (1977), Niedringhaus & Olthoff (1993), Reimer (1992), Remane & Fröhlich (1994b), Walter (1998), Nickel & Achtziger (1999), HN; see also della Giustina & Remane (2001), Nast (1987)
Streptanus marginatus (Kirschbaum, 1858) M IV – E VIII; nymph, 1 gen. In moderately shady, moderately moist to moderately dry sites, mainly in open deciduous and coniferous forests, also along forest margins in dry grassland, heaths and montane meadows. Host plants are fine-leaved grasses, mainly Deschampsia flexuosa and Festuca ovina, perhaps also Nardus stricta and further species. Widespread in Germany and fairly common, found up to at least 1000 m a.s.l. in the Mittelgebirge and the Alps. Achtziger (1991), Bornholdt (1996), Fischer (1972), Nikusch (1976), Reimer (1992), Remane (1987), Schiemenz (1971a), Schiemenz et al. (1996), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN
Streptanus okaensis Zachvatkin, 1948 This taxon is perhaps conspecific with S. ogumae (Matsumura, 1911) described from Sakhalin. The question is left open here until the type material has been revised.
E VI – B X; egg, 1 gen. In Europe known only from Russia, Scandinavia, central Spain and the north German plain, where recorded in c. 20 localities in Niedersachsen, Holstein, Brandenburg and the Oberlausitz, up to at most 150 m a.s.l. It lives on Calamagrostis canescens in acidic, wet, occasionally peaty, sunny to moderately shady sites (mainly fens, intermediate bogs and fen woods). Is probably under-recorded due to secretive life habits. Remane (1995), Remane & Fröhlich (1994b), Niedringhaus & Olthoff (1993), Walter (pers. comm.), HN
Streptanus sordidus (Zetterstedt, 1828) M VI – M X, according to Schiemenz et al. (1996) M V – M XI; egg, 2(?) gen. In moist to peaty or temporarily flooded, sunny to moderately shady sites, mainly lowinput pastures and meadows, fens, intermediate bogs and salt marshes, but also open forests and clearings, rarely in conventionally managed grassland. Host plants are various grasses, preferentially Agrostis stolonifera and A. capillaris, perhaps also Deschampsia cespitosa and others. The species is widespread in Germany and fairly common; it occurs up to at least 1200 m a.s.l. in the Allgäu and Black Forest, but has been found at 1700 m a.s.l. in Styria (Austria). Achtziger (1991), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Nikusch (1976), Reimer (1992), Schiemenz et al. (1996), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN; see also Wagner & Franz (1961)
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Review of species
Artianus interstitialis (Germar, 1821) E VI – B X; egg, 1 gen. In scattered stands of grasses in moderately dry to dry sites, usually on sandy, locally also loamy substrates, notably in ruderal sites, abandoned fields, along waysides and in disturbed patches in dry grassland, occasionally in temporarily dry salt marshes. Lives on various grasses, often Elymus repens, perhaps also Agrostis capillaris, Festuca rubra and Holcus spp. Widespread and locally common in southern and eastern parts of lowland Germany (frequently up to c. 400 m a.s.l., occasionally up to 700 m a.s.l.), at the northwestern edge of the range. Border localities are on a line from Cologne to Marburg, Gotha, Nordhausen, Quedlinburg, Wolfsburg and Lauenburg. Records published from the Netherlands are erroneous. Fröhlich (1996a), Frommer (1996), Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Thüs (1996), Trümbach (1959), Wagner (1935), HN; see also Metcalf (1967), Nast (1987), Reclaire (1944)
Paramesus obtusifrons (Stål, 1853) and P. major Haupt, 1927 Paramesus major Haupt, 1927 has been described from Israel and reported from most parts of the Palearctic since (from eastern Siberia to the eastern Mediterranean region, Poland, the Czech Republic and Hungary). Differences between eastern European populations and P. obtusifrons (Stål), which is confined to central and western Europe, are weak and only concern the shape of the terminal aedeagus. Thus, the specific distinctiveness of these two taxa may be uncertain, although no transitional individuals have been found so far. †† of all German inland populations are not distinct from those living in eastern Europe, which are currently treated as P. major Hpt. (see Anufriev & Emelyanov 1988). From Poland Nast (1976a) mentions P. major Hpt. from inland sites and P. obtusifrons (Stål) from the coast. However, it is unknown whether the ranges overlap along the Mediterranean and the Baltic Sea.
B VII – B X; egg, 1 gen. Monophagous and halobiotic on Bolboschoenus maritimus, usually in reeds in brackish and temporarily flooded sites along the coast and inland. Recorded from all East Frisian Islands, the Weser estuary near Bremerhaven, the island of Sylt, and the coast of the Baltic Sea between Schleswig and Hiddensee (P. obtusifrons Stål), as well as inland salt marshes (rarely in freshwater sites with strongly fluctuating water tables) in the rain shadow area to the east of the Harz Mountains (roughly between Magdeburg, Halle, Erfurt and Sondershausen), in eastern Niedersachsen (Schreyahn), the Werra valley (Ettmarshausen), the Wetterau (Wisselsheim, Münzenberg), and in the northern upper Rhine valley near Speyer (P. major Hpt.). The highest localities are at 250 m a.s.l. only. Published feeding records on Schoenoplectus are probably erroneous. Fröhlich (1996a), Kuntze (1937), Hildebrandt (1995), MTD, Niedringhaus (1991), Schaefer (1973), Schiemenz et al. (1996), Wagner (1935), HN
Parapotes reticulatus (Horváth, 1897) M VII – B X; egg, 1 gen. In tussocks of Schoenoplectus lacustris, usually in permanently flooded reeds along shores of fishponds, glacial and oxbow lakes, a single individual also found in a brackish spring
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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 Lauterberg, Ilmenau and Görlitz. Is probably under-recorded. Has been reported from salt marshes of eastern Austria and Hungary. Höhne (1921), Sander et al. (1999), Schiemenz et al. (1996), Nickel (1997), HN; see also Fröhlich (1996a),
Paralimnus phragmitis (Boheman, 1847) B VII – B X; 1 gen., in southwestern Germany from E V onwards, 2(?) gen. Monophagous on Phragmites australis in mesotrophic to moderately eutrophic, temporarily or non-flooded sites, usually along shores of lakes and ponds, in fens, mining areas, coastal and inland salt marshes. Widespread in Germany, but only scattered; often sampled in low numbers only. The highest localities are at 700 m a.s.l. in the Bavarian foothills of the Alps. Fröhlich (1996a), Heller (1987a), Hoffmann (1980), Niedringhaus (1991), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1939a), Nickel (1999b), HN
Paralimnus rotundiceps (Lethierry, 1885) So far only IX, in Austria M VI (Moosbrugger 1946), egg(?), 1(?) gen. This is a rare species of European distribution, living on Phragmites australis. In Germany it has only recently been discovered in two widely separated localities: coal mine of Mücheln near Merseburg, c. 150 m a.s.l., several records in low-growing and sparse pioneer stands of reed on rather dry, sandy soil, e.g. 03.IX.2001, 2 ††, 1 ‡, and Griesen near Garmisch-Partenkirchen, c. 900 m a.s.l, IX 2000, 1 †, in a Malaise trap on a gravel bank. Further known localities are in Vorarlberg (Austria), the Valais (Switzerland) and southern Scandinavia. Funke (pers. comm.), Al Hussein et al. (1999), Nickel & Voith (unpublished data); see also Cerutti (1939b), Fröhlich (1996a), Moosbrugger (1946), Ossiannilsson (1983)
Metalimnus formosus (Boheman, 1845) Mainly M VI – E X, according to Fischer (1972) also E V; egg, 1 gen., perhaps 2 in lowland sites. Among tall sedges in wet to temporarily flooded, often peaty sites, usually in fens, straw meadows, along shores of oligotrophic to moderately eutrophic lakes and ponds, as well as in floodplain depressions. Host plants are tall-growing species of Carex (notably C. acuta and C. elata, perhaps also C. nigra). The species is widespread in Germany, but occurs only localized, with highest frequencies in river valleys from the Elbe eastward as well as along the edge of the Alps (up to at least 900 m a.s.l., but found at 1500 m a.s.l. in North Tyrol, Austria). Fischer (1972), Fröhlich (1996a), Hoffmann (1980), Remane (1962), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), SMNS, Wagner (1939a), Wagner & Wagner (1938), HN; see also Cobben & Gravestein (1958), Ribaut (1952)
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Review of species
Metalimnus steini (Fieber, 1869) Has been described as a new variety of the preceding taxon, from the surroundings of Berlin, but the whereabouts of the type material are unknown. Later it was treated as a distinct species, and M. tredecimpunctatus (Lindberg, 1929), described from Japan, as a junior synonym. It has been reported from large parts of the Palearctic, including Hungary, Ukraine, Austria, Kazakhstan and far-eastern Russia (Anufriev & Emelyanov 1988; Holzinger et al. 1997; Nast 1972; Remane & Fröhlich 1994a). However, the taxonomic situation is rather critical, since the identity of M. steini (Fieb.) is not clear, and since it is likely to comprise a complex of species.
So far M VI – M X; egg, 2(?) gen. The locus typicus is near Berlin, but there are no more verified records since, although 2 dark-coloured ‡‡ from the collection of Haupt, taken near Aken an der Elbe, 23.VII.1911, are likely to belong to this taxon. Furthermore, there are three very recent records from the upper Rhine plain between Freiburg and Darmstadt (100 – 250 m a.s.l.), in stands of Carex hirta in sunny to moderately shady, moderately wet to moderately dry ruderal habitats and meadows. However, in the Rhine populations conspecificity with the type material has yet to be confirmed. Fieber (1869), MTD, HN
Arocephalus longiceps (Kirschbaum, 1868) M V – M X; egg, 2 gen., perhaps 1 at higher altitudes. In sunny to moderately shady, moderately moist to moderately dry sites, mainly clearings, forest glades, dry grassland, heaths, ruderal sites and abandoned fields, occasionally also open forests and low-input-meadows. Single individuals on dispersal flight are frequently encountered in other habitats. Host plants are various species of Poaceae, notably Holcus mollis, H. lanatus and Bromus erectus. Widespread in Germany and fairly common in middle and southern parts, but more scattered in the north German plain, where living at the edge of the range. Northernmost localities are on the East Frisian Islands, near Pinneberg, Plön and Greifswald (but has recently been found in Skåne, southern Sweden). Found up to at least 1100 m a.s.l. (but single specimens at 1600 m a.s.l.) in the Mittelgebirge and Bavarian Alps, at 1730 m a.s.l. in the Engadine (Switzerland). Achtziger (1991), Fischer (1972), Niedringhaus & Olthoff (1993), Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1939a), Nickel & Achtziger (1999), HN; see also Gillerfors (2002), Günthart (1987a)
Arocephalus languidus (Flor, 1861) M V – B X; egg, 2 gen., 1 at higher altitudes. In moderately dry to dry, sunny to moderately shady sites on basic as well as acidic, gravelly to loamy substrates, usually in various types of xerothermic grassland (also with open stands of pine or oak) as well as in pastures and meadows of the subalpine and alpine belt. Host plants are Sesleria albicans and Stipa spp., probably also Koeleria glauca and other grasses. Rather localized in Germany, at the northwestern edge of the range, and with strongholds in the Alps and their foothills, as well as in warmer regions of southern and eastern parts, often among the dominant species in favourable sites.
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Border localities are on a line from Koblenz to Schlüchtern, Eschwege, Nordhausen, Magdeburg and Rügen. Frequently found up to at least 2000 m a.s.l. in the Bavarian and Allgäu Alps. Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987, and pers. comm.), Schiemenz et al. (1996), Wagner (1939a, 1951), Nickel (1994), HN
Arocephalus punctum (Flor, 1861) B VI – E X; egg, 2 gen., perhaps 1 at higher altitudes. In oligotrophic, sunny to slightly shaded, low-vegetated, dry to moist (rarely wet), acidic as well as basic sites; in the north German plain favouring sandy dry grassland, heaths and forest margins, in the region of the Mittelgebirge mainly on meadows and pastures of the submontane and montane belt, locally also in dry grassland on limestone, sand and other dry substrates at low altitudes. Lives in tussocks of fine-leaved grasses, mainly Festuca ovina, perhaps also F. rubra, Nardus stricta and Corynephorus canescens. Widespread in northern and middle parts of Germany and locally common, but not recorded to the south of the Danube; the highest localities are at 1250 m a.s.l. in the Black Forest. Niedringhaus (1991), Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Schiemenz et al. (1996), Trümbach (1959), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others
Arocephalus sagittarius Ribaut, 1952 So far only M VI, in France also B IX (della Giustina 1989); egg, 2(?) gen. Has very recently been found in the Kaiserstuhl in extreme southwestern Germany: Badberg, 14.VI.2002, small populations in several patches of xerothermic loess grassland, apparently among fine-leaved grasses (perhaps Festuca ovina). Was already reported from adjacent parts of Alsace (France), but is otherwise only known from Switzerland, Spain and Portugal. Lauterer & Malenovsky (pers. comm.), Remane (pers. comm.), Nickel & Remane (2002), HN; see also della Giustina (1989), Remane & Fröhlich (1994b)
Psammotettix maritimus (Perris, 1857) E VI – M IX; egg, 1(?) gen. This species is an endemic of western European coasts and lives monophagously on Ammophila arenaria in calcareous primary and white dunes close to the shoreline. From Germany it is known only from the East and North Frisian Islands (Borkum, Memmert, Juist, Norderney, Baltrum, Langeoog, Spiekeroog, Wangerooge, Amrum and Sylt). Otherwise it is reported only from England, the Netherlands, Belgium, France and Portugal. Niedringhaus (1991), Niedringhaus & Olthoff (1993), Remane (pers. comm.), Wagner (1935); see also Nast (1987)
Psammotettix poecilus (Flor, 1861) M V – B X; egg, 2 gen.
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Review of species
In sandy to gravelly, usually sunny, acidic as well as basic sites with very scattered stands of grass, mainly inland dunes, mining areas, forest clearings and almost bare gravel banks of alpine rivers. Lives on Calamagrostis epigejos in lowland sites, but on C. pseudophragmites in the Alps and their foothills. In Germany at the western edge of the range, widespread and locally fairly common in the sand areas of northeastern parts (westward at least to Greifswald, Brandenburg, Halle, Leipzig and Bautzen), but rather sporadic and in strongly isolated populations elsewhere, notably in the Rhine-Main region, Mittelfranken, and along the upper course of the Isar between Mittenwald and Lenggries, where found up to 900 m a.s.l. There are also single localities in the Kyffhäuser and northern Hessen. Not recorded from Denmark, the Netherlands and Belgium; in France known only from the western Alps (Savoie). Funke & Witsack (1998), Remane (1987), Remane & Fröhlich (1994b), Schiemenz et al. (1996), Wagner (1939a, 1951a), Nickel (1999b), HN; see also della Giustina & Remane (2001)
Psammotettix unciger Ribaut, 1938 So far only B VIII – E VIII; 1(?) gen. This is probably the rarest species treated in this book. Since the original description, it has been recorded only twice. The locus typicus is near Oberstdorf im Allgäu: Iller floodplains; 04. and 07.VIII.1936; also North Tyrol (Austria): Lech valley to the northwest of Reutte, VIII 1973; and Upper Bavaria: Isar floodplains near Vorderriß, 26.VIII.1997, 6 ††, 8 ‡‡. It lives on very scattered grass stems on almost bare gravel banks of unchanneled alpine rivers; specimens taken near Vorderriß were swept from Calamagrostis pseudophragmites. All localities are at c. 800 m a.s.l. The Iller population is probably extinct due to draining of the river bed. Remane & Fröhlich (1994b), Ribaut (1938) (see this paper also for identification), Nickel (1999a)
Psammotettix kolosvarensis (Matsumura, 1908) E V – E IX; egg, 2 gen. Usually in low-growing grass stands in sunny, moderately wet to moderately dry sites with strongly fluctuating levels of fresh or moderately saline water, mainly inland salt marshes, sandy abandoned fields, as well as temporarily dry meadows and pastures, also in more intensively used sites. Single individuals on dispersal flight are frequently encountered in other habitats. Is often found on Puccinellia distans and Elymus repens, but probably feeds also on additional grasses. In Germany largely confined to eastern parts, probably at the edge of the range, with strongholds in the basins of the Elbe and Oder, where locally abundant in riverine grassland, which is frequently flooded during winter, but may become rather dry during summer. Border localities are near Schwedt, Rathenow, Lauenburg, Wolfsburg, Sondershausen, Meißen and Görlitz, up to 200 m a.s.l. To the west of Germany there is only a single record from the Gironde area (France). Fröhlich (1996a), Schiemenz et al. (1996), Rothenbücher (pers. comm.), Walter (1996), Nickel & Achtziger (1999), Nickel (1997), HN; see also Ribaut (1959)
Deltocephalinae Fieber, 1869
211
Psammotettix angulatus (Then, 1899) Regarding genital morphology, this taxon closely resembles Ps. koeleriae Zachvatkin, 1948, which is reported from middle Russia to Mongolia. There are differences, however, in body colouration: Ps. angulatus Then is straw-coloured, Ps. koeleriae Zachv. is green. According to Remane (1965), they should preliminarily be treated as distinct species, until further evidence has been provided.
So far M VI – E VII; egg, 2(?) gen. Known from Germany only from a single site in Brandenburg, on a xerothermic, sandy, south-facing moraine slope: Groß Machnower Weinberg near Zossen, 70 m a.s.l. („Schillergras-Schafschwingelrasen mit Elementen der Silbergrasflur auf einer Waldlichtung“), 12.VI.1964, 1 †, 1 ‡, 27.VII.1964, 1 ‡. Koeleria glauca may have been the host plant. In 1998 the site was studied again, but without a positive record. Hence, the species is perhaps extinct in Germany. Otherwise it is reported only from single localities in Bohemia (Czech Republic), central Poland and northern Italy. Schiemenz (1969) (Remane det.), Schiemenz et al. (1996); see also Nast (1972, 1987); for identification see Remane (1965)
Psammotettix nardeti Remane, 1965 M VII – E IX; egg, 1 gen. This is an endemic species of the Alps, where it is apparently confined to western parts. Lives in meadows and pastures of the subalpine to the upper alpine belt, probably on a specific grass (Nardus stricta has been suggested). In Germany it is known only from 4 records from the Allgäu, the Ammergebirge, and the Berchtesgaden Alps, between 1600 and 2000 m a.s.l. (Untere Gottesackerwände above Oberstdorf, Hochplatte above Füssen, Friederspitz above Garmisch-Partenkirchen, and Funtensee above Berchtesgaden). Otherwise it is reported only from Austria, France and Italy, between 1550 and 2600 m a.s.l. Remane (1965), HN; see also Günthart (1984, 1987a), Leising (1977), Nast (1987); for identification see Remane (1965)
Psammotettix inexpectatus Remane, 1965 So far M VII – B X; egg, 1(?) gen. For a long time, this species was known only from the locus typicus in Thuringia, until there was a second record from Mount Olymp in Greece. In Germany it is restricted to southwest-facing, xerothermic gypsum hillsides in the Kyffhäuser near Bad Frankenhausen, c. 200 to 270 m a.s.l. From this locality there are altogether 5 records: 29.VIII.1956, 1 †, 3 ‡‡; 10.X.1965, 1 †, 2 ‡‡; 17.VII.1991, 1 †, 1 ‡; 06.IX.1996, 2 ††, 1 ‡; 02.IX.2000, 4 ††, 5 ‡‡. No details are known on the biology; the host plant is probably a non-identified species of grass. Fröhlich (pers. comm.), Remane (1965), Schiemenz et al. (1996), Nickel et al. (2001), HN; see also Drosopoulos et al. (1986); for identification see Remane (1965)
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Review of species
Psammotettix alienus (Dahlbom, 1850) Mainly E V – M X, occasionally B V – B XI; egg, 2 gen. This is a eurytopic pioneer species in sunny, dry to wet sites, mainly in fallow and cereal fields, meadows, mining areas and slightly saline sites. Invades all types of tree-less habitats and is among the very few leafhopper species of fertilized meadows and pastures, although individual numbers may fluctuate strongly, and breeding success may be reduced in these places. Host plants are various grasses. Widespread in Germany and particularly common in the southern half, below 900 m a.s.l., but less frequent in the north German plain. Was recorded as a vector of Band mosaic of wheat and rye (BMWR), Wheat dwarf virus (WDV), and perhaps additional plant diseases in various countries. WDV on cultivated barley has recently become a problem in parts of SaxonyAnhalt and also Sweden. Fischer (1972), Müller (1956), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Wagner (1935, 1939a), Manurung et al. (2001), Nickel & Achtziger (1999), HN, and others; see also Brˇcák (1979), Lindblad & Arenö (2002); for nymphal morphology see Guglielmino & Virla (1997)
Psammotettix sabulicola (Curtis, 1837) E V – E IX; egg, 2 gen. On tall but scattered grasses on coastal dunes and in sandy, sun-exposed and open habitats inland. Ammophila arenaria and Elymus spp. are possible host plants, although some specimens have been found among Leymus arenarius and Calamagrostis epigejos. In Germany the species is confined to the northern plains, with strongholds in coastal areas near the North and Baltic Sea, where it is locally fairly common on white, grey and brown dunes, but there is also a number of localities up to c. 70 km inland, notably near Oldenburg, Winsen, Boizenburg and Neuhaus. The range is apparently confined to northern and western European coastal lowlands; records from Italy and Bulgaria may be doubtful. Niedringhaus (1991), Remane (1958, and pers. comm.), Schaefer (1973), Schiemenz et al. (1996), Wagner (1935), HN; see also della Giustina & Remane (2001), Gravestein (1965)
Psammotettix pallidinervis (Dahlbom, 1850) E V – M X; egg, 2 gen. In sparsely vegetated, xerothermic grassland, in northern Germany mainly on sand, in middle and southern parts also on (preferentially grazed) gypsum and limestone sites . Lives on fine-leaved species of fescue (Festuca ovina group), but has been reported from Corynephorus canescens. In Germany the species occurs at the southwestern edge of the range, only very localized, and usually in low abundances, mainly in the north German plain and the rain shadow area to the east of the Harz Mountains; further localities, which appear to be rather isolated, are near Lingen (Ems), Celle, Bad Wildungen and Regensburg, up to at most 400 m a.s.l. It is reported from the Netherlands and Austria, but not from Belgium, France, Switzerland and Italy. Remane & Fröhlich (1994b), Schaefer (1973), Schiemenz et al. (1996), Wagner (1937a), HN; see also Nast (1987)
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Psammotettix cephalotes (Herrich-Schäffer, 1834) E V – M X; egg, 2 gen., at higher altitudes probably only 1. On Briza media in oligotrophic and sunny sites; at lower altitudes the species is largely restricted to grazed, dry grassland on basic soils; at submontane and higher altitudes it is more widespread and common, favouring peaty to moderately dry low-productivity meadows and pastures (also on acidic substrates). Widespread in Germany, but apparently absent from the northwest German plain to the west of the Elbe. Lives up to at least 1500 m a.s.l. in the Bavarian and Allgäu Alps, up to 2200 m a.s.l. in the Engadine (Switzerland) and North Tyrol (Austria). Reimer (1992), Rombach (1999b), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Nickel & Achtziger (1999), Nickel (1994), HN; see also Günthart (1987a), Leising (1977)
Psammotettix helvolus (Kirschbaum, 1868) group According to Remane & Fröhlich (1994a), this is a „Formengruppe mit ungeklärter taxonomischer Wertigkeit der verschiedenen Formen, z.B. P. rhombifer (Fieber, 1869), P. substriatus (Then, 1902), P. obtusiceps (Kirschbaum, 1868)“. Remane (1987) has provisionally named two forms occurring in Germany which are morphologically and ecologically distinct. Accordingly, „helvolus basic“ is macropterous and lives on various grasses, mainly in neutral to basic, and dry sites of lower altitudes (usually below 800 m a.s.l.), whereas „helvolus acidic“ is subbrachypterous and is usually found in acidic sites of higher altitudes (usually above 300 m a.s.l.). The latter prefers Deschampsia flexuosa as host plant. See also Günthart (1987a), Leising (1977), Wagner & Franz (1961). Biosystematic studies are needed to clarify the taxonomic situation.
B V – M X; egg, 2 gen., at higher altitudes only 1. In sunny, occasionally slightly shady, dry to moist sites on basic to acidic substrates. Particularly abundant in low-input meadows, dry grassland and abandoned fields, also found in pastures, forest clearings, open forests, and along waysides, occasionally in fertilized meadows. Widespread and common in southern and middle parts of Germany, but known only from a few localities in the north German plain (recorded near Lüchow, Zossen and Greifswald). It lives up to at least 1900 m a.s.l. in the Allgäu Alps, up to 2600 m a.s.l. in the Engadine (Switzerland). Records from even higher altitudes probably not refer to resident populations, but only to stray individuals. Fischer (1972), Reimer (1992), Remane (1987), Rombach (1999b), Schiemenz et al. (1996), Wagner (1939a, 1951a, 1958), Walter (1996), Nickel & Achtziger (1999), Nickel (1994), HN; see also Dobler (1985), Günthart (1984, 1987a), Leising (1977), Wagner & Franz (1961)
Taxa belonging to the Ps. nodosus group, comprising Ps. nodosus (Rib.), Ps. albomarginatus W.Wg., Ps. putoni (Then), Ps. excisus (Mats.) and Ps. dubius Oss., are morphologically little distinct (in particular considering the aedeagus shape), but show specific habitat requirements and – at least in some cases – also host preferences. Their taxonomic status is not satisfactorily clarified.
Psammotettix albomarginatus W. Wagner, 1941 E V – E X; egg, 2 gen. In oligotrophic, sunny and sparsely vegetated sites, usually with a high proportion of lichens, probably on Agrostis vinealis or Corynephorus canescens, preferentially on sandy substrates, but also found on porphyry. So far known from Germany only from a few
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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, Denmark, the Netherlands and Poland. Remane (1958, and pers. comm.), Schaefer (1973), Schiemenz (1969), Schiemenz et al. (1996), Wagner (1941a), HN; see also Kirby (1992), Nast (1987)
Psammotettix putoni (Then, 1898) E V – B X; egg, 2 gen. Halobiotic in salt marshes; often abundant in salt meadows near the North and Baltic Sea (eastward at least to Kiel), also found in two inland sites of northern Germany (Barnstorf near Braunschweig, and Sülldorf near Magdeburg). Is reported to live on Puccinellia maritima and perhaps P. distans. All localities are below 150 m a.s.l. Otherwise the species is only published from Norway, Sweden, northern Russia, Ireland, Britain, Denmark, the Netherlands, Belgium and France. Records from southeastern Europe require verification. Fröhlich (1996a), Niedringhaus (1991), Remane & Fröhlich (1994b), Tulowitzki (1990), Wagner (1941a), HN; see also Schiemenz et al. (1996),
Psammotettix dubius Ossiannilsson, 1974 In North Tyrol (Austria) according to Leising (1977) M VIII – M IX; egg, 1 gen. So far only known from Norway and Sweden (northward to c. 65o N), middle Russia, North Tyrol and Bavaria. Reported to live in Scandinavia „ ... on grasses on dry, sandy soils, ... also in heather marshes and dry moors“, in North Tyrol in alpine peaty meadows and bogs, between 1900 and 2100 m a.s.l. Has recently been found in the Allgäu Alps: Oberjoch above Hindelang, 1200 m a.s.l., 05.VIII.1995, 4 ††, 2 ‡‡, at the edge of a raised bog. Nickel (1999b), HN; see also Anufriev & Kirillova (1998), Leising (1977), Ossiannilsson (1983)
Psammotettix excisus (Matsumura, 1906) M V – E X; egg, 2 gen. In sandy, xerothermic, sparsely-vegetated and acidic sites, mainly inland dunes, heaths, military training areas and sand pits, preferentially on Corynephorus canescens, perhaps also on other grasses. Widespread and fairly common in the north German plain and locally dominant in favourable sites, but apparently absent or rare on coastal dunes of all kinds; also found in scattered localities in middle and southern parts of Germany, notably in inland dune areas in the northern upper Rhine plain (between Speyer and Darmstadt), in the valleys of the Main (near Karlstadt, Kitzingen, Volkach), Regnitz (near Bamberg, Forchheim, Erlangen) and Danube (Siegenburg near Kelheim), as well as in the basin of Mittelfranken (Altdorf near Nuremberg), up to 450 m a.s.l. Otherwise only reported from Estonia, Lithuania, Poland, Denmark, the Netherlands, France, Portugal, the Czech Republic and Hungary. Remane (1958, 1965), Schiemenz et al. (1996), Trümbach (1959), Wagner (1941a), Nickel (1999b), HN
Deltocephalinae Fieber, 1869
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Psammotettix nodosus (Ribaut, 1925) M V – M X; egg, 2 gen. This is a pioneer species of sunny, dry to damp and sparsely vegetated sites on various substrates, mainly in strongly grazed and oligotrophic pastures, in dunes and ruderal sites, but also in forest clearings, in dried peat diggings, on roadside embankments, etc.; host plants are various, usually low-growing grasses, often Festuca ovina. Widespread in the lowlands of Germany, breeding takes place up to at least 800 m a.s.l., single individuals have been found up to 1700 m a.s.l. in the Bavarian Alps. Widespread in Germany, locally abundant in sandy sites of the north German plain, but more scattered and in lower densities elsewhere. Fischer (1972), Niedringhaus (1991), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Wagner (1939a, 1941a), HN, and others
Psammotettix notatus (Melichar, 1896) Has been misinterpreted for a long time. In fact, almost all published records (see Metcalf 1967; Nast 1987) are in need of revision.
So far only B IX – M IX; egg(?), 1(?) gen. This species is apparently very rare, and almost nothing is known on its life history. There are two German records: Lower Bavaria, Saal an der Donau, c. 400 m a.s.l., 02.IX.1960, 1 †, and Mainfranken, Klotz near Retzbach, c. 250 m a.s.l., 15.IX.1994, 2 ‡‡, both on xerothermic calcareous hillsides. The host plants are presumably grasses. Otherwise, only the locus typicus is known (“Nieder-Oesterreich, am Eichkogel bei Mödling, im August und September”). Most, if not all, other published records are based upon misinterpretation. Remane (1961b, and pers. comm.), HN; see also Melichar (1896), Wagner (1939a) (see this paper also for identification)
Psammotettix confinis (Dahlbom, 1850) Psammotettix remanei Orosz, 1999 (see Nickel & Remane 2002) This taxon shows a considerable variation of colouration, body size and aedeagus shape. Recently the original description of Psammotettix remanei Orosz, 1999 was published, with paratypes from the Oberlausitz. However, the type specimens were clearly individuals whose genital apparatus was not fully developed (perhaps parasitized), notably regarding the genital plates, lateral lobes of the pygophor, as well as the base and shaft of the aedeagus. Moreover, the connective was rather abnormal and did not even show the typical Paralimnini shape. Such individuals of Ps. confinis (Dhlb.) with rudimentary genital apparatus, reduced body size and pigmentation are more or less frequently found among central European populations. Only some of them show visible external or internal signs of parasitation, but their morphology is clearly abnormal.
M V – M X; egg, 2 gen. Eurytopic in grass stands of sunny, dry to moderately wet, preferentially neutral to acidic sites, often abundant in pastures, fertilized meadows and ruderal sites, but also on inland dunes, in wet meadows and moderately brackish sites. Host plants are various grasses, probably Lolium, Poa, Festuca and Agrostis. Widespread all over Germany
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and very common, found up to at least 1200 m a.s.l. in the Bavarian Alps, up to at least 1900 m a.s.l. in subalpine meadows of North Tyrol (Austria). Bornholdt (1996), Emmrich (1966), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Reimer (1992), Remane (1958, 1987), Schiemenz (1969), Schiemenz et al. (1996), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN, and others; see also Leising (1977), Prestidge & McNeill (1983a, 1983b)
Ebarrius interstinctus (Fieber, 1869) The identity of this taxons remained unclear for a long time, until Nast (1977) published a generic revision. Specimens from the French Alps with slight differences compared to the nominate form were described as E. interstinctus sofiae della Giustina, 1989.
M VI – E IX; egg, 2(?) gen. Reported from alpine dwarf shrub heaths of the French and Swiss Alps, between 2100 and 2500 m a.s.l. In Germany, however, there are only two known lowland localities: northwestern slopes of the Swabian Jura: Balingen, c. 500 m a.s.l., 23.VI.1955, a population on a sun-exposed roadside embankment, as well as Nahe valley near Schloßböckelheim, c. 250 m a.s.l., between mid June and end of September 1995, several specimens in Malaise traps, in a xerothermic, abandoned vineyard. Probably lives on grasses. Otherwise reported only from France, Switzerland, Austria, Hungary, northern Italy, former Yugoslavia, Bulgaria and Greece. Remane & Fröhlich (1994b), Fröhlich & Nickel (unpublished data); see also Drosopoulos et al. (1986), della Giustina (1989), Günthart (1987a), Nast (1987)
Ebarrius cognatus (Fieber, 1869) M VII – E IX; egg, 1 gen. In oligotrophic, sunny, short-vegetated sites of the montane to the alpine belt, locally also at lower altitudes along alpine rivers; mainly in grassland with predominating Sesleria albicans, Carex ferruginea or dwarf shrubs. Host plants are grasses. Has been taken on Festuca vivipara in Scotland, which does not occur in central Europe, however. In Germany confined to the Bavarian and Allgäu Alps, where apparently widespread between 1500 and 1850 m a.s.l., occasionally on riverine gravel banks of lower altitudes (Wimbach valley near Ramsau, between 800 and 1400 m a.s.l.). Other lowland records are in need of revision. Was reported from the upper alpine belt (up to 2800 m a.s.l.) of the Tyrolean Alps (Austria). Remane & Fröhlich (1994b), HN; see also Günthart (1984), Leising (1977), Woodroffe (1971)
Adarrus multinotatus (Boheman, 1847) M V – M X, occasionally until XII; egg, 2 gen. On Brachypodium pinnatum in dry to temporarily moist, sunny to moderately shady sites, usually on basic substrates, mainly in various types of rather dry grassland as well as in open forests of pine and oak, occasionally in rather moist straw meadows. Absent or scarce in most parts of the north German plain, but widespread to the south of it, particularly common and abundant in limestone regions. Frequently found above 950 m a.s.l. on the Swabian Jura, up to 1500 m a.s.l. in the Bavarian Alps.
Deltocephalinae Fieber, 1869
217
Achtziger (1991), Fischer (1972), Post-Plangg & Hoffmann (1982), Reimer (1992), Remane (1987), Rombach (1999b), Schiemenz (1969), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1939a), Nickel (1994), HN, and others; see also Cook (1996)
Adarrus bellevoyei (Puton, 1877) So far E VII – B X; egg, 1(?) gen. On Brachypodium pinnatum, usually in temporarily dry to damp, basic sites. In Germany it is found only along the upper course of the Isar and in the Eifel Mountains: near Vorderriß, Wallgau and Krün, between 800 and 900 m a.s.l., in open, cattle-grazed pine and spruce forests on alluvial gravel; near Prüm and Bad Münstereifel, 400 to 500 m a.s.l., in calcareous grassland. Is reported to live in forest clearings between 1300 and 1500 m a.s.l. in the Valais (Switzerland), between 1300 and 1800 m a.s.l. in the French Alps. Bornholdt & Remane (1993), Remane (1961b), Rombach (1999b), HN; see also Cerutti (1938), della Giustina (1989)
Errastunus leucophaeus (Kirschbaum, 1868) Deltocephalus antennalis Haupt, 1924 (see Nickel & Remane 2002)
In the whole range so far found B VI – B IX; egg, 2(?) gen. According to present data this species is a rare endemic of the northern Alps and their foothills, living on almost bare gravel banks of unchanneled rivers. Outside Germany it is only known from the Swiss and Austrian Rhine and the North Tyrolean Lech valley: Rhine near Ragaz (type locality), Rhine delta near Bregenz, 400 m a.s.l., 03.VI.1917, and Lech near Reutte, 800 m a.s.l., VIII 1973. From Germany there are only three records: Iller floodplains near Oberstdorf, 11.VIII.1934 and 20.VIII.1935; Isar floodplains near Ascholding, 10.VII.1959, and near Geretsried, 10.IX.1994, between 600 and 800 m a.s.l. Probably lives on an unidentified grass. The Iller and Rhine populations may be extinct due to channeling. Haupt (1924), Remane (1961b), Remane & Fröhlich (1994b), Wagner (1939a), Nickel (1999a); see also Kirschbaum (1868); for identification see Wagner (1939a)
Errastunus ocellaris (Fallén, 1806) Subalpine populations tend to be more strongly pigmented and have been described as E. ocellaris tatraensis (Heller, 1975) from the Tatra Mountains. However, there is apparently clinal variation towards the nominate form, since specimens from the Bavarian Alps show characters of both taxa. The late Heller (pers. comm.) himself had serious doubts on the distinctiveness.
M V – E X; egg, 2 gen. Eurytopic in sunny to slightly shaded stands of grasses in moderately dry to wet, mesotrophic to eutrophic sites. Usually among the dominant species in intensively managed meadows and pastures of lower altitudes, but also in abandoned fields, ruderal habitats, forest clearings, along waysides, in straw meadows, fens and even brackish sites. Host plants are various, usually taller-growing grasses (species of Holcus and Calamagrostis, Elymus repens, Dactylis glomerata and others). Widespread and very common in Germany, frequently extending up to 1000 m a.s.l.; small populations have also been found in the upper montane and subalpine belt of the Berchtesgaden and Allgäu Alps between 1500 and 2000 m a.s.l.
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Review of species
Achtziger (1991), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991, 1997), Nikusch (1976), Reimer (1992), Remane (1987), Remane & Fröhlich (1994b), Schönitzer & Oesterling (1998b), Wagner (1935, 1939a, 1951a), Nickel & Achtziger (1999), HN, and others
Turrutus socialis (Flor, 1861) M V – E X; egg, 2 gen. (see also Witsack 1985). Usually in sunny and oligotrophic, peaty to moderately dry, basic to acidic sites on various substrates, preferentially low-productivity meadows and pastures, locally ruderal sites, waysides, inland dunes, etc., often abundant in calcareous grassland and submontane meadows. Lives on grasses (Festuca rubra, Bromus erectus, probably Nardus stricta and others; also reared from Helictotrichon pubescens and Brachypodium pinnatum taken from British calcareous grassland). Apparently absent from the northwest German plain, but widespread and common in most other parts of Germany, found up to at least 900 m a.s.l. in the Mittelgebirge and the Bavarian Alps, up to 1500 m a.s.l. in the Engadine (Switzerland). Achtziger (1991), Fischer (1972), Kuntze (1937), Nikusch (1976), Reimer (1992), Remane (1987), Rombach (1999b), Schiemenz (1969), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Wagner (1939a, 1951a), Nickel & Achtziger (1999), HN, and others; see also Cook (1996), Günthart (1987a)
Jassargus pseudocellaris (Flor, 1861) Jassus distinguendus Flor, 1861 Flor (1861) created the name J. distinguendus (Fl.) as a replacement name in a chapter of addenda in the same book, based on the erroneous opinion that the name Jassus distinguendus was preoccupied by Athysanus (= Euscelis) distinguendus Kirschbaum, 1858.
E V – E X; egg, 2 gen. In low-growing grass stands in moist to moderately dry, usually oligotrophic, more or less acidic sites, mainly low-productivity meadows and pastures, heaths and abandoned fields, occasionally also ruderal sites, forest margins and clearings. Host plants are grasses (Agrostis capillaris and Festuca rubra, perhaps also Nardus stricta and Holcus spp.). Widespread in Germany, often among the dominant species of low-productivity grassland of submontane altitudes and in the north German plain, particularly on sand, sandstone, granite and other siliceous substrates, but largely absent from limestone regions. Frequently found up to at least 1400 m a.s.l. in the Alps and in higher parts of the Mittelgebirge, single specimens have been recorded up to 1800 m a.s.l. Achtziger (1991), Fischer (1972), Niedringhaus (1991, 1997), Nikusch (1976), Post-Plangg & Hoffmann (1982), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others
Jassargus obtusivalvis (Kirschbaum, 1868) B V – E X; egg, 2 gen. (see also Witsack 1985). In sunny, moderately dry to dry, basic to acidic sites on various substrates, mainly dry grassland, abandoned vineyards, inland dunes, low-productivity meadows, waysides,
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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), Schiemenz et al. (1996), Schulz (1976), Schwoerbel (1957), Wagner (1939a, 1951a), HN
Jassargus repletus (Fieber, 1869) So far M VII – M IX, in the Austrian Alps from M V onwards (Wagner & Franz 1961); egg, 1(?) gen. On sun-exposed calcareous hillsides and riverine gravel banks, usually with open stands of pine and other trees, probably monophagously on an unidentified grass species. In Germany it is only known from a few localities along the edge of the Alps (Illasberg near Füssen, c. 800 m a.s.l., and upper course of the Isar between Wolfratshausen and Krün, 600 to 1000 m a.s.l.), and the upper Rhine plain (inland dunes near Speyer, c. 100 m a.s.l.). At least small numbers have been found at 1300 m a.s.l. in southern parts of Switzerland. Fischer (1972), Remane & Fröhlich (1994b), Schulz (1976), HN; see also Günthart (1987a, 1997)
Jassargus flori (Fieber, 1869) B VI – B X; egg, 2 gen., perhaps 1 at higher altitudes. Usually in moderately shady, moderately moist to moderately dry sites, preferentially open deciduous and coniferous forests, and clearings of lower altitudes (below c. 700 m a.s.l.), also in sunny and open pastures in the submontane and montane belt of the Alps, found up to at least 1500 m a.s.l. in Upper Bavaria. Host plants are narrow-leaved grasses (perhaps Poa pratensis angustifolia, Agrostis capillaris, Deschampsia flexuosa or Festuca heterophylla). Widespread in most parts of Germany, but uncommon and in low densities only. Fischer (1972), Reimer (1992), Remane (1987), Schiemenz et al. (1996), Schulz (1976), Schwoerbel (1957), Wagner (1935, 1939a), Trümbach (1959), HN
Jassargus alpinus (Then, 1896) M VII – E IX, according to Schiemenz et al. (1996) from E V onwards; egg, 1 gen. In damp to moist, moderately shady to sunny sites, mainly at higher altitudes of the Mittelgebirge and the Alps; usually in open forests and clearings of the submontane to the upper montane belt, higher up also in pastures and meadows. Host plants are various grasses (locally Calamagrostis villosa, probably also C. arundinacea, Deschampsia flexuosa and others). In Germany confined to the Alps and higher parts of the Mittelgebirge, and with distinct subspecies: ssp. alemannicus W.Wg. has been found in the Allgäu Alps
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and the Black Forest, ssp. alpinus Then lives in the Bavarian Alps, Rhön Mountains, Thuringian Forest, Solling and Harz Mountains, between 450 and at least 1900 m a.s.l.; small numbers have been collected along the edge of the Harz Mountains between 150 and 300 m a.s.l., but breeding in these sites may be uncertain. Fischer (1972), Reimer (1992), Schulz (1976), Wagner (1958), Schiemenz et al. (1996), HN
Jassargus allobrogicus (Ribaut, 1936) Deltocephalus bavaricus Ribaut, 1936
M VI – M X, after Schiemenz et al. (1996) from M V onwards; egg, 1 gen., perhaps 2 at low altitudes. Preferentially in open deciduous and coniferous forests on siliceous substrates, and often abundant in stands of Deschampsia flexuosa under beech, spruce and pine; locally and in lower numbers also on basic substrates where probably feeding on Festuca ovina, F. heterophylla and F. rubra. Widespread in Germany and common, also in higher parts of the Mittelgebirge and in the upper montane belt of the Alps, where found up to at least 1800 m a.s.l. Reported from 2200 m a.s.l. in Switzerland. Achtziger (1991), Fischer (1972), Reimer (1992), Schulz (1976), Schiemenz et al. (1996), Wagner (1935, 1939a), HN; see also Günthart (1987a)
Jassargus sursumflexus (Then, 1902) M VI – E X; egg, 1 - 2 gen. Monophagous on Molinia caerulea s.l. in peaty to temporarily moist, sunny to moderately shady sites, mainly in intermediate bogs, straw meadows and open forests. Widespread in Germany, but scattered, with strongholds in the Alps and their foothills, the Mittelgebirge, and remnant bog areas of the northern plains, where locally common and abundant in favourable sites, but absent or rare elsewhere. Recorded up to 1350 m a.s.l. in the Allgäu, at 2000 m a.s.l. in the French Alps. Fischer (1972), Kuntze (1937), Reimer (1992), Remane (1958), Wagner (1935), Schiemenz (1971a, 1975, 1976), Schiemenz et al. (1996), Schulz (1976), Schwoerbel (1957), Nickel & Achtziger (1999), HN
Mendrausus pauxillus (Fieber, 1869) B VII – M X; egg, 1 gen. This species is widespread in the steppe zone between Mongolia and eastern parts of central Europe, with rather isolated populations in eastern Germany. Lives on sunny and sparsely vegetated xerothermic sites with relic steppe vegetation (mainly on gypsum, locally also on limestone), on greyish-green subspecies of Festuca ovina. In Germany it is confined to the rain shadow area to the east of the Harz Mountains, currently known only from altogether 6 localities in the vicinity of Bad Frankenhausen, Artern and Halle, between 100 and 300 m a.s.l. Schiemenz (1969), Schiemenz et al. (1996), HN
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Pinumius areatus (Stål, 1858) Hamilton (1983a) proposed – without presenting further arguments – that European populations belonged to Pinumius nebulicola Emelyanov, 1972, described from high alpine steppes of the Pamir mountains. From a biogeographical point of view, this would seem extremely unlikely. Moreover, specimens recently collected in the Lausitz as well as populations from the European part of Russia and Ukraine clearly belong to P. areatus (Stål) (Emelyanov pers. comm.; Walter pers. comm.).
B VI – M X; egg, 2 gen. This is another eastern steppe species with isolated populations in central Europe. Lives on sunny and sparsely vegetated dry grassland, usually on aeolian or moraine sand on an unidentified grass species, perhaps Festuca ovina or Koeleria spp. In Germany there is only a single recent site in a large coal mining area in the Lausitz (Innenkippe Nochten, 09.VI. 1995, 3 ††, and 27.VI.1995, 1 ‡). Altogether 5 further populations found in the 1930s and 1960s, some of which showed extremely high abundances, are presumably extinct (Mainz, Neustrelitz, as well as three sites on the Oder slopes near Eberswalde and Seelow). All localities are below 150 m a.s.l. Kuntze (1937), Schiemenz (1969, and unpublished data), Wagner (1939a), Walter (pers. comm.), Remane (1987)
Diplocolenus bohemani (Zetterstedt, 1838) Mainly M V – M VII, at higher altitudes VII – X; egg, 1 gen. In tall, but usually scattered stands of grasses in temporarily moist to dry, sunny to moderately shady sites of almost all altitudinal belts. In the lowlands and the Mittelgebirge the species is scattered and largely confined to calcareous and gypsum hillsides as well as forest clearings of various substrates; in the Alps and their foothills it is more eurytopic and widespread in various types of mesotrophic grassland, particularly in montane pastures, and in open forests. Preferentially feeds on species of small-reed (Calamagrostis varia, C. villosa, C. epigejos), but also on other grasses; the main host plant in calcareous grassland of lower altitudes is probably Bromus erectus. Locally common and abundant in the Bavarian and Allgäu Alps (up to at least 1900 m a.s.l., but found up to 2250 m a.s.l. in North Tyrol, Austria), in most limestone regions of southern and eastern Germany (notably Swabian and Franconian Jura, Triassic limestone regions of Baden, Franconia, Thuringia), as well as in the montane belt of most Mittelgebirge). Despite its wide distribution in Scandinavia there are no records from the north German plain; the northernmost German localities are on a line from Siegen to Kassel, Hildesheim, Halberstadt and Cottbus. Fischer (1972), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1951a), Nickel (1994), HN; see also Günthart (1987a), Leising (1977), Ossiannilsson (1983)
Diplocolenus penthopitta (Walker, 1851) Some authors treat D. sudeticus (Kolenati, 1860) which shows slight differences in genital morphology, as a distinct species and consider the identity of Walker’s type material as dubious (Dlabola 1980; Nast 1977, 1987). On the other hand Fieber (1869) has redescribed D. penthopitta (Walk.), his material was revised by Wagner (1939a, 1948a) and declared conspecific with D. sudeticus (Kol.). This opinion is followed by Knight (1974), della Giustina (1989) and Holzinger
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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 Vorarlberg, 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 subspecific 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 pastures among Deschampsia cespitosa and Nardus stricta (but probably feeding on another, rather inconspicuous grass). In the Czech Sudetes it prefers avalanche tracks of the upper montane forest belt. Nickel & Voith (unpublished data), HN; see also della Giustina (1989), Holzinger (1999a), Lauterer (pers. comm.)
Verdanus abdominalis (Fabricius, 1803) In the lowlands mainly M V – VIII, at higher altitudes VII – B X; egg, 1 gen. In sunny, damp to moist, occasionally also wet or moderately dry sites (low-input meadows and pastures, fields, clearings, etc.), on Holcus lanatus and other grasses (notably Festuca rubra, Agrostis capillaris, Deschampsia flexuosa). Widespread in Germany, particularly common in the northern plains and the submontane and montane belt of the Mittelgebirge and the Alps, but absent or uncommon in warmer regions of southern parts (e.g. upper Rhine plain, Main valley). Recorded up to 2050 m a.s.l. in the Bavarian Alps and the Allgäu, up to 2600 m a.s.l. in North Tyrol (Austria) and Switzerland. Achtziger (1991), Fischer (1972), Nikusch (1976), Reimer (1992), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Nickel & Achtziger (1999), HN, and others; see also Dobler (1985), Günthart (1987a), Leising (1977)
Verdanus bensoni (China, 1933) Deltocephalus obenbergeri Dlabola, 1945
So far E VI – B VIII, egg, 1 gen. This species’ range is disjunct and restricted to European mountains. Until recently it was only known from England and Scotland, the Swiss and French Jura, the Bohemian Forest (Czech Republic), the Giant Mountains (Poland) and the Ural Mountains (Russia). During recent years large populations were also recorded in the Bavarian Forest (Falkenstein near Zwiesel, 1130-1150 m a.s.l., in pastures, perhaps living on Nardus stricta and Deschampsia flexuosa), as well as in alpine grassland of the Allgäu and Upper Bavaria (Hochplatte above Füssen, 1850 m a.s.l., and Friederspitz above Garmisch-Partenkirchen, c. 1900 m a.s.l.). Biedermann (1998), Nickel & Voith (unpublished data); see also Emelyanov (2000), Lauterer (pers. comm.), Nast (1976a, 1987), Remane & Fröhlich (1994b), Waloff (1979, 1980)
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Arthaldeus arenarius Remane, 1960 Until the generic revision of Remane (1960) was published, this species was commonly misidentified as A. striifrons (Kbm.). Thus, a considerable part of these records from central Europe – even after Remane’s paper – refer to A. arenarius Rem., which is much more widespread.
E VI – B IX; egg, 1 gen. In sunny to moderately shady, moist to moderately dry sites, mainly in forest clearings, mining areas and various other ruderal habitats, occasionally also in open coppice-withstandards and along forest roads, monophagous on Calamagrostis epigejos. The distribution is only poorly known due to former identification problems (see above). However, the species is widespread in Germany and common in many places, at least to the east and south of a line from Düsseldorf to Hanover and Kiel (but not recorded from Mecklenburg-Vorpommern), up to at least 800 m a.s.l. Otherwise known only from southeastern Poland, the Czech Republic, northern France, Austria, southern Russia and Kazakhstan, although probably more widespread in Europe. The locus typicus is Gauting near Munich. Frommer (pers. comm.), Kolbe & Bruns (pers. comm.), Reimer (1992), Remane (1960), Remane & Fröhlich (1994b), Schiemenz et al. (1996), SMNS, Nickel (1994), HN; see also Emelyanov (1964a), della Giustina & Remane (2001), Lauterer (pers. comm.), Mityaev (1975), Nast (1976a)
Arthaldeus striifrons (Kirschbaum, 1868) B VI – E X; egg, 2 gen. Locally fairly common along the coasts of the North and Baltic Sea, where usually found in sunny, moderately saline and temporarily wet to damp sites, mainly meadows and pastures, but also abandoned fields and ruderal sites. Inland, however, the species is rather localized. Many older records (mainly those from forest and bog habitats, as well as from higher altitudes) refer to the preceding species and are in need of revision. The only verified inland localities are in saltmarshes of Thuringia and Saxony-Anhalt, the lower course of the Oder and Isar, the Lake Constance basin, the upper Rhine plain and the Nahe valley, up to 400 m a.s.l. In these sites, the species lives on moderately saline or compacted substrates on Festuca arundinacea, probably also on F. pratensis or Lolium perenne. The locus typicus is near the salt-works of Bad Dürkheim. Fröhlich (1996a), Niedringhaus (1991), Niedringhaus & Olthoff (1993), Remane (1960), ZIMH, Nickel & Achtziger (1999), Nickel (1999b), HN
Arthaldeus pascuellus (Fallén, 1826) M V – E X; egg, 2 gen. (see also Witsack 1985). In peaty to damp, usually sunny sites, mainly meadows and pastures, also fens, abandoned fields, ruderal sites, clearings, waysides, moderately saline marshes inland and near the coast. Host plants are various grasses (Festuca, Lolium, Poa, Calamagrostis and others ). Widespread and common all over Germany, often dominant in fertilized grassland, frequently found in higher parts of the Mittelgebirge, up to 1500 m a.s.l. in the Bavarian Alps. Achtziger (1991), Emmrich (1966), Fischer (1972), Fröhlich (1996a), Niedringhaus (1991), Reimer (1992), Remane (1958), Schaefer (1981), Schwoerbel (1957), Wagner (1935, 1939a), Nickel & Achtziger (1999), HN, and others
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Sorhoanus assimilis (Fallén, 1806) E VI – M X; egg, 1 gen. In acidic to basic, usually peaty and permanently wet, sunny to moderately shady sites, mainly intermediate and spring mires, calcareous sedge fens and straw meadows. Host plants are sedges (perhaps Carex panicea, C. nigra or C. rostrata). Widespread in Germany, but scattered, locally dominant in favourable sites, with strongholds in the Mittelgebirge, the Alps and their foothills; the highest localities are at 1000 m a.s.l. Fischer (1972), Reimer (1992), Schiemenz et al. (1996), Trümbach (1959), Wagner (1935, 1939a, 1941b, 1951a), Nickel & Achtziger (1999), HN
Sorhoanus schmidti (W. Wagner, 1939) B VII – E IX; egg, 1 gen. In peaty to temporarily wet, basic to acidic sites, mainly straw meadows, calcareous sedge fens and intermediate bogs, occasionally syntopically with S. assimilis (Fall.) or S. xanthoneurus (Fieb.). Appears to be associated with open stands of Molinia caerulea, which may be the host plant. This is an endemic species of the Alps and adjacent areas only known from Austria, Switzerland and southern Germany. In Germany it is largely confined to the Alps and their foothills (mainly Upper Bavaria and the Allgäu), where it occurs locally between 600 and 1200 m a.s.l.; it is also found in the southern Black Forest and the Main valley near Schweinfurt, the latter site is situated at only 210 m a.s.l. The locus typicus is Oberstdorf im Allgäu. Otherwise the species is only reported from altogether 8 localities in the northern and central Alps of Styria (Austria) and the Engadine (Switzerland), where it is found up to 1680 m a.s.l., but it is probably more widespread in the Alps. Remane & Fröhlich (1994b), Fischer (1972), HN; see also Günthart (1987a), Wagner & Franz (1961); for identification see Wagner (1939a)
Sorhoanus xanthoneurus (Fieber, 1869) E VII – M X; egg, 1 gen. On Eriophorum vaginatum in raised and intermediate bogs, usually (but not exclusively) in treeless and sunny sites. Widespread in Germany, but very scattered, although often dominant in remnant bogs of the north German plain, the Mittelgebirge (Harz Mountains, Solling, Thuringian Forest, Vogtland, Erzgebirge, Black Forest), and along the edge of the Bavarian and Allgäu Alps, up to 1350 m a.s.l. Has been found on Trichophorum cespitosum in the Czech Republic, but this is probably not the host plant. Niedringhaus & Olthoff (1993), Schiemenz (1971a, 1975, 1976), Schiemenz et al. (1996), Wagner (1941b), Walter (pers. comm.), Nickel (1999b, 2002), HN; see also Novotný (1995)
Lebradea calamagrostidis Remane, 1959 Is considered as the only leafhopper species apparently endemic to Germany, distinguished from the Holarctic Lebradea flavovirens (Gillette & Baker, 1895) only by the length of the aedeagus appendages. However, in German populations of L. calamagrostidis Rem. there is some variabil-
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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, Meggerdorf, Nettelsee, Plön and Lebrade (locus typicus), on Calamagrostis canescens in fens and intermediate bogs. Remane (1959), Remane & Fröhlich (1994b), Niedringhaus & Olthoff (1993), Schröder & K. Heller (pers. comm.); for identification see Remane (1959); see also Gillerfors (2002, and pers. comm.), Ossiannilsson (1983), Vilbaste (1980)
Cosmotettix caudatus (Flor, 1861) B VI – B IX; egg, 1 gen. On Carex hirta in temporarily moist to wet, usually moderately eutrophic, often disturbed sites on various substrates (low-input pastures and meadows, along ditches and waysides, ruderal habitats). In Germany apparently near the western edge of the range, in low to intermediate frequency on the host. Mainly found to the northeast of a line from the island of Wangerooge to Cologne, Göttingen, Marburg, Aalen, Forchheim and Dippoldiswalde (Erzgebirge, c. 700 m a.s.l.); isolated localities are known from England, western France, Switzerland and the Czech Republic, but there are no records from the Netherlands, Belgium, Austria and Poland. The species is probably much under-recorded due to its secretive and specific life habits. Frommer (1996), Hempel et al. (1971), Niedringhaus & Olthoff (1993), Remane & Fröhlich (1994b), Schiemenz et al. (1996), SMNS, Wagner (1937f), Nickel & Achtziger (1999), HN; see also della Giustína & Remane (2001), Kirby (1992), Nast (1987)
Cosmotettix panzeri (Flor, 1861) B VII – E X; egg, 1 gen. This is a tyrphophilous species which is restricted to northern and central parts of Europe. It lives in intermediate and raised bogs on cottongrass (Eriophorum angustifolium, perhaps also E. vaginatum). In Germany only sporadic, with strongholds in the remnant bog areas of the north German plain; also found in a few sites in the Mittelgebirge and near the edge of the Bavarian Alps up to 900 m a.s.l. Southernmost records are from Wolfratshausen (to the south of Munich) and Titisee-Neustadt (southern Black Forest). Border localities are known from the French Alps, the Czech Republic and Slovakia. Not reported from Austria and Switzerland. Reimer (1992), Remane (1958, and pers. comm.), Schiemenz et al. (1996), Wagner (1935), Nickel & Remane (1996), Nickel (1997), HN; see also della Giustina (1989), Jansky & Okali (1993), Lauterer (1984)
Cosmotettix costalis (Fallén, 1826) E VI – B X, occasionally B VI; egg, 1 gen.
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In wet to temporarily flooded, usually mesotrophic to moderately eutrophic, treeless sites, mainly in fens, straw meadows and depressions of river floodplains. Lives on sedges (Carex acuta, probably also C. elata and C. nigra). In Germany at the southwestern edge of the range, rather scattered and in low individual numbers, although perhaps under-recorded due to secretive life habits. Border localities are on the island of Borkum, near Oldenburg, Göttingen, Gießen, Schweinfurt, Lenzkirch (Black Forest), Immenstadt (Allgäu) and GarmischPartenkirchen, up to 750 m a.s.l. There are only very few isolated records to the west and south of this line, notably from Norfolk (England), near Apeldoorn (Netherlands), Zurich (Switzerland) and Klagenfurt (Austria). Not reported from France. Fröhlich (1996a), Niedringhaus (1991), Nikusch (1976), Remane (1962), Remane & Fröhlich (1994b), Nickel & Achtziger (1999), Nickel & Remane (1996), Nickel (1997), HN; see also Bieman & Rozeboom (1993), Günthart (1987b, and pers. comm.), Holzinger (1995c), Kirby (1992)
Cosmotettix aurantiacus (Forel, 1859) So far M VI – E VIII; egg, 1 gen. In wet, usually peaty sites, mainly straw meadows and fens of the foothills of the Alps, presumably on an unidentified species of Carex. Known in Germany from only 10 localities, in Upper and Lower Bavaria (locally extending northward to the Danube valley), the Allgäu and the southern upper Rhine plain (Burkheim near Freiburg), between 150 and 800 m a.s.l. Otherwise the species is reported from Lithuania, Poland, Switzerland, Austria, eastern France, European and eastern Siberian parts of Russia, as well as Mongolia. Remane (1961d), Remane & Fröhlich (1994b), HN; see also Anufriev & Emelyanov (1988), della Giustina & Remane (2001), Nast (1972, 1987)
Calamotettix taeniatus (Horváth, 1911) So far B VII – B IX; egg, 1 gen. This is another monophagous species on Phragmites australis, preferentially occurring in temporarily flooded freshwater sites as well as inland salt marshes. In Germany at the edge of the range and was recorded only from 9 sites in the southern half: upper Rhine valley (Speyer and Landau), Bodanrück (Konstanz), near Stuttgart (Horrheim), Wetterau (Rockenberg and Selters), Grabfeld (Bad Königshofen), Oberlausitz (Königshain and Nochten) and Geiseltal near Halle, between 100 and 450 m a.s.l. It is probably underrecorded as it is rather wary and thus, difficult to sweep. Fröhlich (1996a), Heller (1987b), Funke (pers. comm.), Remane (1995), Remane & Fröhlich (1994b), Nickel (1999b), HN
Praganus hofferi (Dlabola, 1947) So far B VII – M IX, in Bohemia (Czech Republic) according to Dlabola (1947) also E V; egg, 2 gen. This species is widespread in the Asian steppe zone, with relic populations in central Europe. In Germany it is only known from a single site in the Thuringian Basin:
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Schwellenburg near Kühnhausen, 210 m a.s.l., 15.IX.1963, 36 specimens, in a dry, southsloping pasture on Triassic gypsum with dominating Stipa capillata and Festuca ovina. From this site there are also two recent records: 13.VII.1997, 1 †, 6 ‡‡, and 03.VII.1998, 4 ‡‡. The species is probably associated with Stipa. Schiemenz (1973), HN; also Emelyanov (pers. comm.); for identification see Dlabola (1947, 1954), Mityaev (1971)
Enantiocephalus cornutus (Herrich-Schäffer, 1838) E VI – E IX; egg, 1 gen. In sunny and scarcely vegetated, usually disturbed stands of grasses, mainly in moderately dry to dry (occasionally also temporarily moist and moderately saline), sandy to loamy sites, e.g. waysides, fallows, inland salt marshes and disturbed patches of dry grassland. The main host is Elymus repens, but the species probably lives also on Agrostis capillaris and Festuca rubra. In Germany only locally in lowland areas below 400 m a.s.l., near the northern limit of the range, recorded from the northern upper Rhine plain, the valleys of the Nahe and Tauber, Franconia (locally common in the valleys of Main, Saale and Regnitz, Grabfeld), the rain shadow area to the east of the Harz Mountains, the Thuringian Basin and southeastern Niedersachsen (vicinity of Helmstedt and Wolfenbüttel). Fröhlich (1996a), Reimer (1992), Remane & Fröhlich (1994b), Schiemenz et al. (1996), SMNS, HN
Mocuellus collinus (Boheman, 1850) M V – M X; egg, 2 gen. Often syntopic with the preceding species, but more common and also occurring in the northern plains, with more pronounced preference of sandy soils. In sunny, moderately dry to dry sites with incomplete vegetation cover, mainly abandoned fields, waysides and pastures. Lives on various grasses (Elymus repens, low-growing species of Festuca, Poa compressa and others). In Germany widespread and locally common, particularly in the lowlands of southern and eastern parts, but absent from most parts of the Mittelgebirge as well as to south of the Danube. Highest localities are at 450 m a.s.l. only. Fischer (1972), Post-Plangg & Hoffmann (1982), Remane (1987), Schiemenz (1969), Schiemenz et al. (1996), Schönitzer & Oesterling (1998b), Schwoerbel (1957), Wagner (1935, 1939a, 1951a), HN, and others
Erzaleus metrius (Flor, 1861) B VI – E X; egg, 2 gen., probably 1 in cooler sites. On Phalaris arundinacea in moderately wet to temporarily flooded, often eutrophic sites, mainly along ditches and shores of rivers and lakes, and in abandoned grassland, occasionally on meadows cut at most once a year. Widespread in Germany, usually in intermediate to high frequency and high abundance on the host, found up to at least 1000 m a.s.l. in the Mittelgebirge and the Bavarian Alps. Fischer (1972), Haupt (1925), Mölleken & Topp (1997), Niedringhaus (1991), Reimer (1992), Remane (1958), Schiemenz et al. (1996), Trümbach (1959), Wagner (1935, 1939a), Witsack (1985), HN; see also Lauterer (1986)
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4.3 Dubious records Ribautodelphax pallens (Stål, 1854) Most records of this species are from northern Europe, the Alps (between 1450 and 2250 m a.s.l.) and the Pyrenees; furthermore, it is reported from Poland, the former Yugoslavia and Bulgaria (Nast 1987; Ossiannilsson 1978; Remane & Fröhlich 1994b). From Germany there is only a single record from Thuringia: Sulza near Jena, 26.V.1977, 1 †, 1 ‡ (Coll. Sander, Bieman det.). This is the only central European lowland site. Thus, there may be a possibility of exchanged labels. The species lives monophagously on Festuca ovina (Bieman 1987b). Tettigometra concolor Fieber, 1865 Listed by Nast (1972, but not 1987) for West and East Germany. Varieties bearing the name concolor Fieb. have been described for several taxa of the genus Tettigometra. Identity, taxonomic status, as well as occurrence in Germany must be regarded as dubious. Micrometrina longicornis Signoret, 1866 Listed by Nast (1972, 1987) for West Germany, presumably referring to Melichar (1896), Hüeber (1904) and Haupt (1935), but no specimens have been found in any of the recently revised collections. Therefore, these statements most likely refer to localities outside present-day Germany. Paradorydium paradoxum (Herrich-Schäffer, 1837) This taxon is widespread from central Asia to the Mediterranean region, but perhaps described from Franconia („von Dr. Hahn angeblich im Herbste bei Nürnberg gefunden“, see Herrich-Schäffer 1837), but never recorded again in Germany since. Its occurrence in central Europe may be only erratic. The food plants are grasses in steppe-like habitats (Emelyanov 1964a; Mitjaev 1971; see also D’Urso 1992). Chlorita viridula (Fallén, 1806) Has been listed for Germany in older papers, but was usually confused with Ch. paolii (Oss.). A more recent record was published by Schiemenz (1990): Strausberg near Frankfurt an der Oder, 06.IX.1979, 1 †. However, the specimen is apparently lost. Occasionally, specimens of Ch. paolii (Oss.) showing abnormal genital morphology have been found. Hence, this record is regarded here as dubious. From England Ch. viridula (Fall.) is reported to live in coastal saltmarshes on Artemisia maritima (Le Quesne & Payne 1981; Kirby 1992).
Dubious records
229
Edwardsiana kemneri (Ossiannilsson, 1942) The identity of this taxon is not quite clear. It has been described after a single † from central Sweden. After that, further records from Czechia, Slovakia, Romania, Cyprus and Mongolia have been published (Dlabola 1965, 1967b, 1977). It is, furthermore, reported from the Spitzberg near Tübingen without any further comments (Schwoerbel 1957). However, this record is not found in Wagner’s unpublished files, who has seen Schwoerbel’s material. Due to the widespread occurrence of genitalic abnormalities caused by parasitation in Edwardsiana specimens, this taxon should be omitted from the German list. Populations of E. tersa (Edw.), in particular, have been found to contain †† of the kemneri aedeagus type. Selenocephalus obsoletus (Germar, 1817) There are perhaps two specimens from Germany, the exact origin of which is dubious: Bavaria, Main valley, Retzbach, 09.VI.1964, 1 † (ZMH, leg. H. Lindberg) and “Württemberg, v. Roser, 1872-76” (SMNS). The nearest confirmed localities are in eastern Austria, Slovakia, South Tyrol (Italy) and southern France. In Mediterranean regions the species has been swept from tall herbs, woody species of Fabaceae, and Cistus. Handianus flavovarius (Herrich-Schäffer, 1835) sensu Emelyanov (1964a) This is another species perhaps described from Regensburg, without any further records from Germany. The original diagnosis does not include a description of the genitalia, the type material is lost, and no type locality is given. The species is not mentioned in the ‘Fauna Ratisbonensis’ (Herrich-Schäffer 1935b, 1840). Hence, origin and identity of the holotype as well as the occurrence in Germany must be considered as dubious. According to Nast (1972, 1987), the species is reported from the Altai westward to eastern parts of central Europe. Nearest localities are in southern Poland and Moravia (Nast 1976a; Lauterer 1983). According to various authors, it lives on grasses on sunny, damp to dry sites; in Latvia and Lithuania adults have been found from mid June until mid August (Emelyanov 1964a; Vilbaste 1974). Handianus procerus (Herrich-Schäffer, 1835) sensu Emelyanov (1964a) Often cited as having been described from Regensburg (e.g. Metcalf 1967; Nast 1987). However, Herrich-Schäffer (1835b, 1840) neither gave any information on the locus typicus, nor listed the species in his ‘Fauna Ratisbonensis’ (Herrich-Schäffer 1840); furthermore, the type material is lost. Thus, the occurrence in Germany is not confirmed. According to Nast (1987), the species is widespread from central Asia to western Europe (see also Ribaut 1952), although the identity may be unclear. Senecio and Echinops are reported as food plants from southern Russia (Emelyanov 1964a); in Moravia it has been found on Genista pilosa in xerothermic grassland (P. Lauterer, pers. comm.).
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4.4 Short-term changes 4.4.1 Declines Due to the lack of accurate data, long-term population declines are difficult to substantiate. 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 paradoxum (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 appearance and frequent sampling in the area. The two latter species, however, have been recorded 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). Further, 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 largescale and long-termed decrease of records, or on large-scale habitat changes. Possible factors are taken mostly from Nickel et al. (1999); the present status is mostly taken from Remane 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 & Oesterling 1998a; ZMB). Tettigometra obliqua (Panz.) was particularly numerous and even reported 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 habitat 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 xanthoneurus (Fieb.). In some cases of substantiated local extinctions, notably in Nothodelphax albocarinata (Stål) and Limotettix atricapillus (Boh.), the habitats do still exist, but have become subject 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 created by local farmers is particularly noteworthy. These species originally prefer the vicinity of hollows, which have disappeared from many bogs after draining. In some localities, the
Declines
231
Table 3. Declining Auchenorrhyncha species in Germany. Present status mainly after Remane et al. (1998), possible factors after Nickel et al. (1999). Species
Region of decline
Present status
Possible factors
Cixius stigmaticus (Germ.) Cixius similis Kbm. Myndus musivus (Germ.) Nothodelphax albocarinata (Stål)
Most parts North Germany, Thuringia Rhine, Main, Weser, Elbe Eastern half of Germany
Endangered Vulnerable Critical Critical*
Xanthodelphax flaveola (Fl.)
Most parts
Endangered
Pseudodelphacodes flaviceps (Fieb.) Javesella salina (Hpt.)
Lech floodplains Eastern Germany
Critical Endangered
Tettigometra macrocephala (Fieb.) Tettigometra atra Hag. Tettigometra fusca Fieb. Tettigometra impressopunctata Duf. Tettigometra leucophaea (Preyssl.) Ommatidiotus dissimilis (Fall.) Tibicina haematodes (Scop.)
Thuringia, Franconia Most parts Thuringia, Bavaria, Baden Thuringia, Bavaria Most parts Northern Germany Baden-Württemberg
Critical Critical Endangered Endangered Probably extinct* Endangered Critical
Macropsis impura (Boh.)
Most parts
Vulnerable
Forestry Draining of peatland Channeling of rivers Peatland eutrophication Grassland improvement Channeling of rivers Grassland improvement Unknown Unknown Unknown Unknown Unknown Draining of peatland Viticulture, habitat destruction Wetland improvement
Macropsidius sahlbergi (Fl.) Anoscopus histrionicus (F.) Coryphaelus gyllenhalii (Fall.) Deltocephalus maculiceps Boh. Doratura horvathi W.Wg. Platymetopius undatus (De G.) Mimallygus lacteinervis (Kbm.) Hardya tenuis (Germ.) Sardius argus (Marsh.) Cicadula quinquenotata (Boh.) Mocydiopsis intermedia Rem. Athysanus quadrum Boh. Limotettix atricapillus (Boh.)
Eastern Germany Eastern half of Germany Thuringia, Bavaria Northern Germany Southern Niedersachsen Most parts Iller floodplains Most parts Western parts Northern Germany Thuringia Most parts of Germany Eastern half of Germany
Critical Endangered Critical Critical Vulnerable Endangered Critical Endangered Critical Endangered Vulnerable Endangered Critical
Laburrus impictifrons (Boh.) Psammotettix unciger Rib. Psammotettix pallidinervis (Dhlb.) Errastunus leucophaeus (Kbm.) Pinumius areatus (Stål) Sorhoanus assimilis (Fall.) Sorhoanus xanthoneurus (Fieb.)
Eastern half of Germany Iller floodplains Most parts Iller floodplains Brandenburg and Mainz Most parts Northern Germany
Endangered* Critical Endangered Critical Critical Vulnerable* Endangered
* = not identical with status suggested by Remane et al. (1998)
Habitat destruction Unknown Habitat destruction Draining of peatland Cessation of grazing Unknown Channeling of rivers Unknown Unknown Draining of peatland Unknown Wetland improvement Peatland eutrophication Habitat destruction Channeling of rivers Unknown Channeling of rivers Unknown Wetland improvement Draining of peatland
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Review of species
only populations of Macrosteles fieberi (Edw.) and Limotettix atricapillus (Boh.) were confined to such turbaries, but were absent from other parts of the bog (Nickel 1997, 1999b). Further, the improvement of fen habitats during recent decades was probably equally severe and affected a large number of species, although evidence is only indirect in most cases, except in Macropsis impura (Boh.), Athysanus quadrum Boh. and Sorhoanus assimilis (Fall.). (iii) Species of river banks The regulation of rivers, altering their natural flooding dynamics, has brought some species to the brink of extinction. This is particularly true for Psammotettix unciger Rib. and Errastunus leucophaeus (Kbm.), which are both endemics of the northern Alps, and Pseudodelphacodes flaviceps (Fieb.) and Mimallygus lacteinervis (Kbm.). Direct evidence of decline is lacking for Pentastiridius beieri (W.Wg.), which has probably been overlooked by earlier collectors. All these species live on almost bare gravel banks with extremely scattered vegetation (see Nickel 1999a). Myndus musivus (Germ.) shows similar habitat requirements. 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 grassland 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 horvathi 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 Pinumius 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 preserved, 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) Nannfeldt 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 hybrid U. x hollandica are particularly susceptible, whereas U. laevis is apparently less concerned. 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 European 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 occurred. On the other hand, many of the remaining elm stands still show a considerable infestation of several leafhopper species, and Kyboasca bipunctata (Osh.) even seems to prefer coppices growing after the chopping of dying trees. (vii) Woodland species Cixius stigmaticus (Germ.) and Platymetopius undatus (De G.) suffered a severe decline at least since 1970. The reasons are unknown. They both ascend to the canopy layer after emergence and may require a more complex habitat structure. 4.4.2 Range expansions Statements on range expansions must be drawn with care due to the shortage of faunistic information. Records of new species, which either have been misidentified formerly or which require special habitats or host plants, are constantly being added to many faunal lists even in central Europe. In most cases, this is not caused by range expansions, but rather by the increase of taxonomic and ecological knowledge. Usually, this phenomenon can be definitely proven by the study of older collections. For instance, Kelisia sima Rib., Florodelphax paryphasma (Fl.), Macropsis najas Nast, Alebra coryli Le Q., A. viridis R., Zygina griseombra Rem., Z. nirgitarsis Rem., Balclutha calamagrostis Oss., Streptanus confinis (Reut.) and Arthaldeus arenarius Rem. have all been found to be common and widespread in most parts of central Europe shortly after publication of the relevant information, and most of them also turned up in museum collections. In other cases, however, there are good arguments and sufficient data for a real range expansion. The following assessment is also based on a careful evaluation of negative records in collections and older publications, notably MTD, SMNS, ZIMH, ZMB, ZSM, Haupt (1935), Kirschbaum (1868), Kuntze (1937) and Wagner (1935, 1939, 1941a, 1951). Mere expansions within central Europe are documented for Eurybregma nigrolineata Scott (but see 4.1.2.4), Haematoloma dorsatum (Ahr.), Viridicerus ustulatus (M. & R.), Liguropia juniperi (Leth.), Ribautiana debilis (Dgl.), Zyginella pulchra P. Löw, Zyginidia scutellaris (H.-S.) and Fieberiella septentrionalis W.Wg. (section 4; see also Niedringhaus & Olthoff 1986; Remane 1995; Remane & Fröhlich 1994b). Some of these species prefer anthropogenic or otherwise disturbed habitats, often with introduced plants. E. nigrolineata Scott and Z. scutellaris (H.S.) mainly live on grasses along waysides and on leys, V. ustulatus (M. & R.), R. debilis (Dgl.), Z. pulchra P. Löw and F. septentrionalis W.Wg. occur in natural habitats in southern parts of Germany, but prefer urban sites along the expansion front line. H. dorsatum (Ahr.) is the only expanding species, which avoids anthropogenic sites and prefers closed forests. The Mediterranean Liguropia juniperi (Leth.) lives on the ornamental Chamaecyparis lawsoniana originating from North America and must have invaded only after the host’s introduction. Range expansions have also been suggested for Macropsis megerlei (Fieb.) and Balcanocerus larvatus (H.-S.), which both live on Rosa species and Prunus spinosa. Their host
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Review of species
plants, however, are very difficult to sample with the sweep net. Moreover, there are old records of both species from southern Norway and Mecklenburg, respectively (Kuntze 1937; Ossiannilsson 1981; Remane & Fröhlich 1994b; Nickel 1997). Edwardsiana sociabilis (Oss.) is now common and widespread in urban habitats on ornamental Rosa rugosa and has even been reported as a pest in Finland. Originally, it was probably confined to Filipendula ulmaria in rather moist habitats (Nuorteva 1955; Remane & Fröhlich 1994b). It is not clear, however, whether its host shift caused a real expansion of the geographic range or merely an increase of abundance. 4.4.3 Introductions There is no doubt that Stictocephala bisonia Kopp & Yonke and Graphocephala fennahi Young have been introduced from North America, and that Macropsis elaeagni Em. has been introduced from central Asia. Another interesting example is Opsius stactogalus Fieb., which was formerly confined to Myricaria germanica along banks of unregulated alpine rivers. At least since the second half of the 19th century, it has spread over most lowland areas, living on ornamental Tamarix in urban habitats. Introduction, at least range expansion, is also likely in some Eupteryx species living on archaeophytic spice and medicinal herbs. E. decemnotata R., which was only known from Mediterranean regions of France and Italy a few years ago, has appeared in Switzerland, 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 species 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 (Ocimum). 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 monophagous on native European plants in more or less natural habitats, and their abundances 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 (Wagner & Franz 1961) and has since been found mostly on the native Acer campestre, although 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. Empoasca, 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 widespread 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 typhlocybine leafhopper Kyboasca maligna (Walsh) was found in Alsace (France), close to the German border in 1997 and in the French Ardennes in 2000 (della Giustina & Remane 2001). In recent years, the eastern Palearctic typhlocybine Vilbasteana oculata (Ldb.) has appeared in Moscow on ornamental Syringa (Tishechkin 1989) and may spread westward. 4.4.4 Occasional influxes The Mediterranean issid species Agalmatium bilobum (Fieb.) was common in dune habitats of the East Friesian island of Wangerooge at least in 1960, but was not found afterwards (Harz 1965, 1988; Niedringhaus 1991). Balclutha saltuella (Kbm.) was described from a single female from the vicinity of Wiesbaden in the second half of the 19th century. Only 130 years later, it happened to be recorded again in Germany; but despite three records from widely separated localities, all specimens were apparently vagrants without evidence for reproduction (Nickel 1999a). This species is also known to occur erratically in other parts of Europe. Likewise, an apparently single specimen of Zyginidia pullula (Boh.) found in eastern Bavaria may have immigrated from more southern or eastern parts of Europe. Toya propinqua (Fieb.) breeds in the upper Rhine plain at least in some years, but single specimens have also been found in southern Bavaria. Single individuals of the Mediterranean Cicada orni L. outside their known range were also recorded, but not sampled (P. Dynort, pers. comm.; Heller 1987c). Back (1976) and Hess (pers. comm.) observed single individuals of Tibicina haematodes (Scop.) on dispersal flight in southern Germany. Some older, unconfirmed records of further cicada species (Duffels & van der Laan 1985) may refer to migrating specimens. Autochthonous occurrence in Germany is also uncertain in Zygina tithide Ferr., Fruticidia sanguinosa (R.) and Circulifer opacipennis (Leth.), which may occur only temporarily as vagrants. In contrast, Achorotile albosignata (Dhlb.), Megadelphax haglundi (J. Shlb.) and Megamelodes lequesnei W.Wg., are probably native, although there are only single specimens. Most of these were brachypterous, and in addition, these species either live among thorny scrub or show a hidden way of life, their host plants being unknown.
4.5 General remarks on the German Auchenorrhyncha fauna At present, altogether 620 Auchenorrhyncha species are known and verified from Germany, although in a few cases, there are uncertainties regarding the correct nomenclature. Considering the fact that more than 40 new records, 4 of which were new original 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. Dubious 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., Zyginidia 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 nearest 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 represented (with cicadas and froghoppers interpreted as families, respectively). The Cicadellidae 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, Tropiduchidae 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 (S = 145)
194 158
150
109 100
50
21
2
1
4
C De ixiid lp ha ae ci d Di A c ae ct hi yo lid ph ae ar id ae Te I s tti s go ida m et e rid ae
0
0000000000000000000000000 00000000000000000000 00 000000000000000000 0000000000 000000000000000000 000000000000000000000000000000000 00 000000000000000000 0 00000000000000000000000 0000000000000000000000000
00000000000000000000000 0000000000000000000000000000 000000000000000000 00000000000000000000000 000000000000000000 00000 000000000000000000 000000000000000000000000000000000 000000000000000000 00000000000000000000000 000000000000000000 00000 000000000000000000 000000000000000000000000000000000 000000000000000000 00000000000000000000000 000000000000000000 00000 000000000000000000 000000000000000000000000000000000 000000000000000000 0000000000000000000000000000 000000000000000000 000000000000000000 000000000000000000000000000000000 000000000000000000 0000000000000000000000000000 000000000000000000 000000000000000000 000000000000000000000000000000000 000000000000000000 0000000000000000000000000000 000000000000000000 000000000000000000 000000000000000000000000000000000 00000 00 00 00 00 00 00 00 00 00 00 000000000000000000 00000000000000000000000 00000000000000 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 00000000000000000000000
8
33
17
000 000 000 000 000 000 000 000 000 000 000 000 000 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
3
25
00000000000000000000000 000000000000000000 0000000000000000000000000000000 00000000000000000000000 00 0000000000000000 00000000000000000000000 00000000000000000000000000 00000000000000000000000000 000 0000000000000000 000 0000000000000000 0000000000000000 0000000000000000 0000000000000000000000000000000 00000000000000000000000000 0000000000000000000000000000000 00 0000000000000000 00000000000000000000000 00 0000000000000000 0000000000000000000000000 00000000000000000000000 0 000000000000000000000 0000000000000000000000000 0000000000000000000000000 000000000000000000000000 000000000000000000000 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 00000000000000000000000 0000000000000000000000000
3
3
2
1
8
00000000000000000000000 00000000000000000000 0 00 00000000000000000 0000000000000000000 00 0 00000000000000000 00 0000000000000000000 00 0000000000000000000 00 00 00000000000000000 000000000000000000 0 00 00000000000000000 00 00000000000000000 00000000000000000000 000 00 00000000000000000 0 000000000000000000 0 00 00000000000000000 00 00000000000000000 00000000000000000000 000 00 00000000000000000 0 000000000000000000 0 00 00000000000000000 00 00000000000000000 00000000000000000000 000 00 00000000000000000 0 000000000000000000 0 00 00000000000000000 00 00000000000000000 00000000000000000000 000 00 00000000000000000 0 000000000000000000 0 00 00000000000000000 00 00000000000000000 00000000000000000000 000 00 00000000000000000 0000000000000000000 00 000 00000000000000000 00000000000000000 00000000000000000000 000 00 00000000000000000 0000000000000000000 00 000 00000000000000000 00000000000000000 00000000000000000000 000 00 00000000000000000 0 00000000000000000 0000000000000000000 00 00000000000000000000000 0 000000000000000000 0 0000 0 0000000000000000000000000000 0000000000000000000000000 00 00000000000000000 00000000000000000000 000 000000000000000000 00000000000000000 0000000000000000000000000 00000 0000000000000000000000000 0 00000000000000000000000 00000000000000000000000 0 000000000000000000 0 0000000000000000000000000 00000000000000000000000 00000000000000000000000 0000000000000000000000000 0000000000000000000000000 000000000000000000 00000000000000000000000
4
16
1
1
6
0000000000000000000000000 0 00000000000000000000000 00000000000000000000 00000 00000000000000000000 000000000000000000000000000000 00000000000000000000 00000 00000000000000000000 000000000000000000000000000000 00000000000000000000 000000000000000000000000000000 00000000000000000000 00000 00000000000000000000 000000000000000000000000000000 00000000000000000000 000000000000000000000000000000 00000000000000000000 00000 00000000000000000000 00000000000000000000000000000000000 00000000000000000000 0000000000000000000000000 00000000000000000000 00000 00000000000000000000 00000000000000000000000000000000000 00000000000000000000 0000000000000000000000000 00000000000000000000 00000 00000000000000000000 00000000000000000000000000000000000 00000000000000000000 0000000000000000000000000 00000000000000000000 00000 00000000000000000000 00000000000000000000000000000000000 00000000000000000000 0000000000000000000000000 00000000000000000000 00000 00000000000000000000 00000000000000000000000000000000000 00000000000000000000 0000000000000000000000000 00000000000000000000 00000 00000000000000000000 00000000000000000000000000000000000 00000000000000000000 000000000000000000000000000000 00000000000000000000 00000000000000000000 00000000000000000000000000000000000 00000000000000000000 000000000000000000000000000000 00000000000000000000 00000000000000000000 00000000000000000000000000000000000 00000000000000000000 000000000000000000000000000000 00000000000000000000 00000000000000000000 00000000000000000000000000000000000 00000000000000000000 0000000000000000000000000 0000000000000000000000000
Ci ca Ce did rc ae M em opid br ae ac i M eg Ul dae op op i ht na ha e lm in Le ae M dr ac in ro ae ps A ina ga e Id lliin io c e ae rin Ia ae Pe ss i Do nth nae ry im c e iin ph ae A alin ph a e r Ci odin ca Ty d ae ph elli De loc nae lto y b ce ina ph e al in ae
Species number
200
Cicadomorpha (S = 475)
Fig. 2: Species numbers and systematic affiliations of the Auchenorrhyncha fauna of Germany
General remarks and problems
237
5 Utilization of plant resources
5.1 Plant taxa 5.1.1 General remarks and problems In view of more than 600 species of Auchenorrhyncha and roughly 3000 plants, resulting in 1.8 million possible associations, the approach of assessing the overall relationships between a rather diverse insect group and a whole flora in a large geographical area such as Germany may seem out of reach. But in fact, only a relatively small proportion of the taxonomic plant diversity is exploited, and the data base on the more relevant groups is often sufficient or at least moderate, though not in all parts of the study area. Nevertheless, there are substantial gaps in the knowledge of plant preferences of many polyphages and of all species spending the nymphal stage in the soil, such as the Cixiidae, Cicadidae and Cercopinae. Furthermore, there is a general scarcity of specific data from plant species confined to southern parts of Germany, notably the alpine and subalpine belt as well as viticultural regions of the Rhine valley and its tributaries. For instance, alpine grasses, sedges and willows, as well as extensive forests of Castanea sativa along the upper and middle Rhine may provide ample resources, which are not yet studied. Further sampling is also needed on a number of more widespread plants, such as Abies alba, Ulmus laevis and even Betula pubescens. In contrast, the lack of Auchenorrhyncha data from rare or localized plants is probably of minor significance, since almost all species live on more widespread and abundant plants. Exceptions from this rule are few, and comprise, for instance, the delphacids Kelisia minima Rib., living on Carex distans, and Nothodelphax albocarinata (Stål), found on Carex limosa. Another difficulty is the evaluation and classification of food plant records of phytophagous insects in general and the resulting problems for analysis, which were discussed by Ward (1988) and, more extensively, by Schoonhoven et al. (1998). The major problems will be discussed in the following sections. (i) Uncertainties regarding the feeding substrate Although the majority of central European Auchenorrhyncha species are rather homogeneous in feeding, there are some exceptions. The Achilidae are considered to suck on fungi mycelium. At least in Empoasca decipiens Paoli feeding is known on phloem as well as on mesophyll. Prestidge (1982), without experimental studies, reported xylemfeeding in Euscelis incisus (Kbm.). However, deltocephaline leafhoppers and Euscelis spp. in particular, which were reared by Müller, Strübing and Witsack for decades, are
238
Utilization of plant resources
widely considered as phloem-feeders. In fact, systematic experimental studies on the type of substrate have been done only in a few species, most of which are of economic importance, although generally, there is no reason to question classifications made by Müller and co-workers, who determined food substrates after the type of excrements: greenish (due to chlorophyll) in mesophyll feeders, watery both in xylem and phloem feeders, but viscous due to the surplus of sugar in the latter (Müller 1956). (ii) Dependence on developmental stage and season Some species utilize different plants for oviposition, larval development, adult feeding, wintering, or even in the first and the second generation. These shifts can be either facultative or obligate. Muellerianella fairmairei (Perr.) develops on Holcus species, where at least some individuals of the first generation lay their eggs. Oviposition of the second generation, however, obligatorily takes place on Juncus effusus (Drosopoulos 1977, and pers. comm.). Many vertical migrants feed on grasses and other herbaceous plants as nymphs, and on various trees and shrubs as adults, e.g. Haematoloma dorsatum (Ahr.), Stictocephala bisonia Kopp & Yonke, Allygus spp., Allygidius spp. and Hesium domino (Reut.). Many Zygina species, being monophagous as nymphs on deciduous woody plants, feed on conifers as wintering adults and on various woody Rosaceae in spring (mainly Crataegus spp., Prunus spinosa and P. padus). Also, Balclutha punctata (F.), Empoasca vitis (Göthe) and probably some species of Idiocerinae overwinter on conifers. B. punctata (F.) develops on Poaceae, the others on deciduous trees and shrubs of various taxa. Claridge & Wilson (1978a) and Vidano et al. (1990) demonstrated food plant shifts between the first and the second generation, which may be due to the plant’s suitability for egg overwintering. In Wales (and probably in central Europe, too), Lindbergina aurovittata (Dgl.) migrates from evergreen Rubus species to Quercus and other deciduous trees, and back. In some species of Eupteryx, the first generation is monophagous, the second, however, is oligophagous or even polyphagous. Thus, nymphs of the first generation of Eupteryx aurata (L.) are apparently confined to Urtica dioica, whereas the emerging adults colonize dicotyledonous plants of numerous families. Perennial populations of Eupteryx filicum (Newm.) only occur on the evergreen fern Polypodium, but adults of the first generation may found a second generation also on hemicrytophytic fern species. In Eupteryx thoulessi Edw., most populations are found on Mentha aquatica, but at least summer individuals (including nymphs) also occur on Lycopus europaeus, where both plants grow together. A possible explanation for this host shift (or more precise: diet width expansion) may be differing suitability of plant species for egg overwintering. This seems rather evident in Eupteryx filicum (Newm.), for which hemicryptophytic ferns can not provide substrate for winter eggs. (iii) Subterranean feeding With few exceptions, e.g. Haematoloma dorsatum (Ahr.) – see above – the host plants of subterranean nymphs of Cixiidae, Cicadidae, and Cercopinae are completely unknown. There is evidence, that at least some species feed on woody plants. But if this is true, more tight host relations are possible or even likely, simply because of the limited ability of small insects to move through the soil for longer distances and into another tree’s rhizosphere. So far, most of these species are tentatively considered as polyphagous. (iv) ‘Vagrants’, ‘tourists’, ‘migrants’ Migrating and drifted individuals, as well as those fallen down from the canopy above have caused many erroneous food plant records in the literature. For a long time,
General remarks and problems
239
such records concealed the monophagy of a number of species, for instance in most idiocerine leafhoppers. (v) Geographic differences Many examples of stenophagous insects feeding on different plant species in different 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 species occurs. For instance, Eurhadina kirschbaumi W.Wg. is strictly monophagous on Quercus 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 oligophagous. 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 caerulea, 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 require 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 phenomenon 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 introduced 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 Rhododendron 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 maritima 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 geographical 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 Ribautodelphax (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 feeding 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.). However, 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 species, respectively, on the substantiated host. Nevertheless, these single occasions may be more widespread and deserve further attention. In combination with results of laboratory 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 between 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 insects 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 entomologists. 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, specificity 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 phylogenetic 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 European Auchenorrhyncha, both terms can be used as synonyms, except for those migrating 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-mining 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 comprising 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. Extensive 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 Bieman (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 always preferred. But in general, there is still a considerable deficit in understanding food plant relationships of phytophagous insects, in particular of those feeding on Poaceae 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 associated guilds. Informations on distribution and ecology of plant species were combined after Benkert et al. (1996), Ellenberg (1996), Haeupler & Schönfelder (1989) and Oberdorfer (2001). Species groups (“agg.” sensu Wisskirchen & Haeupler 1998) were treated as a single taxon, e.g. Ranunculus auricomus, Alchemilla spp. and Rubus fruticosus. Taxonomy and systematic order follow Wisskirchen & Haeupler (1998), although phylogenetic aspects will be discussed on the basis of Judd et al. (1999) (see chapter 5.1.3). The classification of the diet breadth applied here is shown in Table 4 and follows Schaefer (1992, slightly modified). Like other classifications, it is unable to solve the problem of
242
Utilization of plant resources
Table 4. Classification of diet breadth applied in this book, after Schaefer (1992), modified Diet breadth
Classification
Abbreviation
1 host species 1 host genus 1 host family 2 food plant families or up to 4 species which belong to no more than 4 plant families More
1st degree (= strictly) monophagous 2nd degree monophagous 1st degree oligophagous 2nd degree oligophagous
m1 m2 o1 o2
Polyphagous
po
inadequate definitions of supraspecific plant taxa. A host record is defined here as at least three nymphs or at least 10 adults, respectively, on an identified plant species. Records are treated as uncertain, if they have been gathered under unfavourable conditions, particularly after windy weather or late in the season, or if they largely consisted of parasitized individuals. An insect – plant relationship is assumed here, if there are at least 3 host records (or, alternatively, food plant records in vertical migrants), marked with “x” (or “X”, for at least 10 records) in the following tables. Less than 3 records are treated as uncertain relationships, marked with “?”. Parantheses denote host relations during the nymphal stage only (e.g. in species leaving the nymphal host after emergence). Feeding records from outside Germany were also treated as uncertain due to the possibility of regional changes. The assumption of an insect to be monophagous on the basis of three records may appear critical, but it should be noted that for the majority of species the record numbers are much higher, and that during recent years, almost all species, which were only rarely reported in the literature, turned out to be monophagous on little-studied or neglected plants. 5.1.2 Plant groups and their associated Auchenorrhyncha guilds 5.1.2.1 Pteridophyta The pteridophytes represent a geologically very old group and occur in considerable species numbers in many forest ecosystems worldwide. Ferns, in particular, can reach a high dominance in the herbaceous layer of central European forests, notably where air humidity is high. Their Auchenorrhyncha guilds, however, comprise only very few species (Table 5). Apart from 4 specialists, non of which occurs in high frequency, utilization 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 represent 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
x* x* X ?*
?*
?*
?*
?*
?*
X
0
0
0
0
0
1
x* A A ? ? 2
1
3
Overw intering stage
Substrate
D iet breadth
P. int erject um
Poly podium v ulgare
Poly st ichum spp.
Dry opt eris spp.
At hy rium filix-femina
A. scolopendrium
Asplenium spp.
Pt eridium aquilinum
E. sy lv at icum
Species Javesella stali (Metc.) X M acrosteles frontalis (Scott) X Ditropis pteridis (Spin.) Eupteryx filicum (N ew m .) Lepyronia coleoptrata (L.) ?* Cercopis vulnerata Rossi Centrotus cornutus (L.) Javesella pellucida (F.) ? Philaenus spumarius (L.) Empoasca decipiens Paoli M acrosteles sexnotatus (Fall.) ? Total 2
E. palust re
Equiset um arv ense
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).
m 1 P ny m 2 P eg Monophages m 1 P ny ?* o1 M eg Oligophages po X eg Polyphages po X ny Vertical po P ny m igrants po P ny. po X eg Unpo M? ad certain po P eg 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 oligophagous 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 Cercopis 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 defence 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 deciduous 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 species of gymnosperms have been introduced and are usually grown as ornamental trees in gardens and parks, except the western North American douglas fir (Pseudotsuga menziesii), 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 overview 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 Pinaceae, 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 specialists, two of them – Cixidia confinis (Zett.) and Aphrophora corticea Germ. – show typical pine bark colouration. C. confinis (Zett.) does not feed on phloem, but on undetermined 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 Switzerland. 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
A
W W W W W W W W W
? ?
A A A W W
?** ?** ? ?
?
W W W W W W ? ?**
W W W? ? ?** W
? ? ? ?** W? W? W? W? ? 4
?**
0
? ? 6
0
1
0
0
1
D iet breadth
Overw intering stage
X* X ?** X ?** X ?**
m 1? m 1? m 1? m 1? m 1? m1 m2 m 1? (o1) o2 o2? W (o1) W po m2 (m2) m1 (m1) m2 (m2) (m 1) o2 o2? o2? m 2? (m2?) po po W po o1 m 1? o1? m2 m2 o1 m2 po po 0
Substrate
X X X X
Taxus baccat a
Chamaecy paris law soniana
Juniperus communis
P. nigra
P. mugo
P. cembra
Pinus sy lv est ris
Larix decidua
Species Liguropia juniperi (Leth.) Pithyotettix abietinus (Fall.) ? Perotettix pictus (Leth.) ? Colobotettix morbillosus (Mel.) ? Cixidia confinis (Zett.) Grypotes puncticollis (H .-S.) W agneripteryx germari (Zett.) Cixius beieri W.Wg. ? Haematoloma dorsatum (Ahr.) A phrophora corticea Germ . Cixius similis Kbm . Balclutha punctata (F.) Empoasca vitis (Göthe) ? M etidiocerus rutilans (Kbm .) Tremulicerus fulgidus (F.) Z ygina lunaris (M. & R.) Z ygina nigritarsis Rem . Z ygina angusta Leth. Z ygina flammigera (Geoffr.) Z ygina rosincola (Cer.) Z ygina rosea (Fl.) ?** Tettigometra impressopunctata Duf. Tettigometra virescens (Panz.) Issus coleoptratus (F.) N eophilaenus campestris (Fall.) Dryodurgades reticulatus (H .-S.) Emposaca apicalis (Fl.) Linnavuoriana decempunctata (Fall.) Z yginella pulchra P. Löw Z ygina suavis R. Z ygina tiliae (Fall.) A llygus mixtus (F.) Thamnotettix confinis (Zett.) Total (excl. overw intering species) 0 0
Picea abies
Abies alba
Pseudot suga menz iesii
Table 6. Auchenorrhyncha species utilizing Gymnospermae in Germany. Explanations see Table 5.
M? P P P F P M P X X P P P P P M M M M M M P P P X P M? P P M M P P
ad ny ny ny ny eg eg? ny ny eg ny ad ad ad ad ad ad ad ad ad ad ad ad ny eg ad ad ad ad ad ad eg ny
Monophages
Vertical m igrants
Overw intering
Uncertain
* = 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 addition, 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 mentioned 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 northern 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 monotypic 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 specialist 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 (Ranunculus 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 predominate. In general, the Ranunculaceae play only a minor role as resources for Auchenorrhyncha, 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
X ?
? x* x*
? ? 1
1
2
1
0
1
? 0
Overw intering stage
S x
Substrate
Th. flav um
Thalict rum minus
R. mont anus
R. bulbosus
Ranunculus repens
? X
D iet breadth
M icantulina micantula (Zett.) Eupteryx vittata (L.) Issus coleoptratus (F.) Philaenus spumarius (L.) Erythria manderstjernii (Kbm .) Hyalesthes obsoletus Sign. Cercopis sanguinolenta (Scop.) A phrophora alni (Fall.) Centrotus cornutus (L.) Empoasca decipiens Paoli Eupteryx atropunctata (Goeze) Total
Clemat is v it alba
Species
Calt ha palust ris
Table 7. Auchenorrhyncha species utilizing Ranunculaceae in Germany. Explanations see Table 5.
m 1? o2 po po po po po po po po po
M M P X M P X X P M? M
ad eg ny eg ad ny ny eg ny ad eg
Monophages Oligophages Polyphages Vertical m igrants Uncertain
* = 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 European 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 proportion 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 species in the 2nd degree, namely Edwardsiana ulmiphagus Wils. & Clar., E. ishidai (Mats.),
248
Utilization of plant resources
6
15
U. glabra ? ? 9
Overw intering stage
x ? ? ? ?
? X X X X x ? x x x x
Substrate
? ? X X X X
X X ? X X X X X X x X x x x A A ? ? ? ? ? ? ?
D iet breadth
X ?
U. minor*
Species Ribautiana ognevi (Zachv.) M acropsis glandacea (Fieb.) Iassus scutellaris (Fieb.) Edwardsiana smreczynskii Dw or. Kyboasca bipunctata (Osh.) Edwardsiana ulmiphagus Wils. & Clar. Edwardsiana ishidai (Mats.) Edwardsiana plebeja (Ed w .) Ribautiana ulmi (L.) Z onocyba bifasciata (Boh.) Issus coleoptratus (F.) A lebra wahlbergi (Boh.) Empoasca vitis (Göthe) Fagocyba cruenta (H .-S.) A lnetoidia alneti (Dhlb.) Lamprotettix nitidulus (F.) A llygidius atomarius (F.) Cixius nervosus (L.) Empoasca affinis N ast Edwardsiana frustrator (Ed w .) Edwardsiana lethierryi (Ed w .) Fieberiella florii (Stal) å A llygus modestus Scott A llygidius commutatus (Fieb.) Hesium domino (Reut.) Total
Ulmus laev is
Table 8. Auchenorrhyncha species utilizing Ulmaceae in Germany. Explanations see Table 5.
m1 m 1? m 1? m 2? m2 m2? m2 m2 m2 m2 o2 po po po po po po o2? po po po o2 po po po po
M P P M M M M M M M P M P M M P P P M? M M P P P P
eg eg eg eg eg eg eg eg eg eg ny eg ad eg eg eg eg eg ad eg eg eg eg eg eg
Monophages
Oligophages
Polyphages
Vertical m igrants
Unconfirm ed
* = 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 Glycyrrhiza in eastern parts of Europe (Dworakowska 1973; Lauterer 1984). Zonocyba bifasciata (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 Lamprotettix 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 common, 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 perhaps 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 Macropsis 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 churchyards, 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 polyphagous Eupteryx aurata (L.). Furthermore, it harbours breeding populations of the broadly polyphagous species Philaenus spumarius (L.), Aphrodes makarovi Zachv. and Evacanthus interruptus (L.). Finally, nymphal development is documented for the vertical migrants 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 distribution (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 mammals, 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 community 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 suitable host for the spring generation of Eupteryx aurata (L.), because the additional summer 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 preferred 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 differential utilization of the Urticaceae. Finally, it is noteworthy that Zabel & Tscharntke (1998), who studied Auchenorrhyncha, Heteroptera and Coleoptera on Urtica dioica, could demonstrate that habitat fragmentation affected species numbers of monophages more negatively 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 Germany. 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 submontane 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 ornamental 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 emphasis 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 unconfirmed 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 pronounced, 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 deciduous 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 dominance 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 confined to southern Europe (Lang 1994). Important aspects of this question could be answered 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
? ? ?
Cast anea sat iv a
Q u. pubescens ?*
?* ?* ?*
?* ?* ? ?* ?*
?* ?* ?* ?*
?* ?*
?* ?
Overw intering stage
? ? x x X x x ?*
?*
Substrate
?
X X X X X x X ?* x X x ? ? ? x ?* x ?* x x x A A A A A A A ? A A A A A A A A ? ? ?
? X X X X X X ? X X ?* x X x ? ? ? x ?* x ?* x x x ? ? A A ? ? ? ? A A A A A ? A A ?
D iet breadth
X X ?
Q u. pet raea
Q uercus robur
Species Eurhadina saageri W.Wg. A lebra albostriella (Fall.) A lebra viridis R. Eurhadina kirschbaumi W.Wg. Iassus lanio (L.) Fagocyba carri (Ed w .) Eurhadina ribauti W.Wg. Eurhadina pulchella (Fall.) Ribautiana alces (Rib.) Ribautiana scalaris (Rib.) A rboridia velata (Rib.) A rboridia spathulata (Rib.) Typhlocyba quercus (F.) Eurhadina concinna (Germ .) Z ygina angusta Leth. Edwardsiana flavescens (F.) Fagocyba cruenta (H .-S.) A lnetoidia alneti (Dhlb.) Empoasca vitis (Göthe) Lindbergina aurovittata (Dgl.) Issus coleoptratus (F.) Issus muscaeformis (Schrk.) Penthimia nigra (Goeze) Ledra aurita (L.) Edwardsiana frustrator (Ed w .) Cixius distinguendus Kbm . Cixius dubius W.Wg. Cixius stigmaticus (Germ .) Tachycixius pilosus (Ol.) Centrotus cornutus (L.) Platymetopius major (Kbm .) Platymetopius guttatus Fieb. Lamprotettix nitidulus (F.) A llygus communis (Ferr.) A llygus mixtus (F.) A llygus maculatus Rib. A llygus modestus Scott A llygidius commutatus (Fieb.) A llygidius atomarius (F.) Thamnotettix confinis (Zett.) Thamnotettix dilutior (Kbm .) Cixidia pilatoi D’Urso & Gugl. Tettigometra impressopunctata Duf. Edwardsiana ampliata (W.Wg.)
Fagus sy lv at ica
Table 9. Auchenorrhyncha species utilizing Fagaceae in Germany. Explanations see Table 5.
m 1? m 1? m 1? m1 m2 m2 m2 m2 m 2? m 2? m2 m2 o2? o2? o2 o2 po po po po po po? po po po po po po po po po po? po o2? po? o2 po? po po po po po? po? o2?
M M M M P M M M M M M M M M M M M M P M P P P P M P P P P P P P P P P P P P P P P F P M
eg eg eg eg eg eg eg eg eg eg ad ad eg eg ad eg eg eg ad eg ny ny ny ny eg ny ny ny ny ny eg eg eg eg eg eg eg eg eg ny ny ny ad eg
1st d egree m onophages
2nd d egree m onophages
Oligophages
Polyphages
Vertical m igrants
Unconfirm ed
252
Utilization of plant resources
5 8
?*
33 22
25 23
0 10
?* ?* 0 11
Overw intering stage
? ? ? ? ? ?
Substrate
?*
Cast anea sat iv a
Q u. pubescens
Q u. pet raea
?* ?* ? ? ? ? ? ? ? ? ?
D iet breadth
?
Q uercus robur
Species Edwardsiana lamellaris (Rib.) Ribautiana debilis (Dgl.) Z yginella pulchra P. Löw A rboridia parvula (Boh.) A rboridia ribauti (Oss.) Ribautiana tenerrima (H .-S.) Z ygina rosincola (Cer.) Platymetopius undatus (De G.) Fruticidia bisignata (M. & R.) Z ygina flammigera (Geoffr.) A lebra wahlbergi (Boh.) A rboridia erecta (Rib.) Total substantiated Total unconfirm ed
Fagus sy lv at ica
Table 9. (continued):
o2? po? o2 o2 o2 o2? o1? po? o2 o1? po o2?
M M M M M M M P M M M M
eg eg ad ad ad eg ad eg ad ad eg ad
Unconfirm ed
* = 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 between 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 family, although there is a considerable overlap. In general, most mesophyll feeders are monophagous and overwinter as egg, whereas phloem feeders tend to be host generalists and comprise about equal numbers of species wintering in the egg stage and as nymphs, respectively. 5.1.2.7 Betulaceae (incl. Corylaceae) The birch family includes 4 native genera with altogether 9 species, all of which are woody. Hornbeam (Carpinus betulus), hazel (Coylus avellana), black alder (Alnus glutinosa) and silver birch (Betula pendula) are all widespread and common in low and submontane altitudes, the latter species reaching up to the alpine tree line. Carpinus betulus is a climax species on locations of intermediate humidity and fertility. Corylus avellana is essentially a heliophilous underwood shrub of open forests. Betula pendula is a pioneer tree occupying a wide ecological amplitude on soils with extremely varying supply of humidity and nutrients. In contrast, Alnus glutinosa is confined to wet and temporarily flooded locations, often along streams and rivers. Downy birch (B. pubescens) is widespread in the north German plain; in central and southern parts, however, it prefers higher and cooler sites on wet and acidic soils. Grey and green alder (Alnus incana and A. alnobetula) grow in higher altitudes, the former mainly along streams and on seepy mountain slopes (although now often planted in lower altitudes). The latter is a dominant shrub of the subalpine „krummholz“ belt, with scattered occurrence as a pioneer species in lower altitudes and in the Black Forest. Tree hazel and filbert (Corylus colurna and C. maxima), both of southwest European and western Asian origin, are commonly planted in parks and gardens. Finally, dwarf and shrub birch (Betula nana and B. humilis) occur very rarely in bogs of north Germany and the foothills of the Alps. The data base of Auchenorrhyncha is quite extensive for Carpinus betulus, Corylus avellana, Alnus glutinosa and Betula pendula, but only moderate and incomplete for B. pubescens, Alnus incana and A. alnobetula. There are a few samples from Corylus colurna and C. maxima, but none from Betula nana and B. humilis. Altogether, 61 species of Auchenorrhyncha are confirmed feeders, further 27 are unconfirmed (Table 10). Thus, the total number is among the highest of all central European plant families, surpassed only by the Salicaceae, Cyperaceae and Poaceae. Similarly, the ratio Auchenorrhyncha species number : plant species number is the second highest (6.8 vs. 8.4 in the Fagaceae). Betula pendula, Alnus glutinosa and Corylus avellana, with 28, 19 and 16 species, respectively, are among the most-favoured plants. On the other hand, low species numbers are found on the localized Alnus alnobetula as well as on the introduced Corylus colurna and C. maxima. Low species numbers can also be predicted for Betula nana and B. humilis. The ratio specialists : generalists is < 1 (except in A. alnobetula and the introduced hazels, which are insufficiently studied) indicating that plant defence mechanisms may either be less effective or less specific. As in other tree families,
254
Utilization of plant resources
Cory lus av ellana
C. colurna
C. maxima
Bet ula pendula
B. pubescens
Alnus glut inosa
A. incana
A. alnobet ula
D iet breadth
Substrate
Overw intering stage
Species Oncopsis carpini (J. Shlb.) Z ygina griseombra Rem . Oncopsis avellanae Ed w . Edwardsiana avellanae (Ed w .) Edwardsiana spinigera (Ed w .) Edwardsiana stehliki Laut. A lebra coryli Le Q. Oncopsis appendiculata W.Wg. Kybos calyculus (Cer.) Z ygina rosea (Fl.) Oncopsis tristis (Zett.) Oncopsis flavicollis (L.) Oncopsis subangulata (J. Shlb.) Kybos lindbergi (Lnv.)** Linnavuoriana decempunctata (Fall.) Kybos mucronatus (Rib.) Edwardsiana gratiosa (Boh.) Eupterycyba jucunda (H .-S.) Kybos strobli (W.Wg.) Edwardsiana soror (Lnv.) Linnavuoriana intercedens (Lnv.) Oncopsis alni (Schrk.) Kybos smaragdula (Fall.) Edwardsiana alnicola (Ed w .) Edwardsiana geometrica (Schrk.) Z ygina tiliae (Fall.) Edwardsiana plurispinosa (W.Wg.) Edwardsiana bergmani (Tull.) A lebra neglecta W.Wg. Edwardsiana flavescens (F.) Z onocyba bifasciata (Boh.) A rboridia ribauti (Oss.) Eurhadina concinna (Germ .) Typhlocyba quercus (F.) Edwardsiana lanternae (W.Wg.) Empoasca vitis (Göthe) Fagocyba cruenta (H .-S.) A lnetoidia alneti (Dhlb.) A lebra wahlbergi (Boh.) Edwardsiana frustrator (Ed w .) Issus coleoptratus (F.) Ledra aurita (L.) A guriahana stellulata (Burm .) Cixius nervosus (L.) Cixius cunicularius (L.) Cixius dubius W.Wg.
Carpinus bet ulus
Table 10. Auchenorrhyncha species utilizing Betulaceae in Germany. Explanations see Table 5.
X X . . . . . . . . . . . . . . . . . . . . . . . . . . X X X X ? ?* . x X X X ? . . . . . .
. . X X X X X . . . . . . . . . . . . . . . . . . . X . . . . . . ?* . x x X ? x x ? . A A A
. . . . . . x . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? ? x . . . . . . . .
. . . . . . x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . X . . X X X X X . . . . . . . . . . . . X . ? . . ?* . . x x ?* x ? x x x A A .
. . . . . . . ? ?* x X X x* ?* X X . . . . . . . . . . . . X . . . . ? . . x ? ?* . . . . . . . .
. . ?* . . . . . . . . . . . . ?* X X . . . X X x X X X x . . ? . ?* . ?* x x X x . . ? . A A .
. . . . . . . . . . . . . . . . . . ?* x x X X X X x ? . . . . . . . . x ? X . . . . . A . .
. . . . . . . . . . . . . . . . . . . ? ?* . ?* . ? . x x . . . . . . . ? ? ? . . . . . ? . .
m1 (m 1) m1 m1 m1 m 1? m2 m 1? m 1? m 1? m2 m2 m 2? m2 m2 m 1? m1 m1 m 1? m 1? m 1? m2 m2 m2 m2 o2 o1 o1 o2 o2 o2 o2 o2? o2? o2? po po po po po po po po po po po
P M P M M M M P M M P P P M M M M M M M M P M M M M M M M M M M M M M P M M M M P P M P P P
eg ad eg eg eg eg eg eg eg ad eg eg eg eg ad eg eg eg eg eg ad eg eg eg eg ad eg eg eg eg eg ad eg eg eg ad eg eg eg eg ny ny eg ny ny ny
Monophages
Oligophages
Polyphages
Vertical m igrants
Betulaceae (incl. Corylaceae)
255
Cory lus av ellana
C. colurna
C. maxima
Bet ula pendula
B. pubescens
Alnus glut inosa
A. incana
A. alnobet ula
D iet breadth
Substrate
Overw intering stage
Species Cixius stigmaticus (Germ .) Cixius similis Kbm . Tachycixius pilosus (Ol.) A phrophora alni (Fall.) A phrophora major Uhler Platymetopius major (Kbm .) Platymetopius guttatus Fieb. Lamprotettix nitidulus (F.) A llygus communis (Ferr.) A llygus mixtus (F.) A llygus modestus Scott A llygidius commutatus (Fieb.) Speudotettix subfusculus (Fall.) Hesium domino (Reut.) Thamnotettix confinis (Zett.) Cixius cambricus China Cixius heydenii Kbm . Cixius simplex (H .-S.) Pentastiridius beieri (W.Wg.) Issus muscaeformis (Schrk.) Lepyronia coleoptrata (L.) Philaenus spumarius (L.) Centrotus cornutus (L.) A lebra albostriella (Fall.) A lebra viridis R. Empoasca affinis N ast Empoasca decipiens Paoli Edwardsiana ampliata (W.Wg.) Edwardsiana ishidai (Mats.) Lindbergina aurovittata (Dgl.) Ribautiana debilis (Dgl.) Ribautiana tenerrima (H .-S.) Eurhadina ribauti W.Wg. Eurhadina pulchella (Fall.) A guriahana pictilis (Stal) å Z yginella pulchra P. Löw Z ygina angusta Leth. Z ygina rosincola (Cer.) A rboridia erecta (Rib.) Platymetopius undatus (De G.) Idiodonus cruentatus (Panz.) Colladonus torneellus (Zett.) Total substantiated Total unconfirm ed
Carpinus bet ulus
Table 10. (continued):
. . . . . . . . . ? . . A . ? . . . . . . . . . . ? . . . . . . ? ? . . ? . . . . . 11 9
. . . A . . . . . . . . A . ? . . . . ? ?* ?* ? . . . ? ? ? ?* ?* ?* . . . . . . ?* . ? . 16 16
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0
A . A A A A A ? A A A A A A A . . ? . . . ?* ? ? . . ? . . . ?* ? . ? . . . ? . ? ? ? 28 17
. A . A A . . . . ? . . . A . . . ? . . ?* ?* . . . . . . . ?* . . . . ? . . . . . . . 12 11
? . . A . . . A . A A . ? ? ? . . ? . . . ?* ? ? . . . . . ?* ?* ? ? . . ? . . . . . ? 19 20
. . . A . . . . . ? . . . . . . . ? ?* . . . . . . . . . . . . . . . . . . . . . . . 10 7
. . . ? . . . . . . . . ? . . ?* ?* . . . . . . . ? . . . . . . . . . . ? . . . . . . 2 14
po o2? po po po po po po o2? po? po? po? po o2? po po? po? po o2? po? po po po m 1? m 1? po po o2? m 2? po po? o2? m2 m2 m 1? m 2? o2 o1? o2? o2? po po
P P P X X P P P P P P P P P P P P P P P X X P M M M? M? M M M M M M M M M M M M P P P
ny ny ny eg eg eg eg eg eg eg eg eg ny eg eg ny ny ny ny ny eg eg ny eg eg ad ad eg eg eg eg eg eg eg eg? ad ad ad ad eg eg ny
Vertical m igrants
Unconfirmed Unconfirm ed
* = 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 prevalence 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 hibernation and to shrub species of Rosaceae in spring. Alebra neglecta W.Wg., Edwardsiana flavescens (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 plurispinosa (W.Wg.), which reproduces both on Corylus avellana and Alnus spp. Among polyphages, Alnetoidia alneti (Dhlb.) is clearly the most abundant species, accompanied by smaller numbers of Empoasca vitis (Göthe), Fagocyba cruenta (H.-S.), Edwardsiana frustrator (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 pendula 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 (although 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 decempunctata (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 stellulata (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, Platymetopius, Allygus and others. Adults of Platymetopius major (Kbm.), P. guttatus Fieb., Allygus 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 reproduce 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.), Empoasca vitis (Göthe), Fagocyba cruenta (H.-S.) and Alnetoidia alneti (Dhlb.) are polyphagous.
Tiliaceae
257
Vertical migrants include Cixius nervosus (L.), Aphrophora alni (Fall.), Lamprotettix nitidulus (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 oakhornbeam 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 vertical migrant and 10 unconfirmed species (Table 11). The degree of specificity is low, without
* = after Chudzicka (1986a, 1986b)
? ?
? ? 6
Overw intering stage
x X x X
Substrate
X x
D iet breadth
X X x x X X X X X A ? ? ? ? ?* ?* ? ? ? ?* 10
T. plat y phy llos
Species Pediopsis tiliae (Germ .) Edwardsiana lethierryi (Ed w .) Z ygina tiliae (Fall.) Issus coleoptratus (F.) A lebra wahlbergi (Boh.) Empoasca vitis (Göthe) Fagocyba cruenta (H .-S.) A lnetoidia alneti (Dhlb.) A guriahana stellulata (Burm .) Cixius nervosus (L.) Ledra aurita (L.) Edwardsiana flavescens (F.) Edwardsiana frustrator (Ed w .) Eurhadina concinna (Germ .) Edwardsiana ampliata (W.Wg.) Typhlocyba quercus (F.) Z onocyba bifasciata (Boh.) A rboridia ribauti (Oss.) Lamprotettix nitidulus (F.) A llygus mixtus (F.) Total
Tilia cordat a
Table 11. Auchenorrhyncha species utilizing Tiliaceae in Germany. Explanations see Table 5.
m2 o2 o2 po po po po po po po po o2 po o2 o2? o2? o2 o2 po po
P M M P M P M M M P P M M M M M M M P P
eg eg ad ny eg ad eg eg eg ny ny eg eg eg eg eg eg ad eg eg
Monophages Oligophages
Polyphages
Vertical m igrants
Unconfirm ed
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 oligophages, 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 glutinosa 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 preferring 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 unintentionally been sampled for their Auchenorrhyncha. Although most low-growing willows (e.g. Salix retusa, S. herbacea, S. reticulata) are supposed to be devoid of Auchenorrhyncha, 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 species 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 Germany, 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 specialists 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 altogether 12 monophages in the 1st degree and 24 in the 2nd degree. The following section treats (i) 1st degree monophages and their hosts, (ii) monophages specific to narrow-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 species perhaps breeds also on S. pentandra. Finally, Kybos digitatus (Rib.) is probably strictly 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, Macropsis, 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 specific. Thus, Metidiocerus impressifrons (Kbm.), Kybos limpidus (W.Wg.) and Edwardsiana tersa (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 sallows. 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 families 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 Auchenorrhyncha 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 include Idiocerus lituratus (Fall.), Kybos strigilifer (Oss.), Macropsis prasina (Boh.), Metidiocerus elegans (Fl.), Populicerus confusus (Fl.), Edwardsiana salicicola (Edw.) and Linnavuoriana 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 somewhat 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 willow and sallow species. These include Aphrophora pectoralis Mats., A. salicina (Goeze), Macropsis cerea (Germ.), Idiocerus stigmaticalis Lew., Metidiocerus rutilans (Kbm.) and Kybos virgator (Rib.). (v) Oligo- and polyphages, and vertical migrants In contrast, numbers and proportions of oligophages and polyphages utilizing Salicaceae 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 include 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 monophages on Populus alba, P. nigra and Salix purpurea are among the highest of all central European 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, abundance) and plant physiology (chemistry). Thus the high content of phenolic heterosides, which are toxic to many insects and which are largely restricted to the Salicaceae, is likely to play an important role.
Salicaceae
261
X X X X . . . . . . . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . . .
. . . . X X X X X X ? . . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . . .
. . . . . . . . . . . X X X X X X X . . . . . . . . . . . . X . . . . . . . . . . . . . .
. . . . . . . . . . . . . ? x x x x . . . . . . . . . . . . x . . . . . . . . . . . . . .
. . . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . . x X X x ? X . x x ? .
. . . . . . . . . . . . . . . . . . X X X . . . . . . . . . . . . ? X X X X X x x X X ? .
. . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . ? . ? . . ? x . . . . x ? ?
. . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . ? x x . . x x X x . x X X 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 . . .
Substrate Overw intering stage
D iet breadth
Species M acropsis fuscinervis (Boh.) Tremulicerus tremulae (Estl.) Populicerus laminatus (Fl.) Populicerus populi (L.) M acropsis vicina (H orv.) Tremulicerus distinguendus (Kbm .) V iridicerus ustulatus (M. & R.) Populicerus albicans (Kbm .) Edwardsiana candidula (Kbm .) Z ygina nivea (M. & R.) Z ygina tithide Ferr. Stenidiocerus poecilus (H .-S.) Kybos abstrusus (Lnv.) Tremulicerus fulgidus (F.) Tremulicerus vitreus (F.) M acropsis graminea (F.) Rhytidodus decimusquartus (Schrk.) Populicerus nitidissimus (H .-S.) M acropsis albae W.Wg. M acropsis gravesteini W.Wg. M acropsis najas N ast M acropsis viridinervis W.Wg. M acropsis remanei N ick. M acropsis marginata (H .-S.) M acropsis haupti W.Wg. Idiocerus similis Kbm . Kybos rufescens Mel. Idiocerus vicinus Mel. M imallygus lacteinervis (Kbm .) Kybos digitatus (Rib.) Kybos populi (Ed w .) Edwardsiana tersa (Ed w .) M etidiocerus impressifrons (Kbm .) Kybos limpidus (W.Wg.) Idiocerus herrichii (Kbm .) Z ygina lunaris (M. & R.) Kybos virgator (Rib.) Idiocerus stigmaticalis Lew . M etidiocerus rutilans (Kbm .) M acropsis cerea (Germ .) A phrophora pectoralis Mats. A phrophora salicina (Goeze) Z ygina ordinaria (Rib.) Sagatus punctifrons (Fall.) Kybos butleri (Ed w .)
Populus t remula P. alba P. nigra Populus hybrids Salix fragilis S. alba S. pent andra S. t riandra S. v iminalis S. eleagnos S. purpurea S. repens S. aurit a S. cinerea S. caprea S. appendiculat a S. my rsinifolia
Table 12. Auchenorrhyncha species utilizing Salicaceae in Germany. Explanations see Table 5.
. . m1 P . . m1 P . . m1 P . . m1 P . . m1 P . . m1 P . . m1 P . . m1 P . . m1 M . . m 1? M . . m 1? M . . m1 P . . m1 M . . m1 P . . (m 1) P . . (m 1) P . . (m 1) P . . (m 1) P . . m1 P . . m1 P . . m1 P . . m 1? P . . m1 P . . m1 P . . m1 P . . m1 P . . m1 M . . m 1? P . . m 1? P . . m 1? M . . m2 M . . m 2? M . . m2 P . . m2 M . . m2 P . . m 2? M . . m2 M . . m2 P . . m2 P . x m2 P ? x m2 X . . m 2? X . . m2 M . . m2 P . . m2 M
eg eg eg eg eg eg ad eg eg ad ad ad eg ad eg eg ad eg eg eg eg eg eg eg eg eg eg ad eg eg eg eg ad eg ad ad eg eg ad eg eg eg ad eg eg
1 st d egree m onophages
2nd d egree m onophages
262
Utilization of plant resources
. . X X X X X X . . x x . A . . A . . . ? . . ?* ?* . . . . ? ? . . . . 17 6
. . m2 P . . m2 P . x m2 M . . m2 P . . m2 P x x m2 P x . m2 M . . m2 M . . m2 P . x m2 P . . o2 M . . po P . . po M . ? po P . . m 2? P . . o2? P . ? po X . . po X . . po P . . po P . . po P . . po P . . po P . . po X . . po X . . po P . . po P . . po P . . po M . . po M? . . o2 M . . o2? M . . po? P . . po M . . po P 2 5 1 2
x X x X X X X X . . x ? . . . . A . . ? . . . . . . . . . . . . . . . 16 4
. X X X X X X X . X x x . A . . A . . . . . . . ?* . . . . . . ? ? . . 17 4
Substrate Overw intering stage
x* X . . . . . . . . . . . . . . . . . . . . . ?* ?* . . . . . . . . . . 6 2
D iet breadth
S. appendiculat a S. my rsinifolia
. . . . ? . . . . . . x . . ? A A A . A . . . . . . . . . . . . . . ? 19 5
S. cinerea S. caprea
S. eleagnos S. purpurea
. . . . . . . . . . . . ? . . . . . . . . . . . . . . . . . . . . ? . . ? ? . ? X . . . . . . ? . . . ? . . X x . x . . X . . . . . . . . . . . . . . x . ? . x ? x x x . . . . . . . . A A . A A . . . . . A A . . . . . . . . ? A A . A A A . . A . . . . . . . . . . . . . . . . . ? . ? . . . . . . . . . . . . . . . . . . . . . . ?* . . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . ? . . . . . . . . . . . 5 10 16 2 16 18 7 3 5 5 8 2 5 8
S. repens S. aurit a
. . . . . . . . . . . x x . . . A . . A A . . . . . . . ? ? . . . . . 12 3
S. t riandra S. v iminalis
. . . . . . . . . . . x . . . . A . . A . . . . . . . . . . . . . . . 10 1
S. alba S. pent andra
. . . . . . . . . . . x . . . . A . ? . . . . ?* ?* ? ? ? ? . . . . . . 7 7
Salix fragilis
P. alba P. nigra Populus hybrids
Species M acropsis impura (Boh.) Idiocerus lituratus (Fall.) Kybos strigilifer (Oss.) M acropsis prasina (Boh.) M etidiocerus elegans (Fl.) Populicerus confusus (Fl.) Edwardsiana salicicola (Ed w .) Linnavuoriana sexmaculata (H ard y) M acropsis notata (Proh.) M acropsis infuscata (J. Shlb.) Edwardsiana prunicola (Ed w .) Empoasca vitis (Göthe) A guriahana stellulata (Burm .) Cixius nervosus (L.) M yndus musivus (Germ .) Pentastiridius beieri (W.Wg.) A phrophora alni (Fall.) A phrophora major Uhler Centrotus cornutus (L.) Stictocephala bisonita Kopp & Yonke A llygus mixtus (F.) Hyalesthes obsoletus Sign. Issus coleoptratus (F.) Lepyronia coleoptrata (L.) Philaenus spumarius (L.) Ledra aurita (L.) Penthimia nigra (Goeze) Speudotettix subfusculus (Fall.) Fagocyba cruenta (H .-S.) Empoasca affinis N ast Edwardsiana flavescens (F.) Ribautiana tenerrima (H .-S.) Colladonus torneellus (Zett.) A lnetoidia alneti (Dhlb.) Fieberiella florii (Stål) Total substantiated Total unconfirm ed
Populus t remula
Table 12. (continued):
eg eg eg eg eg? eg eg ad eg eg eg ad eg ny ny ny eg eg ny eg eg ny ny eg eg ny ny ny eg ad eg eg ny eg eg
2nd d egree m onophages
Oligo-/ Polyphages
Vertical migrants
Unconfirm ed
* = 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, usually 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 degradation 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 habits. The genus Rhododendron, in particular, includes numerous ornamental species and varieties 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 phytophagous 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 Andromeda, 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 Germany (Table 13), plus 8 which need further confirmation. Heather (Calluna vulgaris), being
?
x x A x A
? ?
?
x*
? ?
X X X X X ?* ? ? X ?
?
x ?
?
? ?* ?* ?*
?* ?*
?* ?
?* ? ?
?
?
1
?* 5
0
0
1
W? ?* 6
0
Overw intering stage
X
Substrate
X
D iet breadth
E. carnea
Erica t et ralix
Calluna v ulgaris
V. oxy coccos
V. v it is-idaea
V. uliginosum
Vaccinium my rt illus
Species Graphocephala fennahi Young Ulopa carneae W.Wg. Ulopa reticulata (F.) Z ygina rubrovittata (Leth.) Ophiola cornicula (Marsh.) Ophiola russeola (Fall.) Erythria aureola (Fall.) Philaenus spumarius (L.) Erythria manderstjernii (Kbm .) Cixius similis Kbm . A phrophora corticea Germ . Idiodonus cruentatus (Panz.) Fieberiella septentrionalis W.Wg. Lepyronia coleoptrata (L.) A guriahana pictilis (Stål) Colladonus torneellus (Zett.) Ommatidiotus dissimilis (Fall.) Planaphrodes trifasciata (Geoffr.) Empoasca vitis (Göthe) A rboridia ribauti (Oss.) Ophiola decumana (Kontk.) Total
Rhododendron cultivated
Table 13. Auchenorrhyncha species utilizing Ericaceae in Germany. Explanations see Table 5.
(m 2)? m1 m1 m1 o1? o1 o2 po po o2 (po) po po po m 1? po? m 1? o2? po o2 o1?
X P P M P P M X M P X P P X M P P P P M P
eg ny/ ad ny/ ad ad eg eg eg eg ad ny eg eg eg eg eg? ny eg eg ad ad eg
2
* = after Günthart (1987a), Holgersen (1945), Ossiannilsson (1981), Strübing (1955)
Monophages
Oligophages Polyphages Vertical m igrants
Unconfirm ed
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 monophagous 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 sporadic 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 crossleaved 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 properly studied. The only feeding record is of Ophiola russeola (Fall.) on cranberry (V. oxycoccos). 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 include 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 monophagous (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 basic 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 polyphagous 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 hillsides in southern Germany had no positive results, but this statement should be considered 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 Auchenorrhyncha on cultivated apple has been done in eastern Germany (Lehmann 1973a) and in Switzerland (Günthart 1971). Edwardsiana crataegi (Dgl.), f. froggatti (Bak.), clearly dominated 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 subalpine belt of the Alps. Finally, sorb (S. domestica) occurs rather localized in thermophilous 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 dominating. 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
Sorbus aucuparia
S. aria
S. int ermedia
S. t orminalis
Crat eagus spp.
Py racant ha coccinea
Amelanchier spp.
Rubus idaeus
R. frut icosus
R. caesius
. . . ? . . . . . . ? . . . . .
Malus domest ica
. . . . . . . . . . . . . X x . . . . . . ? . . .
Py rus communis
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . x . . . . .
Cot oneast er cultivated
Species M acropsis brabantica W.Wg. M acropsis scotti Ed w . Edwardsiana rosaesugans (Cer.) Edwardsiana rhodophila (Cer.) Balcanocerus larvatus (H .-S.) Balcanocerus pruni (Rib.) Z ygina nigritarsis Rem . A rboridia kratochvili (Lang) Eupteryx signatipennis (Boh.) M acrosteles septemnotatus (Fall.) M acropsis fuscula (Zett.) M acropsis megerlei (Fieb.) A rboridia simillima (W.Wg.) Edwardsiana rosae (L.) Z ygina schneideri (Günth.) Z ygina flammigera (Geoffr.) Edwardsiana crataegi (Dgl.) Fruticidia bisignata (M. & R.) Ribautiana debilis (Dgl.) Z ygina rosincola (Cer.) Edwardsiana sociabilis (Oss.) Typhlocyba quercus (F.) Edwardsiana prunicola (Ed w .) Z ygina angusta Leth. A lebra neglecta W.Wg. Ribautiana tenerrima (H .-S.) A rboridia parvula (Boh.) Fagocyba cruenta (H .-S.) A lnetoidia alneti (Dhlb.) Empoasca vitis (Göthe) Ledra aurita (L.) A lebra wahlbergi (Boh.) Issus coleoptratus (F.) Empoasca decipiens Paoli Philaenus spumarius (L.) A guriahana stellulata (Burm .) Fieberiella septentrionalis W.Wg. Fieberiella florii (Stål) Lindbergina aurovittata (Dgl.) Cixius sticticus R. Tachycixius pilosus (Ol.) Reptalus panzeri (P. Löw )
Spiraea spp.
Table 14. Auchenorrhyncha species utilizing Rosaceae in Germany. Explanations see Table 5.
. . . . . . . . . . . . . ? . . . . ?* . . ? . . . ? . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . X ?* X X ?* ?* . . . ? ? . . . x X X x . . . . . . . . . . .
. . . . . . . . . . . . . ?* . ? . . . ?* . . . . . . . x ?* X . ? . . . . . . . . . .
. . . . . . . . . . . . . x . ? . . . . . x . . . . . x x ? . x x . . . . . . . . .
. . . . . . . . . . . . . x . x . . . . . . . . . . . x . X . x . . . . . . . . . .
. . . . . . . . . . . . . . . . ? . . . . . . . . . . x ? ? . x . . . . . . . . . .
. . . . . . S . . . . . . x S x X x . S . ? ? X X . . x x x . . . . . . . . . . ? .
. . . . . . . . . . . . . . x . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . x ? . . . . . . . ? . . . . . . . . . . . . . . . . . .
. . . . . . . . . . X . . ? . . ? . . . . . ? . . X ? ? ? X . . . ? . . X x ?* . . .
. X . . . . . . . . X . . ? . . . . X . . . . . . X . . . X . . . ? . . x ? ?* . . .
X . . . . . . . . . X . . . . . . . X . . . . . . X . ? . X . . . ? . . . . ?* . . .
Rosaceae
267
R. cory mbifera
R. glauca
R. pendulina
R. rubiginosa
R. rugosa
R. spinosissima
Prunus av ium
P. cerasus
P. domest ica
P. padus
P. spinosa
D iet breadth
Substrate
Overw intering stage
. . . . . . . . . . . . . . . . . . . . ? . . . . .
Rosa canina
. . . . . . . . . . . . . . x
Filipendula ulmaria
. . . . . . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Fragaria spp.
P. t abernaemont ani
. . . . . . . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . . . .
P. frut icosa
Pot ent illa incana
Table 14. (continued):
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ?* . . . . . . .
. . . . . . . . X X . . . . . . . . . . X . . . . . ? . . . . . . . X . . . . . . .
. . ? . . . . . . . . ? . X ? ? . . ? ? . ? ? x . ? . ? ? x . . . . . . x x . A ? A
. . . . . . . . . . . . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . ? . ? . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . X . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . . . .
. . . X . . . . . . . X ? X ? ? . . . . . . ? ? . . . ? . x . . . . . . x . . ? . ?
. . . . . . . . . . . ? . X X ? . . . . X ? . . . . . ? ? x . . . . . . x . . . . .
. . . . . . . . . . . X X . ? . . . . . . . . . . . . ? . x . . . . . . ? . . . . .
. . . . . . . . . . . . . x ? X x . . . . x ? ? ? ? . x X x . x . ? . x . . . . . .
. . . . . . . . . . . . . . . x* ?* . . . . x* . ?* . . . ?* x* . . ?* . ?* . ?* . . . . . .
. . . . . . . . . . . . . x ? X x . . . . x X ? . . . ? X x . x . ? . . . . . . . .
. . . . . . X . . . . . . . . X . . . ? . X . . X . ? x X x . . . ? . . . . . . . .
. . . . X X . . . . . . . ? X X x . . ? . x X x . ? . ? x x . ? ? ? . x X x . ? A A
m1 m1 m 1? m1 m1 m1 (m 1) m 1? m1 m1 m2 m2 m2 o1 o1 o1? o1 o1 o1? o1? o1 o2? o2? o2 o2 o2? o2? po po po po po po po po po po po po o2? po o2
P P M M P P M M M P P P M M M M M M M M M M M M M M M M M P P M P M? X M P P M P P P
eg eg eg eg eg ad ad ad eg eg eg eg ad eg ad ad eg ad eg ad eg eg eg ad eg eg ad eg eg ad ny eg ny ad eg eg eg eg eg ny ny ny
1st d egree monophages
2nd d egree m onophages
1st d egree oligophages
2nd d egree oligophages
Polyphages
Vertical m igrants
268
Utilization of plant resources
S. t orminalis
. . . . . . . . . . . . . . . . . ? . . . . . . . . . 6 3
. . . . . . . . . . . . . . . . . ? . . . . . . . . . 5 1
. . . . . . . . . . . . . . . . . . . . . . . . . . . 2 3
. . . . ?* . . . . . . . . ? ?* . ? . . . . . . . . . 12 7
R. caesius
S. int ermedia
. . . . . . . . . . . . . . . . . ?* . ?* . . . . . . . 2 7
R. frut icosus
S. aria
. . . . ?* . . . . . . . . . . . . . . . . . . . . . . 7 6
Rubus idaeus
Sorbus aucuparia
. . . . ?* . . . . . . . . . . . . . . . . . . . . . . 0 5
Amelanchier spp.
Malus domest ica
. . . . . . . . . . . . . . . . . . . . . . . . . . . 2 3
Py racant ha coccinea
Py rus communis
. . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0
Crat eagus spp.
Cot oneast er cultivated
Species Tibicina haematodes (Scop.) Cicadetta montana (Scop.) A phrophora alni (Fall.) Centrotus cornutus (L.) 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 Total unconfirm ed
Spiraea spp.
Table 14. (continued):
. . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0
. . . . . . . . . . . . . . . . . . . . . . . . . . . 1 2
. . . A . . ? . ? . . . . . . . . ? . . . ? ? . ? ?* . 6 15
. . . . . . . . . . ? . . . . . . . . . . . . . . . . 6 5
. . . . . . . . . . ? . . . . . . . . . . . . . . . . 5 4
* = after Claridge & Wilson (1981), Günthart (1980, 1987b), Hoffrichter & Tröger (1973), Lauterer & MalenVasarainen (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 imperfectly 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
Filipendula ulmaria
Rosa canina
R. cory mbifera
R. glauca
R. pendulina
R. rubiginosa
R. rugosa
R. spinosissima
Prunus av ium
P. cerasus
P. domest ica
P. padus
. . ?* . . . . . . ?* . . . . . . . . . . . . . . . . . 0 3
. . . . ?* A . . . ?* . . . . ?* . . . . . . . . . . . . . . ?* . . . . . . . ? . . . . . . . . . . . . ?* . ? . 4 8 5 15
. . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0
. . . . . . . . . . . . . . . . . . . . . . . . . . . 0 2
. . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1
. . ? . . . . . . . . . . . . . . . . . . . . . . . . 5 9
. . ? . . . . . . . . . . . . . . . . . . . . . . . . 5 6
. . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3
. . . . ? . . . . . . ? . . . . . . . . . . . . . . . 9 8
. . . . . . . . . . . ?* . . . . . . . . . . . . . . . 3 7
. . . . . . . . . . . ? . . . ?* ? . . . . . . . . . . 8 7
. A . A . A . ? . ? . ? . . . . . . . . . ? . ? ?* . . . . . . . ? ? . ? . . . . . ? . . . . . ? . . . . . . 7 18 5 17
po po po po po po po po po po po po o2? m2 m 2? o1? m 2? po o2? o2 po po po? po po m 1? m 1?
Overw intering stage
Fragaria spp.
. . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1
Substrate
P. frut icosa
. . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0
D iet breadth
P. t abernaemont ani
. . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0
P. spinosa
Pot ent illa incana
Table 14. (continued):
X ny X ny X eg P ny P eg P ny P ny X eg X eg M eg M? ad M? ad ? M? ad M ad M ad M ad M ad M eg M eg M eg P eg P eg P ny P ny P ny P eg P eg
Vertical m igrants
Unconfirm ed
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 Zygina 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 growing 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 undergrowth 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 serviceberry (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 species 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. brabantica 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 polyphagous 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 oligophages and polyphages, like Empoasca vitis (Göthe), Zygina schneideri (Günth.) and Fieberiella septentrionalis W.Wg. Among herbaceous species, only meadowsweet (Filipendula ulmaria), cinquefoil (Potentilla 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 native 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. potentillae (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 probably higher. Only 6 species, all of which are easy to identify, have been sampled for leafhoppers. Sampling of less known roses may reveal further, perhaps even undescribed species. Edwardsiana rhodophila (Cer.) and E. rosaesugans (Cer.) are both strictly monophagous 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 Edwardsiana 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 Japanese 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, including 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 perhaps 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 (Prunus cerasus), plum (P. domestica), peach (P. persica) and almond (P. dulcis), but also neophytes 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 altogether 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 quercus (F.), Zygina flammigera (Geoffr.) and Z. schneideri (Günth.). Other oligophages and polyphages include Edwardsiana rosae (L.), E. crataegi (Dgl.), Alebra wahlbergi (Boh.), Empoasca 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 exceptions, notably Potentilla and Filipendula, herbs remain almost completely uninfested. There is also a clear tendency of native species harbouring more phytophages than introduced ones (including archaeophytes). The high proportion of 1st degree oligophages 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 widespread. Most species overwinter as egg, and those wintering as adults migrate to evergreen 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 phytophages. The Auchenorrhyncha data, which are mainly based on studies by Claridge & Wilson (1976, 1981), seem roughly comparable to those from Germany. As shown in Fig. 3, British species numbers are generally lower on all studied Prunus species (which 25 P. avium
P. c erasus
P. domestic a
P. padus
P. spinosa
S Auchenorr hyncha
20
15
10
5
0
000000000000000000000000000000000000000000000 0000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000000 0000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000000 0000000000000000000000000000000000000000 000000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000000 0000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000000 0000000000000000000000000000000000000000 000000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000000 0000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000000 0000000000000000000000000000000000000000 000000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000000 0000000000000000000000000000000000000000 000000000000000000000000000000000000000000000 000000000000000000000000000000000000000000000
0000000000 00000000000000000000000000 000000000000000000000000 000000000000000000000000000 000 000 000 000 000 000000 000 000 000 000000000000000000000000 0000000000000000000000000 0 0 0 0 0 00 0 0 00000000000000000000000000 00 00 00 00 00 0000 00 00 000 000000000000000000000000 00000000000000000000000000 00 00 00 00 00 0000 00 00 00 000000000000000000000000 0000000000000000000000000 0 0 0 0 0 00 0 0 00000000000000000000000000 00 00 00 00 00 0000 00 00 000 000000000000000000000000 00000000000000000000000000 00 00 00 00 00 0000 00 00 00 000000000000000000000000 0000000000000000000000000 0 0 0 0 0 00 0 0 00000000000000000000000000 00 00 00 00 00 0000 00 00 000 000000000000000000000000 00000000000000000000000000 00 00 00 00 00 0000 00 00 00 000000000000000000000000 0000000000000000000000000 0 0 0 0 0 00 0 0 00000000000000000000000000 00 00 00 00 00 0000 00 00 000 000000000000000000000000 00000000000000000000000000 00 00 00 00 00 0000 00 00 00 000000000000000000000000 0000000000000000000000000 0 0 0 0 0 00 0 0 00000000000000000000000000 00 00 00 00 00 0000 00 00 000 000000000000000000000000 0000000000000000000000000000000000000000 00 00 0000000000000000000000000
00000000000000000000000 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 00000000000000000000000
0000000000 00000000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 00000000000000000 00000000000 00000000000 000000000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 00000000000000000 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 000000000000 0000000000
0000000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 000 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 00 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 000 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 00 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 000 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 00 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 000 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 00 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 000 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000000
00000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000 0 000000000000000000000000000000000000000000 000000000000000000000000000000000000000000 000 000000000000000000000000000000000000000000 000000000000000000000000000000000000000000 00 000000000000000000000000000000000000000000 000000000000000000000000000000000000000000 000000000000000000000000000000000000000000 000 000000000000000000000000000000000000000000 000000000000000000000000000000000000000000 00 000000000000000000000000000000000000000000 000000000000000000000000000000000000000000 0 000000000000000000000000000000000000000000 000000000000000000000000000000000000000000 0 00000000000000000000000000000000000000000000000000 00 000000000000000000000000000000000000000000 000000000000000000000000000000000000000000 0000000000000000000000000000000000000000 0 0000000000000000000000000000000000000000000 0 000000000000000000000000000000000000000000 00 00000000000000000000000000000000000000 0000000000 00000000000000000000000000000000000000000000 00000000000000000000000000000000000000 0 0000000000000000000000000000000000000000000 000000000000000000000000000000000000000000000
0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000 00 00 00000000000000000000000000000000000000 000000000000000000000000000000000000000 0 0 0000000000000000000000000000000000000000 00 00 00000000000000000000000000000000000000 0000000000000000000000000000000000000000 00 00 00000000000000000000000000000000000000 000000000000000000000000000000000000000 0 0 0000000000000000000000000000000000000000 00 00 00000000000000000000000000000000000000 000000000000000000000000000000000000000 0 0 0000000000000000000000000000000000000000000
000000000000000000000000000000000000000000000 00 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 00 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 0 0000000000000000000000000000000000000000000 0 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 0 0000000000000000000000000000000000000000000 0 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 0 0000000000000000000000000000000000000000000 0 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 0 0000000000000000000000000000000000000000000 0 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 00 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 00 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 00 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 00 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 00 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000 0000000000000000000000000000000000000000000
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 taxonomic 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 graminicolous 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 Auchenorrhyncha, 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 Auchenorrhyncha, 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 individuals. Genista is furthermore favoured by Stictocoris picturatus (C. Shlb.), which also lives on restharrow (Ononis spp.) and winged broom (Chamaespartium sagittale). Gargara genistae (F.) is apparently a less specific feeder to various woody Fabaceae, including restharrow Ononis spp., bladder senna (Colutea arborescens) and even herbaceous 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
. . . . . . . . . . . . . . . . . ?* . . . ?* . . . . ? . . 0
m1 m1 m 2? o1 o1 o1 o1 o1 o1? o1? o2 o2 o2? o2? o2? po po po po po po po po? o1? o2? po o1? o2? o2?
Overw intering stage
Lat hy rus spp. spp.
. . . . . . . . . ?* . . . . . . ? ? . . . ?* . . . ?* . . . 0
Substrate
Vicia spp.
X . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
D iet breadth
Vicia t enuifolia
. . . . . . . . . . . . . . . . . . . . x x . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . x* . . . x* . . x* . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . . . 1 1 1 2
Coronilla spp.
O nobry chis spp.
Hippocrepis comosa
Colut ea arborescens
Lot us corniculat us
Trifolium spp.
Medicago spp.
O nonis spinosa
. . . . . . . . X . . . . . . . x ? . . . . . . X . x . . . . . X . x . . . . . X ?* x . x ?* ?* . . . X x X . ? . . . . . . x x x ? . . . . ?* X . . . . . . . ?* . . . . . . x* x* ?* . . . . . ? X . . . . . . x* x* . . . . . . . ? . . . . . . . . x ? . . . . ?* . . . . . . . x ? . . . . . . . ?* x* . . . . . . . . A . . . . . ?* ?* . . . . . A . . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . ? . 6 0 4 1 2 5 5 3
Genist a t inct oria
Cy t isus scoparius
. . . . . . . . . . . . . . . . . . A . . . . . . . . . . 1
C. nigricans
Lupinus poly phy llus
Species Dryodurgades reticulatus (H .-S.) Dryodurgades antoniae (Mel.) Handianus ignoscus (Mel.) Batracomorphus allionii (Turt.) Euscelis ohausi W.Wg. Gargara genistae (F.) Stictocoris picturatus (C. Shlb.) M egophthalmus scanicus (Fall.) A phrodes bicincta (Schrk.) A phrodes diminuta Rib. Euscelis lineolatus Br. Euscelis incisus (Kbm .) A naceratagallia ribauti (Oss.) A gallia brachyptera (Boh.) A naceratagallia venosa (Geoffr.) Empoasca decipiens Paoli Empoasca pteridis (Dhlb.) Philaenus spumarius (L.) Cercopis vulnerata Rossi A phrophora alni (Fall.) Stictocephala bisonia K. & Y. Lepyronia coleoptrata (L.) Utecha trivia (Germ .) M egophthalmus scabripennis Ed w . A naceratagallia frisia (W.Wg.) Eupteryx atropunctata (Goeze) A thysanus quadrum Boh. Handianus procerus (H .-S.) Euscelis venosus (Kbm .) Total
Chamaespart ium sagit t ale
Table 15. Auchenorrhyncha species utilizing Fabaceae in Germany. Explanations see Table 5.
P ad MonoP eg phages P eg P eg P eg P eg P eg P eg P eg OligoP eg phages P eg P ny/ eg P ad P eg P eg M? ad PolyM? ad ? phages X eg X ny Vertical X eg m igrants P eg X eg P ad P eg P ad UnconP eg firm ed P eg P eg P eg
* = 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 monophages and 1st degree oligophages are identical. Introduced ornamentals include ashleaf, sugar, silver and Japanese maple (Acer negundo, A. saccharum, A. saccharinum, A. palmatum) 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 wellstudied. Sycamore (Acer pseudoplatanus) has its strongholds in rather cool, hilly to mountainous 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 forests, usually below 1000 m a.s.l. Aceraceae provide important leafhopper food plants, altogether harbouring 17 confirmed plus 10 unconfirmed species (Table 16). Acer pseudoplatanus is of particular importance and holds the only 1st degree monophages, notably Ossiannilssonola callosa (Then), Edwardsiana nigriloba (Edw.), Eurhadina loewii (Then) and perhaps Acericerus heydenii (Kbm.). Along with Zyginella pulchra P. Löw, which also breeds on other maples, 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 lethierryi (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 wahlbergi (Boh.), Fagocyba cruenta (H.-S.) and Alnetoidia alneti (Dhlb.), as well as the planthopper 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 Japananus 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 feeding 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
?
A. palmat um
A. campest re
A. monspessulanum ? x
? x ? ?
? ? ? ? ?* ? ? ? ? 13
7
? ?* ? ?* ? ? 12
2
Overw intering stage
A
? x X X X X x X x x x ? A A ?
Substrate
x x x X X
? x x ? ? ? x X ? X X x
D iet breadth
X X X X X x x ?
A. plat anoides
Species Ossiannilssonola callosa (Then) Edwardsiana nigriloba (Ed w .) Eurhadina loewii (Then) A cericerus heydenii (Kbm .) Z yginella pulchra P. Löw A cericerus ribauti N ick. & Rem . A cericerus vittifrons (Kbm .) Japananus hyalinus (Osb.) Edwardsiana lethierryi (Ed w .) Issus coleoptratus (F.) A lebra wahlbergi (Boh.) Empoasca vitis (Göthe) Fagocyba cruenta (H .-S.) A lnetoidia alneti (Dhlb.) A guriahana stellulata (Burm .) Tachycixius pilosus (Ol.) A llygus mixtus (F.) A phrophora alni (Fall.) Ledra aurita (L.) Empoasca decipiens Paoli Edwardsiana flavescens (F.) Edwardsiana ampliata (W.Wg.) Edwardsiana frustrator (Ed w .) Ribautiana tenerrima (H .-S.) A rboridia erecta (Rib.) Fieberiella septentrionalis W.Wg. Platymetopius major (Kbm .) Total
Acer pseudoplat anus
Table 16. Auchenorrhyncha species utilizing Aceraceae in Germany. Explanations see Table 5.
m1 m1 m1 m 1? m2 m2 m2 m2 o2 po po po po po po po po po po po o2 o2? po o2? o2? po po
M M M P M P P P M P M P M M M P P X P M? M M M M M P P
eg eg eg ad ad ad ad eg eg ny eg ad eg eg eg ny eg eg ny ad eg eg eg eg ad eg eg
Monophages
Oligo-/ Polyphages
Vertical m igrants
Unconfirm ed
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
?
X
x
x A* A* ?* ? ?* 0
2
0
1
0
1
1
?* 0
1
1
Overw intering stage
x
Substrate
x
D iet breadth
H. mant egaz z ianum
Heracleum sphondy lium
Past inaca sat iv a
Angelica sy lv est ris
Falcaria v ulgaris
Daucus carot a
Aegopodium podagraria
X
Ant hriscus sy lv est ris
?
Ch. aromat icum
Ch. hirsut um
Species Eupteryx heydenii (Kbm .) Eupteryx atropunctata (Goeze) Eupteryx aurata (L.) Philaenus spumarius (L.) Hyalesthes obsoletus Sign. Cercopis sanguinolenta (Scop.) Emelyanoviana mollicula (Boh.) Cercopis vulnerata Rossi Lepyronia coleoptrata (L.) A phrophora alni (Fall.) Total
Chaerophy llum aureum
Table 17. Auchenorrhyncha species utilizing Apiaceae in Germany. Explanations see Table 5.
m 2? po po po po po po po po po
M M M X P X M X X X
eg eg eg eg ny ny eg ny eg eg
Monophage Polyphages Vertical m igrants Uncertain
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 Lamiaceae and Asteraceae, or to xylem-feeding cercopids (Table 17). Thus, only Philaenus spumarius (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 important 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 attacking 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 grassland), 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 Auchenorrhyncha 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 attacked, 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 feeding on Ocimum and Satureja in gardens by oligo- and polyphages is likely to occur, provided 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 exploit 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 latter is probably host in the second generation only. Among all plant families, the Lamiaceae are outstanding in showing the highest proportion of mesophyll feeders, whereas exploitation of xylem and phloem is rare. Similarly high values are only found in some further groups of dicotyledonous herbs, such as the Ranunculaceae, Geraniaceae, Malvaceae, Clusiaceae, Scrophulariaceae and Dipsacaceae, although these groups are infested only by very few species. Stewart (1988) has compiled the British feeding data of the genus Eupteryx and suggested 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. within the Salicaceae or the Poaceae, the utilization of Lamiaceae appears to be well-balanced, mostly with no more than three leafhopper species per plant species. Even on much-favoured hosts, like wood sage (Teucrium scorodonia), ground-ivy (Glechoma hederacea), hedge woundwort (Stachys sylvatica) or horse mint (Mentha longifolia), there is either only one dominating leafhopper species, or the leafhopper species belong to taxonomically 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 relevant 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 families, 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 species 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 Europe, 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 (Artemisia 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 Artemisia 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 Asteraceae 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 feeders of mesophyll and xylem are slightly overrepresented (see section 5.4). Specialists among Auchenorrhyncha appear to prefer members of the subfamily Asteroideae, with Laburrus pellax (Horv.) living on goldilocks (Aster linosyris), both Eupteryx tenella (Fall.) and Ophiola transversa (Fall.) on yarrow (Achillea millefolium), both Macropsidius sahlbergi (Fl.) and Laburrus impictifrons (Boh.) on field southernwood (Artemisia 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 probably 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
Lamium album
L. galeobdolon
L. maculat um
Ballot a nigra
St achy s palust ris
S. sy lv at ica
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ?* . ?* . . . . . . . . . . . . . . x* . . . . . . . . . . . . . . . . . . . . . . ?* . . . . . . . 0 1 3 0
. . . . . . . . . . X . . . . . . . . . . ? . . . . . . . . . . . . . . . . . 1 1
. . . . . . . ? X . . . . . . . . . . . . . . . . . . . . . . . ? . . . . . . 1 2
X . . . . . . . . . . . . . . . . . . . . ? . . ? . . . . . . x X . . . . . . . . . . x . . . . . . . . . . ? . . ? . . ?* . ?* . . ? X ?* . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . . . ? . . . ? . . 2 2 1 7 1 3
. . . . . . . . . . x . . . . . . . . . . x . . . . . . . . . . . . . . . . . 2 0
. . . . . . . ? X ? X . . . . . . . ? . . . . . . . . . . . . . . . . . . . . 2 3
Leonurus cardiaca
Galeopsis angust ifolia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . x . . x . . . . . . . . . . X . . . . . ? . ? x ?. . . . . . x . . . . . . . . . . . . . . . . . . . ?* x . . x . . . . X . . . . . . . . . ?* . . . . . . . . . . ? . . . . . . . . . . x . . . . . . . ?* ?* . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . . . 0 1 0 3 0 2 3 12 0 1 1 2 3 1 * = after Halkka et al. (1977), Remane (pers. comm.), Stewart (1988),
Prunella v ulgaris
Glechoma hederacea
Nepet a cat aria
Marrubium v ulgare
T. scorodonia
Teucrium mont anum
Ajuga rept ans
Species Eupteryx immaculatifrons (Kbm .) Eupteryx lelievrei (Leth.) Eupteryx origani Zachv. Eupteryx collina (Fl.) Chlorita pusilla Mats. Chlorita dumosa (Rib.) Goniagnathus brevis (H .-S.) A gallia consobrina Curt. Eupteryx stachydearum (H ard y) Eupteryx curtisii (Fl.) Eupteryx florida Rib. Eupteryx decemnotata R. Eupteryx salviae Arz. & Vid . Eupteryx thoulessi Ed w . Eupteryx melissae Curt. Eupteryx vittata (L.) Eupteryx notata Curt. Erythria aureola (Fall.) Evacanthus acuminatus (F.) Evacanthus interruptus (L.) Emelyanoviana mollicula (Boh.) Eupteryx atropunctata (Goeze) Eupteryx aurata (L.) Hyalesthes obsoletus Sign. 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 Total unconfirm ed
O cimum basilicum
Table 18. Auchenorrhyncha species utilizing Lamiaceae in Germany. Explanations see Table 5.
Vidano & Arzone (1978)
Asteraceae
281
Ly copus europaeus
Ment ha aquat ica
M. longifolia
M. x piperit a
D iet breadth
. . X . . . . . . . ? ? . . . . . . . . ? X . . . . . . . . . . . . . . . . . 2 2
. . . . . x x . . . . . . . . . ?* ? . . . . . . . . . . ? ? . . . . . . . . . 2 4
. . . . . ? x . . . . x . . . . ?* ? . . x . . . . . . . ? ? ? ? . ? . ? . . . 3 9
. . . . X x ? . . . . . . . . . . . . . ?* . . . . . . . . . . . . ? . . . . . 2 3
. . . . . . . . . . . x . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0
. . . . . . . . . . . . . x . . . . . . . . . . . . ?* . . . . . . . . . . . . 1 1
. . . . . . . . ? . . . . X . . . . . . ?* x ?* . . . . ? . . . . . . . . . . . 2 4
. . . X . . . . . . x . . . . . . . . . ? . x . . . . ? . . . . . . . . . . . 3 2
. . . . . . . . . . . . . ?* ?* . . . . . . x . . . . . . . . . . . . . . . . . 1 2
m1 m1 m1 m 1? m1 m2 m 2? o1? o1 o1? o1 o1 o1? o1 o2 o2 o2 o2 po po po po po po po po po po o2? o2? o2? o2? o1? o2? po po? m 1? po? po
Overw intering stage
Th. v ulgaris
. . . . . . . . ? ? X . . . . . . . . . x ? . . . . . . . . . . . . . . . . . 2 3
Substrate
Th. serpy llum
?* . . . . . ? x . . . . . . . . . . . . ?* . . . . 2 5
Th. pulegioides
X . . . . . x x x . . . . . . . . . . . . . . . . 6 2
. . . ?* . . . . . . X ? .
Thy mus praecox
. . . . . . . . . . . . . . . . . . . . ? x . . . . . . . . . . . . . . . . . 1 1
O riganum v ulgare
. . . . . . . . . . . . . . . . . . . . X . . . x* . . . . . . . . . . . . . . 2 0
Clinopodium v ulgare
. . . ?* . . . . . . X X ? ?*
Melissa officinalis
S. v ert icillat a
x . . . . . . . . . . . . . . . . . 1 0
S. prat ensis
. . . . . . . . . . . . . . . . . . . .
S. officinalis
. X . . . . . . . . . . . . . . . . . ? ? x . . . . . . . . . . . . . . . . . 2 2
Salv ia nemorosa
Bet onica officinalis
Table 18. (continued):
M eg M eg M eg M eg M eg M eg P ad P ad M eg M eg M eg M eg M eg M eg M eg M eg M eg M eg X eg X eg M eg M eg M eg P ny X ny X ny X eg X eg P ad P ad P ad P eg P eg P eg M? ad M ad M eg M ad ? P eg
1st d egree m onophages 2nd d egree m onophages
1st d egree oligophages
2nd d egree oligophages
Polyphages
Vertical m igrants
Unconfirm ed
282
Utilization of plant resources
Solidago spp.
Ast er linosy ris
Helichry sum arenarium
Heliant hus annuus
Dahlia pinnat a
Achillea millefolium
Chry sant hemum v ulgare
Art emisia abrot anum
A. v ulgaris
A. absint hium
A. marit ima
A. campest ris
Pet asit es spp.
Senecio spp. spp.
Species Laburrus pellax (H orv.) Eupteryx tenella (Fall.) Ophiola transversa (Fall.) M acropsidius sahlbergi (Fl.) Laburrus impictifrons (Boh.) Eupteryx adspersa (H .-S.) A ustroasca vittata (Leth.) Eupteryx artemisiae (Kbm .) Euscelis venosus (Kbm .) N eoaliturus fenestratus (H .-S.) Chlorita paolii (Oss.) Euscelis distinguendus (Kbm .) A gallia brachyptera (Boh.) Eupteryx notata Curt. Eupteryx vittata (L.) Lepyronia coleoptrata (L.) Philaenus spumarius (L.) A phrodes makarovi Zachv. Evacanthus interruptus (L.) Erythria manderstjernii (Kbm .) Empoasca decipiens Paoli Empoasca pteridis (Dhlb.) Empoasca vitis (Göthe) Eupteryx atropunctata (Goeze) Eupteryx aurata (L.) Hyalesthes obsoletus Sign. Cercopis sanguinolenta (Scop.) Cercopis vulnerata Rossi Stictocephala bisonia K. & Y. Centrotus cornutus (L.) A phrophora alni (Fall.) Hephathus nanus (H .-S.) Eupteryx heydenii (Kbm .) Fieberiella septentrionalis W.Wg. A thysanus quadrum Boh. Total substantiated Total unconfirm ed
Eupat orium cannabinum
Table 19. Auchenorrhyncha species utilizing Asteraceae in Germany. Explanations see Table 5.
. . . . . . . . . . . . . . . x* . . x ? . . . . ?* . . . . . . . . . . 2 2
. . . . . . . . . . . . . . . x* x* ?* . . ? ?* . . . . . A A . . . . ? . 4 4
X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0
. . . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . . . 0 1
. . . . . . . . . . . . . . . . . . . . . . ? ?* ? . . . . . . . . . . 0 3
. . . . . . . . . . . . . . . . . . . . . . . . x . . . . . . . . . . 1 0
. X x x* . . . . . . . X . ?* . . . x . . . . . . . . . . . . . . . . . . 4 1
. . . . . . . . . . . . . . . ? x . . . ? . . ?* . x* . ? . . . . . . . 2 4
. . . . . . . x . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0
. . . . . . . . . . x . . . . x* x . . . . . . . . x* . ? ?* . . . . . . 4 2
. . . . . X X . . . x . . . . . . . . . ?* . . . . . . . . . . . . . . 3 1
. . . . . . x X . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 0
. . . X X . . . . . X . . . . . . . . . . . . . . . . . . . . . . . . 3 0
. . . . . . . . . . . . . . . . ? . x . . . ? ?* X . . . . . . . ?* . . 2 4
. . . . . . . . . . . . . . . . ? . x ? . . . . X x . . . . . . . . . 3 2
* = after Biedermann (1998a), Bopp (1997), Kuntze (1937), Nickel (1979), Remane & Fröhlich (1994b), Stewart
Asteraceae
283
Carduus spp.
Cirsium acaule
C. arv ense
C. oleraceum
O nopordum acant hium
Lapsana communis
Hy pochaeris spp.
Leont odon hispidus
Picris hieracioides
Taraxacum spp.
Hieracium pilosella
Hieracium spp.
Crepis spp.
. . . . . . . . . . . . . . . . ? . x . . . . ?* X . . . . . . . . . . 2 2
. . . . . . . . . . . . . . . . . x . . . . . . . . . A . A . . . . . 3 0
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . 0 1
. . . . . . . . . . . . . . . ?* X ? . . . ? . ? . . . . . ? . . . . . 1 5
. . . . . . . . . . . . . . ? . . . x . . . . . x . . . . . . . . . . 2 1
. . . . . . . . . . . . . . . . . . . . . . . . . . x* . . . . . . . . 1 0
. . . . . . . . . . . . . . . . . . . . . . . . x . . . . . . . . . . 1 0
. . . . . . . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . 0 1
. . . . . . . . . ? . . . x . . . . . . . . . . . . . . . . . . . . . 1 1
. . . . . . . . . . . ? . . . . . . . . . . . . . . . . . . . . . . . 0 1
. . . . . . . . . ? . ? x . . ?* ? X . . . ? . . . . . . . . . . . . . 2 5
. . . . . . . . . . . . . X . . . . . . . . . . . . . . . . . . . . . 1 0
. . . . . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . 0 1
. . . . . . . . . . . . . . x . x* . . ?* . . . . . . . . . . . . . . . 2 1
(1988), Strübing (pers. comm.), Vidano (1959a), Vidano & Arzone (1978)
Overw intering stage
Arct ium spp.
. . . . . . . . ? . . . . . . . . . . . . . . . . . . . . . . . . . . 0 1
Substrate
C. v ulgaris
. . . . . . . . ? . . . . . . . . . . . . . . . . . . . . . . . . . . 0 1
D iet breadth
Carlina acaulis
Table 19. (continued):
m1 m1 m1 m 1? m1 m 1? m2 m2 m 2? m 2? o1 o1? o2? o2 o2 po po po po po? po po po po po po po po po po po m 1? m 2? po m 1?
P M P P P M M M P P M P P M M X X P X M M? M? P M M P X X P P X P M P P
eg eg eg eg eg eg eg eg eg ad eg eg eg eg eg eg eg eg eg ad ad ad ? ad eg eg ny ny ny eg ny eg eg eg eg eg
1st d egree m onophages
2nd d egree m onophages
Oligophages
Polyphages
Vertical m igrants
Unconfirm ed
284
Utilization of plant resources
other Asteraceae. Among oligophages, Chlorita paolii (Oss.) is confined to both wormwood 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-growing 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 mountain 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 entomologists collecting Auchenorrhyncha. Thus, the majority of data refers to only a few widespread 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.), Florodelphax leptosoma (Fl.), Macrosteles horvathi (W.Wg.) and probably Conomelus lorifer Rib. are 2nd degree monophagous on rushes, whereas Cicadella viridis (L.) and Conosanus obsoletus (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 velvetgrass (Holcus spp.) immediately after hatching, where they usually build up a summer 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
X X
X X
? x*
X X
? ?
X
?
?
? ?* ?*
?
? ?
?* ?*
?* ? ?* ?
0
?* ?* ? ? 2
3
3
5
1
?* 4
0
D iet breadth
x
Luz ula spp.
x
J. effusus
X
m2 m 2? m2 m2 x* o2 x o2 o2 po x po o2 m 2? po po? o1? po ?* po po po? po m 1? o1? 3
Overw intering stage
x
?
Substrate
? X
X x x
J. acut iflorus
x
J. art iculat us
X
J. subnodulosus
?
J. conglomerat us
?*
J. inflexus
J. gerardii
Species Conomelus anceps (Germ .) Conomelus lorifer Rib. Florodelphax leptosoma (Fl.) M acrosteles horvathi (W.Wg.) Criomorphus albomarginatus Curt. M acustus grisescens (Zett.) Conosanus obsoletus (Kbm .) Cicadella viridis (L.) Forcipata forcipata (Fl.) M egamelodes quadrimaculatus (Sign.) M egamelodes lequesnei W.Wg. Javesella pellucida (F.) Javesella obscurella (Boh.) Javesella salina (H pt.) Lepyronia coleoptrata (L.) N eophilaenus lineatus (L.) M acrosteles laevis (Rib.) M acrosteles ossiannilssoni Ld b. M acrosteles sexnotatus (Fall.) M acrosteles sordidipennis (Stål) Limotettix striola (Fall.) Total
Juncus bufonius
Table 20. Auchenorrhyncha species utilizing Juncaceae in Germany. Explanations see Table 5.
P P P P P P P X M P P P P P X X P P P P P
eg eg ny eg ny. ny eg eg eg ad ad ny ny ny eg eg eg eg eg eg eg
Monophages
Oligophages Polyphages
Unconfirm ed
* = after Halkka et al. (1977), Novotný (1995), Remane (pers. comm.), Witsack (1985)
versity. Cyperaceae biomass and dominance are particularly high in open, moist to flooded sites, notably bogs, fens and fen woods, marshes and lake shores. They are also common, 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 nonflowering 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 observation. 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 separately 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 geographic 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 monophagous, 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. Specific 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 cottongrasses 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 cottongrass (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 populations of Limotettix striola (Fall.), and has been reported to be utilized by Sorhoanus xanthoneurus (Fieb.) in the Czech Republic (Novotný 1995). Alpine deergrass (Trichophorum 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 superspecies, 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 Schoenoplectus 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 preferably 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.), respectively. 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 (= Holoschoenus), 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 harbour 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 additionally lives on pill sedge (C. pilulifera). Dwarf sedge is the exclusive host of the recently 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
C. remot a
C. elongat a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? .
C. paniculat a
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . x? X . . . . . . . X . . . . . . X . . . . . . . X . . . . . . . X . . . . . . . X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . ? x . . . . . . .
C. dist icha
. . . . . . . . . . . . . . . . . x . . . . . . . . . . . . . . . . . . . . . . . .
C. arenaria
Carex muricat a
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
C. briz oides
Rhy nchospora alba
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
C. v ulpina
Sch. nigricans
. . . . . . . . . . . . . . . . . . . . . . . . . . . . x X X ? x X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Schoenus ferrugineus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
E. uniglumis
Eleocharis palust ris
Scirpus sy lv at icus
Sch. t abernaemont ani
. . . . . . . . . . . . . . . . ?* . . . . X x . . . . . X . . . . . . X . . . . . . X . . . . . . x . . . . . . . . . X . . . . . . X ? . . . . . X x. x . . . . . . . X . . . . . . X . . . . . . . X . . . . . . . . . . . . . . . . ? . ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
S. radicans
Bolboschoenus marit imus
Schoenoplect us lacust ris
E. angust ifolium
X X X X . ?* . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
E. lat ifolium
Eriophorum v aginat um
Species N othodelphax distincta (Fl.) Ommatidiotus dissimilis (Fall.) Sorhoanus xanthoneurus (Fieb.) Kelisia vittipennis (J. Shlb.) M acrosteles fieberi (Ed w .) Cosmotettix panzeri (Fl.) Delphacodes capnodes (Scott) Javesella simillima (Lnv.) Coryphaelus gyllenhalii (Fall.) Parapotes reticulatus (H orv.) Calligypona reyi (Fieb.) Paramesus major H pt. Paramesus obtusifrons (Stål) Cicadula albingensis W.Wg. Euconomelus lepidus (Boh.) M acrosteles lividus (Ed w .) Limotettix striola (Fall.) Limotettix atricapillus (Boh.) Kelisia monoceros Rib. Cicadula rubroflava Lnv. Kelisia praecox H pt. Kelisia sabulicola W.Wg. Florodelphax paryphasma (Fl.) Stenocranus longipennis (Curt.) 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 .) Kelisia minima Rib.
Trichophorum cespit osum
Table 21. Auchenorrhyncha species utilizing Cyperaceae in Germany. Explanations see Table 5.
Cyperaceae
289
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . ? . . . ?* . . . . . . . . . . . . . . . . ? ? . ?* . . . . . . . . . . . ? . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . ?* . . . . . . ? . . . . . . . . . . ? . . . . ? . . . . . . . . . . . .
. . . . . . ?* . . . . . . ?* . . . . . . . . . . . . . ?* . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
m1 m 1? m 1? m2 m1 m 1? o1 o1? m1 m 1? m2 m1 m1 o1? m 2? m 2? o1? m 1? m2 m1 m2 m1 m1 m1 m 2? m 2? m 2? m 2? m2 m 2? m1 m1 m1 m2 m1 m1 m1 m 2? m1 m 2? m2 m 1?
Overw intering stage
D iet breadth
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Substrate
C. hirt a
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X ? . x
C. riparia
. . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . . . . . . . . . . . . .
C. acut iformis
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
C. v esicaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X ? . .
C. rost rat a
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
C. pseudocy perus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
C. flav a
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X . X . X . X ? . . . . . . . .
C. dist ans
C. humilis
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
C. sy lv at ica
C. cary ophy llea
. . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . X X x X . . . . . . . .
C. semperv irens
C. flacca
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
C. ferruginea
C. mont ana
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . x . . . . . . . .
C. panicea
C. pilulifera
. . . . . . . . . . . . . . . . . . . . . . . . ? . . ? x ? . . . . . . . . . . . .
C. limosa
C. elat a
. . . . . . ?* . . . . . . ? . . . . . . . . . . ? . . X X X . . . . . . . . . . . .
C. pilosa
C. acut a
. . . . . . . . . . . . . ? . . . . . . . . . . X X x ? ?* ?* . . . . . . . . . ? . .
C. alba
C. nigra
Table 21. (continued):
P ny P eg P eg P eg P eg P eg P ad P ny P eg P eg P ny P eg P eg P eg P eg P eg P eg P eg P eg P eg N arrow P ad ly stenoP ad phagous P ny P eg P ny P eg X eg P eg P eg P eg P eg P eg M eg? P eg P eg P eg P eg M eg? P eg P eg P eg P eg
290
Utilization of plant resources
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . . . . x . . . . . X . . . . ?* . X x . . x x ? X x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . x . . . . . . . . . ?* . . x . . . . . . x . . . . . . . . . . ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
C. elongat a
C. remot a
C. paniculat a
C. dist icha
C. arenaria
C. briz oides
C. v ulpina
Carex muricat a
Sch. nigricans
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ?* . . . . . . . . . . . . . . ?* . . . . . . . . ? x . . . . . . ?* ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . .
E. uniglumis
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Rhy nchospora alba
Schoenus ferrugineus
Eleocharis palust ris
. . . . . . . . . . . . . . . . . . . . ? . . . . . . . . ? . . ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ?* . . . . . . . . ?* . . ?* . x . . . . . . . . . . . . . . . . . . . . . . . ? . .
S. radicans
. . . . . . . . . . . . . . . . . . . . . . . . . . x . . . . . . . . . . . . .
Scirpus sy lv at icus
Schoenoplect us lacust ris
. . . . . . . . . . . . . ? . . . . . . ? . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . ?* . . ? . . ? . . . ? . . . . . . . . . . . . . . . . . . .
Bolboschoenus marit imus
Trichophorum cespit osum
. . . . . . . . . . . . . . . . . . . . . . . . . . x . . . . . . . . . . . . .
Sch. t abernaemont ani
E. lat ifolium
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) M egamelus notula (Germ .) N otus flavipennis (Zett.) Cicadula quadrinotata (F.) 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. M acustus grisescens (Zett.) Cicadella viridis (L.) N eophilaenus lineatus (L.) Forcipata forcipata (Fl.) Speudotettix subfusculus (Fall.) Lepyronia coleoptrata (L.) Philaenus spumarius (L.) M acrosteles sexnotatus (Fall.) 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.)
Eriophorum v aginat um E. angust ifolium
Table 21. (continued):
. . . . . . . . . . . . . . . . . . . . . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cyperaceae
291
C. dist ans
C. flav a
C. pseudocy perus
C. rost rat a
C. v esicaria
C. acut iformis
C. riparia
C. hirt a
D iet breadth
. . . . . . . . . . . . . . . . . . . . . . . . . . . . x . . . . . . . . . . .
. . . . . . . . . . . . . . ? . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . x . . . . . . . ? . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . ? . ? . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
X . . . . . . . . . . . . . . . . . ? . . . . . . . . . . . . . . . . . . . . .
. . ? . . . x . ? ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . X ?* X . . ?* . ? ?* . . . . . . . ? x ?* . . . . . . . . . . . . .
. . . . . . . . x X x . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. X X ? . . X ? X X X . ? . . . . . . ? . . . . x ? . ? . . . . . . . . . . . .
. . X X . . . . ? ? ? . ?* . . . . . . . . . . . . x* . . . . . . . . . . . . . .
. . . . X x . . . . X . . . . . . . . . . . . . . x* . . . . . . . . . . . . . .
m1 m 1? m2 m 2? m1 m 1? m 2? m2 m2 o1 m2 m 1? m 2? m 2? m 1? m 2? m 2? m 2? m 2? m 2? m 2? m 2? m 1? po? o2 po po po? po po po po m 2? m 2? po? po? po? o1? po po
Overw intering stage
C. sy lv at ica
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Substrate
C. semperv irens
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ?* . ? . . . . . . . . . . . . . . . . . . . . . . . . . . ? . x . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
C. ferruginea
C. flacca
. . . . . . . . . . . . . . . . . ? . . . . . . . . . ? . . . . . . . . . . . .
C. panicea
C. mont ana
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
C. limosa
C. pilulifera
. . . . . . ? x x x ? ? . . . . . . . . . . . . . x x* . . . . . . . . . . . . .
C. pilosa
C. elat a
. . ? ? . . X ? X X ? . . . . . . . . ? . . . . ? ? . . . . . . . . . . . . . .
C. alba
C. acut a
. . . . . . x . x x X . . ? . . . . ? ?* . . . . ? x ?* . . ?* . . . . . . . . . .
C. humilis
C. nigra
C. cary ophy llea
Table 21. (continued):
P eg P eg N arrow P eg ly stenoP eg phagous P eg P eg? P eg P ad Broad ly P eg stenoM eg phagous P eg P eg? P eg P eg P ny P ny on unP ad know n M eg? CyperaCarex M eg ceae P eg P eg P eg P eg P eg P ny X eg X eg Oligo-/ M eg polyP ny phagous X eg X eg P eg P ny P ad P ny UnconP ny Unconfirm ed P ny firmed P eg P eg P eg
292
Utilization of plant resources
. . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . . . ? ? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5 1 1 4 1 2 1 0 4 2 0 0 2 0 2 2 4 2 2 1 5 3 0 1 4 1 1 1 0 1 1 7 9 3 3 5 6 5 1 1 8 3 1 1 2 1 3
. . . . 3 4 7
C. elongat a
C. briz oides C. arenaria C. dist icha C. paniculat a C. remot a
Schoenus ferrugineus Sch. nigricans Rhy nchospora alba Carex muricat a C. v ulpina
E. uniglumis
Bolboschoenus marit imus Scirpus sy lv at icus S. radicans Eleocharis palust ris
Species M acrosteles oshanini Razv. M acrosteles viridigriseus (Ed w .) Deltocephalus maculiceps Boh. Sorhoanus schmidti (W.Wg.) Total substantiated Total unconfirm ed Substantiated plus unconfirm ed
Eriophorum v aginat um E. angust ifolium E. lat ifolium Trichophorum cespit osum Schoenoplect us lacust ris Sch. t abernaemont ani
Table 21. (continued):
. . . . . . . . . . . . . . . . . . . . 1 4 6 1 0 1 2 1 0 2 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), distant 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 European Cyperaceae, with 8 definitely and 11 possibly associated Auchenorrhyncha species. 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 Wagneriala incisa (Then), while C. limosa has been found to be infested by Nothodelphax albocarinata (Stål) in two cases. C. distans has in a few occasions been found to harbour populations of the rare Kelisia minima Rib. For C. rostrata only three rather euryphagous sedgefeeding 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 frontalis (H.-S.) also exploits the following sedge. C. riparia is the only verified host of Anakelisia fasciata (Kbm.), although a few specimens have been taken on C. acuta and
Cyperaceae
293
C. alba
C. pilosa
C. limosa
C. panicea
C. ferruginea
C. semperv irens
C. sy lv at ica
C. dist ans
C. flav a
C. pseudocy perus
. . . . 5 0 5
. . . . 0 1 1
. . . . 1 0 1
. . . . 0 1 1
. . . . 2 2 4
. . . . 0 1 1
. . . . 0 2 2
. . . . 1 2 3
. . . . 1 1 2
. . . . 1 1 2
. . . . . . . . . . . . . . . . . . . . 1 3 3 7 3 3 12 0 10 7 4 15 3 17 10
. ? . o2? . m 1? . m 1? 4 0 4
Overw intering stage
C. humilis
. . . . 0 1 1
Substrate
C. cary ophy llea
. . . . 4 3 7
D iet breadth
C. flacca
. . . . 0 2 2
C. hirt a
C. mont ana
. . . . 1 0 1
C. riparia
C. pilulifera
. . . . 6 6 12
C. acut iformis
C. elat a
. . . . 6 10 16
C. v esicaria
C. acut a
. . . . 8 11 19
C. rost rat a
C. nigra
Table 21. (continued):
P P P P
eg Unconeg firm ed Unconeg firmed eg
Nickel (1979), Novotný (1995), Schiemenz (1987), Vidano (1965), Wagner & Franz (1961), Witsack (1985)
C. acutiformis (Lauterer 1984). In Greece C. riparia is reported to be infested by Kelisia confusa Lnv. (Drosopoulos et al. 1983). (iii) Carex species with substantiated 2nd degree monophages Further sedges are frequently attacked, but without showing exclusive associations. False fox sedge (C. vulpina ssp. otrubae) and prickly sedge (C. muricata) are the main hosts of Kelisia monoceros Rib. in the lowlands, but not in the Alps, where perhaps evergreen sedge (C. sempervirens) is preferred. Kelisia guttulifera (Kbm.) usually lives on remote and wood Sedge (C. remota and C. sylvatica), but on single occasions, reproducing populations have also been found on elongated sedge (C. elongata), false fox sedge (C. vulpina ssp. otrubae), greater tussock sedge (C. paniculata) and distant sedge (C. distans). Cicadula flori (J. Shlb.), Metalimnus formosus (Boh.) and Cosmotettix costalis (Fall.) all favour slender tufted sedge (C. acuta) in many lowland floodplain sites, along with some less specific species, such as Megamelus notula (Germ.), Stroggylocephalus agrestis (Fall.), Notus flavipennis (Zett.) and Cicadula quadrinotata (F.). On acidic substrates Anakelisia perspicillata (Boh.) lives on pill sedge (C. pilulifera), but in more basic sites it is clearly associated with glaucous sedge (C. flacca). Other infested, but little studied sedges include cyperus and bladder sedge (C. pseudocyperus, C. vesicaria). (iv) Carex species probably with 2nd degree monophages There is also some field evidence, that spring sedge (C. caryophyllea), white-flowered sedge (C. alba), rusty sedge (C. ferruginea), ciliated sedge (C. pilosa) and evergreen sedge (C. sempervirens) may be attacked, but more samples are required for confirmation. Furthermore, it should be noted, that in some Auchenorrhyncha species, the host relationships of which are little known, an association with unidentified species of Carex is highly likely. This is true in Kelisia confusa Lnv., K. ribauti W.Wg., Stroggylocephalus livens (Zett.), Forcipata citrinella (Zett.), Cicadula quinquenotata (Boh.), C. ornata (Mel.),
294
Utilization of plant resources
Cosmotettix aurantiacus (For.) and perhaps Sorhoanus schmidti (W. Wg.), Macrosteles oshanini 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 Fagaceae, 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 creating 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 maintaining 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 pratensis, 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 relationships 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 Auchenorrhyncha 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 specifically. 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. Altogether, 43 Auchenorrhyncha species are confirmed feeders on Festuca, with a high proportion 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, pastures, 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 subspecies. 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., Rhytistylus proceps (Kbm.), Hardya signifer (Then), Rhopalopyx vitripennis (Fl.), Mocydiopsis longicauda Rem., Psammotettix pallidinervis (Dhlb.), Mendrausus pauxillus (Fieb.) and perhaps 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 Eupelix cuspidata (F.) and Arocephalus punctum (Fl.). Dicranotropis divergens Kbm. is apparently 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 arundinacea), 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
Calamagrostis villosa
A . stolonifera
A . capillaris
A . vinealis
A grostis canina
H. mollis
Holcus lanatus
Corynephorus canescens
D. flexuosa
Deschampsia cespitosa
Trisetum flavescens
Koeleria glauca
H. pratense
Helictotrichon pubescens
A rrhenatherum elatius
A vena spp.
Phleum spp.
A lopecurus spp.
A nthoxanthum odoratum
Phalaris arundinacea
M olinia caerulea
Phragmites australis
Cynodon dactylon
Setaria spp.
Z ea mays
Table 22. Auchenorrhyncha species utilizing Poaceae in Germany. Explanations see Table 5.
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.
.
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x
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.
Jassargus sursumflexus (Then)
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X
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X anthodelphax xantha Vilb.
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?
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.
Sorhoanus schmidti (W.Wg.)
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?
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.
Stenocranus major (Kbm .)
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X
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.
Paraliburnia adela (Fl.)
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X
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Balclutha rhenana W.Wg.
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X
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.
Erzaleus metrius (Fl.)
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X
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Ribautodelphax angulosa (Rib.)
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X
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M egadelphax sordidula (Stål)
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X
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Doratura horvathi W.Wg.
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X
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Z yginidia viaduensis (W.Wg.)
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X
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Psammotettix angulatus (Then)
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?
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A canthodelphax denticauda (Boh.)
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X
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M uellerianella brevipennis (Boh.)
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X
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Streptanus confinis (Reu t.)
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X
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Psammotettix excisus (Mats.)
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X
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Psammotettix albomarginatus W.Wg.
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?
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Recilia horvathi (Then)
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M uellerianella fairmairei (Perr.)
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X
X
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M ocydiopsis monticola Rem .
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X
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Ribautodelphax vinealis Biem .
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X
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Doliotettix lunulatus (Zett.)
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?
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297
Calamagrostis varia
C. arundinacea
C. canescens
C. epigejos
C. pseudophragmites
A mmophila arenaria
Stipa spp.
N ardus stricta
Glyceria maxima
G. fluitans
M elica uniflora
Sesleria albicans
Puccinellia distans
P. maritima
Briza media
Dactylis glomerata
Festuca altissima
F. pratensis
F. arundinacea
F. heterophylla
F. rubra
F. ovina
Lolium perenne
Poa annua
P. pratensis
P. trivialis
P. nemoralis
Bromus erectus
B. inermis
Brachypodium pinnatum
Elymus repens
Leymus arenarius
Hordeum spp.
Secale cereale
Triticum spp.
Poaceae
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298
Utilization of plant resources
A rrhenatherum elatius
Helictotrichon pubescens
H. pratense
Koeleria glauca
Trisetum flavescens
Deschampsia cespitosa
D. flexuosa
Corynephorus canescens
Holcus lanatus
H. mollis
A . vinealis
A grostis canina
A . capillaris
A . stolonifera
Calamagrostis villosa
.
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X
.
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X
.
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X anthodelphax straminea (Stål)
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?*
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X
X
x
.
Elymana kozhevnikovi (Zachv.)
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Phleum spp.
.
.
A lopecurus spp.
.
.
Phalaris arundinacea
.
.
M olinia caerulea
.
.
Phragmites australis
.
.
Cynodon dactylon
.
M ocydiopsis parvicauda Rib.
Setaria spp.
Ribautodelphax collina (Boh.)
Z ea mays
A vena spp.
A nthoxanthum odoratum
Table 22. (continued):
Paraliburnia clypealis (J. Shlb.)
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Criomorphus moestus (Boh.)
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Streptanus okaensis Zachv.
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Lebradea calamagrostidis Rem .
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.
Criomorphus borealis (J. Shlb.)
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.
X
Paluda flaveola (Boh.)
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?
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?
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X
Eurysula lurida (Fieb.)
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M irabella albifrons (Fieb.)
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Endria nebulosa (Ball)
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Doratura impudica H orv.
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A rthaldeus arenarius Rem .
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Balclutha calamagrostis Oss.
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Psammotettix poecilus (Fl.)
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Pseudodelphacodes flaviceps (Fieb.)
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Psammotettix unciger Rib.
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Gravesteiniella boldi (Scott)
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Psammotettix maritimus (Perr.)
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Doratura littoralis Ku ntze
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Praganus hofferi (Dlab.)
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Psammotettix nardeti Rem .
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Struebingianella lugubrina (Boh.)
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Chlorionidea flava P. Löw
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Z yginidia mocsaryi (H orv.)
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Z yginidia franzi (W.Wg.)
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Psammotettix putoni (Then)
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M acrosteles sordidipennis (Stål)
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A noscopus limicola (Ed w .)
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Psammotettix cephalotes (H .-S.)
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Cicadula persimilis (Ed w .)
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Stenocranus minutus (F.)
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Ribautodelphax imitans (Rib.)
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A rthaldeus striifrons (Kbm .)
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M egamelodes quadrimaculatus (Sign.)
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Dicranotropis divergens Kbm .
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A canthodelphax spinosa (Fieb.)
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299
Calamagrostis varia
C. arundinacea
C. canescens
C. epigejos
C. pseudophragmites
A mmophila arenaria
Stipa spp.
N ardus stricta
Glyceria maxima
G. fluitans
M elica uniflora
Sesleria albicans
Puccinellia distans
P. maritima
Briza media
Dactylis glomerata
Festuca altissima
F. pratensis
F. arundinacea
F. heterophylla
F. rubra
F. ovina
Lolium perenne
Poa annua
P. pratensis
P. trivialis
P. nemoralis
Bromus erectus
B. inermis
Brachypodium pinnatum
Elymus repens
Leymus arenarius
Hordeum spp.
Secale cereale
Triticum spp.
Poaceae
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X
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X
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X
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.
300
Utilization of plant resources
Z ea mays
Setaria spp.
Cynodon dactylon
Phragmites australis
M olinia caerulea
Phalaris arundinacea
A nthoxanthum odoratum
A lopecurus spp.
Phleum spp.
A vena spp.
A rrhenatherum elatius
Helictotrichon pubescens
H. pratense
Koeleria glauca
Trisetum flavescens
Deschampsia cespitosa
D. flexuosa
Corynephorus canescens
Holcus lanatus
H. mollis
A grostis canina
A . vinealis
A . capillaris
A . stolonifera
Calamagrostis villosa
Table 22. (continued):
Delphacinus mesomelas (Boh.)
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Eupelix cuspidata (F.)
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.
Rhopalopyx adumbrata (C. Shlb.)
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.
M ocydiopsis attenuata (Germ .)
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A rocephalus punctum (Fl.)
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?
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.
.
Kosswigianella exigua (Boh.)
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.
Rhytistylus proceps (Kbm .)
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Rhopalopyx vitripennis (Fl.)
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Jassidaeus lugubris (Sign.)
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.
M etropis latifrons (Kbm .)
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.
M etropis inermis W.Wg.
.
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.
N eophilaenus infumatus (H pt.)
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.
.
Doratura exilis H orv.
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.
Hardya signifer (Then)
.
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.
.
.
M ocydiopsis longicauda Rem .
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.
.
.
.
Psammotettix pallidinervis (Dhlb.)
.
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.
?
.
.
.
.
.
.
.
M endrausus pauxillus (Fieb.)
.
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.
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.
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.
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.
.
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.
.
.
.
A chorotile albosignata (Dhlb.)
.
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.
.
.
.
A rocephalus sagittarius Rib.
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.
.
Ebarrius cognatus (Fieb.)
.
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Ribautodelphax albostriata (Fieb.)
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X anthodelphax flaveola (Fl.)
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Rhopalopyx preyssleri (H .-S.)
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M ocydiopsis intermedia Rem .
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M uirodelphax aubei (Perr.)
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Jassargus flori (Fieb.)
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Criomorphus williamsi China
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Ditropsis flavipes (Sign.)
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Ribautodelphax pungens (Rib.)
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N eophilaenus albipennis (F.)
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A darrus multinotatus (Boh.)
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A darrus bellevoyei (Pu t.)
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Unkanodes excisa (Mel.)
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Eurysa lineata (Perr.)
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x
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Eurysella brunnea (Mel.)
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Eurybregma nigrolineata Scott
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x
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Stiroma affinis Fieb.
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x
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Stiroma bicarinata (H .-S.)
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x
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X
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X
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Laodelphax striatella (Fall.)
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x
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?*
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.
301
Calamagrostis varia
C. arundinacea
C. canescens
C. epigejos
C. pseudophragmites
A mmophila arenaria
Stipa spp.
N ardus stricta
Glyceria maxima
G. fluitans
M elica uniflora
Sesleria albicans
Puccinellia distans
P. maritima
Briza media
Dactylis glomerata
Festuca altissima
F. pratensis
F. arundinacea
F. heterophylla
F. rubra
F. ovina
Lolium perenne
Poa annua
P. pratensis
P. trivialis
P. nemoralis
Bromus erectus
B. inermis
Brachypodium pinnatum
Elymus repens
Leymus arenarius
Hordeum spp.
Secale cereale
Triticum spp.
Poaceae
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X
X
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?
X
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X
x
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x
X
X
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?* X
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?* X
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?* X
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X
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X
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X
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X
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X
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x
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X
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X
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X
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X
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x
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X
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x
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x
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x
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X
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x
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X
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x
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x
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x
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x
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x
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?
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x
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x
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?
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?
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?
?
x
302
Utilization of plant resources
Cynodon dactylon
Phragmites australis
M olinia caerulea
Phalaris arundinacea
A nthoxanthum odoratum
A lopecurus spp.
Phleum spp.
A vena spp.
H. pratense
Koeleria glauca
Trisetum flavescens
Deschampsia cespitosa
D. flexuosa
Corynephorus canescens
Holcus lanatus
H. mollis
A grostis canina
A . vinealis
A . capillaris
A . stolonifera
Calamagrostis villosa
.
.
.
.
x
.
.
.
.
.
?* ?*
.
.
.
.
X
.
.
x
.
.
.
.
.
Delphacodes venosus (Germ .)
.
.
.
.
x
.
.
.
.
.
.
.
.
.
.
X
.
.
.
.
x
.
.
x
.
Dicranotropis hamata (Boh.)
.
.
.
.
.
.
.
.
.
.
?
.
.
.
.
.
.
.
X
X
.
.
.
.
.
Dicranotropis montana (H orv.)
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
X
.
.
.
.
.
.
.
.
.
Scottianella dalei (Scott)
.
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.
.
.
.
.
.
.
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.
.
.
.
.
.
.
.
?*
.
.
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
.
Helictotrichon pubescens
Setaria spp.
Hyledelphax elegantula (Boh.)
A rrhenatherum elatius
Z ea mays
Table 22. (continued):
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.)
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?*
.
N eophilaenus campestris (Fall.)
.
.
.
.
.
.
.
.
.
.
?
.
.
.
.
.
.
.
.
.
.
.
?
.
.
N eophilaenus exclamationis (Thnbg.)
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?
.
.
.
.
.
.
.
.
x* ?*
N eophilaenus lineatus (L.)
.
.
.
x*
?
?
.
.
.
.
.
.
.
.
.
X
X
.
.
x
.
.
x
N eophilaenus minor (Kbm .)
.
.
.
.
.
.
.
.
.
.
.
.
.
x
.
.
.
X
.
.
.
.
.
?* X .
.
A phrophora major Uhl.
.
.
.
x*
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
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.
.
.
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
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
303
C. pseudophragmites
A mmophila arenaria
Stipa spp.
N ardus stricta
Glyceria maxima
G. fluitans
M elica uniflora
Sesleria albicans
Puccinellia distans
P. maritima
Briza media
F. heterophylla
F. rubra
F. ovina
Lolium perenne
Poa annua
P. pratensis
P. trivialis
P. nemoralis
Bromus erectus
B. inermis
Brachypodium pinnatum
Elymus repens
Leymus arenarius
Hordeum spp.
Secale cereale
Triticum spp.
.
.
.
.
.
.
.
.
.
.
.
?*
.
.
.
.
.
x
?*
.
.
.
.
x
?*
.
x
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?
.
?
x
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
X
.
.
x
.
.
.
.
.
.
.
.
.
?
.
X
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
x
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
X
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
x
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
x
x
x
.
?
.
.
.
.
.
.
x
.
.
X
.
?* X
.
x
?
X
X
X
X
.
?
?
.
X
.
?* ?* ?*
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?
?
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
x
.
.
.
.
.
.
.
.
.
.
.
.
?
?
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?
.
?
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?*
.
?
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
x*
.
.
x*
.
.
.
.
x*
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?*
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
x
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?
.
X
.
.
.
.
.
.
.
.
.
.
.
.
.
x
x
x
X
.
?
.
x
.
.
.
.
.
.
.
?*
?
.
.
.
X
x
.
.
.
.
.
.
.
.
.
?
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
X
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?*
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?* ?*
.
.
.
.
?* ?*
.
.
.
.
.
.
.
x
.
.
.
.
.
.
.
.
?
.
.
.
.
.
.
.
.
.
.
.
x
.
.
?
?
?
.
.
.
.
.
.
.
.
.
x
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?
.
.
x
.
.
?
.
?
.
?
?
.
?
.
.
.
.
x
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
x
x
x
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?*
.
.
.
.
.
.
.
.
.
?
.
.
.
.
.
.
.
.
.
.
?* X
.
?*
?
.
?
x
?
?
?
.
.
?
?
?*
?
.
.
.
x
?
?
x
X
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?*
.
.
.
.
.
.
.
.
.
?
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?
?
.
.
.
.
.
.
.
.
?* ?* ?*
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?*
.
.
.
.
?*
.
?
?
.
.
.
.
.
.
.
.
?
?*
?
.
.
.
.
.
.
.
.
.
.
.
.
x
.
.
x*
.
.
.
.
.
.
?
?
.
.
.
.
.
.
?
.
?
?
?
.
.
.
.
.
.
.
.
.
.
.
.
?
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
x
.
.
.
.
.
.
x
x
.
?
.
.
.
.
x
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
F. arundinacea
C. epigejos
.
.
F. pratensis
C. canescens
x
.
Festuca altissima
C. arundinacea
.
Dactylis glomerata
Calamagrostis varia
Poaceae
.
304
Utilization of plant resources
Z ea mays
Setaria spp.
Cynodon dactylon
Phragmites australis
M olinia caerulea
Phalaris arundinacea
A nthoxanthum odoratum
A lopecurus spp.
Phleum spp.
A vena spp.
A rrhenatherum elatius
Helictotrichon pubescens
H. pratense
Koeleria glauca
Trisetum flavescens
Deschampsia cespitosa
D. flexuosa
Corynephorus canescens
Holcus lanatus
H. mollis
A grostis canina
A . vinealis
A . capillaris
A . stolonifera
Calamagrostis villosa
Table 22. (continued):
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 .
.
.
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.
.
.
.
.
.
.
.
.
.
.
.
.
X
.
.
.
.
.
.
.
.
Psammotettix nodosus (Rib.)
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
x
?
?
.
.
.
?
.
.
Psammotettix confinis (Dhlb.)
.
.
.
.
.
.
.
?*
.
.
.
.
.
.
.
.
?
?
.
.
.
.
x
?*
. .
Errastunus ocellaris (Fall.)
.
.
.
.
.
.
.
.
.
.
?
.
.
.
.
.
.
.
X
X
.
.
.
?
Turrutus socialis (Fl.)
.
.
.
.
.
.
.
.
.
.
.
?*
.
.
.
.
.
.
.
.
.
.
.
.
.
Jassargus pseudocellaris (Fl.)
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?
.
.
x
.
.
Jassargus obtusivalvis (Kbm .)
.
.
.
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.
.
.
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.
.
.
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.
.
Jassargus alpinus (Then)
.
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.
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.
.
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.
.
?
.
.
.
.
.
.
.
X
Jassargus allobrogicus (Rib.)
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
X
.
.
.
.
.
.
.
?*
Diplocolenus bohemani (Zett.)
.
.
.
.
.
.
.
.
.
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.
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.
.
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.
.
X
V erdanus abdominalis (F.)
.
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.
.
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.
.
?*
.
.
.
.
?
.
x
?
.
.
?
.
.
V erdanus bensoni (China)
.
.
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.
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.
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.
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.
?*
.
.
?*
.
.
.
.
.
305
Calamagrostis varia
C. arundinacea
C. canescens
C. epigejos
C. pseudophragmites
A mmophila arenaria
Stipa spp.
N ardus stricta
Glyceria maxima
G. fluitans
M elica uniflora
Sesleria albicans
Puccinellia distans
P. maritima
Briza media
Dactylis glomerata
Festuca altissima
F. pratensis
F. arundinacea
F. heterophylla
F. rubra
F. ovina
Lolium perenne
Poa annua
P. pratensis
P. trivialis
P. nemoralis
Bromus erectus
B. inermis
Brachypodium pinnatum
Elymus repens
Leymus arenarius
Hordeum spp.
Secale cereale
Triticum spp.
Poaceae
.
.
.
.
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?
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.
X
x
.
.
?
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x
.
x
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.
?
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x
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?
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x
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X
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?
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x
x
.
X
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?
.
x
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?
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X
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.
X
?
X
x
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?
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?
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.
.
.
.
x
.
x
.
.
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.
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.
.
.
.
x
.
.
x
.
.
.
.
.
.
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.
.
?
.
x
.
.
.
.
.
.
x
X
.
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.
.
.
.
.
.
X
.
.
x
.
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.
?
.
X
.
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.
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x
.
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?
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.
?
.
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.
X
.
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.
.
.
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.
.
?
.
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.
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.
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.
X
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?
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X
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.
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.
.
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.
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.
X
.
.
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.
.
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.
X
.
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.
X
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.
X
.
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.
x
.
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.
?
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?
?
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?
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?
.
X
?*
?
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?
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?
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?
?
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?
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?
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?
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?
?
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?
?
?*
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?*
.
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?
?
?
?
?
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.
x
X
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.
x
.
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?
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X ?*
.
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?*
.
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.
x
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?
?
?
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.
X
?
.
X
.
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.
.
.
.
.
.
.
.
.
.
?
.
.
.
.
.
.
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.
x
.
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.
.
.
.
.
?*
.
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?
?*
.
.
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.
.
?*
.
?*
.
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.
?*
.
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.
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.
.
.
.
.
.
.
.
?
.
X
.
.
.
.
.
.
.
.
.
X
.
?*
.
.
.
.
.
306
Utilization of plant resources
Corynephorus canescens
Holcus lanatus
H. mollis
A grostis canina
A . capillaris
A . stolonifera
Calamagrostis villosa
.
x
.
.
.
?
x
.
.
X
?
?
.
?
x
?*
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?
.
.
M ocuellus collinus (Boh.)
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
?
.
.
M egadelphax haglundi (J. Shlb.)
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
Ederranus discolor (J. Shlb.)
.
.
.
?*
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
Psammotettix dubius Oss.
.
.
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.
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.
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.
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.
.
.
.
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.
.
.
.
.
.
.
.
.
Psammotettix inexpectatus Rem .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
A . vinealis
Deschampsia cespitosa
.
.
D. flexuosa
Trisetum flavescens
.
.
H. pratense
Koeleria glauca
Helictotrichon pubescens
A rrhenatherum elatius
.
.
A lopecurus spp.
.
.
Phleum spp.
.
.
Phalaris arundinacea
.
.
M olinia caerulea
.
.
Phragmites australis
.
.
Cynodon dactylon
.
Enantiocephalus cornutus (H .-S.)
Setaria spp.
A rthaldeus pascuellus (Fall.)
Z ea mays
A vena spp.
A nthoxanthum odoratum
Table 22. (continued):
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.)
.
.
.
.
.
.
.
.
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M acrosteles ossiannilssoni Ld b.
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A llygus communis (Ferr.)
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Hesium domino (Reu t.)
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Thamnotettix confinis (Zett.)
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Euscelis incisus (Kbm .)
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3
12 17
Euscelis lineolatus Br.
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Feeding substantiated
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2
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18
8
4
1
3
2
0
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0
1
3
0
13 16
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3
1
13
7
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Feeding unconfirmed
1
4
1
3
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2
17
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0
3
6
4
13
6
11 10
2
1
18 10
4
Substantiated plus unconfirmed
2
6
4
21 16
8
2
9
6
2
26
7
1
6
6
17 29
9
23 27
5
2
31 17
9
x* = after literature from Germany, ?* = after literature from neighbouring countries.
307
Poa annua
P. pratensis
P. trivialis
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2
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0
11 18
10
8
. 4
10 16 19 4
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2
1
1
1
2
2
3
4
1
1
15
2
1
10
2
14 27
Triticum spp.
Lolium perenne
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Secale cereale
F. ovina
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.
Hordeum spp.
F. rubra
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Leymus arenarius
F. heterophylla
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Elymus repens
F. arundinacea
Brachypodium pinnatum
F. pratensis
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.
B. inermis
Festuca altissima
?*
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Bromus erectus
Dactylis glomerata
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P. nemoralis
Briza media
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P. maritima
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Puccinellia distans
N ardus stricta
X
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Sesleria albicans
Stipa spp.
x
.
M elica uniflora
C. epigejos
?
.
G. fluitans
C. canescens
?
Glyceria maxima
C. arundinacea
A mmophila arenaria
Calamagrostis varia
C. pseudophragmites
Poaceae
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1
1
0
2
3
5
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
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 obsoletus (Kbm.). Various-leaved and wood fescue (F. heterophylla and F. altissima), both growing 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 scutelaris (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 northeastern Germany, is still almost unknown. Altogether, Calamagrostis is attacked by 32 Auchenorrhyncha feeders, perhaps plus further 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 usually 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 associated with C. canescens, whereas Endria nebulosa (Ball), Arthaldeus arenarius Rem. and probably 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 confined 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 Xanthodelphax xantha Vilb. utilize several Calamagrostis species. Further common, but less specific feeders on Calamagrostis include Neophilaenus lineatus (L.), Balclutha punctata (F.), Allygidius commutatus (Fieb.), Elymana sulphurella (Zett.), Mocydia crocea (H.-S.), Macustus grisescens (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 localized 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 localized endemics of Lake Constance and the lower Elbe, respectively. Only tufted and wavy hair-grass (Deschampsia cespitosa and D. flexuosa) are both widespread and common. 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 Auchenorrhyncha 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 nemoralis, 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 grasses occur. Furthermore, D. flexuosa is preferentially utilized by Javesella discolor (Boh.), Neophilaenus lineatus (L.), Anoscopus albifrons (L.), A. flavostriatus (Don.), Balclutha punctata (F.), Streptanus marginatus (Kbm.), Jassargus allobrogicus (Rib.), the acidophilic form of Psammotettix helvolus (Kbm.) and some others, and nymphs of the cercopid Haematoloma dorsatum (Ahr.) live on its roots. Published records of further species, e.g. Dicranotropis 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 studied 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 Auchenorrhyncha species have been found to feed, but as on other low-growing grasses, the
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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 pellucida (F.), Neophilaenus lineatus (L.), Deltocephalus pulicaris (Fall.), Streptanus sordidus (Zett.), Psammotettix confinis (Dhlb.) and Jassargus pseudocellaris (Fl.). Noteworthy among uncertain 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 Ribautodelphax vinealis Biem., which was undescribed until then, and which is exclusively associated 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 forests 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. Altogether, 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 Delphacodes 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).
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Less specialized feeders include Delphacodes venosus (Germ.), Macrosteles viridigriseus (Edw.) and perhaps Deltocephalus pulicaris (Fall.) and Arthaldeus pascuellus (Fall.). (v) Velvet-grass (Holcus) This genus includes only two species, notably Yorkshire fog and creeping soft-grass (H. lanatus and H. mollis). Both are widespread and common, the former is mainly found on moist meadows and pastures of lower altitudes, the latter more on acidic substrates and in open forests and on clearings. 36 Auchenorrhyncha species have been recorded altogether, 21 of which definitely breed. With 18 species alone, H. mollis is among the most-favoured plants in the study area (see Fig. 14). Mocydiopsis monticola Rem. is apparently restricted to H. mollis, in moderately shaded sites, and is absent from frequently managed grassland, where H. lanatus occurs. Muellerianella fairmairei (Perr.) lives on both species of Holcus, mainly in meadows and pastures, but needs Juncus effusus for the oviposition of winter eggs (Drosopoulos 1977). Numbers of species and individuals of further less specific Auchenorrhyncha are often high, with a marked overlap between the two grasses. For instance, Dicranotropis hamata (Boh.), Javesella pellucida (F.), Elymana sulphurella (Zett.), Athysanus argentarius Metc., Streptanus aemulans (Kbm.), Arocephalus longiceps (Kbm.), Errastunus ocellaris (Fall.) and Arthaldeus pascuellus (Fall.) all attack both species. Thus, most differences between the two guilds may rather be explained by requirements of specific microclimatic conditions than by host preferences. For instance, the more sciophilous or hygrophilous species Stiroma bicarinata (H.-S.), Hyledelphax elegantula (Boh.), Javesella forcipata (Boh.), Anoscopus albifrons (L.), Balclutha punctata (F.) and Recilia coronifer (Marsh.) have only been found on H. mollis, whereas the more heliophilous or thermophilous species Eurybregma nigrolineata Scott, Mocydia crocea (H.-S.), Verdanus abdominalis (F.) and Arthaldeus pascuellus (Fall.) clearly favour H. lanatus. The relationship of further species such as Criomorphus albomarginatus Curt., Javesella dubia (Kbm.), Zyginidia scutellaris (H.-S.), Macustus grisescens (Zett.) and Euscelis spp. has yet to be confirmed. (vi) Common reed (Phragmites australis) Phragmites australis forms large monospecific stands in temporarily or permanently flooded sites, mainly near standing or slowly running water. It also occurs in saltmarshes, straw meadows and fen woods. This plant is outstanding in harboring the maximum number of strictly monophagous Auchenorrhyncha species in central Europe, and probably also on a larger geographical scale (see Fig. 14). In Germany alone these include the leafhoppers Paralimnus phragmitis (Boh.), P. rotundiceps (Leth.), Calamotettix taeniatus (Horv.), and the planthoppers Delphax crassicornis (Panz.), D. pulchellus (Curt.), Euides basilinea (Germ.), Chloriona unicolor (H.-S.), Ch. sicula Mats., Ch. dorsata Edw., Ch. glaucescens Fieb., Ch. smaragdula (Stål), Ch. stenoptera (Fl.), Ch. vasconica Rib. and Pentastiridius leporinus (L.). The latter species perhaps feeds on other plant species at least in the nymphal stage. It has also been suggested that Ederranus discolor (J. Shlb.) lives on reed, but host reports for this species are contradictory. Additional feeders occur in more southern parts of Europe and Scandinavia (see Ossiannilsson 1978; Drosopoulos et al. 1983; Nickel et al. 2002a). Most stands are infested by several species, with up to 7 monophages that have been recorded to live syntopically. The decisive factors controlling the distribution of these species are probably water table and salinity. For instance Chloriona glaucescens Fieb. is restricted to saltmarshes (see
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also Hildebrandt 1999), whereas Ch. stenoptera (Fl.) was found to occur only on freshwater sites. Most of the remaining species on reed, however, apparently prefer a low or intermediate 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 smaragdula (Stål). At least for epigeic species (see below), such habitats are probably not suitable. Delphax crassicornis (Panz.) and D. pulchellus (Curt.) both prefer low and loosely growing 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 oviposits into the stem, the latter into the terminal leaf sheath. Vogel (1981) studied the distribution of three Chloriona species and Euides basilinea (Germ.) in middle Hessen (Schweinsberger 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 geminipunctata Haw., a noctuid moth), thus altering the resource availability for other groups, referred 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 production 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
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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 Auchenorrhyncha. Hairy couch (E. hispidus), Mainz couch (E. arenosus) and the recently introduced blunt-flowered couch (E. obtusiflorus) are all rare or localized. Sea couch (E. athericus) 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, common 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 species show a clear preference for this grass, notably Eurybregma nigrolineata Scott, Dicranotropis hamata (Boh.), Athysanus argentarius Metc., Artianus interstitialis (Germ.), Errastunus 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 kolosvarensis (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 meadow 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-
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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 moderately 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 loosely-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 regarding moisture, insolation and perhaps soil (see chapter 4). Further less specific Auchenorrhyncha 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 perennial; thus, hibernation of Auchenorrhyncha is possible, although not yet documented. The situation of Auchenorrhyncha assessment is rather similar as in P. pratensis regarding the total numbers, with altogether 15 species recorded. But only two of these, notably 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 pellucida (F.) and Arthaldeus pascuellus (Fall.) are the only breeding species found so far, although there is evidence for Javesella forcipata (Boh.), Anoscopus serratulae (F.), Deltocephalus 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
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three oligophagous grass-feeding species, namely Neophilaenus campestris (Fall.), Doratura homophyla (Fl.) and Mocuellus collinus (Boh.), but a host association has yet to be confirmed. Swamp meadow-grass (P. palustris), growing in wet and rather eutrophic sites, has been sampled only in a few occasions. It is probably the host of the rarely found planthopper Criomorphus williamsi China. (x) Tor-grass (Brachypodium pinnatum) Brachypodium pinnatum which forms dense, often monospecific swards on calcareous hillsides and in open forests, is another grass heavily infested by Auchenorrhyncha, with 11 confirmed plus 8 unconfirmed species. Ribautodelphax pungens (Rib.), Neophilaenus albipennis (F.), Adarrus multinotatus (Boh.) and A. bellevoyei (Put.) are all strictly monophagous, and all except the localized A. bellevoyei (Put.) are widespread and fairly abundant. Furthermore, tor-grass is an important host of Eurysella brunnea (Mel.), Stiroma affinis Fieb., Mocydia crocea (H.-S.), Elymana sulphurella (Zett.), Allygidius spp. and some others. Cook (1996) presented a detailed study of the guilds of Brachypodium pinnatum and other grasses in a British chalk grassland, including choice experiments in the field and in the greenhouse. Accordingly, additional species on this grass include Zyginidia scutellaris (H.-S.), Turrutus socialis (Fl.), Verdanus abdominalis (F.) and also Planaphrodes bifasciata (L.). Rock tor-grass (B. rupestre) is considered to belong to the same species group or as a subspecies. It is confined to limestone regions of the Alps and the Swabian and Franconian Jura, and has not been sampled separately. Surprisingly, the infestation of the closely related false-brome (B. sylvaticum), growing in open deciduous woodland usually on moist soils, is extremely low, despite extensive sampling. So far, not a single breeding species has been found. There are only a few adult records of Mocydia crocea (H.-S.) and Adarrus multinotatus (Boh.), which may have been vagrants. Recently, it has been demonstrated, that the endophytic fungus Epichloë sylvatica Leuchtmann & Schardl (Ascomycota, Clavicipitaceae), which lives specifically in B. sylvaticum, may reduce noctuid herbivory through fungal alkaloids (Brem & Leuchtmann 2001). In contrast, B. pinnatum is only rarely infected by the closely related E. typhina (Pers.) Tul. & C. Tul. (Choke) (A. Leuchtmann, pers. comm.). Further studies of this subject may reveal interesting insights into host relationships of phytophages. (xi) False Oatgrass (Arrhenatherum elatius) Arrhenatherum elatius has been a dominant grass on many lowland meadows, but is now less common due to intensive fertilization. Its occurrence in central Europe is perhaps only caused by human influence, except on some montane rocky slopes in southern parts (Körber-Grohne 1990). Much specific sampling has been done, but due to the loosely tufted physiognomy, particularly after cutting, it is often difficult to decide if sampled insects really feed on this grass. Thus, the total number of recorded Auchenorrhyncha species is 26, and includes results of field and greenhouse choice experiments conducted in Britain (Cook 1996), but only 9 of these can be considered as confirmed feeders or breeders in Germany. Megadelphax sordidula (Stål) is the only species, which has been found exclusively on this plant, although it has been reported from Scandinavia to transmit cereal viruses on Triticum and Avena. The rare Criomorphus williamsi China is perhaps another host specialist, but more field data are needed to confirm its plant relationship. Oligo- and poly-
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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 sanguinolenta (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 probably reveal more breeding species. Thus records of adult Eurysella brunnea (Mel.), Laodelphax striatella (Fall.), Dicranotropis hamata (Boh.), further species of Anoscopus, Mocydia 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. Specific 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 prefers 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 grisescens (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), Hungarian 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 greenhouse 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. Ditropsis 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
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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 reported 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 phosphate works). Macrosteles sordidipennis (Stål) has been found to breed, being perhaps strictly monophagous. Further feeders include Javesella pellucida (F.), Macrosteles sexnotatus (Fall.), Psammotettix kolosvarensis (Mats.) and perhaps Javesella salina (Hpt.), Anoscopus 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 monophagous. Other host specialists, at least on generic level, perhaps include Anoscopus limicola (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 confirmed 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.
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(xvi) Grass genera with at least one host specialist Vernal-grass (Anthoxanthum) is represented by three species. Annual vernal-grass (A. aristatum) is a psammophilous plant of the northern plains, growing in open pastures and fallow fields. Alpine vernal-grass (A. alpinum) is restricted to alpine and subalpine grassland of the Alps, the Bavarian Forest and the Harz Mountains. Both species have not been sampled for Auchenorrhyncha. In contrast, sweet vernal-grass (A. odoratum) is common and widespread, though rarely dominant, in nutrient-poor meadows, pastures and open forests. It has been sampled in many locations, but was only found to be attacked by the monophagous and rare Ribautodelphax angulosa (Rib.) and perhaps by the oligophagous Graphocraerus ventralis (Fall.). Oat-grass (Helictotrichon) (sensu Conert 1998) is a diverse group comprising c. 80 species, most of which occur in temperate Eurasia, with 4 species found in Germany. Variegated and alpine oat-grass (H. versicolor and H. parlatorei) are both rare and confined to the subalpine and alpine belt of Upper Bavaria and the Allgäu. Downy oat-grass (H. pubescens) is an inconspicuous, loosely tufted species of rather nutrient-poor meadows and pastures, usually on damp to moderately dry soils. Specific sampling is very difficult due to its scattered and low growth. Thus, there are no specific Auchenorrhyncha records from Germany, although many sites have been sampled where the grass occurred. In Britain, however, field choice experiments as well as subsequent rearings were conducted with some positive results (Cook 1996). Meadow oat-grass (H. pratense) is a locally dominant plant of rather dry pastures (never found on meadows, as the name suggests), with strongholds in limestone and basalt regions, although not actually calciphilous. Due to its low-growing tufts and isolated stems, insect records are difficult to assign. Direct sampling in some southern Lower Saxonian locations has revealed large populations of the localized Doratura horvathi W.Wg., which is apparently strictly monophagous. Further search may yield more species records. Most hair-grasses (Koeleria) are xerophilous, low-growing and loosely tufted species mainly of dry pastures and dunes. 5 species occur in Germany. Dune hair-grass (K. arenaria) is a localized western European endemic usually found on grey dunes of the East Frisian Islands. The only German location of rock hair-grass (K. vallesiana) is a single xerothermic site in the upper Rhine plain. Pyramidal hair-grass (K. pyramidata) is the most widespread and common representative of its genus in central Europe, being absent only from most parts of the northern plains, but one of the dominating grasses of grazed calcareous hillsides in middle and southern parts. Occasionally, it is also found in open xerothermic pine forests, on inland dunes or in temporarily dry meadows. Crested hair-grass (K. macrantha) shows similar habitat requirements, but is more confined to dry and basic substrates in more continental climates. For all 4 species there are no specific data on the associated Auchenorrhyncha fauna due to their low and inconspicuous growth. Glaucous hair-grass (Koeleria glauca) forms small tufts on dry and open, usually basic sands of northeast Germany and the northern upper Rhine plain. The few sampled sites have revealed the strictly monophagous Zyginidia viaduensis (W.Wg.). Furthermore, the only German record of the rare Psammotettix angulatus (Then) is from a site with dominating K. glauca, and from Russia, the closely related, perhaps even conspecific Ps. koeleriae Zachv. is reported to live on this grass. Further breeding species include Neophilae-
Poaceae
319
nus minor (Kbm.), Zyginidia scutellaris (H.-S.), perhaps also Arocephalus languidus (Fl.) and the rare Pinumius areatus (Stål). Grey hair-grass (Corynephorus canescens) is the only central European representative of its genus, growing as a perennial pioneer plant on open desalinated and decalcified sand, mainly in the north German plain and in inland dune areas further south. It is often dominant along with Festuca ovina s.l., forming low and compact tufts. In these sites, the two grasses are usually the only Auchenorrhyncha hosts. Both the monophagous Psammotettix excisus (Mats.) and the oligophagous Neophilaenus minor (Kbm.) often 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 primary 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 Doratura 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 epigejos, 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 harbouring 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 monophagous, 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 nutrient-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 preference 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, although there are many uncertain records. Dicranotropis divergens Kbm., Xanthodelphax flaveola (Fl.), Neophilaenus lineatus (L.), Streptanus marginatus (Kbm.), Arocephalus punctum (Fl.), the acidophilic form of Psammotettix helvolus (Kbm.), Ps. nardeti Rem., Turrutus socialis (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 sanguinalis 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 cereals, 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.), finger-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 physiognomy. 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 sampled, 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 Netherlands (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) occur 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), perennial rye-grass (Lolium perenne) and yellow oat-grass (Trisetum flavescens). Most feed-
322
Utilization of plant resources
ing records only refer to the most abundant and eurytopic grassland Auchenorrhyncha such as Javesella spp., Anoscopus serratulae (F.), Zyginidia scutellaris (H.-S.), Macrosteles spp., Psammotettix spp., Errastunus ocellaris (Fall.) and Arthaldeus pascuellus (Fall.). Most other members of these grass genera, like purple-stem, alpine and rough cat’s-tail (Phleum phleoides, Ph. alpinum, Ph. hirsutum), orange foxtail and black grass (Alopecurus aequalis and A. myosuroides) have not been specifically sampled. At least locally, marsh foxtail (A. geniculatus) is the main host of Javesella obscurella (Boh.). (xviii) Grasses without positive feeding records Finally, there are no feeding records for the following grasses, all of which are either annuals or rare or form only very loose stands: Millet (Panicum spp.) Smooth finger-grass (Bothriochloa ischaemum) Cut grass (Leersia oryzoides) Love-grass (Eragrostis spp.) Cord-grass (Spartina spp.) Burdock grass (Tragus racemosus) Holy grass (Hierochloe spp.) Early sand-grass (Mibora minima) Soft-bearded oat-grass (Ventenata dubia) Aira hair-grass (Aira spp.) Silky-bent (Apera spp.)
Wood millet (Milium effusum) Sheath grass (Coleanthus subtilis) Swamp grass (Scolochloa festucacea) Whorl grass (Catabrosa aquatica) Crested dog’s-tail (Cynosurus cristatus) Fern grass (Catapodium rigidum) Tough grass (Sclerochloa dura) Gravel fescue (Micropyrum tenellum) Vulpia fescue (Vulpia spp.) Hard grass (Parapholis strigosa) Wood barley (Hordelymus europaeus)
5.1.2.20 Further plant groups The remaining plants exploited by Auchenorrhyncha shall be grouped in (i) woody host plants harbouring breeding species, (ii) woody plants found to be utilized by adults or in adjacent countries only, (iii) herbaceous plants harbouring specific feeders, and (iv) herbaceous plants utilized by polyphages only. It should be noted, that this chapter deals with some of the most diverse plant taxa in central Europe, such as the Brassicaceae, Caryophyllaceae and Scrophulariaceae, with more than 100 plant species each. Further diverse groups almost neglected as food resources include the Chenopodiaceae, Primulaceae, Geraniaceae, Onagraceae, Euphorbiaceae, Boraginaceae, Campanulaceae, Liliales, Asparagales and Orchidaceae. (i) Remaining woody plants harbouring breeding species Host specialists on the remaining woody plants are very few (Table 23). There are only three monophages, notably Macropsis mulsanti (Fieb.) on sea-buckthorn (Hippophae rhamnoides), the central Asian neozoon Macropsis elaeagni Em. on cultivated Russian-olive (Elaeagnus angustifolia) and Edwardsiana diversa (Edw.) on both dogwood (Cornus sanguinea) and Cornelian cherry (C. mas). Oligophages include Opsius stactogalus Fieb. on both German false tamarisk (Myricaria germanica) and cultivated tamarisks (Tamarix spp.), and Zygina suavis R. on both common and alder buckthorn (Rhamnus cathartica and Frangula alnus). In the last case the weak point of the classification of any diet breadth becomes evident, since this species may be treated either as 2nd degree monophagous or 1st degree oligophagous, depending on the question, whether alder buckthorn is placed into
Further plant groups
323
X X x X X X x ? x ? X X x ? ? X
x x ? x
x
? ? x ? ?
?
? x ? ? x ? ? x x ?
?
A
? A A A
A
A ? ?* ? ?*
? ?
?
? ?*
? ?
?* 1
? 4 2 2 2 3 1 1 1 4
Overw intering stage
X
Substrate
X
D iet breadth
C. mas Fraxinus excelsior Ligust rum v ulgare Sambucus nigra Viburnum spp.
Aesculus hippocast anum Rhamnus cat hart ica Frangula alnus Vit is v inifera Hippophae rhamnoides Elaeagnus angust ifolia Tamarix spp. My ricaria germanica Cornus sanguinea
Species M acropsis mulsanti (Fieb.) M acropsis elaeagni Em . Edwardsiana diversa (Ed w .) Z ygina suavis R. Opsius stactogalus Fieb. Edwardsiana lethierryi (Ed w .) Empoasca decipiens Paoli Empoasca vitis (Göthe) Fagocyba cruenta (H .-S.) A lnetoidia alneti (Dhlb.) Fieberiella florii (Stål) Fieberiella septentrionalis W.Wg. Tachycixius pilosus (Ol.) Hyalesthes obsoletus Sign. A phrophora alni (Fall.) Centrotus cornutus (L.) Issus coleoptratus (F.) Stictocephala bisonia K. & Y. A lebra wahlbergi (Boh.) Empoasca affinis N ast Empoasca pteridis (Dhlb.) Edwardsiana frustrator (Ed w .) Edwardsiana prunicola (Ed w .) A guriahana stellulata (Burm .) A rboridia ribauti (Oss.) Total
Plat anus spp.
Table 23. Auchenorrhyncha species utilizing further woody host plants in Germany. Explanations see Table 5.
m1 m1 m2 o1 o1 o2 po po po po po po po po po po po po po po po po o2 po o2
P P P M P M M? P M M P P P P X P P P M M? M? M M M M
eg eg eg ad eg eg ad ad eg eg eg eg ny ny eg ny ny eg eg ad ad ? eg eg eg ad
Monophages Oligophages
Polyphages
Vertical m igrants
Unconfirm ed
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 arrowwood (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 perhaps 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 Empoasca 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 differing plant families, including both dicots and monocots. From other European countries it is furthermore reported from water-nut (Trapa natans, Trapaceae) and even from the primitive clover fern Marsilea quadrifolia (Pteridophyta, Marsileaceae).
? ? ? ? ? ? ?
?
?
D iet breadth
po po o1? o2 (m 1) po po po po po po o2 po po ?* o1? po ?
?* ?* ? ?
Lonicera spp.
Forsy t hia spp.
Sy ringa v ulgaris
? W W? ? W W? W?
Overw intering stage
?
Substrate
? S S S ?
Hedera helix
?
Euony mus europaeus
?
Buxus semperv irens
Berberis v ulgaris
?
Robinia pseudoacacia
Ficus carica
?
Ribes spp.
My rica gale
Species Issus coleoptratus (F.) Empoasca vitis (Göthe) Z ygina flammigera (Geoffr.) Z ygina angusta Leth. Z ygina nigritarsis Rem . A phrophora alni (Fall.) Lepyronia coleoptrata (L.) Philaenus spumarius (L.) Fagocyba cruenta (H .-S.) A lnetoidia alneti (Dhlb.) Empoasca decipiens Paoli Typhlocyba quercus (F.) Empoasca pteridis (Dhlb.) Fieberiella septentrionalis W.Wg. Emposaca apicalis (Fl.) Fieberiella florii (Stål)
Juglans regia
Table 24. Further woody plant species in Germany utilized by adult Auchenorrhyncha or in adjacent countries only. Explanations see Table 5.
P P M M M X X X M M M? M M? P M? P
ny ad ad ad ad eg eg eg eg eg ad eg ad ? eg ad eg
Winter/ spring
Unconfirm ed
* = after Günthart (1987a), Halkka et al. (1977), Lauterer (1980), Nuorteva (1952b), Smreczynski (1954)
Species M acropsis scutellata (Boh.) Eupteryx calcarata Oss. Eupteryx cyclops Mats. Eupteryx urticae (F.) M acrosteles variatus (Fall.) M acrosteles maculosus (Then) Z ygina hypermaculata Rem . & H lzg. Z ygina hyperici (H .-S.) Batracomorphus irroratus Lew . Sonronius binotatus (J. Shlb.) Sonronius dahlbomi (Zett.) A rboridia pusilla (Rib.) M icantulina stigmatipennis (M. & R.) Eupteryx austriaca (Metc.) Ophiola decumana (Kontk.) A gallia brachyptera (Boh.) Erotettix cyane (Boh.) Eupteryx vittata (L.) Eupteryx melissae Curt. Platymetopius undatus (De G.) A naceratagallia ribauti (Oss.) Eupteryx notata Curt.
Nuphar lut ea . . . . . . . . . . . . . . . . X . . . . .
Ny mphaea alba . . . . . . . . . . . . . . . . x . . . . .
Table 25. Auchenorrhyncha species utilizing further dicotyledonous herbs as main host or food plants in Germany. Explanations see Table 5.
Conv olv ulus arv ensis
Solanum t uberosum
Vincet oxicum hirundinaria
Epilobium angust ifolium Geranium sanguineum
Heliant hemum nummularium
Alt haea rosea
A. officinalis
Hy pericum perforat um
Poly gonum av iculare H. maculat um
Rumex acet osella
Urt ica dioica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X . . . . . . . . . . . . . . . . . . ?* . . . . . . . . . X x ? . ?* . ?* .
P. major
. . . . . . . . . . . . . . . . . . . . . .
P. lanceolat a
. . . . . . . . . . . . . . . . . . . . . .
Plant ago media
. . . . . . . . . . . . . . . . . . . . . .
Verbascum ly chnit is
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X . . . . X . . . x* . . . . X . . . . . . . . . . . . . . . . . . . . . . . . ?* . . . . x* . . . . . . . . . .
. . . . . . . . . . . . . X . . . . . . . .
Knaut ia dipsacifolia
. . . . . . . . . . . . . . . . . . x . . .
. . . . . . . . . . . . . . . . X . . . . .
Pot amoget on nat ans
. . . . . . . X . . . . . . . . . . . . . .
m1 m 1? m1 m 1? m 1? m 1? m1 m1 m 1? m 2? m 1? m1 m 1? m 1? o1 o2? o2 o2 o2 o2? o2? o2
D iet breadth
. . . . . . X . . . . . . . . . . . . . . .
P eg M eg M eg M eg P eg P eg M eg MonoM eg phages P eg P eg P eg M ad M eg? M eg P eg P eg P eg M eg OligoM eg phages P eg P ad M eg
Substrate
. . . . . . . . . . . X . . . . . . . . . . . . . . . . X X ?* . . . . . . . . . . . . .
Overw intering stage
X X X X X . . . . . . . . . . . . . . . . .
Further plant groups
325
Nuphar lut ea . . . . . . . . . . . . . . . . . . . .
Ny mphaea alba
. . . . . . . . . . . . . . . . . . . .
Vincet oxicum hirundinaria
Geranium sanguineum
Epilobium angust ifolium
Heliant hemum nummularium
Alt haea rosea
A. officinalis
Hy pericum perforat um
H. maculat um
Poly gonum av iculare
Rumex acet osella
Urt ica dioica . . . . . . . . . . . . . . . . . . . ?* . X . . . ? . . . X . . . x* . . . . X* . . . . . x* . . . . . . . . . . ?* . . . . . . . . . . . . . . . . . . . . . . . .
Solanum t uberosum
. . . . . . . . . . . . . . . . . . . .
Conv olv ulus arv ensis
?* . . x . . . . . . . A . . . . . . . .
P. lanceolat a
. . . . . . . . . . . . x . . . . . . .
?* . . . . . . . . . . . . . . . . . . .
P. major
. . . . . . x ?* x . . . . . . . . ? . .
. . . . ?* . . . . . . . . . . . . . . .
Plant ago media
. x . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . ?
Verbascum ly chnit is
?* . . . . . . . . . ?* . . . . . . . . .
. . . . . . . . . . . . . . . . . . . .
Knaut ia dipsacifolia
?* . . . . . . . . . ?* . . . . . . . . .
. . . . . . . . . . . . . . . . . . . .
Pot amoget on nat ans
. . . . . . . . . . . . . . . . . . . .
po po po po po po po po po po po po po m 1? o1? o2? po po po po
D iet breadth
?* . . . . . . . . . . . . . . ?* . . . .
X X P X M M? P M M P X P P P P P M M? P P
Substrate
. X X x . . . ? X x* . x* . . ? . . . ? ?
eg eg eg eg ad ad ? ad eg eg ny eg ny eg ad ad eg eg ad eg eg
Overw intering stage Unconfirm ed
Vertical m igrants
Polyphages
* = after Le Quesne & Payne (1981), Morris (1972), Müller (1984a), Ossiannilsson (1981, 1983), Remane (1987, and pers. comm.), Stewart (1988), Vidano (1959a), Weber (pers. comm.)
Species Lepyronia coleoptrata (L.) Philaenus spumarius (L.) A phrodes makarovi Zachv. Evacanthus interruptus (L.) Erythria manderstjernii (Kbm .) Empoasca pteridis (Dhlb.) Empoasca vitis (Göthe) Eupteryx atropunctata (Goeze) Eupteryx aurata (L.) Hyalesthes obsoletus Sign. A phrophora alni (Fall.) Centrotus cornutus (L.) Platymetopius major (Kbm .) Utecha trivia (Germ .) A gallia consobrina Curt. A gallia brachyptera (Boh.) Emelyanoviana mollicula (Boh.) Empoasca decipiens Paoli Lamprotettix nitidulus (F.) Euscelidius schenckii (Kbm .)
Table 25. (continued):
326 Utilization of plant resources
Further plant groups
327
Among the Polygonaceae, both sheep’s sorrel (Rumex acetosella) and knotgrass (Polygonum aviculare) are the main hosts of Ophiola decumana (Kontk.). The latter plant is also the only host of the strictly monophagous Macrosteles maculosus (Then). The subgenus 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’swort (Hypericum perforatum), whereas the recently described Zygina hypermaculata Rem. & Hlzg. lives in subalpine meadows on imperforate St John’s-wort (Hypericum maculatum). 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 Platymetopius, adults of which usually ascend up to the tree canopy. Among the Onagraceae, only fireweed (Epilobium angustifolium) harbours Auchenorrhyncha feeders. Both native 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 xerothermic 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 atropunctata (Goeze) and E. aurata (L.). Field bindweed (Convolvulus arvensis, Convolvulaceae) has been found to be an important food plant of the endogeic nymphs of the cixiid planthopper Hyalesthes obsoletus Sign., whereas plantain (notably Plantago lanceolata, 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 countries and include Utecha trivia (Germ.), Erythria manderstjernii (Kbm.), Eupteryx vittata (L.) and E. notata Curt. White mullein (Verbascum lychnitis), utilized by Micantulina stigmatipennis (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 dipsacifolia), 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 Cardamine pratensis and Silene flos-cuculi, the German names of which (‘WiesenSchaumkraut’, ‘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.
Humulus lupulus . . X . . . . . . . . . . . . . . . .
Veronica spp.
Echium v ulgare
Caly st egia sepium
Solanum nigrum
Geranium robert ianum
G. pusillum
Ly simachia v ulgaris
Lagenaria siceraria Cardaria draba
Viola spp.
Silene flos-cuculi
Bet a v ulgaris
Pariet aria judaica . . ? . . X ? . . ?* . . . . . . . . . x* . ?* . . . . ?* . . . . . ? . . . . .
Verbascum t hapsus
. . ? . . . ?* ?* . . . . . . . . . ?* .
Galium spp.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . x* x* . . . . . . . . . . . . ?* ?* . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . .
Valerianella locust a
. . . . . . . . . . . . ?* . . . . . .
? . . . . . . . . . . . . . . . . . ?
Sambucus ebulus
. . x . . . . . . . . . . . . . . . .
. . ? . . . . x . . . . . . . . ? . .
Valeriana dioica
. . . . . . . . . x* . . . . . . . . .
. . . . . . . . . . . . . . . ?* . . .
Succisa prat ensis
. . . . . . . . . . . . ? . . . . . .
po po po po po po o2? o2 po po po po? m 1? po? m 1? m 1? po m 1? o1?
D iet breadth
. . . . . . . . . . ?* . . . . . . . .
M? M X M X M P M P X X P M M M M X M M?
Substrate
. . X . . . . . . . . . . . . . . . .
ad ? eg eg eg eg eg ad eg ny ny eg ad ad ad ? eg eg eg eg ad
Overw intering stage Unconfirm ed
Vertical m igrants
Oligo-/ Polyphages
* = after Biedermann (1998a), Billen (pers. comm.), Cerutti (1939a), Haupt (1935), Kuntze (1937), Le Quesne & Payne (1981), Ossiannilsson (1981), Schrameyer (pers. comm.), Wagner (1935), Weber (pers. comm.)
Species Empoasca pteridis (Dhlb.) . Eupteryx atropunctata (Goeze) . Philaenus spumarius (L.) . Emelyanoviana mollicula (Boh.) . Lepyronia coleoptrata (L.) . Eupteryx aurata (L.) ?* A naceratagallia ribauti (Oss.) . Eupteryx vittata (L.) . Hyalesthes obsoletus Sign. . Cercopis sanguinolenta (Scop.) . A phrophora alni (Fall.) . Utecha trivia (Germ .) . Hauptidia distinguenda (Kbm .) . Hauptidia provincialis (Rib.) . Eupteryx urticae (F.) . Eupteryx austriaca (Metc.) . Evacanthus acuminatus (F.) . Eupteryx origani Zachv. . Emposaca apicalis (Fl.) .
Cardamine prat ensis
. x . x* . . ?* . . . . . . . . . . . . . . . . . . . ?* . ?* . . . . . . . . .
Table 26. Auchenorrhyncha species utilizing further dicotyledonous herbs in Germany. Explanations see Table 5.
328 Utilization of plant resources
Major plant clades
329
5.1.3 Differences between plant taxa 5.1.3.1 Major plant clades The distribution of the utilization of 7 major plant clades by Auchenorrhyncha in Germany is highly uneven, with strong preferences of some groups (Table 27). In general, dicotyledonous and monocotyledonous angiosperms are by far the most-favoured plants, holding more than 90% of the Auchenorrhyncha species total. Among these groups, it is particularly the graminoid monocots (Poaceae, Cyperaceae and Juncaceae, but not the Typhaceae) and the higher dicots, or tricolpates (eudicots) sensu Judd et al. (1999), which account for almost all feeding records. Even within these, the distribution is rather non-uniform (see below). In contrast, the primitive dicots (“non-monocot paleoherbs” and the “magnoliid complex”) and the non-graminoid monocots are only utilized by very few broadly polyphagous species (included in row “more than one”). Utilization of bryophytes (mosses) is not documented, although it has been suggested in a few cases of epigeic or tyrphobiotic species. Only 13 species (equalling 2%) are specialised on fungi, pteridophytes or gymnosperms. In none of these, there is any evidence of a phylogenetically old relationship, since all non-angiosperm feeders belong to more advanced Auchenorrhyncha taxa, most of which predominantly feed on more recent plant groups such as the Poaceae and Lamiaceae. In Achilidae, which largely feed on fungi in decaying wood (i.e. organisms depending, in turn, on higher plants) during the nymphal stage, adults are believed to suck on vascular plants. Likewise, the primitive dicots, comprising non-monocot paleoherbs and magnoliids (see Judd et al. 1999) are neglected, except by the polyphagous Erotettix cyane (Boh.) (see below). On the other hand, in Germany these plant groups include only the Nymphaeaceae, Ceratophyllaceae and Aristolochiaceae, with altogether only 8 species. Fungi
Pteridophyta
Unknown
Several
Higher dicots
Gymnospermae
Graminoid monocots
Fig. 4: Utilization of major plant clades by Auchenorrhyncha species in Germany (S Auchenorrhyncha = 620)
330
Utilization of plant resources
In the remaining 7% of the Auchenorrhyncha species, food plants belong to more than one clade, or are unknown. Further species utilize non-angiosperms only (i) casually, e.g. some pronounced generalists such as Philaenus spumarius (L.), or (ii) in winter, e.g. some idiocerine and typhlocybine leafhoppers, which breed on deciduous woody dicots, or (iii) in the adult stage, e.g. Haematoloma dorsatum (Ahr.) and Aphrophora corticea Germ., which are associated with Pinus, but frequently feed on angiosperms in the nymphal stage. The deltocephaline leafhopper Erotettix cyane (Boh.) is remarkable in feeding on a few plant species belonging to almost all major groups, notably the nonmonocot paleoherbs, non-graminoid monocots, pteridophytes and higher dicots (the two latter in other countries). Thus, it becomes evident that feeding on primitive plants is not likewise a primitive trait of some Auchenorrhyncha groups, but has evolved secondarily and repeatedly by host shift from both advanced dicots or monocots. Such host shifts have occurred in single species, like Ditropis pteridis (Spin.), Javesella stali (Metc.) and Eupteryx filicum (Newm.), the relatives of which live on grasses, sedges or Lamiaceae, and is also reported from aphids (Eastop 1973). In other cases, such as the Achilidae, the nymphs of which are mycophagous, host associations may have phylogenetic implications, since they are shared only with the closely related Derbidae, and hence, can be interpreted as synapomorphic (see Bourgoin et al. 1997). Furthermore, host shifts between major plant clades may perhaps be facilitated by initial acceptance of pteridophytes as additional hosts by polyphages such as Philaenus spumarius (L.) or of coniferous winter hosts, for instance in Zygina. In general, sap-sucking on plants probably dates back as far as the Carboniferous (Strong et al. 1984) and is perhaps even the oldest mode of herbivory, whereas chewing, mining and gall-making are believed to have evolved later (Shear 1991). Hemipteralike insects are known from not later than the Permian (Hennig 1969). Thus, it is rather surprising, that fungi and pteridophytes as primitive plants are not attacked by diversiTable 27. Species numbers of plants and Auchenorrhyncha of major plant clades in Germany.
Plants
Auchenorrhyncha Fulgorom orpha Cicad om orpha
Plant group Fungi Pterid ophyta Gym nosperm ae N on-m onocot paleoherbs + m agnoliid s* Gram inoid m onocots N on-gram inoid m onocots H igher Dicotyled onae More than one Unknow n Total
n ? 76 12 8 424 222 2155 – – 2897**
% ? 2.6 0.4 0.3 14.6 7.7 74.4 – – 100**
n 2a 2 1 0 108 0 18 4b 10 145
% 1.4 1.4 0,7 0 74.5 0 12.4 2.8 6.9 100
n 0 2 6 0 169d 0 270c 24e,f 4 475
% 0 0.4 1.3 0 35.6 0 56.8 5.1 0.8 100
Total n 2 4 7 0 277 0 289 27 14 620
% 0.3 0.6 1.1 0 44.7 0 46.6 4.4 2.3 100
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, pteridophytes 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/semisubterranean 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 fulgoromorphans 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 acquisition 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 provided by Sequeira & Farrell (2001), who studied DNA sequences of scolytid beetles living on the Australian and South American conifer Araucaria. 5.1.3.2 Plant families An overview of the Auchenorrhyncha of the most important vascular plant taxa in Germany and some of their life history traits is presented in Table 28. Considered plant taxa harbour at least 5 Auchenorrhyncha species or comprise at least 100 plant species. Differences in selected parameters are shown in Fig. 5, Fig. 6, Fig. 7 and Fig. 8. As in major plant clades, Auchenorrhyncha infestation is rather uneven (Fig. 5), with roughly two third of the species total feeding on only 6 families, notably the Poaceae, Cyperaceae, Fagaceae, Betulaceae, Salicaceae and Rosaceae. Thus, almost 70% of the Auchenorrhyncha species attack less than 20% of the plant species. All the remaining families are of negligible significance as food plants, and are – if at all – attacked only by extremely polyphagous species such as Philaenus spumarius (L.), Lepyronia coleoptrata
81 7
Pinaceae
8 10a
6 1 1 2 0
1,3 4
6 2 1 1
3 7 0
4
Diet bread th 1st d egree m onophagous 2nd d egree m onophagous 1st d egree oligophagous 2nd d egree oligophagous polyphagous
Ratios S Auchenorrhyncha/ S Plants S Specialists*/ S Generalists**
Substrate Phloem Xylem Mesophyll Fungi
Overw intering stage Egg N ym ph Ad ult
Vertical m igrants
2
15 1 1
6 0 11 0
2
9 2 0
5 2 4 0
17
26 12 4
22 0 19 1
7 0,4
18
45 9 7
26 2 33 0
6,8 1
0
1 0 0
0 1 0 0
0 0
0 0 0 0 1
1
6 2 2
4 0 6 0
5 0,1
0 1 0 2 7
8
48 4 14
44 4 18 0
2,1 6
30 26 0 2 8
1
1 1 0
0 2 0 0
0 0
0 0 0 0 2
3
7 3 2
6 3 3 0
0,4 1
3 1 2 3 3
8
27 8 12
17 4 26 0
0,4 0,8
10 3 8 8 18
3
15 2 4
16 3 2 0
0,2 0,9
2 1 7 5 6
2
10 2 5
8 0 9 0
2,8 0,9
5 3 0 1 8
6 17
a
0
4 0 0
0 1 3 0
0 0,3
0 1 0 0 3
93 4
2
21 2 3
3 3 20 0
0,3 1
5 2 7 4 8
82 26
1
4 0 0
0 1 3 0
1,3 0,3
1 0 0 0 3
5
21 4 5
13 5 12 0
0,1 0,7
6 4 2 3 15
130 457 4 30
0
6 3 0
7 1 1 0
0,2 0,8
0 4 0 3 2
47 9
0
60 9 5
63 5 6 0
0,5 7,2
30 30 5 1 8
0
123 66 21
192 13 6 0
0,9 10
98 20 68 4 14
144 225 74 211bc
* = 1st and 2nd degree monophages plus 1st degree oligophages, ** = 2nd degree oligophages plus polyphages; = excluding overwintering species; b = diet breadth unknown in 7 species; c = 1 species probably semivoltine, overwintering both as nymph and adult
2
2 3 2
Ulm aceae
2 2 3 0
Urticaceae 1,8 0,8
Fagaceae
4,3 1,1
Betulaceae
0,1 0,2
Caryophyllaceae
15 9 3 11 23
Tiliaceae
4 8 0 6 24
Salicaceae
5 0 0 0 6
Brassicaceae
4 5 0 1 7
Ericaceae 124 126 47 21
Rosaceae
29 12
Fabaceae
139 2
Aceraceae
32 66
Apiaceae
2 10
Lam iaceae
119 1
Scrophulariaceae
9 61
Asteraceae
6 42
Juncaceae
6 11
Cyperaceae
4 17
Poaceae
1 0 0 1 5
Ranunculaceae
Param eter Species total Plants Auchenorrhyncha
Table 28. Auchenorrhyncha species numbers of important angiosperm taxa and selected characteristics of their life histories
332 Utilization of plant resources
Plant families
333
(L.), Aphrophora alni (Fall.) and Empoasca vitis (Göthe). These plant groups include, for instance, the Papaveraceae, Caryophyllaceae, Chenopodiaceae, Amaranthaceae, Saxifragaceae, Euphorbiaceae, Violaceae, Brassicaceae, Primulaceae, Boraginaceae, Gentianaceae, Campanulaceae, Liliaceae and Orchidaceae. Most of them are also largely avoided 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 overwintering 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 numerous 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 members 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 exceptional in holding the highest number of 1st degree monophages among all central European, 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 engineer’ 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, including 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, Xanthodelphax, Muellerianella, Ribautodelphax, Wagneriala, Doratura, Rhopalopyx, Cicadula, Mocydiopsis, Streptanus, Psammotettix and Cosmotettix. It is mostly those genera in which species 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 similarities. It should also be noted that parapatric speciation, i.e. speciation promoted by geographic isolation, is unlikely to play a role in many, if not most, of the genera men-
334
Utilization of plant resources 250
S Auchenorrhyncha
200
150
100
50
al ae ic ac ea ss e ic ac e E ric a e ac e R os ae ac Fa ea ba e ce A ae ce ra ce ae A pi ac L ea S cr am e op ia hu c e a la r ia e ce A st ae er a J u ce n c ae ac C yp e a e er ac ea P oa e ce ae
e
S
ra
B
ea
ce
lia
ac yll
ph
yo
Ti
ae
ae
ce
ce
la
ga
C
R
ar
Be
tu
e
ae ce
rti ca
Fa
U
U
lm
ac ea
ce a la
na Pi
an
un
cu
ce a
e
e
0
Fig. 5: Species numbers of Auchenorrhyncha guilds on the most important 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.
8
S Auchenorrhyncha / S Plants
7
6
5
4
3
2
1
e ce ae ga ce B et C ae ar ul a yo ph c e a e yl la ce a Ti lia e ce S ae al ic B ra a ce ss i c ae ac e E ric a e ac e R os ae ac Fa ea ba e ce A ae ce ra ce ae A pi ac L ea S cr am e op ia hu c e a la ria e ce A st ae er a J u ce n c ae ac C yp e a e er ac ea P oa e ce ae
e
rti ca
Fa
lm
ac ea U
ea U
la c
cu
P R
an
un
in a
ce
ae
0
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
S Auchenorrhyncha (%)
80 70 60 50 40 30 20 10
R
an
un
P
in ac e c u ae la ce U lm a e ac ea U rti ca e ce Fa a ga e Be c e C t u ae ar y o l ac ea ph e yl la ce ae Ti lia c S al eae ic B a ra c ss ea e ic ac e E r ic a e ac e R o s ae ac e Fa a ba e ce A ae ce ra ce ae A pi a L a ce a S cr m e op ia ce hu ae la ri a ce A st ae er a J u c ea nc e C a ce yp ae er ac ea P oa e ce ae
0
1st degree monophagous
2nd degree monophagous
1st degree oligophagous
2nd degree oligophagous
polyphagous
Fig. 7: Diet breadth of Auchenorrhyncha guilds on 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.
100 90
S Auchenorrhyncha (%)
80 70 60 50 40 30 20 10
R
an
Pi
na u n ce c u ae la ce U lm a e ac ea U rti ca e ce Fa a ga e Be c e C t u ae ar y o lac ea ph e yll ac ea Ti lia e c S al e a e ic B a ra ce ss i c ae ac e E ric a e ac e R os ae ac e Fa a ba e ce A ae ce ra ce ae A pi ac L ea S cr am e op ia c hu ea la r ia e ce A st ae er ac Ju ea nc e ac C yp e a e er ac ea P oa e ce ae
0
Phloem
Xylem
Mesophyll
Fungi
Fig. 8: Auchenorrhyncha food resources of vascular plant families in Germany (excluding overwintering species on Pinaceae). Only plant families with more than 5 Auchenorrhyncha species or more than 100 plant species are shown.
336
Utilization of plant resources
tioned, since there are numerous examples of largely overlapping or even identical geographic 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, Araliaceae, Platanaceae, Myricaceae, Tamaricaceae, Cupressaceae, Cornaceae, Rhamnaceae, Oleaceae and Equisetaceae, but the overall importance of these families as Auchenorrhyncha 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, Caryophyllaceae, Brassicaceae, Fabaceae, Apiaceae, Scrophulariaceae and Tiliaceae. Most remaining groups of broad-leaved trees show intermediate values, notably the Ulmaceae, Fagaceae, Betulaceae and Aceraceae. This is also true in the Poaceae, which are further characterised by a large proportion of 1st degree oligophages. Thus, regarding specificity up to plant family level, the grasses are inhabited by the highest proportion of specialists, 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 compounds such as phenolic heterosides (salicin, populin) in the Salicaceae, and specific ethereal oils and resins in the Pinaceae. Similarly, the Poaceae and Cyperaceae are phylogenetically 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 polyphagous generalists and are unlikely to be quickly colonized by specialists (Novotný 1994a, 1995). Despite the overall dominance of host specialists among graminicoles, the species 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 Auchenorrhyncha, and perhaps, in insects in general, the degree of host specificity of graminoid feeders is clearly higher than previously thought.
Plant families
337
In general, there is a positive correlation between species numbers of Auchenorrhyncha and plants in the vascular plant families of Germany (Fig. 9), i.e. diverse plant families are exploited by more diverse guilds of Auchenorrhyncha than species-poor plant families, but only 24% of the variation is explained (P < 0.01). There is a number of species-rich plant families such as the Poaceae, Cyperaceae and Rosaceae, which harbour numerous Auchenorrhyncha species, but there are also many exceptions such as the Brassicaceae, Caryophyllaceae, Scrophulariaceae, Asteraceae as well as the entire non-graminoid monocots (e.g. the Orchidaceae, Liliaceae). These groups are altogether attacked by only 31 Auchenorrhyncha species (equalling roughly 5% of the Auchenorrhyncha species total), although they account for almost 40% of the phytodiversity. In conclusion, there is a general tendency for higher phytophage diversity on more diverse plant families, as was also shown in curculionoid and chrysomeloid beetles (e.g. Farrell 1998). However, exceptions indicate that other factors must be strongly involved. A second, more plausible correlation can be discerned when focusing on those plant families, which show rather high (> 10) Auchenorrhyncha species numbers (Fig. 9). Their common trait is a pronounced apparency in space and time, showing an exceptionally high dominance among the vegetation all over central Europe. They include three groups:
Fig. 9: Relationship between species numbers of Auchenorrhyncha and plants in the plant families of Germany. Only families with more than two Auchenorrhyncha species or more than 40 plant species are labelled: 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 = Malvaceae, 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 Salicaceae, 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 widespread 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 insects, without any host specialists, e.g. most species of Bromus, Vulpia and Aira. Similarly, 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 resources 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 either treat only small regions or species-poor groups, or they utilize different classifications 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 Auchenorrhyncha due to their more northern temperate distribution. Accordingly, aphids share the general pattern of a strong infestation of highly apparent plants, but with a pronounced preference 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 Hamamelidae sensu Cronquist (1968), i.e. on Fagaceae, Betulaceae, Juglandaceae, Ulmaceae and related families, or on some extinct group of gymnosperms. For 25 German tree genera, Brändle & Brandl (2001) found positive correlations between herbivore species numbers and present plant abundance (based on grid occupancy), pollen abundance, and tree height.
5.2 General conclusions on food plant choice The following conclusions can be drawn from these patterns: sap-sucking herbivores favour spatially and temporally highly apparent plant species, notably phanerophytes
General conclusions on food plant choice
339
and perennial graminoids, which are tall or form large, contiguous stands. In contrast, annuals, bienniels and geophytes are largely neglected, and infestation of chamaephytes and most non-graminoid hemicryptophytes is low to moderate. Further, primitive plants, such as pteridophytes and gymnosperms, have only secondarily been colonized by host shift from more advanced plant groups. The same is true for fungi, whereas bryophytes are not exploited at all. The large majority of species feed on a few plant taxa of higher monocots and dicots, notably the clades of the Poales, Juncales, rosids and asterids (sensu Judd et al. 1999). Apparently, this situation differs markedly from patterns found in Aphidoidea 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 temporarily 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 defended 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 unstable 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 substances, 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 Auchenorrhyncha 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 relatively few plant groups and an avoidance of most of the existing taxonomic plant diversity. 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 importance. 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 herbivore 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 rhizome grasses, are another group of host plants offering rather adverse conditions, since many species have green parts only in the first half of spring, when temperatures are still relatively low. As a consequence, most of them are totally avoided.
5.3 Stratification and plant architecture Auchenorrhyncha utilize all strata of central European terrestrial ecosystems. Four layers will be distinguished here (see Table 29): (i) the soil, with the mineral layer and organic litter, (ii) the herb layer, comprising the soil surface and the herbaceous vegetation (including 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 referred 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 arboricolous Macropsinae, Idiocerinae and Typhlocybinae on the one hand, and the vertically Table 29. Utilization of strata of the German Auchenorrhyncha fauna
Auchenorrhyncha total
Fulgorom orpha
Cicad om orpha
Layer
n
%
n
%
n
%
Tree
107
17.3
0
0
107
22.6
Shrub
63
10.2
0
0
63
13.3
H erb
377
60.8
112
77.2
265
55.9
Soil
1
0.2
1
0.7
0
0
Facultative m ulti-layer
23
3.7
10
6.9
13
2.7
Obligate m ulti-layer
49
7.9
22
15.2
26
5.5
Total
620
100
145
100
469
100
Stratification and plant architecture
341
migrating Cixiidae, most of which perform an obligate shift from the soil to the canopy, on the other hand. Whereas the evolution of arboricolous leafhoppers is certainly a relatively recent development, which must be seen in connection with the Tertiary rise of angiosperm trees (see section 5.4.3), the soil association of most planthopper groups is subject to a controversial debate. After Wilson et al. (1994), there is a significant tendency for primitve 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 comparative analysis of habitats (as well as ant-attendance) among Hemiptera, and concludes 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 Deltocephalinae, 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 species have expanded their range or even invaded central Europe. According to KörberGrohne (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 Megadelphax 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 other hand, some open forests with a well-developed grass layer, e.g. pine forests, coppiced 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, accounting for 27% of the Auchenorrhyncha total, and 36% of the cicadomorphans (see above). Reliable field data on canopy abundance in central Europe do not exist, but there is some indirect evidence. In a single tree-eclector in a 30-year old monospecific,
342
Utilization of plant resources
uniform stand of Populus tremula in Thuringia, c. 33,000 individuals of the idiocerine leafhopper Populicerus laminatus (Fl.) were collected within one season. The abundance in this site was estimated to be more than 5000 ind./m2, which is the same order of magnitude as in grasslands (see above). In this particular case, the real abundance was probably even much higher, since P. laminatus (Fl.) is a permanently arboricolous species, and only a certain proportion of the total number of individuals is likely to have fallen down due to leaf-trembling caused by wind (which is certainly an aspen pecularity). More exact figures of canopy abundance may be gained by tree fogging, but this method has only rarely been applied in European forests (see section 3.5.2). The diversity of arboricolous leafhoppers is positively correlated with the large amount of biomass and the structural complexity of woody plants, which facilitated sympatric speciation without host shift (see section 5.1.3.2). Among native species, Fraxinus excelsior is the only one, which is almost devoid of Auchenorrhyncha. Most of the remaining tree species harbour diverse and specific guilds (see section 5.1.2). Among the most important host families, the ratio of species numbers Auchenorrhyncha : plants is clearly highest in trees, and declines rapidly in the following order: Fagaceae > Betulaceae > Tiliaceae > Ulmaceae > Aceraceae > Salicaceae > Urticaceae > Pinaceae > Poaceae > Cyperaceae (see section 5.1.3.2). This pattern of higher insect species numbers on structurally complex plants is also evident within the herb layer. For instance, among grasses, the most diverse guilds are found on tall (e.g. Phragmites), tussock- forming (e.g. Festuca ovina, Deschampsia cespitosa) or turf-forming (e.g. Elymus repens, Calamagrostis epigejos) species, whereas loosely growing species such as Milium effusum, Festuca gigantea, Bromus ramosus, Hordelymus europaeus and Helictotrichon pubescens, are only rarely attacked. Further uninfested grasses are also found in the genera Bromus, Aira, Vulpia, Hordeum and Eragrostis, but in these cases, architectural simplicity and small size are combined with a therophytic growth form, resulting in a reduced temporal persistence (see below). High Auchenorrhyncha species numbers on structurally complex plants are in accordance with predictions made by Lawton (1983) and Strong & Levin (1979), who demonstrated that plant architecture is a major determinant of insect phytophage diversity, based on data sets on various insect groups mainly from Britain and North America. The most important components of plant architecture include size and resource diversity. For Auchenorrhyncha, as well as other homogeneous feeding guilds, the latter may be only of minor significance, since even the smallest vascular plant stems and leaves provide the basic resources of phloem, xylem, mesophyll and oviposition sites. Other resources are only utilized by very few species, such as Ledra aurita (L.) and some Idiocerinae, which are often found on older bark. In the data base presented here, the statement of diverse herbivore guilds on structurally complex plants is clearly corroborated concerning plant family level (see Fig. 6), but must be modified when considering plant species level. Fig. 14 shows the most diverse guilds in the study area, with at least 18 Auchenorrhyncha species per plant species. Seven out of 12 of these guilds live on trees or shrubs, whereas the remaining 5 live on grasses. Although maximum species numbers are found on Quercus and Betula, species numbers on Festuca ovina and Calamagrostis epigejos are also considerable. The degree of host specificity on the grasses is higher than on the trees, despite their lower
343
Stratification and plant architecture
architectural complexity. Moreover, data on insects feeding on grasses are fewer compared 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 Fulgoromorpha and the Cicadomorpha, which perform obligate vertical migrations from one stratum 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 Aphrophorinae, 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, Thamnotettix 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 specific during oviposition and larval development, but become oligophagous or even polyphagous 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 genTable 30. Species numbers of obligate vertical migrants and their utilization of strata
Auchenorrhyncha total Layers
n
%
Soil - H erb
5
10,2
Soil - Shrub
16
32,7
Soil - Tree
8
16,3
H erb - Shrub
3
H erb - Tree Total
Fulgorom orpha n
Cicad om orpha
%
n
%
2
9,1
3
11,5
13
59,1
2
7,7
7
31,8
1
3,8
6,1
0
0
3
11,5
17
34,7
0
0
17
65,4
49
100
22
100
26
100
344
Utilization of plant resources
eral, nymphal diet width and other ecological traits of vertically moving species are poorly 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 development (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 differentiation 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 discussion (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 utilization of the upper soil perhaps also occurs in a number of epigeic taxa such as Eupelix, Aphrodinae and Errhomenus, which may suck on fine roots in the top soil layer.
5.4 Utilized substrates Table 31 shows the utilization of different types of food source. The type is largely combined with phylogenetic affiliations (see below). It should be noted, however, that most feeding experiments focused on relatively few species of economic importance, and that opinions on whole subgroups gained by mere generalisation may have to be revised in future. The differential exploitation of resources of the most important plant families in the study area is shown in Fig. 8. Important reviews on these resources have been presented by Raven (1983) and Tonkyn & Whitcomb (1987) and will be discussed below. Table 31. Substrate utilization of the Auchenorrhyncha species of Germany
Phloem
345
5.4.1 Phloem Phloem-feeding is found in all the fulgoromorphans, except the mycophagous Achilidae 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 morphological 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, Membracidae, 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 constant, 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 dicotyledonous 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 Lamiaceae. 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 brachypterus Fieb. must be considered as tentative due to its unclear systematic position. Only the cicadas, Cercopidae and Cicadellinae are involved. Examples include Philaenus spumarius (L.), which is certainly one of the world’s most polyphagous insect herbivores, as well as some other pronounced generalists such as Lepyronia coleoptrata (L.), Aphrophora alni (Fall.) and Cercopis spp. Xylem feeders, notably the Cicadidae and Cercopinae, 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 utilized resources in different plant families (Fig. 8) shows that xylem-feeding species dominate 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 content 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 dilatory cibarial muscles. Novotný & Wilson (1997) demonstrated that simply for physical reasons, 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 species 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 S xylem feeding species
12
y = 0,0219x + 0,0713 2
10
R = 0,5396***
8 Ran
6 4
Sal Eri
2
Lam Ros
Rub Pol
Api Car
0 0
100
Ast
Cyp Fab Bra Scr 200
300
400
500
S plan ts Fig. 10: Species numbers of xylem-feeders on the vascular plant families in Germany. Ast = Asteraceae, Bra = Brassicaceae, Car = Caryophyllaceae, Cyp = Cyperaceae, Eri = Ericaceae, Fab = Fabaceae, Lam = Lamiaceae, Pol = Polygonaceae, Poa = Poaceae, Ran = Ranunculaceae, Rub = Rubiaceae, Sal = Salicaceae, Scr = Scrophulariaceae.
Xylem
347
notably Aphrophora pectoralis Mats., A. salicina (Goeze), Neophilaenus albipennis (F.) and probably N. infumatus (Hpt.) and N. exclamationis (Thnbg.). These species are accociated with willows or grasses, respectively, i.e. members of plant groups, which generally hold high proportions of host specialists (Fig. 7). Therefore, the dominance of xylem feeders on plants largely avoided by the remaining 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 towards 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 strictly associated with rather unusual and toxic host plants such as Asphodelus (Asphodelaceae) 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 phloemfeeders. Due to the much diluted and unbalanced soluables, larger body size and locally high abundance (up to 1280 ind./m2 in Michigan, USA – see Wiegert 1964), the removed 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) experimentally 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 measurable in the aphid. Thus, the turnover of assimilates and the functional significance of xylem-feeders may in fact be higher than previously thought.
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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 containing 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 Empoasca 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 reevaluate applied aspects as well as the phylogenetic significance of mesophyll-feeding 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 venation (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 Typhlocybinae is likely to constitute a more recent, though widespread evolutionary trend, which has independently occurred within the Erythroneurini and Typhlocybini. On the other 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-hemipterologists, the characteristic stippling of leaves, particularly visible late in the season, 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 sampling 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 almost 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 dominance of mesophyll-feeding species on some higher dicotyledonous herb families, no-
Fungi
349
tably the Lamiaceae, Asteraceae, Ranunculaceae, Apiaceae and Scrophulariaceae, although species numbers are generally low on the last three mentioned families. It should also be noted here, that mesophyll-feeding typhlocybids have managed to perform some rather spectacular host shifts. Examples include Eupteryx filicum (Newm.) on Polypodium and other ferns, although most congeners live on Lamiaceae. Zygina, subgenus Hypericella, is the only Auchenorrhyncha taxon specialised on Hypericum, a member of the Clusiaceae, which is otherwise only (rarely) exploited by Philaenus spumarius (L.). Most relatives live on woody trees or shrubs (see below for further examples). The second feature of mesophyll-feeders is their high proportion on most woody plants, notably the Ulmaceae, Fagaceae, Betulaceae, Tiliaceae, Rosaceae and Aceraceae, which offer enormous quantities of resources. Mesophyll-feeding is also found in the members of the Dikraneurini, all of which live on leaves of sedges, grasses and forbs, except the Mediterranean Liguropia juniperi (Leth.), which is found on ornamental Chamaecyparis (Gymnospermae, Cupressaceae). Among the Empoascini, Chlorita, Austroasca and some species of Empoasca live on leaves of herbaceous plants, and are considered to feed on mesophyll, although no experimental studies have been conducted. The Typhlocybini are largely arboricolous on broadleaved trees, except Wagneripteryx germari (Zett.), which has secondarily shifted to gymnosperms (Pinus spp.). Further exceptions are found in Eupteryx and Edwardsiana sociabilis (Oss.). The latter has colonized ornamental Rosa rugosa, but originally lives on the herb Filipendula ulmaria. The genus Eupteryx is exclusively found on herbs and is a good example of a secondary host shift towards the herb layer in an arboricolous group. Preferred host plants mainly belong to the rather advanced Lamiaceae and Asteraceae, many members of which originate from semi-arid parts of southern Europe and Asia. Similarly, among the largely arboricolous Erythroneurini, the genus Zyginidia is purely graminicolous, whereas Zygina species of the subgenus Hypericella are specialised on herbaceous Hypericum spp. (see above), and Z. rubrovittata (Leth.) lives on the dwarf shrub Calluna vulgaris. Further, among the genus Arboridia, arboricolous species are found along with feeders on shrubs, as well as on the herbaceous Geranium and Potentilla. Finally, Hauptidia comprises species reported from a number of rather unusual food plants, including Silene (Caryophyllaceae), Primula (Primulaceae), Digitalis (Scrophulariaceae), Geranium (Geraniaceae), Galium (Rubiaceae) and Ecballium (Cucurbitaceae) (Dworakowska 1970d; della Giustina 1989). 5.4.4 Fungi Among Auchenorrhyncha, feeding on fungal mycelium is rather exceptional, and only found in nymphs of the fulgoromorphan groups of the Achilidae and Derbidae (Wilson et al. 1994). Usually, they live on decaying wood, often under bark, but details on their biology are unknown. Adults show morphological traits of typical corticoles, i.e. their body is dorsoventrally flattened, and their colouration resembles the bark of their preferred tree species. As in mesophyll and xylem feeding, this type of nutrition is highly likely to have evolved from phloem feeding, because it is peculiar to two groups treated as adelphotaxa by Bourgoin et al. (1997), and also because their food apparently depends on higher plants. Both groups, however, are mainly found in more southern countries (Fennah 1950; O’Brien
350
Utilization of plant resources
& Wilson 1985), and only two species of Achilidae occur in Germany. Due to their life habits, they escape most hemipterologists, and thus, only a handful of records is known from both species, without any data on their feeding habits.
5.5 Pests on cultivated plants Planthoppers and leafhoppers play an important role as pests on various cultivated crops by transmitting plant diseases as well as by direct feeding. Transmitted pathogens include viruses, bacteria and mycoplasma-like organisms (MLO), some of which have not yet been properly identified (Maramorosch & Harris 1979; Nault & Rodriguez 1985). Although much research has been done on etiology and vector control, knowledge of causal connections is often poor. Thus, the capability of transmitting pathogens has been demonstrated for many species under laboratory conditions, but the real amount of resulting crop damage has rarely been quantified. There is no doubt that some species are serious pests of rice, maize, legumes and other crops in tropical and subtropical latitudes. Important planthopper pests include the delphacids Nilaparvata lugens (Stål) and Sogatella furcifera (Horv.) on rice, Peregrinus maidis (Ashmead) on maize, and Perkinsiella species on sugarcane. Deltocephaline leafhoppers of the genera Dalbulus, Nephotettix, Circulifer and Macrosteles are vectors of diseases of maize, rice and other cereals as well as beets (Müller 1956; Nault & Rodriguez 1985; Wilson & Claridge 1991). In central Europe, however, there are only few recent cases where damage has been proved, most of which has occurred in cultivations of perennial plants and in greenhouses. In vineyards of southwestern Germany the cixiid planthopper Hyalesthes obsoletus (Sign.) has been found to transmit an MLO-associated grapevine yellows disease locally known as ‘Vergilbungskrankheit’ (Maixner et al. 1995). More recently, Psammotettix alienus (Dhlb.) has been assumed to cause damage in barley fields of Sachsen-Anhalt by transmission of wheat dwarf virus (WDV) (Manurung et al. 2001). Graphocephala fennahi Young is suspected to transmit Pycnostysanus azaleae (Peck.), a fungus causing bud blast on ornamental Rhododendron, although the etiology is not properly known and evidence is ambiguous (Vidano et al. 1987b). Scaphoideus titanus Ball is another Nearctic species, belonging to the Deltocephalinae and potentially being noxious to grapevines (Maixner et al. 1993). First recorded in the Gironde area, France, in 1958, it has spread over most French viticultural areas and has invaded northern Italy and Ticino (della Giustina 1989; Maixner et al. 1993; Bosco et al. 1997), but so far, it has not yet reached Germany. Direct damage by sucking plays a more important role, with most species involved belonging to the Typhlocybinae. During recent years, Empoasca decipiens Paoli has caused losses of cucumber and sweet pepper yields in greenhouses in the Lake Constance area (Schmidt & Rupp 1997; K. Raupach, pers. comm.). The closely related E. vitis (Göthe) has periodically become noxious in vineyards of southwestern Germany (Louis & Schirra 1997; Schruft 1983; see also Günthart & Günthart 1967). More recently, Eupteryx decemnotata Rey, E. melissae Curt. and perhaps some congeners have attacked various officinal herbs in gardens and fields, notably sage, marsh-mallow and balm. However, the extent of damage is usually limited. Locally, Edwardsiana rosae (L.), attacks ornamental roses and is able to cause damage, but can easily be con-
Pests on cultivated plants
351
trolled by insecticides. At the beginning of the 20th century, Tettigometra obliqua (Panz.) was reported to have invaded fields of rye and oats, with losses of at least some plants (Torka 1905). Nowadays, this species appears to have completely vanished from most parts of central Europe. Similarly, Empoasca pteridis (Dhlb.) and Eupteryx atropunctata (Goeze) are both found on potatoes, but damages are apparently exceptional, although leaf-rolling after feeding has been confused with symptoms of virus diseases (Müller 1956). Very recently, the Mediterranean typhlocybid Hauptidia provincialis (Rib.) has turned up in a greenhouse of Bavarian Swabia causing feeding damage on corn salad and perennial basil. The introduced treehopper Stictocephala bisonia Kopp & Yonke has become a pest in Italy on grapevine and various fruit trees. After oviposition into the bark of young twigs, the phloem stream may become interrupted, resulting in necrosis (Vidano 1963a, 1963b; Günthart 1980). From Germany, however, no damages have been reported yet, although the species is locally common in southwestern parts of the country. Apart from Graphocephala fennahi Young, xylem suckers have become noxious only in forest cultivations. Schmutterer (1953) observed a mass attack of Cicadella viridis (L.) in nurseries of black alder, with twigs and young trees dying after extensive oviposition. In the Netherlands and the Mediterranean region, Haematoloma dorsatum (Ahr.) is causing 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 herbaceous 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 reported as pests from other countries, but not from Germany, notably Megadelphax sordidula (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 comprised almost exclusively migrants (Raatikainen 1971, 1972). Moreover, a considerable proportion was likely to feed on weeds, such as Phleum pratense, Elymus repens, Deschampsia 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.
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6 Life strategies
6.1 Host specificity and its possible causes Resource specialization is a central issue in evolutionary ecology (e.g. Begon et al. 1996; Futuyma & Moreno 1988) and is thought to be correlated with the dramatic radiation of phytophagous insects and mites and their angiosperm hosts (Hutchinson 1959; Strong et al. 1984; Farrell 1998; Schoonhoven et al. 1998; Walter & Proctor 1999). Concerning insects, the discussion on diet width for a long time focused on plant defence and reciprocal evolutionary effects between insects and plants, initiated by the classical paper of Ehrlich & Raven (1964) (see below). Crawley (1983) summarized advantages of polyphagy and monophagy. Accordingly, food is easier to find for polyphages, and its availability is temporally more constant. Energetic costs and the risk of predation during search are low, and shortage of certain nutritious elements in a single plant species can be balanced. Moreover, toxic compounds of certain host species may be tolerated in low concentrations. In contrast, monophages can develop special adaptations in morphology, physiology and behaviour. Thus, their feeding efficiency is high, and they may avoid interspecific competition. Therefore, monophagy is likely to be advantageous in small and short-lived animals such as insects, which need only a few plant individuals for feeding. The following sections will present broad patterns found in the studied Auchenorrhyncha guilds and discuss the relevant theories. 6.1.1 Diet width in Auchenorrhyncha The classification of diet width used here is shown in Table 4 (see also Fig. 11). Altogether 5 categories are distinguished, which must – at the present stage of knowledge – deal with inconsistencies caused by different taxonomic categories of plants, and also, if comparative studies are intended, with diverging classifications of diet width of other authors (see Symons & Beccaloni 1999). However, the data matrix presented here (sections 4 and 5.1.2) should allow an alignment with categories of other authors. Species were preliminarily assigned (marked with “?”) to one of the categories of diet width after at least 3 host (i.e. breeding) records (or alternatively, feeding records in vertical migrants and polyphages). Substantiated assignment is assumed here after at least 10 records. These criteria may seem rather speculative, but will be maintained here based upon the following arguments:
Diet width in Auchenorrhyncha
353
(i) In most species, the number of host records is clearly higher than three. The total number of own insect – host records collected for the present study is 8,600. Thus, the average number of positive plant records per Auchenorrhyncha species is only 14, but these data are further supported by numerous negative plant records and literature data. (ii) Field experience has clearly shown that the bulk of previously only little known species shows specific host requirements, and consequently, the occurrence is often rather patchy, although the overall distribution may be wide. There are numerous examples of species formerly thought to be rare, which turned out to be fairly common, as soon as their hosts had been identified. This is particularly true for Kelisia sima Rib., K. guttulifera (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 Germany 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 locality or region, may show a broader host range in breeding experiments and in other places. Thus, the term ‘monophagy’ needs not necessarily imply exclusive morphological or physiological adaptations, but may simply involve a functional or regional specialization. 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. However, 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 pronounced 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
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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
Diet w id th 1st d egree m onophagous 1st d egree m onophagous? 2nd d egree m onophagous 2nd d egree m onophagous? 1st d egree oligophagous 1st d egree oligophagous? 2nd d egree oligophagous 2nd d egree oligophagous? Polyphagous Polyphagous? Unknow n Total
Auchenorrhyncha total n % 138 22.3 101 16.3 84 13.5 44 7.1 69 11.1 41 6.6 18 2.9 22 3.5 43 6.9 46 7.4 14 2.3 620 100
Fulgorom orpha n % 41 28.3 25 17.2 17 11.7 11 7.6 9 6.2 6 4.1 0 0 4 2.8 3 2.1 26 17.9 3 2.1 145 100
Cicad om orpha n % 97 20.5 76 16.0 67 14.1 33 7.0 60 12.7 35 7.4 18 3.8 18 3.8 40 8.4 19 4.0 11 2.3 475 100
half of this number, respectively. Higher numbers of monophages are also found on the families of dominating woody plants, notably the Betulaceae, Rosaceae, Fagaceae, Ulmaceae, 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
355
Plant apparency 140 1 15
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S
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120
Fig. 12: Species numbers of 1st and 2nd degree monophagous Auchenorrhyncha on the plant families of Germany
toxic secondary compounds. Noteworthy are also 5 monophages on the Urticaceae, which are restricted to Urtica dioica. The situation is somewhat different when we regard plant genera (Fig. 13) Carex (true sedges) is by far the most preferred taxon, with 46 specialists, followed by Salix (willows and sallows), Festuca (fescue), Populus (poplar), Calamagrostis (small-reed) and Phragmites (common reed). The latter comprises only a single plant species and is notewothy in harbouring the highest number of 1st degree monophages all over central Europe and probably all over the western Palearctic. Among the graminoids, Agrostis, Eriophorum, Brachypodium, Molinia, Phalaris and Juncus are noteworthy. Among deciduous trees, Quercus, Ulmus, Alnus, Betula, Acer, Corylus, Rosa and Picea are the most species-rich. Only Urtica and Artemisia are noteworthy among the remaining genera. The former is purely herbaceous, and only a single species is attacked. Among the latter, only dwarf shrubs are exploited by specialists. The following sections will discuss possible explanations for these patterns against the background of more general theories. 6.1.2 Plant apparency High absolute numbers of monophages on highly apparent trees, graminoids and Urtica provide evidence, that first of all, plant apparency promotes herbivore specialization. Plants with many specialist herbivores are tall or grow in large stands, and they are perennial and widespread (see Fig. 12, Fig. 13, Fig. 14). A similar preference of apparent host plants is also found in Sternorrhyncha (based on worldwide data). The majority of host-specific species of Aphidoidea, Psylloidea,
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Life strategies 50 45
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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 Aphidoidea, the Asteraceae, which are largely herbaceous, are the most favoured plant family (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 regarding 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 climate and geology, whereas spatial plant apparency comprises range size, abundance, as well as plant size and architectural complexity. Some of these factors are intercorrelated 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 classical 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.
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6.1.3 Plant diversity Species-rich plant groups like the Poaceae, Cyperaceae and Salicaceae (and Carex, Salix, Festuca in particular), tend to be attacked by more herbivore specialists than species-poor groups (Fig. 13). Moreover, these specialists tend to be closely related and are often congeneric, for instance in Kelisia, Ribautodelphax, Macropsis, Idiocerinae, Kybos, Mocydiopsis, Psammotettix and Cosmotettix. Many of these hosts show a similar geographic distribution, and thus, parapatric speciation is unlikely. However, the significance of secondary plant compounds may be low at least on graminoids, which are more physically defended against chewing herbivores. One of the best documented examples of sympatric host-associated race formation with evidence for incipient speciation is found in the Nearctic membracid treehopper Echenopa binotata Say (Wood & Guttman 1981, 1982; Wood 1987). In this species complex, different host races are found on 7 plant genera belonging to 6 taxonomically rather distinct families (notably the Magnoliaceae, Rutaceae, Juglandaceae, Celastraceae, Caprifoliaceae, Fabaceae). Interestingly, speciation appears to be mediated rather by host-specific water contents causing differences in phenology of egg hatch, and perhaps the impact of a myrmarid egg parasitoid, rather than by plant defence (Wood 1987, 1988). Among central European Auchenorrhyncha, such examples of sympatric speciation on different hosts are most likely to exist in guilds living on grasses, sedges, Salicaceae and perhaps Lamiaceae. 40
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1st degree monophagous
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Fig. 14: Species numbers and diet width of the most diverse Auchenorrhyncha guilds (> 17 species) in Germany
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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 abiotic parameters in the canopy may be less variable, irrespective of site conditions. 6.1.4 Plant defence The evidence for surface defence in central European plants against Auchenorrhyncha is rather limited. High contents of silica in epidermal tissues of Equisetum spp., which may deter cattle and other chewing herbivores, are apparently of little efficiency against specialized sap-sucking insects, like Javesella stali (Metc.) and Macrosteles frontalis (Scott) (see section 5.1.2.1). Similarly, trichomes are found in many dicotyledonous plants heavily attacked by mesophyll-sucking typhlocybine leafhoppers, such as Urtica dioica and numerous species of Lamiaceae. The same is true in some woody species showing hairy leaf undersides, e.g. among the Salicaceae and Rosaceae. In fact, the most heavily attacked plants are largely glabrous (notably most species of Poaceae, Cyperaceae and Fagaceae) or lose pubescence after a while (at least on the upper leaf surface, for instance some species of Salix, Populus and Betula). In plant families or genera with both glabrous and hairy species, there appears to be no difference in species richness of Auchenorrhyncha guilds, e.g. in Populus and Salix (see section 5.1.2.9). As for chemical plant defence, the data presented here can only offer some general clues. Overall, 75% of the Auchenorrhyncha species of the German fauna feed on phloem or xylem. Unlike leaf parenchyma, these plant saps generally contain low levels of digestability-reducing compounds, which are generally of a high molecular weight. In contrast, concentrations and significance of toxins, which tend to consist of smaller molecules, are only poorly studied (Raven 1983), but may be more important than previously thought (see Dixon 1998). Noxious compounds are likely to play a role in some rather diverse plant taxa which are entirely avoided by Auchenorrhyncha. This may be the case in most ferns (particularly the Aspleniaceae and Dryopteridaceae), the Ranunculaceae, Chenopodiaceae, Caryophyllaceae, Brassicaceae, Onagraceae, Euphorbiaceae, Apiaceae, Gentianaceae, Boraginaceae, Scrophulariaceae, Campanulaceae, Rubiaceae, Asteraceae, the former Liliaceae (now split into several families) and the Orchidaceae. In some of these groups, however, for instance in the Chenopodiaceae, Caryophyllaceae, Brassicaceae and Onagraceae, reduced apparency may offer an alternative explanation because they comprise numerous annuals, biennials and rare species. Further groups, which are known for their toxic compounds, such as Hypericum (Clusiaceae), Chaerophyllum (Apiaceae) and Knautia (Dipsacaceae) are attacked only by a small number of specialised, monophagous mesophyll feeders.
Plant defence
359
The Salicaceae are probably the best example of a plant group showing chemical compounds 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. Similarly, 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 specifically studied (Pollard 1968, 1969). Thus, detoxification or sequestering mechanisms are likely to occur in this taxon. Conversely, Fraxinus excelsior (Oleaceae), is the only central European 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 remain 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 welldocumented 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 (Crawley 1983; Futuyma 1983). Principally, two categories of chemical defence have been distinguished (Feeny 1976; Rhoades & Cates 1976; see also Howe & Westley 1988). Qualitative defences, i.e. toxins, such as alkaloids, cyanogens, glucosinolates, non-protein amino acids and terpenes, comprise small molecules and cause direct poisonous effects. 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 assimilation, 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 broadly 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 properties of secondary compounds may include repelling, growth inhibiting, reducing fecundity and delaying of development (see Hartley & Jones 1997). Further ideas stressed
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a possible synergistic interplay of different substances and their chemical distinctiveness, although a study of the British insect fauna on umbellifers did not provide positive evidence (Jones & Lawton 1991). However, there is a number of shortcomings in explaining observed patterns by these theories of plant defence. Thus, ideas stressing mutual effects on the ecology and evolution of insect herbivores and plants have been challenged by the theory of sequential evolution, which generally questions the impact of insects as a driving force in plant evolution, thereby regarding most insect – plant relationships as rather asymmetric. Instead, climate, soil, interspecific competition, pathogens and other factors are likely to be much more relevant for plants, whilst the overall impact of insects on their hosts is negligible (Jermy 1976). Concerning Auchenorrhyncha and other sap-sucking insects, published evidence of specific plant defence mechanisms is rather scarce compared to leaf-chewing groups. For instance, pubescence of cotton, soybean, alfalfa, potato and bean has been found to act as an effective physical barrier against leafhopper probing and feeding, particularly in the nymphs of Empoasca fabae (Harr.) (Tingey 1985). On the other hand, Lovinger et al. (2000) also found evidence for negative effects of pubescence on parasitoid searching behaviour (see also Eisner et al. 1998). Plant defence through protection by animals has long been known mainly as plant-ant mutualism (Janzen 1966; Price et al. 1980). Since the 1980s, however, much research work has focused on indirect plant defence through various kinds of organisms, including parasitoids, predators and endophytic fungi (Sabelis et al. 1999; Clay 1996; Leuchtmann et al. 2000; Brem & Leuchtmann 2001). Effects on Auchenorrhyncha have not been subject to study so far. The significance of secondary plant metabolites for phloem and xylem feeders was discussed by Raven (1983). He suggested that quantitative defences are of limited efficiency due to their larger molecular size and reduced solubility compared to mostly monomeric and oligomeric nutrients. Moreover, they are toxic to the symplast and thus, must be sequestered by the plant. In contrast, the toxicity of qualitative defences to plant tissues is generally less severe, because they are often activated or released only where and after tissue has been damaged by herbivores. Although toxins appear to be widespread in the phloem sap of plants, their content is considerably lower than in other tissues. As for xylem sap, the concentration of toxins may be potentially high, but the evidence suggests that this is only exceptionally the case. Instead, the low and unbalanced nutritional value combined with the strong negative pressure in itself renders xylem sap a rather unattractive resource. Thus, Raven (1983) states, that there is no reason to assume major negative effects of digestability-reducing compounds on sap-sucking Hemiptera, while evidence for toxins as deterrents is inconclusive. Regarding Asian planthoppers of rice and some aphids on plants of economic importance, there is some scattered evidence of negative response to toxins and even more complex polymers, but the present data do not suggest a major significance (Risebrow & Dixon 1987; Cook & Denno 1994). In contrast, mesophyll-feeding typhlocybine leafhoppers ingest cells mainly of the palisade and spongy parenchyma, and principally have to cope with secondary metabolites. Their contact may be reduced, since glands and other storage organs are more common in or on epidermic tissue, although vacuoles, which also contain secondary metabolites, are mechanically broken before passing the food canal (Pollard 1968). Thus, we may conclude, that the relationship of mesophyll-feeding leafhoppers and chemical plant com-
Nitrogen
361
pounds is largely unknown, but the dramatic relatively recent radiation of Typhlocybinae suggests, that plant chemistry was not a major obstacle for detoxification. There is indirect 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 nitrogen. 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 utilization 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 eurytopic, polyphagous and bivoltine species, which a priori show two annual peaks of abundance (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 results). This phenology probably reflects the overall dominance of monovoltine Auchenorrhyncha 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 species, which favour certain life strategies, but should not be generalized. Climate is likely to play a much more important role for insect phenology than nitrogen content. Furthermore, in phloem and xylem feeders, gain of nitrogen in summer can simply be achieved by spending more time on sucking, even without moving to other plants or feeding sites. An important clue to assess the significance of nitrogen for herbivore host specificity may be provided by correlating the Auchenorrhyncha species numbers on different grasses to phloem nitrogen content. 6.1.6 Interspecific competition The role of interspecific competition on resource specialization has been a classical topic in evolutionary ecology. Direct competition is most likely to occur, if two or more species utilize a similar niche, and if densities are high in at least one of them. Central European Auchenorrhyncha species tend to concentrate on relatively few plant fami-
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Life strategies
lies and species (Fig. 9, Fig. 12, Fig. 13, Fig. 14). Thus, the potential for competition should be rather high. However, on a closer look, niche parameters like host plant species, 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 Lamium 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 existTable 33. Some potential model guilds for further studies of interspecific competition
H ost plant Populus nigra Populus tremula Salix alba Salix caprea Salix spp. Corylus avellana Betula pendula A lnus glutinosa Quercus robur Quercus petraea Ulmus spp. A cer pseudoplatanus A cer campestre Stachys sylvatica Salvia officinalis Juncus effusus Phragmites australis
Potential com petitors Taxon Species num ber Id iocerinae 5 Id iocerinae ≥3 M acropsis 3 Id iocerinae 5 A phrophora spp. 2 Edwardsiana 5 Oncopsis ≥4 Typhlocybinae ≥ 12 Typhlocybinae ≥ 13 Typhlocybinae ≥ 12 Edwardsiana 4 Typhlocybinae 8 A cericerus ≥2 Eupteryx ≥3 Eupteryx ≥5 Conomelus 2 Chloriona 7
Param eters potentially allow ing niche partitioning Tim e, bod y size Bod y size, sun exposure Geographical range, bod y size Tim e, bod y size, alternative hosts Altitud e, alternative hosts Geographical range, sun exposure Altitud e, bod y size, m oisture Moisture, alternative hosts? Moisture, alternative hosts Alternative hosts, sun exposure? Geographical range?, sun exposure? Moisture ? Geographical range, m oisture, alternative hosts Geographical range Geographical range, m oisture Geographical range, m oisture, salinity
Interspecific competition
363
ence’, and later, Gause’s (1936, 1937) principle of competitive exclusion. In the presence of potential competitors, specialization would occur, eventually leading to character displacement (Futuyma & Moreno 1988). Consequently, closely related phytophagous insects should avoid competition by utilizing different host plants or parts of them. During the second half of the 20th century serious doubts about the overall significance of interspecific competition became evident, particularly for herbivore communities, and alternative models were proposed (e.g. Hairston et al. 1960; Strong et al. 1984). These authors 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 below 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 reviewed (Denno et al. 1995; see also Stewart 1996). Accordingly, the bulk of experimental studies carried out mainly through the 1980s revealed negative effects between potential competitors, particularly among closely related, introduced, sessile and aggregative 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 conclusive, since researchers are likely to select systems with a higher probability of positive 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 smallscale 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 interspecific 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 intraand interspecific competition. However, densities in these experiments were high, ranging 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 monophagously 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 & Tscharntke 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 southeast 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 densities. 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 competition is found in xylem-feeding cercopids, which often form mono- and multispecific aggregations in shared spittle (Halkka et al. 1977; McEvoy 1986). Apparently, possible negative effects are outweighed by the advantages of aggregation. Unlike in leafchewing 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 occidentalis 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 severe infestation of most central European deciduous trees by mesophyll-feeding leafhoppers 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 selection in Auchenorrhyncha, must be considered as weak. Whereas classical notions of interspecific competition usually referred to more apparent 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 interaction between the aphid Rhopalosiphum padi (L.) and the leafhopper Psammotettix alienus (Dhlb.), resulting in negative effects of an aphid extract on the leafhopper nymphs. Clearly, this field deserves further attention. It should also be taken into consideration that intra- and interspecific crowding effects by accumulating honeydew, e.g. of aphids, and excrements of mesophyll feeders are likely to have the potential to reduce densities by deteriorating physical conditions, e.g. adhesive properties of the plant surface, or by
Resource predictability
365
favouring pathogens. Experimental studies should focus on more discrete interactions, particularly diffuse competition (including inter-guild interactions), indirect effects of predation and parasitoid attacks mediated by plant semiochemicals, and perhaps soundtransmitting properties of the host, which may offer a considerable potential for explaining patterns in Auchenorrhyncha community structure. 6.1.7 Resource predictability Predictable resources, involving habitat persistence as well as temporal and spatial host plant apparency, are correlated with larger numbers of monophagous Auchenorrhyncha species, which prefer old-aged plants (notably trees) or plants forming large stands (notably graminoids, Urtica dioica) (see chapters 5.1.3.2; 6.1.2). Conversely, most unpredictable habitats select for host generalists (see the more detailed discussion in section 6.2.3). 6.1.8 Neural constraints Concerning the prevalence of monophagy in Auchenorrhyncha, many of these concepts fail to offer a satisfactory explanation. While chemical plant defence is likely to play a role in some broadly avoided groups, like Brassicaceae and Boraginaceae, the scarcity of potentially noxious compounds in phloem and xylem sap suggest alternative mechanisms. In particular, one of the major questions raised in this book is the high proportion of host specialists on graminoid plants, which are generally thought to be poor in secondary compounds (Frohne & Jensen 1998). None of the previous chapters offered the potential to explain, why stems or leaves of grasses and sedges, which are almost indistinguishable for humans, attract different and highly specific insect herbivores. A potentially powerful explanation is the neural hypothesis proposed by Bernays (1998, 1999a), although experimental confirmation for Auchenorrhyncha has not yet been presented. Accordingly, polyphagous grasshoppers, whiteflies and aphids are significantly less efficient in foraging, ovipositing and probably escaping from predators, compared to specialists, since they have to cope with ambiguous stimuli from various potential host plants (Bernays 1996, 1998, 1999b; Bernays & Funk 1999). Due to the limited neural capacity of processing and analyzing these stimuli, the effects of plant chemistry on host choice would rather be a facilitation and stimulation instead of a restriction or inhibition. The principal autonomy of the evolution of ecological traits in herbivorous insects from plant properties was also stressed by Schoonhoven et al. (1998). This hypothesis is indirectly supported by experimental evidence from breeding of monophagous species of the delphacid genera Muellerianella (Booij 1982) and Ribautodelphax (Bieman 1987b), which could successfully reproduce over several generations on several grasses, which have never been found to be the host in the field. 6.1.9 Regional monophagy Regional monophagy has been found in the delphacid Hyledelphax elegantula (Boh.), which feeds on numerous grasses (including Calamagrostis arundinacea, Brachypodium pin-
366
Life strategies
natum, Holcus mollis, Molinia caerulea, Deschampsia flexuosa) in the lowlands of southern and middle parts of Germany. At higher altitudes and in the north German plain, however, 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 Germany. 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 specialists on Phragmites australis additionally attack the giant grass Arundo donax in southern 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 Germany. However, feeding data in southern Europe are rather scarce. The possible mechanism is a reduced acceptance of potential host plants towards the border of the insect’s range, ultimately allowing reproduction only on a single host species, whereas other hosts grow in microclimatically and otherwise unsuitable sites. Regional monophagy is also reported from British populations of the swallowtail butterfly Papilio machaon L. (Strong et al. 1984). 6.1.10 Endophytic fungi Finally, toxins produced by endophytic fungi might play a role for the unequal Auchenorrhyncha infestation of the closely related grasses Brachypodium sylvaticum and B. pinnatum. The latter supports at least 11 substantiated feeders, 4 of which are monophagous, whereas the former is entirely unattacked. Recently, Brem & Leuchtmann (2001) found that an endophytic fungus specific to B. sylvaticum may reduce noctuid herbivory through fungal alkaloids. In contrast, B. pinnatum is only rarely infected by another fungus species (A. Leuchtmann, pers. comm.). These organisms are widespread, particularly in grasses, and may turn out to offer a considerable potential in explaining differential attack in sap-sucking insects on grasses (Clay 1990, 1996, 1997; Saikkonen et al. 1999).
6.2 Dispersal 6.2.1 Wing length Wing length is a major determinant of flight capability, and hence, dispersal and population dynamics (e.g. Denno & Roderick 1990; Johnson 1969; Taylor 1985). Macroptery is considered to be the primitive condition in Auchenorrhyncha and other Hemiptera (Hennig 1969), but wing dimorphism, subbrachyptery or brachyptery prevail in several groups of both Fulgoromorpha and Cicadomorpha. Brachypterous individuals in planthoppers and other insect groups have been shown to start reproducing earlier
Wing length
367
and to have a higher fecundity than macropterous individuals, because they do not spend resources for the development of the wing apparatus (e.g. Denno & Roderick 1990; Denno et al. 1991; Roff 1990; Andersen 1997). Flight capability, but not necessarily flight activity, can be expressed as the relation of fore and hind wing length, of hind wing and abdomen length or of wing surface area and body weight. An overview of wing length distribution in central European Auchenorrhyncha, however, is complicated by the fact, that only few data are published, particularly 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 knowledge, a simple distinction between macroptery and wing dimorphism is the most reliable 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 dimensionality, geographic distribution (altitude and latitude, in particular), parthenogenesis, gender 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 Auchenorrhyncha 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, macropters 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 Deltocephalinae, the genera Fieberiella, Grypotes, Japananus, Opsius, Balclutha, Sagatus, Platymetopius, Idiodonus, Colladonus, Lamprotettix, Allygus, Allygidius, Speudotettix, Hesium, Thamnotettix, Pithyotettix, Perotettix and Colobotettix are exclusively long-winged. Prevailing macroptery in arboreal Hemiptera was already demonstrated and discussed for the British fauna, and was explained by the fact that woody plants are architecturally 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 brachypterous insects, and consequently, the possibility of heavy population losses due to epigeic predators, and missing the proper host when climbing up again. The latter phenomenon can be found in tree eclector catches containing numerous nymphs of species being monophagous on other trees, perhaps due to nymphal inability of host recognition through the thick bark of basal trunks. In contrast, wing dimorphism prevails in species living among herbaceous vegetation. In these cases, macroptery does occur at least in a low proportion of the population and may increase, if densities are high or if host plant quality deteriorates (see Denno 1976; Denno et al. 1991). In general, males are often longer-winged than females, for
368
Life strategies
instance in many species of Deltocephalinae, where macroptery in male is often combined with female subbrachyptery. Reduction is most pronounced in epigeic species, with brachyptery prevailing in Delphacidae, except in Stenocranus, Laodelphax striatella (Fall.) and Javesella pellucida (F.), as well as in males of reed-dwelling species, like Delphax, Euides and Chloriona. Females of the latter are the only brachypterous Auchenorrhyncha in central Europe, which spend their whole life cycle high above the ground. Among Cicadomorpha, brachyptery is found in Ulopa, Errhomenus, Doratura and a few others, all of which are usually found on the soil surface. For the bulk of brachypterous Auchenorrhyncha species, it can be concluded that they show epigeic habits. As phytophages, they are unlikely to be limited by shortage of food, since much of the plant biomass in many ecosystems is left for consumption to detritivores (e.g. Hairston et al. 1960; White 1993). Thus, mobility is less important, except if habitat conditions deteriorate. Moreover, phloem- and xylem-feeding species do not consume or destroy living plant biomass and may potentially spend their whole life on a single plant stem. Thus, selective pressure for higher reproduction rates may usually be stronger than for macroptery and dispersal ability. Among holometabolous insect clades, only the Siphonaptera and the Mecoptera, notably the Boreidae, show a high degree of secondary wing reduction (Roff 1990). Like in brachypterous Auchenorrhyncha, they live in a more or less two-dimensional habitat without food limitation. In most other groups of terrestrial Holometabola, however, adults live in the tree or shrub layer or among tall herbaceous vegetation. In those groups utilizing epigeic resources (e.g. in saprophagous Diptera and predatory Coleoptera), food is often scattered, unpredictable or mobile. Hence, macroptery in insects has been suggested to be the crucial adaptation for the utilization of taller vegetation and predacious life habits (Hennig 1969; Snodgrass 1958; Swain 1978). (ii) Habitat persistence Pioneer species, like Laodelphax striatella (Fall.), Javesella pellucida (F.) and some species of Macrosteles, are the most striking exception to the rule of prevailing brachyptery in two-dimensional habitats (see section 6.2.3; Table 35). They show maximum densities in disturbed or unpredictable habitats, like high-intensity meadows and pastures, ruderal sites and temporarily dry banks near standing or running water (Novotný 1994a; Achtziger & Nickel 1997). Even in wing-dimorphic species, brachypters are rare, being usually confined to habitats, where disturbance dates far back, e.g. old fallows or abandoned fields. Novotný (1995) discussed prevailing macroptery in temporary habitats, combined with high dispersal ability, wide host plant range, wide geographic distribution and increased annual generation numbers in context with the r and K concept (see Pianka 1970) and the C-S-R (competitive, stress-tolerant, ruderal) system (Grime 1979), and suggested the term ‘colonization syndrome’ for the combination of these trails. It should be noted, however, that Auchenorrhyncha communities on highly disturbed alpine river banks, which are subject to frequent flooding and translocation, are characterized by a high proportion of host specialists, monovoltine species and European or even Alpine endemics (Nickel 1999a). (iii) Alpine habitats Increased brachyptery in Auchenorrhyncha is evident above the alpine tree line, where habitat dimensionality is reduced. The same should be true in higher geographic lati-
Migration
369
tudes. For further conclusions on the relationship of wing reduction and geographic altitude and latitude, a careful analysis of faunistic data (e.g. Günthart 1987a, 1997; Leising 1977; Ossiannilsson 1978, 1981, 1983; Wagner & Franz 1961) along a continuous gradient is required. Hence, the prevalence of brachyptery in alpine and tundra habitats 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 delphacid 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 brachypterous) have been studied so far. Evidence for a correlation of parthenogenesis and flightlessness was demonstrated and discussed by Bell (1982). As for Auchenorrhyncha, more karyological studies are needed, but the current impression is, that parthenogenesis 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 paurometabolous 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, smallscale migration can be performed by the relatively mobile nymphs, and – if habitat conditions 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, Cicadidae 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 controlled by synoptic weather conditions, thus creating the basis for predicting the occurrence 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 southwestern 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, Errastunus ocellaris (Fall.) was found to fly only in the first generation, and also Hyledelphax elegantula (Boh.), Streptanus sordidus (Zett.), Jassargus pseudocellaris (Fl.) and Arthaldeus 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 potential 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 species (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 completely altered vegetation. Funke & Witsack (1998) recorded altogether 141 species in surface coal mines of eastern Germany, equalling one third of the regional species diversity. 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 northern 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 planthopper, 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 abundant in most of the French traps, accounting for 42% of the total catch of these sites.
371
Migration
Surprisingly, both Macrosteles sexnotatus (Fall.) and M. laevis (Rib.), which are considered 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 remaining species preferred grassland habitats. From Germany, there is only a single small-scale study on Auchenorrhyncha dispersal 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 dominated (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.), Allygus 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 considerable potential for dispersal is also likely for larger species of Cixiidae, Cercopidae, Idiocerinae and Deltocephalinae.
Species
Rennes
Angers
Caen
Poitiers
Orléans
Versailles
Lens
Auxerre
Montpellier
Reim s
Valence
Colm ar
Silw ood 1970
Silw ood 1971
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)
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
0
6.3
0
0
0
0
0
0
23.8
0
4.7
0.3
0
0
1.1
4.1
70.3 57.6 82.5
45
0
91.5
0.2
0.1
0
1.5
1.8
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 Empoasca spp.
Toya propinqua (Fieb.) Javesella pellucida (F.)
13.7 21.9
J. dubia (Kbm .)
3.3
18.7
7.9
6.9
M acrosteles laevis (Rib.)
0
0
0
M . sexnotatus (Fall.)
0
0
0
Others Total Total (n individuals)
5.6
1860 347
* = could not be related to the total
177
0.1
2.1
56.7 73.4
-*
-*
1.8
3.5
2.6
8.4
10
4.4
10.6
9,8
-*
-*
0
0
0
0
0
0.2
0
0.1
1
6.4
8.3
0
0
0
0
0
0
0.1
0
0
27
25.5
20.9 15.5
23.3 20.9
0.3
22.4 12.8
12
10.6 17.1
2.6
13.7 40.1
950
444
876
666
316
1.4
14.1 12.5 44.2 53.4
429 1455 1892 1321 1276 1090
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., usually 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 scutellaris (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 higher 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 mowing (Andrzejewska 1979a; Novotný 1994a; Achtziger & Nickel 1997). Although exclosure experiments are still to be done to asess survival rates and subsequent immigration, the herbaceous 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 surrounding 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 successional stages, but densities are usually higher in other habitats, notably grassland, rocky
Colonization
373
Table 35: Auchenorrhyncha species of early successional and disturbed habitats and their life history traits. Diet width: po = polyphagous, o2 = 2nd degree oligophagous, o1 = 1st degree oligophagous, m2 = 2nd degree monophagous, m1 = 1st degree monophagous (see Table 4 for definitions). Substrate: P = phloem, X = xylem, M = mesophyll. Wing length: d = wing-dimorphic, m = macropterous.
Species
Diet w id th
Substrate
Wing length
Overw intering stage
Voltinism
Geographic range
Pioneer species Laodelphax striatella (Fall.) Javesella pellucida (F.) Empoasca pteridis (Dhlb.) Z yginidia scutellaris (H .-S.) M acrosteles cristatus (Rib.) M acrosteles laevis (Rib.) M acrosteles quadripunctulatus (Kbm .) M acrosteles sexnotatus (Fall.) M acrosteles viridigriseus (Ed w .) Ophiola decumana (Kontk.) Psammotettix alienus (Dhlb.) Psammotettix confinis (Dhlb.)
po? po? po o1 po po po? po o2? o1? o1 o1
P P M? M P P P P P P P P
d d m m m m m m m m m d
nym ph nym ph egg? ad ult egg egg egg egg egg egg egg egg
2 2 2? 1? 2 (1-?)2 2 2 2 2 2 2
Palearctic Palearctic Western Palearctic Med iterranean Eurosiberian Palearctic? Palearctic? Palearctic European Eurosiberian? Palearctic Eurosiberian
Specialists of river banks M yndus musivus (Germ .) Pentastiridius beieri (W.Wg.) Pseudodelphacodes flaviceps (Fieb.) Javesella stali (Metc.) Opsius stactogalus Fieb. M imallygus lacteinervis (Kbm .) Psammotettix unciger Rib. Errastunus leucophaeus (Kbm .)
m 2? o2? m1 m1 o1 m 1? m 1? ?
P P P P P P P P
m m d d m d m? d
nym ph nym ph nym ph nym ph egg egg egg egg
1 1 2 2 1 1 1? 2?
Western Palearctic European-alpine Western Palearctic Siberian Western Palearctic European-alpine Alpine Alpine
Species of d isturbed habitats A siraca clavicornis (F.) Dicranotropis hamata (Boh.) Toya propinqua (Fieb.) Javesella obscurella (Boh.) Ribautodelphax albostriata (Fieb.) Dictyophara europaea (L.) M egophthalmus scanicus (Fall.) A naceratagallia venosa (Geoffr.) A phrodes makarovi Zachv. A noscopus albiger (Germ .) N eoaliturus fenestratus (H .-S.) M acrosteles frontalis (Scott) M acrosteles horvathi (W.Wg.) M acrosteles lividus (Ed w .) Deltocephalus pulicaris (Fall.) Doratura homophyla (Fl.) Limotettix striola (Fall.) Euscelidius schenckii (Kbm .) Euscelidius variegatus (Kbm .) Euscelis incisus (Kbm .) Euscelis lineolatus Br. Psammotettix nodosus (Rib.) A rthaldeus pascuellus (Fall.)
po o1 o1? o1? m1 po o1 o2? po o1? m 2? m2 m2 m 2? o1 o1 o1? po po o2 o2 o1 o1
P P P P P P P P P P P P P P P P P P P P P P P
d d d d d m d d d d d m m m d d m m m d d m d
ad ult nym ph nym ph nym ph nym ph egg egg egg egg egg ad ult egg egg egg egg egg egg egg egg nym ph/ egg egg egg egg
1 1-2 2? 2 2 1 1 1 1 1 (1-?)2 (1-?)2 1-2 2 1-2 2 (1-?)2 1 1 1-2 1 2 2
Palearctic Palearctic Cosm opolitan Palearctic Western Palearctic Western Palearctic Med iterranean Eurosiberian? European? European? Palearctic Eurosiberian Eurosiberian Siberian Palearctic Palearctic Palearctic Western Palearctic Western Palearctic Palearctic Med iterranean? European Eurosiberian
374
Life strategies
slopes or salt marshes. Principally, however, this list is rather arbitrary, since any Auchenorrhyncha species may turn up in any habitat in small numbers, depending on the distance 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 river banks, where just the opposite is true. A third group of species, which prefer disturbed 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 experiments on newly established study plots. After a single field experiment at Silwood (Britain), it has been suggested that selection in early successional habitats favours species overwintering in the adult or nymphal 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 Zyginidia 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 uncertain, although there is weak evidence for species proportions shifting from overwintering as nymph towards overwintering as adult, with ongoing succession.
6.3 Life cycle 6.3.1 Voltinism Annual generation numbers have been published for numerous species (Leising 1977; Müller 1957, 1978, 1981, 1984a; Peter 1978, 1981; Remane 1958, 1987; Schiemenz 1969, 1987, 1988, 1990; Schiemenz et al. 1996; Strübing 1960; Witsack 1985 and others). The current information on the voltinism of the Auchenorrhyncha fauna of Germany has been compiled by Nickel & Remane (2002). The most reliable data have been obtained by laboratory rearings with manipulation of length of photoperiod and temperature. This tool, however, allows the identification of the mechanisms, but not necessarily the voltinism in the field. The latter is best studied by quantitative or semiquantitative sampling every 2 to 4 weeks throughout the season. This allows a clear documentation of the sequential appearance of nymphs, †† and ‡‡ (in this order) of each generation. Uncertainties remain, if late nymphs are found, which may be-
375
Voltinism
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 voltinism 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 European 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 September 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 maturity peak before August. Furthermore, hygric aestivation quiescence, extending the interval 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 confirmation), 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 conTable 36. Annual generation numbers of the Auchenorrhyncha species of Germany (1?/2? = probably mono/bivoltine; 1(-2?)/2(-3?) = usually mono-/bivoltine, perhaps bi-/trivoltine after early spring or in southern regions; 1-2 = mono- or bivoltine, depending on regional, altitudinal or weather conditions; (1-?)2 = usually bivoltine, perhaps monovoltine in higher altitudes or in northern parts)
Generation num ber
Auchenorrhyncha total
Fulgorom orpha
Cicad om orpha
n
%
n
%
n
%
1
343
55,3
90
62,1
253
53,3
1?
55
8,9
8
5,5
47
9,9
1(-2?)
5
0,8
0
0
5
1,1
1-2
25
4,0
12
8,3
13
2,7
(1-?)2
23
3,7
4
2,8
19
4,0
2
126
20,3
29
20,0
97
20,4
2?
26
4,2
2
1,4
24
5,1
2(-3?)
8
1,3
0
0
8
1,7
<1 Total
9
1,5
0
0
9
1,9
620
100
145
100
475
100
376
Life strategies
trol and a conservative selective force, acting both on Fulgoromorpha and Cicadomorpha (see below). However, there is evidence that at least 53 species may have either one or two generations 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 annual 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, wintering 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 univoltine at least from the montane belt upwards (see Leising 1977). Variation in voltinism is also influenced by geographical latitude. Kontkanen (1953) showed that Megadelphax sordidula (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.), Zyginidia pullula (Boh.) and others have been reported to show up to 4 generations in Mediterranean 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 (‘probably 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. Unfortunately, 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 reduction 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 fecundity increases and the mortality is low at least in older instars (Karban 1986). This nutritional 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, probably feed on phloem sap. Apart from geographic and climatic conditions, habitat disturbance is likely to be another important factor for the selection of voltinism, favouring bi- and polyvoltine species, 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 nearnatural 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 comparison between different sites, a correlation between annual generation numbers and certain habitat conditions (e.g. disturbance, successional stage and geographical position) 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 animals in temperate latitudes. It is of major significance not only for the escape from adverse climatic conditions, but also for resource utilization and intraspecific synchronization (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 (Migration)
Dormancy
Consecutive dormancy Quiescence
Oligopause
Prospective dormancy Parapause
Eudiapause
Hyperpause
Fig. 15: Dormancy phenomena in arthropods, after Müller (1970, 1992) and Witsack (1981, 1985, 2002)
Emancipation includes all kinds of responses to adverse environmental conditions. The organism may migrate to more favourable localities (recession), or delay its development (dormancy). For purpose of hibernation, dormancy plays the most important role in central European Auchenorrhyncha. In addition, some species, e.g. Empoasca spp., Zygina spp., Balclutha spp., show a small-scale migration to other food plants or other strata (see section 6.2). Consecutive dormancy is purely facultative, and is induced only after the onset of adverse conditions. In quiescence, induction as well as termination happen immediately after environmental conditions, usually temperature or moisture, have changed. Examples include embryos and occasional adults of Anakelisia fasciata (Kbm.) and nymphs and occasional embryos of Euscelis incisus (Kbm.), both species overwintering in thermic quiescence. Further, thermic quiescence plays a major role – also among other insect groups – as a secondary dormancy after the termination of oligopause, parapause or eudiapause, e.g. in Philaenus spumarius (L.), Turrutus socialis (Fl.) and Muellerianella brevipennis (Boh.). In oligopause, the response happens only gradually, often induced and terminated cumulatively by the same factor. Examples include Euides basilinea (Germ.) and Chloriona smaragdula (Stål). Noteworthy is the fact that there are transitions towards quiescence (quiescent oligopause, not yet found in Auchenorrhyncha) and diapausic oligopause, found in Muellerianella brevipennis (Boh.). Prospective dormancy starts before the onset of adverse conditions. Parapause is ontogenetically triggered, but terminated by an external factor, e.g. cold, and often followed by thermic quiescence, which in turn can be immediately terminated after the onset of spring. Primary parapause, e.g. found in the oocytes of Stenocranus minutus (F.), St. major (Kbm.) and Mocydia crocea (H.-S.), can be experimentally prevented by exposure to terminating factors. Secondary parapause is rather fixed and will definitively progress. It is often found in monovoltine species overwintering in the embryonic stage, e.g. in Agallia brachyptera (Boh.), Anaceratagallia venosa (Geoffr.) and Elymana sulphurella (Zett.), or potentially bivoltine, e.g. in Philaenus spumarius (L.), Cicadella viridis (L.) and Erzaleus metrius (Fl.). Eudiapause is externally controlled and usually induced by photoperiod, but terminated by low temperature. In 4 out of 5 cases studied, species were bivoltine, and overwintering took place in the embryonic stage of eggs laid by the second generation. Examples include Macrosteles sexnotatus (Fall.), Turrutus socialis (Fl.),
379
Dormancy
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 threeyear 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 overwintering 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 planthoppers, it is 78.4% in leafhoppers. Conversely, only 4.3% of leafhoppers, but 67.5% of Table 37. Overwintering stages of the Auchenorrhyncha species of Germany
Overw intering
Auchenorrhyncha total
Fulgorom orpha
Cicad om orpha
stage
n
%
n
%
n
%
Egg
379
61,1
27
18.6
352
74.1
Egg?
24
3.9
2
1.4
22
4.6
N ym ph
110
17.7
89
61.4
21
4.4
N ym ph?
7
1.1
7
4.8
0
0
Ad ult
82
13.2
18
12.4
64
13.5
Ad ult?
13
2.1
2
1.4
11
2.3
Mixed
5
0.8
0
0
5
1.1
Total
620
100
145
100
475
100
380
Life strategies
planthoppers, overwinter as nymphs. Differences are only slight in proportions of species wintering as adults, whereas overwintering in more than one stage is found in the Cicadomorpha only. The present data do not allow an explanation for this difference. Thus, taxon-specific constraints may be involved. There is a number of ambiguous cases among those species marked as uncertain (‘?’). For instance, during mild and even moderately cold winters, adults of both Eupteryx melissae Curt. and E. florida Rib. have been found to survive. The same is true for Anakelisia fasciata (Kbm.), Megamelus notula (Germ.) and probably some other species. Ribautiana tenerrima (H.-S.) and R. debilis (Dgl.) are frequently found until the onset of frost. The ecological significance of occasional overwintering as adults in these species has not been studied, although most individuals probably die without reproduction. In Mediterranean regions, where occasional warm winter days may allow feeding and other activities, overwintering as adults may play a more important role. Thus, a number of Mediterranean species reaching central Europe, clearly favour this overwintering stage, for instance most species of Tettigometridae, Hauptidia, Arboridia and Mocydiopsis. Müller (1992) reviewed dormancy and other life cycle traits in arthropods. He listed numerous examples of allopatric and sympatric variation of life cycles and discussed possible implications on ecology and speciation. Differences in overwintering stages between Fulgoromorpha and Cicadomorpha have been interpreted as a temporal niche segregation. For nymphal development, which is supposed to be the most crucial period with maximum demand of nutrients, the Fulgoromorpha utilize much more the autumn and early spring, whereas the Cicadomorpha rather utilize the late spring and early summer. Müller (1978), Peter (1981) and Waloff (1979, 1980) discussed niche partitioning among grassland Auchenorrhyncha, assuming that potential interspecific competition should be markedly reduced by temporal segregation. Other authors suggested an adaptation of Auchenorrhyncha phenology to seasonal fluctuations of plant nitrogen content, which is highest in spring and autumn, but relatively low in summer (Hill 1982; McNeill & Southwood 1978), as was found in aphids (Dixon 1985; Risebrow & Dixon 1987; Wellings et al. 1980). In grassland communities of Auchenorrhyncha in the Leutra valley (Thuringia) and Silwood (England), however, peaks of total individual numbers were subject to conspicuous annual variation (Waloff 1979; Müller 1978). The reduction of densities during summer in some years were rather explained by drought instead of response to nutrient conditions. Peter (1981) showed that the maturity peaks of the dominating species in the Leutra valley were evenly spread over the whole season, ranging from the beginning of April until the end of August. As a consequence, total individual numbers reached their maximum in early and high summer. The proportion of monovoltine species wintering in the egg stage, and reaching their maturity peak in June and July was rather high. Thus, phenological adaptations to nutrient contents of plants may occur in some species of Auchenorrhyncha, but the majority shows maturity peaks in early and mid summer, when nitrogen contents in above-ground parts of plants are rather low, suggesting that temperature has a much stronger impact on phenology than nitrogen fluctuations. Hill (1982) demonstrated that the leafhopper Recilia coronifer (Marsh.), which develops in summer, shows a different nitrogen utilization strategy and uric acid storage, compared with two species developing in spring. Further research in this field is needed to detect the mechanisms enabling summer species to compensate the reduced nitrogen content of their hosts.
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7 Summary
This work presents an overview of the fauna of leafhoppers and planthoppers (Hemiptera, Cicadomorpha and Fulgoromorpha = Auchenorrhyncha) of Germany and their life strategies, with special reference to insect – plant relationships. The data base included 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 methods, 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 altitudinal 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 differences between main plant clades and families, between different vegetation layers and different plant substrates. A short overview is given on pests on cultivated plants. Further, 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 Fagaceae, Betulaceae, Salicaceae and Rosaceae are attacked by approximately 50 species, respectively. Some highly diverse plant groups are not or only rarely attacked, nota-
382
Summary
bly the Caryophyllaceae, Brassicaceae, Apiaceae and Scrophulariaceae. Due to much differing plant species numbers per family, the ratio of Auchenorrhyncha species : plant species per plant family is high on woody plants, but relatively low on graminoids. Thus, herbivore species richness is highest on plants, which are rich in species, or biomass, or both. It can be concluded that apparent plants attract highest numbers of herbivores. Roughly two third of the total number of Auchenorrhyncha species live in the herb layer, but only one third lives in the canopies of woody plants. Thus, despite the fact that central Europe is a woodland country, where biomass is thought to be more concentrated in the canopy layer, speciation in Auchenorrhyncha was more pronounced in the herbacous vegetation. Ten percent of the species are vertical migrants, i.e. they perform an obligate or facultative migration from one stratum to another. Some species groups are particularly interesting due to their endogeic nympal life habits, raising the question for the evolutionary significance of a diverging morphology and ecology of immature and adult stages in hemimetabolous versus holometabolous insects. More than 70% of the total number of Auchenorrhyncha species feed on phloem, which is considered as the primitive substrate of Hemiptera. Secondarily, some groups have tapped xylem, mesophyll and fungi mycelium. Species numbers of xylem feeders on plant families are more strongly correlated with plant species number per family, than all feeding groups together, providing evidence that plant chemistry of xylem may play only a negligible role for host choice. Thus, the Caryophyllaceae and Brassicaceae, which are avoided by phloem and mesophyll feeders, are attacked only by a few species of xylem feeders, which generally tend to be more polyphagous. Mesophyll feeders, which account for one fourth of the total fauna, show a high proportion of arboricolous species. Feeding on leaf parenchyma is only found in the Typhlocybinae and has probably evolved along with the rise of angiosperm trees, but out of this group, a number of species has secondarily performed a host shift back to the herbaceous vegetation, and colonized primitive or even toxic plants. Despite widespread crop damage by feeding and transmission of plant diseases in more southern countries, Auchenorrhyncha play only a minor role as pests in central Europe, although a few recent cases are known, notably Psammotettix alienus (Dhlb.) transmitting wheat dwarf virus (WDV), Hyalesthes obsoletus Sign. transmitting grapevine yellows (FD), and Empoasca vitis (Göthe) causing ‘hopperburn-like symptoms’ in vineyards. Other examples are more localized and include Empoasca decipiens Paoli on greenhouse vegetables, and Eupteryx spp. on medical herbs and spices. Further, the Nearctic Graphocephala fennahi Young has been suspected to facilitate infection of ornamental Rhododendron by a noxious fungus causing bud burst. Invasion of further pest species from Mediterranean countries, such as Metcalfa pruinosa (Say) and Scaphoideus titanus Ball may occur in near future. In general, the relationship between plants and Auchenorrhyncha appears to be rather asymmetrical. Whereas plant apparency (and in particular, distribution, abundance and size) and plant chemistry are likely to be crucial factors for Auchenorrhyncha occurrence, reverse effects are apparently discrete or weak. These effects, however, may have been underestimated. On the one hand, vascular feeding does not remove living plant tissue, but, on the other hand, constant removal of resources and transmission of
Summary
383
pathogens (or at least facilitation of infection) certainly have the potential of causing damages and persistent weakening of hosts. Thus, even if direct effects may be insignificant, 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 sufficient 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 Urticaceae. 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 concluded, 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 radiation by host shift from one grass to another. Phragmites australis, the common reed, and Urtica dioica, the stinging nettle, are exceptional among herbaceous plants for their high Auchenorrhyncha species numbers, including 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 specialists 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 dominate in herbaceous vegetation, whereas all arboricolous species and vertical migrants are macropterous. However, one particular group of herb-dwelling species is also monomorphic macropterous, and is largely confined to disturbed habitats and early successional stages. In this group, most species are bivoltine and polyphagous, and their geographic ranges are large, whereas mesophyll-feeding and overwintering in the adult stage are underrepresented.
384
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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 overwinter 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 taken 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 diverse not only in Auchenorrhyncha, but also in other insect taxa such as Orthoptera, Diptera, Hymenoptera and Lepidoptera.
8 References
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Appendix
Index of Auchenorrhyncha genera and species Page numbers are given only for chapter 4. abbreviatus (Leth.), Allygidius 183 abdominalis (F.), Verdanus 222 abietinus (Fall.), Pithyotettix 195 abstrusus (Lnv.), Kybos 118 Acanthodelphax 53 Acericerus 100 achilleae Mity., Hephathus 91 Achorotile 42 acuminatus (F.), Evacanthus 109 Adarrus 216 f. adela (Fl.), Paraliburnia 47 adspersa (H.-S.), Eupteryx 144 adumbrata (C. Shlb.), Rhopalopyx 187 aemulans (Kbm.), Streptanus 204 aestuarina (Edw.), Aphrodes 103 affinis Fieb., Stiroma 41 affinis Nast, Empoasca 121 Agallia 91 f. Agalmatium 70 agrestis (Fall.), Stroggylocephalus 108 Aguriahana 151 albae W.Wg., Macropsis 84 albicans (Kbm.), Populicerus 99 albifrons (Fieb.), Mirabella 49 albifrons (L.), Anoscopus 106 albiger (Germ.), Anoscopus 106 albingensis W.Wg., Cicadula 189 albipennis (F.), Neophilaenus 74 albocarinata (Stål), Nothodelphax 53 albomarginatus Curt., Criomorphus 58
albomarginatus W.Wg., Psammotettix 213 albosignata (Dhlb.), Achorotile 42 albostriata (Fieb.), Ribautodelphax 62 albostriella (Fall.), Alebra 112 alces (Rib.), Ribautiana 138 Alebra 111-113 alemannicus W.Wg., Jassargus alpinus 219 alienus (Dhlb.), Psammotettix 212 allionii (Turt.), Batracomorphus 101 allobrogicus (Rib.), Jassargus 220 Allygidius 182 f. Allygus 181 f. alneti (Dhlb.), Alnetoidia 152 Alnetoidia 152 alni (Fall.), Aphrophora 76 alni (Schrk.), Oncopsis 82 alnicola (Edw.), Edwardsiana 127 alpestris W.Wg., Cixius 24 alpicola (Cer.), Zyginidia 153 alpicola W.Wg., Neophilaenus exclamationis 76 alpina Mel., Aphrophora 76 alpinus (Then), Jassargus 219 alpinus (W.Wg.), Anoscopus 107 alpinus (Zett.), Macrosteles 169 alta Walk., Ceresa 79 alticola Rib., Eupteryx 146 ampliata (W.Wg.), Edwardsiana 130 Anaceratagallia 92 f. Anakelisia 36 anceps (Germ.), Conomelus 38
Index of Auchenorrhyncha genera and species angulatus (Then), Psammotettix 211 angulosa (Rib.), Ribautodelphax 63 angusta Leth., Zygina 156 Anoscopus 106-108 antennalis (Hpt.), Errastunus 217 antoniae (Mel.), Dryodurgades 94 Aphrodes 103 f. aphrodoides Hpt., Thamnotettix 186 Aphrophora 76-78 apicalis (Fl.), Empoasca 122 appendiculata W.Wg., Oncopsis 82 Arboridia 160-162 arcuata Fieb., Cercopis 72 areatus (Stål), Pinumius 221 arenarius Rem., Arthaldeus 223 argentarius Metc., Athysanus 196 argus (Marsh.), Sardius 186 Arocephalus 208 f. artemisiae (Kbm.), Eupteryx 145 Arthaldeus 223 Artianus 206 Asiraca 29 assimilis (Fall.), Sorhoanus 224 assimilis (Sign.), Anoscopus 107 aterrimus (J. Shlb.), Neophilaenus lineatus 75 Athysanus 196 f. atomarius (F.), Allygidius 183 atra Hag., Tettigometra 66 atrata Fieb., Tettigometra 66 atricapillus (Boh.), Limotettix 199 atropunctata (Goeze), Eupteryx 142 attenuata (Germ.), Mocydiopsis 192 aubei (Perr.), Muirodelphax 52 aurantiacus (Forel), Cosmotettix 226 aurata (L.), Eupteryx 142 aureola (Fall.), Erythria 113 aurita (L.), Ledra 82 aurovittata (Dgl.), Lindbergina 137 austriaca (Metc.), Eupteryx 142 austriaca W.Wg., Anaceratagallia 92 austriacus W.Wg., Cixius 25 Austroasca 124 avellanae (Edw.), Edwardsiana 127 avellanae Edw., Oncopsis 83 Balcanocerus 101
Balclutha 167 f. barbata (Rib.), Edwardsiana 134 basilinea (Germ.), Euides 43 Batracomorphus 101 f. bavaricus Rib., Jassargus 220 beieri (W.Wg.), Pentastiridius 27 beieri W.Wg., Cixius 25 bellevoyei (Put.), Adarrus 217 bensoni (China), Verdanus 222 bergmani (Tull.), Edwardsiana 127 betulicola (W.Wg.), Kybos 119 bicarinata (H.-S.), Stiroma 41 bicincta (Schrk.), Aphrodes 104 bifasciata (Boh.), Zonocyba 140 bifasciata (L.), Planaphrodes 105 bilobum (Fieb.), Agalmatium 70 binotatus (J. Shlb.), Sonronius 174 bipunctata (Osh.), Kyboasca 124 bisignata (M. & R.), Fruticidia 163 bisonia Kopp & Yonke, Stictocephala 79 bohemani (Zett.), Diplocolenus 221 boica W.Wg., Balclutha 167 boldi (Scott), Gravesteiniella 51 borealis (J. Shlb.), Criomorphus 58 borussicus W.Wg., Cixius 25 brabantica W.Wg., Macropsis 89 Brachyceps 66 brachyptera (Boh.), Agallia 91 brachypterus Fieb., Errhomenus 109 brevipennis (Boh.), Muellerianella 51 brevis (H.-S.), Goniagnathus 164 brunnea Mel., Eurysella 40 bubalus (F.), Ceresa 79 butleri (Edw.), Kybos 117 calamagrostidis Rem., Lebradea 224 calamagrostis Oss., Balclutha Calamotettix 226 calcarata Oss., Eupteryx 145 Calligypona 49 callosa (Then), Ossiannilssonola 126 calyculus (Cer.), Kybos 120 cambricus China, Cixius 25 campestris (Fall.), Neophilaenus 74 candidula (Kbm.), Edwardsiana 127 capnodes (Scott), Delphacodes 50
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432
Appendix
carneae W.Wg., Ulopa 80 carniolicus W.Wg., Cixius 24 carpini (J. Shlb.), Oncopsis 83 carri (Edw.), Fagocyba 125 caudatus (Fl.), Cosmotettix 225 centrorossica Zachv., Aphrodes 104 Centrotus 79 cephalotes (H.-S.), Psammotettix 213 Ceresa 79 Cercopis 72 cerea (Germ.), Macropsis 86 Chloriona 44-46 Chlorionidea 52 Chlorita 124 f., 228 Cicada 70 Cicadella 110 Cicadetta 71 Cicadula 189-191 Circulifer 166 citrinella (Zett.), Forcipata 116 Cixidia 64 f. Cixius 22-26 clavicornis (F.), Asiraca 29 clypealis (J. Shlb.), Paraliburnia 48 cognatus (Fieb.), Ebarrius 216 coleoptrata (L.), Lepyronia 74 coleoptratus (F.), Issus 69 Colladonus 180 collina (Boh.), Ribautodelphax 63 collina (Fl.), Eupteryx 146 collinus (Boh.), Mocuellus 227 Colobotettix 196 communis (Ferr.), Allygus 181 commutatus (Fieb.), Allygidius 182 concinna (Germ.), Eurhadina 140 concolor Fieb., Tettigometra 228 confinis (Dhlb.), Psammotettix 215 confinis (Reut.), Streptanus 204 confinis (Zett.), Cixidia 64 confinis (Zett.), Thamnotettix 194 confusa Lnv., Kelisia 34 confusus (Fl.), Populicerus 99 Conomelus 38 Conosanus 201 consobrina Curt., Agallia 92
contraria (Rib.), Emelyanoviana 114 cornicula (Marsh.), Ophiola 198 cornicula (Meusn.), Zyginidia 154 cornutus (H.-S.), Enantiocephalus 227 cornutus (L.), Centrotus 79 coronifer (Marsh.), Recilia 176 corticea Germ., Aphrophora 77 coryli Le Q., Alebra 112 coryli (Toll.), Alnetoidia alneti 152 Coryphaelus 166 Cosmotettix 225 f. costalis Mats., Aphrophora 77 costalis (Fall.), Cosmotettix 225 crassicornis (Panz.), Delphax 43 crataegi (Dgl.), Edwardsiana 128 Criomorphus 58 f. cristatus (Rib.), Macrosteles 169 crocea (H.-S.), Mocydia 192 cruenta (H.-S.), Fagocyba 126 cruentatus (Panz.), Idiodonus 180 cunicularius (L.), Cixius 23 curtisii (Fl.), Eupteryx 147 cuspidata (F.), Eupelix 103 cyane (Boh.), Erotettix 174 cyclops Mats., Eupteryx 145 dahlbomi (Zett.), Sonronius 175 dalei (Scott), Scottianella 56 dealbata Cer., Empoasca 123 debilis (Dgl.), Ribautiana 138 decemnotata R., Eupteryx 148 decempunctata (Fall.), Linnavuoriana 136 decimusquartus (Schrk.), Rhytidodus 94 decipiens Paoli, Empoasca 122 decumana (Kontk.), Ophiola 198 dehneli Nast, Conomelus lorifer 38 Delphacinus 39 Delphacodes 50 Delphax 43 Deltocephalus 175 denticauda (Boh.), Acanthodelphax 53 depressa Fieb., Tettigometra 66 Dicranotropis 54 f. Dictyophara 65 digitatus (Rib.), Kybos 119 Dikraneura 115
Index of Auchenorrhyncha genera and species dilutior (Kbm.), Thamnotettix 195 diminuta Rib., Aphrodes 104 Diplocolenus 221 discolor (Boh.), Javesella 60 discolor (J. Shlb.), Ederranus 204 dissimilis (Fall.), Ommatidiotus 69 distincta (Fl.), Nothodelphax 54 distinguenda (Kbm.), Hauptidia 152 distinguendus (Fl.), Jassargus 218 distinguendus (Kbm.), Euscelis 202 distinguendus (Kbm.), Tremulicerus 98 distinguendus Kbm., Cixius 23 Ditropis 39 Ditropsis 47 divergens Kbm., Dicranotropis 54 diversa (Edw.), Edwardsiana 128 Doliotettix 196 domino (Reut.), Hesium 194 douglasi (Edw.), Fagocyba cruenta var. 126 Doratura 177-179 dorsata Edw., Chloriona 44 dorsatum (Ahr.), Haematoloma 73 Dryodurgades 94 dubia (Kbm.), Javesella 61 dubius Oss., Psammotettix 214 dubius W.Wg., Cixius 26 duffieldi (Le. Q.), Anoscopus 107 dumosa (Rib.), Chlorita 124 Ebarrius 216 Ederranus 204 Edwardsiana 127-136, 229 elaeagni Em., Macropsis 89 elegans (Fl.), Metidiocerus 96 elegantula (Boh.), Hyledelphax 48 elongata W.Wg., Rhopalopyx 187 Elymana 188 Emelyanoviana 114 emmeae Fieb., Trigonocranus 27 Empoasca 121-123 Enantiocephalus 227 Endria 176 erecta (Rib.), Arboridia 160 Erotettix 174 Errastunus 217 Errhomenus 109
Errhomenellus 109 Erythria 113 Erzaleus 227 Euconomelus 43 Euides 43 Eupelix 103 Eupterycyba 136 Eupteryx 142-150 Eurhadina 140 f. europaea (L.), Dictyophara 65 Eurybregma 41 Eurysa 40 Eurysella 40 Eurysula 40 Euscelidius 201 Euscelis 202 f. Evacanthus 109 excisa (Mel.), Unkanodes 46 excisus (Mats.), Psammotettix 214 exclamationis (Thnbg.), Neophilaenus 75 exigua (Boh.), Kosswigianella 55 exilis Horv., Doratura 177 extrusa (Scott), Muellerianella 52 Fagocyba 125 f. fairmairei (Perr.), Muellerianella 51 fallax Rib., Kelisia 33 fasciata (Kbm.), Anakelisia 36 fenestratus Em., Delphacodes 59 fenestratus (H.-S.), Neoaliturus 165 fennahi Young, Graphocephala 111 fieberi (Edw.), Macrosteles 169 fieberi (Scott), Megamelodes 49 Fieberiella 163 f. filicum (Newm.), Eupteryx 150 flammigera (Geoffr.), Zygina 156 flava P. Löw, Chlorionidea 52 flaveola (Boh.), Paluda 186 flaveola (Fl.), Xanthodelphax 56 flavescens (F.), Edwardsiana 129 flavescens (F.), Empoasca 123 flaviceps (Fieb.), Pseudodelphacodes 59 flavicollis (L.), Oncopsis 83 flavipennis (Zett.), Notus 117 flavipes (Sign.), Ditropsis 47 flavopunctatus Mel., Errhomenellus 109
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flavostriatus (Don.), Anoscopus 107 flavovarius (H.-S.), Handianus 229 flavovirens (Gill. & Bak.), Lebradea 224 flori (Fieb.), Jassargus 219 flori (J. Shlb.), Cicadula 190 florida Rib., Eupteryx 147 florii (Stål), Fieberiella 163 Florodelphax 55 Forcipata 116 f. forcipata (Boh.), Javesella 62 forcipata (Fl.), Forcipata 117 formosus (Boh.), Metalimnus 208 forneri Hpt., Aphrophora salicis 77 franzi (W.Wg.), Zyginidia fratercula (Edw.), Edwardsiana 129 frisia (W.Wg.), Anaceratagallia 92 froggatti (Bak.), Edwardsiana crataegi var. 128 frontalis (H.-S.), Cicadula 191 frontalis (Scott), Macrosteles 170 frustrator (Edw.), Edwardsiana 130 Fruticidia 163 fuchsii (Kbm.), Tremulicerus 98 fulgidus (F.), Tremulicerus 98 fusca Fieb., Tettigometra 66 fuscinervis (Boh.), Macropsis 88 fuscovittatus (Stål), Stenocranus 37 fuscula (Zett.), Macropsis 89 Gargara 78 genistae (F.), Gargara 78 geometrica (Schrk.), Edwardsiana 130 germari (Zett.), Wagneripteryx 150 glandacea (Fieb.), Macropsis 89 glaucescens Fieb., Chloriona 45 Goniagnathus 164 graminea (F.), Macropsis 88 Graphocephala 111 Graphocraerus 183 gratiosa Dlab., Dicranotropis 55 gratiosa (Boh.), Edwardsiana 131 gravesteini Dlab., Delphacodes 56 gravesteini W.Wg., Macropsis 85 Gravesteiniella 51 grisea Hpt., Aphrophora 77 griseola Fieb., Tettigometra 67 griseombra Rem., Zygina 159
grisescens (Zett.), Macustus 196 Grypotes 164 guentharti Dlab., Edwardsiana 135 guttatus Fieb., Platymetopius 180 guttula (Germ.), Kelisia 30 guttulifera (Kbm.), Kelisia 33 gyllenhalii (Fall.), Coryphaelus 166 haagii Kbm., Jassus 166 haematoceps (M. & R.), Circulifer 166 haematodes (Scop.), Tibicina 70 Haematoloma 73 hagemini Rem. & Jung, Kelisia 31 haglundi (J. Shlb.), Megadelphax 46 halpina Rem. & Jung, Kelisia 32 hamata (Boh.), Dicranotropis 54 Handianus 197, 229 Hardya 185 hassicus Nikusch, Cixius haupti 25 haupti Dlab., Cixius 25 haupti W.Wg., Kelisia 31 haupti W.Wg., Macropsis 87 Hauptidia 152 f. helvolus (Kbm.), Psammotettix 213 henribauti Dlab., Platymetopius 179 Hephathus 91 herrichii (Kbm.), Idiocerus 96 Hesium 194 heydenii (Kbm.), Acericerus 100 heydenii (Kbm.), Eupteryx 143 heydenii Kbm., Cixius 24 hippocastani (Edw.), Edwardsiana 131 histrionicus (F.), Anoscopus 107 hofferi (Dlab.), Praganus 226 homophyla (Fl.), Doratura 178 horvathi (Then), Recilia 176 horvathi (W.Wg.), Macrosteles 170 horvathi W.Wg., Doratura 178 Hyalesthes 29 hyalinus (Osb.), Japananus 165 Hyledelphax 48 hyperici (H.-S.), Zygina 159 hypermaculata Rem. & Hlzg., Zygina 160 Iassus 102 Idiocerus 95 f. Idiodonus 180
Index of Auchenorrhyncha genera and species ignoscus (Mel.), Handianus 197 imitans (Rib.), Ribautodelphax 63 immaculatifrons (Kbm.), Eupteryx 145 impictifrons (Boh.), Laburrus 200 impressifrons (Kbm.), Metidiocerus 96 impressopunctata Duf., Tettigometra 65-69 impudica Horv., Doratura 178 impura (Boh.), Macropsis 88 incisa (Then), Wagneriala 116 incisus (Kbm.), Euscelis 202 inconstans (Rib.), Zygina 157 inermis W.Wg., Metropis 42 inexpectatus Rem., Psammotettix 211 infumatus (Hpt.), Neophilaenus 75 infuscata (J. Shlb.), Macropsis 86 inquinata (Rib.), Fagocyba cruenta var. 126 intercedens (Lnv.), Linnavuoriana 136 intermedia (Boh.), Cicadula 192 intermedia Rem., Mocydiopsis 192 intermedius Fieb., Cixius 23 interruptus (L.), Evacanthus 109 interstinctus (Fieb.), Ebarrius 216 interstitialis (Germ.), Artianus 206 intractabilis Kontk., Limotettix 198 intricatus (H.-S.), Phlepsius 184 irregulata Hpt., Kelisia 31 irroratus Lew., Batracomorphus 102 ishidai (Mats.), Edwardsiana 132 Issus 69 Japananus 165 Jassargus 218-220 Jassidaeus 38 Javesella 60 jucunda (H.-S.), Eupterycyba 136 juniperi (Leth.), Liguropia 114 Kelisia 30-35 kemneri (Oss.), Edwardsiana 229 kirschbaumi W.Wg., Eurhadina 140 koeleriae Zachv., Psammotettix 211 kolosvarensis (Mats.), Psammotettix 210 Kosswigianella 55 kozhevnikovi (Zachv.), Elymana 188 kratochvili (Lang), Arboridia 160 Kyboasca 124 Kybos 117-121
Laburrus 200 lacteinervis (Kbm.), Mimallygus 184 laeta H.-S., Tettigometra 66, 67 laetifica Metc., Tettigometra atra 66, 67 laeva (R.), Planaphrodes 106 laevis (Rib.), Macrosteles 170 lamellaris (Rib.), Edwardsiana 131 laminatus (Fl.), Populicerus 99 Lamprotettix 181 languidus (Fl.), Arocephalus 208 lanio (L.), Iassus 102 lanternae (W.Wg.), Edwardsiana 132 Laodelphax 47 larvatus (H.-S.), Balcanocerus 101 lasiocarpae Oss., Cicadella 110 latifrons (Kbm.), Metropis 42 Lebradea 224 Ledra 82 lelievrei (Leth.), Eupteryx 143 lepidus (Boh.), Euconomelus 43 leporinus (L.), Pentastiridius 28 leptosoma (Fl.), Florodelphax 55 Lepyronia 74 lequesnei W.Wg., Megamelodes 48 lethierryi (Edw.), Edwardsiana 131 leucophaea (Preyssl.), Tettigometra 68 leucophaeus (Kbm.), Errastunus 217 Liguropia 114 limicola (Edw.), Anoscopus 106 Limotettix 198, 199 f. limpidus (W.Wg.), Kybos 118 lindbergi (Lnv.), Kybos 119 Lindbergina 137 lineata (Perr.), Eurysa 40 lineatus (L.), Neophilaenus 75 lineolata (Horv.), Balclutha 167 lineolatus Br., Euscelis 202 Linnavuoriana 136 f. Litemixia 52 littoralis Kuntze, Doratura 179 lituratus (Fall.), Idiocerus 95 livens (Zett.), Stroggylocephalus 108 lividus (Edw.), Macrosteles 171 loewii (Then), Eurhadina 141 loginovae (Em.), Arboridia 161
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longicauda Rem., Mocydiopsis 193 longiceps (Kbm.), Arocephalus 208 longicornis Sign., Micrometrina 228 longipennis (Curt.), Stenocranus 36 longispinus W.Wg., Cixius nervosus 22 longiventris J. Shlb., Cicadula 189 lorifer Rib., Conomelus 38 ludus (Dav. & De L.), Kybos 120 lugens Germ., Ulopa 81 lugubrina (Boh.), Struebingianella 56 lugubris (Sign.), Jassidaeus 38 lunaris (M. & R.), Zygina 155 lunulatus (Zett.), Doliotettix 196 lurida (Fieb.), Eurysula 40 macrocephala Fieb., Tettigometra 68 Macropsidius 91 Macropsis 84-91 Macrosteles 169-174 maculatus Rib., Allygus 182 maculiceps Boh., Deltocephalus 175 maculosus (Then), Macrosteles 171 Macustus 196 major (Kbm.), Platymetopius 179 major (Kbm.), Stenocranus 37 major Hpt., Paramesus 206 major Uhl., Aphrophora 76 makarovi Zachv., Aphrodes 104 mali (Edw.), Alnetoidia alneti 152 maligna (Walsh), Kyboasca 235 manderstjernii (Kbm.), Erythria 113 marginata (H.-S.), Macropsis 86 marginatus (Kbm.), Streptanus 205 marginicollis (Spin.), Cixidia 65 maritimus (Perr.), Psammotettix 209 martigniaca Cer., Edwardsiana candidula 127 Megadelphax 46 Megamelodes 48 f. Megamelus 38 megerlei (Fieb.), Macropsis 90 Megophthalmus 81 melanopsis (Hardy), Hardya 185 melissae Curt., Eupteryx 148 mendax (Fieb.), Macropsis 89 Mendrausus 220 meridionalis Bonf., Japananus 165
mesomelas (Boh.), Delphacinus 39 mesopyrrhus (Kbm.), Tremulicerus 98 Metalimnus 207 f. Metcalfa 235 Metidiocerus 96 metrius (Fl.), Erzaleus 227 Metropis 42 micantula (Zett.), Micantulina 115 Micantulina 115 microcera Vilb., Macropsis 85 Micrometrina 228 Mimallygus 184 minima (J. Shlb.), Wagneriala 116 minima Rib., Kelisia 34 minor (Kbm.), Neophilaenus 76 minutus (F.), Stenocranus 37 Mirabella 49 Mitricephalus 66 mixtus (F.), Allygus 181 mocsaryi (Horv.), Zyginidia 153 Mocuellus 227 Mocydia 192 Mocydiopsis 192 f. modestus Scott, Allygus 182 moestus (Boh.), Criomorphus 58 mollicula (Boh.), Emelyanoviana 114 monoceros Rib., Kelisia 35 montana (Horv.), Dicranotropis 55 montana (Scop.), Cicadetta 71 montanus Geb. & Szwed., Speudotettix 194 monticola Rem., Mocydiopsis 193 monticola W.Wg., Neophilaenus exclamationis 76 morbillosus (Mel.), Colobotettix 196 mucronatus (Rib.), Kybos 120 Muellerianella 51 f. Muirodelphax 52 mulsanti (Fieb.), Macropsis 91 multinotatus (Boh.), Adarrus 216 muscaeformis (Schrk.), Issus 69 musivus (Germ.), Myndus 27 Myndus 27 najas Nast, Macropsis 87 nanus (H.-S.), Hephathus 91 nardeti Rem., Psammotettix 211 nebulicola Em., Pinumius 221
Index of Auchenorrhyncha genera and species nebulosa (Ball), Endria 176 neglecta W.Wg., Alebra 111 Neoaliturus 165, 166 Neophilaenus 74-76 nervosa Vilb., Kelisia 34 nervosus (L.), Cixius 22 nigra (Goeze), Penthimia 103 nigriloba (Edw.), Edwardsiana 133 nigrita (Kbm.), Planaphrodes 105 nigritarsis Rem., Zygina 159 nigrolineata Scott, Eurybregma 41 nitidissimus (H.-S.), Populicerus 99 nitidulus (F.), Lamprotettix 181 nivea (M. & R.), Zygina 155 nodosus (Rib.), Psammotettix 215 notata (Proh.), Macropsis 86 notata Curt., Eupteryx 150 notatus (Mel.), Psammotettix 215 Nothodelphax 53 f. notula (Germ.), Megamelus 38 Notus 117 nubilus Oss., Macrosteles 170 obenbergeri (Dlab.), Verdanus 222 obliqua (W.Wg.), Stiromella 39 obliqua (Panz.), Tettigometra 68 obscurella (Boh.), Javesella 61 obsoletus (Germ.), Selenocephalus 229 obsoletus (Kbm.), Conosanus 201 obsoletus Sign., Hyalesthes 29 obtusiceps (Kbm.), Psammotettix 213 obtusifrons (Stål), Paramesus 206 obtusivalvis (Kbm.), Jassargus 218 ocellaris (Fall.), Errastunus 217 ocellaris (Leth.), Ophiola 198 ocellata Prov., Macropsis 84 ognevi (Zachv.), Ribautiana 139 ogumae (Mats.), Streptanus 205 ohausi W.Wg., Euscelis 203 okaensis Zachv., Streptanus 205 Ommatidiotus 69 Oncodelphax 57 Oncopsis 82-84 opacipennis (Leth.), Circulifer 166 Ophiola 198 f. Opsius 165
ordinaria (Rib.), Zygina 156 orichalceus Thoms., Jassus 198 origani Zachv., Eupteryx 144 ornata (Mel.), Cicadula 191 ornatus (Perr.), Phlepsius 184 orni L., Cicada 70 oshanini Razv., Macrosteles 171 ossiannilssoni Ldb., Macrosteles 172 ossiannilssoni Nuort., Empoasca 123 Ossiannilssonola 126 pallens (Stål), Ribautodelphax 228 pallidus (Hpt.), Neophilaenus lineatus 75 pallidinervis (Dhlb.), Psammotettix 212 pallidula (Boh.), Kelisia 33 Paluda 186 paludosa (Fl.), Paradelphacodes 57 pandellei (Leth.), Lindbergina 137 panzeri (Fl.), Cosmotettix 226 panzeri (P. Löw), Reptalus 28 paolii (Oss.), Chlorita 125 Paradelphacodes 57 Paradorydium 228 paradoxum (H.-S.), Paradorydium 228 Paraliburnia 47 f. Paralimnus 207 paraltaicus (Orosz), Kybos 120 Paramesus 206 Parapotes 206 parvicauda Rib., Mocydiopsis 193 parvula (Boh.), Arboridia 161 paryphasma (Fl.), Florodelphax 55 pascuellus (Fall.), Arthaldeus 223 pauxillus (Fieb.), Mendrausus 220 pectoralis Mats., Aphrophora 77 Pediopsis 84 pellax (Horv.), Laburrus 200 pellucida (F.), Javesella 60 Pentastiridius 27 f. Penthimia 103 penthopitta (Walk.), Diplocolenus 221 Perotettix 195 perplexus (Rib.), Kybos 120 persimilis (Edw.), Cicadula 190 perspicillata (Boh.), Anakelisia 36 petryi (Schum.), Cicadetta 71
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Philaenus 78 Phlepsius 184 phragmitis (Boh.), Paralimnus 207 pictilis (Stål), Aguriahana 151 picturatus (C. Shlb.), Stictocoris 197 pictus (Leth.), Perotettix 195 pilatoi D’Urso & Gugl., Cixidia 65 pilosus (Ol.), Tachycixius 26 Pinumius 221 Pithyotettix 195 Planaphrodes 105 planicollis (Thoms.), Macropsis 87 Platymetopius 179 f. plebeja (Edw.), Edwardsiana 133 plurispinosa (W.Wg.), Edwardsiana 131 poecilus (Fl.), Psammotettix 209 poecilus (H.-S.), Stenidiocerus 97 populi (Edw.), Kybos 118 populi (L.), Populicerus 100 Populicerus 99 f. potentillae (Mor.), Arboridia 161 praecox Hpt., Kelisia 33 Praganus 226 prasina (Boh.), Macropsis 85 preyssleri (H.-S.), Rhopalopyx 187 proceps (Kbm.), Rhytistylus 184 procerus (H.-S.), Handianus 229 propinqua (Fieb.), Toya 59 provincialis (Rib.), Hauptidia 153 pruinosa (Say), Metcalfa 235 pruni (Rib.), Balcanocerus 101 prunicola (Edw.), Edwardsiana 134 Psammotettix 209-215 pseudocellaris (Fl.), Jassargus 218 Pseudodelphacodes 59 pteridis (Dhlb.), Empoasca 123 pteridis (Spin.), Ditropis 39 pulchellus (Curt.), Delphax 43 pulchella (Fall.), Eurhadina 141 pulchra P. Löw, Zyginella 151 pulchripennis Asche, Litemixia 52 pulicaris (Fall.), Deltocephalus 175 pullula (Boh.), Oncodelphax 57 pullula (Boh.), Zyginidia 154 punctata (F.), Balclutha 167, 168
puncticollis (H.-S.), Grypotes 164 punctifrons (Fall.), Sagatus 175 punctulum (Kbm.), Kelisia 32 punctum (Fl.), Arocephalus 209 pungens (Rib.), Ribautodelphax 64 pusilla (Mats.), Chlorita 125 pusilla (Rib.), Arboridia 162 putoni (Then), Psammotettix 214 quadrimaculatus (Sign.), Megamelodes 48 quadrinotata (F.), Cicadula 191 quadripunctulatus (Kbm.), Macrosteles 172 quadrum Boh., Athysanus 197 quercus (F.), Typhlocyba 139 quinquecostatus (Duf.), Reptalus 28 quinquenotata (Boh.), Cicadula 190, 192 Recilia 176 remanei Nick., Macropsis 87 remanei Orosz, Psammotettix 215 repletus (Fieb.), Jassargus 219 Reptalus 28 reticulata (F.), Ulopa 80 reticulatus (H.-S.), Dryodurgades 94 reticulatus (Horv.), Parapotes 206 reyi (Fieb.), Calligypona 49 rhamnicola Horv., Zygina 157 rhenana W.Wg., Balclutha 167, 168 rhodophila (Cer.), Edwardsiana 129 rhombifer (Fieb.), Psammotettix 213 Rhopalopyx 187 f. Rhytidodus 94 Rhytistylus 184 ribauti (Oss.), Anaceratagallia 93 ribauti (Oss.), Arboridia 161 ribauti Nick. & Rem., Acericerus 100 ribauti W.Wg., Eurhadina 141 ribauti W.Wg., Kelisia 34 Ribautiana 138 f. Ribautodelphax 62-64, 228 rosae (L.), Edwardsiana 134 rosaesugans (Cer.), Edwardsiana 134 rosea (Fl.), Zygina 157 roseipennis (Toll.), Zygina 157 rosincola (Cer.), Zygina 158 rossica Zachv., Edwardsiana solearis 130 rostrata Rib., Eupteryx 234
Index of Auchenorrhyncha genera and species rotundiceps (Leth.), Paralimnus 207 rotundifrons Kbm., Idiocerus 100 rubroflava Lnv., Cicadula 189 rubrovittata (Leth.), Zygina 157 rufescens Mel., Kybos 118 russeola (Fall.), Ophiola 199 rutilans (Kbm.), Metidiocerus 96 saageri W.Wg., Eurhadina 141 sabulicola (Curt.), Psammotettix 212 sabulicola W.Wg., Kelisia 35 Sagatus 175 sagittarius Rib., Arocephalus 209 sahlbergi (Fl.), Macropsidius 91 salicicola (Edw.), Edwardsiana 134 salicina (Goeze), Aphrophora 78 salina (Hpt.), Javesella 62 saltuella (Kbm.), Balclutha 168 salviae Arz. & Vid., Eupteryx 148 sanguinolenta (Scop.), Cercopis 72 sanguinosa (R.), Fruticidia 163 Sardius 186 sardus Rib., Macrosteles 172 saturata (Edw.), Cicadula 190 scabripennis Edw., Megophthalmus 81 scalaris (Rib.), Ribautiana 138 scanicus (Fall.), Megophthalmus 81 schenckii (Kbm.), Euscelidius 201 schmidtgeni (W.Wg.), Recilia 176 schmidti (W.Wg.), Sorhoanus 224 schneideri (Günth.), Zygina 158 Scleroracus 198 scotti Edw., Macropsis 90 Scottianella 56 scutellaris (Fieb.), Iassus 102 scutellaris (H.-S.), Zyginidia 154 scutellata (Boh.), Macropsis 90 Selenocephalus 229 septemnotatus (Fall.), Macrosteles 173 septentrionalis W.Wg., Fieberiella 164 serratulae (F.), Anoscopus 108 sexmaculata (Hardy), Linnavuoriana 137 sexnotatus (Fall.), Macrosteles 173 sicula Mats., Chloriona 44 signatipennis (Boh.), Eupteryx 144 signifer (Then), Hardya 185
sima Rib., Kelisia 30 similis Kbm., Cixius 26 similis Kbm., Idiocerus 95 simillima (Lnv.), Javesella 60 simillima (W.Wg.), Arboridia 161 simplex (H.-S.), Cixius 23 sinuata (Then), Wagneriala 116 smaragdula (Fall.), Kybos 119 smaragdula (Stål), Chloriona 45 smreczynskii Dwor., Edwardsiana 135 sociabilis (Oss.), Edwardsiana 129 socialis (Fl.), Turrutus 218 Sonronius 174 f. sordidipennis (Stål), Macrosteles 173 sordidula (Stål), Megadelphax 46 sordidus (Zett.), Streptanus 205 Sorhoanus 224 soror (Lnv.), Edwardsiana 133 sororcula (Oss.), Edwardsiana 129 Sotanus 204 spathulata (Rib.), Arboridia 162 speciosa (Boh.), Euides 43 Speudotettix 194 spinigera (Edw.), Edwardsiana 135 spinosa (Fieb.), Acanthodelphax 53 spoliata (Horv.), Lindbergina 137 spumarius (L.), Philaenus 78 stachydearum (Hardy), Eupteryx 146 stactogalus Fieb., Opsius 165 stali (Metc.), Javesella 62 staminata (Rib.), Edwardsiana 127 stehliki Laut., Edwardsiana 135 steini (Fieb.), Metalimnus 208 stellulata (Burm.), Aguriahana 151 Stenidiocerus 97 Stenocranus 36 f. stenoptera (Fl.), Chloriona 45 sticticus R., Cixius 24 Stictocephala 79 Stictocoris 197 stigmaticalis Lew., Idiocerus 95 stigmaticus (Germ.), Cixius 25 stigmatipennis (M. & R.), Micantulina 115 Stiroma 41 Stiromella 39
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straminea (Stål), Xanthodelphax 57 Streptanus 204 f. striatella (Fall.), Laodelphax 47 striatula Fall., Cicada 198 strigilifer (Oss.), Kybos 120 striifrons (Kbm.), Arthaldeus 223 striola (Fall.), Limotettix 200 strobli (W.Wg.), Kybos 121 Stroggylocephalus 108 Struebingianella 56 stylata (Boh.), Doratura 177 suavis R., Zygina 157 subangulata (J. Shlb.), Oncopsis 84 subfusculus (Fall.), Speudotettix 194 substriatus (Then), Psammotettix 213 sudeticus (Kol.), Diplocolenus 221 sulphurella (Zett.), Elymana 188 sursumflexus (Then), Jassargus 220 Tachycixius 26 taeniatus (Horv.), Calamotettix 226 taimyrica Vilb., Hardya 185 tatraensis (Hell.), Errastunus ocellaris 217 tenella (Fall.), Eupteryx 149 tenerrima (H.-S.), Ribautiana 139 tenuis (Germ.), Hardya 185 tersa (Edw.), Edwardsiana 136 Tettigometra 65-68, 228 Thamnotettix 194 f. thenii (P. Löw), Sotanus 204 thoulessi Edw., Eupteryx 149 Tibicina 70 tiliae (Fall.), Zygina 158 tiliae (Germ.), Pediopsis 84 tithide Ferr., Zygina 155 torneellus (Zett.), Colladonus 180 Toya 59 transversa (Fall.), Ophiola 199 tredecimpunctatus (Ldb.), Metalimnus 208 tremulae (Estl.), Tremulicerus 97 Tremulicerus 97 f. tricolor (Gmel.), Balclutha 167 tricolorata Dlab., Calligypona 59 tridentata Edw., Anomia 128 trifasciata (Geoffr.), Planaphrodes 105 Trigonocranus 27
tristis (Zett.), Oncopsis 82 trivia (Germ.), Utecha 81 Turrutus 218 Typhlocyba 139 ulmi (Scott), Macropsis 89 ulmi (L.), Ribautiana 139 ulmiphagus Wils. & Clar., Edwardsiana 132 Ulopa 80 unciger Rib., Psammotettix 210 undatus (De G.), Platymetopius 179 unicolor Hpt., Aphrophora salicis 77 unicolor (H.-S.), Chloriona 44 Unkanodes 46 untica Dlab., Eurhadina 141 urticae (F.), Eupteryx 146 ustulatus (M. & R.), Viridicerus 98 Utecha 81 variata Hardy, Dikraneura 115 variatus (Fall.), Macrosteles 173 variegatus (Kbm.), Euscelidius 201 vasconica Rib., Chloriona 46 velata (Rib.), Arboridia 162 venosa (Geoffr.), Anaceratagallia 93 venosus (Germ.), Delphacodes 50 venosus (Kbm.), Euscelis 203 ventralis (Fall.), Graphocraerus 183 verbae Zachv., Kybos mucronatus 120 f. Verdanus 222 versuta (Mel.), Arboridia 162 viaduensis (W.Wg.), Zyginidia 154 vicina (Horv.), Macropsis 88 vicinus Mel., Idiocerus 95 vinealis Biem., Ribautodelphax 64 virescens (Panz.), Tettigometra 68 virgator (Rib.), Kybos 121 Viridicerus 98 viridigriseus (Edw.), Macrosteles 174 viridinervis W.Wg., Macropsis 85 viridis (L.), Cicadella 110 viridis R., Alebra 113 viridula (Fall.), Chlorita 228 vitis (Göthe), Empoasca 123 vitreus (F.), Tremulicerus 97 vitripennis (Fl.), Rhopalopyx 187, 188 vittata (L.), Eupteryx 149
Index of Auchenorrhyncha genera and species vittata (Leth.), Austroasca 124 vittifrons (Kbm.), Acericerus 100 vittipennis (J. Shlb.), Kelisia 32 volgensis Vilb., Kybos 121 vulnerata Rossi, Cercopis 72 wagneri China, Cixius 23 Wagneriala 116 Wagneripteryx 150 wahlbergi (Boh.), Alebra 112
williamsi China, Criomorphus 59 xantha Vilb., Xanthodelphax 57 Xanthodelphax 56 f. xanthoneurus (Fieb.), Sorhoanus 224 Xerochlorita Zachv. 124 Zonocyba 140 Zygina 155-160 Zyginella 151 Zyginidia 153 f.
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Index and vernacular names of plants Taxonomy and German vernacular names after Wisskirchen & Haeupler (1998), English vernacular names mainly after Clapham et al. (1987). Page numbers are given only for chapter 5. Bold numbers indicate tables. Abies alba Mill. – Silver Fir, Weiß-Tanne 237, 244, 245 Acer L. – Maple, Ahorn 275 f. Aceraceae 275 f., 332 ff., 342, 349 Acer campestre L. – Field Maple, Feld-Ahorn 275, 276, 323 Acer monspessulanum L. – Montpelier Maple, Französischer Ahorn 275, 276 Acer negundo L. – Ash-leaved Maple, Eschen-Ahorn 275 Acer opalus Mill. – Italian Maple, Frühlings-Ahorn 275 Acer palmatum Thunb. – Japanese Maple, Fächer-Ahorn 275, 276 Acer platanoides L. – Norway Maple, Spitz-Ahorn 275, 276 Acer pseudoplatanus L. – Sycamore, Berg-Ahorn 275, 276 Acer saccharum Marsh. – Sugar Maple, Zucker-Ahorn 275 Acer saccharinum L. – Silver Maple, Silber-Ahorn 275 Achillea millefolium L. s.l. – Yarrow (group), Artengruppe Wiesen-Schafgarbe 279 f., 282 f. Achnatherum P. Beauv. – Needlegrass, Rauhgras 321 Acinos alpinus (L.) Moench [= Calamintha alpina (L.) Lam.] – Alpine Calamint, Alpen-Steinquendel Acinos arvensis (Lam.) Dandy [= Calamintha acinos (L.) Clairv.] – Basil Thyme, Feld-Steinquendel Aegopodium podagraria L. – Goutweed, Giersch 277 Aesculus hippocastanum L. – Horse-chestnut, Rosskastanie 323 Agrostis L. – Bent-grass, Straußgras 309 f. Agrostis agrostiflora (Beck) Rauschert – Slender Bent-grass, Zartes Straußgras 310 Agrostis alpina Scop. – Alpine Bent-grass, Alpen-Straußgras 310 Agrostis canina L. – Velvet Bent-grass, Hunds-Straußgras 296 ff., 310 f. Agrostis capillaris L. – Common Bent-grass, Rotes Straußgras 296 ff., 310 f. Agrostis castellana Boiss. & Reut. – Castilian Bent-grass, Kastilisches Straußgras 310 Agrostis rupestris All. – Rock Bent-grass, Felsen-Straußgras 310 Agrostis scabra Willd. – Rough Bent-grass, Rauhes Straußgras 310 Agrostis stolonifera L. s.l. [incl. A. gigantea Roth] – Creeping Bent-grass (group), Artengruppe Weißes Straußgras 240, 296 ff., 310 Agrostis vinealis Schreb. [= A. coarctata Ehrh. ex Hoffm..] – Brown Bent-grass, Sand-Straußgras 296 ff., 310, 319 Aira L. – Hair-grass (partim), Haferschmiele 322, 342 Ajuga reptans L. – Bugle, Kriechender Günsel 280 f. Alnus Mill. – Alder, Erle 253 ff., 342 Alnus alnobetula (Ehrh.) K. Koch [= A. viridis (Chaix) DC.] – Green Alder, Grün-Erle 253 f., 254 f. Alnus glutinosa (L.) P. Gaertn. – Black Alder, Schwarz-Erle 253 ff., 254 f., 258, 351 Alnus incana (L.) Moench – Grey Alder, Grau-Erle 253 f., 254 f. Alopecurus L. – Foxtail, Fuchsschwanz 296 ff., 322 Alopecurus aequalis Sobol. – Orange Foxtail, Rotgelber Fuchsschwanz 322 Alopecurus geniculatus L. – Marsh Foxtail, Knick-Fuchsschwanz 322
Index and vernacular names of plants
443
Alopecurus myosuroides L. – Black Grass, Acker-Fuchsschwanz 322 Alopecurus pratensis L. – Meadow Foxtail, Wiesen-Fuchsschwanz 294, 321 Althaea officinalis L. – Marsh Mallow, Echter Eibisch 325 f., 327, 350 Althaea rosea (L.) Cav. [= Alcea rosea L.] – Hollyhock, Stockrose 325 f., 327 Amaranthaceae 333 Amelanchier lamarckii F.G. Schroed. – Juneberry, Kupfer-Felsenbirne 266 ff., 270 Amelanchier ovalis Medik. – European Serviceberry, Mitteleuropäische Felsenbirne 266 ff., 270 Amelanchier spicata (Lam.) K. Koch – Low Serviceberry, Besen-Felsenbirne 266 ff., 270 Ammophila arenaria (L.) Link – Marram Grass, Gewöhnlicher Strandhafer 296 ff., 319 Andromeda polifolia L. – Bog Rosemary, Rosmarinheide 263 Angelica L. – Angelica, Engelwurz 277 Anthoxanthum alpinum A. Löve & D. Löve – Alpine Vernal-grass, Alpen-Ruchgras 318 Anthoxanthum aristatum Boiss. – Annual Vernal-grass, Grannen-Ruchgras 318 Anthoxanthum odoratum L. – Sweet Vernal-grass, Gewöhnliches Ruchgras 296 ff., 318 Anthriscus Pers. – Chervil, Kerbel 277 Apera Adans. – Silky-bent, Windhalm 322 Apiaceae 276 f., 332 ff., 349 Araliaceae 336 Arctium L. – Burdock, Klette 282 f. Arctostaphylos Adans. – Bearberry, Bärentraube 263 Aristolochiaceae 329 Arrhenatherum elatius (L.) P. Beauv. ex J. Presl & C. Presl – False Oatgrass, Glatthafer 294, 296 ff., 315 f., 341 Artemisia L. – Wormwood, Beifuß 279 f. Artemisia abrotanum L. – Lad’s Love, Eberraute 239, 279, 282 f. Artemisia absinthium L. – Wormwood, Wermut 282 f. Artemisia campestris L. – Field Southernwood, Feld-Beifuß 279, 282 f. Artemisia maritima L. – Sea Wormwood, Strand-Beifuß 239, 279, 282 f. Artemisia pontica L. – Roman Wormwood, Pontischer Beifuß Artemisia vulgaris L. – Mugwort, Gewöhnlicher Beifuß 282 f. Arundo donax L. – Giant Reed, Pfahlrohr 239 Asparagales 322 Asphodelus L. – Asphodel, Affodil 347 Asplenium L. – Spleenwort, Streifenfarn 243 Asplenium scolopendrium L. [= Phyllitis scolopendrium (L.) Newman] – Hart’s-tongue Fern, Hirschzunge 243 Asteraceae 279 ff., 332 ff., 346, 348 f. Aster linosyris (L.) Bernh. – Goldilocks, Gold-Aster 279, 282 f. Athyrium filix-femina (L.) Roth – Lady-fern, Wald-Frauenfarn 243 Avena L. – Oat, Hafer 296 ff., 321, 351 Ballota nigra L. – Black Horehound, Schwarznessel 280 f. Berberis vulgaris L. – Barberry, Berberitze 324 Beta vulgaris L. – Beet, Runkelrübe 328 Betonica officinalis L. – Betony, Heil-Ziest 278, 280 f. Betula L. – Birch, Birke 253 ff., 342
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Betulaceae 253 ff., 331 ff., 342, 343, 349 Betula humilis Schrank – Shrub Birch, Strauch-Birke 253 Betula pendula Roth – Silver Birch, Hänge-Birke 253 f., 254 f. Betula pubescens Ehrh. – Downy Birch, Moor-Birke 237, 253 f., 254 f. Betula nana L. – Dwarf Birch, Zwerg-Birke 253 Blysmus Panz. ex Schult. – Flat Sedge, Quellbinse 287 Bolboschoenus maritimus (L.) Palla – Sea Club-rush, Gewöhnliche Strandsimse 287, 288 ff. Boraginaceae 322, 333 Botriochloa ischaemum (L.) Keng – Smooth Finger-grass, Bartgras Brachypodium P. Beauv. – False-brome, Zwenke 315 Brachypodium pinnatum (L.) P. Beauv. – Tor-grass, Fieder-Zwenke 239, 296 ff., 315 Brachypodium rupestris (Host) Roem. & Schult. – Rock False-brome, Felsen-Zwenke 315 Brachypodium sylvaticum (Huds.) P. Beauv. – Slender False-brome, Wald-Zwenke 315 Brassicaceae 332, 333 ff., 345 f. Briza media L. – Quaking Grass, Mittleres Zittergras 296 ff., 320 Bromus L. – Brome, Trespe 316, 342 Bromus erectus Huds. – Upright Brome, Aufrechte Trespe 296 ff., 316 f., 341 Bromus inermis Leyss. – Hungarian Brome, Wehrlose Trespe 296 ff., 316 f. Bromus ramosus Huds. – Hairy Brome, Wald-Trespe 316 f., 342 Bryophyta 329, 339 Buxus sempervirens L. – Box, Buchsbaum 324 Calamagrostis Adans. – Small-reed, Reitgras 308 Calamagrostis arundinacea (L.) Roth – Rough Small-reed, Wald-Reitgras 239, 296 ff., 308 Calamagrostis canescens (Weber) Roth – Purple Small-reed, Sumpf-Reitgras 296 ff., 308 Calamagrostis epigejos (L.) Roth – Wood Small-reed, Land-Reitgras 296 ff., 308, 319, 342 Calamagrostis pseudophragmites (Haller F.) Koel. – Bank Small-reed, Ufer-Reitgras 296 ff., 308 Calamagrostis stricta (Timm) Koeler – Bog Small-reed, Moor-Reitgras 308 Calamagrostis varia (Schrad.) Host – Mountain Small-reed, Buntes Reitgras 296 ff., 308 Calamagrostis villosa (Chaix ex Vill.) J.F. Gmel. – Shaggy Small-reed, Wolliges Reitgras 296 ff., 308 x Calammophila baltica (Flüggé ex Schrad.) Brand – Hybrid Marram, Baltischer Strandhafer 319 Calluna vulgaris (L.) Hull – Heather, Heidekraut 262 ff., 263 Caltha palustris L. – Marsh Marigold, Sumpfdotterblume 247 Calystegia sepium L. s.l. – Hedge Bindweed (group), Artengruppe Gewöhnliche Zaunwinde 328 Campanulaceae 322, 333 Cannabis sativa L. – Hemp, Hanf 248 Caprifoliaceae 337 Capsicum annuum L. – Sweet Pepper, Paprika 350 Caragana arborescens Lam. – Siberian Pea Shrub, Gewöhnlicher Erbsenstrauch Cardamine pratensis L. – Cuckooflower, Wiesen-Schaumkraut 328 Cardaria draba (L.) Desv. – Hoary Cress, Pfeilkresse 328 Carduus L. – Thistle, Distel (partim) 279, 282 f. Carex acuta L. [= Carex gracilis Curtis] – Slender Tufted Sedge, Schlank-Segge 288 ff., 292 f., 343 Carex acutiformis Ehrh. – Lesser Pond Sedge, Sumpf-Segge 288 ff., 292 f., 343 Carex alba Scop. – White-flowered Sedge, Weiße Segge 287, 288 ff., 293 Carex arenaria L. – Sand Sedge, Sand-Segge 287, 288 ff.
Index and vernacular names of plants
445
Carex brizoides L. – Quaking Sedge, Zittergras-Segge 287, 288 ff. Carex caryophyllea Latourr. [= Carex verna Chaix] – Spring Sedge, Frühlings-Segge 288 ff., 293 Carex distans L. – Distant Sedge, Entferntährige Segge 237, 288 ff., 292, 293 Carex disticha Huds. – Brown Sedge, Zweizeilige Segge 287, 288 ff. Carex elata All. – Tufted Sedge, Steife Segge 288 ff., 292 Carex elongata L. – Elongated Sedge, Walzen-Segge 288 ff., 293 Carex ferruginea Scop. – Rusty Sedge, Rost-Segge 288 ff., 293 Carex flacca Schreb. – Glaucous Sedge, Blaugrüne Segge 287, 288 ff., 293 Carex flava L. s.l. – Yellow Sedge (group), Artengruppe Gelb-Segge 287, 288 ff. Carex hirta L. – Hairy Sedge, Behaarte Segge 288 ff., 292 Carex humilis Leyss. – Dwarf Sedge, Erd-Segge 287, 288 ff. Carex limosa L. – Mud Sedge, Schlamm-Segge 237, 288 ff., 292 Carex montana L. – Soft-leaved Sedge, Berg-Segge 288 ff., 292 Carex muricata L. s.l. – Prickly Sedge (group), Artengruppe Sparrige Segge 288 ff., 293 Carex nigra (L.) Reichard – Common Sedge, Braune Segge 288 ff., 292, 343 Carex panicea L. – Carnation Sedge, Hirsen-Segge 288 ff., 292 Carex paniculata L. – Greater Tussock Sedge, Rispen-Segge 287, 288 ff., 293 Carex pilosa Scop. – Ciliated Sedge, Wimper-Segge 288 ff., 293 Carex pilulifera L. – Pill Sedge, Pillen-Segge 287, 288 ff., 293 Carex pseudocyperus L. – Cyperus Sedge, Scheinzypergras-Segge 288 ff., 293 Carex remota L. – Remote Sedge, Winkel-Segge 288 ff., 293 Carex riparia Curtis – Great Pond Sedge, Ufer-Segge 288 ff., 292 f. Carex rostrata Stokes – Beaked Sedge, Schnabel-Segge 286, 288 ff., 292 Carex sempervirens Vill. – Evergreen Sedge, Horst-Segge 288 ff., 293 Carex sylvatica Huds. – Wood Sedge, Wald-Segge 288 ff., 293 Carex vesicaria L. – Bladder Sedge, Blasen-Segge 288 ff., 293 Carex vulpina L. s.l. [incl. C. otrubae Podp.] – Fox Sedge (group), Artengruppe Fuchs-Segge 288 ff., 293 Carlina acaulis L. – Greater Carline Thistle, Silberdistel 282 f. Carlina vulgaris L. – Lesser Carline Thistle, Golddistel 282 f. Carpinus betulus L. – Hornbeam, Hainbuche 253 f., 254 f. Caryophyllaceae 327, 332 ff., 345 f., 349 Castanea sativa Mill. – Sweet Chestnut, Esskastanie 239, 250, 251 f. Catabrosa aquatica (L.) P. Beauv. – Whorl Grass, Quellgras 322 Catapodium rigidum (L.) C.E. Hubb. ex Dony – Fern Grass, Steifgras 322 Ceratophyllaceae 329 Chaerophyllum aromaticum L. – Carrot Chervil, Gewürz-Kälberkropf 277 Chaerophyllum aureum L. – Golden Chervil, Gold-Kälberkropf 277 Chaerophyllum hirsutum L. – Hairy Chervil, Rauhhaariger Kälberkropf 277 Chamaecyparis lawsoniana (A. Murr.) Parl. – Lawson’s False Cypress, Lawson-Scheinzypresse 240, 244, 245, 349 Chamaespartium sagittale (L.) Gibbs [= Genista sagittalis L.; = Genistella sagittalis (L.) Gams] – Winged Broom, Gewöhnlicher Flügelginster 273, 274 Chenopodiaceae 322, 333, 337 Chrysanthemum vulgare L. – Tansy, Rainfarn 282 f.
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Cirsium acaule Scop. – Dwarf Thistle, Stengellose Kratzdistel 279, 282 f. Cirsium arvense (L.) Scop. – Creeping Thistle, Acker-Kratzdistel 282 f. Cirsium oleraceum (L.) Scop. – Cabbage Thistle, Kohldistel 282 f. Cistaceae 327 Cistus L. – Cistus, Zistrose Cladium mariscus (L.) Pohl – Great Fen Sedge, Schneidried 287 Clematis vitalba L. – Traveller’s Joy, Gewöhnliche Waldrebe 247 Clinopodium vulgare L. – Wild Basil, Wirbeldost 280 f. Clusiaceae 327, 349 Coleanthus subtilis (Tratt.) Seidl – Sheath-grass, Scheidenblütgras 322 Colutea arborescens L. – Bladder Senna, Gewöhnlicher Blasenstrauch 273, 274 Convolvulaceae 327 Convolvulus arvensis L. – Field Bindweed, Acker-Winde 326, 327 Corispermum leptopterum (Aschers.) Iljin – Narrow-winged Bugseed, Schmalflügeliger Wanzensame Cornaceae 336 f. Cornus mas L. – Cornelian Cherry, Kornelkirsche 322, 323 Cornus sanguinea L. – Dogwood, Blutroter Hartriegel 322, 323 Coronilla L. – Crownvetch, Kronwicke 273, 274 Corylus avellana L. – Hazel, Gewöhnliche Hasel 253 f., 254 f. Corylus colurna L. – Tree Hazel, Baum-Hasel 253, 254 f. Corylus maxima Mill. – Filbert, Lambertsnuss 253, 254 f. Corynephorus canescens (L.) P. Beauv. [= Weingaertneria canescens (L.) Bernh.] – Grey Hair-grass, Silbergras 296 ff., 319 Cotoneaster Medik. – Cotoneaster, Zwergmispel 264 f., 266 ff. Crataegus L. – Hawthorn, Weißdorn 266 ff., 269 f., 272 Crepis L. – Hawk’s-beard, Pippau 279, 282 f. Crepis paludosa (L.) Moench – Marsh Hawk’s-beard, Sumpf-Pippau 284 Cucumis sativus L. – Cucumber, Gurke 350 Cucurbitaceae 349 Cupressaceae 244 ff., 336 f., 349 Cupressus sempervirens L. – Italian Cypress, Italienische Zypresse Cydonia oblonga Mill. – Quince, Quitte 270 Cynodon dactylon (L.) Pers. – Bermuda-grass, Hundszahngras 294, 296 ff., 320 f. Cynosurus cristatus L. – Crested Dogstail, Wiesen-Kammgras 294, 322 Cyperaceae 284 ff., 329, 331 ff., 342, 345 f. Cyperus L. – Galingale, Zypergras 287 Cyperus longus L. – Tall Galingale, Hohes Zypergras Cytisus nigricans L. [= Lembotropis nigricans (L.) Griseb.] – Black Broom, Schwarzwerdender Ginster 273, 274 Cytisus scoparius (L.) Link [= Sarothamnus scoparius (L.) W.D. J. Koch] – Broom, Besenginster 273, 274 Dactylis glomerata L. – Cock’s-foot, Wiesen-Knäuelgras 296 ff., 313 Dactylis polygama Horv. – Wood Cock’s-foot, Wald-Knäuelgras 313 Dahlia pinnata Cav. – Dahlia, Dahlie 282 f. Danthonia decumbens (L.) DC. [= Sieglingia decumbens (L.) Bernh.] – Heath Grass, Dreizahn 321
Index and vernacular names of plants
447
Daucus carota L. – Wild Carrot, Wilde Möhre 277 Dennstaedtiaceae 336 f. Deschampsia P. Beauv. – Hair-grass (partim), Schmiele 308 f. Deschampsia cespitosa (L.) P. Beauv. – Tufted Hair-grass, Rasen-Schmiele 266 ff., 309, 342, 351 Deschampsia flexuosa (L.) Trin. [= Avenella flexuosa (L.) Drejer] – Wavy Hair-grass, SchlängelSchmiele 239, 240, 266 ff., 309 Deschampsia littoralis (Gaudin) Reut. – Lake Constance Hair-grass, Bodensee-Schmiele 309 Deschampsia media (Gouan) Roem. & Schult. – Intermediate Hair-grass, Binsen-Schmiele 309 Deschampsia setacea (Huds.) Hack. – Bog hair-grass, Borst-Schmiele 309 Deschampsia wibeliana (Sond.) Parl. – Mud Hair-grass, Elbe-Schmiele 309 Digitalis L. – Foxglove, Fingerhut 349 Digitaria sanguinalis (L.) Scop. – Hairy Finger-grass, Blutrote Fingerhirse 294, 320, 321 Dipsacaceae 327 Dryopteris Adans. – Male-fern, Wurmfarn 243 Ecballium (L.) A. Rich. – Squirting Cucumber, Spritzgurke 349 Echinochloa crus-galli (L.) P. Beauv. – Cockspur, Gewöhnliche Hühnerhirse 321 Echinops L. – Globe Thistle, Kugeldistel Echium vulgare L. – Viper’s Bugloss, Gewöhnlicher Natternkopf 328 Elaeagnaceae 336 f. Elaeagnus angustifolia L. – Russian-olive, Schmalblättrige Ölweide 322, 323 Eleocharis palustris (L.) Roem. & Schult. – Common Spike-rush, Gewöhnliche Sumpfbinse 287, 288 ff. Eleocharis uniglumis (Link) Schult. – Slender Spike-rush, Einspelzige Sumpfbinse 287, 288 ff. Elymus L. – Couch-grass, Quecke 312 f., 319 Elymus arenosus (Spenn.) Conert – Mainz Couch, Sand-Quecke 313 Elymus athericus (Link) Kerguélen [= Agropyron pungens auct. non (Pers.) Roem. & Schult.] – Sea Couch, Dünen-Quecke 313 Elymus caninus (L.) L. – Bearded Couch, Hunds-Quecke 313 Elymus farctus (Viv.) Runemark ex Melderis – Sand Couch, Binsen-Quecke 313 Elymus hispidus (Opiz) Melderis – Hairy Couch, Graugrüne Quecke 313 Elymus obtusiflorus (DC.) Conert – Blunt-flowered Couch, Stumpfblütige Quecke 313 Elymus repens (L.) Gould [= Agropyron repens (L.) P. Beauv.] – Couch-grass, Kriech-Quecke 296 ff., 312 f., 342, 351 Epilobium angustifolium L. [= Chamaenerion angustifolium (L.) Scop.] – Fireweed, Schmalblättriges Weidenröschen 325 f., 327 Equisetaceae 336 Equisetum arvense L. – Field Horsetail, Acker-Schachtelhalm 243 Equisetum palustre L. – Marsh Horsetail, Sumpf-Schachtelhalm 243 Equisetum sylvaticum L. – Wood Horsetail, Wald-Schachtelhalm 243 Eragrostis Wolf – Love-grass, Liebesgras 322, 342 Ericaceae 262 ff., 332 ff., 345 f. Erica carnea L. – Spring Heath, Schnee-Heide 263, 264 Erica tetralix L. – Cross-leaved Heath, Glocken-Heide 263, 264 Eriophorum angustifolium Honck. – Common Cottongrass, Schmalblättriges Wollgras 286, 288 ff. Eriophorum gracile W.D.J. Koch ex Roth – Slender Cottongrass, Schlankes Wollgras 286
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Eriophorum latifolium Hoppe – Broad-leaved Cottongrass, Breitblättriges Wollgras 286, 288 ff. Eriophorum scheuchzeri Hoppe – White Cottongrass, Scheuchzers Wollgras 286 Eriophorum vaginatum L. – Hare’s-tail Cottongrass, Scheiden-Wollgras 286, 288 ff. Eryngium L. – Eryngo, Mannstreu 347 Euonymus europaeus L. – Spindle Tree, Europäisches Pfaffenhütchen 324 Eupatorium cannabinum L. – Hemp Agrimony, Gewöhnlicher Wasserdost 282 f. Euphorbiaceae 322, 333 Fabaceae 273 f., 332 ff., 345 f. Fagaceae 250 ff., 331 ff., 342, 343, 349 Fagus sylvatica L. – Beech, Rotbuche 250, 251 f. Falcaria vulgaris Bernh. – Longleaf, Sichelmöhre 277 Festuca L. – Fescue, Schwingel 295, 308 Festuca altissima All. – Wood Fescue, Wald-Schwingel 296 ff., 308, 319 Festuca arundinacea Schreb. – Tall Fescue, Rohr-Schwingel 295, 296 ff. Festuca gigantea (L.) Vill. – Giant Fescue, Riesen-Schwingel 308, 342 Festuca heterophylla Lam. – Various-leaved Fescue, Verschiedenblättriger Schwingel 296 ff., 308 Festuca ovina L. s.l. [incl. F. pallens Host, F. rupicola Heuff., F. valesiaca Schleich. ex Gaudin, etc.] – Sheep’s Fescue (group), Artengruppe Schaf-Schwingel 295, 296 ff., 308, 319, 320, 342 Festuca pratensis Huds. – Meadow Fescue, Wiesen-Schwingel 296 ff., 308 Festuca puccinellii Parl. – Dark-violet Fescue, Dunkelvioletter Schwingel 295 Festuca pulchella Schrad. – Beautiful Fescue, Schöner Schwingel 295 Festuca quadriflora Honck. – Four-flowered Fescue, Niedriger Schwingel 295 Festuca rubra L. s.l. – Red Fescue (group), Artengruppe Rot-Schwingel 295, 296 ff., 308, 319, 320 Festuca rupicaprina (Hack.) A. Kern. – Chamois Fescue, Gemsen-Schwingel 295 Festuca vivipara (L.) Sm. – Viviparous Fescue, Knollen-Schwingel Ficus carica L. – Common Fig, Feigenbaum 324 Filipendula ulmaria (L.) Maxim. – Meadowsweet, Echtes Mädesüß 239, 266 ff., 270 f., 349 Forsythia Vahl – Forsythia, Goldflieder 324 Fragaria L. – Strawberry, Erdbeere 266 ff., 270 Frangula alnus Mill. [= Rhamnus frangula L.] – Alder Buckthorn, Faulbaum 241, 322, 323 Fraxinus excelsior L. – Ash, Gewöhnliche Esche 323, 342 Fungi 329 ff. Galeopsis angustifolia Hoffm. – Red Hemp-nettle, Schmalblättriger Hohlzahn 278, 280 f. Galium L. – Bedstraw, Labkraut 328, 349 Galium saxatile L. [= Galium harcynicum Weigel] – Heath Bedstraw, Harzer Labkraut Galium verum L. s.l. – Lady’s Bedstraw (group), Artengruppe Echtes Labkraut Genista anglica L. – Petty Whin, Englischer Ginster 273 Genista pilosa L. – Hairy Whin, Behaarter Ginster 273, 274 Genista tinctoria L. – Dyer’s Greenweed, Färber-Ginster 273, 274 Gentianaceae 333 Geraniaceae 322, 327, 349 Geranium pusillum Burm. F. – Small-flowered Crane’s-bill, Kleiner Storchschnabel 328 Geranium robertianum L. – Herb-Robert, Stink-Storchschnabel 328 Geranium sanguineum L. – Bloody Crane’s-bill, Blutroter Storchschnabel 325, 327 Glechoma hederacea L. – Ground-ivy, Gundermann 278, 280 f.
Index and vernacular names of plants
449
Globularia cordifolia L. – Heart-leaf Globularia, Herzblättrige Kugelblume Glyceria R. Br. – Sweet-grass, Schwaden 320 Glyceria declinata Bréb. – Small Sweet-grass, Blaugrüner Schwaden 320 Glyceria fluitans L. s.l. – Floating Sweet-grass (group), Artengruppe Flutender Schwaden 296 ff., 320 Glyceria maxima (Hartm.) Holmb. [= Poa aquatica L.; = Glyceria spectabilis Mert. & W.D.J. Koch] – Reed Sweet-grass, Wasser-Schwaden 296 ff., 320 Glyceria nemoralis (Uechtr.) Uechtr. & Körn. – Wood Sweet-grass, Hain-Schwaden 320 Glyceria notata Chevall. – Plicate Sweet-grass, Falt-Schwaden 320 Glyceria striata (Lam.) Hitchc. – Striated Sweet-grass, Streifen-Schwaden 320 Glycyrrhiza glabra L. – Licorice-root, Süßholz 274 Graminoid monocots 329 f. Gymnospermae 244 ff., 329 ff., 349 Hedera helix L. – Ivy, Efeu 246, 324, 340 Helianthemum alpestre (Jacq.) DC – Mountain Rock-rose, Alpen-Sonnenröschen Helianthemum nummularium (L.) Mill. – Common Rock-rose, Gewöhnliches Sonnenröschen 325 f., 327 Helianthus annuus L. – Common Sunflower, Sonnenblume 282 f. Helichrysum arenarium (L.) Moench – Strawflower, Sand-Strohblume 282 f. Helictotrichon s.l. Besser – Oat-grass (partim), Wiesen-/Flaumhafer 318 Helictotrichon parlatorei (Woods) Pilg. – Alpine Oat-grass, Parlatores Wiesenhafer 318 Helictotrichon pratense (L.) Besser [= Avena pratensis L.; = Avenochloa pratensis (L.) Holub; = Avenula pratensis (L.) Dum.] – Meadow Oat-grass, Gewöhnlicher Wiesenhafer 296 ff., 318 Helictotrichon pubescens (Huds.) Pilg. – Hairy Oat-grass, Flaumhafer 296 ff., 318, 342 Helictotrichon versicolor (Vill.) Pilg. – Variegated Oat-grass, Bunter Wiesenhafer 318 Heracleum mantegazzianum Sommier & Levier – Giant Hogweed, Riesen-Bärenklau 277 Heracleum sphondylium L. – Cow Parsnip, Wiesen-Bärenklau 277 Hieracium L. – Hawkweed, Habichtskraut 279, 282 f. Hieracium pilosella L. – Mouse-ear Hawkweed, Mausohr-Habichtskraut 282 f., 284 Hierochloe R. Br. – Holy grass, Mariengras 322 Hippocastanaceae 336 f. Hippocrepis comosa L. – Horseshoe Vetch, Gewöhnlicher Hufeisenklee 274 Hippophae rhamnoides L. – Sea-buckthorn, Sanddorn 322, 323 Holcus L. – Velvetgrass, Honiggras 284, 311 Holcus lanatus L. – Yorkshire Fog, Wolliges Honiggras 296 ff., 311 Holcus mollis L. – Creeping Soft-grass, Weiches Honiggras 239, 296 ff., 311 Hordelymus europaeus (L.) Jessen ex Harz – Wood Barley, Waldgerste 322, 342 Hordeum L. – Barley, Gerste 296 ff., 321, 342, 350 Horminum L. – Drachenmaul, Dragon’s-mouth 278 Humulus lupulus L. – Hop, Gewöhnlicher Hopfen 328 Hypericum L. – St John’s-wort, Joahnniskraut 349 Hypericum maculatum Crantz s.l. – Imperforate St John’s-wort (group), Artengruppe Geflecktes Johanniskraut 325 f., 327 Hypericum perforatum L. – Perforate St John’s-wort, Tüpfel-Johanniskraut 325 f., 327 Hypochaeris L. – Cat’s Ear, Ferkelkraut 282 f. Isolepis R. Br. – Club-rush (partim), Moorbinse 287
450
Appendix
Juglans regia L. – Walnut, Echte Walnuss 324 Juncaceae 284 f., 329, 332 ff., 345 Juncus articulatus L. – Jointed Rush, Glieder-Binse 284, 285 Juncus bufonius L. – Toad Rush, Kröten-Binse 285 Juncus conglomeratus L. – Compact Rush, Knäuel-Binse 285 Juncus effusus L. – Soft Rush, Flatter-Binse 284, 285 Juncus filiformis L. – Thread Rush, Faden-Binse Juncus gerardii Loisel. – Saltmarsh Rush, Salz-Binse 285 Juncus inflexus L. – Hard Rush, Blaugrüne Binse 284, 285 Juncus subnodulosus Schrank – Blunt-flowered Rush, Stumpfblütige Binse 285 Juniperus L. – Juniper, Wacholder 239 Juniperus communis L. – Common Juniper, Gewöhnlicher Wacholder 244, 245 Juniperus sabina L. – Savin Juniper, Sadebaum 244 Kalmia L. – Laurel, Lorbeerrose 263 Knautia dipsacifolia Kreutzer – Teasel-leaved Knautia, Wald-Witwenblume 325, 327 Kobresia myosuroides (Vill.) Fiori – Bellard’s Kobresia, Europäisches Nacktried 287 Koeleria Pers. – Hair-grass (partim), Schillergras 318 Koeleria arenaria (Dumort.) Conert – Dune Hair-grass, Sand-Schillergras 318 Koeleria glauca (Spreng.) DC. – Glaucous Hair-grass, Blaugrünes Schillergras 296 ff., 318 f. Koeleria macrantha (Ledeb.) Schult. – Crested Hair-grass, Zierliches Schillergras 318 Koeleria pyramidata (Lam.) P. Beauv. – Pyramidal Hair-grass, Pyramiden-Schillergras 318 Koeleria vallesiana (Honck.) Gaudin – Rock Hair-grass, Walliser Schillergras 318 Lagenaria siceraria (Mol.) Standl. – Calabash, Kalebasse 328 Lamiaceae 277 ff., 332 ff., 345 f., 348 f. Lamium album L. – White Dead-nettle, Weiße Taubnessel 280 f. Lamium galeobdolon L. s.l. – Yellow Archangel (group), Artengruppe Goldnessel 280 f. Lamium maculatum L. – Spotted Dead-nettle, Gefleckte Taubnessel 278, 280 f. Lapsana communis L. – Nipplewort, Rainkohl 282 f. Lathyrus L. – Sweet-pea, Platterbse 274 Larix decidua Mill. – European Larch, Europäische Lärche 244, 245 Ledum palustre L. – Labrador Tea, Sumpf-Porst 263 Leersia oryzoides (L.) Sw. – Cut Grass, Reisquecke 324 Leontodon hispidus L. – Rough Hawkbit, Rauher Löwenzahn 279, 282 f. Leonurus cardiaca L. – Motherwort, Herzgespann 280 f. Leymus arenarius (L.) Hochst. [= Elymus arenarius L.] – Lyme-grass, Strandroggen 296 ff., 320 Ligustrum vulgare L. – Common Privet, Liguster 246, 323 Liliaceae 333, 337 Liliales 322 Loiseleuria procumbens Desv. – Alpine Azalea, Alpenazalee 263 Lolium perenne L. – Rye-grass, Ausdauerndes Weidelgras 296 ff., 321 Lonicera L. – Honeysuckle, Heckenkirsche 324 Lonicera nigra L. – Black Honeysuckle, Schwarze Heckenkirsche Lonicera xylosteum L. – Fly Honeysuckle, Rote Heckenkirsche Lotus corniculatus L. s.l. – Common Birdsfoot-trefoil (group), Artengruppe Gewöhnlicher Hornklee 273, 274
Index and vernacular names of plants
451
Lotus pedunculatus Cav. [= Lotus uliginosus Schkuhr] – Greater Birdsfoot-trefoil, Sumpf-Hornklee Lupinus polyphyllus Lindl. – Common Lupin, Vielblättrige Lupine 274 Luzula DC. – Woodrush, Hainsimse 284, 285 Luzula luzuloides (Dam.) Dandy & Wilm. [= Luzula albida (Hoffm.) DC.] – White Woodrush, Weißliche Hainsimse Luzula sylvestris (Huds.) Gaudin s.l. – Great Woodrush (group), Artengruppe Wald-Hainsimse Lycopus europaeus L. – Gipsywort, Ufer-Wolfstrapp 278, 280 f. Lysimachia vulgaris L. – Yellow Loosestrife, Gewöhnlicher Gilbweiderich 328 magnoliid complex 329 Malus Mill. – Apple, Apfel 265, 266 ff. Malvaceae 327, 337 Marrubium vulgare L. – White Horehound, Gewöhnlicher Andorn 280 f. Marsileaceae 324 Marsilea quadrifolia L. – Water-clover, Vierblättriger Kleefarn 324 Medicago lupulina L. – Black Medick, Hopfenklee 273, 274 Medicago sativa L. s.l. – Alfalfa (group), Artengruppe Saat-Luzerne 273, 274 Melica L. – Melick, Perlgras 321 Melica ciliata L. – Hairy Melick, Wimper-Perlgras 321 Melica nutans L. – Mountain Melick, Nickendes Perlgras 321 Melica picta K. Koch – Variegated Melick, Buntes Perlgras 321 Melica transsilvanica Schur – Transsylvanian Melick, Siebenbürger Perlgras 321 Melica uniflora Retz. – Wood Melick, Einblütiges Perlgras 296 ff., 321 Melilotus Mill. – Melilot, Steinklee Melissa officinalis L. – Balm, Zitronen-Melisse 278, 280 f., 350 Melittis melissophyllum L. – Bastard Balm, Immenblatt 278 Mentha aquatica L. – Water Mint, Wasser-Minze 278, 280 f. Mentha longifolia (L.) Huds. – Horse Mint, Ross-Minze 278, 280 f. Mentha x piperita L. – Peppermint, Pfeffer-Minze 280 f. Mespilus L. – Medlar, Mispel 270 Mibora minima (L.) Desv. – Early Sand-grass, Zwerggras 322 Micropyrum tenellum (L.) Link – Gravel Fescue, Kies-Dünnschwingel 322 Milium effusum L. – Wood Millet, Flattergras 322, 342 Molinia caerulea (L.) Moench s.l. – Purple Moor-grass (group), Artengruppe Pfeifengras 239, 240, 296 ff., 316 Myricaceae 336 Myricaria germanica (L.) Desv. – German False Tamarisk, Deutsche Tamariske 239, 322, 323 Myrica gale L. – Bog Myrtle, Gagelstrauch 324 Nardus stricta L. – Mat-grass, Borstgras 296 ff., 320 Nasturtium W.T. Aiton – Water-cress, Brunnenkresse Nepeta cataria L. – Cat-mint, Gewöhnliche Katzenminze 280 f. non-monocot paleoherbs 329 Nicotiana tabacum L. – Tobacco, Tabak 351 Nothofagus Blume – Southern Beech, Südbuche Nuphar lutea (L.) Sibth. & Sm. – Yellow Water-lily, Gelbe Teichrose 324, 325 Nymphaea alba L. – White Water-lily, Weiße Seerose 324, 325
452
Appendix
Nymphaeaceae 324, 329 Ocimum basilicum L. – Basil, Basilikum 278, 280 f., 351 Oleaceae 323, 336 f. Onagraceae 322, 327, 337 Onobrychis Mill. – Sainfoin, Esparsette 273, 274 Ononis spinosa L. – Spiny Restharrow, Dornige Hauhechel 273, 274 Onopordum acanthium L. – Scotch Thistle, Eselsdistel 282 f. Orchidaceae 322, 333, 337 Origanum vulgare L. – Marjoram, Gewöhnlicher Dost 278, 280 f. Orobanche L. – Broomrape, Sommerwurz 346 Paeonia L. – Peony, Pfingstrose Panicum L. – Millet (partim), Rispenhirse 322 Papaveraceae 333 Parapholis strigosa (Dumort.) C.E. Hubb. – Hard Grass, Gekrümmter Dünnschwanz 322 Parietaria judaica L. – Common Pellitory-of-the-Wall, Mauer-Glaskraut 248 f., 328 Parietaria officinalis L. – Upright Pellitory-of-the-Wall, Aufrechtes Glaskraut 248 Pastinaca sativa L. – Wild Parsnip, Pastinak 277 Petasites Mill. – Butterbur, Pestwurz 279, 282 f. Phalaris arundinacea L. [= Typhoides arundinacea (L.) Moench; = Baldingera arundinacea (L.) Dumort.] – Reed-grass, Rohr-Glanzgras 296 ff., 317 Phleum L. – Cat’s-tail, Lieschgras 294, 296 ff., 321, 351 Phleum pratense L. – Timothy, Wiesen-Lieschgras 321 Phleum phleoides (L.) H. Karst. – Purple-stem Cat’s-tail, Steppen-Lieschgras 322 Phleum alpinum L. – Alpine Cat’s-tail, Alpen-Lieschgras 322 Phleum hirsutum Honck. – Rough Cat’s-tail, Matten-Lieschgras 322 Phlomis L. – Jerusalem Sage, Brandkraut 278 Phragmites australis (Cav.) Trin. ex Steud. [= Phragmites communis Trin.] – Common Reed, Gewöhnliches Schilf 239, 260, 296 ff., 311 f., 333, 342 Picea abies (L.) H. Karst. – Norway Spruce, Gewöhnliche Fichte 244, 245, 246, 259 Picea omorika (Pancic) Purkyne – Serbian Spruce, Serbische Fichte Picris hieracioides L. – Hawkweed Oxtongue, Gewöhnliches Bitterkraut 282 f. Pinaceae 332 ff., 342 Pinus L. – Pine, Kiefer 244, 330, 343, 349, 351 Pinus cembra L. – Stone Pine, Zirbel-Kiefer 244, 245 Pinus mugo Turra – Mountain Pine, Latschen-Kiefer 244, 245 Pinus nigra J.F. Arnold – Black Pine, Schwarz-Kiefer 239, 244, 245 Pinus sylvestris L. – Scots Pine, Wald-Kiefer 239, 244, 245, 246 Plantaginaceae 327 Plantago lanceolata L. – Ribwort Plantain, Spitz-Wegerich 325 f., 327 Plantago major L. – Common Plantain, Breit-Wegerich 325 f., 327 Plantago media L. – Hoary Plantain, Mittlerer Wegerich 325 f., 327 Platanaceae 336 Platanus L. – Plane, Platane 323 Poa L. – Meadow-grass, Rispengras 313 f., 351 Poa alpina L. – Alpine Meadow-grass, Alpen-Rispengras 314
Index and vernacular names of plants
453
Poa annua L. – Annual Meadow-grass, Einjähriges Rispengras 296 ff., 314 Poa badensis Haenke ex Willd. – Baden Meadow-grass, Badener Rispengras 314 Poaceae 294 ff., 329, 331 ff., 342, 343, 345 f. Poa chaixii Vill. – Broad-leaved Meadow-grass, Wald-Rispengras 314 Poa compressa L. – Flattened Meadow-grass, Platthalm-Rispengras 314 Poa hybrida Gaudin – Hybrid Meadow-grass, Großes Rispengras 314 Poa nemoralis L. – Wood Meadow-grass, Hain-Rispengras 296 ff., 314 Poa palustris L. – Swamp Meadow-grass, Sumpf-Rispengras 315 Poa pratensis L. s.l. [incl. P. angustifolia L.] – Smooth Meadow-grass (group), Artengruppe WiesenRispengras 296 ff., 314 Poa supina Schrad. – Prostrate Meadow-grass, Läger-Rispengras 314 Poa trivialis L. – Rough Meadow-grass, Gewöhnliches Rispengras 296 ff., 314 Polygonaceae 327, 337, 346 Polygonum aviculare L. s.l. – Knotgrass (group), Artengruppe Gewöhnlicher Vogelknöterich 325, 327 Polypodium interjectum Shivas – Western Polypody, Gesägter Tüpfelfarn 243, 349 Polypodium vulgare L. – Common Polypody, Gewöhnlicher Tüpfelfarn 243, 349 Polystichum Roth. – Shield-fern, Schildfarn 243 Populus L. – Poplar, Pappel 258 ff. Populus alba L. – White Poplar, Silber-Pappel 258, 260, 261 f. Populus nigra L. [incl. Populus nigra “italica” (Duroi) Moench] – Black Poplar [Lombard Poplar], Schwarz-Pappel [Pyramiden-Pappel] 258 ff, 261 f.. Populus tremula L. – Aspen, Zitter-Pappel 258, 261 f., 340 Potamogetonaceae 324 Potamogeton natans L. – Broad-leaved Pondweed, Schwimmendes Laichkraut 324, 325 Potentilla L. – Cinquefoil, Fingerkraut 349 Potentilla fruticosa L. – Shrubby Cinquefoil, Strauch-Fingerkraut 266 ff., 270 Potentilla incana P. Gaertn., B. Mey. & Scherb. [= Potentilla arenaria P. Gaertn., B. Mey. & Scherb.] – Sand Cinquefoil, Sand-Fingerkraut 266 ff., 271 Potentilla tabernaemontani Asch. [= Potentilla neumanniana Rchb.] – Spring Cinquefoil, Gewöhnliches Frühlings-Fingerkraut 266 ff., 271 Primulaceae 322, 333, 349 Prunella vulgaris L. – Selfheal, Kleine Braunelle 280 f. Prunus L. – Cherry, Plum, Kirsche, Pflaume 271 ff. Prunus avium L. [= Cerasus avium Moench] – Wild Cherry (incl. Sweet Cherry), Vogel-Kirsche (incl. Süß-Kirsche) 266 ff., 271 f. Prunus cerasus L. s.l. – Sour Cherry (group), Artengruppe Sauerkirsche 266 ff., 271 Prunus domestica L. – Plum, Zwetschge, Pflaume 266 ff., 271 Prunus dulcis (Mill.) D.A. Webb – Almond, Mandel 271 Prunus fruticosa Pall. – Dwarf Cherry, Zwerg-Kirsche 271 Prunus laurocerasus L. – Laurel Cherry, Lorbeer-Kirsche 271 Prunus mahaleb L. – Mahaleb Cherry, Felsen-Kirsche 271 Prunus padus L. – Bird Cherry, Trauben-Kirsche 266 ff., 271 ff. Prunus persica (L.) Batsch – Peach, Pfirsich 271 Prunus serotina Ehrh. – Black Cherry, Späte Traubenkirsche 271 Prunus spinosa L. – Blackthorn, Schlehe 266 ff., 271 ff.
454
Appendix
Pseudotsuga menziesii (Mirb.) Franco – Douglas Fir, Douglasie 244, 245 Pteridium aquilinum (L.) Kuhn – Bracken, Gewöhnlicher Adlerfarn 243 Pteridophyta 242 f., 324, 329 ff. Puccinellia Parl. – Saltmarsh-grass, Salzschwaden 317 Puccinellia capillaris (Lilj.) Jansen – Northern Saltmarsh-grass, Haar-Salzschwaden 317 Puccinellia distans (Jacq.) Parl. – Reflexed Saltmarsh-grass, Gewöhnlicher Salzschwaden 296 ff., 317 Puccinellia limosa (Schur) E. Holmb. – Mud Saltmarsh-grass, Sumpf-Salzschwaden 317 Puccinellia maritima (Huds.) Parl. – Common Saltmarsh-grass, Andel 296 ff., 317 Pyracantha coccinea Roem. – Fire Thorn, Feuerdorn 266 ff., 270 Pyrus communis L. s.l. - Pear (group), Artengruppe Birnbaum 265, 266 ff., 351 Quercus L. – Oak, Eiche 250 ff., 342 Quercus cerris L. – Turkey Oak, Zerr-Eiche 250 Quercus petraea Liebl. – Durmast Oak, Trauben-Eiche 239, 250 ff., 251 f. Quercus pubescens Willd. – Downy Oak, Flaum-Eiche 250, 251 f. Quercus robur L. – Common Oak, Stiel-Eiche 250 ff., 251 f. Quercus rubra L. – Northern Red Oak, Rot-Eiche 250 Ranunculaceae 246, 332 ff., 345 f., 349 Ranunculus bulbosus L. – Bulbous Buttercup, Knolliger Hahnenfuß 247 Ranunculus montanus Willd. s.l. – Mountain Buttercup, Berg-Hahnenfuß 247 Ranunculus repens L. – Creeping Buttercup, Kriechender Hahnenfuß 246, 247 Rhamnaceae 336 f. Rhamnus cathartica L. – Buckthorn, Purgier-Kreuzdorn 322, 323 Rhododendron L. – Rhododendron, Alpenrose 239, 263, 264, 350 Rhodothamnus Rchb. – Dwarf-rhododendron 263 Rhynchospora alba (L.) Vahl – White Beak-sedge, Weißes Schnabelried 287, 288 ff. Ribes L. – Currant, Johannisbeere 324 Robinia pseudoacacia L. – Acacia, Robinie 324 Rosaceae 264 ff., 331 ff., 349 Rosa L. – Rose, Rose 271, 350 Rosa canina L. – Dog Rose, Hunds-Rose 266 ff., 271 Rosa corymbifera Borkh. – Corymb Rose, Hecken-Rose 266 ff., 271 Rosa glauca Pourr. – Glaucous Rose, Bereifte Rose 266 ff. Rosa montana Chaix – Mountain Rose, Berg-Rose Rosa pendulina L. – Alpine Rose, Alpen-Rose 266 ff., 271 Rosa rubiginosa L. – Sweet Briar, Wein-Rose 266 ff., 271 Rosa rugosa Thunb. – Japanese Rose, Kartoffel-Rose 239, 266 ff., 271, 349 Rosa spinosissima L. [= Rosa pimpinellifolia L.] – Burnet Rose, Bibernell-Rose 266 ff., 271 Rubiaceae 337, 346, 349 Rubus L. – Bramble, Raspberry, Brombeere, Himbeere 340 Rubus caesius L. – Dewberry, Kratzbeere 266 ff., 270 Rubus chamaemorus L. – Cloudberry, Moltebeere 270 Rubus fruticosus L. s.l. – Bramble (group), Artengruppe Echte Brombeere 241, 266 ff., 270 Rubus idaeus L. – Raspberry, Himbeere 239, 266 ff., 270 Rubus saxatilis L. – Stone Bramble, Steinbeere 270 Rumex acetosella L. – Sheep’s Sorrel, Kleiner Sauerampfer 325 f., 327
Index and vernacular names of plants
455
Salicaceae 258 ff., 331 ff., 342, 345 f. Salix L. – Willow, Weide 258 ff. Salix alba L. – White Willow, Silber-Weide 258 ff., 261 f., 340 Salix appendiculata Vill. – Large-leaved Willow, Großblättrige Weide 258, 260, 261 ff. Salix aurita L. – Eared Willow, Ohr-Weide 258 ff., 261 f. Salix bicolor Willd. – Tea Willow, Zweifarbige Weide 258 Salix caprea L. – Goat Willow, Sal-Weide 258 ff., 261 f. Salix cinerea L. – Grey Willow, Grau-Weide 258 ff., 261 f. Salix daphnoides Vill. – Violet Willow, Reif-Weide 258 Salix eleagnos Scop. [= Salix incana Schrank] – Olive Willow, Lavendel-Weide 258 ff., 261 f. Salix fragilis L. – Crack Willow, Bruch-Weide 258 ff., 261 f., 340 Salix hastata L. – Halberd Willow, Spieß-Weide 258 Salix herbacea L. – Forb Willow, Kraut-Weide 258 Salix myrsinifolia Salisb. [= Salix nigricans Sm.] – Dark-leaved Willow, Schwarzwerdende Weide 258 ff., 261 f. Salix pentandra L. – Bay Willow, Lorbeer-Weide 258 ff., 261 f. Salix purpurea L. – Purple Willow, Purpur–Weide 258 ff., 261 f. Salix repens L. s.l. – Creeping Willow (group), Artengruppe Kriech-Weide 258 ff., 261 f. Salix reticulata L. – Reticulate Willow, Netz-Weide 258 Salix retusa L. – Blunt-leaved Willow, Stumpfblättrige Weide 258 Salix triandra L. – Almond Willow, Mandel-Weide 258 ff., 261 f., 340 Salix viminalis L. – Osier, Korb-Weide 258 ff., 261 f. Salvia officinalis L. – Sage, Echter Salbei 278, 280 f., 350 Salvia nemorosa L. – Violet Sage, Steppen-Salbei 280 f. Salvia pratensis L. – Meadow Clary, Wiesen-Salbei 280 f. Salvia verticillata L. – Whorled Clary, Quirlblütiger Salbei 280 f. Sambucus ebulus L. – Danewort, Zwerg-Holunder 328 Sambucus nigra L. – Elder, Schwarzer Holunder 323 Sanguisorba minor Scop. – Salad Burnet, Kleiner Wiesenknopf Satureja L. – Savory, Bohnenkraut 278 Saxifragaceae 333 Schoenoplectus lacustris (L.) Palla – Bulrush, Gewöhnliche Teichsimse 286, 288 ff. Schoenoplectus tabernaemontani (C.C. Gmel.) Palla – Grey Club-rush, Salz-Teichsimse 286, 288 ff. Schoenus ferrugineus L. – Brown Bog-rush, Rostrotes Kopfried 287, 288 ff. Schoenus nigricans L. – Black Bog-rush, Schwarzes Kopfried 287, 288 ff. Scirpoides holoschoenus (L.) Soják – Round-headed Club-rush, Kugelbinse 287 Scirpus radicans Schkuhr – Creeping Club-rush, Wurzelnde Simse 287, 288 ff. Scirpus sylvaticus L. – Wood Club-rush, Wald-Simse 287, 288 ff. Sclerochloa dura (L.) P. Beauv. – Tough Grass, Hartgras 322 Scolochloa festucacea (Willd.) Link – Swamp Grass, Schwingelschilf 322 Scrophulariaceae 327, 332 ff., 345 f., 349 Scutellaria L. – Scullcap, Helmkraut 278 Secale cereale L. – Rye, Roggen 296 ff., 321, 351 Senecio L. – Ragwort, Greiskraut 279, 282 f. Senecio erucifolius L. – Hoary Ragwort, Rautenblättriges Greiskraut
456
Appendix
Sesleria Scop. – Moor-grass, Blaugras 319 Sesleria albicans Kit. ex Schult. [= Sesleria caerulea auct. non (L.) Arduino; = Sesleria varia auct.] – Blue Moor-grass, Kalk-Blaugras 296 ff., 319 Sesleria disticha (Wulf.) Pers. – Two-rowed Moor-grass, Zweizeiliges Blaugras 319 Sesleria ovata (Hoppe) Kern.– Moraine Moor-grass, Eiköpfiges Blaugras 319 Setaria P. Beauv. – Bristle-grass, Borstenhirse 294, 296 ff., 321 Silene L. – Campion, Leimkraut 349 Silene flos-cuculi (L.) Clairv. [= Lychnis flos-cuculi L.] – Ragged Robin, Kuckucks-Lichtnelke 328 Solanaceae 327, 337 Solanum melangena L. – Eggplant, Aubergine Solanum nigrum L. – Black Nightshade, Schwarzer Nachtschatten 328 Solanum tuberosum L. – Potato, Kartoffel 326, 327, 351 Solidago L. – Goldenrod, Goldrute 279, 282 f., 347 Sorbus L. – Whitebeam, Rowan, Mehlbeere, Eberesche 285 f. Sorbus aria L. (Crantz) s.l. – Common Whitebeam (group), Artengruppe Gemeine Mehlbeere 265 f., 266 ff. Sorbus aucuparia L. – Rowan, Eberesche 265 f., 266 ff. Sorbus chamaemespilus (L.) Crantz – Dwarf Whitebeam, Zwerg-Mehlbeere 265 Sorbus domestica L. – True Service-tree, Speierling 265 Sorbus intermedia (Ehrh.) Pers. – Swedish Whitebeam, Schwedische Mehlbeere 265, 266 ff. Sorbus torminalis (L.) Crantz – Wild Service-tree, Elsbeere 265, 266 ff. Sorghum Moench – Millet (partim), Mohrenhirse Spartina Schreb. – Cord-grass, Schlickgras 322 Spiraea L. – Bridewort, Spierstrauch 266 ff., 270 Stachys palustris L. – Marsh Woundwort, Sumpf-Ziest 280 f. Stachys sylvatica L. – Hedge Woundwort, Wald-Ziest 278, 280 f. Stipa L. – Feather-grass, Federgras 296 ff., 321 Stipa capillata L. – Hairy Feathergrass, Haar-Pfriemengras 319, 321 Succisa pratensis Moench – Devil’s-bit Scabious, Gewöhnlicher Teufelsabbiss 328 Syringa vulgaris L. – Lilac, Gewöhnlicher Flieder 324 Tamaricaceae 336 Tamarix L. – Tamarisk, Tamariske 239, 322, 323 Taraxacum F.H. Wigg. – Dandelion, Löwenzahn 279, 282 f. Taxaceae 244, 336 f. Taxus baccata L. – Yew, Europäische Eibe 244, 245 Tetraclinis articulata (Vahl) Mast. – Arartree, Gliederzypresse Teucrium montanum L. – Mountain Germander, Berg-Gamander 280 f. Teucrium scorodonia L. – Wood Sage, Salbei-Gamander 278, 280 f. Thalictrum flavum L. – Common Meadow Rue, Gelbe Wiesenraute 247 Thalictrum minus L. – Lesser Meadow Rue, Kleine Wiesenraute 246, 247 Thuja L. – Arborvitae, Lebensbaum Thymus L. – Thyme, Thymian 278 Thymus praecox Opiz – Wild Thyme, Frühblühender Thymian 280 f. Thymus pulegioides L. – Large Thyme, Feld-Thymian 280 f. Thymus serpyllum L. – Breckland Thyme, Sand-Thymian 278, 280 f.
Index and vernacular names of plants
457
Thymus vulgaris L. – Garden Thyme, Echter Thymian 280 f. Tiliaceae 257 f., 332 ff., 342, 349 Tilia L. – Lime, Linde 257, 323 Tilia cordata Mill. – Small-leaved Lime, Winter-Linde 257 Tilia petiolaris DC. – Weeping Silver Lime, Hänge-Silber-Linde 257 Tilia platyphyllos Scop. – Large-leaved Lime, Sommer-Linde 257 Tragus racemosus (L.) All. – Burdock-grass, Traubiges Klettengras 322 Trapaceae 324 Trapa natans L. – Water-nut, Wassernuss 324 Trichophorum alpinum (L.) Pers. – Alpine Deergrass, Alpen-Rasenbinse 286 Trichophorum cespitosum (L.) Hartm. – Deergrass, Gewöhnliche Rasenbinse 286, 288 ff. “Tricolpates” 329 Trifolium campestre Schreb. – Larger Hop Clover, Feld-Klee 273, 274 Trifolium dubium Sibth. – Suckling Clover, Kleiner Klee 273, 274 Trifolium repens L. – White Clover, Weiß-Klee 273, 274 Trisetum flavescens (L.) P. Beauv. – Yellow Oat-grass, Wiesen-Goldhafer 296 ff., 321 Triticum L. – Wheat, Weizen 296 ff., 321 Typhaceae 329 Ulmaceae 246 ff., 332 ff., 342, 349 Ulmus L. – Elm, Ulme 246 ff. Ulmus glabra Huds. [= Ulmus scabra Mill.; = Ulmus montana With.] – Wych Elm, Berg-Ulme 247 f., 248 Ulmus laevis Pall. [= Ulmus effusa Willd.] – European White Elm, Flatter-Ulme 237 247, 248 Ulmus minor Mill. [= Ulmus glabra Mill.; = Ulmus carpinifolia Gled.; = Ulmus campestris auct. non L.] – Smooth-leaved Elm, Feld-Ulme 247 f., 248 Ulmus x hollandica Mill. [= U. minor Mill. x U. glabra Huds.] – Dutch Elm, Bastard-Ulme 247 f., 248 Ulmus pumila L. – Siberian Elm, Sibirische Ulme Urticaceae 248 ff., 332 ff., 342, 345 Urtica dioica L. – Stinging Nettle, Große Brennessel 248 ff., 325 f., 333 Urtica kiovensis Rogow. – Kievian Nettle, Röhricht-Brennessel 248 Urtica urens L. – Small Nettle, Kleine Brennessel 248 f. Vaccinium myrtillus L. – Bilberry, Heidelbeere 263, 264 Vaccinium oxycoccos L. – Cranberry, Gewöhnliche Moosbeere 263, 264 Vaccinium vitis-idaea L. – Foxberry, Preiselbeere 263, 264 Vaccinium uliginosum L. – Bog Bilberry, Rauschbeere 263, 264 Valeriana dioica L. – Marsh Valerian, Kleiner Baldrian 328 Valerianella locusta (L.) Laterr. – Corn Salad, Echter Feldsalat 328, 351 Ventenata dubia (Leers) Coss. – Soft-bearded Oat-grass, Zweifelhafter Grannenhafer 322 Verbascum lychnitis L. – White Mullein, Mehlige Königskerze 325 f., 327 Verbascum thapsus L. – Great Mullein, Kleinblütige Königskerze 328 Veronica L. – Speedwell, Ehrenpreis 328 Veronica chamaedrys L. – Germander Speedwell, Gamander-Ehrenpreis Veronica montana L. – Wood Speedwell, Berg-Ehrenpreis Viburnum L. – Arrow-wood, Schneeball 323 Viburnum lantana L. – Wayfaring-tree, Wolliger Schneeball Vicia tenuifolia Roth – Fine-leaved Vetch, Feinblättrige Vogel-Wicke 273, 274
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Vicia faba L. – Horse Bean, Saubohne Vincetoxicum hirundinaria Medik. – Angle Pod, Schwalbenwurz 326 Violaceae 333 Viola L. – Violet, Veilchen 328 Viscum L. – Mistle, Mistel 346 Vitis vinifera L. – Grape Vine, Weinrebe 323, 350 f. Vulpia C. C. Gmel. – Vulpia Fescue, Federschwingel 322, 342 Xanthium albinum (Widder) H. Scholz – Cocklebur, Ufer-Spitzklette Zea mays L. – Maize, Mais 296 ff., 320, 321
Acknowledgements
459
Acknowledgements This work would not have been possible without the help of a great many friends and colleagues. I am particularly indebted to my academic teacher Prof. Dr. Matthias Schaefer (Göttingen), who offered a laboratory, logistic and financial support, as well as manifold discussion. Prof. Dr. Reinhard Remane (Marburg) and Dr. Pavel Lauterer (Brno) were open for my visits and confirmed many difficult species. They were also open for endless discussions on taxonomic problems and insect – plant relations. Dr. Michael Sayer, Dr. Michael Judas, and Dr. Jürgen Schauermann (all Göttingen) provided substantial help in logistics and software problems. Dr. Alan Stewart (Brighton), Dr. Stephen Wilson (Warrensburg) 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 Achtziger (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 (Oldenburg), Stuart McKamey (Washington, D.C.), Ute Oesterling (Mammendorf), Ing. András Orosz (Budapest), Katharina Raupach (Hanover), Gerd Reder (FlörsheimDalsheim), Dr. Ralf Rombach (Rech), Judith Rothenbücher (Göttingen), Prof. Dr. Wolfgang 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 discussion on various topics.
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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 during 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 identification. 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. Herbert Zettel (Vienna) generously offered access to museum or private collections or provided help in revision. Wolfgang Frommer (Cologne), Dr. Wilhelm Kolbe (Burscheid) and Holger Thüs (Frankfurt) provided unpublished manuscripts. Elena Nickel (Göttingen) 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 (Augsburg) stimulated and promoted research and interest in Auchenorrhyncha. I am also much indebted to my friend Anja Ritzel (Göttingen) as well as to my neighbours 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 particularly Dr. Lyubomir Penev and Teodor Georgiev, who did much effort to get this book into shape.
