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Species richness of gall midges (Diptera, Cecidomyiidae) in Europe (West Palaearctic): biogeography and coevolution with host plants

January 2010

January 2010
73:87-156

Authors:

Europe and its boundary (black line) as defined in the Fauna Europaea (2007) and its main geographical features shown in different colours: lowlands from sea level to 200 m a. s. l. (green); hilly country from 200 to 500 m a. s. l. (yellow ); uplands from 500 to 900 m a. s. l. (pale brown); lower mountains from 900 to 1800 m a. s. l. (brown); upper mountains from 1800 to 2400 m a. s. l. and over (dark brown). Important islands: 1 ? Iceland; 2 ? Faroe Islands; 3 ? Azores; 4 ? Madeira; 5 ? Canary Islands; 6 ? Cyprus.

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. Species numbers of Cecidomyiidae in Europe in 2009 and comparison with species numbers known in 1984

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Biological groups of gall midges of the family Cecidomyiidae in Europe.

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+29

. Species density of gall-inducing and plant-inhabiting members of the subfamily Cecidomyiinae in countries and islands of Europe

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Acta Soc. Zool. Bohem. 73: 87–156, 2009

ISSN 1211-376X

Species richness of gall midges (Diptera: Cecidomyiidae) in Europe (West

Palaearctic): biogeography and coevolution with host plants

Marcela SKUHRAVÁ & Václav SKUHRAVÝ

Bítovská 1227, CZ–140 00 Praha 4, Czech Republic; skuhrava@quick.cz

Received 25 September 2009; accepted 21 December 2009

Published 2 December 2010

Abstract. The present fauna of the family Cecidomyiidae Macquart, 1838 in Europe includes 1800 valid species in 270

genera. It is divided into three subfamilies: Lestremiinae Rondani, 1840 with 280 species in 33 genera, Porricondylinae

Enderlein, 1936 with 310 species in 62 genera and Cecidomyiinae Rondani, 1840 with 1210 species in 175 genera.

Only the last subfamily includes gall-causing and plant-inhabiting species. For the family Cecidomyiidae, Europe

is the most species-rich continent of the world. On average, each gall midge genus contains 6,6 species. The most

species-rich genus is Dasineura Rondani, 1840, with 230 species, followed by Contarinia Rondani, 1860, with 135

species. On the basis of the larval feeding habits, the family Cecidomyiidae may be divided into three biological

groups: 51% of species are phytophagous, 39% mycophagous, 8% zoophagous and the biology of about 2% remains

unknown. All species of the subfamilies Lestremiinae and Porricondylinae are mycophagous.

Fundamental sources for evaluation of the family Cecidomyiidae in Europe are 40,000 records about their

occurrence and distribution gathered by the present authors at more than 1850 localities in 17 countries of mainland

Europe and nine islands in the Mediterranean at altitudes from sea level up to 2645 m a. s. l. in the Alps in the period

1955–2009. All samples were obtained using the time and area unit collection method and results were enriched

by data collected by earlier researchers. Evaluation was based on identiﬁed and valid species, not on unidentiﬁed

and unnamed gall-inducing organisms.

This abundant data enabled production of maps of occurrence of individual gall midge species of the Czech

Republic, Slovakia, France, Spain, Portugal, Poland and Austria, and also graphs showing the vertical occurrences

of some species in several countries of Europe. These were published in several summarizing articles (Skuhravá

1991, 1994a, b, Skuhravá & Skuhravý 2009a, Skuhravá et al. 2005, 2006). The accumulated data makes it possible

to determine zoogeographical characteristics of single species occurring in Europe.

Here we summarize information about the occurrence of gall midge species of the main genera in Europe and

present results in maps of the following genera: Dasineura Rondani, 1840, Contarinia Rondani, 1860, Asphondylia

Loew, 1850, Rhopalomyia Rübsaamen, 1892, Lasioptera Meigen, 1818, Stefaniola Kieffer, 1913, Rabdophaga

Westwood, 1847, Jaapiella Rübsaamen, 1915, Resseliella Seitner, 1906, Macrolabis Kieffer, 1892, Baldratia

Kieffer, 1897, and Mayetiola Kieffer, 1896.

Evaluation of species richness using the average species number per locality showed that Hungary, with

29 species, France with 28 species and the Czech Republic with 26 species are the countries with the highest gall

midge species richness in Europe. The average species number decreases to the north and to the south of Europe.

Evaluation of species richness using the species density calculated for country or island unit area of 1000 km2

makes it possible to divide countries into ﬁve density groups. TheCzech Republic and South Tyrol (northern Italy)

are the countries with the highest species density, each with 189 gall midge species occurring per 1000 km2. Gall

midges of Europe belong to the following zoogeographic or biogeographic units: 71% are European species, 12%

Eurosiberian, 9.5% Mediterranean including Submediterranean, 1.3% Palaearctic or Euroasian, 4.5% Holarctic,

0.4% cosmopolitan and 1.3% are alien species.

Gall midge species are not uniformly distributed in Europe. Many species were found only once at the type

locality, some species are rare, others occur moderately frequently and only a few species occur very frequently.

It is possible to divide all species found in the area under study into six frequency groups by using the ﬁrst six

members of the geometrical progression with quotient “2” and coefﬁcient “a”, as follows: a.20 + a.21 + a.22 + a.23

+ a.24 + a.25. The most frequent species in the whole of Europe is Jaapiella veronicae (Vallot, 1827) causing galls

on Veronica chamaedrys; in northern Europe it is Iteomyia capreae (Winnertz, 1853) on Salix caprea; in southern

Europe both Phyllodiplosis cocciferae (Tavares, 1901) and Dryomyia lichtensteinii (Löw, 1878) causing galls on

Quercus coccifera, Q. ilex and Q. suber are equally frequent. Each part of Europe has its own gall midge species

which is the most frequent in the area characterized by particular conditions of the environment. Species number

changes with geographic position. The highest species number occurs in the temperate part of Europe covered

with mixed forests between latitudes 45° and 52° North. Species number decreases to the north and to the south

and is inﬂuenced importantly by climatic factors. Central Europe has favourable weather with mild temperatures

and with a relatively sufﬁcient amount of rainfall. Species number changes with increasing altitude of localities

together with changes in composition of vegetation: in lower parts the gall midge species number is relatively high

and it decreases with increasing altitude. Rhopalomyia luetkemuelleri (Thomas, 1893) causing galls on Artemisia

spicata and Jaapiella alpina (Löw, 1885) inducing galls on Silene acaulis are two gall midge species found at the

highest altitudes in Europe – at elevations of 2500–2700 m a. s. l. in the Alpine and subnivale zones. Oligotrophus

juniperinus (Linnaeus, 1758) inducing galls on Juniperus communis, occurs over a large altitudinal span including

sites in the planare zone at 190 m a. s. l. up to 2348 m a. s. l. in the Alps. It has considerable ecological tolerance

and has been recorded over a vertical range of more than 2000 m. Gall-inducing gall midges of Europe are one of

the best known gall-causing groups of animals in the world. They have to be the most suitable group for prognosis

of gall midge species numbers and their galls in the future. On the basis of data on the occurrence and distribution

of gall midge species gathered from the times of Linnaeus to the present it may be predicted that in Europe the

number of valid species in the family Cecidomyiidae may be increased by about four hundred to 2200 valid species

and the number of galls caused by gall midges by about two hundred to 1400 sorts of galls by the year 2050.

In Europe 915 gall-causing and plant-inhabiting species of the subfamily Cecidomyiinae are associated with 280

genera of host plants that belong in 72 plant families. On average one plant genus hosts 3,26 species of gall midges.

Only two gall midge species are associated with the Pteridophyta, 27 species with the Gymnospermae, 886 species

with the Angiospermae, of which 109 species are on Monocotyledones, and 777 species on Dicotyledones. Gall

midges are more closely associated with some plant families in which they ﬁnd more favourable host plant species

and genera for their development. The family Fabaceae hosts the highest number of gall midge species – 123 species

– and the family Asteraceae hosts 104 gall midge species. In Europe about 70% of gall-inducing species and plant

inhabiting species of the subfamily Cecidomyiinae are associated with herbaceous plants and only 30% with trees

and shrubs. For the comparison of relationships between gall midges and host plants we introduce the formula

– the index of mutual gall midge species and plant species occurrence:

IGS-PSO= NPS/NGMS where NPS is the number of plant species in a particular country and NGMS is the

number of gall midge species occurring in the country. This index indicates the degree of mutual occurrence of

gall midges on host plants in particular parts of Europe. The analysis of relationships between gall midges and host

plants produced the surprising ﬁnding that although the species number of ﬂowering plants in Europe increases

towards the south, the species number of gall midges decreases.

Galls of gall midge species are not equally distributed on the organs of their host plants. Species of the genus

Dasineura Rondani, 1840 cause galls mainly on buds and leaves; of the genus Contarinia Rondani, 1860 on

ﬂowers; of Asphondylia Loew, 1850 on ﬂowers and fruits, of Rabdophaga Westwood, 1847 on stems and buds;

of Rhopalomyia Rübsaamen, 1892 on buds; of Jaapiella Rübsaamen, 1915 on buds and ﬂowers, species of the

genus Macrolabis Kieffer, 1892 are mainly inquilines in galls of other gall midges and the species of Mayetiola

Kieffer, 1896, Lasioptera Meigen, 1818, Planetella Westwood, 1840 and Resseliella Seitner, 1906 develop in

galls mainly on stems. In Europe 94 gall midge species are associated with twelve main broad-leaved tree species

and 23 gall midge species with six coniferous tree species. Quercus robur is the host tree species with the highest

species gall midge number (15 species), followed by Populus tremula and Quercus cerris (each with 13 species),

Betula pendula and Tilia platyphyllos (each with 10 species).

Coevolution of gall-inducing and plant inhabiting species of the family Cecidomyiidae developed in parallel

with two groups of vascular plants, the Pteridophyta and the Spermatophyta, a long time ago in the geological

history of the Earth. One fossil record of Sequoiomyia kraeuseli (Möhn, 1960) found in fossil cones of Sequoia

langsdorﬁi (Taxodiaceae) supports the early coevolution of gall midges with conifers. Fossil galls of ten gall midge

species provide evidence of the coevolution of gall midges with ﬂowering plants in the Miocene and Pliocene

of the Tertiary in various parts of Europe. The gall-inducing and plant inhabiting gall midges of the subfamily

Cecidomyiinae probably started to develop at the end of the Mesozoic when they entered into interactions with

developing groups of ﬂowering plants. The main development of gall-inducing and plant inhabiting gall midges

came in the Pliocene and Miocene of the Tertiary.

This article is illustrated with 40 ﬁgures, including maps of geographical distribution of the main genera of

the subfamily Cecidomyiinae, maps of distribution areas of several interesting gall midge species and graphs

illustrating summaries of results obtained during investigations in Europe.

Distribution, Diptera, Cecidomyiidae, Lestremiinae, Porricondylinae, Cecidomyiinae, Palaearctic, West

Palaearctic, Europe, species number, species richness, frequency, geographic gradients, biogeography,

zoogeography, plant-animal relations, coevolution with host plants.

CONTENTS

Abstract .......................................................................................................................................... 87

Introduction ......................................................................................................................................... 89

Material and methods ........................................................................................................................................................ 92

Results ............................................................................................................................................................................... 95

Fauna ........................................................................................................................................................................... 95

Species richness .............................................................................................................................................. 97

Species density .......................................................................................................................................................... 103

Species richness of the main genera .......................................................................................................................... 106

Geographic distribution ............................................................................................................................................. 113

Geographic position and geographic gradients ......................................................................................................... 123

Climatic factors ......................................................................................................................................................... 125

Vertical distribution ................................................................................................................................................... 126

Frequency .................................................................................................................................................................. 130

Possibilities of ﬁndingadditional gall midge species number in Europe .................................................................. 132

Gall midges and host plants ....................................................................................................................................... 134

Mutual occurrence of gall midges and host plants .................................................................................................... 142

Distribution of galls on host plants ............................................................................................................................ 144

Abundance of gall midge species on host plant species ............................................................................................ 144

Coevolution of the family Cecidomyiidae with plants ............................................................................................. 145

Factors affecting gall midge species richness ........................................................................................................... 148

Acknowledgements ......................................................................................................................................................... 150

References ....................................................................................................................................................................... 150

INTRODUCTION

The Cecidomyiidae is one of the most species-rich families of Diptera and also contains the

largest group of gall-inducing animals in the world. Gagné (2004), in his World Catalog, listed

5451 species in 598 genera of living and fossil gall midges. Altogether 3113 species in 344 genera

occur in the Palaearctic Region of which more than 1800 species in 270 genera occur in Europe

(Skuhravá 2006a). The family Cecidomyiidae Macquart, 1838 is currently classiﬁed under three

subfamilies: Lestremiinae Rondani, 1840, Porricondylinae Enderlein, 1936 and Cecidomyiinae

Rondani, 1840. Only larvae of Cecidomyiinae are able to induce galls on host plants, larvae of

three subfamilies are mycophagous or mycosaprophagous. Adults are usually very small, incon-

spicuous ﬂies but the galls (in Latin: cecidium) that are caused by their larvae on various organs

of host plants are strange and striking formation (hence the common name “gall midges”). Larvae

of some species live free in ﬂower heads or stems of plants without making galls and others are

mycophagous or saprophagous. Zoophagous larvae are predators of larvae of other gall midges,

of aphids, mites, coccids, or other small arthropods and some of them may be used in biological

control of pests. Larvae of some species live as inquilines in galls of other gall midges or other

insects. The biology of many species of gall midges caught as adults in nature is completely unk-

nown (Skuhravá 1986, 1997a, Skuhravá et al. 1984a, b). In Europe about sixty gall midge species

are economically important, of them forty species cause damage to agricultural plants (Darvas et

al. 2000) and twenty to forest trees (Skuhravá & Roques 2000).

The Palaearctic Region is the largest zoogeographic or biogeographic region of the world oc-

cupying an area of about 52 millions km2 (about 39% of the Earth surface). It includes arctic and

temperate Eurasia and all islands surrounding the continent in the Arctic, in the sea of Japan and

the eastern half of the North Atlantic, the Mediterranean North Africa and the northern and central

parts of the Arabian Peninsula. In Asia the customary boundary has the High Himalayas chain as

main divider and the boundary in China and Japan runs at Northern Latitude 30° (Udvardy 1975,

Soós & Papp 1986). The Palaearctic Region is usually divided into two parts, namely the West

Palaearctic, and the East Palaearctic (sensu lato), that includes an extensive area from the Ural

Mountains across Siberia to the Kamchatka Peninsula in the most eastern part of Russia, or into

four subregions, viz. Euro-Siberian, Mediterranean, Central-Asian, and East-Palaearctic (sensu

stricto), situated in eastern Asia and including Japan and its islands, the Korean Peninsula, part

of Russia and part of China.

The West Palaearctic consists of the western part of the Euro-Siberian subregion (i.e. Europe)

and of the Mediterranean subregion, together including 48 countries and several islands that differ

substantially from each other by the geographic position and mainly by various areas. It is very

difﬁcult to compare gall midge species numbers of these countries and to evaluate them. For

example, France is the largest European country, occupying an area of 551,255 km2 and the gall

midge fauna includes 668 species of which 581 belong to Cecidomyiinae including gall-inducing

species (Skuhravá et al. 2005). On the other hand, Liechtenstein situated between Austria and

Switzerland is the smallest country occupying an area of 157 km2 and its gall midge fauna inclu-

des 65 species (Skuhravá & Skuhravý 1993a). It was therefore necessary to ﬁnd a method that

would make it possible to compare these values. We used two methods for evaluation of species

numbers of individual country: species number per unit area and average species number per

locality of area under study.

Europe is one of the ﬁve continents of the Earth covering an area of about 10,180,000 km2. It

is bounded to the north by the Arctic Ocean, to the west by the Atlantic Ocean, to the south by

the Mediterranean Sea, to the southeast by the Caucasus Mountains and the Black Sea and the

waterways connecting the Black Sea to the Mediterranean. To the east, Europe is divided from

Asia by the water divide of the Ural Mountains, the Ural River, and by the Caspian Sea. For the

project Fauna Europaea, the external geographic boundaries are deﬁned as follows: the deﬁned

area consists of the European mainland and Mediterranean islands, plus the Macaronesian islands

(Azores, Canary Islands, Madeira; excluding Cape Verde Islands), Franz Josef Land and Novaya

Zemlya, with Western Kazakhstan excluded (Fig. 1).

In Europe the main natural vegetation is forest. In temperate Europe, mixed forests with both

broadleaved and coniferous trees dominate. The most important species in central and western

Europe are broadleaved trees of Fagus sylvatica, Quercus robur and Q. petraea. In the north,

the taiga is a mixed forest with dominant trees of Picea abies, Pinus sylvestris and Betula spp.;

further north within Russia and extreme northern Scandinavia, the taiga changes to tundra. In the

Mediterranean the dominant tree is Olea europaea which is very well adapted to its arid climate.

In the semi-arid Mediterranean region scrub forests occur. a narrow east-west oriented strip of

Eurasian grassland, the steppe, extends eastwards from Ukraine and southern Russia as far as Hun-

gary. In Europe animals and plants have been substantially affected by the presence and activities

of man since ancient times. With the exception of Scandinavia and northern Russia, few areas of

untouched wilderness and original nature are at present found here, except for various national

parks. In the past, most (about 90%) of Europe was covered by forests. They extended from the

Mediterranean to the Arctic Ocean in the north. About half of these original forests disappeared

through the centuries owing to deforestration.

Southern parts of Europe, forming a zoogeographic sub-unit called the Mediterranean subre-

gion partly borders the Mediterranean Sea and also includes the northern part of Africa and part

of western Asia. The climate of this part is mild, winters are rainy and summers are hot and dry.

The landscape is covered with forests, woodlands and scrub. This part is one of the world‘s most

endangered biogeographic regions. Only about 4% of the original vegetation remains. Man and

his activities, including overgrazing, deforestation and conversion of lands for pasture, agriculture,

or urbanization, have degraded much of the region. Formerly this region was mostly covered with

forests and woodlands, but heavy human use has reduced much of the region to the sclerophyll

shrublands known as chaparral, matorral, maquis and garrigue. At present, 48 countries of vari-

ous size comprise Europe. Of them, the European part of Russia is the largest both in area and

population. The level of knowledge about galls on plants and gall-inducing organisms is very

uneven in Europe. Some countries have a long tradition and many researchers contributed to the

knowledge which resulted in high species numbers of the family Cecidomyiidae being recorded

in Germany, France, United Kingdom, Czech Republic, Poland and Austria. Up to the end of

the twentieth century only a few records of gall midge species and their galls were known from

Greece and islands in the Mediterranean Sea. At present we have little information about gall

midges and their galls in Albania and Moldova.

Gall midges are the richest group of gall-inducing animals in Europe. Houard (1908–1909) in

his two-volume book on plant galls of Europe and the Mediterranean included 6239 sorts of galls

that are induced by 1450 species of animals belonging to various taxonomical groups. Gall midges

(Diptera: Cecidomyiidae) totalled 420 species are the richest group of all gall-inducing animals,

constituting 29%. Buhr (1964–1965), in identiﬁcation keys for galls on the plants of Central and

Northern Europe, recorded gall midges as the richest group, including 580 species, i.e. 36% of

all insect species inducing galls. Kolomoets et al. (1989) recorded 328 gall-inducing species of

Diptera associated with plants in the European part of the USSR. Of those, 301 species (92%)

belong to Cecidomyiidae and 25 species to six other dipteran families. In Japan, situated in the

Eastern Palaearctic, the gall midges with 628 species (44%), including described and undescribed

species, are the richest group of gall-inducing arthropods (Yukawa & Masuda 1996). Of those,

only 176 species are described species.

Fig. 1. Europe and its boundary (black line) as deﬁned in the Fauna Europaea (2007) and its main geographical features

shown in different colours: lowlands from sea level to 200 m a. s. l. (green); hilly country from 200 to 500 m a. s. l. (yel-

low); uplands from 500 to 900 m a. s. l. (pale brown); lower mountains from 900 to 1800 m a. s. l. (brown); upper moun-

tains from 1800 to 2400 m a. s. l. and over (dark brown). Important islands: 1 – Iceland; 2 – Faroe Islands; 3 – Azores;

4 – Madeira; 5 – Canary Islands; 6 – Cyprus.

We summarized knowledge about gall midges in several European countries from various

points of view, including a review of the history of research, evaluation of contributions of earlier

researchers to the progress of studies, the biology, zoogeography, relations of gall midges to host

plants and economic importance of gall midges. We published such summarizing articles about

gall midges of Slovakia (Skuhravá 1991), of the Czech Republic (Skuhravá 1994a, b), of Greece

(Skuhravá & Skuhravý 1997), of France (Skuhravá et al. 2005), of Denmark (Skuhravá et al.

2006), of the Iberian Peninsula (Skuhravá et al. 2006), of Poland (Skuhravá et al. 2008) and of

Austria (Skuhravá & Skuhravý 2009a).

The aim of our present article is to show the species richness of gall midges in Europe and the

West Palaearctic, the abundance, density and frequency of gall midges and factors inﬂuencing

them, including geographic distribution and the effect of geographic gradients for distribution of

species numbers, the inﬂuence of climatic factors on species distribution and to predict increa-

sing of species number in the future. We analyse relations of gall midges to host plants and their

botanical families, try to elucide causes of their diversity and to indicate the probable process of

coevolution of the family Cecidomyiidae with plants in the history of the Earth.

MATERIAL AND METHODS

During the period 1955–2009 we investigated gall midges in 17 countries and nine islands at more than 1850 localities

spread throughout Europe from the northernmost situated locality at Harstad in the island Hinnoya in northern Norway

to the southernmost situated locality in Malta in the Mediterranean. Localities for our investigations were selected from

the sea level – some at the seaside of the Atlantic Ocean and many along the coast of the Mediterranean, and at various

altitudinal zones starting with lowlands up to mountains with the highest situated locality at an altitude of 2645 m a. s. l.

at Col du Galibier in Hautes Alpes in France. We gathered about 40,000 records about the occurrence of gall midge spe-

cies in Europe and in nine islands in the Mediterranean. These records are a unique collection of such data dealing with

a group of gall-inducing arthopods and they make it possible to evaluate the Cecidomyiidae from several points of view

and to determine factors that inﬂuence their species richness in Europe. Important is the fact that our considerations on

species richness and evaluations of factors inﬂuencing them are based on valid species, e.i. on identiﬁed species, not on

unidentiﬁed and unnamed gall-inducing organisms (Fig. 2).

During our faunistic investigations in various countries of Europe and in Asia we used at each locality the time and

area unit collection method that we established in 1955. This involves surveying each selected locality visually by walking

through it slowly over a period of at least one up to three hours. Two persons walk through various biotopes at the locality

and search for gall midges galls and collect them on herbs, shrubs and trees and examine visually buds, leaves, ﬂowers,

fruits and stems of herbaceous plants, shrubs and trees. The altitude and short characteristics of biotopes are noted. All

ﬁndings are recorded, including notes about the local abundance of species. Each locality was investigated only once by

this method. During excursions, several specimens of each host plant with galls were put in separate small plastic bags.

All species that were found were recorded in the protocol and all collected material was prepared. Several specimens of

each host plant with galls were preserved as herbarium items, several plants with galls were kept in plastic bags to obtain

living larvae, several plants with galls were placed in small emergence cages to obtain adults and ﬁnally several galls

with larvae were put into vials with 75% alcohol for future morphological studies or for studies of the digestive system

of larvae. Such samples taken from various habitats or microhabitats are very important.

The localities where we investigated gall midges in the Czech Republic, Slovakia and Austria are given in Fig. 3.

Localities were selected throughout the whole territory using the map with Ehrendorfer’s network (Ehrendorfer 1973), as

far as possible examining one locality in each oblong of a size 11×12 km. In a similar way we prepared our investigations

in Italy, Switzerland, Greece and other countries.

All data obtained during our investigations of gall midge faunas in Europe, was augmented with data of earlier authors,

and evaluated and analysed from two points of view, viz. from the geographic and from the zoological points of view

using the methods of Skuhravá (1987, 1994a, b, 1997b). From the geographic point of view, the horizontal occurrence is

presented by various species numbers (absolute species numbers found) and by average species numbers in countries of

Europe and by numbers of species related to area unit of 1000 km2. The vertical occurrence is shown as average species

numbers in the rising altitudinal zones. From the zoological point of view, it is possible to elaborate the characterization

of a particular species (zoogeographic diagnosis) and elaborate maps of distribution areas of a particular species which

may be compared with distribution areas of pertinent host plant species.

Fig. 2. Distribution of sites in Western Europe where the investigations of the family Cecidomyiidae were carried out in

countries and islands by the present authors (black circles) in the period 1955–2009, or where summaries of information

were published by other authors (white circles). See Table 1 for the abbreviations for countries; numbers for islands:

1 – Faroe Islands, 2 – Mallorca, 3 – Corsica, 4 – Sardinia, 5 – Sicily, 6 – Malta, 7 – Corfu, 8 – Samos, 9 – Crete.

We gathered not only our own data on occurrence of gall midges in European countries that we obtained in the course

of our expeditions but we collected also data on occurrence of gall midge species in countries of Europe found by earlier

researchers from the middle of the nineteenth century, when various researchers started investigations of galls, up to

the present. These data about occurrence were included in the part devoted to distribution of each gall midge species in

the Catalogue of Palaearctic Diptera (Skuhravá 1986). The paucity of records of gall midge species in some countries

motivated us to investigate in countries where the species number was low, as for example in Norway, Greece, northern

Italy (South Tyrol) and the islands in the Mediterranean.

Fig. 3. A–C. Distribution of localities in Central Europe where planned systematic faunistic investigations of the family

Cecidomyiidae were carried out by the present authors: a – in the Czech Republic investigations were made at 670

localities at altitudes from 116 m a. s. l. to 1603 m a. s. l. at Mount Sněžka, in the Giant Mountains, the highest point of

the country, in 1957–1982 (Skuhravá 1994a, b); b – in Slovakia investigations were made at 336 localities at altitudes

from 110 m a. s. l. to 2000 m a. s. l. in the High Tatras in 1969–1976 (Skuhravá 1991); c – in Austria investigations were

made at 274 localities situated at altitudes from 115 m a. s. l. up to 2500 m a. s. l. in the Alps in 1967 and 1991–1993

(Skuhravá & Skuhravý 2009a).

In Table 1, we summarized fundamental data and source of information on the Cecidomyiidae in alphabetical order of

countries and islands of Europe, abbreviations of countries as used in the maps; total gall midge species of Cecidomyiidae,

followed totals for the three subfamilies, the Lestremiinae, Porricondylinae and Cecidomyiinae; number of localities

examined by the present authors and the references. Islands belonging to Europe are here considered to be independent

geographic units, although some of them are components of continental countries as indicated in parentheses; for Spain

and Portugal, see the chapter “Area of study” in the catalog of Carles-Tolrá Hjorth-Andersen (2002). Cyprus and Malta are

independent island countries. The present level of knowledge in countries of Europe and in islands in the Mediterranean,

given as species numbers of Cecidomyiidae, is shown in Fig. 4.

It is necessary to emphasize that we comprehend and interpret the family Cecidomyiidae as a whole mainly in the

introductory parts of our article. In the parts devoted to evaluation of results we deal mainly with part of it – the subfamily

Cecidomyiinae – that includes gall-inducing and plant-inhabiting species which are associated with host plants. Members

of the subfamilies Lestremiinae and Porricondylinae are mycophagous and are not associated directly with host plants.

RESULTS

Fauna

The fauna includes all the species of animals found in a particular region, period, or special en-

vironment. The following six faunal regions or realms are recognized on the basis of terrestrial

animal species. The fauna of the Diptera is known from each of these regions and relevant cata-

logues have been published. We give in the following part the authors that prepared the family

Cecidomyiidae for catalogues of various parts of the world.

The fauna of Cecidomyiidae of the Palaearctic region including Eurasia, i.e.Europe and Asia

north of the tropics, and northern Africa, was prepared by Skuhravá (1986); the fauna of the Nearctic

region consisting of North America except for tropical Mexico by Foote (1965) and Gagné (1989a);

of the Neotropical region (situated in South and Central America including tropical Mexico) by

Gagné (1968, 1994); of the Afrotropical or Ethiopian region of Africa south of the Sahara and

part of southern Arabia by Harris (1980); of the Indomalayan or Oriental region covering tropical

Asia and associated continental islands by Gagné (1973) and Grover (1981); of the Austral-Asian

and Oceanian regions including Australia, New Guinea and islands in Oceania by Gagné (1989b).

In addition, Nearctic and Palaeartic (including Eurasia and northern Africa) regions are usually

combined into the Holarctic region. Many changes have occurred in the composition of faunas

of all regions in the last twenty years. They are reﬂected in the world catalog of Gagné (2004)

and in the analysis of world faunas in the article by Skuhravá (2006a).

The present or contemporary or extant fauna of the family Cecidomyiidae known in Europe

totals 1800 species in 270 genera (Table 2). Europe, a small continent if it is classiﬁed by size, is

the most species-rich continent of the world in terms of known species of gall midges. On average

each gall midge genus includes 6,6 species. Most genera are monotypic, containing only a single

species. On the other hand there are several large genera with many species, as follows: the species

richest genus Dasineura (230 species), followed by Contarinia (135 species), Asphondylia (50

species) and Rabdophaga (40 species) (see Table 7). The distribution of gall midge species in

Europe where they form single small faunas of countries and islands is given in Fig. 4.

The gall midges occurring in Europe may be divided into three main biological groups on the

basis of the larval feeding habits. If we evaluate the family Cecidomyiidae as the whole, we obtain

the following data: 51% of species are phytophagous, larvae of which feed on or in plant tissue,

forming galls or inhabiting plant organs without making galls; 39% are mycophagous, larvae of

which are associated with various fungi; in this group are usually included xylophagous species;

8% are zoophagous, larvae of which feed as predators on small animals, especially insects, and

the biology of about 2% remains unknown. See Table 2 and Figs 5, 6. If we evaluate the subfa-

mily Cecidomyiinae as separate unit that contains 1210 species, the separation of species into

Table 1. Sources of information on the family Cecidomyiidae in Europe. Abbreviations: CA – country abbreviation;

T – total species number; L – Lestremiinae; P – Porricondylinae; C – Cecidomyiinae; n – number of localities

country CA T L P C n references

Andorra AND 30 2 28 Skuhravá et al. 1996, 2002, 2006

Albania AL 19 19 Skuhravá 1986 (including 8 alien species)

Austria A 396 31 13 352 274 Skuhravá & Skuhravý 1967, 1992c, 1995, 2009a,

Skuhravá & Franz 1989

Belarus BY 38 0 0 38 Fiedorowiczówna 1931, Perepeczko-Baumanowa

1935

Belgium B 140 5 5 130 Lambinon 1957, 1958, 1976, Gosseries 1991

Bosnia and BH 169 169 Simova-Tošič et al. 2007

Herzegovina

Bulgaria BG 240 240 46 Skuhravá et al.1991

Canary Is. (Spain) 3 3 Skuhravá et al. 2002

Corfu (Greece) 49 49 8 Skuhravá & Skuhravý 2006b

Corsica (France) 60 60 10 Skuhravá & Skuhravý (in prep.)

Crete (Greece) 38 38 10 Skuhravá & Skuhravý 1997a

Croatia CR 232 232 Simova-Tošič et al. 2004

Cyprus CY 32 32 17 Skuhravá & Skuhravý 2004a

Czech Republic CZ 590 13 13 564 670 Skuhravá 1994a, 1994b, 2006b, 2009

Denmark DK 307 18 14 275 22 Skuhravá et al. 2006

Estonia 93 48 45 0 Spungis 1998, Spungis & Jaschhof 2000

Faroe Is. (Denmark) 5 5 5 Skuhravá & Skuhravý 2009c

Finland FIN 135 4 27 104 Hackman 1980

France F 668 33 54 581 17 Skuhravá & Skuhravý 2004c, d, Skuhravá et al. 2005

Germany D 836 125 58 653 26 Meyer & Jaschhof 1999

Greece GR 167 56 Skuhravá & Skuhravý 1997a, 2009b

211 Skuhravá & Skuhravý (unpubl.)

Hungary H 332 332 22 Skuhravá & Skuhravý 1999

Ireland IRL 87 87 O’Connor et al. 1999

Italy I 324 7 2 315 Skuhravá & Skuhravý 1994, Skuhravá 1995

508 27 10 471 14 Skuhravá & Skuhravý (unpubl.)

Italy, South Tyrol 312 312 124 Skuhravá et al. 2001, 2002, Skuhravá & Skuhravý

2003, 2005a, b, 2006a, 2007b, 2009d

Latvia LV 506 113 129 264 Spungis 2002, 2003, Spungis & Jaschhof 2000

Liechtenstein FL 65 65 9 Skuhravá & Skuhravý 1993a

Lithuania LI 224 25 35 164 Pakalniškis et al. 2000, Spungis & Jaschhof 2000

Luxembourg L 75 75 Lambinon et al. 2001

Macedonia MK 148 148 Simova-Tošič et al. 2007

Madeira (Portugal) 18 18 Skuhravá 2008

Mallorca (Spain) 33 33 7 Skuhravá & Skuhravý 2004b

Malta M 36 36 17 Skuhravá et al. 2002

Moldova MD 5 5

Montenegro MNE 85 85 Simova-Tošič & Skuhravá 2001

Netherlands NL 340 20 15 305 Nijveldt & Beuk 2002

Norway N 170 48 4 112 15 Skuhravá & Skuhravý (in preparation)

Poland PL 320 17 397 Skuhravá & Skrzypczyńska 1983, Skuhravá et al.

463 49 2008

Portugal P 122 122 Skuhravá et al. 1996, 2002, 2006

Romania RO 315 2 9 304 23 Skuhravá et al. 1972

Russia (European) RU 650 122 128 400 Skuhravá, Skuhravý & Prziboro (in prep.)

Samos (Greece) 34 34 8 Skuhravá & Skuhravý 2006b

Sardinia (Italy) 44 44 15 Skuhravá & Skuhravý 2002

Serbia SE 283 283 Simova-Tošič et al. 2000, Skuhravá & Skuhravý 1964

Sicily (Italy) 89 89 Skuhravá et al. 2007

Slovakia SK 362 2 5 355 336 Skuhravá 1991, 2006b, 2009

biological groups is quite different: phytophagous gall midges comprise 76%, zoophagous 12%,

mycophagous including xylophagous 9% and for 3% the larval biology remains unknown.

Species richness

The term species richness is used here for the records of all species occurring in Europe, com-

pared with species numbers of other parts of Palaearctic Region or with other continents. The

term species density is used for the number of species calculated for each country on a unit area

of 1000 km2 (i.e. 31.62×31.62 km).

A total of 1800 gall midge species (i.e. valid, described species) of the family Cecidomyiidae

are known from the whole territory of Europe (Table 2). Species are not equally distributed in

European countries. The occurrence of gall midge species in a country may be inﬂuenced by se-

veral factors, mainly by the geographic position, size of the country, composition of vegetation,

environmental changes and also by investigations that were done in a country by researchers in the

past and by investigations that are carried out at present. We tried to analyse some of these factors

on the basis of our rich experience obtained during long-term studies of this group in Europe.

In Table 3 we compare species numbers in countries of Europe ranked according to geogra-

phic area and simultaneously we evaluate the level of knowledge and level of recent reseach

of the family in each country. In the column called level of knowledge we evaluate results of

investigations of the family obtained by researchers in the past which are expressed in the pre-

sent species number of the family. For evaluation of the level of knowledge we use the adapted

classiﬁcation scale given in the publication of Merz et al. (1998) where they evaluate the quality

of knowledge of dipteran families in Switzerland, viz. as very good, good, medium, poor or very

poor. The level of recent research in the last column of the table is evaluated for the period of

Table 1. (continued)

country CA T L P C n references

Slovenia SLO 219 219 Simova-Tošič et al. 1996, Skuhravá & Skuhravý 1964

Spain E 229 17 0 212 Skuhravá et al. 1996, 2002, 2006, Skuhravý &

Skuhravá 1999

Sweden S 378 102 7 266 Skuhravá & Skuhravý (in prep.)

Switzerland CH 237 3 6 228 56 Skuhravá & Skuhravý 1997b, Merz et al. 1998

Turkey TR 70 70 Skuhravá et al. 2005

Ukraine UA 298 93 39 166 Skuhravá, Skuhravý & Berest (in prep.)

United Kingdom UK 620 62 32 526 7 Harris 1976, Chandler 1998, Skuhravá & Skuhravý

2007b

Table 2. Number of species and genera of Cecidomyiidae in Europe (Skuhravá 2010, unpublished data) and separation of

gall midge species into biological groups. Abbreviations: n – number of, myco – mycophagous, phyto – phytophagous,

xylo – xylophagous, zoo – zoophagous, un – unknown biology

subfamily n genera n species myco xylo phyto zoo un

Lestremiinae 33 280 280 0 0 0 0

Porricondylinae 62 310 310 0 0 0 0

Cecidomyiinae 175 1210 75 35 915 150 35

total 270 1800 665 35 915 150 35

% 100 37 2 51 8 2

Fig. 4. Distribution of species of the family Cecidomyiidae in countries and islands of Europe in 2009; islands belonging

to Europe are here considered to be independent geographic units although some of them are components of continental

countries. Cyprus and Malta are independent island countries.

about the last twenty years (1989–2009). We use the following three categories: no, few or many

investigations (Table 3).

The species number does not depend directly on the size of the country. European part of

Russia has the largest area but from the point of view of its species richness (600 species) is in

third place. Ukraine is second in area but with about 300 species only in the sixteenth place from

the point of species richness. Two large countries, France and Germany, are among the ﬁrst ﬁve

countries both from the point of view of area and species numbers. Finland with a large area of

337,030 km2 has only 135 species, in contrast to Bulgaria occupying a much smaller area (about

one third the area of Finland) where 240 species are known to occur. On the other hand, from

Slovenia, a relatively small country situated in the western part of the Balkan Peninsula, with an

area of 20,273 km2, a relatively high species number is known (219 species). Species numbers of

countries occupying quite similar areas may differ substantially, as for example 590 gall midge

species are known to occur in the Czech Republic with an area of 78,900 km2 (Skuhravá 2006b),

283 species in Serbia with an area of 77,500 km2 (Simova-Tošić et al. 2000) and only 87 species

in Ireland with an area 70,280 km2 (O’Connor et al. 1999).

Species richness of countries in Europe is importantly inﬂuenced by the level of knowledge,

by the number of researchers who contributed to increasing known species numbers in the past

and also by the level of recent research. Germany, France, United Kingdom and other countries

given in the upper part of Table 3 are countries where investigations of gall midges have a long

tradition and where many researchers contributed to the high level of knowledge of this group.

On the other hand, Albania, Finland, Ireland and Norway may be counted as countries where

gall midge faunas are poorly known and only a few researchers have investigated gall midges.

In Belarus and Moldova gall midges are very poorly known. Lestremiinae and Porricondylinae

of Estonia are well known, in contrast to Cecidomyiinae the number of which are not known to

us at present.

An important measurement for evaluating of species numbers of areas under study is the

average species number per locality. It may be calculated for a particular area of the country

where investigations were carried out at particular number of localities and gall midge galls were

collected by the time and area unit collection method. The average species number per locality may

be calculated according to the formula: ASN=sf/nl, where ASN is the average species number per

locality, sf is the total of all species found at all examined localities in the course of investigati-

ons in this area and nl is the number of localities where investigations were carried out. Average

species numbers of Cecidomyiidae in countries of Europe where investigations of present authors

Fig. 5. Biological groups of gall midges of the family Cecidomyiidae in Europe.

100

have been done are shown in Fig. 7. The highest value of average gall midge species numbers

was found in central and western Europe. In central Europe the highest value was calculated for

Hungary (29 species) and the Czech Republic (26), in western Europe for central France (28

species) and south-eastern France (25). This fact may be conditioned or resulted from the highly

Table 3. Species numbers of Cecidomyiidae related to country area and evaluation of the level of knowledge and recent

research. For distribution of the Cecidomyiidae in three subfamilies see Table 1

country country area (km2) number of level of level of recent

species knowledge research

Russia (European) 3,960,000 650 medium few

Ukraine 603,700 300 medium many

France 547,030 668 very good many

Spain 505,992 229 medium few

Sweden 449,964 378 good few

Norway 385,155 170 poor few

Germany 357,050 836 very good many

Finland 338,145 135 poor no

Poland 312,685 463 very good many

Italy 301,318 508 very good many

United Kingdom 244,820 620 very good many

Romania 238,392 315 medium no

Belarus 207,600 38 poor no

Greece 131,940 211 good many

Bulgaria 110,910 240 good few

Iceland 103,000 0 unknown no

Hungary 93,030 332 very good few

Portugal 91,568 122 medium no

Austria 83,858 396 very good few

Czech Republic 78,866 590 very good few

Serbia 77,474 283 good many

Ireland 70,280 87 poor few

Lithuania 65,200 224 medium many

Latvia 64,589 506 very good many

Croatia 56,542 232 good many

Bosnia and Herzegovina 51,129 169 good many

Slovakia 48,845 362 very good few

Estonia* 45,226 93 see the note*

Denmark 43,094 307 good few

Netherlands 41,526 340 very good few

Switzerland 41,290 273 very good few

Moldova 33,843 8 very poor no

Belgium 30,510 140 medium few

Albania 28,748 19 poor no

Macedonia 25,333 148 medium many

Slovenia 20,273 219 good many

Montenegro 13,812 85 medium few

Cyprus 9,251 32 good no

Luxembourg 2,586 75 medium few

Andorra 468 30 good no

Malta 316 36 good no

Liechtenstein 160 65 good no

* Note. Estonia: Level of knowledge: medium for Lestremiinae and Porricondylinae, unknown for Cecidomyiinae; no

investigations of the subfamily Cecidomyiinae have been done.

101

varied topography and ecology of these parts of Europe. Many varied factors inﬂuence the bi-

odiversity of organisms, both plants and animals. Central Europe has heterogeneous geological

substrata and considerable differences in composition of natural vegetation. In addition, Central

Europe is situated at the crossroads of routes of plant species that penetrate to this part from all

directions, viz. from southern and northern Europe, from western and eastern parts. The average

species numbers decreases to the north and also to the south of Europe. In the northern part of

the Scandinavian Peninsula the average species number was found to be from 1 to 5, in southern

parts of Europe, and in some islands in the Mediterranean (Corsica, Crete, Mallorca) the average

species number is around 9, in Malta 4.5 and in Cyprus 3 species.

In Table 4 the countries of Europe are arranged according to the present species numbers

and these numbers are compared with data obtained 25 years ago during the preparation of the

section on the family Cecidomyiidae for the Catalogue of Palaearctic Diptera (Skuhravá 1986,

Skuhravá et al. 1984).

At present, Germany is the country with the highest species number, followed by France, Eu-

ropean part of Russia, United Kingdom, Czech Republic, Italy, Latvia and Poland. Seven other

countries (Austria, Sweden, Slovakia, the Netherlands, Hungary, Romania, Denmark) have a re-

lative high species numbers from 307 to 396 species, nine countries (Ukraine, Serbia, Bulgaria,

Switzerland, Croatia, Spain, Lithuania, Slovenia and Greece) have medium species number from

211 to 298 species, six other countries (Norway, Bosnia and Herzegovina, Macedonia, Belgium,

Finland and Portugal) are species poor having from 122 to 170 species and the remaining countries

characterized by decreasing species numbers from 93 to 11 per country may be considered to be

species poor within the framework of Europe. It is clear that it is necessary to include into the

evaluation of species numbers not only simple species numbers but also other criteria.

We included in evaluation of species numbers of Europe also the historical aspect. We com-

pared species numbers of countries in Europe and their changes during the last 25 years (Table 4).

Fig. 6. Biological groups of gall midges belonging to the subfamily Cecidomyiinae in Europe.

102

Additions of recorded species are expressed in percentages of the species numbers given in 1984.

Greece and Cyprus are two countries with the highest additional species numbers during the period

1984–2009. Greece was the country where only 19 species were known in 1984. All species were

discovered only by chance. In 1994 we started our systematic faunal investigations at localities

Table 4. Species numbers of Cecidomyiidae in Europe in 2009 and comparison with species numbers known in 1984

country number of species number of species change

1984 2009 1984–2009 [%]

1 Germany1 586, 406 836 21

2 France 523 668 28

3 Russia (European Part)2 524 650 24

4 United Kingdom 603 620 3

5 Czech Republic 504 590 17

6 Italy 334 508 52

7 Latvia 506

8 Poland 360 463 29

9 Austria 256 396 55

10 Sweden 261 378 45

11 Slovakia 350 362 3.5

12 Netherlands 295 340 15

13 Hungary 240 332 38

14 Romania 294 315 7

15 Denmark 160 307 92

16 Ukraine 298

17 former Yugoslavia3 287 400 39

18 Serbia 283

19 Bulgaria 104 240 130

20 Switzerland 81 237 192

21 Croatia 232

22 Spain 80 229 186

23 Lithuania 224

24 Slovenia 219

25 Greece 19 211 1000

26 Norway 65 170 162

27 Bosnia and Herzegovina 169

28 Macedonia 148

29 Belgium 89 140 57

30 Finland 135 135 0

31 Portugal 118 122 3

32 Estonia 93

33 Ireland 87

34 Montenegro 85

35 Turkey 34 70 105

36 Liechtenstein 65

37 Belarus 38

38 Malta 36

39 Andorra 30

40 Cyprus 9 32 255

41 Albania 11 19 72

Notes: 1) In 1984 the German Federal Republic included 586 species and the German Democratic Republic 406 species;

2) In 1984 the European part of the Soviet Union was formed of the following parts using the recent country names: Russia,

Ukraine, Belarus, Moldova, Estonia, Latvia, Lithuania; each of them is now an independent country;

3) The former Yugoslavia is formed at present of six independent countries: Bosnia and Herzegovina, Croatia, Macedonia,

Montenegro, Serbia, Slovenia.

103

spread over the whole of Greece and on several islands. In the course of several years we raised

species number to 211 species, i.e. more than ten times (1000%). Only nine species were known

in Cyprus in 1984. After our investigations the species number increased to 32 species (an increase

of 255%). Bulgaria, Switzerland, Spain, Norway and Denmark are countries where the progress

in investigations of gall midges is expressed in increasing species numbers from 90 to 200%.

Other countries with lower increasing additional species during this time period usually posses

a relatively high species number that were gathered mainly in previous time periods. The species

number of Finland remains at the same level as before 25 years.

Species density

Species density refers to the number of species per unit area. We use this term for the number of

gall midge species calculated for each country or island per unit of 1000 km2, that is for a square

with four equal sides each long 31.62 km. This method makes it possible to compare species

numbers occurring in various parts of Europe, determine where the area or areas with the highest

species numbers occur and to elucidate reasons that cause inequalities in distribution of species

numbers in Europe.

Species density for gall midge species occurring per 1000 km2 may be calculated using the

formula of MacArthur & Wilson (1967): S = x/a0.25 where S is the number of species per area of

Figs. 7, 8. Evaluation of gall midge species richness. 7 (left) – average species numbers of the family Cecidomyiidae

in countries of Europe where systematic faunal investigations have been made during 1955–2009. 8 (right) – species

density of gall-inducing and plant-inhabiting species of the family Cecidomyiidae in Europe calculated per unit area of

1000 km2 for each country and island.

104

Table 5. Species density of gall-inducing and plant-inhabiting members of the subfamily Cecidomyiinae in countries

and islands of Europe

country or island area (km2) n species species density per 1000 km2

Albania 28,748 19 8.2

Andorra 468 30 37.5

Austria 83,858 352 116.5

Belgium 30,510 130 55.3

Bosnia and Herzegovina 51,129 169 63.3

Bulgaria 110,910 240 74.1

Canary Islands (Spain) 7,273 3 1.9

Corfu (Greece) 592 49 56.3

Corsica (France) 8,680 60 35.1

Crete (Greece) 8,260 38 22.3

Croatia 56,542 232 84.7

Cyprus 9,251 32 18.3

Czech Republic 78,866 564 189.2

Denmark 43,094 275 107.4

Faroe Islands (Denmark) 1,399 5 4.6

Finland 336,593 104 24.2

France 547,030 581 120.3

Germany 357,021 653 150.5

Greece 131,940 211 62.1

Hungary 93,030 332 107.1

Ireland 70,280 87 29.0

Italy 301,230 471 114.9

Italy, South Tyrol 7,400 312 189.1

Latvia 64,589 264 94.1

Liechtenstein 160 65 103.0

Lithuania 65,200 164 57.7

Luxembourg 2,586 75 57.7

Macedonia 25,333 148 66.1

Madeira (Portugal) 740 18 21.7

Mallorca (Spain) 3,411 33 24.4

Malta 316 36 48.6

Moldova 33,843 5 2.1

Montenegro 13,812 85 44.7

Netherlands 41,524 305 120.1

Norway 324,220 112 26.4

Poland 312,685 397 94.5

Portugal 91,568 122 39.4

Romania 238,391 304 76.0

Russia (European) 3,960,000 400 50.6

Samos (Greece) 479 34 42.5

Sardinia (Italy) 23,813 44 20.0

Serbia 88,361 283 91.3

Sicily (Italy) 25,426 89 40.1

Slovakia 48,845 355 134.5

Slovenia 20,273 219 104.2

Spain 504,851 212 42.9

Sweden 449,964 266 46.0

Switzerland 41,290 228 90.1

Ukraine 603,700 166 33.8

United Kingdom (GB) 244,820 526 132.8

105

1000 km2, x is the number of gall midge species found in the country and a is the whole area of

the country expressed in 1000 km2.

Results and calculations are given in Tables 5, 6 and in Fig. 8. Only countries of Europe whe-

re investigations were carried out or the summary of knowledge is available are given. Species

numbers in the third column involves only the species of the subfamily Cecidomyiinae includ-

ing gall-inducing and plant-inhabiting gall midges and inquilines, not the species of subfamilies

Lestremiinae and Porricondylinae, larvae of which do not induce galls.

On the basis of our calculations, two countries situated in Central Europe, the Czech Republic

and South Tyrol, the autonomous province in northern Italy, have the highest species density, each

with 189 gall midge species occurring per 1000 km2. Germany with 150 species in an area of

1000 km2 is at third place. United Kingdom, France, the Netherlands, situated in western Europe,

and Slovakia in Central Europe, also have a high level of species density, including 120–150 species

per area of 1000 km2. The countries of Central and western Europe have a higher species density

than the countries situated to the north and to the south. The very low species density have been

recorded in countries situated in northern Europe, viz. Finland, Norway, Sweden, and in the Faroe

Islands, situated in the Atlantic Ocean far from the European continent, and on the other hand the

countries and islands in southern Europe, viz. Madeira, Sardinia, Crete and Cyprus.

Species richness of gall midges in Europe is inﬂuenced by many factors. Above all by the

geographic position on the continent with different natural conditions of the realm. The number

of gall midges is much lower in the tundra and taiga covered with species-poor vegetation. In

countries of northern Europe, i.e. in Finland, Sweden and Norway, the relatively low gall midge

species number is the result of lower occurrence and poorer species composition of plants. In

temperate parts of Europe, i.e. in United Kingdom, Germany, France, Czech Republic, Slovakia,

Poland, Hungary, Romania and countries of the former Yugoslavia (Bosnia-Herzegovina, Croatia,

Serbia, Slovenia, Macedonia, Montenegro) broad-leaved and mixed forests cover large areas and

these forests contain many plant species which may be hosts of gall midges.

Table 6. Evaluation of gall-inducing and plant-inhabiting species of the family Cecidomyiinae occurring in countries of

Europe into ﬁve density groups according to the species density

density groups

very high high medium low very low

150 and more species 120–150 species 60–120 species 30–60 species 1–30 species

Czech Republic France Austria Andorra Albania

South Tyrol Netherlands Denmark Belgium Canary Is.

Germany Slovakia Hungary Corfu Crete

United Kingdom Italy Corsica Cyprus

Latvia Lithuania Faroe Islands

Liechtenstein Luxembourg Finland

Poland Malta Ireland

Serbia Montenegro Madeira

Slovenia Portugal Mallorca

Switzerland Samos Moldova

Bosna and Herzegovina Sicily Norway

Bulgaria Spain Sardinia

Croatia Sweden Belarus

Greece Ukraine

Macedonia

Romania

Russia

106

Species numbers per 1000 km2 in the southern parts of Europe, the so called Mediterranean

subregion, including southern parts of Greece, southern parts of Italy, southern part of the Iberi-

an Peninsula and islands in the Mediterranean Sea, are much lower than in Central Europe. For

example, the total value of species density for nine islands in the Mediterranean Sea is seven. Large

parts of these countries and islands were deforested in ancient times and at present the landscape

is covered mainly with sclerophyll vegetation (garrigue or maquis). Relations between plants

bearing galls to plants without galls are from 1:50 up to 1:80. We can conﬁrm that low numbers

of galls occurs in these areas on the basis of our own experience. During our investigations when

we used the time-area collecting method, we could speak about a success when we found 5 to

10 galls of gall midges at one locality, whereas in Central Europe we usually found galls of 15

to 30 species at one locality.

Species richness of a country, zoogeographic region, continent or their parts is expressively

inﬂuenced by the level of knowledge and degree of exploration of such area. Germany, France,

United Kingdom, Italy, Poland, Czech Republic and Austria are countries with a relative long tra-

dition in the study of gall midges and with many researchers who devoted their efforts to the study

of gall midges and their galls in the past. These countries have also the highest species numbers

of gall midges. On the other hand there are many countries where the study of gall midges does

not have so long a tradition or is in the initial phase.

Gall midge species numbers in Greece, South Tyrol and Austria were relatively low before

our investigations. Only 20 gall midge species were known from Greece before 1994. After our

exploration the species number increased to 210. In South Tyrol the species number increased

from 24 to 312 and in Austria from 170 to 396. The number of gall midge species in Greece and

South Tyrol increased more than ten times after using the time-area collecting method.

Surprising is the high abundance of gall midges in Kazakhstan. Fedotova (2000) gave 804 gall

midge species occurring on the area of 2,780,000 km2 where the exploration was made at more

than 60 localities. Kazakhstan with 112 gall midge species occurring in the area unit per 1000 km2

is one of the very well explored countries in Central Asia, the western part of which is adjacent

to eastern Europe. Richness of gall midge species in that area is connected with relatively species

rich vegetation. The ﬂora of Kazakhstan includes about 6000 plant species in 1118 genera and

161 families (Ryabushkina et al. 2008).

If we compare the species density of Europe with other parts of the Palaearctic region, species richness of the family

Cecidomyiidae in the East Palaearctic Region is known mainly owing to the studies of Yukawa (1971). He treated in his

revision 167 named and 17 unnamed gall midge species, forming a collection of 184 gall-inducing species of the family

Cecidomyiidae. After twenty ﬁve years of intensive studies Yukawa and his students (so called Yukawa’s Cecidological

School) were able to broaden the knowledge of Cecidomyiidae in Japan and to increase the species number from 184 to

628 (which is the number of sorts of galls caused by gall midges, including described and undescribed species), that is

3.4 times (Yukawa & Masuda 1996, Paik et al. 2004). Japan extending over an area of 377,835 km2 in the temperate to

subtropical zones of the East Palaearctic, is of quite similar size to Norway in the West Palaearctic where only 170 gall

midge species were found. In contrast, Norway is situated in the boreal part of northern Europe. Its northernmost part

has a mostly maritime subarctic climate and arctic tundra with species poor vegetation that does not give enough host

plants for gall midges.

Also of interest is the comparison of species numbers found in Europe with species richness of gall midges occurring

in North America. It occupies a territory of 24,490,000 km2, of which the USA has an area of 9,400,000 km2 and Canada

9,976,139 km2. Gagné (1989) in his book gives 891 gall midge species associated with 350 plant genera of 87 plant

families. If we count relations between gall midge species number known and the territory according to the formula of

MacArthur and Wilson, we obtain an average of 76 gall midge species per 1000 km2.

Species richness of the main genera

Numbers of species assigned to the genera of gall-inducing gall midges differ in the countries of

western and eastern Europe. In Table 7 we give species numbers of twelve of the most species-

107

Figs. 9–12. Distribution of species of the main genera of the family Cecidomyiidae in countries of Europe. 9 – Dasineura

Rondani, 1840. 10 – Contarinia Rondani, 1860. 11 – Asphondylia Loew, 1850. 12 – Rhopalomyia Rübsaamen, 1892.

910

11 12

108

-rich genera of the world and compare these numbers with numbers in the Palaearctic, Western

and Eastern Europe and in Kazakhstan, situated in south-western Asia, the area of which is greater

than western Europe.

Dasineura is the largest genus of the family Cecidomyiidae containing 490 species spread

over all the world. Most of these species – 335 – occur in the Palaearctic Region, 112 species in

the Nearctic Region and only a few species in other biogeographic regions. Larvae induce galls

mainly on leaves, on leaf and ﬂower buds on host plants of various plant families. In Europe

most species are associated with Fabaceae, Rosaceae, Asteracae, Brassicaceae and Lamiaceae.

Dasineura is the most species-rich genus in the Palaearctic Region and is abundant in Western

Europe where 230 species occur. The species number is much lower in Eastern Europe (Fig. 9).

In Kazakhstan it is also the most species-rich genus and is in ﬁrst position, together with the

genus Halodiplosis.

Contarinia with 340 species described in the world is in second place. In Western Europe

135 species were found. Larvae induce galls mainly on ﬂowers, ﬂower and leaf buds and leaves

or live freely in ﬂower heads of many host plants of various plant families. In Europe most spe-

cies are associated with Fabaceae, Asteraceae and Poaceae. Species are abundant in Western and

Central Europe, their number decreases to the east, southeast and southwest but they are relati-

vely abundant in Kazakhstan. Of 230 species that are distributed mainly in Central Europe, the

highest species numbers are found in Germany, Czech Republic, France and United Kingdom in

the latitudinal belt between 350 and 540 North. Lower species numbers were found in Norway,

Sweden and Finland and, on the other hand, species are relatively abundant in eastern parts of

southern Europe (Fig. 10).

Asphondylia with 270 species occurring in the world is in third place. Of these, 50 species

occur in Central and Western Europe (Fig. 11). Species number decreases to the south and east. In

Kazakhstan only nine species were found. Larvae induce galls on ﬂoweror leaf budsand fruits on

host plants of various plant families. Galls are usually associated with fungal mycelia (so called

ambrosia galls). In Europe the galls are induced on host plants belonging mainly to Fabaceae

and Lamiaceae. Asphondylia is considered to have its origin in the Neotropical Region and then

to have spread throughout the world (Möhn 1961, Tokuda & Yukawa 2007). We suppose that at

least one centre of the origin of the genus Asphondylia is in the Mediterranean area in southern

Europe in the western part of the Palaearctic Region. This area is considered to be one of the cen-

Table 7. Species numbers of the main genera of gall-inducing Cecidomyiinae in the world, in the Palaearctic Region,

Western and Eastern Europe in comparison with Kazakhstan (Central Asia)

genus World Palaearctic Europe Kazakhstan

Western Eastern

Dasineura Rondani, 1840 490 335 230 61 92

Contarinia Rondani, 1860 340 225 135 42 79

Asphondylia Loew, 1850 270 66 50 20 9

Rhopalomyia Rübsaamen, 1892 260 161 38 9 60

Lasioptera Meigen, 1818 125 50 27 9 3

Stefaniola Kieffer, 1913 108 108 6 0 74

Rabdophaga Westwood, 1847 80 40 40 18 11

Jaapiella Rübsaamen, 1915 80 80 35 9 51

Resseliella Seitner, 1906 47 27 23 9 1

Macrolabis Kieffer, 1892 50 50 35 5 22

Baldratia Kieffer, 1897 38 36 3 1 20

Mayetiola Kieffer, 1896 31 30 28 1 1

109

tres of origin of plant and animal species in Europe. In this area 50 described species and several

undescribed species of the genus Asphondylia are distributed which are associated with 13 plant

families. Interestingly, most species of this genus are associated with herbaceous plants, only

a few with trees and shrubs. No species have been found in Norway and Finland and the species

numbers in northern parts of Europe are low. Species of the genus Asphondylia occur abundantly

in southern Europe. This fact supports our hypothesis that at least one of the centres of diversity

of the genus Asphondylia is situated in Europe. In the Fig. 21 there are shown distribution areas

of several species of this genus.

Rhopalomyia, including 260 species in the world, is relatively species-rich in central, southeast

and western Europe where it contains 38 species (Fig. 12). Its number decreases markedly to the

south. No species were found in the islands of the Mediterranean Sea. Many Rhopalomyia species

were found and occur in Central Asia, mainly in Kazakhstan. Larvae induce galls on stems, buds

and leaves of host plants mainly of the family Asteraceae. Galls often occur on several organs

of the same host plant species. Alternation of plant organs is connected with the development of

several generations per year.

The genus Lasioptera includes 50 species in the Palaearctic Region and 27 of them occur in

Western Europe (Fig. 13). Larvae induce galls on stems of host plants of various families, mainly

on Poaceae and Apiaceae, or develop in fruits without making galls. Galls are usually associated

with fungal mycelia (so called ambrosia galls). Larvae of Lasioptera buhri develop in stems of

host plants belonging to several genera of three different families. Species of the genus Lasioptera

are relatively abundant in Central and Western Europe and numbers decreases northwards and

southwards. Also in Central Asia its numbers are low – only three species occur in Kazakhstan.

Stefaniola contains 108 species and is a Palaearctic genus. Species are distributed in Central

Asia, mainly in Kazakhstan and adjacent countries, and in North Africa. Only six species reach

to Europe where they occur in southwestern parts, in Spain and Portugal (ﬁve species) and in

Italy (one species), all very rare. Larvae induce leaf and bud galls on various species of Che-

nopodiaceae. Four species are associated with Salsola vermiculata, viz. Stefaniola bilobata, S.

gloma, S. parva and S. salsolae, S. vastita cause galls on Anabasis. All these species occur in the

Iberian Peninsula. Stefaniola mediterranea, causing galls on stems of Salicornia europaea and

Halocnemum strobilaceum, was found in Italy (Fig. 14).

Rabdophaga is a Holarctic genus with 40 species occurring in the Palaearctic Region and with

the same species number occurring in North America. Larvae cause stem, bud and leaf galls on

various species of Salix and one species, Rabdophaga giraudiana, is known to be associated with

Populus tremula and P. alba. Both genera of host plants belong to the family Salicaceae. The

species are abundant in central and northern parts of Western Europe and their number decreases

to the south and southwest (Fig. 15). Only one species was found in Sardinia and Mallorca in the

Mediterranean and no species was found in the remaining seven islands in the Mediterranean.

a relatively low species number was found in Kazakhstan.

Jaapiella is a Palaearctic genus with 80 species of which 35 species occur in Western Europe

(Fig. 16). Larvae induce galls on leaf and ﬂower buds and leaves of host plants of various plant

families. Some species are inquilines in galls of other gall midges. Higher species numbers occur

in Central and Western Europe and species numbers reduces towards the marginal parts of Europe.

Only a few species were found in northern and southern Europe, on islands in the Mediterranean

and no species was found in Mallorca in the Mediterranean Sea. a relatively large number of

species occurs in Kazakhstan.

Macrolabis is also a Palaearctic genus with 35 species distributed in Western Europe (Fig. 17).

Larvae induce galls on leaf and ﬂower buds on host plants of various families. About one half

of species are inquilines in galls of other gall midges. Species numbers decrease from Central

110

Figs. 13–16. Distribution of species of the main genera of the family Cecidomyiidae in countries of Europe. 13 – Lasioptera

Meigen, 1818. 14 – Stefaniola Kieffer, 1913. 15 – Rabdophaga Westwood, 1847. 16 – Jaapiella Rübsaamen, 1915.

13 14

15 16

111

and Western Europe to the south and no species was found in Macedonia and Montenegro in the

Balkan Peninsula and in the islands in the Mediterranean, with the exception of Corsica. Species

of this genus were found abundantly in Kazakhstan.

Baldratia is a Palaearctic genus with 36 species and with two species occurring in Africa. Larvae

induce galls on stems of various Chenopodiaceae in Central Asia and only three species occur in

Western Europe. Only one species, Baldratia salicorniae, causing galls on Salicornia fruticosa,

is native to Europe and occurs scattered along the coast of the Black Sea, along the coasts of

the Mediterranean Sea and in several islands in this sea. Galls of B. salicorniae were found also

at several localities in southern France, Spain, Portugal and in southern parts of England on the

coast of the Atlantic Ocean. Baldratia suaedae and B. similis are species which have their origin

in the Middle East. Galls of Baldratia suaedae on Suaeda vera, described from material found in

Israel, were discovered at two localities in Europe, viz. on the Canary Islands and in Monegros in

north-eastern Spain. Galls of Baldratia similis causing galls on Anabasis aphylla, described from

material found in Iran, were found at one locality in south-eastern Russia (Fig. 18).

Mayetiola is a Palaearctic genus containing 30 species with only one species native to North

America. Larvae induce galls on stems of various species and genera of Poaceae. Mayetiola de-

structor, a serious pest of cereals that occurred abundantly in Europe in the past, is considered to

have its origin in south-eastern Europe, probably in the Ukraine. It has been accidentally introduced

with cereals into other biogeographic regions where it causes serious damage. Most species were

described from western Europe, only one species occurs in eastern Europe and in Kazakhstan.

Species of this genus occur abundantly in Central and Western Europe, their number reduces

northwards and southwards and no species were found in Croatia and Slovenia in the Balkan Pe-

ninsula or on seven islands in the Mediterranean. Mayetiola destructor is the only species known

to occur in Sardinia and Malta. It has been probably introduced there with cereals (Fig. 19).

Resseliella is a genus with cosmopolitan distribution and diverse biology. Of 43 species occur-

ring in the Holarctic, 27 occur in the Palaearctic and 16 in the Nearctic region. Larvae of many

Palaearctic species live under the bark of trees and shrubs, causing them damage, and some species

are considered to be pests. Several species are associated with resin of coniferous trees or with

cambium liquid ﬂowing from stems of young broad-leaved trees that were primarily wounded

Table 8. Distribution of species of the family Cecidomyiidae occurring in Europe and in zoogeographical units of the

world

taxon zoogeographical unit

European Euro- Mediter- Palaearctic Holarctic cosmopolitan alien

siberian ranean

Cecidomyiidae

(1800 species) 1278 216 170 25 81 7 23

% 71.0 12.0 9.5 1.3 4.5 0.4 1.3

Lestremiinae

(280 species) 181 54 0 0 43 2 0

% 64.6 19.3 15.4 0.7

Porricondylinae

(310 species) 280 12 0 5 13 0 0

% 90.3 3.9 1.6 4.2

Cecidomyiinae

(1210 species) 817 150 170 20 25 5 23

% 67.5 12.5 14.0 1.7 2.0 0.4 1.9

112

Figs. 17–20. Distribution of species of the main genera of the family Cecidomyiidae in countries of Europe. 17 – Macrolabis

Kieffer, 1892. 18 – Baldratia Kieffer, 1897. 19 – Mayetiola Kieffer, 1896. 20 – Resseliella Seitner, 1906.

17 18

19 20

113

by animals (Dengler 2004). European species are associated with 11 plant families: six species

with Pinaceae and four with Rosaceae, other families each hosting one gall midge species. The

biology of several species remains unknown. Most species are distributed in Central and Western

Europe, their number decreases in southern Europe and no species was found in Mallorca, Corsica

and Sardinia and Cyprus (Fig. 20).

Geographic distribution

The gall midge species occurring in Europe may be divided, according to their overall distribution

in the world, into six zoogeographic units: European, Eurosiberian, Mediterranean (including

Submediterranean), Palaearctic or Euroasian, Holarctic and cosmopolitan (Table 8). In addition,

species that penetrated into Europe from other continents and are members of the present European

Fig. 21. Distribution areas of ﬁve species of the genus Asphondylia Loew, 1850 in Europe shown by lines of their limits:

A. calycotomae Kieffer, 1912 causing galls on Calycotome spinosa (Fabaceae) (blue line); A. melanopus Kieffer, 1890

inducing galls on Lotus corniculatus (Fabaceae) (red line); A. punica Marchal, 1897 evoking galls on Atriplex halimus

(Chenopodiaceae) (brown line); A. sarothamni (Loew, 1850) producing galls on Cytisus (Sarothamnus) scoparius

(Fabaceae) (green line); A. ulicis Trail, 1873 causing galls on Ulex europaeus (Fabaceae) (yellow line). A. calycotomae

and A. punica are Mediterranean species, A. ulicis is an Atlantic species, A. sarothamni is a sub-Atlantic species and A.

melanopus a European species.

114

gall midge fauna are designated as alien. On the other hand, some European gall midge species

that spread into other continents are considered to be immigrants there and some European gall

midge species were introduced in other continents of the world as agents for biological control

of pests or of weeds. The distribution of species of the subfamily Cecidomyiinae including the

gall-causing and plant inhabiting species is better known than the distribution of species belonging

to subfamilies Lestremiinae and Porricondylinae that require different sampling methods.

Fig. 22. Distribution area of Mikiola fagi (Hartig, 1839) in countries of Europe including its occurrence in single countries

(black circles) and its absence (white circles), and distribution area of its host plant, Fagus sylvatica L. (Fagaceae) (black

line). Distribution area of the host plant after Meusel et al. (1965–1992).

115

Of 1800 species of the family Cecidomyiidae forming the present fauna of Europe, 71% are

European, 12% Eurosiberian, 9.5% Mediterranean and Submediterranean, 4.5% Holarctic, 1.3%

Palaearctic, 0.4% cosmopolitan and 1.3% alien species. In the best known group of gall midges,

in the subfamily Cecidomyiinae, we found all types of distribution. Of interest is the fact that in

the subfamilies Lestremiinae and Porricondylinae there were not found gall midge species with

Mediterranean or Submediterranean types of distribution and only a few European species occur

or have been found beyond the boundary of Europe, in Asia or in North America.

European species occur only in Europe and are considered to have the centres of their origin

in Europe. Altogether 1278 species (71%) of the family Cecidomyiidae known to occur in Europe

belong in this group, i.e. nearly three quarters of all species. Areas of their distribution may range

from very small to large. Most of them are known only from the type locality in some country of

Europe and have never been recorded again after the original description. Several species have

small areas of distribution, occurring only in the country where they were discovered and in one or

two adjacent countries. Some species occur abundantly in several countries of Central Europe and

some of them extend southwards to the Mediterranean and even to North Africa, others extend to

south-east or to western Turkey or the Caucasus. Typical representatives of European gall midges

are Mikiola fagi galling leaves of Fagus sylvatica (Fig. 22), Macrodiplosis pustularis with galls

on leaves of Quercus robur and its relatives, Acericecis vitrina causing galls on leaves of Acer

pseudoplatanus, Zygiobia carpini inducing galls on leaves of Carpinus betulus, and Taxomyia

taxi causing galls on buds of Taxus baccata. Most species of the subfamily Porricondylinae and

about two thirds of species of the subfamily Lestremiinae have European distributions.

Several gall midge species that are members of the European fauna have a particular or

strange distribution area. Gall midge species associated with Cytisus (Sarothamnus) scoparius,

Fig. 23. Distribution area of Kochiomyia kochiae (Kieffer, 1909) (black circles) on Bassia prostrata (L.) Scott (= Kochia

prostrata (L.) Schrad.) (Chenopodiaceae) (black line), a Pontic-Pannonian species. Distribution area of the host plant

after Meusel et al. (1965–1992).

116

viz. Asphondylia sarothamni, Contarinia scoparii, Dasineura tubicoloides, Jaapiella sarothamni

and Janetiella tuberculi, are European and sub-Atlantic species occurring abundantly along the

shores of the Atlantic Ocean, penetrating inland and reaching, with their host plant, eastwards to

the Czech Republic. Kochiomyia kochiae is a Pontic-Pannonian species causing galls on Bassia

prostrata (= Kochia prostrata) (Fig. 23). It occurs abundantly along the shores of the Black Sea

(Pontus Euxinus) and in the Pannonian lowlands in Hungary (Skuhravá & Skuhravý 1999). The

galls were found also in southern France and in north-eastern Spain where this species reaches the

Fig. 24. Disjunct area of distribution of Jaapiella alpina (Löw, 1877) (black circles) and its host plant Silene acaulis (L.)

Jacq. (Caryophyllaceae) (black line) in Europe. Distribution area of the host plant after Meusel et al. (1965–1992).

117

most western limits of its remarkable distribution area. The gall midge Jaapiella alpina causing

galls on Silene acaulis, the host plant species of typical arctic-alpine distribution, has a disjunct

distribution area with scattered occurrence at very high elevations in the mountains of Europe

(Fig. 24).

Eurosiberian species inhabit the Eurosiberian subregion of the Palaearctic region. About one

eighth, i.e. 216 species (12%) of the Cecidomyiidae known to occur in Europe belong in this

group. They have centres of origin in Europe where they occur usually abundantly and extend at

least to Western Siberia, with some of them reaching to Central Siberia and only few reaching

to Eastern Siberia and to the most eastern part of the Palaearctic Region, to the Far East. Typical

representatives of Eurosiberian species are Harmandiola cavernosa, H. globuli, H. populi causing

galls on leaves of Populus tremula, Lasioptera rubi inducing galls on stems of Rubus idaeus and

its relatives, Iteomyia capreae on Salix caprea (Fig. 26), Geocrypta galii on Galium mollugo and

Dasineura urticae on Urtica dioica. Thecodiplosis brachyntera galling needles of Pinus sylvestris

and Pinus mugo occurs abundantly in Central Europe where it is classed as a pest causing serious

outbreaks from time to time (Skuhravý 1991). Its occurrence decreases eastwards and the galls

were found in southern part of Central Siberia (Skuhravá & Skuhravý 1993b).

Mediterranean species have centres of origin in the Mediterranean area. About 170 species, i.e.

9.5% of the family Cecidomyiidae known to occur in Europe belong in this group. They occur along

the shores of the Mediterranean and are associated with Mediterranean host plant species. Some

of them occur over greater areas and a few species reach the northern limits of their distribution

Fig. 25. Mediterranean distribution area of Dasineura oleae (Angelini, 1831) (black circles) and its host plant, Olea

europaea L. (Oleaceae) (black line) in the southern part of Europe and westernmost Asia. Distribution area of the host

plant after Meusel et al. (1965–1992).

118

areas in Central Europe. In such case they are deﬁned as Submediterranean species. Nine gall

midges associated with Quercus cerris, viz. Dryomyia circinans, Contarinia quercicola, C. subu-

lifex, Dasineura tubularis, Janetia cerris, J. homocera, J. nervicola, J. pustularis and J. szepligetii

that occur abundantly in the Mediterranean area reach the northern limits of their distribution in

the eastern part of Austria, southern part of the Czech Republic and of the Slovak Republic.

Typical Mediterranean species are Dasineura oleae inducing galls on leaves of Olea europaea

(Fig. 25), Phyllodiplosis cocciferae and Dryomyia cocciferae causing galls on leaves of Quercus

coccifera; four species associated with Quercus ilex, viz. Contarinia ilicis, C. luteola, Dasineura

ilicis and Dryomyia lichtensteinii; Dasineura asparagi causing galls on Asparagus aphyllus;

four species associated with Artemisia herba-alba, viz. Rhopalomyia hispanica, R. navasi, R.

producticeps and R. tavaresi; Braueriella phillyreae, Dasineura rufescens and Probruggmanniella

phillyreae which are associated with various species of Phillyrea. Also six gall midge species

associated with various Erica spp. have Mediterranean type distributions. Dasineura broteri,

D. elegans and D. zimmermanni occupy only small distribution areas. In contrast, Dasineura

ericaescopariae, Myricomyia mediterranea and Wachtliella ericina occupy larger areas in the

Mediterranean. W. ericina reaches up to southern England, to southern Denmark and to the

southern part of the Czech Republic in Central Europe. Asphondylia coronillae causing galls on

buds and pods of Coronilla emerus is a Mediterranean species that occurs abundantly in southern

Europe and penetrates up to Switzerland and northern Italy. Lasioptera eryngii causing large

galls on stems of Eryngium campestre occurs scattered in the whole Mediterranean and occupies

a large area reaching up to Turkey. It penetrates northwards up to the Czech and Slovak Republics

where the most northern boundary of its distribution area runs. It may be designated as a typical

Submediterranean species.

Palaearctic or Euro-Asian species are species that inhabit Europe or the Eurosiberian subregion

and at least one of the other Palaearctic subregions, i.e. Central Asian or East Palaearctic subregion.

Only about 25 species (1.3%) of the family Cecidomyiidae known to occur in Europe belong in

this group, mainly species caught as adults, viz. Trisopsis abdominalis, T. karelini, T. punctiven-

Fig. 26. Eurosiberian distribution area of Iteomyia capreae (Winnertz, 1853) (black circles) and its host plant, Salix

caprea L. (Salicaceae) (black line) including Europe, Siberia and the East-Palaearctic subregion. Distribution area of the

host plant after Meusel et al. (1965–1992).

119

tris, T. tyroglyphi, Stomatosema kamali of the subfamily Cecidomyiinae and Peromyia truncata

and Polyardis bispinosa of the subfamily Lestremiinae. Of gall-causing species of the subfamily

Cecidomyiinae, Etsuhoa sabinae and Xerephedromyia ustjurtensis, have disjunct Euro-Asian areas

of distribution. Galls of Etsuhoa sabinae on Juniperus sabina occur rarely in mountains of Europe

and were also found in mountains of eastern Kazakhstan. Galls of Xerephedromyia ustjurtensis on

Ephedra distachya were discovered in western Kazakhstan near the Caspian Sea, in Turkmenistan,

Jordan, in southern Ukraine, northern Spain and southern France and Italy. Such types of disjunct

distribution may be denominated as Mediterraneo-Turanian. Also Lasioptera umbelliferarum

causing galls on Hippomarathrum (formerly Seseli) has a disjunct distribution area. Galls were

found in Sicily (southern Italy), in southern Russia, Georgia, Azerbaijan, Iran and Israel.

Holarctic species occur simultaneously in the Palaearctic and in the Nearctic regions. 81 species

(4.5%) of the family Cecidomyiidae known to occur in Europe belong in this group. The most

abundant are the Lestremiinae with 43 species, followed by the Cecidomyiinae with 25 species

and at the last position are the Porricondylinae with only 13 species. Camptomyia multinoda and

Porricondyla nigripennis of the subfamily Porricondylinae, Aphidoletes aphidimyza, A. thompsoni

and A. urticaria, Coquillettomyia dentata and C. lobata, Mycodiplosis coniophaga and M. inimica

of the subfamily Cecidomyiinae may be considered to be truly Holarctic species. These species

are mycophagous or zoophagous and are not associated with speciﬁc host plants. On the other

hand, several gall midge species associated with their host plants, mainly with cereals, fruit trees

and fodder crops, that occur in both regions, may be classed as secondarily Holarctic because

their host plants are of European origin and gall midges were imported from Europe into North

America with their host plants. For example: Contarinia pyrivora and Dasineura pyri causing

galls on Pyrus communis; Dasineura mali on Pyrus malus (Fig. 28); Dasineura alopecuri devel-

oping in inﬂorescences ofAlopecurus pratensis; D. gentneri and D. leguminicola associated with

Fig. 27. Cosmopolitan distribution of the gall midge Lestremia cinerea Macquart, 1826 (black circles) in the biogeogra-

phical regions of the world: 1 – Palaearctic, 2 – Nearctic, 3 – Afrotropical, 4 – Indomalayan, 5 – Oceanian, 6 – Australian,

7 – Neotropical, 8 – Antarctic regions.

120

Trifolium spp. In some cases it is not quite clear where a particular gall midge species is native

and to which part it was introduced.

Cosmopolitan species are species that occur on all continents of the Earth having a world-wide

distribution. Only seven species of the family Cecidomyiidae known to occur in Europe may be

assigned to this group: Anarete johnsoni, Lestremia cinerea (Fig. 27) and Monardia modesta (Les-

tremiinae), Didactylomyia longimana, Feltiella acarisuga, Stomatosema obscura and Silvestriola

cincta (Cecidomyiinae) were found on several continents and are considered to be cosmopolitan

in distribution. The origin of most of them is unknown. Mayetiola destructor developing on cere-

als is considered to be a primarily European species that has been imported into other continents

probably with straw.

Alien species (non-native, exotic, non-indigenous, foreign) are species introduced outside their

normal distribution. Invasive alien species are alien species whose establishment and spread

modify ecosystems, habitats, or species (DAISIE 2009). In the fauna of gall midges of Europe

23 species were found that are noted as alien or invasive species (Table 9). Most of them are

associated with various plant families and four species are predators. They were introduced into

Table 9. Alien gall midge species occurring in Europe, their host species, plant family and origin; zoophagous species are

marked by an asterisk before the species name

gall midge species host species plant family origin

Asphondylia buddleia Felt, 1935 Buddleia racemosa Loganiaceae North America

Clinodiplosis cattleyae (Molliard, 1903) Cattleia Orchidaceae Central and South

Americas

Contarinia citri Barnes, 1944 Citrus spp. Rutaceae Africa

Contarinia quinquenotata (Löw, 1888) Hemerocallis fulva Liliaceae temperate Asia

Dasineura gibsoni Felt, 1911 Cirsium Asteraceae North America

Dasineura gleditchiae (Osten Sacken, 1866) Gleditchia triacanthos Caesalpiniaceae North America

Dasineura oxycoccana (Smith, 1890) Vaccinium spp. Ericaceae North America

*Dicrodiplosis pseudococci (Felt, 1914) Planococcus citri Africa

(Pseudococcidae)

*Epidiplosis ﬁlifera (Nijveldt, 1965) Ceratoplates ﬂoridensis Eastern Asia

(Coccidae)

*Feltiella acarisuga (Vallot, 1827) Tetranychidae (Acari) unknown

Horidiplosis ﬁcifolii Harris, 2003 Ficus benjamina Moraceae tropical Asia

Janetiella siskiyou Felt, 1917 Chamaecyparis Cupressaceae North America

lawsoniana

*Lestodiplosis aonidiellae Harris, 1968 Aonidiella aurantii Africa

(Diaspidae)

Obolodiplosis robiniae (Haldeman, 1847) Robinia pseudoacacia Fabaceae North America

Orseolia cynodontis Kieffer

et Massalongo, 1902 Cynodon dactylon Poaceae Africa

Procontarinia matteiana Kieffer

et Cecconi, 1906 Mangifera indica Anacardiaceae tropical Asia

Prodiplosis vaccinii (Felt, 1926) Vaccinium spp. Ericaceae North America

Prodiplosis violicola (Coquillett, 1900) Viola spp. Violaceae North America

Resseliella conicola (Foote, 1956) Picea sitchensis Pinaceae North America

Rhopalomyia chrysanthemi (Ahlberg, 1939) cultivated Asteraceae temperate Asia

Chrysanthemum

Rhopalomyia grossulariae Felt, 1911 Ribes grossularia Grossulariaceae North America

Stenodiplosis panici Plotnikov, 1926 Panicum spp. Poaceae temperate Asia

Stenodiplosis sorghicola (Coquillett, 1899) Sorghum spp. Poaceae Africa

121

Europe unintentionally from other parts of the world. Typical representatives are Obolodiplosis

robiniae causing galls on leaﬂets of Robinia pseudoacacia (Fig. 29), and Dasineura gleditchiae,

causing galls on leaﬂets ofGleditchia triacanthos, both originally Nearctic species that have been

imported into Europe unintentionally, probably with seedlings of their host plant. Ten species came

to Europe from North America, one species from Central and South America, six from Asia, ﬁve

from Africa and the origin of one species is unknown (Skuhravá et al. 2007, 2010).

Fig. 28. Distribution of Dasineura mali (Kieffer, 1904) (black circles) on apple trees Malus sylvestris Mill. and Malus

domestica Borkh. (Rosaceae) in Europe (black line). D. mali probably originates, as do its host plants, from southern

Europe and western Asia which are primary areas of its distribution, and it was imported secondarily to other parts of the

world. Distribution area of the host plant after Meusel et al. (1965–1992).

122

Five European gall midge species were intentionally introduced into other continents for bi-

ological control of weeds: Cystiphora sonchi against Sonchus arvensis in Canada (Peschken

1982); Cystiphora schmidti against skeleton weed (Chondrilla juncea) in Australia, California

and Argentina; Zeuxidiplosis giardi against St. John’s wort (Hypericum perforatum) in Hawaii,

New Zealand and Australia; Spurgia euphorbiae against leafy spurge (Euphorbia esula) in North

America (North Dakota) (Sobhian et al. 2000); Rhopalomyia tripleurospermi against scentless

chamomile (Tripleurospermum perforatum) in Canada (Skuhravá & Hinz 2000). On the other

hand, the European species Gephyraulus raphanistri which was tested as a potential control agent

of Raphanus raphanistrum, one of the most important weeds of crops in Australia, was found to

have unsuitable phenological characteristics (Vitou et al. 2008).

Endemic species form a special category. Such species are native to a particular geographical

region or their occurrence is restricted to such region. In Europe it is possible to class thirty eight

gall midge species as endemic. They are known so far only from the Iberian Peninsula. They

have not been recorded in any other country and seem to be restricted to the Iberian Peninsula. It

is necessary to emphasize that they may be revealed in some of other countries in the future. As

Fig. 29. Occurrence of Obolodiplosis robiniae (Haldeman, 1847) (black circles) causing galls on Robinia pseudoacacia

L. (Fabaceae) and its spread throughout Europe. O. robiniae is a Nearctic species, alien to Europe. It has been imported

into Europe, probably with seedlings, and its galls were discovered for the ﬁrst time in 2003 in northern Italy. Since then

it has spread quickly over large areas of Europe up to Doneck in Ukraine in 2006, to southern England in 2007, northern

Spain and southern Sweden in 2008.

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a true endemic may be regarded only such species which are associated with host plants endemic

to the Iberian Peninsula, as for example Dasineura halimii and D. herminii causing galls on

Halimium spp. (Skuhravá et al. 2006).

We have prepared maps showing the distribution areas of more than 500 gall midge species.

Some of them were published in our articles and the larger part await publication in the future.

In addition, we have drawn several hundreds of maps showing the occurrence of gall midge spe-

cies in several countries of Europe where we summarized previous knowledge, i.e. of the Czech

Republic (Skuhravá 1994a, b), Slovakia (Skuhravá 1991), France (Skuhravá et al. 2005), Spain

and Portugal (Skuhravá et al. 2006), Poland (Skuhravá et al. 2008) and Austria (Skuhravá &

Skuhravý 2009a).

Geographic position and gradients

Gall midge species richness changes with composition of natural vegetation and with geographic

position, mainly along a north-south axis (Skuhravá & Skuhravý 1998). We tested both geogra-

phic gradients, latitudinal and longitudinal, using the average species numbers per locality

obtained from samples taken by the time unit area collection method at localities with precisely

determined geographic positions. Fundamental data for the analysis are given in Table 10 and

results are shown in Figs 30 and 31.

The occurrence of gall midges given by average species numbers along the latitudinal gradient

produces interesting results (Fig. 30). The average species numbers of gall-inducing Cecidomyiidae

Fig. 30. Average species numbers of gall-inducing and plant inhabiting species of the subfamily Cecidomyiinae in coun-

tries of Europe along the latitudinal gradient of 3800 km from the northernmost investigated area in northern Norway

(Harstad) to the southernmost investigated area in the island of Malta in the Mediterranean Sea. All samples have been

taken by the time and area unit collection method.

124

Table 10. European countries where systematic faunal investigations have been undertaken by the present authors during the period 1955–2009, important characte-

ristics of countries and average species numbers of Cecidomyiidae. Geographic coordinates (latitude and longitude) are related as far as it is possible to the area or

town situated nearest to the study area. Altitudes include the lowest and the highest elevations of examined localities. Biogeographic provinces are given according

to Udvardy (1975), types of natural vegetation according to Noirfalise (1987). At the end of the table there are added data obtained in the course of investigations in

the southern part of Central Siberia (Asia) for comparison with data obtained in Europe. ASN – average species number

country lat. N long. E altitude biogeographic provinces type of natural vegetation ASN references

m a.s.l.

Austria, Wien 48°12’ 16°22’ 240 Central European Highlands mixed forest, conifer forests 17.0 Skuhravá & Skuhravý 2009a

115–2500

Bulgaria, Soﬁa 42°40’ 23°19’ 1–2000 Balkan Highlands mixed forest, oak forest 14.0 Skuhravá et al. 1991

Bulgaria, Varna 43°13’ 27°55’ 1–180 Black Sea Coast Pontic beech forest 14.0 Skuhravá et al. 1991

Corfu (Greece) 39°37’ 20°02’ 1–250 Mediterranean Sclerophyll Hellenic oakwood 12.6 Skuhravá & Skuhravý 2006b

Corsica (France) 41°55’ 08°44’ 1–1000 Mediterranean sclerophyll 9.8 Skuhravá & Skuhravý (in prep.)

Crete (Greece) 35°20’ 28°08’ 1–800 Mediterranean sclerophyll 9.2 Skuhravá & Skuhravý 1997a

Cyprus 35°00’ 33°00’ 1–150 Mediterranean sclerophyll 3.1 Skuhravá & Skuhravý 2004a

Czech Republic, Prague 50°05’ 14°25’ 120–1603 Central European Highlands mixed forest 26.0 Skuhravá 1994a, b

Denmark, Sjæland 55°66’ 12°34’ 1–75 Atlantic Province

(Copenhagen) Boreo-nemoral beech forest 22.0 Skuhravá et al. 2006

Faroe Islands, Tórshavn 62°00’ 6°46’W 0–300 Arctic-Alpine blanket bogs 1.0 Skuhravá & Skuhravý 2009

(Denmark)

France, Orleans 47°54’ 01°54’ 110–630 Atlantic Province oak forest 28.0 Skuhravá & Skuhravý 2004d

France, Hautes Alpes,

Grenoble 45°10’ 05°43’ 850–2645 Central European Highlands beech-ﬁr forest 25.0 Skuhravá & Skuhravý 2004c

Germany, Bayerischer 48°32’ 09°03’ 650–1450 Middle European Forest coniferous, mixed forest 13.0 Skuhravá & Skuhravý 1992b

Wald

Germany, Harz 51°41’ 10°58’ 450–550 Middle European Forest Hercynian beech forest 20.0 Skuhravá & Skuhravý 1988

Germany, Fichtelgebirge 50°07’ 12°00’ 528–875 Middle European Forest Hercynian beech forest 13.0 Skuhravá & Skuhravý 1992a

Greece, Athens 38°03’ 23°40’ 140 Balkan Highlands, oak forest, sclerophyll 9.3 Skuhravá & Skuhravý 1997a

1–1100 Mediterranean

Hungary, Budapest 47°30’ 19°04’ 116–1015 Pannonian Danubian thermophilous 29 Skuhravá & Skuhravý 1999

oak forest

Italy, Central part (Roma) 41°58’ 12°40’ 10–13 Mediterranean Quercus ilex forest 8.0 Skuhravá & Skuhravý (unpubl.)

Italy, Calabria, Cetraro 39°31’ 15°56’ 1–560 Mediterranean Quercus ilex forest 14.0 Skuhravá & Skuhravý (in prep.)

Italy, South Tyrol, Bolzano 46°20’ 11°14’ 224–2340 Central European Highlands spruce forest 15.0 Skuhravá & Skuhravý

2001–2009

Liechtenstein, Vaduz 47°07’ 09°31’ 476–620 Central European Highlands spruce forest 19.5 Skuhravá & Skuhravý 1993a

Mallorca (Spain) 39°33’ 02°44’ 1–400 Mediterranean sclerophyll 9.5 Skuhravá & Skuhravý 2004b

Malta, La Valetta 35°54’ 14°30’ 1–240 Mediterranean sclerophll 4.5 Skuhravá et al. 2002

Norway, Harstad 68°48’ 06°30’ 1–3 West European taiga blanket bogs 5 Skuhravá & Skuhravý (in prep.)

Norway, Trondheim 63°25’ 10°25’ 5–560 West European taiga boreal spruce forest 14.0 Skuhravá & Skuhravý (in prep.)

125

Samos (Greece) 37°41’ 27°00’ 1–600 Mediterranean sclerophyll 8.8 Skuhravá & Skuhravý 2006b

Sardinia (Italy) 40°55’ 09°00’ 0-700 Mediterranean sclerophyll 6.5 Skuhravá & Skuhravý 2002

Slovakia, Bratislava 48°08’ 17°07’ 110–2000 Middle European forest, Carpathian forest, Danubian 19.5 Skuhravá 1991

Pannonian thermophilous oak forest

Sweden, Kiruna 67°51’ 20°13’ 330 West European taiga subarctic pine-birch woodland 1.0 Skuhravá (unpubl.)

Sweden, Gävle 60°40’ 17°10’ 1–10 West European taiga subarctic pine-birch woodland 1.0 Skuhravá (unpubl.)

Switzerland, Zurich 47°23’ 08°33’ 208–2044 Central European Highlands montane Scotts pine forest, 16.0 Skuhravá & Skuhravý 1997b

beech forest

United Kingdom, 51°30’ 00°12’ 20–150 British Islands beech and oak forest 14.2 Skuhravá & Skuhravý 2007b

Ripley, Surrey (GB)

Asia: Siberia, Krasnojarsk 56°00’ 92°00’ 350– 1400 Central Asian taiga East Siberian taiga 13.5 Skuhravá & Skuhravý 1993b

in countries of Europe arranged along the latitudinal gradient

over a distance of about 3800 km from the northernmost

investigated area in Norway (Harstad) to the southernmost

investigated area in the island of Malta shows the highest

value between latitudes 45° and 52° North. Species num-

bers in the northernmost localities were low, mainly in tundral

parts of Scandinavia. In contrast, gall midge species numbers

found at the coast in northern Norway were relatively high

owing to the inﬂuence of the warm Gulf Stream. Average

species numbers decrease rapidly to the south and in Malta

the average species number per locality was only 5.

On the other hand, the occurrence of gall midge species

along the longitudinal gradient does not produce such

clear results (Fig. 31). The average species numbers of

gall-inducing Cecidomyiidae in countries of Europe along

the longitudinal gradient of 2000 km from the westernmost

investigated area in England to the easternmost area in

Bulgaria at the Black Sea coast in comparison with average

species number in the southern part of Central Siberia seems

to be nearly equal, with two higher values (peaks), viz. in

France, where average species number was 29 owing to the

inﬂuence of theAtlantic Ocean, and in Central Europe where

the highest average species number was found in Hungary

situated in the warmer part of the Pannonian lowland.

Climatic factors

Climate and climatic factors have an important inﬂuence

on the occurrence and distribution of gall midges in Euro-

pe. Climate includes temperature, humidity, atmospheric

pressure, wind, rainfall and some other elements in a given

region over long periods of time. The climate of a locality is

affected by its latitude, terrain, altitude, ice or snow cover,

as well as nearby water bodies and their currents. Climates

can be classiﬁed accordingto the average and typical ranges

of different variables, most commonly temperature and pre-

cipitation which may be shown in graphs, so called clima-

tograms. They represent the sum of two climatic variables,

e.g. temperature and rainfall for a particular geographical

locality and for a speciﬁed period, with measurements taken

at regular intervals, usually monthly. In the climatogram

are given the long-term monthly average temperatures (in

a graph on the left side) and the monthly precipitations (in

a graph on the right side) in relation 10° C = 20 mm. When

the precipitation curve falls below the temperature curve, it

is a dry period.

The average species number of gall midges is highest

in Central Europe and decreases towards the south, as was

described in the previous chapters. We compared decreasing

126

average gall midge species numbers per locality with climatograms of several parts of Europe and

found that this phenomenon is correlated with climatic conditions and with the temperature and

rainfall (Fig. 32). Central Europe has favourable weather during the year which is characterized

with mild, i.e. not too high temperatures and with a relative sufﬁcient amount of rainfall. The

average gall midge species number in Central Europe is very high, i.e. 26 species in the Czech

Republic and 29 species in Hungary.

The average temperature increses to the south of Europe and, on the other hand, the average

rainfall decreases, which is shown on climatograms for southern Europe where the middle part

of a year with hot and dry weather is ﬁgured in black. Hot and dry weather is not too suitable for

the development of gall midges and their galls on host plants. It may be shown that low average

gall midge species numbers occur in the islands of the Mediterranean, viz. only 9.5 species on

average in Mallorca, 9.2 in Crete and even only 4.5 species in Malta.

Long winter periods of cold with low temperatures and short summer periods in northern Europe

do not give suitable conditions for the development of gall midges and their galls on host plants.

On the other hand, a narrow strip (several kilometres only) along the coast in northern Europe

which is under the inﬂuence of the Gulf Stream provides suitable conditions for development of

gall midges on host plants and the gall midge species number is there much higher than in inland

parts at the same latitude.

Vertical distribution

The composition of vegetation, including availability of various plant species as potential hosts

for gall-inducing gall midge species, changes with increasing altitude. It is an important value for

zoogeographic analysis. Europe is very varied; the conﬁguration of terrain changes very quickly

Fig. 31. Average species numbers of gall-inducing and plant inhabiting species of the subfamily Cecidomyiinae in coun-

tries of Europe along the longitudinal gradient of 2000 km from the westernmost investigated area in southern England

to the eastermost area in Bulgaria at the Black Sea compared with average species number in the southern part of Central

Siberia. All samples have been taken by the time and area unit collection method. For abbreviations of country names

see Table 1.

127

in some parts and large differences in altitudes occur in relatively small areas. We analyzed the

vertical distribution of the gall midge species from localities in several parts of Europe accor-

ding to the altitudinal zones proposed by Ellenberg (1978) that we adapted for different areas. In

Central Europe we usually use eight zones for the area under study, viz. planare zone (from the

sea level–200 m a. s. l.), colline (200–500 m a. s. l.), submountain (500–900 m a. s. l.), mountain

(900–1500 m a. s. l.), sub-Alpine (1500–1700 m a. s. l.), Alpine (1700–2400 m a. s. l.), subnivale

and nivale zones (over 2400 m a. s. l.). Each of these zones is indicated by different abiotic factors,

mainly climatic, and by characteristic composition of vegetation in which there may be present

plant species suitable as hosts for gall-inducing Cecidomyiidae. Altitudinal range of these zones

differ in slopes of mountains or mountain ranges oriented south and north, and with geographic

position.

Elevations of localities at which particular species have been found are an essential basis for

zoogeographic evaluation. For example, in Austria the lowest point is at Neusiedler See at 115 m

a. s. l. and the highest point is the mount Grossglockner at 3797 m a. s. l. We collected galls of

Fig. 32. Europe and its climatic conditions shown in several climatograms. Mild weather conditions in Central Europe are

suitable for development of gall midges. In southern Europe, on islands in the Mediterranean Sea and in northern Africa

dry and hot climates prevail which are not suitable for development of gall midges and for their galls on host plants.

128

gall midges at localities situated in planare zone at 120 m up to the Alpine zone at 2400 to 2500

m. Only a few gall midge species and their associated host plants occur in narrow altitudinal

spans of one zone. Most species occupy larger areas of distribution both in horizontal and vertical

directions and usually are able to occur with their host plants over a relatively large altitudinal

range (Skuhravá & Skuhravý 2009a).

Gall midge species numbers in the planare zone of Europe are usually low and numbers are

usually much higher in colline, submontane and montane zones. Above the boundary of forests,

the so-called timber line (this altitude differs in various parts of Europe in connection with geo-

graphic position), the gall midge species number falls abruptly and only two to four species occur

in the higher zones, i.e. in the sub-Alpine and Alpine zones. The number of gall midge species

therefore decreases with rising altitude.

Of 396 gall midge species forming the fauna of Austria (Skuhravá & Skuhravý 2009a), the

majority occur in colline, submontane and montane zones, only 20 species occur in the sub-Al-

pine zone and 13 species in the Alpine zone. Jaapiella alpina causing galls on Silene acaulis,

Dasineura bistortae on Polygonum viviparum and Hygrodiplosis vaccinii on Vaccinium uligi-

Fig. 33. Vertical distribution of gall midges (Cecidomyiidae) shown as average species numbers in belts of 100 meters in

altitudinal zones of Austria in Central Europe. The average species number falls with rising elevation from the planare

zone up to the subnivale zone where only one gall midge species, Jaapiella alpina causing galls on Silene acaulis (Cary-

ophyllaceae) has been found at altitudes of 2500 m a. s. l. Span of altitudinal zones according to Ellenberg (1978).

129

nosum, found at elevations from 2300–2500 m a. s. l., are true Alpine species associated with

Alpine host plants. In South Tyrol, where intensive investigations were carried out in the Alpine

zone (Skuhravá & Skuhravý, in press) 33 species were found in the Alpine zone. The galls of

Rhopalomyia luetkemuelleri on Artemisia spicata and Jaapiella alpina on Silene acaulis were

found in the subnivale zone at altitudes of 2700 m a. s. l. and are gall midge species found at the

highest altitudes in Europe.

The vertical distribution of gall midges in Europe is shown by average species numbers

recorded at localities in separate successive altitudinal belts of 100 meters (Fig. 33). The basis of

calculations are totals of all species numbers obtained during our own investigations at localities

situated in particular altitudinal zones.

We recognized twenty three types of vertical distribution in the gall midge fauna of Austria

on the basis of their altitudinal span which is given by elevations of the lowest and the highest

situated locality where a particular species was found (Fig. 34). Two fundamental patterns of

vertical distribution were found in each of the lower zones, viz. the occurrence of a species only

inside a particular zone and the occurrence of a species exceeding this zone and reaching higher

zones or running across them to higher altitudinal zones. Few gall midge species and their asso-

ciated host plants occur in narrow altitudinal spans of one zone. These may be named monozonal

species. Most species occupy larger areas of distribution both in horizontal and vertical directions

and usually are able to occur with their host plants in relative large altitudinal range. They may

be named bi-, tri-, or polyzonal species.

Fig. 34. Diagram of twenty-three types of vertical distribution in seven altitudinal zones of Europe recognized in the gall

midge fauna in the course of investigations carried out by present authors at 1821 localities in the years 1955–2009.

130

For example, the gall midges associated with Quercus cerris in Central Europe may be conside-

red to be monozonal and colline species, Macrodiplosis pustularis causing galls on Quercus robur

and Q. petraea is a bizonal species occurring in colline and submontane zones and Oligotrophus

juniperinus inducing galls on Juniperus communis a polyzonal species. It occurs over a large

altitudinal span, including sites in the planare zone up to sites in the Alpine zone. Bisamberg,

190 m a. s. l. in north-eastern Austria was the lowest situated locality where galls were found

and Mount Kanzel, 2348 m a. s. l. in the Alps in northern Italy is the highest point of its known

vertical occurrence (Skuhravá 1987, 1997, Skuhravá & Franz 1989, Skuhravá et al. 2002). O.

juniperinus has a large ecological tolerance and is able to inhabit places that are distant vertically

from each other by more than 2000 meters. O. juniperinus may be considered to be a sub-boreal

and Alpine species.

The occurrence of gall midges in single altitudinal zones may change with geographical position

in Europe. For example, Asphondylia verbasci causing ﬂower gallson various Verbascum species

is in the Czech Republic a colline species occurring from 198 m a. s. l. to 510 m a. s. l., but in

southern Europe, in Bulgaria, it occurs at altitudes of 1400 m a. s. l. in the montane zone.

We prepared detailed analyses of the vertical distribution of gall midge species occurring in

the Czech Republic, Slovakia and Austria and expressed them in graphs (Skuhravá 1991, 1994a,

b, Skuhravá & Skuhravý 2009a).

Frequency

Frequency in ecology is the number of individuals of a species within a given area. In our case,

the frequency is the number of occurrences (given as the number of localities where the species

was found) of a particular gall midge species within a given area. The area under study may be of

varied size from small, including e.g. district, county, region, mount, mountains, island, protected

part of landscape, up to large, e.g. country, Western Europe, Central Europe, zoogeographic or

biogeographic subregion and region, continent. The basis for such analysis is the number of loca-

lities at which a particular species was found, without determining the local abundance of species.

The frequency of animals is usually expressed only by words: very scarce, scarce, moderately

frequent, frequent and very frequent.

For example, if a particular species was found in the area under study at only one locality out

of 60 localities that had been investigated, it is designated as very rare and a species found at 30

localities is designated as very frequent. Species between these two value are denominated by the

terms: scarce, moderately frequent or frequent based only on the experience of the researcher.

On the basis of our rich experiences with the evaluation of frequency of gall midge species

in areas of various size in Europe we developed a uniﬁed method for evaluation of frequency of

a particular species in a given area where a particular number of localities was investigated.

If we plotted the occurrence of all species found in the area at localities under study, we received

always the same conﬁguration in the graph, as in Fig. 35. The highest column on the left inclu-

des species each of which was found at only one locality in the course of our investigations, the

number of localities per species reduces very quickly, only a few species occur at more localities

and usually one or two species occur very frequently, each being found at many localities (small

column on the right) (Skuhravá & Skuhravý 2009a). It is possible to divide all species found in

the area under study into six frequency groups by using the ﬁrst six members of the geometrical

progression with quotient “2” and coefﬁcient “a”:

a.20 + a.21 + a.22 + a.23 + a.24 + a.25 = 63.a (a sum of the progression).

1a + 2a + 4a + 8a + 16a + 32 a = 63 a (simpliﬁed)

131

The value of the coefﬁcient may be determinated froma sum of the geometrical progression. Ifwe

investigate an area at 63 localities, the coefﬁcient “a” has the value 1. In other cases it is possible

to count the coefﬁcient “a” from the number of localities that we investigated which is equal to

63a. The extent of each of six frequency group is given by one member of this progression. De-

tails are given in the article of Skuhravá (1994a, b). It is possible to assign verbal denominations

to each of the six group. As an example we give here the calculation of the frequency groups for

Austria in Table 11.

The frequency of gall midge species in Europe is noted for the following shared characters

(Fig. 35). On one side is the group of many very scarcely occurring species that were found only

once in the area under study, some of them were found only at the type locality and have not been

Table 11. Frequency groups of gall midges found in Austria (after Skuhravá & Skuhravý 2009a)

frequency group 1 2 3 4 5 6

members of geometrical progression a.20 a.21 a.22 a.23 a.24 a.25

simpliﬁed 1a 2a 4a 8a 16a 32a

number of localities (if a=300) 1–5 6–14 15–33 34–71 72–147 148≤

verbal denomination very scarce medium frequent very most

scarce frequent frequent frequent

Fig. 35. Species of the family Cecidomyiidae found in Austria in the period 1834–2004 arranged in six frequency groups

according to the number of localities at which each species was found. The ﬁrst column on the left includes 145 gall

midge species which have been found at only one locality. The last column on the right (in the form of a black point)

shows one species, Macrolabis heraclei (Kaltenbach, 1862), causing galls on Heracleum sphondylium (Apiaceae), the

most frequent species in Austria, which was found at 159 localities.

132

found again since that time. They are ﬁgured in the graph on the left side. On the opposite side is

the group of a small number of species, sometimes only one or two species, that were found many

times (= at many localities) and are the most frequent species in the area under study. In the graph

they are ﬁgured at the right side. Each of four groups between these two extreme groups includes

decreasing numbers of species each of which was found at more and more localities.

We have determined the frequency of gall midges occurring in Europe in some of the countries

where we have carried out investigations, viz. in Slovakia (Skuhravá 1991), Bulgaria (Skuhravá

et al. 1991), the Czech Republic (Skuhravá 1994a, b), Switzerland (1997), Hungary (Skuhravá &

Skuhravý 1999), Denmark (Skuhravá et al. 2006) and Austria (Skuhravá & Skuhravý 2009a). If

there are not such data at our disposal, it is possible to use the number of citations in the literature

to evaluate the frequency of occurrence, as we have done for Portugal and Spain (Skuhravá et al.

2006) and for Poland (Skuhravá et al. 2008), or to combine both methods, as we have done for

France (Skuhravá et al. 2005).

Each part of Europe is characterized by the occurrence of the different groupings of the most

frequent gall midge species. The most frequent gall midge species in the whole of Europe is

Jaapiella veronicae causing galls on leaf buds of Veronica chamaedrys. The galls of this species

were found at 501 localities (of 670 explored) in the Czech Republic (Central Europe) during

investigations in the years 1957–1982 (Fig. 36). In Northern Europe, in Norway, the most fre-

quent species is Iteomyia capreae causing galls on leaves of Salix caprea and related species, in

Denmark Mikiola fagi inducing galls on leaves of Fagus sylvatica and in Poland Lasioptera rubi

causing galls on stems of Rubus idaeus and related species. In France the most frequent species

is Dasineura rosae causing galls on various species of Rosa, in Switzerland Dasineura fraxinea

producing pustule galls on leaves of Fraxinus excelsior, in Austria Macrolabis heraclei causing

galls on leaves of Heracleum sphondylium, in Slovakia Geocrypta galii evoking galls on stems of

Galium mollugo and related species, in Hungary Lasioptera rubi and Dasineura rosae, in Bulgaria

also D. rosae and D. hyperici which causes galls on Hypericum perforatum. All above mentioned

gall midge species belong to faunal elements of the Middle European Forest Province (Fig. 37).

The most frequent gall midge species in the western part of the Mediterranean area – in the

Iberian Peninsula, that is in Spain and Portugal – are Phyllodiplosis cocciferae and Dryomyia

lichtensteinii causing galls on leaves and buds of south-European oaks, viz. Quercus coccifera,

Q. ilex and Q. suber, whereas in the eastern part of this area – in Greece – the most frequent spe-

cies is Dasineura turionum changing stems of Asparagus acutifolius into galls. These gall midges

belong to faunal elements of the Mediterranean Sclerophyll Province.

Possibilities of ﬁnding additional gall midge species in Europe

Gall-inducing gall midges of Europe are one of the best known gall-causing group of animals in

the world. They can be used therefore as the most suitable group for estimating the number of gall

midges and their galls that may be found in future studies. We try to determine this prognosis using

data from the literature and also data obtained in the course of our investigations in Europe.

We use as the starting point for our prognosis of the increasing species number of the family

Cecidomyiidae in Europe ﬁve gall midge species given in Linnaeus (1758) whose 10th edition of

Systema Naturae is the starting point of zoological nomenclature. Bergenstamm & Löw (1876)

summarized knowledge of that time and gave 606 species of the family Cecidomyiidae from all

the world (Table 12). For Europe they gave 492 species. In the subsequent period of 120 years

the number of known gall midges was increased by about ﬁve hundred species. Of those, 45 spe-

cies belonged to the subfamily Lestremiinae, 357 to the subfamily Cecidomyiinae and 90 to

undescribed species causing galls. The two last groups form together a group of 447 gall causing

and plant inhabiting species.

133

Kieffer (1898) gave 650 species in his list of the family Cecidomyiidae of Europe and Algeria

where Lestremiinae included 88 species and Cecidomyiinae 562 species, of which 492 species cau-

sed galls or inhabited plants without making galls. From 1876 to 1898 the number of Lestremiinae

increased importantly from 45 to 88 species, i.e. by about 100%, on the other hand the number of

gall-causing species was increased only moderately from 447 to 492, i.e. by about 10%.

Fig. 36. Occurrence of Jaapiella veronicae (Vallot, 1827), causing galls on Veronica chamaedrys L. (Scrophulariaceae),

the most frequent species in Central Europe (black circles). Its galls were found at 510 localities in the Czech Republic.

The density of its occurrence decreases southwards and this species does not occur in the Mediterranean area. White circles

indicate absence of the species. Distribution area of the host plant (black line) after Meusel et al. (1965–1992).

134

Kieffer (1901) summarized knowledge of galls in Europe and gave in his synopsis 1067 spe-

cies of animals belonging to various groups as causers of galls. The family Cecidomyiidae with

364 species was the most species-rich group of gall-inducing animals.

Houard (1908–1909) recorded from Europe and the Mediterranean area 6239 sorts of galls

caused by 1450 species belonging to various groups of animals of which 420 were gall-causing

species of the family Cecidomyiidae (i.e. valid, described species). In addition, he listed about

300 galls, the causers of which were unknown, on various host plants.

Buhr (1964–1965) included 580 valid species (65%) of the family Cecidomyiidae as causers

of galls on various host plants in Central and Northern Europe and about 320 undescribed species

(35%) the causers of which had not been identiﬁed because larvae were not known and adults had

not been reared at that time. The relation between described species and undescribed species is

2:1. It is possible to assume that these unknown causers will be found, reared from galls, identiﬁed

and described in the future. The gall midge fauna of Europe will thus be enriched in the future by

about 300 species known at the time of Buhr and by another 100 undescribed species associated

with various plant species discovered mainly in the southern parts of Europe from the second

half of 20th century until the present. It is therefore possible to expect about a 50% increase in

the number of valid gall midge species and their galls in the future.

Skuhravá (1986) recorded in the Catalogue of Palaearctic Diptera 1500 species in the family

Cecidomyiidae in Europe of which 170 belong to the subfamily Lestremiinae, 215 to the subfamily

Porricondylinae and 1115 to the subfamily Cecidomyiinae. After 25 years, the number has increased

to 1800 species (Skuhravá 2010, unpublished). The subfamily Lestremiinae includes 280 species,

the subfamily Porricondylinae 310 and the subfamily Cecidomyiinae 1210 species.

In Fig. 38 we show the increasing species numbers of gall midges and increasing number of

galls caused by gall midges during the last 150 years and the lines show the trends of increase.

On the basis of summarized facts we predict that in Europe between the years 2010 and 2050 the

number of gall midges species recorded will increase by about four hundred to 2200 valid species

and the number of galls caused by gall midges by about two hundred to 1400 sorts of galls caused

by gall midges. This is based on the assumption that in the future most of the unidentiﬁed causers

of galls will be recognized as good species and described and that new galls and new gall midge

species will be found by future researchers.

Gall midges and host plants

The vegetation covering Europe is relatively species-rich. Tutin et al. (1964–1980) give the

following numbers of plant taxa: 11,557 species, 1541 genera and 203 families. 915 gall midges

species (i.e. valid, described species) forming the subfamily Cecidomyiinae are associated with

280 host plant genera belonging to 72 plant families, i.e. with 18% of plant genera and 35% of plant

families known to occur in Europe. Undescribed gall midge species and species identiﬁed to the

Table 12. Number of species of the family Cecidomyiidae in various parts of the world, as given by Bergenstamm &

Löw (1876)

Cecidomyiidae number of species total

Europe N America S America Africa Orient Australia

Lestremiinae 45 1 1 0 0 1 48

Cecidomyiinae 357 42 4 6 4 2 415

unidentiﬁed species 90 45 0 7 1 0 143

total 492 88 5 13 5 3 606

135

genus level that are known on the morphology of the gall only are not included in our consideration.

On average one plant genus hosts 3,26 species of gall midges. We analyse relations between gall

midges and their host plant from several points of view, viz. from the phylogenetical system of

plants, abundance of gall midges associated with host plant families and relations between gall

midges and their host plants in different countries of Europe, and the distribution and abundance

of galls on different organs of host plants. See tables 13, 14 and 15.

In Europe only two gall midge species are associated with the Pteridophyta and 27 species

with the Gymnospermae. The majority of gall midges, 886 species, are associated with the Angi-

Fig. 37. The most frequent species of the family Cecidomyiidae in several countries of Europe. See comments in the

text p. 132.

136

Table 13. Gall-inducing and plant inhabiting species of the subfamily Cecidomyiinae associated with plant families and

plant genera in Europe, arranged according to the phylogenetical system of plants according to Tutin et al. (1964–1980)

in Flora Europaea; n1 – number of plant genera; n2 – number of gall midge species

host plants taxa n1 n2 host plants taxa n1 n2

Pteridophyta Dennstaedtiaceae 1 2 Rhamnales Rhamnaceae 2 5

Spermatophyta Vitaceae 1 3

Gymnospermae Malvales Tiliaceae 1 8

Coniferales Hypericales Hypericaceae 1 6

Pinaceae 4 17 Thymelaeales Thymelaeaceae 1 2

Cupressaceae 2 8 Violales Violaceae 1 5

Taxales Taxaceae 1 1 Cistaceae 2 4

Gnetales Ephedraceae 1 1 Tamaricaceae 1 5

Angiospermae Cucurbitales Cucurbitaceae 1 2

Dicotylenodes Myrtales Lythraceae 1 1

Salicales Salicaceae 2 64 Onagraceae 1 2

Fagales Betulaceae 2 14 Umbelliﬂorae Araliaceae 1 1

Corylaceae 2 10 Cornaceae 1 1

Fagaceae 2 49 Apiaceae 18 28

Urticales Ulmaceae 1 4 Ericales Pyrolaceae 1 1

Moraceae 1 1 Ericaceae 3 15

Cannabaceae 1 1 Primulales Primulaceae 1 1

Urticaceae 1 3 Oleales Oleaceae 4 1

Santalales Loganiaceae 1 1 Gentianales Gentianaceae 1 1

Polygonales Polygonaceae 2 10 Apocyanaceae 1 1

Centrospermae Chenopodiaceae 7 21 Asclepiadaceae 1 2

Caryophyllaceae 9 23 Rubiaceae 2 14

Ranales Ranunculaceae 7 13 Tubiﬂorae Boraginaceae 4 8

Berberidaceae 1 2 Lamiaceae 20 42

Rhoeadales Papaveraceae 1 1 Solanaceae 1 3

Capparidaceae 1 1 Scrophulariaceae 7 16

Brassicaceae 20 28 Plantaginales Plantaginaceae 1 1

Rosales Saxifragaceae 2 1 Dipsacales Caprifoliaceae 3 11

Grossulariaceae 1 5 Valerianaceae 1 2

Rosaceae 11 49 Dipsaceae 3 4

Fabaceae 28 123 Campanulales Campanulaceae 2 11

Geraniales Geraniaceae 1 2 Asteraceae 25 104

Linaceae 1 1 Monocotyledones

Euphorbiaceae 1 9 Liliiﬂorae Liliaceae 7 10

Rutales Rutaceae 2 2 Dioscoreaceae 1 1

Polygalaceae 2 1 Juncales Juncaceae 1 1

Sapindales Anacardiaceae 2 3 Graminales Poaceae 31 69

Aceraceae 1 9 Cyperales Cyperaceae 2 27

Celastrales Buxaceae 1 1 Microspermae Orchideaceae 1 1

total 72 plant families, 280 host plant genera, 915 gall midge species

ospermae, of which a smaller part, including 109 species, cause galls or develop in plant organs

of host plants belonging to the Monocotyledones and a larger part, including 777 species, develop

on Dicotyledones.

The Pteridophyta in Europe comprise 25 families of which only one family, Dennstaedtiaceae

(Polypodiaceae), contains only one species – Pteridium aquilinum – that is the host plant of two

gall midge species, Dasineura pteridis and D. pteridicola. Larvae of both species cause simple

galls on leaf margins (bent or rolled parts of leaf margins). These two gall midge species form

only 0,21% of all gall midge species associated with host plants in Europe.

137

The Gymnospermae in Europe comprise ﬁve families ofwhich four include host plant species

for 27 gall midge species, i.e.2,95% of all gall midge species associated with host plants in Europe.

The family Pinaceae hosts 17 gall midge species which are associated with four plant genera of

coniferous trees (Abies, Larix, Picea and Pinus). The family Cupressaceae hosts eight gall midge

species associated with two plant genera, viz. Juniperus, hosting seven gall midge species, and

Chamaecyparis, hosting only one gall midge species, which is of North American origin. The

family Taxaceae hosts only one gall midge species, Taxomyia taxi, developing in galls on Taxus

baccata and the family Ephedraceae includes the host plant for only one species, Xerephedromyia

ustjurtensis, causing galls on Ephedra distachya.

About one half of gall midge species developing on host plants of Gymnospermae do not cause

galls at all: two species develop inside seed, ﬁve species in cones and ﬁve species are associated

with resin. Fifteen gall midge species developing in various plant organs cause galls: two species

induce stem galls, four species cause galls on needles and nine species induce galls in buds. The

galls on Gymnospermae are of various shape. The most primitive seem to be simple swellings

on the bark of Picea abies caused by larvae of Dasineura abietiperda. More complicated are the

galls of Xerephedromyia ustjurtensis caused on stems of Ephedra distachya, which are formed

by swellings of stems with large central chambers containing the larva of the causer. The larvae

of Paradiplosis abietispectinatae induce swellings of needles on Abies alba and the larvae of

Thecodiplosis brachyntera cause swellings at the base of needle pairs on Pinus sylvestris and P.

mugo. Seven species of gall midges cause bud galls on various species of Juniperus, the larvae of

Dasineura kellneri induce galls on buds of Larix decidua and the larvae of Taxomyia taxi change

leaf buds of Taxus baccata into galls.

Fig. 38. Increasing species number of the family Cecidomyiidae during the 19th and 20th centuries and the prognosis

for the future.

138

Of the ﬂowering plants, the Angiospermae are represented by 173 families in Europe, 68 of

which provide host plants for 886 gall midge species, i.e. 96.84% of all gall midge species asso-

ciated with host plants in Europe.

Of the Monocotyledonae, that contain 34 families, only six provide host plants for 109 gall

midge species, i.e. 11.92% of all gall midge species associated with host plants in Europe. Most

of them, totalling 69 species, are associated with various species and genera of Poaceae, 27 gall

midge species with Cyperaceae and 10 gall midge species with Liliaceae. Each of three families,

viz. Dioscoreaceae, Juncaceae and Orchideaceae, hosts only one gall midge species. Most gall

Table 14. Species numbers of gall-inducing and plant-inhabiting gall midges of the subfamily Cecidomyiinae associated

with host plant families in Europe and comparison with occurrence of gall midge species numbers of the same fami-

lies in Kazakhstan (Central Asia). Abbreviations: WE – Western Europe; EE – Eastern Europe; MD – Mediterranean;

KZ – Kazakhstan

plant family number of gall midge species plant family number of gall midge species

WE EE MD KZ WE EE MD KZ

Fabaceae 123 60 22 114 Vitaceae 3 0 2 0

Asteraceae 104 33 8 139 Urticaceae 3 1 2 1

Poaceae 69 14 6 16 Solanaceae 3 1 1 3

Salicaceae 64 26 3 18 Anacardiaceae 3 0 1 0

Fagaceae 49 9 18 2 Asclepiadaceae 2 0 0 0

Rosaceae 49 30 16 39 Berberidaceae 2 0 0 2

Lamiaceae 41 15 5 25 Cucurbitaceae 2 0 1 1

Apiaceae 28 7 6 33 Dennstaedtiaceae 2 0 1 0

Brassicaceae 28 8 5 44 Geraniaceae 2 0 1 6

Cyperaceae 27 3 1 1 Onagracceae 2 2 0 1

Caryophyllaceae 23 4 2 15 Rutaceae 2 0 1 0

Chenopodiaceae 21 2 8 90 Thymelaeaceae 2 1 0 0

Pinaceae 17 7 1 2 Valerianaceae 2 0 0 0

Scrophulariaceae 16 9 2 11 Apocyanaceae 1 0 0 0

Ericaceae 15 3 4 0 Araliaceae 1 0 0 0

Oleaceae 15 3 9 0 Buddlejaceae 1 0 0 0

Betulaceae 14 5 0 2 Buxaceae 1 1 0 0

Rubiaceae 14 4 4 14 Cannabaceae 1 0 0 0

Ranunculaceae 13 5 1 14 Capparidaceae 1 0 1 1

Caprifoliaceae 11 3 1 7 Cornaceae 1 0 0 0

Campanulaceae 11 1 0 2 Dioscoreaceae 1 0 0 0

Corylaceae 10 0 2 0 Ephedraceae 1 0 0 9

Polygonaceae 10 1 2 28 Gentianaceae 1 0 0 1

Liliaceae 10 0 2 1 Juncaceae 1 0 0 0

Aceraceae 9 4 3 0 Linaceae 1 0 0 2

Euphorbiaceae 9 3 2 14 Lythraceae 1 0 0 0

Tiliaceae 8 5 0 0 Loganiaceae 1 0 0 0

Cupressaceae 8 2 1 6 Moraceae 1 0 0 0

Boraginaceae 8 1 1 4 Orchideaceae 1 0 0 0

Hypericaceae 6 1 3 0 Papaveraceae 1 0 1 2

Grossulariaceae 5 3 0 3 Plantaginaceae 1 1 0 1

Rhamnaceae 5 1 1 1 Polygalaceae 1 2 0 1

Tamaricaceae 5 0 1 12 Primulaceae 1 1 0 1

Violaceae 5 1 1 1 Pyrolaceae 1 0 0 0

Cistaceae 4 0 0 1 Saxifragaceae 1 0 0 0

Dipsacaceae 4 2 0 0 Taxaceae 1 0 0 0

Ulmaceae 4 2 0 0 total families 73 41 40 54

139

midges totalling 777 species, i.e. 84.92% of all gall midge species associated with host plants in

Europe, are associated with 62 families of the Dicotyledones that contains a total of 139 plant

families.

Gall midges are more diverse on some plant families in which they ﬁnd more favourable host

plant species and genera for their development. Of the Dicotyledones, the family Fabaceae hosts

the highest number of gall midges – 123 species, followed by Asteraceae with 104 gall midge

species, Salicaceae with 64 species, Fagaceae and Rosaceae, each with 49 species, and Lamiaceae

with 41 species. About two thirds of gall midge species are associated with 12 large plant families

and the remaining third of gall midge species with 61 plant families.

The family Fabaceae hosts 123 gall midge species on plant species belonging to 28 genera,

for example Lathyrus hosts 17 gall midge species, Vicia 14 species, Medicago 11 species, Ge-

nista 10 species, and Trifolium 8 species. The family Asteraceae hosts 104 gall midge species on

25 genera of host plants, for example Artemisia hosts 24 gall midge species, Achillea 7 species,

Hieracium 7 species, Chrysanthemum 6 species, Erigeron 5 species. The family Poaceae hosts 69

gall midge species on 31 genera of host plants, for example Poa 9 gall midge species, Dactylis and

Table 15. Relations of gall-inducing and plant-inhabiting gall midges of the subfamily Cecidomyiinae to host plant spe-

cies in countries of Europe given as the index of mutual gall midge species and plant species occurrence (IGS-PSO); ng

– number of gall midge species; np – number of plant species; index – index of mutual occurrence

country ng np index references to botanical articles

Austria 352 3400 9 Tutin et al. 1964–1980

Belgium 130 1400 11 Tutin et al. 1964–1980

Bulgaria 240 3560 15 Jordanov 1976

Corsica 60 1625 27 Rikli 1946

Crete 38 2200 58 Rikli 1946, Sﬁkas 1987

Cyprus 32 1800 56 Sinclair 1991

Czech Republic 564 2200 4 Kubát 2002

Faroe Islands 5 400 80 Fosaa 2000

France 581 4500 8 Tutin et al. 1964–1980, Rikli 1946

Germany 653 2800 4 Jäger & Werner 2007

Greece 211 6700 32 Rikli 1946

Hungary 332 4200 13 Soó 1973

Italy 315 5600 18 Pignatti 1982

Italy (South Tyrol) 312 2361 7 Wilhalm & Hilpold 2006

Latvia 264 1385 5 Flora of Latvia 1953–1959

Mallorca 33 1250 37 Tutin et al. 1964–1980

Malta 36 1250 35 Haslam et al. 1977

Netherlands 305 1500 5 Tutin et al. 1964–1980

Poland 397 2500 6 Szafer et al. 1986, Mirek et al. 2002

Portugal 122 2735 22 Rikli 1946, Castroviejo 1986–2005

Romania 304 3600 12 Hayek 1927–1933, Savulescu 1952

Russia (European) 400 6500 16 Tutin et al. 1964–1980

Sardinia 44 2000 45 Tutin et al. 1964–1980

Sicily 89 2200 25 Tutin et al. 1964–1980

Slovakia 355 3352 10 Baláž et al. 2001

Slovenia 219 3200 15 Martinčič & Sušnik 1999

Spain 212 6500 30 Castroviejo 1986–2005

Sweden 266 2000 8 Karlsson 1998

Switzerland 228 3500 15 Lauber & Wagner 2001

Ukraine 166 3700 22 Bordzilowskij 1938

United Kingdom 526 2200 4 Fitter & Peat 1994

140

Festuca each 5 species, Alopecurus, Bromus, Calamagrostis and Phragmites each 4 gall midge

species. In contrast, the family Salicaceae hosts 64 gall midge species which are associated with

only two plant genera: Populus with 14 gall midge species and Salix with 50 gall midge species.

Also in the family Fagaceae only two plant genera are associated with gall midges: Fagus hosts

6 gall midge species and Quercus 43 gall midge species.

Most gall midges that are associated with Angiospermae induce galls on various plant organs

and only a few develop on or in organs of host plants without inducing gall development. Larvae

of plant-inhabiting, i.e. not gall-inducing gall midges, live under the leaf sheaths of grasses, inside

plant stems, under the bark of living, injured or dying trees, in resin masses, in inﬂorescences, in

or on seeds and in cones. About one half of gall-inducing gall midge species cause galls on ﬂowers

and leaf buds, others induce galls on leaves and only a small part develop inside fruits.

In Western Europe 915 gall midge species are associated with 72 plant families and 12.5 is the

average number of gall midge species per plant family; in Eastern Europe 287 gall midge species

are associated with 41 plant families and 7 is the average number of gall midge species per plant

family; in the Mediterranean 153 gall midge species are associated with 40 plant families and

3.8 is the average number of gall midge species per plant family. As it is shown, the western part of

Europe is more species rich than the eastern part and the Mediterranean is notably species poorer.

Surprisingly, the gall midge fauna of Kazakhstan in Central Asia is more species rich including

714 gall midge species that are associated with 52 families (according to Fedotova 2000), and

13.5 is the average gall midge species number per family.

In Western Europe 115 gall midge species are associated with Fabaceae and 100 species with

Asteraceae. Host plants of the family Fabaceae, with 60 gall midge species, are also in ﬁrst place

in Eastern Europe and in the Mediterranean. On the other hand, in Kazakhstan the highest species

number of gall-inducing gall midges – 139 species – is associated with the family Asteraceae

the species of which form the main plant associations in desert and semidesert areas of Central

Asia. In Western Europe a large amount – 69 species – are associated with Poaceae, 64 species

with Salicaceae and 49 species with Fagaceae. Gall midge species associated with plant families

including trees and shrubs (Salicaceae, Fagaceae, Pinaceae, Oleaceae, Betulaceae, Corylaceae,

Aceraceae, Tiliaceae, Cupressaceae, Rhamnaceae, Tamaricaceae and Ulmaceae) and including

about 200 gall-inducing species markedly predominate gall midge species associated with her-

baceous plants.

The gall midge fauna of Eastern Europe including 287 gall-inducing species that are associated

with 41 plant families is similar to that occurring in Western Europe but the species number as-

sociated with trees and plants is much lower. For example, only 26 gall midge species are known

to occur on Salicaceae, in contrast to 64 species occurring in Western Europe and 9 species on

Fagaceae in contrast to 49 species in Western Europe.

The gall midge fauna occurring in nine islands situated in the Mediterranean Sea and including

153 gall midge species that are associated with 40 plant families is relatively poor.

Gall midge species cause galls mainly on herbaceous plants of the family Fabaceae and on trees

and shrubs of the family Fagaceae in second place. Islands in the Mediterranea area are densely

covered with broad-leaved forests with dominant tree species of the genus Quercus including

many species that are host plants of many gall midge species.

Kazakhstan is a large country situated in Central Asia and with only a small part in Eastern

Europe. It occupies an area of 2,727,300 km² which is greater than Western Europe. It includes very

diverse types of landscape: lowlands with steppes, taigas, rock-canyons, hills, deltas, mountains,

snow-capped mountains and deserts. The ﬂora of Kazakhstan is rich and includes about 6000 plant

species in 1118 genera and 161 families (Ryabushkina et al. 2008). Gall midge fauna occurring in

Kazakhstan is also rich: 714 gall-inducing species are associated with 54 plant families, i.e. with

141

33% of all plant families known in Kazakhstan (Fedotova 2000). Asteraceae with 139 gall-inducing

species is the richest family, followed by Fabaceae hosting 114 species, Chenopodiaceae with 90

species, Brassicaceae with 44 species and Rosaceae with 39 gall midge species. In Kazakhstan,

compared with Western and Eastern Europe, the majority of gall-inducing gall midge species are

associated with herbaceous plants and only a few species cause galls on trees and shrubs.

If we compare the species richness of gall midges in the four mentioned areas using the average

species number per one host plant family, Western Europe with 12.5 gall midge species associa-

Fig. 39. Relations of gall-inducing and plant-inhabiting gall midges of the subfamily Cecidomyiinae to host plant species

in countries of Europe given by the index of mutual occurrence.

142

ted with one plant family is in ﬁrst place, followed by Eastern Europe with 7 gall midge species

associated with one plant family and the Mediterranean with 3.8 gall midge species associated

with one plant family in last place in Europe. Of interest is the comparison of data from Europe

with data obtained in Kazakhstan situated in Central Asia where it has been found that an average

13.5 gall midge species are associated with one plant family.

Mutual occurrence of gall midges and host plants

As we have shown in previous sections, gall midges are not equally distributed in countries

of Europe. We try to analyse the relations between the species numbers of gall midges and the

numbers of plant species in particular countries and islands of Europe (Table 15). It is necessary

to mention that the numbers of species of ﬂowering plants occurring in particular country given

by various authors sometimes differ importantly. For some countries we were not able to obtain

recent data of species numbers of ﬂowering plants.

For the comparison of relations between gall midges and host plants we introduce here the

formula – the index of mutual gall midge species and plant species occurrence (IGS-PSO),

as follows:

IGS-PSO= NPS/NGMS

where NPS is the number of plant species in a particular country and NGMS is the number of gall

midge species occurring in the same part. This index indicates the degree of mutual occurrence

of gall midges on host plants in particular parts of Europe (Table 15, Fig. 39).

For example. If 5000 plant species have been recorded in a particular country and the fauna

of gall midges of that country includes 500 gall midge species, the index is IGS-PSO=5000/500

and the result after calculating is 10. This number indicates a high value of mutual occurrence of

gall midge species and host plants in that country. It means that we may ﬁnd in such a country

an average of one gall midge species on every tenth plant species. If in such a country the gall

midge fauna includes only 100 gall midge species, the index IGS-PSO=5000/100 and the result

after calculating is 50. In such a case we should ﬁnd one gall midge species on every 50th plant

species. Such a result indicates that the fauna of gall midges associated with plant species in

a particular country is poor.

Table 16. Distribution of galls of eleven major genera of gall-inducing Cecidomyiinae on different organs of host plants

in Europe

gall midges n species organs of host plants inquiline biology

genus stem bud leaf ﬂower fruit unknown

Dasineura 230 20 62 62 56 19 17

Contarinia 135 6 18 17 80 9 3

Asphondylia 50 0 13 1 20 19 0

Rabdophaga 40 20 16 4 0 1 1

Rhopalomyia 38 4 18 7 5 0 4

Jaapiella 35 0 10 8 12 1 4

Macrolabis 35 0 9 5 5 0 15

Mayetiola 28 27 0 0 0 0 0 1

Lasioptera 27 16 0 2 0 0 5 4

Planetella 27 8 0 6 0 0 0 13

Resseliella 23 13 1 0 0 4 2 3

total 668 114 147 112 178 53 51 21

143

It is necessary to note that the value of the index is greatly inﬂuenced by the intensity of inves-

tigation of the gall midge faunas of the various countries and islands as well as by the diversity

of the ﬂora. For example, the UK low ﬁgure results from the combination of a depauperate post-

-glacial ﬂora and long-term study of galls (K. M. Harris, pers. comm.).

Countries in Europe with an index value from 1 to 4 are noted for the highest mutual occurrence

of gall midges and host plants. The United Kingdom, Germany and the Czech Republic having

an index number of 4 are the countries with the highest mutual occurrence of gall midges and

host plants in Europe. Countries characterized by an index value from 5 to 10 are noted for high

mutual occurrence, viz. Latvia, the Netherlands, Poland, South Tyrol, France, Sweden, Austria

and Slovakia. Eight countries, viz. Belgium, Romania, Hungary, Bulgaria, Slovenia, Switzerland,

European part of Russia and Italy having been characterized by the index numbers from 11 to 20

may be evaluated as countries with medium mutual occurrence of gall midges and host plants.

Five countries, viz. Spain and Portugal, Corsica, Sicily and Ukraine, characterized by the index

numbers from 21 to 30, are countries of weak mutual occurrence of gall midges and host plants.

Greece and ﬁve islands in the Mediterrranean Sea (Mallorca, Sardinia, Crete, Malta and Cyprus)

with index numbers from 31 to 60 are countries with poor mutual occurrence of gall midges and

host plants. The Faroe Islands situated in the Atlantic Ocean in northern Europe have a very poor

mutual occurrence of gall midges and host plants characterized with the index number 80.

Although Greece is in Europe the country with the richest ﬂora, including 6700 species of

ﬂowering plants, the gall midge fauna of Greece, including 211 species, may be classed as poor.

Table 17. Species numbers of gall-inducing Cecidomyiinae associated with main broad-leaved and coniferous trees in

Europe. Abbreviations of countries: A – Austria, CH – Switzerland, CZ – Czech Republic, D – Denmark, F – France,

P – Poland (data from Skuhravá & Skuhravý 1997a, b, 2009a, Skuhravá et al. 2002, 2005, 2006, 2007, 2009, Skuhravý

& Skuhravá 1998)

gall midge species number

Europe F A CH CZ P D Sicily Malta Crete

broad-leaved trees

Fagus sylvatica 4 4 4 3 4 4 4 2 0 0

Quercus robur 15 14 8 3 6 7 7 0 0 0

Acer pseudoplatanus 4 4 4 4 4 4 3 1 0 0

Fraxinus excelsior 5 4 4 3 4 4 3 0 0 0

Tilia platyphyllos 8 5 6 5 5 5 5 0 0 0

Betula pubescens 10 6 4 2 5 4 5 0 0 0

Carpinus betulus 6 5 4 4 5 4 3 0 0 0

Populus tremula 13 11 9 8 9 10 7 0 0 0

Salix caprea 7 13 6 2 7 9 9 0 0 0

Olea europaea 4 2 0 0 0 0 0 4 1 2

Quercus cerris 13 2 9 0 8 1 0 1 0 0

Quercus ilex 5 5 0 0 0 0 0 4 0 0

total 94 75 58 34 57 52 46 12 1 2

coniferous trees

Picea abies 5 4 4 3 4 5 5 0 0 0

Pinus sylvestris 5 3 4 3 3 3 4 0 0 0

Abies alba 3 2 2 0 2 2 3 0 0 0

Larix decidua 3 2 2 1 2 3 2 0 0 0

Juniperus communis 7 4 2 2 3 4 3 0 0 0

Taxus baccata 1 1 1 1 1 1 1 0 0 0

total 23 16 15 10 15 18 18 0 0 0

144

We obtain similar relations between the rich ﬂora of Spain including 6500 species of ﬂowering

plants and 212 gall midge species found. Similar relations between rich ﬂoras and poorgall midge

faunas were found in islands in the Mediterranean Sea, viz. in Mallorca, Corsica, Sardinie, Sicily,

Malta, Corfu, Crete and Cyprus.

The analysis of the relationship between gall-inducing and plant-inhabiting gall midges of the

subfamily Cecidomyiinae and potential host plant species in the countries of Europe produced

a surprising result: although number of species of ﬂowering plants increases from north to south,

the species number of gall midges decreases.

Distribution of galls on host plants

Galls of gall-inducing species of the subfamily Cecidomyiinae are not equally distributed on

organs of their host plants. We analysed eleven genera of gall-inducing Cecidomyiinae occurring

in Europe on the basis of which organ of host plant is used for development of their larvae (Table

16). Of 668 species belonging to these eleven genera, 90% induce galls on various plant organs,

7% are inquilines in galls of other gall midges or insects and the biology of 3% is completely

unknown. Species of the genus Dasineura cause galls mainly on buds and leaves, of the genus

Contarinia on ﬂowers, of Asphondylia on ﬂowers and fruits, of Rabdophaga on stem and buds,

of Rhopalomyia on buds, of Jaapiella on buds and ﬂowers, species of the genus Macrolabis are

mainly inquilines in galls of other gall midges and species of Mayetiola, Lasioptera, Planetella

and Resseliella develop in galls mainly on stems. Some species that have two or more generations

per year may cause galls on two or more different organs of their host plant and therefore the sum

of inhabited plant organs in the table does not equal the number of gall midge species.

Abundance of gall midge species on host plant species

We use the term abundance here for the number of species associated with one host plant species.

Different numbers of species or genera of gall-inducing gall midges may develop on one host plant

species. The occurrence of gall midges (Cecidomyiidae) on host plants, mainly on trees, is not so

abundant as that of gall wasps (Cynipidae, Hymenoptera). For example, about 130 different gall-

-inducing species of gall wasps, belonging to several genera, are associated with Quercus robur

and Quercus petraea (Fagaceae) in Europe but only ﬁfteen gall-inducing species of gall midges,

also belonging to several genera (Skuhravá et al. 1998).

If we analyse the gall midge faunas of European countries from the point of view of association

with host plants, we ﬁnd that herbaceous plants are more frequently host plants than trees and

shrubs. Although trees and shrubs host fewer gall midge species than herbaceous plants, their

galls on leaves are usually more conspicuous and, therefore, much better known than small and

inconspicuous galls on herbaceous plants that are hidden among grasses near the soil.

For example: the gall midge fauna of Poland includes 397 species of the subfamily Cecido-

myiinae that are associated with 257 species of host plants belonging to 48 plant families of which

197 species (76%) are herbaceous plants and 60 species (24%) trees and shrubs (Skuhravá et al.

2008). Similar relationships between number of herbaceous plants and trees/ shrubs as host plants

were found in the gall midge fauna of Austria (Skuhravá & Skuhravý 2009a) and of Denmark

(Skuhravá et al. 2006). In Europe about 70% of gall-inducing species of the subfamily Cecido-

myiinae are associated with herbaceous plants and only 30% with trees and shrubs.

Many tree species are hosts of one, two, three or more gall midge species. Some gall midge

species are monophagous and are associated with only one host plant species, some are oligo-

phagous having been associated with several species of one host plant genus and only a few gall

midge species are polyphagous and are associated with species of several genera belonging to

145

one or more plant families. Gall midge species associated with twelve main broad-leaved tree

species and six coniferous trees in Europe are given in the Table 17.

Altogether one hundred and seventeen gall-inducing and plant inhabiting species are associated

with eighteen main tree species in Europe. Ninety four gall midge species are associated with

twelve main broad-leaved host tree species and twenty three gall midge species with six coni-

ferous trees. These trees are not distributed equally in Europe and, naturally, gall midge species

coincide with their host plants in distribution. Quercus robur is the host tree species with the

highest species number of Cecidomyiidae (15 species), followed by Populus tremula and Quercus

cerris, each with thirteen gall midge species, then Betula pendula with ten gall midge species and

Tilia platyphyllos with ten gall midge species. Olea europaea is the south-European plant species

and the boundary of its distribution area are considered to be the boundary of the Mediterranean

subregion. Gall midge species associated with broad-leaved trees occurring in Sicily, Malta and

Crete, parts of the Mediterranean subregion, form a characteristic species composition which

differs substantially from the gall midge species composition in other parts of Europe.

France, situated in the most western part of Europe bordering the Atlantic Ocean has the richest

gall midge species composition including 75 species that are associated with broad-leaves trees

and also the highest species numbers of gall midges associated with one species of host plant. It is

composed not only of gall midge species abundantly distributed in Central Europe but also includes

nine gall midge species associated with Quercus cerris, Q. ilex and Olea europaea originating

from the Mediterranean area and distributed only in the southernmost part of France. Some of

these gall midges reach to the southern parts of Austria and of the Czech Republic.

In Poland and Germany, situated in the northern part of continental Europe, there was found

the highest gall midge species number associated with coniferous trees. This is in harmony with

character of vegetation in this part of Europe. No gall midge species associated with coniferous

tree was found in Sicily, Malta and Crete that are situated in the Mediterranean area.

Coevolution of the family Cecidomyiidae with plants

Coevolution is deﬁnedby Lincoln etal. (1982) as the independentevolution of two ormore species

having an obvious ecological relationship, usually restricted to cases in which the interactions

are beneﬁcial to both species; it is also used for the evolutionary interactions between species

with some degree of independence, such as parasites and their hosts. Such relationships may be

observed between the gall midges and their host plants, host animals or other host organisms in

the course of their evolution.

No members of the family Cecidomyiidae are known to be associated with the most primitive

groups of organisms, i.e. with Prokaryota and Algae, and no gall midges have been found to be

directly associated with lichens (Lichenes) (Table 18, Fig. 40). It seems that at least two main

developmental lines of gall midges appeared during their evolution. In the past, members of the

family Cecidomyiidae were probably involved in interactions with three groups of fungi, i.e. with

Ascomycetes, Basidiomycetes and Deuteromycetes. This is supported by the fact that in the recent

time we have evidence that the larvae of gall midges develop in association with various groups

of fungi (Nijveldt 1969, Skuhravá et al. 1984, Yakovlev 1994, Ševčík 2006).

The current associations may result from more than one coevolutionary event. We know that

the ability of organisms to induce galls has arisen on many separate occasions so the same may

be true within cecidomyiid lineages. Coevolution of gall-inducing and plant inhabiting species of

the family Cecidomyiidae with two groups of vascular plants, Pteridophyta and Spermatophyta,

is shown on Fig. 40. Plant groups marked by red frames indicate host plant species suitable for

development of gall midges.

146

The ferns (Pteridophyta) ﬁrst appeared in the fossil record in the early-Carboniferous period.

By the Triassic, the ﬁrst evidence of ferns related to several modern families appeared. The“great

fern radiation” occurred in the late-Cretaceous, when many modern families of ferns ﬁrst appeared.

There is no fossil evidence of galls of gall midges on ferns and only two extant gall midge species

are known to occur on ferns in Europe, viz Dasineura pteridis and D. pteridicola, both causing

galls on Pteridium aquilinum of the family Dennstaedtiaceae.

The seed plants (Spermatophyta, Gymnospermae) originated in the late Carboniferous Period.

Early characteristics of seed plants were evident in fossil Progymnospermes of the late Devonian

period around 380 million years ago. Conifers are by far the most abundant gymnosperms with

around 600 plant species. Only one fossil record supports the coevolution of gall midges with

conifers – Sequoiomyia kraeuseli found in fossil cones of Sequoia langsdorﬁi of the family Ta-

xodiaceae. A relatively low species number of twenty six gall midge species currently occurring

in Europe indicates a low degree of coevolution.

Cycads are the next most abundant group with about 130 plant species and without fossil evi-

dence of coevolution with gall midges. Gnetales included approximately 75–80 species. There

is no fossil evidence about the galls of gall midges on Gnetales. Only one recent gall midge spe-

cies is known – Xerephedromyia ustjurtensis inducing galls on Ephedra distachya of the family

Ephedraceae in Europe.

Fig. 40. Coevolution of gall-inducing and plant inhabiting species of the family Cecidomyiidae with two groups of vascular

plants, Pteridophyta and Spermatophyta, according to the phylogenetic system proposed by Kenrick & Crane (1997) and

Smith et al. (2006). Plant groups marked by red frames indicate host plant species of gall midges.

147

The ﬂowering plants (Magnoliophyta, Angiospermae) are the most abundant group of plants,

including about 260,000 recent and many fossil species. Most gall midges known to occur in

Europe, including 870 recent species, are associated with this group of plants. Fossil galls of ten

gall midge species provide evidence of their coevolution with ﬂowering plants in the Miocene

and Pliocene of the Tertiary in various parts of Europe.

Adults of eighty-nine fossil species of 54 genera of the family Cecidomyiidae are known from

various parts of the world (Evenhuis 1994, Skuhravá 2006). Fossil gall midge adults were found

mainly in amber and copal and are dated from the Upper Jurassic, Lower Cretaceous, Upper

Cretaceous, Eocene, Oligocene, Miocene, Pliocene and Holocene. Fossil gall midge adults were

found scattered in all biogeographic regions except the Australasian and Oceanian regions. The

oldest fossil gall midge species, Catotricha mesozoica Kovalev, 1990, found in Siberia from the

Upper Jurassic and Lower Cretaceous, provides evidence that members of the Lestremiinae and

Porricondylinae probably originated and developed in the Mesozoic (Kovalev 1990).

In Europe, fossil gall midge adults were found mainly in amber of the Baltic Region. They

belong mainly to the subfamilies Porricondylinae and Lestremiinae and only a very few species

are Cecidomyiinae. Single species, sometimes identiﬁed to the genus level only, were found

in the UK, Denmark, Germany, France, Austria and Italy. Fossil adults of several gall midge

species were discovered in the earliest Eocene amber in northern France (Nel & Prokop 2006)

and in later Eocene amber in western Ukraine (Fedotova & Perkovsky 2004, 2005, Perkovsky

& Fedotova 2004).

There are only a few data about fossil galls induced by gall midges on organs of fossil host

plants in the world (Larew 1992). In Europe the galls of ten gall-causing species were found,

mainly in the form of the leaf impressions of host plant trees from the Tertiary Upper Miocene

and Pliocene (Trotter 1903, Rübsaamen 1929). Möhn (1960) discovered the fossil gall midge

species Sequoiomyia kraeuseli in the cone seeds of the fossil plant Sequoia langsdorﬁi (Taxodi-

aceae) in Rheinland (Germany) from the Miocene. Straus (1977) mentioned among fossil galls

Table 18. Recent species numbers of the family Cecidomyiidae associated with the main groups and subgroups of or-

ganisms in Europe. Groups of vascular plants are given according to the phylogenetical system proposed by Kenrick &

Crane (1997) and Smith et al. (2006)

phylogenetical group or subgroup of organisms and plants recent species number of gall midges

Prokaryota Cyanophyta 0

Bacteriophyta 0

Algae 0

Mycophyta (fungi) Ascomycetes 7

Basidiomycetes 70

Deuteromycetes 0

Lichenes 0

Bryophyta (mosses) 0

Vascular plants Pteridophyta (ferns) Pteridopsida (true ferns) 2

(897 gall midge species) Spermatophyta Cycadophyta 0

(Gymnospermae) Pinophyta (conifers) 26

(seed plants) Ginkgophyta 0

Gnetophyta 1

Magnoliophyta (Angiospermae)

(ﬂowering plants) 870

148

found at Willershausen in Harz (Germany) in the Pliocene also six fossil galls that were probably

caused by species of the family Cecidomyiidae. Villalta (1957) found in Spain traces of galls

which he identiﬁed as Mikiola pontiensis. It was a misidentiﬁcation. On the basis of their form,

these galls were caused by another gall midge species – Phegomyia fagicola (Kieffer, 1913), an

extant species (Skuhravá 2006). Information about fossil galls caused by gall midge species is

summarised in Table 19.

Fossil galls of ten gall midge species were found on nine genera of host plants belonging to

eight plant families. Most gall midges induced galls on leaves and only one species inhabited

cones. Fagaceae are the plant family hosting the highest number of gall midges – four species,

and each of the other families hosts only one gall midge species.

It is necessary to note that three gall midge species given in Table 19 were incorrectly identi-

ﬁed in the past. Identiﬁcation of the species causing galls on leaves of Fagus castaneifolia was

corrected by Skuhravá (2006). The galls on leaves of Betula are caused by another species, not

by Contarinia carpini which is speciﬁcally associated with Carpinus. Also the galls on leaves

of Quercus given as Phegomyia fagicola were incorrectly identiﬁed. P. fagicola is speciﬁcally

restricted to Fagus and could not cause galls on leaves of Quercus.

During evolution, the gall-inducing and plant inhabiting gall midges of the subfamily Ceci-

domyiinae probably started to develop at the end of the Mesozoic when they entered into inte-

ractions with developing groups of plants, started to inhabit suitable parts of plants, their organs

Table 19. Fossil galls induced by gall midges of the subfamily Cecidomyiinae on host plants found in Europe

gall midge species host plant host family host organ locality age reference

Sequoiomyia Sequoia langsdorﬁi Taxodiaceae cones Germany: Duren Miocene? Möhn 1960

kraeuseli

Asphondylia Sarothamnus Fabaceae leaf Europe, not precised tertiary Rübsaamen

sarothamni 1929

Cecidomyia sp. Populus Salicaceae leaf Switzerland: Oeningen tertiary Trotter 1903

Contarinia Betula Betulaceae leaf Germany: Willershausen pliocene Straus 1977

carpini (misidentiﬁcation)

Dasineura Fraxinus Oleaceae leaf Germany: Willershausen pliocene Straus 1977

fraxini

Dasineura Carpinus Corylaceae leaf Germany: Willershausen pliocene Straus 1977

ruebsaameni?

Didymomyia Tilia Tiliaceae leaf Germany: Willershausen pliocene Straus 1977

tiliacea

Mikiola fagi Fagus pliocenica Fagaceae leaf France: pliocene Marty 1894

Pas-de-la-Mougudo

Mikiola fagi Fagus Fagaceae leaf Europe, not precised tertiary Rübsaamen

1929

Phegomyia Fagus Fagaceae leaf Spain: Cerdãna miocene Villalta

fagicola castaneaefolia 1957,

(misidentiﬁed) Skuhravá

as Mikiola? pontiensis 2006

Phegomyia Quercus Fagaceae leaf Germany: Willershausen pliocene Straus 1977

fagicola

(misidentiﬁcation)

149

and tissues inducing there changes that initiated the formation of galls. The main development of

gall-inducing and plant inhabiting gall midges came in the Pliocene and Miocene of the Tertiary,

together and in parallel with the development of ﬂowering plants, as is supported by the evidence

of several fossil galls of gall midges.

Factors affecting gall midge species richness

From our observations and literature research we conclude that gall midge species richness is

inﬂuenced mainly by abiotic factors, i.e. by geographic and altitudinal position of the area under

study that are associated with changing climatic variables including sunshine, temperature and

rainfall; by biotic factors, i.e. mainly by the type of vegetation cover and composition of natural

vegetation; and by anthropic factors, involving various human activities. The present species

richness and distribution of gall midge species in Europe are results of all these inﬂuences which

are brieﬂycommented on below.In most cases detailedinformation is provided in earliersections

of this article.

Geographic position. In Europe the most suitable area for gall midges is the temperate zone

situated between latitudes 45° and 52° North, which is covered with broadleaved, coniferous and

mixed forests. Gall midge species number changes with geographical position and decreases both

northwards and southwards. On the other hand, species numbers of gall midges are relatively

stable along the west-east axis. The exceptionally low gall midge species number in the Faroe

Islands, situated in the Atlantic Ocean far off the European continent, is the result of geographical

isolation.

Altitude. Generally speaking, the number of gall midge species decreases with rising altitude.

Species numbers in the planare zone of Europe are usually low and are much higher in colline,

submontane and montane zones. Above the timber line, usually at an altitude of about 1500 m

in Europe (this altitude differs in various parts of Europe with geographic position) the species

number falls abruptly and only a few species occur in the sub-Alpine and Alpine zones.

Climatic factors. Gall midge species number is relative high in Central, Western and Eastern

Europe and falls to the south. This corresponds with changing climatic factors that are shown in

climatograms: the temperature increases to the south and, on the other hand, the rainfall decreases

and the hot and dry weather in southern Europe may be unsuitable for the development of gall

midges and their galls on host plants.

Composition of vegetation. Richness of plant species, providing plenty of potential hosts for

gall midges, is the most important prerequisite of richness of gall midges in the area. When we

compared the number of plant species in various parts of Europe with the number of gall midge

species, we discovered a very surprising fact that the number of gall midge species decreases to

the south of Europe although the number of plant species increases.

Association with host plant species. Most gall-inducing gall midges are speciﬁcallyrestricted to

their host plants and usually mirror the distribution of their host plants in their native environments,

but in a few cases their distribution may be extended into new areas when their host plants become

established in distant areas, such as Siberia, the Far East or even North America.

Anthropic factors. People have changed landscapes in Europe for a long time. Mediterranean

areas have been exploited in ancient times by clearing, burning and grazing and natural forests

have been replaced by olive trees, carob, wine grapes and citrus trees. Also landscapes in other

parts of Europe changed under inﬂuence of man which gradually decreased the number of gall

150

midge species. Some species that were abundant in the past disappeared as a result of the degra-

dation and destruction of biotopes, the use of chemical agents in plant protection against pests and

intensive fertilization of ﬁelds, cuttingand burning of grasson headlands andfallows, destruction

of small forest areas, use of land for new building activities, park and orchard arrangements with

regular cutting of grass, cutting of grass on slopes, formerly left without human interventions,

cutting of reeds and adjustment of littoral growths, amelioration of wetlands and wet meadows.

In the course of all these activities, the natural growth and multiplication of host plants of gall

midges is restricted and thus, the populations of plants and animals that are dependent on them

(including Cecidomyiidae) are progressively reduced (Skuhravá 2005).

Intensity of investigations. Results of gall midge species numbers in the area under study are

affected also by the number of investigations and by selection of suitable sites for collection of

gall midges. More investigations and more samples usually result in increased species numbers,

new records and also discovery of new species. It is necessary also to emphasize the importance

of the researchers, their ability, experience, knowledge and enthusiasm that make it possible

to discover gradually additional gall midge species and hidden factors inﬂuencing gall midge

species richness.

A c k n o w l e d g e m e n t s

We would like to express our thanks to Keith M. Harris (Ripley, Woking, Surrey, England) for valuable comments on the

manuscript and for improvements of the English text. We thank Jan Zárybnický, Agency for Nature Conservation and

Landscape Protection of the Czech Republic, Prague, for all advice and for preparing maps according to the map basis of

species numbers of Sweden. We are grateful to our daughter, Ivana Vaněčková, and our granddaughter, Pavla Kosová,

for their help in preparing computer version of graphs.

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Gall midges (Diptera: Cecidomyiidae) of Algeria

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Full-text available

Dec 2020

The known fauna of gall midges in Algeria is composed of 109 valid species belonging to 43 genera, and four undescribed species. It is the richest fauna of five countries spread along the Mediterranean Sea in North Africa. Twenty five species of gall midges were decribed on the basis of material collected in Algeria and have there their type localities. The two most abundant species of gall midges in Algeria are Asphondylia punica Marchal, 1897, causing galls on Atriplex halimus, and Rhopalomyia navasi Tavares, 1904, inducing galls on Artemisia herba-alba. Galls of R. navasi were recorded at two localities in northwestern Algeria recently, in the year 2015. the most species rich genus is Dasineura Rondani, 1840, with twenty five species, followed by Asphondylia Loew, 1850 (ten species), Contarinia Rondani, 1860 (eight species), and Stefaniola Kieffer, 1913 (seven species). On the basis of larval feeding habits most species are phytophagous, ten species are mycophagous and Aphidoletes aphidimyza (Rondani, 1847) and Arthrocnodax clematitis Marchal, 1897 are zoophagous. Phytophagous species are associated with 83 host plant species belonging to 65 genera of 29 plant families. Chenopodiceae is the plant family hosting the highest number of gall midges-sixteen species of gall midges. Three quarter of host plant species are herbaceous plants and one quarter trees or shrubs. Usually only one species of gall midge is associated with one host plant species. Four host plants-Atriplex halimus, Fraxinus excelsior, Phillyrea latifolia, and Salsola vermiculata-are associated with three species of gall midges. Erica arborea is the host plant with the highest number of gall midges: it hosts Dasineura ericaescopariae (Dufour, 1837), D. grasseti Barnes, 1935, Myricomyia mediterranea (Löw, 1885), and Wachtliella ericina (Löw, 1885). Zoogeographical analysis: 62 species (57%) are Mediterranean, 33 species (30%) European, ten species (8%) Eurosiberian; Micromya lucorum rondani, 1840, Cedrocrypta montana Kieffer, 1919 and Aphidoletes aphidimyza (Rondani, 1847) are Holarctic species; Mayetiola destructor (Say, 1817) is Palaearctic species. Economic importance: no significant pests in agriculture and forestry were recorded. Mayetiola destructor, M. hordei Kieffer, 1909, and M. avenae (Marchal, 1895) are potential pests of cereals.

Ecology and Evolution of Gall-Inducing Arthropods: The Pattern From the Terrestrial Fossil Record

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Jun 2021

Conrad Labandeira

Insect and mite galls on land plants have a spotty but periodically rich and abundant fossil record of damage types (DTs), ichnotaxa, and informally described gall morphotypes. The earliest gall is on a liverwort of the Middle Devonian Period at 385 million years ago (Ma). A 70-million-year-long absence of documented gall activity ensues. Gall activity resumes during the Pennsylvanian Period (315 Ma) on vegetative and reproductive axial organs of horsetails, ferns, and probably conifers, followed by extensive diversification of small, early hemipteroid galler lineages on seed-plant foliage during the Permian Period. The end-Permian (P-Tr) evolutionary and ecological crisis extinguished most gall lineages; survivors diversified whose herbivore component communities surpassed pre-P-Tr levels within 10 million years in the mid-to late Triassic (242 Ma). During the late Triassic and Jurassic Period, new groups of galling insects colonized Ginkgoales, Bennettitales, Pinales, Gnetales, and other gymnosperms, but data are sparse. Diversifying mid-Cretaceous (125–90 Ma) angiosperms hosted a major expansion of 24 gall DTs organized as herbivore component communities, each in overlapping Venn-diagram fashion on early lineages of Austrobaileyales, Laurales, Chloranthales, and Eurosidae for the Dakota Fm (103 Ma). Gall diversification continued into the Ora Fm (92 Ma) of Israel with another 25 gall morphotypes, but as ichnospecies on a different spectrum of plant hosts alongside the earliest occurrence of parasitoid attack. The End-Cretaceous (K-Pg) extinction event (66 Ma) almost extinguished host–specialist DTs; surviving gall lineages expanded to a pre-K-Pg level 10 million years later at the Paleocene-Eocene Thermal Maximum (PETM) (56 Ma), at which time a dramatic increase of land surface temperatures and multiplying of atmospheric pCO2 levels induced a significant level of increased herbivory, although gall diversity increased only after the PETM excursion and during the Early Eocene Climatic Optimum (EECO). After the EECO, modern (or structurally convergent) gall morphotypes originate in the mid-Paleogene (49–40 Ma), evidenced by the Republic, Messel, and Eckfeld floras on hosts different from their modern analogs. During subsequent global aridification, the early Neogene (20 Ma) Most flora of the Czech Republic records several modern associations with gallers and plant hosts congeneric with their modern analogs. Except for 21 gall DTs in New Zealand flora, the gall record decreases in richness, although an early Pleistocene (3 Ma) study in France documents the same plant surviving as an endemic northern Iran but with decreasing associational, including gall, host specificity.

Pharmacological Properties of Four Plant Species of the Genus Anabasis, Amaranthaceae

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May 2023
MOLECULES

The genus Anabasis is a member of the family Amaranthaceae (former name: Chenopodiaceae) and includes approximately 102 genera and 1400 species. The genus Anabasis is one of the most significant families in salt marshes, semi-deserts, and other harsh environments. They are also renowned for their abundance in bioactive compounds, including sesquiterpenes, diterpenes, triterpenes, saponins, phenolic acids, flavonoids, and betalain pigments. Since ancient times, these plants have been used to treat various diseases of the gastrointestinal tract, diabetes, hypertension, and cardiovascular diseases and are used as an antirheumatic and diuretic. At the same time, the genus Anabasis is very rich in biologically active secondary metabolites that exhibit great pharmacological properties such as antioxidant, antibacterial, antiangiogenic, antiulcer, hypoglycemic, hepatoprotective, antidiabetic, etc. All of the listed pharmacological activities have been studied in practice by scientists from different countries and are presented in this review article to familiarize the entire scientific community with the results of these studies, as well as to explore the possibilities of using four plant species of the genus Anabasis as medicinal raw materials and developing medicines based on them.

Shedding Light on Dasineura oleae Parasitoids: Local and Landscape Effects

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Feb 2023

Dasineura oleae was considered a minor pest in olive orchards. However, in the last decade, outbreaks have been reported all over its distribution area. Little is known about D. oleae biological control strategies; therefore, investigations into the biology and ecology of D. oleae parasitoids are urgently needed. In this scenario, the present field study reported the flight period of D. oleae parasitoids, evaluating their relative abundance over other parasitoids living in olive orchards. Furthermore, it estimated the effect of local and landscape features on D. oleae parasitoids within the frame of the overall parasitoid community. Lastly, we aimed to provide useful insights into the effectiveness of parasitoids for D. oleae population management. Hymenopteran parasitoids were sampled using Malaise traps in six sampling sites in central Italy. Results showed that Platygaster demades was the most abundant D. oleae parasitoid. Its presence was associated with high rates of D. oleae parasitism. The abundance of this parasitoid was influenced by the abundance of seminatural habitats.

Additions to the fauna of gall midges (Diptera, Cecidomyiidae) in Iran

Article

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Dec 2022

Detection of a Diverse Endophyte Assemblage within Fungal Communities Associated with the Arundo Leaf Miner, Lasioptera donacis (Diptera: Cecidomyiidae)

Article

Full-text available

Apr 2023
Diversity

The larvae of Lasioptera donacis Coutin feed on fungal communities lining galleries within the mesophyll of leaf sheaths of Arundo donax in an aggregative manner. It has been stated that L. donacis could have established a fundamental symbiotic relationship with one fungus, although the fungal composition of these communities remains unsettled. Using a culture-dependent approach and ITS sequencing, the present work characterizes and compares the fungal communities associated with L. donacis in Eurasia with the endophytes of A. donax in Texas where L. donacis is absent. The 65 cultivable isolates obtained from L. donacis fungal communities were sorted into 15 MOTUs, among which Fusarium and Sarocladium predominated. No particular MOTU was systematically recovered from these communities regardless of the sites. The 19 isolates obtained in Texas were sorted into 11 MOTUs. Sarocladium and Fusarium were commonly found in Texas and Eurasia. Our finding indicate that the communities were composed of a diverse assemblage of non-systemic endophytes, rather than an exclusive fungal symbiont. From ovipositors and ovarioles of L. donacis emerging from plants in France, we opportunistically isolated the endophyte Apiospora arundinis, which lies at the origin of further research pertaining to its role in the feeding and oviposition of L. donacis.

Fauna, trophic connections, and morpho-functional adaptations of gall midges (Diptera, Cecidomyiidae) forming parenchymal leaf galls of arboreal and shrubby plants. Izvestia Sankt-Peterburgskoj Lesotehniceskoj Akademii

Article

Full-text available

Nov 2022

Zoya Alexandrovna Fedotova

For the first time, a review of gall midges-inquilines, developing together with gall-formers, among which gall midges dominate is provided. In the world, 197 species of gall midges of 41 genera have been identified, in the galls of which 177 species of inquiline gall midges from 27 genera have been found. They are found on plants of 243 species of 160 genera of 53 families and 20 orders. The core of the gall-forming fauna is Cecidomyiinae – 118 species (59.9%) from 26 genera (63.4%), and the core of the inquiline fauna is Lasiopterinae: 63 species (32.0%) from 15 genera (36.6%). Ten common genera were identified, in which there are both inquilines and gall-formers. The proportion of inquilines in these genera is more than a third of the species, for Macrolabis – 39.1% (25 out of 64) and Camptoneuromyia – 86.7% (13 out of 15). Gall-formers and inquilines are predominantly narrow oligophages, specific in respect to the genus or family of the host plant, predominantly Fabaceae. Among gall-formers, the share of specific genera is 61.0% (25 out of 41), while among inquilines is 37.0% (10 out of 27). Host plants of the Rosids subclass are represented by 124 species (51.0%), 88 genera (55.0%) from 28 families (46.2%), most of which are trees and shrubs. Inquiline gall midges have been found in all zoogeographic regions; however no common species have been found. They dominate in the Palaearctic (118 species, 66.7% of 177) and Neotropical (40 species, 22.6%) regions. In the Nearctic region there are only 15 (8.5%) species. In the Palaearctic, inquiline species have been identified in 19 genera (70.4% of 27), of which 7 are endemic. In the Neotropical region, the inquilines belong to 7 genera (25.9%), with no endemics found. The core of the fauna with an abundance of endemic and widespread genera of inquilines formed in the Palaearctic region. In the gall midges, inquiline gall midges predominate, but inquilines develop from other taxonomic groups: insects (with a predominance of Cynipidae) and fungi. Inquilines present the potential for their gradual transition to gall formation and possible speciation during the assimilation of galls and plants of other species. In the galls of gall midges, inquilines actively influence the formation of galls and the development of the host larvae, contribute to their feeding, inhibit or lead to death. Key words : complexes of gall midges, specific genera, inquilines, gall-formers, endemics, host plants, evolution.

Haplodiplosis marginata (saddle gall midge)

Article

Jan 2022

This datasheet on Haplodiplosis marginata covers Identity, Overview, Distribution, Dispersal, Hosts/Species Affected, Diagnosis, Biology & Ecology, Environmental Requirements, Natural Enemies, Impacts, Uses, Prevention/Control, Further Information.

New records and expansion of the geographic distribution of gall inducers of the family Cecidomyiidae (Diptera) associated to Andira Lam. (Fabaceae) species in Brazil

Article

Full-text available

Oct 2021

Galls are structures of vegetal tissue modified by several biological agents, mainly dipterans of the Cecidomyiidae family. The galls are indicators of the gall inducers’ presence, because of the species-specific interaction between the gall inducer and its host plant species. The aim of this study was to expand the geographical distribution of cecidomyiids – usually known only to the type-locality – in species of Andira Lam. (Fabaceae) distributed in Brazil, through the presence of their gall morphotypes in host plants. We searched for records of gall morphotypes in Andira species in all inventories of insect galls in Brazilian biomes and in virtual herbaria in Brazilian and abroad institutions. We found six species of Andira with forty-nine registers of 20 morphotypes of galls induced by cecidomyiids. They were from 35 localities; of those, 15 Municipalities, five States and the Federal District are new points of occurrence. We found three new registers for the fusiform gall in A. fraxinifolia, four registers for the fusiform gall in A. humilis, two new registers for the lenticular gall in A. nitida, and six new registers for three morphotypes in A. vermifuga. No new records were found for the morphotypes of A. cujabensis and A. surinamensis. These results indicate that digital tools, combined with well-illustrated inventory data, are capable to expand knowledge about the distribution of cecidomyiids through the recognition of their specific marker, the morphotypes. Keywords: Andira fraxinifolia; Andira humilis; Andira nítida; Andira vermífuga; Insect-plant interaction

Sampling Galls and Galling Arthropods

Chapter

Full-text available

Nov 2020

Gall-inducing insects and mites are able to manipulate plant tissues to induce galls. As galling species are sessile, their sampling is relatively easy, but many aspects must be taken into account in order to adequately characterize galling fauna. Aspects related to the adoption of the methods for field sampling, standardization of gall morphotypes, and adequate photographic record of the galls are essential for a good sampling of galling arthropods. The adequate laboratory processing of host plants, as well as the treatment of galls to obtain gallers, and the subsequent identification of galling taxa are also important aspects for studies involving plant-galling interactions.

New gall midges (Diptera, Cecidomyiidae) from the Rovno Amber: Subfamily porricondylinae (Tribes Bryocryptini and Winnertziini) and subfamily lasiopterinae (Tribes Brachineurini and Oligotrophini)

Article

Full-text available

Jan 2005

Eight new species of gall midges of the genera Bryocrypta, Winnertzia, Brachineurina, Jaapiella, and Didymomyia are described from the Eocene Rovno amber.

Species richness of gall midges (Diptera: Cecidomyiidae) in the main biogeographical regions of the world

Article

Full-text available

Jan 2006

Marcela Skuhravá

Gall midges of the subfamily Lestremiinae (Diptera, Cecidomyiidae) in Estonia, Lithuania and Latvia: a check-list and the description of new species.

Article

Full-text available

Jan 2000

Voldemars Spungis

Invasive spread of the gall midge Obolodiplosis robiniae in Europe

Article

Full-text available

Jan 2006

(A review of gall-midges ( Cecidomyiidae, Diptera) of Poland).

Article

Jan 1983

Contains a review of 320 gall-midge species ascertained in Poland. From the zoogeographical veiwpoint the most interesting species occur in the High Tatra regions on both the Polish and Czechoslovak sides: Jaapiella alpina (on Silene acaulis), Dasyneura alpestris (on Arabis alpina) and Dasyneura phyteumatis (on Phyteuma orniculare and Phyteuma spiceatum).-from English summary

Synopsis des zoocécidies d'Europe

Article