Horse-chestnut leaf miner
Encyclopedia
The horse-chestnut leaf miner (Cameraria ohridella) is a leaf-mining
moth
of the Gracillariidae
family
. The horse-chestnut leaf miner was first observed in Macedonia
in 1984, and was described as a new species in 1986. Its larva
e are leaf miner
s on the Common Horse-chestnut
(Aesculus hippocastanum). The horse-chestnut leafminer was first collected and inadvertently pressed in herbarium sheets by the botanist T.Heldreich
in central Greece in 1879.
The larva feeds in a mine in the leaves of the tree, damaging the leaves and stunting growth. Infected leaves are covered in small brown patches which spread rapidly across the entire tree, giving an autumnal appearance. Eventually the leaves die and drop off; when new ones grow they are again infected. This cycle can repeat itself several times in one season.
e develop through five feeding phases (or instar
s) and two prepupal (spinning) phases before the pupal phase. The first stage creates a small cavity (or mine) parallel to a vein in the leaf and is "sap
-sipping" rather than "tissue
-feeding". By the third instar, the larva creates a mine approximately 8mm in diameter; this is further expanded by later instars until one mine can cover several square centimeters. The larva starts to pupa
te around four weeks after the egg hatches and except when hibernating as a pupa in the mine, the adult emerges around two weeks later. In severe infestations, the mines of individuals can merge and almost the entire leaf area may be utilised. When this occurs it may lead to high moth mortality as the larvae compete
for space and food. The moth is able to go through up to five generations each year, if the weather is hot and dry; on average in western Europe, the moth goes through three generations each year. The last generation of the year pupates for over six months so as to survive the winter. The pupae are extremely frost tolerant and have been recorded to survive temperatures as low as -23 C. This allows its populations to increase even after hard winters.
The dead patches that the horse-chestnut leaf miner causes on leaves are similar to damage caused to horse-chestnut trees by the fungus
Guignardia
aesculi, but can be distinguished by the fungal infection often being outlined by a conspicuous yellow band which the mines lack. The pupae can be mistaken for pupae of the Phyllonorycter
genus but can be distinguished because the C. ohridella pupae do not have a cremaster
and the first five abdominal sections have strong lateral spines on them.
, Austria
, Belgium
, Belarus
, Bosnia and Herzegovina
, Bulgaria
, Croatia
, the Czech Republic
, Denmark
, England
and Wales
, Finland
, France
, Germany
, Greece
, Hungary
, Kosovo
, Italy
, Latvia
, Liechenstein, Lithuania
, Luxembourg
, Macedonia
, Moldova
, Montenegro
, the Netherlands
, Poland
, Serbia
, Romania
, Russia
, Serbia
, Slovakia
, Slovenia
, Spain
, Sweden
, Switzerland
, Turkey
and Ukraine
. Although horse-chestnut occurs naturally mostly above the 700–1000 m. contours the moth does well in well-watered places such as parks in cities and at low elevation but not well in the hotter parts of Europe e.g. Spain
. Probably aided greatly by vehicular transport, the moth has attained a very rapid dispersal rate across Europe of 60 km. per year.
in 1984, and was described as a new species by Deschka and Dimić in 1986. A likely Balkan origin for this moth was evidenced from a decrease in genetic diversity from natural towards artificial horse-chestnut stands that were planted around Europe since around 1600. This Balkan origin is further documented by numerous herbarium samples that date back to 1879. These include an outbreak which occurred in horse-chestnut specimens collected by F.K. Meyer in 1961 in Albania. Of the 30 known mitochondrial haplotype
s for the species only three (known as A, B and C) have invaded the rest of Europe since 1989, and only A is dominant. It is likely that the frequency of haplotype A has been increasing even in Balkan natural sites, aided by the late development of roads in the region.
) has also been found to predate C. ohridella, consuming around 10 larvae per day. Overall the predation by the southern oak bushcricket is insignificant compared to that by birds however. Experiments with the predatory mite
Euseius finlandicus, bush crickets (Phaneroptera
sp.), ladybirds and lacewings found that none predate C. ohridella.
is usually banned as it kills bees. Fenoxycarb
causes up to 100% pupal mortality, has low environmental toxicity and can be combined successfully with manual leaf removal. A synthetic pheromone can be used to trap males, but effective control may be hard to thus achieve. In any case, infestation levels could diminish over time as Cameraria ohridella starts to recruit generalist members of the local parasitoid wasp community.
-wide multidisciplinary project, CONTROCAM ("Control of Cameraria") and the HAM-CAM Project.
Leaf miner
Leaf miner is a term used to describe the larvae of many different species of insect which live in and eat the leaf tissue of plants. The vast majority of leaf-mining insects are moths , sawflies and flies , though some beetles and wasps also exhibit this behavior.Like Woodboring beetles, leaf...
moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...
of the Gracillariidae
Gracillariidae
Gracillariidae is an important family of insects in the order Lepidoptera and the principal family of leaf miners that includes several economic, horticultural or recently invasive pest species such as the horse-chestnut leaf miner, Cameraria ohridella....
family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
. The horse-chestnut leaf miner was first observed in Macedonia
Macedonia (Greece)
Macedonia is a geographical and historical region of Greece in Southern Europe. Macedonia is the largest and second most populous Greek region...
in 1984, and was described as a new species in 1986. Its larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
e are leaf miner
Leaf miner
Leaf miner is a term used to describe the larvae of many different species of insect which live in and eat the leaf tissue of plants. The vast majority of leaf-mining insects are moths , sawflies and flies , though some beetles and wasps also exhibit this behavior.Like Woodboring beetles, leaf...
s on the Common Horse-chestnut
Aesculus
The genus Aesculus comprises 13-19 species of woody trees and shrubs native to the temperate northern hemisphere, with 6 species native to North America and 7-13 species native to Eurasia; there are also several hybrids. Species are deciduous or evergreen...
(Aesculus hippocastanum). The horse-chestnut leafminer was first collected and inadvertently pressed in herbarium sheets by the botanist T.Heldreich
Theodor von Heldreich
Theodor Heinrich Hermann von Heldreich was a German botanist, who was born on 3 March 1822 in Dresden and died on 7 September 1902 in Athens....
in central Greece in 1879.
Damage
Cameraria ohridella causes significant damage, mainly late summer browning, to the appearance of horse-chestnut trees. Despite the poor appearance of these infested trees, there is no evidence that damage by the moth leads to tree death. Seed weight and photosynthetic storage and reproductive capacity may however be reduced. Trees survive repeated infestations and re-flush normally in the following year. It appears that most of the damage caused by the moth occurs too late in the growing season to greatly affect tree performance. Consequently, there is no reason to fell and remove trees just because they are attacked by C. ohridella.Description
The moth is to 5mm long, with shiny, bright brown forewings with thin, silvery white stripes. The hindwings are dark grey with long fringes. Each female moth lays between 20 and 40 eggs singly on the upper surface of leaves, and once these hatch 2–3 weeks later, the larvaLarva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
e develop through five feeding phases (or instar
Instar
An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...
s) and two prepupal (spinning) phases before the pupal phase. The first stage creates a small cavity (or mine) parallel to a vein in the leaf and is "sap
Plant sap
Sap is a fluid transported in xylem cells or phloem sieve tube elements of a plant. It transports water and nutrients throughout the plant....
-sipping" rather than "tissue
Tissue (biology)
Tissue is a cellular organizational level intermediate between cells and a complete organism. A tissue is an ensemble of cells, not necessarily identical, but from the same origin, that together carry out a specific function. These are called tissues because of their identical functioning...
-feeding". By the third instar, the larva creates a mine approximately 8mm in diameter; this is further expanded by later instars until one mine can cover several square centimeters. The larva starts to pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
te around four weeks after the egg hatches and except when hibernating as a pupa in the mine, the adult emerges around two weeks later. In severe infestations, the mines of individuals can merge and almost the entire leaf area may be utilised. When this occurs it may lead to high moth mortality as the larvae compete
Competition (biology)
Competition is an interaction between organisms or species, in which the fitness of one is lowered by the presence of another. Limited supply of at least one resource used by both is required. Competition both within and between species is an important topic in ecology, especially community ecology...
for space and food. The moth is able to go through up to five generations each year, if the weather is hot and dry; on average in western Europe, the moth goes through three generations each year. The last generation of the year pupates for over six months so as to survive the winter. The pupae are extremely frost tolerant and have been recorded to survive temperatures as low as -23 C. This allows its populations to increase even after hard winters.
The dead patches that the horse-chestnut leaf miner causes on leaves are similar to damage caused to horse-chestnut trees by the fungus
Fungus
A fungus is a member of a large group of eukaryotic organisms that includes microorganisms such as yeasts and molds , as well as the more familiar mushrooms. These organisms are classified as a kingdom, Fungi, which is separate from plants, animals, and bacteria...
Guignardia
Guignardia
Guignardia is a genus of fungi in the family Botryosphaeriaceae.-Species:*Guignardia abeana*Guignardia abietella-sibirica*Guignardia acaciae*Guignardia adeana*Guignardia adianti*Guignardia adriatica...
aesculi, but can be distinguished by the fungal infection often being outlined by a conspicuous yellow band which the mines lack. The pupae can be mistaken for pupae of the Phyllonorycter
Phyllonorycter
Phyllonorycter is a genus of moths in the family Gracillariidae.-Diversity:The genus comprises about 400 species, with a worldwide distribution. The vast majority of species are found in the temperate regions, with about 257 species described from the Palaearctic region and 81 from the Nearctic...
genus but can be distinguished because the C. ohridella pupae do not have a cremaster
Cremaster
Cremaster is a term derived from the Greek verb κρεμάννυμι = "I hang ", not from Latin cremare = "to burn". It may refer to:* The cremaster muscle, part of genital anatomy in human males** Cremaster reflex, a reflex in the muscle...
and the first five abdominal sections have strong lateral spines on them.
Distribution and dispersal
Cameraria ohridella has now been found in AlbaniaAlbania
Albania , officially known as the Republic of Albania , is a country in Southeastern Europe, in the Balkans region. It is bordered by Montenegro to the northwest, Kosovo to the northeast, the Republic of Macedonia to the east and Greece to the south and southeast. It has a coast on the Adriatic Sea...
, Austria
Austria
Austria , officially the Republic of Austria , is a landlocked country of roughly 8.4 million people in Central Europe. It is bordered by the Czech Republic and Germany to the north, Slovakia and Hungary to the east, Slovenia and Italy to the south, and Switzerland and Liechtenstein to the...
, Belgium
Belgium
Belgium , officially the Kingdom of Belgium, is a federal state in Western Europe. It is a founding member of the European Union and hosts the EU's headquarters, and those of several other major international organisations such as NATO.Belgium is also a member of, or affiliated to, many...
, Belarus
Belarus
Belarus , officially the Republic of Belarus, is a landlocked country in Eastern Europe, bordered clockwise by Russia to the northeast, Ukraine to the south, Poland to the west, and Lithuania and Latvia to the northwest. Its capital is Minsk; other major cities include Brest, Grodno , Gomel ,...
, Bosnia and Herzegovina
Bosnia and Herzegovina
Bosnia and Herzegovina , sometimes called Bosnia-Herzegovina or simply Bosnia, is a country in Southern Europe, on the Balkan Peninsula. Bordered by Croatia to the north, west and south, Serbia to the east, and Montenegro to the southeast, Bosnia and Herzegovina is almost landlocked, except for the...
, Bulgaria
Bulgaria
Bulgaria , officially the Republic of Bulgaria , is a parliamentary democracy within a unitary constitutional republic in Southeast Europe. The country borders Romania to the north, Serbia and Macedonia to the west, Greece and Turkey to the south, as well as the Black Sea to the east...
, Croatia
Croatia
Croatia , officially the Republic of Croatia , is a unitary democratic parliamentary republic in Europe at the crossroads of the Mitteleuropa, the Balkans, and the Mediterranean. Its capital and largest city is Zagreb. The country is divided into 20 counties and the city of Zagreb. Croatia covers ...
, the Czech Republic
Czech Republic
The Czech Republic is a landlocked country in Central Europe. The country is bordered by Poland to the northeast, Slovakia to the east, Austria to the south, and Germany to the west and northwest....
, Denmark
Denmark
Denmark is a Scandinavian country in Northern Europe. The countries of Denmark and Greenland, as well as the Faroe Islands, constitute the Kingdom of Denmark . It is the southernmost of the Nordic countries, southwest of Sweden and south of Norway, and bordered to the south by Germany. Denmark...
, England
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...
and Wales
Wales
Wales is a country that is part of the United Kingdom and the island of Great Britain, bordered by England to its east and the Atlantic Ocean and Irish Sea to its west. It has a population of three million, and a total area of 20,779 km²...
, Finland
Finland
Finland , officially the Republic of Finland, is a Nordic country situated in the Fennoscandian region of Northern Europe. It is bordered by Sweden in the west, Norway in the north and Russia in the east, while Estonia lies to its south across the Gulf of Finland.Around 5.4 million people reside...
, France
France
The French Republic , The French Republic , The French Republic , (commonly known as France , is a unitary semi-presidential republic in Western Europe with several overseas territories and islands located on other continents and in the Indian, Pacific, and Atlantic oceans. Metropolitan France...
, Germany
Germany
Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...
, Greece
Greece
Greece , officially the Hellenic Republic , and historically Hellas or the Republic of Greece in English, is a country in southeastern Europe....
, Hungary
Hungary
Hungary , officially the Republic of Hungary , is a landlocked country in Central Europe. It is situated in the Carpathian Basin and is bordered by Slovakia to the north, Ukraine and Romania to the east, Serbia and Croatia to the south, Slovenia to the southwest and Austria to the west. The...
, Kosovo
Kosovo
Kosovo is a region in southeastern Europe. Part of the Ottoman Empire for more than five centuries, later the Autonomous Province of Kosovo and Metohija within Serbia...
, Italy
Italy
Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...
, Latvia
Latvia
Latvia , officially the Republic of Latvia , is a country in the Baltic region of Northern Europe. It is bordered to the north by Estonia , to the south by Lithuania , to the east by the Russian Federation , to the southeast by Belarus and shares maritime borders to the west with Sweden...
, Liechenstein, Lithuania
Lithuania
Lithuania , officially the Republic of Lithuania is a country in Northern Europe, the biggest of the three Baltic states. It is situated along the southeastern shore of the Baltic Sea, whereby to the west lie Sweden and Denmark...
, Luxembourg
Luxembourg
Luxembourg , officially the Grand Duchy of Luxembourg , is a landlocked country in western Europe, bordered by Belgium, France, and Germany. It has two principal regions: the Oesling in the North as part of the Ardennes massif, and the Gutland in the south...
, Macedonia
Republic of Macedonia
Macedonia , officially the Republic of Macedonia , is a country located in the central Balkan peninsula in Southeast Europe. It is one of the successor states of the former Yugoslavia, from which it declared independence in 1991...
, Moldova
Moldova
Moldova , officially the Republic of Moldova is a landlocked state in Eastern Europe, located between Romania to the West and Ukraine to the North, East and South. It declared itself an independent state with the same boundaries as the preceding Moldavian Soviet Socialist Republic in 1991, as part...
, Montenegro
Montenegro
Montenegro Montenegrin: Crna Gora Црна Гора , meaning "Black Mountain") is a country located in Southeastern Europe. It has a coast on the Adriatic Sea to the south-west and is bordered by Croatia to the west, Bosnia and Herzegovina to the northwest, Serbia to the northeast and Albania to the...
, the Netherlands
Netherlands
The Netherlands is a constituent country of the Kingdom of the Netherlands, located mainly in North-West Europe and with several islands in the Caribbean. Mainland Netherlands borders the North Sea to the north and west, Belgium to the south, and Germany to the east, and shares maritime borders...
, Poland
Poland
Poland , officially the Republic of Poland , is a country in Central Europe bordered by Germany to the west; the Czech Republic and Slovakia to the south; Ukraine, Belarus and Lithuania to the east; and the Baltic Sea and Kaliningrad Oblast, a Russian exclave, to the north...
, Serbia
Serbia
Serbia , officially the Republic of Serbia , is a landlocked country located at the crossroads of Central and Southeast Europe, covering the southern part of the Carpathian basin and the central part of the Balkans...
, Romania
Romania
Romania is a country located at the crossroads of Central and Southeastern Europe, on the Lower Danube, within and outside the Carpathian arch, bordering on the Black Sea...
, Russia
Russia
Russia or , officially known as both Russia and the Russian Federation , is a country in northern Eurasia. It is a federal semi-presidential republic, comprising 83 federal subjects...
, Serbia
Serbia
Serbia , officially the Republic of Serbia , is a landlocked country located at the crossroads of Central and Southeast Europe, covering the southern part of the Carpathian basin and the central part of the Balkans...
, Slovakia
Slovakia
The Slovak Republic is a landlocked state in Central Europe. It has a population of over five million and an area of about . Slovakia is bordered by the Czech Republic and Austria to the west, Poland to the north, Ukraine to the east and Hungary to the south...
, Slovenia
Slovenia
Slovenia , officially the Republic of Slovenia , is a country in Central and Southeastern Europe touching the Alps and bordering the Mediterranean. Slovenia borders Italy to the west, Croatia to the south and east, Hungary to the northeast, and Austria to the north, and also has a small portion of...
, Spain
Spain
Spain , officially the Kingdom of Spain languages]] under the European Charter for Regional or Minority Languages. In each of these, Spain's official name is as follows:;;;;;;), is a country and member state of the European Union located in southwestern Europe on the Iberian Peninsula...
, Sweden
Sweden
Sweden , officially the Kingdom of Sweden , is a Nordic country on the Scandinavian Peninsula in Northern Europe. Sweden borders with Norway and Finland and is connected to Denmark by a bridge-tunnel across the Öresund....
, Switzerland
Switzerland
Switzerland name of one of the Swiss cantons. ; ; ; or ), in its full name the Swiss Confederation , is a federal republic consisting of 26 cantons, with Bern as the seat of the federal authorities. The country is situated in Western Europe,Or Central Europe depending on the definition....
, Turkey
Turkey
Turkey , known officially as the Republic of Turkey , is a Eurasian country located in Western Asia and in East Thrace in Southeastern Europe...
and Ukraine
Ukraine
Ukraine is a country in Eastern Europe. It has an area of 603,628 km², making it the second largest contiguous country on the European continent, after Russia...
. Although horse-chestnut occurs naturally mostly above the 700–1000 m. contours the moth does well in well-watered places such as parks in cities and at low elevation but not well in the hotter parts of Europe e.g. Spain
Spain
Spain , officially the Kingdom of Spain languages]] under the European Charter for Regional or Minority Languages. In each of these, Spain's official name is as follows:;;;;;;), is a country and member state of the European Union located in southwestern Europe on the Iberian Peninsula...
. Probably aided greatly by vehicular transport, the moth has attained a very rapid dispersal rate across Europe of 60 km. per year.
Origin and epidemiology
Cameraria ohridella was first noticed from outbreaks near Ohrid Lake, MacedoniaMacedonia (Greece)
Macedonia is a geographical and historical region of Greece in Southern Europe. Macedonia is the largest and second most populous Greek region...
in 1984, and was described as a new species by Deschka and Dimić in 1986. A likely Balkan origin for this moth was evidenced from a decrease in genetic diversity from natural towards artificial horse-chestnut stands that were planted around Europe since around 1600. This Balkan origin is further documented by numerous herbarium samples that date back to 1879. These include an outbreak which occurred in horse-chestnut specimens collected by F.K. Meyer in 1961 in Albania. Of the 30 known mitochondrial haplotype
Haplotype
A haplotype in genetics is a combination of alleles at adjacent locations on the chromosome that are transmitted together...
s for the species only three (known as A, B and C) have invaded the rest of Europe since 1989, and only A is dominant. It is likely that the frequency of haplotype A has been increasing even in Balkan natural sites, aided by the late development of roads in the region.
Host plants
As well as colonising the leaves of the common horse-chestnut, C. ohridella is also able to feed on Aesculus pavia, Acer platanoides and Acer pseudoplatanus, on which in particular one mitochondrial race, haplotype B, seems to develop successfully when nearby horse-chestnut leaves are exhausted, but is not thought to pose such a strong risk to these species unlike to the common horse-chestnut.Parasitoids
Over 60 generalist parasitoids have been recorded. However, for biological control a highly specialist parasitoid still needs to be found.Predators
A number of natural predators of the larval stages of C. ohridella have been recorded. Observations have shown that blue tits (Parus caeruleus), great tits (Parus major) and marsh tits (Parus palustris) feed on the larvae. Between them, three tit species are thought to predate between 2 and 4% of the larvae. The southern oak bushcricket (Meconema meridionaleMeconema meridionale
Meconema meridionale is an insect in the family Tettigoniidae, commonly known as the southern oak bush cricket. It is carnivorous, arboreal and nocturnal. It has a more southerly distribution than its relative, Meconema thalassinum, being found in southern France, south-western Germany, Italy,...
) has also been found to predate C. ohridella, consuming around 10 larvae per day. Overall the predation by the southern oak bushcricket is insignificant compared to that by birds however. Experiments with the predatory mite
Mite
Mites, along with ticks, are small arthropods belonging to the subclass Acari and the class Arachnida. The scientific discipline devoted to the study of ticks and mites is called acarology.-Diversity and systematics:...
Euseius finlandicus, bush crickets (Phaneroptera
Phaneroptera
Phaneroptera is an Old World genus of katydids in the sub-family Phaneropterinae.It contains the following species: *Phaneroptera acaciae Chopard, 1954*Phaneroptera adusta *Phaneroptera albida Walker, F., 1869...
sp.), ladybirds and lacewings found that none predate C. ohridella.
Procedures
Inadvisably, trees can be removed, or better, leaves cleared and burned before adult emergence by the end of March. Use of the systemic insecticide imidaclopridImidacloprid
Imidacloprid is a nicotine-based, systemic insecticide, which acts as a neurotoxin and belongs to a class of chemicals called the neonicotinoids. Although it is now off patent, the primary manufacturer of this chemical is Bayer CropScience,...
is usually banned as it kills bees. Fenoxycarb
Fenoxycarb
Fenoxycarb is a carbamate insect growth regulator. It has a low toxicity for bees, birds, and humans, but is toxic to fish. The oral LD50 for rats is greater than 16,800 mg/kg....
causes up to 100% pupal mortality, has low environmental toxicity and can be combined successfully with manual leaf removal. A synthetic pheromone can be used to trap males, but effective control may be hard to thus achieve. In any case, infestation levels could diminish over time as Cameraria ohridella starts to recruit generalist members of the local parasitoid wasp community.
Projects
A number of projects have been launched to investigate the biology and biological control of Cameraria ohridella and its impact since 2001, for example, an EUEuropean Union
The European Union is an economic and political union of 27 independent member states which are located primarily in Europe. The EU traces its origins from the European Coal and Steel Community and the European Economic Community , formed by six countries in 1958...
-wide multidisciplinary project, CONTROCAM ("Control of Cameraria") and the HAM-CAM Project.
External links
- Taxon page for Cameraria ohridella Deschka & Dimic 1986. In: EOLspecies in English
- HAMburger-CAMeraria-Projekt - Films Photos incl. REM in German
- Cameraria Homepage in German and English
- Czech Academy of Sciences Cameraria homepage in English
- Forestry Commission Research page in English
- Mactode Publications - Educational Resources on CD/DVD in English
- HAMburger-CAMeraria-Projekt in German
- Abstract of HAM-CAM-Projekt, 2003
- BBC News report on infestation in Leicester, England, September 2010