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''Cryptococcus fagisuga'', commonly known as the beech scale or woolly beech scale, is a
felted A felted material is a hairy or Wikt:filamentous, filamentous (hairy-like) fibre that is densely packed or tangled, forming felt or felt-like structures. Advantages of felted tissue The dermis is described in Gray's Anatomy as "felted connect ...
scale insect Scale insects are small insects of the Order (biology), order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient g ...
in the superfamily
Coccoidea Scale insects are small insects of the Order (biology), order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient g ...
that infests beech trees of the genus ''
Fagus Beech (genus ''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to subtropical (accessory forest element) and temperate (as dominant element of Mesophyte, mesophytic forests) Eurasia and North America. There are 14 accepted ...
''. It is associated with the transmission of
beech bark disease Beech bark disease is a disease that causes mortality and defects in beech trees in the eastern United States, Canada and Europe.Tainter, Frank H., and Fred A. Baker . Principles of Forest Pathology. New York, NY: John Wiley & Sons, Inc., 1996.Sin ...
because the puncture holes it makes in the
bark Bark may refer to: Common meanings * Bark (botany), an outer layer of a woody plant such as a tree or stick * Bark (sound), a vocalization of some animals (which is commonly the dog) Arts and entertainment * ''Bark'' (Jefferson Airplane album), ...
allow entry of
pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...
ic
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
which have been identified as '' Nectria coccinea var. faginata'' and sometimes '' Nectria galligena''.U.S. Department of Agriculture, Forest Service
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History and distribution

''Cryptococcus fagisuga'' aka Beech bark disease has been recorded as affecting common beech trees, ''
Fagus sylvatica ''Fagus sylvatica'', the European beech or common beech, is a large, graceful deciduous tree in the Fagaceae, beech family with smooth silvery-gray bark, large leaf area, and a short trunk with low branches. Description ''Fagus sylvatica'' i ...
'', in
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
since before 1849. Until 1914 it was thought that the beech scale insect itself was responsible for the disease. Subsequently it was discovered that a fungus, then identified as '' Neonectria ditissima'', was in fact killing the trees infested by the scale. Around 1890 the scale insect was accidentally introduced into
Nova Scotia Nova Scotia is a Provinces and territories of Canada, province of Canada, located on its east coast. It is one of the three Maritime Canada, Maritime provinces and Population of Canada by province and territory, most populous province in Atlan ...
. By 1932, the scale and its associated ''Neonectria'' fungus had spread to many areas of the
Maritime Provinces The Maritimes, also called the Maritime provinces, is a region of Eastern Canada consisting of three provinces: New Brunswick, Nova Scotia, and Prince Edward Island. The Maritimes had a population of 1,899,324 in 2021, which makes up 5.1% of ...
and parts of eastern and south central
Maine Maine ( ) is a U.S. state, state in the New England region of the United States, and the northeasternmost state in the Contiguous United States. It borders New Hampshire to the west, the Gulf of Maine to the southeast, and the Provinces and ...
. It continues to spread in North America and is now found in
Quebec Quebec is Canada's List of Canadian provinces and territories by area, largest province by area. Located in Central Canada, the province shares borders with the provinces of Ontario to the west, Newfoundland and Labrador to the northeast, ...
,
Ontario Ontario is the southernmost Provinces and territories of Canada, province of Canada. Located in Central Canada, Ontario is the Population of Canada by province and territory, country's most populous province. As of the 2021 Canadian census, it ...
,
New England New England is a region consisting of six states in the Northeastern United States: Connecticut, Maine, Massachusetts, New Hampshire, Rhode Island, and Vermont. It is bordered by the state of New York (state), New York to the west and by the ...
,
New York New York most commonly refers to: * New York (state), a state in the northeastern United States * New York City, the most populous city in the United States, located in the state of New York New York may also refer to: Places United Kingdom * ...
,
New Jersey New Jersey is a U.S. state, state located in both the Mid-Atlantic States, Mid-Atlantic and Northeastern United States, Northeastern regions of the United States. Located at the geographic hub of the urban area, heavily urbanized Northeas ...
,
Pennsylvania Pennsylvania, officially the Commonwealth of Pennsylvania, is a U.S. state, state spanning the Mid-Atlantic (United States), Mid-Atlantic, Northeastern United States, Northeastern, Appalachian, and Great Lakes region, Great Lakes regions o ...
and
West Virginia West Virginia is a mountainous U.S. state, state in the Southern United States, Southern and Mid-Atlantic (United States), Mid-Atlantic regions of the United States.The United States Census Bureau, Census Bureau and the Association of American ...
. Attempts are being made to discover the geographic origin of beech scale in order to try to identify any natural enemies that might hold promise for its
biological control Biological control or biocontrol is a method of controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or o ...
. From these studies and associated
ribosomal DNA The ribosomal DNA (rDNA) consists of a group of ribosomal RNA encoding genes and related regulatory elements, and is widespread in similar configuration in all domains of life. The ribosomal DNA encodes the non-coding ribosomal RNA, integral struc ...
analysis, it seems likely that the pest originated in the region of northeast Greece, northern Iran, the Caucasus and the Black Sea drainage basin on the host beech subspecies ''F. sylvatica orientalis''.


Description

Adult scales are elliptical and about one millimetre long and are covered by a cream coloured, cottony wax secretion. They have reddish-brown eyes, no wings, rudimentary antennae and legs, and numerous minute wax-secreting glands. The stylet through which they suck sap can be up to two millimetres long.


Life cycle

There are no male beech scale insects and the females reproduce by
parthenogenesis Parthenogenesis (; from the Greek + ) is a natural form of asexual reproduction in which the embryo develops directly from an egg without need for fertilization. In animals, parthenogenesis means the development of an embryo from an unfertiliz ...
. From midsummer onwards, they deposit four to eight pale yellow eggs, attached to each other in strings end to end, on the bark of the trees where they have been feeding. The wingless larvae that hatch out about twenty days later are known as crawlers or
nymphs A nymph (; ; sometimes spelled nymphe) is a minor female nature deity in ancient Greek folklore. Distinct from other Greek goddesses, nymphs are generally regarded as personifications of nature; they are typically tied to a specific place, land ...
. They have well-developed legs and short antennae. Some remain concealed under the female, which dies after the eggs are laid, and others disperse to cracks and crevices on the tree. A few get washed down or fall to the ground and most of these perish. Occasionally one may find its way to another beech tree, perhaps wafted there by the wind or on the foot of a bird, and founds a new colony. Having found a location on a suitable tree, the crawler forces its tubular stylet into the bark and starts to feed. At this stage, it
moult In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is a process by which an animal casts off parts of its body to serve some beneficial purpose, either at ...
s and becomes a second-stage, legless nymph, and will remain sedentary for the rest of its life. It secretes wax from glands and is soon covered in a protective coating of wool-like material. After overwintering it completes a second moult in the spring to become a mature female.


Biology

The beech scale is monophagous, being found only on beech trees, and sucks sap from the parenchyme tissue of the bark. The small wounds produced when it feeds allow the ''Nectria'' fungi to invade infested trees. The insect colonises beech trees that are aged at least thirty years. Studies have shown that younger trees produce defensive chemicals which deter infestation. A ladybird beetle '' Chilocorus stigma'' feeds on this scale and is common throughout most of the Eastern United States, but this
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
does not reduce scale populations sufficiently to control infestations. Persistent severely cold weather may kill beech scale and air temperatures of -37 °C have been shown to be lethal to insects not protected by snow. Although trees are weakened when supporting scale colonies, this does not usually cause mortality, which only occurs after the trees have been invaded by the ''Nectria'' fungi. It usually takes three to six years of infestation by the scale insect before the fungus reaches critical levels. Where beech bark disease becomes established, most of the larger trees will die. Some trees seem to be partially resistant to the disease and a small number seem to be completely resistant. This may be partly due to the fact that trees with smooth bark provide fewer cracks and crevices in which the scale insect can flourish.Forest Encyclopedia Network
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References

{{Taxonbar, from=Q998347 Insect vectors of plant pathogens Hemiptera of North America Insects described in 1936 Eriococcidae