Coccus Fagi
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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 filamentous (hairy-like) fibre that is densely packed or tangled, forming felt or felt-like structures. Anatomy and zoology The dermis is described in Gray's Anatomy as "felted connective tissue, with a vary ...
scale insect Scale insects are small insects of the order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient grouping than the ...
in the
superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
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 (''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to temperate Europe, Asia, and North America. Recent classifications recognize 10 to 13 species in two distinct subgenera, ''Engleriana'' and ''Fagus''. The ''Engle ...
''. 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.Houston, David R., and James T. O'Brien. "Beech Bark Disease." Forest Insect and Disease Leaflet 75(1998) 1-7. 17 Mar 2 ...
because the puncture holes it makes in the
bark Bark may refer to: * 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) Places * Bark, Germany * Bark, Warmian-Masurian Voivodeship, Poland Arts, ...
allow entry of
pathogen In biology, a pathogen ( el, πάθος, "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 ...
ic
fungi A fungus ( : fungi or funguses) is any member of the 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, separately from ...
which have been identified as '' Nectria coccinea var. faginata'' and sometimes ''
Nectria galligena ''Neonectria ditissima'' (syn. ''Neonectria galligena'') is a fungal plant pathogen. It causes cankers that can kill branches of trees by choking them off. Apple and beech trees are two susceptible species. Host range ''Neonectria ditissima ...
''.U.S. Department of Agriculture, Forest Service
/ref>


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 deciduous tree belonging to the beech family Fagaceae. Description ''Fagus sylvatica'' is a large tree, capable of reaching heights of up to tall and trunk diameter, though mor ...
'', in
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
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 ''Neonectria ditissima'' (syn. ''Neonectria galligena'') is a fungal plant pathogen. It causes cankers that can kill branches of trees by choking them off. Apple and beech trees are two susceptible species. Host range ''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 one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. Nova Scotia is Latin for "New Scotland". Most of the population are native Eng ...
. 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 Ca ...
and parts of eastern and south central
Maine Maine () is a state in the New England and Northeastern regions of the United States. It borders New Hampshire to the west, the Gulf of Maine to the southeast, and the Canadian provinces of New Brunswick and Quebec to the northeast and north ...
. It continues to spread in North America and is now found in
Quebec Quebec ( ; )According to the Canadian government, ''Québec'' (with the acute accent) is the official name in Canadian French and ''Quebec'' (without the accent) is the province's official name in Canadian English is one of the thirtee ...
,
Ontario Ontario ( ; ) is one of the thirteen provinces and territories of Canada.Ontario is located in the geographic eastern half of Canada, but it has historically and politically been considered to be part of Central Canada. Located in Central Ca ...
,
New England New England is a region comprising 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 to the west and by the Canadian provinces ...
,
New York New York most commonly refers to: * New York City, the most populous city in the United States, located in the state of New York * New York (state), a state in the northeastern United States New York may also refer to: Film and television * '' ...
,
New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north and east by the state of New York; on the east, southeast, and south by the Atlantic Ocean; on the west by the Delaware ...
,
Pennsylvania Pennsylvania (; ( Pennsylvania Dutch: )), officially the Commonwealth of Pennsylvania, is a state spanning the Mid-Atlantic, Northeastern, Appalachian, and Great Lakes regions of the United States. It borders Delaware to its southeast, ...
and
West Virginia West Virginia is a state in the Appalachian, Mid-Atlantic and Southeastern regions of the United States.The Census Bureau and the Association of American Geographers classify West Virginia as part of the Southern United States while the Bur ...
. 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, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also invo ...
. From these studies and associated
ribosomal DNA Ribosomal DNA (rDNA) is a DNA sequence that codes for ribosomal RNA. These sequences regulate transcription initiation and amplification, and contain both transcribed and non-transcribed spacer segments. In the human genome there are 5 chromos ...
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

Beech scale nymph There are no male beech scale insects and the females reproduce by
parthenogenesis Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and development ...
. 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 ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are typ ...
. 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 the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
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 A generalist species is able to thrive in a wide variety of environmental conditions and can make use of a variety of different resources (for example, a heterotroph with a varied diet). A specialist species can thrive only in a narrow range of env ...
, 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 ''Chilocorus stigma'', commonly known as the twice-stabbed ladybug, is a native resident of the United States and Canada but does not live west of the Sierra Nevada.. It also has been introduced to Hawaii. It is shiny black, and there is one red ...
'' feeds on this scale and is common throughout most of the Eastern United States, but this
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
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