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The white-faced darter or small whiteface (''Leucorrhinia dubia'') is a dragonfly belonging to the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
'' Leucorrhinia'' in the family Libellulidae, characterised by red and black markings and a distinctive white patch on the head. It is found in
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The p ...
s and
peat bog A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main types of wetlands. Other names for bogs include mire, mosses, quagmire, and muskeg; a ...
s from northern
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 subcontinent of Eurasia and it is located entirel ...
eastwards to
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive region, geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a ...
, and the adults are active from around April till September, which is known as the "flight period". It breeds in acidic bodies of water, laying its eggs in clumps of
sphagnum moss ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, peat moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store w ...
that provide a safe
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
for larval development. The
larva A larva (; plural larvae ) 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 particularly vulnerable to
predation 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 ...
by fish, and so are usually found in lakes where fish are not present. ''L. dubia'' is listed as a species of
least concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
(LC) by the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
, however, it is potentially threatened by
habitat destruction Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
, pollution, and climate change.


Description

This species has a predominantly black body, usually in length - the
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
is long and the hindwing is long. Mature males have striking red and orange markings on the
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
and
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
which become darker with age, whereas young males and females have pale yellow markings. All individuals have a conspicuous white
frons Insect morphology is the study and description of the physical form of insects. The terminology used to describe insects is similar to that used for other arthropods due to their shared evolutionary history. Three physical features separate insec ...
at the front of the head. The wings have a brown patch at the base and noticeable black pterostigma. It may be confused with similar species, including the black darter (''Sympetrum danae''), ruddy darter (''Sympetrum sanguineum'') and common darter (''Sympetreum striolatum''), however is usually distinguishable by its white face patch.


Distribution

This dragonfly's range extends from western
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 subcontinent of Eurasia and it is located entirel ...
to Japan, and it is commonly found in western, northern and eastern Europe at higher altitudes, but is rare in southern Europe and the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the European mainland, continental mainland. It comprises England, Scotlan ...
. In recent decades, this species has been observed as far afield as
Russia Russia (, , ), or the Russian Federation, is a transcontinental country spanning Eastern Europe and Northern Asia. It is the largest country in the world, with its internationally recognised territory covering , and encompassing one-eig ...
and China, however the vast majority of sightings have occurred in central and northern Europe. In Britain, the majority of individuals of this species are found in the highlands of
Scotland Scotland (, ) is a Countries of the United Kingdom, country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a Anglo-Scottish border, border with England to the southeast ...
, with key populations located in
Inverness-shire Inverness-shire ( gd, Siorrachd Inbhir Nis) is a historic county, registration county and lieutenancy area of Scotland. Covering much of the Highlands and Outer Hebrides, it is Scotland's largest county, though one of the smallest in popula ...
and
Ross-shire Ross-shire (; gd, Siorrachd Rois) is a historic county in the Scottish Highlands. The county borders Sutherland to the north and Inverness-shire to the south, as well as having a complex border with Cromartyshire – a county consisting o ...
. In England, a few isolated populations occur in Cheshire and
Cumbria Cumbria ( ) is a ceremonial and non-metropolitan county in North West England, bordering Scotland. The county and Cumbria County Council, its local government, came into existence in 1974 after the passage of the Local Government Act 1972. C ...
, where biodiversity action plans have been set up to protect them, and as far south as Chartley Moss National Nature Reserve in Staffordshire. However, when British populations are considered overall, this species has been in decline over the past 35 years.


Habitat

Adult individuals of ''L. dubia'' can utilise scrub and
woodland A woodland () is, in the broad sense, land covered with trees, or in a narrow sense, synonymous with wood (or in the U.S., the ''plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunlight and limited shade (se ...
habitat for roosting and foraging. The larvae require terrestrial areas of water, such as
marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found a ...
es,
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The p ...
s and
peat bogs A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main types of wetlands. Other names for bogs include mire, mosses, quagmire, and muskeg ...
, that generally have
vegetation Vegetation is an assemblage of plant species and the ground cover they provide. It is a general term, without specific reference to particular taxa, life forms, structure, spatial extent, or any other specific botanical or geographic characte ...
growing at the water's edge. Peat bogs form a particularly important habitat, since they provide acidic conditions necessary for the growth of
sphagnum moss ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, peat moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store w ...
, which provides a source of food and shelter.


Life history


Larval stage

Like all other species of dragonfly, the larvae of this species grow by moulting (in which the
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
is shed) several times during development. In the early stages of development,
larva A larva (; plural larvae ) 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 preferentially inhabit
sphagnum moss ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, peat moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store w ...
, which may provide a reliable source of food because the moss can trap organic matter that would otherwise sink to the bottom of the water. As well as fish, many waterfowl, amphibians, and other
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s will prey upon dragonfly larvae, so the mats likely also provide a safe hiding place from
predators 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 ...
. The larvae are able to change colour depending on whether the moss they are inhabiting is brown or green, which would help to visually disguise them from predators. The acidification of ponds and lakes due to airborne
pollution Pollution is the introduction of contaminants into the natural environment that cause adverse change. Pollution can take the form of any substance (solid, liquid, or gas) or energy (such as radioactivity, heat, sound, or light). Pollutants, the ...
may in fact benefit ''L. dubia'' and other species that prefer acidic conditions, as this can result in a greater occurrence of sphagnum moss. Larvae will actively forage throughout both the day and night, however have been shown to capture more
prey 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 ...
during the night. Larvae will also cannibalise their
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organis ...
s, particularly in the absence of an abundant food source (
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
), and can be threatened by predation from other species of dragonflies. Larger individuals will generally cannibalise smaller individuals, and it is believed that this functions as a way of controlling
population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction using a ...
numbers. The larvae generally do not thrive in environments where fish predators are present, and in some case appear to be more vulnerable to predation compared to other species of dragonflies, possibly due to their active foraging behaviour. An active foraging strategy means the larvae purposely swim to different parts of their habitat in search of prey, as opposed to a passive “sit and wait” strategy exhibited by some other species of dragonfly. They rely primarily upon visual cues to locate prey, and will consume anything that isn’t too large or powerful for them to handle. They are therefore more likely to occur in areas of water where fish are absent. Since fish are less likely to occur in relatively smaller bodies of water, since these are at greater risk of becoming
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as ...
deficient during the winter, it has been suggested that adult dragonflies may choose where to lay their eggs by judging the size of a lake. The larvae have, however, been shown to exhibit
phenotypic plasticity Phenotypic plasticity refers to some of the changes in an organism's behavior, morphology and physiology in response to a unique environment. Fundamental to the way in which organisms cope with environmental variation, phenotypic plasticity encompa ...
in the presence of predators, which means that larvae in danger of predation are able to grow longer dorsal and lateral spines as a
defence mechanism In psychoanalytic theory, a defence mechanism (American English: defense mechanism), is an unconscious psychological operation that functions to protect a person from anxiety-producing thoughts and feelings related to internal conflicts and o ...
.


Reproductive stage

Individuals of this species typically take 1–3 years to reach adulthood. The adults breed in acidic pools where
sphagnum ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, peat moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store wa ...
moss is present. The male holds a small
territory A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or a ...
near a body of water, and
copulation Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetra ...
with the female often begins over the water before they settle on the ground for about 30 minutes. The female drops the
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
amongst submerged moss or stems of
cottongrass ''Eriophorum'' (cottongrass, cotton-grass or cottonsedge) is a genus of flowering plants in the family Cyperaceae, the sedge family. They are found throughout the arctic, subarctic, and temperate portions of the Northern Hemisphere in acid bo ...
that grow along the edge of the water. Adult dragonflies emerge between May and early July in Great Britain; the exact timing depends on the
latitude In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north pol ...
and weather. When the larvae have developed sufficiently and are ready to emerge as adults, they climb out of the water up a plant stem and shed their exoskeleton one final time. Males become
sexually mature Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definiti ...
4–12 days after emergence, and females a few days later; can generally be seen during their "flight period" from April till September.


Conservation


Status

''L. dubia'' is listed as a species of
least concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
(LC) by the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
, meaning that it is not currently considered to be threatened by
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
. This categorisation applies to the species when all populations are considered across its entire range. However, it is
red-listed The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biologi ...
in a number of European countries within its range, including
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
,
Austria Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...
, Switzerland and the UK. Greater research and monitoring is currently required on population distribution and trends, to better understand how this species will fare in the future. Most of the recent scientific research has been conducted in central and northern Europe, and less is known about eastern populations in Siberia, where ''L. dubia'' is rarely seen.


Threats

''L. dubia'' is vulnerable to alteration and destruction of its
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
, primarily due to water pollution, industrialisation, and development for agriculture. It is protected by the
Wildlife and Countryside Act 1981 The Wildlife and Countryside Act 1981 is an Act of Parliament in the United Kingdom implemented to comply with European Council Directive 79/409/EEC on the conservation of wild birds. In short, the act gives protection to native species (especia ...
in the United Kingdom, where 95% of lowland
peat bogs A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main types of wetlands. Other names for bogs include mire, mosses, quagmire, and muskeg ...
have been destroyed, and it is also covered by Biodiversity Action Plans in some British counties. Conservation reintroduction programmes have been shown to be a successful method for restoring populations to suitable sites, and in 2010 it was reintroduced to
Witherslack Mosses Witherslack Mosses are a fragmented wetland west of the Kent estuary in Cumbria, England, within the Lake District National Park. They consist of three raised bogs, the remnants of a formerly extensive, estuarine bog, which have been protected ...
in Cumbria, after 13 years of restoration management. Isolated, fragmented populations are at an increased risk of local extinction due to a lack of exchange of genetic variation, resulting in potential
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...
and increased impact of sudden, random events such as disease outbreaks. Populations of ''L. dubia'' in the UK, the
Pyrenees The Pyrenees (; es, Pirineos ; french: Pyrénées ; ca, Pirineu ; eu, Pirinioak ; oc, Pirenèus ; an, Pirineus) is a mountain range straddling the border of France and Spain. It extends nearly from its union with the Cantabrian Mountains to ...
and the
Alps The Alps () ; german: Alpen ; it, Alpi ; rm, Alps ; sl, Alpe . are the highest and most extensive mountain range system that lies entirely in Europe, stretching approximately across seven Alpine countries (from west to east): France, Swi ...
have been found to be more genetically distinct than other European populations, and may therefore be considered priorities for conservation, as they are likely to retain important genetic variation for adapting to changing climate conditions. A rise in average annual temperatures caused by global warming could negatively impact ''L. dubia'' by allowing invasive species such as the scarlet dragonfly ''(Crocothemis erythraea)'' to occupy habitats further north of their natural range. Although the growth rate of ''L. dubia'' doesn’t appear to be affected by temperature, ''C. erythraea'' can grow faster at higher temperatures, causing ''L. dubia'' to suffer higher levels of
competition Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indiv ...
and mortality. Due to the particular sensitivity of the larval stage to the presence of fish, this species could be threatened by the practice of artificially stocking lakes with fish for the purpose of recreational fishing. The process of liming acidified lakes, to increase their pH and restore fish populations, also poses a threat by reducing the amount of sphagnum moss present. Acidic lakes where fish are naturally absent could be of high conservation value for this species.


References


External links


NBN Gateway: map of UK records of white-faced darterBiopix: ''Leucorrhinia dubia''Map of the global distribution of ''L. dubia''
{{Taxonbar, from=Q1746499 Libellulidae Dragonflies of Europe Insects described in 1825