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''Boloria selene'', known in Europe as the small pearl-bordered fritillary and in North America as the silver-bordered fritillary, is a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...
of the family Nymphalidae. It is found across
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 ...
,
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
and
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
, and feeds exclusively on
violets Violet identifies various plant taxa, particularly species in the genus ''Viola'', within which the common violet is the best known member in Eurasia and the common blue violet and common purple violet are the best known members in North America ...
in its
larval 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. The l ...
stages.Silver-bordered Fritillary
CBIF
This species prefers wet grassland habitats, where its larval food source, violets, are found. It
overwinters Overwintering is the process by which some organisms pass through or wait out the winter season, or pass through that period of the year when "winter" conditions (cold or sub-zero temperatures, ice, snow, limited food supplies) make normal activi ...
in its larval stage, and eggs hatch in the late summer to early autumn. Members of this species are prey for multiple types of birds and other insects. Due to modern agriculture, most of the grassland habitats that sustain ''Boloria selene'' are fragmented or lost all together in favor of farmland. Because of this, the small pearl-bordered fritillary has seen a serious drop in population across Europe, in some places as much as 80%. Factors including limited habitat range, low dispersal rate, and strong food specialization also contribute to population loss. Despite modern conservation efforts, the number of small pearl-bordered fritillaries is still declining. The North American populations appear to be affected in the same way, at least in the continental United States.


Appearance

The small pearl-bordered fritillary is similar to the pearl-bordered fritillary but has black chevrons on the edge of its wings, a large central black dot on each wing, and white pearls on the underside. Males tend to be smaller than females, with a wingspan of 35 to 41 mm. Females are 38 to 44 mm in length. Though the small pearl-bordered fritillary is similar to the pearl-bordered fritillary it is much brighter in color because the pearl-bordered fritillary emerges from its pupa earlier. Seitz - A. ''selene'' Schiff. (= ''euphrosyne'' Bgstr., ''euphrasia'' Lew., ''silene'' Haw.) (67g). Forewing above and beneath very similar to tliat of the preceding 'selenis'' Hindwing beneath with the median band distinct but not broad, the costal spot of the band, the large tooth above the apex of the cell and the one below the cell-end being vividly silvery like the marginal lunules; the broad interspace between the median band and the marginal lunules is leatheryellow and bears below the apex and above the anal angle two large dark cinnamon clouds; obsolescent silvery selenia bands extend from the costal and abdominal margins towards the centre of the wing. File:Petit collier argenté MHNT CUT 2013 3 21 Sornac Dos.jpg, Dorsal side File:Petit collier argenté MHNT CUT 2013 3 21 Sornac Ventre.jpg, Ventral side


Distribution

The species is widespread across central and 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 Continent#Subcontinents, subcontinent of Eurasia ...
,
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
, and through
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
to
Korea Korea ( ko, 한국, or , ) is a peninsular region in East Asia. Since 1945, it has been divided at or near the 38th parallel, with North Korea (Democratic People's Republic of Korea) comprising its northern half and South Korea (Republic o ...
. Specifically, it can be found in places like
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 ...
, 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 continental mainland. It comprises England, Scotland, Wales and North ...
, and
Sweden Sweden, formally the Kingdom of Sweden,The United Nations Group of Experts on Geographical Names states that the country's formal name is the Kingdom of SwedenUNGEGN World Geographical Names, Sweden./ref> is a Nordic country located on ...
and in the Midwestern United States in places like
Iowa Iowa () is a state in the Midwestern region of the United States, bordered by the Mississippi River to the east and the Missouri River and Big Sioux River to the west. It is bordered by six states: Wisconsin to the northeast, Illinois to the ...
and the
Dakotas The Dakotas is a collective term for the U.S. states of North Dakota and South Dakota. It has been used historically to describe the Dakota Territory, and is still used for the collective heritage, culture, geography, fauna, sociology, econom ...
. In the United Kingdom, ''Boloria selene'' is widespread across upland and western Britain, but is not found in central, eastern England or Ireland. Its range appears to be stable through much of Europe but declines have been reported in at least nine countries, such as England. This species is found across Europe and North America in grassland environments where native violets grow. It occurs in damp, grassy habitats, woodland clearings and
moorland Moorland or moor is a type of habitat found in upland areas in temperate grasslands, savannas, and shrublands and montane grasslands and shrublands biomes, characterised by low-growing vegetation on acidic soils. Moorland, nowadays, generally ...
, but has also been found in
dune A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, f ...
slacks and coastal cliffs.


Life cycle


Larva

The larvae of this species hatch in the late summer to early fall. They feed exclusively on violets and are active mostly at night. In drier areas, dog violets are used while in wetter areas the species feeds on bog violets (''
Viola palustris ''Viola palustris'' (marsh violet, or alpine marsh violet) is a perennial forb of the genus ''Viola''. It inhabits moist meadows, marshes, and stream banks in northern parts of North America and Eurasia. The species epithet ''palustris'' is Latin ...
'') or
marsh violet ''Viola palustris'' (marsh violet, or alpine marsh violet) is a perennial forb of the genus ''Viola''. It inhabits moist meadows, marshes, and stream banks in northern parts of North America and Eurasia. The species epithet ''palustris'' is Lat ...
s growing among
purple moor-grass ''Molinia caerulea'', known by the common name purple moor-grass, is a species of grass that is native to Europe, west Asia, and north Africa. It grows in locations from the lowlands up to in the Alps. Like most grasses, it grows best in acid so ...
or
tufted hair-grass ''Deschampsia cespitosa'', commonly known as tufted hairgrass or tussock grass, is a perennial tufted plant in the grass family Poaceae. Distribution of this species is widespread including the eastern and western coasts of North America, parts ...
. They prefer the damper areas. They also occur among
bracken Bracken (''Pteridium'') is a genus of large, coarse ferns in the family Dennstaedtiaceae. Ferns (Pteridophyta) are vascular plants that have alternating generations, large plants that produce spores and small plants that produce sex cells (eggs ...
, which provides shade for the appropriate violet species. The caterpillars overwinter by hibernating, and then reemerge in the spring to finish growth and pupate. However, recent studies have shown that times of laying, hatching, and reemerging vary by temperature, and all life cycle stages appear to be strongly linked to the timing of the seasons. This effect is a suspected form of plastic behavior seen in many species of insects, especially ones in temperate climates that experience large temperature changes due to a change in seasons.


Pupa

Caterpillars pupate between mid May and August. When the larvae forms the pupa, or
chrysalis A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
, it is formed with its head down. This transformative stage lasts about two to three weeks.


Adult

Adults mate and lay eggs in the spring on or in the near vicinity of violets. A common violet selected for oviposition is the
Marsh Violet ''Viola palustris'' (marsh violet, or alpine marsh violet) is a perennial forb of the genus ''Viola''. It inhabits moist meadows, marshes, and stream banks in northern parts of North America and Eurasia. The species epithet ''palustris'' is Lat ...
. Adult ''Boloria selene'' have been observed to fly in a brood once a year sometime in the summer. Adult small pearl-bordered fritillaries feed on the following plants: * Bramble *
Thistle Thistle is the common name of a group of flowering plants characterised by leaves with sharp prickles on the margins, mostly in the family Asteraceae. Prickles can also occur all over the planton the stem and on the flat parts of the leaves. ...
* Bugle (''
Ajuga reptans ''Ajuga reptans'' is commonly known as bugle, blue bugle, bugleherb, bugleweed, carpetweed, carpet bugleweed, and common bugle, and traditionally but less commonly as St. Lawrence plant. It is an herbaceous flowering plant in the mint family Lamia ...
'')


Migration habits

This species is well known to migrate short distances during its reproductive stage, but does not appear to make any long distance migrations that cross over unsuitable habitats such as farmland and urban areas.


Predators

Small birds appear to be major predators of both larval and adult stages, including introduced ground birds such as pheasants that were introduced in the UK for sport hunting (in quantities of almost 40 million birds according to a study done on ground birds' impact on local wildlife). However, no major decline was detected from the introduced predators alone, as there was no significance between reduced adult emergence and increased ground birds. In a study of ambush bugs, ''Boloria selene'' was identified as a prey, although predation rates were not studied.


Genetics and local adaptation

A study using North American populations found that sibling mating events of this species almost always result in unviable offspring. Even hybrids that were crossed with individuals from parent populations often had unviable offspring. This indicates that this species requires non-kin mating to survive, which may help with decreasing inbreeding events. Certain larvae did better in their home environments and struggled in foreign environments, indicating potential
local adaptation Local adaptation is a mechanism in evolutionary biology whereby a population of organisms evolves to be more well-suited to its local environment than other members of the same species that live elsewhere. Local adaptation requires that different po ...
. There were also slight coloration changes between males in
Massachusetts Massachusetts (Massachusett language, Massachusett: ''Muhsachuweesut assachusett writing systems, məhswatʃəwiːsət'' English: , ), officially the Commonwealth of Massachusetts, is the most populous U.S. state, state in the New England ...
(a “warm leathery tone”) and males in South Dakota (a “chestnut brown” color), indicating potential genetic differences between the two populations, which are separated by unsuitable habitat and by a distance of 2000 miles (3218.688 km). However, due to the age of the study, more work is needed to confirm that local adaptation is indeed taking place in North American populations of small pearl-bordered fritillaries instead of individual plasticity. Another study found ''Boloria selene'' are quite plastic in their response to temperature variation, and will adjust much of their life cycle in response to temperature changes such as an earlier warming that comes with an early spring. The same study also analyzed among and between population data, and suggested that these animals quickly capitalize on early warming and late cooling regardless of location (in the US at least), and seem to also know when to wait longer or shorter before beginning reproduction.


Conservation

There appears to be no place (at least in Europe) that is not suffering from at least light population degradation. Many studies in the UK (where population loss is most severe) and several studies from other areas of Europe state that the small pearl-bordered fritillary is rapidly declining in numbers. This is likely from indirect harm due to
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 ...
and
fragmentation Fragmentation or fragmented may refer to: Computers * Fragmentation (computing), a phenomenon of computer storage * File system fragmentation, the tendency of a file system to lay out the contents of files non-continuously * Fragmented distributi ...
. Since this species is primarily attached to violet that grows in wild grasslands, modern agricultural practices which have caused most of Europe's grasslands to become fragmented or destroyed for farmland have a severe effect on the butterfly. Without native violets to host their larvae, this species is facing decline across Europe. This species also struggles with breeding across habitat fragments as it must lay eggs exclusively on violets in order to have any viable offspring. However, adults seem to be quite flexible in their movement patterns, with one study stating that only 45% of released ''Boloria selene'' specimens were able to be found and recaptured. This indicates that farmland itself appears to more strongly damage reproduction rates instead of general adult survival. Another study found that this species has another characteristic which makes it vulnerable to extinction: it is not only a dietary specialist, also migrates only small distances. This creates problems for the species, since fragmentation most strongly affects animals that migrate short distances and are unable to cross the gaps made by human
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 ...
. While long distance migratory species are often unaffected and sedentary species are often able to better use the fragments they are in, short distance migrators like the small pearl-bordered fritillary are unable to properly disperse and are hit hardest from environmental damage. A study in Germany found that over 40 years (1971-2011), populations of adults dropped rapidly in agricultural land, while dropping slightly less severely in native grasslands (although still decreasing by almost 83%). The article stated that fragmented habitat protection does a poor job of securing biodiversity and protecting many species that are in peril, as it does not account for many species that cannot seek extra fragmented breeding sites. Many species, including ''Boloria selene'', fail to find good nesting sites as they cannot pass through the human habitat that divides suitable fragments. These drastic changes stem from the continued expansion of agriculture over time that continued to damage and degrade the remaining native breeding habitat for ''Boloria selene''. However, the species seems to fare well as adults in a variety of artificial and natural habitats, with the same study showing slightly larger numbers of the species in farmland than in wild habitat in 1971. Another big issue is that ''Boloria selene'' struggles with sibling mating, as shown when almost no viable offspring came from sibling mating. This could be detrimental to a population that already has reduced individual genetic diversity, as sibling mating will become more common as fragmentation chips away at short distance migration and gene flow in this species. Butterflies of the United States Boloria.selene.1.jpg, Dorsal view Boloria.selene.2.jpg, Ventral view Butterfly Boloria selene (Linnaeus, 1758).jpg, Mating Britishentomologyvolume5Plate386.jpg, Illustration from John Curtis's ''
British Entomology ''British Entomology'' is a classic work of entomology by John Curtis, FLS. It is subtitled ''Being Illustrations and Descriptions of the Genera of Insects found in Great Britain and Ireland: Containing Coloured Figures from Nature of the Most ...
'', Volume 5 File:Silver-bordered Fritillaries, Quetico.jpg, ''B. s. atrocostalis'', mating,
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 ...
,
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
Image:The larvæ of the British butterflies and moths BHL41107817.jpg, Figs 1 larva after 2nd moult, 1a larva after 3nd moult; 1b, 1c, 1d, 1e larva after 4th moult; 1f pupa side view; 1g pupa front view.


References


External links


Butterfly Conservation website descriptionButterfly Conservation management Fact SheetBracken for butterflies fact-sheet
by
Butterfly Conservation Butterfly Conservation (BC) is a UK-wide nonprofit environmentalist organization and charity dedicated to conserving butterflies, moths, and the environment. The charity uses its research to provide advice on how to conserve and restore butterfly ...
* NBN Gateway distribution profile can be accessed throug
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{{Taxonbar, from=Q901373 Boloria Butterflies described in 1775 Butterflies of Asia Butterflies of Europe Butterflies of North America Taxa named by Michael Denis Taxa named by Ignaz Schiffermüller