Glanville fritillary
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The Glanville fritillary (''Melitaea cinxia'') is a
butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprise ...
of the family
Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a red ...
. It is named for the naturalist who discovered it and the checkerboard pattern on its wings. These butterflies live in almost all of Europe, especially
Finland Finland ( fi, Suomi ; sv, Finland ), officially the Republic of Finland (; ), is a Nordic country in Northern Europe. It shares land borders with Sweden to the northwest, Norway to the north, and Russia to the east, with the Gulf of B ...
, and in parts of northwest Africa. They are absent from the far north of Europe and parts of the Iberian Peninsula. To the east they are found across the Palearctic (in Turkey, Russia, northern Kazakhstan and Mongolia). It has been discovered that this butterfly only mates one time in June or July and lays its eggs. It does not provide any protection to these eggs or care for the offspring. As adults, the Glanville fritillaries are short-lived; they spend most of their lives as caterpillars. As caterpillars, Glanville fritillaries enter a stage of
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press I ...
, which is a period of suspended development, during the winter time. The spiked speedwell and ribwort plantain are the Glanville fritillary's preferred plants to lay eggs and to eat as larvae. Female butterflies will show a preference for one plant species over the other when deciding where to lay their eggs, but the caterpillars have no preference once they hatch. After entering the adult phase the fritillaries feed on nectar of the spiked speedwell and ribwort plantain, among others. This species of butterfly is at risk of population decline because it is not a migratory species. Though widespread, populations in Finland are at risk because they are not able to travel great distances as easily as other species, such as monarchs, if their environment should suddenly become unsuitable.


Taxonomy

The word ''fritillary'' refers to the checkered pattern of the butterfly's wings, which comes from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word ''fritillus'' which means "dicebox". The name "Glanville" comes from the naturalist who discovered it, Lady Eleanor Glanville, who was an eccentric 17th- and 18th-century English butterfly enthusiast – a very unusual occupation for a woman at that time. She was the first to capture British specimens in
Lincolnshire Lincolnshire (abbreviated Lincs.) is a Counties of England, county in the East Midlands of England, with a long coastline on the North Sea to the east. It borders Norfolk to the south-east, Cambridgeshire to the south, Rutland to the south-we ...
during the 1690s. A contemporary wrote


Geographic range and habitat

The Glanville fritillary is found across Europe and temperate Asia. It is most commonly found on
Åland Åland ( fi, Ahvenanmaa: ; ; ) is an autonomous and demilitarised region of Finland since 1920 by a decision of the League of Nations. It is the smallest region of Finland by area and population, with a size of 1,580 km2, and a populat ...
(Finland), which host a network of about 4,000 dry meadows, the fritillary's ideal habitat. These butterflies commonly inhabit open grassland at an elevation of above sea level.


Glanville fritillaries in the UK

In the UK the Glanville fritillary occurs only on soft
undercliff The Undercliff is the name of several areas of landslip on the south coast of England. They include ones on the Isle of Wight; on the Dorset-Devon border near Lyme Regis; on cliffs near Branscombe in East Devon; and at White Nothe, Dorset. All aro ...
and
chine A chine () is a steep-sided coastal gorge where a river flows to the sea through, typically, soft eroding cliffs of sandstone or clays. The word is still in use in central Southern England—notably in East Devon, Dorset, Hampshire and the Is ...
grassland A grassland is an area where the vegetation is dominated by grasses ( Poaceae). However, sedge ( Cyperaceae) and rush ( Juncaceae) can also be found along with variable proportions of legumes, like clover, and other herbs. Grasslands occur na ...
and where its main larval food plant '' Plantago lanceolata'' (English plantain) occurs in abundance on sheltered, south facing slopes. The Glanville fritillary is a highly restricted species within the UK, being confined to the
Isle of Wight The Isle of Wight ( ) is a Counties of England, county in the English Channel, off the coast of Hampshire, from which it is separated by the Solent. It is the List of islands of England#Largest islands, largest and List of islands of England#Mo ...
and even there being largely limited to the southern coast. It also occurs in the
Channel Islands The Channel Islands ( nrf, Îles d'la Manche; french: îles Anglo-Normandes or ''îles de la Manche'') are an archipelago in the English Channel, off the French coast of Normandy. They include two Crown Dependencies: the Bailiwick of Jersey, ...
, and since 1990 there has been a mainland site on the Hampshire coast, possibly the result of an introduction. There are small introduced populations on the
Somerset ( en, All The People of Somerset) , locator_map = , coordinates = , region = South West England , established_date = Ancient , established_by = , preceded_by = , origin = , lord_lieutenant_office =Lord Lieutenant of Somerset , lord_ ...
coast and two in Surrey: one near
Wrecclesham Wrecclesham is a village on the southern outskirts of the large town of Farnham in Surrey, England. Its local government district is the Borough of Waverley. History It was once in the estate of Henry of Westminster and Blois the powerful 13th ...
, and one at a nature reserve in Addington, near
Croydon Croydon is a large town in south London, England, south of Charing Cross. Part of the London Borough of Croydon, a local government district of Greater London. It is one of the largest commercial districts in Greater London, with an extensi ...
. Historic UK records suggest a distribution which went as far north as
Lincolnshire Lincolnshire (abbreviated Lincs.) is a Counties of England, county in the East Midlands of England, with a long coastline on the North Sea to the east. It borders Norfolk to the south-east, Cambridgeshire to the south, Rutland to the south-we ...
. However, by the middle of the 19th century the Glanville fritillary was known only from the Isle of Wight and the coast of
Kent Kent is a county in South East England and one of the home counties. It borders Greater London to the north-west, Surrey to the west and East Sussex to the south-west, and Essex to the north across the estuary of the River Thames; it faces ...
between Folkestone and Sandwich. It became extinct in Kent by the mid-1860s.


Description

''Melitaea cinxia'' has a
wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingspan o ...
of about . These medium-sized butterflies have orange, black and white "checkerspot" forewings. On the upper side of the hindwings they have a row of black dots. The hindwings have white and orange bands and a series of black dots inside them, also clearly visible on the reverse. Females are usually more dull than males with more developed black dots. In Seitz it is described - ''M. cinxia'' L. (65 e, f). Above uniformly pale yellowish red, marked with black, somewhat recalling a chess-board, the white fringes being checkered. A row of heavy black dots in the submarginal row of spots on the hindwing is characteristic. Excepting the pale yellow black-dotted apex, the forewing beneath uniformly reddish leather-yellow, with dispersed blackspots, which vary in number. The species though of wide distribution, has not developed into many races. Caterpillars are about 25 mm long with a reddish-brown head and a spiny black body with small white dots. ''Melitaea cinxia'' is rather similar to the heath fritillary ('' Melitaea athalia''), but the beige and orange bands on the underwings are distinctive. Moreover, the latter one has no spots on the upperside of the hindwings. Melitaea cinxia MHNT CUT 2013 3 27 Avilly-Saint-Léonard dorsal.jpg, Upperside Melitaea cinxia MHNT CUT 2013 3 27 Avilly-Saint-Léonard ventral.jpg, Underside


Food Resources


Caterpillars

After hatching, Glanville fritillary caterpillars live in gregarious sibling groups. They feed on their host plant, either '' Plantago lanceolata'' (ribwort plantain) or '' Veronica spicata'' (spiked speedwell). Adult females prefer one plant over the other when choosing where to lay their eggs, but the larvae do not have a feeding preference for either plant when they are born.


Spiked speedwell Defense

As an attempt to fend off predators and parasitoids, the spiked speedwell emits
volatile organic compound Volatile organic compounds (VOCs) are organic compounds that have a high vapour pressure at room temperature. High vapor pressure correlates with a low boiling point, which relates to the number of the sample's molecules in the surrounding air, a ...
s (VOCs) when threatened. Some researchers found that this plant species has two different defenses for when it is being fed on and when a butterfly is in oviposition. The oviposition of the butterfly on this plant was able to induce the increase of two ketones (6-methyl-5-hepten-2-one and t-geranylacetone) and the suppression of green leaf volatiles (GLVs).


Adults

Adult Glanville fritillaries have a diet that consists solely of nectar. It has been shown that larvae with higher amounts of
iridoid Iridoids are a type of monoterpenoids in the general form of cyclopentanopyran, found in a wide variety of plants and some animals. They are biosynthetically derived from 8-oxogeranial. Iridoids are typically found in plants as glycosides, mos ...
glycoside In chemistry, a glycoside is a molecule in which a sugar is bound to another functional group via a glycosidic bond. Glycosides play numerous important roles in living organisms. Many plants store chemicals in the form of inactive glycoside ...
s have better defense against parasitoids and bacterial pathogens.


Mating

Adult Glanville fritillaries take flight, mate, and lay their eggs from June to early July. During this time it has been found that females only mate once. Females also tend to mate in their natal groups before dispersing with a mate to lay their eggs in a different population. This dispersal helps induce genetic flow between populations on the fragmented meadows of the Åland Islands.


Life history


Life cycle (egg, caterpillar, pupa, adult)

A female Glanville fritillary will lay as many as 10 clutches of eggs in her lifetime. These clutches can range in size from 50 up to 300 eggs and are laid on the underside of the larval food plant of either '' Plantago lanceolata'' (ribwort plantain) or '' Veronica spicata'' (spiked speedwell). After hatching, the caterpillars feed on their host plants until the end of the summer where they spin a "winter nest" to
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press I ...
. When winter is over they emerge from their nest to feed again before pupating around the beginning of May. A month later, the adult fritillaries emerge and go about their adult lives of feeding, mating, and laying eggs.


Clutch size

In the Glanville fritillary, clutch size is highly important as larval survival depends on groups size. Thus, understanding the different factors that contribute to clutch size is important in developing an understanding of population dynamics and life cycle. Clutch size is inversely related to the size of the clutches previously laid, meaning that the more eggs a female lays in years prior, the fewer eggs will be in the present clutch. Further, it is known that clutch sizes decrease with the age of the female. This is thought to be due to resource depletion over time in the female, potentially from high investment in earlier clutches. Clutch size also increases with increased intervals between oviposition. Typical Glanville fritillary oviposition occurs once every two days.


PGI genotypes

Glanville fritillary females vary in their PGI, a glycolytic enzyme, genotype. The genotype of the female influences the time of day that she is active, which also influence her ability to lay larger clutches. Heterozygote females are able to operate earlier in the day compared to all other genotypes. Further, females with a ''Pgi-f'' allele are able to begin oviposition earlier in the day and are also able to lay larger clutch sizes. The ''Pgi-f'' allele contributes to a higher metabolic rate and the ability for females to take advantage of the climate of the early day - both of which factors allow for increased clutch size. File: Ausgewachsene Raupe des Wegerich - Scheckenfalters.jpg, Caterpillar File:Wegerich Scheckenfalter Puppen.jpg, Pupa File: Nymphalidae - Melitaea cinxia.jpg, Imago


Ecology


Parasites

Glanville fritillary eggs and pupae are often parasitized by several species of parasitoids. Two specialist species: * '' Cotesia melitaearum'' * '' Hyposoter horticola'' And several generalist species: * '' Pterolmalus apum'' * '' Ichneumon gracilicornis'' * '' Pteromalus apum'' * ''
Pteromalus puparum ''Pteromalus'' is a genus of pteromalids in the family Pteromalidae. There are at least 430 described species in ''Pteromalus''. See also * List of Pteromalus species This is a list of 432 species in ''Pteromalus'', a genus of pteromalids in t ...
'' * '' Coelopisthia caledonica'' And has several
hyperparasitoid A hyperparasite, also known as a metaparasite, is a parasite whose host, often an insect, is also a parasite, often specifically a parasitoid. Hyperparasites are found mainly among the wasp-waisted Apocrita within the Hymenoptera, and in two ...
s * '' Mesochorus stigmaticus'' (a specialist) * ''
Gelis agilis ''Gelis agilis'' is a tiny wingless hyperparasitoid wasp that attacks the parasitoid wasp ''Cotesia glomerata'' and other parasitoids, like ''Dinocampus coccinellae''. References

{{Taxonbar, from=Q2751372 Ichneumonidae Hyperparasites ...
'' * '' Gelis ruficornis'' * ''
Gelis acarorum ''Gelis'' is a genus of ichneumon wasps in the family Ichneumonidae The Ichneumonidae, also known as the ichneumon wasps, Darwin wasps, or ichneumonids, are a family of parasitoid wasps of the insect order Hymenoptera. They are one of the mos ...
''


''Cotesia melitaearum''

The parasitoid '' C. melitaearum'' can be extremely detrimental to Glanville fritillary populations if it is able to become well established. This species is able to increase its own parasitism if the fritillary population is also increasing in size and age, but will decrease if said population becomes isolated. Thus, in well established fritillary populations there is a risk of local extinction by the ''C. melitaearum'' parasitoid.


Genetics


Subspecies

Subspecies include: * ''Melitaea cinxia cinxia'' * ''Melitaea cinxia amardea'' Grum-Grshimailo, 1895 * ''Melitaea cinxia atlantis'' Le Cerf, 1928 * ''Melitaea cinxia clarissa'' Staudinger, 1901 * ''Melitaea cinxia heynei'' Rühl, 1895 * ''Melitaea cinxia karavajevi'' Obraztsov, 1936 * ''Melitaea cinxia oasis'' Huang & Murayama * ''Melitaea cinxia sacarina'' Fruhstorfer, 1917 * ''Melitaea cinxia tschujaca'' Seitz, 1908


Conservation


Migration Issues

As the global temperature of the Earth warms each year, many butterfly species are forced to shift northward in order to keep living in their preferred climates. Scientists have found that migratory species, such as monarch butterflies, are better able to adapt to the rising temperatures than sedentary species, such as Glanville fritillaries. Glanville fritillaries will migrate around the Åland Islands to different habitat patches, but they will not embark on great transcontinental seasonal migrations like other butterflies do. Because of this they are very susceptible to climate change, which is something to keep in mind when considering plans to aid in the conservation of this species.


Status

This butterfly is not currently listed as threatened in Europe, but its UK BAP status is Priority Species. The NERC act of England lists it as species of principal importance. Its Butterfly Conservation priority is high, so this is a butterfly likely to be increasingly threatened in the coming years.


References


General references

* * * * *


External links


Lepiforum
* Paolo Mazzei, Daniel Morel, Raniero Panfil

{{Taxonbar, from=Q938020 Melitaea Butterflies of Asia Butterflies of Europe Butterflies described in 1758 Taxa named by Carl Linnaeus Environment of the Isle of Wight