Heath Fritillary
Encyclopedia
The Heath Fritillary is a butterfly
of the Nymphalidae
family. It is found throughout the Palaearctic from Western Europe to Japan
, in heathland, grassland, and in coppiced woodland. Its association with coppiced woodland earned it the name "Woodman's Follower" in parts of the UK. It is considered a threatened species in the UK and Germany
, but not Europe-wide or globally.
Heath Fritillaries have a wing-span of 39–47 mm. The upperside is predominantly dark brown and orange-brown, with the orange-brown spots delineated by dark brown (along and across the wing-veins); there is a white fringe to the wings through which the dark brown extends. The upperside of the body is a similar dark brown to the colour on the wing, and the base of both wings is dark brown. The underside shows bands of red and (off-)white, again with each vein dark brown and each colour delineated by dark brown. The pattern of white spots at the base of the hindwing (visible at rest) is diagnostic for identification.
The Heath Fritillary ranges throughout the Palaearctic region from Western Europe
to Japan
. In Europe, it is absent from Iceland, Ireland, Scotland, Wales, southern Portugal, southern Spain, the Mediterranean islands and southern Greece.
In the UK, it is restricted to Cornwall
and Devon
grasslands (abandoned hay-meadows); Exmoor
(heathland); and Kent
and Essex
(coppiced woodland on acid soils). The populations in Essex are the result of reintroduction
s, and further reintroductions are ongoing in various other parts of the UK. In the late 1990s, the species was estimated to occupy just 0.2% within the 10-km grid squares over the whole of its UK range.
Its altitudinal range is from sea-level to 2600 m.
, suggesting that globally it is not considered threatened.
In the UK, however, the species was "considered to be the most endangered British butterfly" after a nationwide survey in 1980 found only 31 surviving colonies. Consequently, it was given protection under the Wildlife and Countryside Act 1981
. Despite subsequent widespread efforts to conserve and manage the species' habitats, it has continued to decline: distribution has declined 25% since the 1970s, and abundance-wise it declined by 46% in the 10 years 1995–2004. Woodland sites in Kent and Essex are actively managed (coppiced) for the conservation of this species. It has been a "high priority" species in the UK Biodiversity Action Plan
(BAP) since 1995 and has its own Species Action Plan.
Heath Fritillary is also on the "Red List" in Germany
.
However, the species is considered of "least concern" on a European scale.
, the Heath Fritillary occupies a diversity of grassy, flowery habitats—dry or damp, upland or lowland, with or without shrubs or trees, including woodland clearings and heathland.
More specifically, in England
, this species occupies three distinct habitats:
In France
, this species also occurs on unimproved hay-meadows and pastures.
Eggs are oval spheroids with flattened bases, about 0.5 mm high. They are ribbed (longitudinally, i.e. from top to bottom) and striated (transversely, i.e. around the egg). Pale cream when laid, eggs darken to pale yellow within 2 days, and then dark grey a few days before hatching. Eggs mature in 2–3 weeks.
caterpillar
s (or larvae) eat their eggshells. The caterpillars from a clutch initially stay together, feeding in a small, unobtrusive web. Second or third instar caterpillars disperse into smaller groups. Then the third instars tend to feed and rest solitarily; they rest beneath dead leaves at night and during bad weather. The caterpillar hibernates for the winter in a hibernaculum
, made from a curled dead leaf by spinning its edges together. Hibernacula are usually close to the ground. Although most caterpillars hibernate singly, they sometimes group in twos and threes, although 15–20 caterpillars have been found in single hibernaculum.
Caterpillars re-emerge in early spring. When it is warm, they feed a little, but most of the time is spent basking in the sun. There are six instars in total. The full-grown sixth instar caterpillar is 22–25 mm long, and predominantly black; it has pale (yellow-orange) spines and (greyish-white) spots.
A study in Cornwall (England) recorded mortality rates in the region of 50%, mostly from predation by small mammals, but including predation by beetles, and parasitism
.
Fennoscandian subspecies norvegica flies in June–July, precise timing being affected by the season.
In southern Europe, subspecies celadussa flies in a single brood at high altitude in June–July. Below the subalpine level, however, it is bivoltine
, flying in May–June and late July–August—except for f. nevadensis in the Sierra Nevada
, which is univoltine.
Additional host-plants used elsewhere in Europe are:
are recognized in Europe:
Many forms and subspecies of M. a. athalia have been described, but are best regarded as "ecological variants" and intermediates in clinal variation. Bulgaria
has f. boris Frühstorfer, with heavier marginal borders, alongside the typical nominate form.
In central Sweden
and Finland
, M. a. norvegica f. lachares Frühstorfer has finer black markings; transitional forms between f. lachares and typical M. a. norvegica occur in southern Sweden.
In Portugal
and north-west Spain
, M. a. celadussa f. biedermanni Querci is larger than typical M. a. celadussa, with finer black lines in the outer half of the wing, but a thicker discal line; transitional forms occur in central west Spain. In Sierra Nevada, f. nevadensis Verity
is golden-yellow with fine black markings (it is also univoltine
, whereas typical M. a. celadussa is bivoltine at low altitude). In "southern" Europe, f. tenuicola Verity occurs in late broods—it is small and its black markings reduced.
Butterfly
A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...
of the Nymphalidae
Nymphalidae
The Nymphalidae is a family of about 5,000 species of butterflies which are distributed throughout most of the world. These are usually medium sized to large butterflies. Most species have a reduced pair of forelegs and many hold their colourful wings flat when resting. They are also called...
family. It is found throughout the Palaearctic from Western Europe to Japan
Japan
Japan is an island nation in East Asia. Located in the Pacific Ocean, it lies to the east of the Sea of Japan, China, North Korea, South Korea and Russia, stretching from the Sea of Okhotsk in the north to the East China Sea and Taiwan in the south...
, in heathland, grassland, and in coppiced woodland. Its association with coppiced woodland earned it the name "Woodman's Follower" in parts of the UK. It is considered a threatened species in the UK and Germany
Germany
Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...
, but not Europe-wide or globally.
Description
- See glossaryGlossary of Lepidopteran termsThis glossary describes the terms used in the formal descriptions of insect species, jargon used mostly by professionals or entomologist....
for terms used
Heath Fritillaries have a wing-span of 39–47 mm. The upperside is predominantly dark brown and orange-brown, with the orange-brown spots delineated by dark brown (along and across the wing-veins); there is a white fringe to the wings through which the dark brown extends. The upperside of the body is a similar dark brown to the colour on the wing, and the base of both wings is dark brown. The underside shows bands of red and (off-)white, again with each vein dark brown and each colour delineated by dark brown. The pattern of white spots at the base of the hindwing (visible at rest) is diagnostic for identification.
Range
The Heath Fritillary ranges throughout the Palaearctic region from Western Europe
Western Europe
Western Europe is a loose term for the collection of countries in the western most region of the European continents, though this definition is context-dependent and carries cultural and political connotations. One definition describes Western Europe as a geographic entity—the region lying in the...
to Japan
Japan
Japan is an island nation in East Asia. Located in the Pacific Ocean, it lies to the east of the Sea of Japan, China, North Korea, South Korea and Russia, stretching from the Sea of Okhotsk in the north to the East China Sea and Taiwan in the south...
. In Europe, it is absent from Iceland, Ireland, Scotland, Wales, southern Portugal, southern Spain, the Mediterranean islands and southern Greece.
In the UK, it is restricted to Cornwall
Cornwall
Cornwall is a unitary authority and ceremonial county of England, within the United Kingdom. It is bordered to the north and west by the Celtic Sea, to the south by the English Channel, and to the east by the county of Devon, over the River Tamar. Cornwall has a population of , and covers an area of...
and Devon
Devon
Devon is a large county in southwestern England. The county is sometimes referred to as Devonshire, although the term is rarely used inside the county itself as the county has never been officially "shired", it often indicates a traditional or historical context.The county shares borders with...
grasslands (abandoned hay-meadows); Exmoor
Exmoor
Exmoor is an area of hilly open moorland in west Somerset and north Devon in South West England, named after the main river that flows out of the district, the River Exe. The moor has given its name to a National Park, which includes the Brendon Hills, the East Lyn Valley, the Vale of Porlock and ...
(heathland); and Kent
Kent
Kent is a county in southeast England, and is one of the home counties. It borders East Sussex, Surrey and Greater London and has a defined boundary with Essex in the middle of the Thames Estuary. The ceremonial county boundaries of Kent include the shire county of Kent and the unitary borough of...
and Essex
Essex
Essex is a ceremonial and non-metropolitan county in the East region of England, and one of the home counties. It is located to the northeast of Greater London. It borders with Cambridgeshire and Suffolk to the north, Hertfordshire to the west, Kent to the South and London to the south west...
(coppiced woodland on acid soils). The populations in Essex are the result of reintroduction
Reintroduction
Reintroduction is the deliberate release of a species into the wild in zones formerly inhabited by said species but where it has disappeared from for a number of reasons, from captivity or relocated from other areas where the species still survives in...
s, and further reintroductions are ongoing in various other parts of the UK. In the late 1990s, the species was estimated to occupy just 0.2% within the 10-km grid squares over the whole of its UK range.
Its altitudinal range is from sea-level to 2600 m.
Status
Heath Fritillary is not listed on the IUCN Red ListIUCN Red List
The IUCN Red List of Threatened Species , founded in 1963, is the world's most comprehensive inventory of the global conservation status of biological species. The International Union for Conservation of Nature is the world's main authority on the conservation status of species...
, suggesting that globally it is not considered threatened.
In the UK, however, the species was "considered to be the most endangered British butterfly" after a nationwide survey in 1980 found only 31 surviving colonies. Consequently, it was given protection under the Wildlife and Countryside Act 1981
Wildlife and Countryside Act 1981
The Wildlife and Countryside Act 1981 is an Act of Parliament in the United Kingdom and was implemented to comply with the Directive 2009/147/EC on the conservation of wild birds...
. Despite subsequent widespread efforts to conserve and manage the species' habitats, it has continued to decline: distribution has declined 25% since the 1970s, and abundance-wise it declined by 46% in the 10 years 1995–2004. Woodland sites in Kent and Essex are actively managed (coppiced) for the conservation of this species. It has been a "high priority" species in the UK Biodiversity Action Plan
Biodiversity Action Plan
A Biodiversity Action Plan is an internationally recognized program addressing threatened species and habitats and is designed to protect and restore biological systems. The original impetus for these plans derives from the 1992 Convention on Biological Diversity...
(BAP) since 1995 and has its own Species Action Plan.
Heath Fritillary is also on the "Red List" in Germany
Germany
Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...
.
However, the species is considered of "least concern" on a European scale.
Habitat
Within EuropeEurope
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...
, the Heath Fritillary occupies a diversity of grassy, flowery habitats—dry or damp, upland or lowland, with or without shrubs or trees, including woodland clearings and heathland.
More specifically, in England
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...
, this species occupies three distinct habitats:
- Unimproved grassland with abundant short (5–15 cm) or sparse swards of Ribwort Plantain or Germander Speedwell (or both) on stony soils – sometimes in the form of abandoned hay-meadows
- Sheltered heathland with Common Cow-wheat scattered among BilberryBilberryBilberry is any of several species of low-growing shrubs in the genus Vaccinium , bearing edible berries. The species most often referred to is Vaccinium myrtillus L., but there are several other closely related species....
-dominated vegetation – valleys with mineral soils - Coppiced woodland (especially in clearings) with Common Cow-wheat on acid soils.
In France
France
The French Republic , The French Republic , The French Republic , (commonly known as France , is a unitary semi-presidential republic in Western Europe with several overseas territories and islands located on other continents and in the Indian, Pacific, and Atlantic oceans. Metropolitan France...
, this species also occurs on unimproved hay-meadows and pastures.
Habits
Heath Fritillaries typically fly close to the ground, with characteristic "flits" and glides. Colonies tend to be compact, centred on favoured breeding areas. Heath Fritillaries are highly sedentary for the most part, adults rarely moving more than 100 m; however, some have been recorded dispersing up to 2 km. For a species of often short-lived habitats, it has remarkably limited colonizing ability; suitable habitats over 600 m from an existing colony are colonized slowly.Egg
The female Heath Fritillary lays its eggs (or ova) in batches of (15–)80–150 on the underside of leaf of a larval food plant or on a plant adjacent to the larval foodplant.Eggs are oval spheroids with flattened bases, about 0.5 mm high. They are ribbed (longitudinally, i.e. from top to bottom) and striated (transversely, i.e. around the egg). Pale cream when laid, eggs darken to pale yellow within 2 days, and then dark grey a few days before hatching. Eggs mature in 2–3 weeks.
Caterpillar
Upon emergence, first-instarInstar
An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...
caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...
s (or larvae) eat their eggshells. The caterpillars from a clutch initially stay together, feeding in a small, unobtrusive web. Second or third instar caterpillars disperse into smaller groups. Then the third instars tend to feed and rest solitarily; they rest beneath dead leaves at night and during bad weather. The caterpillar hibernates for the winter in a hibernaculum
Hibernaculum
Hibernaculum can refer to:* Hibernaculum , the location chosen by an animal for hibernation...
, made from a curled dead leaf by spinning its edges together. Hibernacula are usually close to the ground. Although most caterpillars hibernate singly, they sometimes group in twos and threes, although 15–20 caterpillars have been found in single hibernaculum.
Caterpillars re-emerge in early spring. When it is warm, they feed a little, but most of the time is spent basking in the sun. There are six instars in total. The full-grown sixth instar caterpillar is 22–25 mm long, and predominantly black; it has pale (yellow-orange) spines and (greyish-white) spots.
Pupa
The pupae are 12.4–12.8 cm long and last 15–25 days (early May to late June in the UK). They are white with black and orange-brown blotches. Pupae are usually found close to the ground in or beneath dead leaves.A study in Cornwall (England) recorded mortality rates in the region of 50%, mostly from predation by small mammals, but including predation by beetles, and parasitism
Parasitism
Parasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host . These are now called macroparasites...
.
Imago
Imagines probably live for 5–10 days. Males are active on warm sunny days. Females mate once shortly after emerging; they lay their eggs only during warm weather, spending most of the time either basking or hiding in vegetation.Flight period
Across its range (see "Subspecies and variation" below), subspecies athalia shows a protracted flight period from mid-May to mid-August. In favourable localities and/or favourable seasons, a partial second brood has been recorded from mid- or late August to September. In the UK, the flight period is from the end of May to the beginning of July (in the south-west) and early June to early August (in the south-east).Fennoscandian subspecies norvegica flies in June–July, precise timing being affected by the season.
In southern Europe, subspecies celadussa flies in a single brood at high altitude in June–July. Below the subalpine level, however, it is bivoltine
Voltinism
Voltinism is a term used in biology to indicate the number of broods or generations of an organism in a year. The term is particularly in use in sericulture, where silkworm varieties vary in their voltinism....
, flying in May–June and late July–August—except for f. nevadensis in the Sierra Nevada
Sierra Nevada (Spain)
The Sierra Nevada is a mountain range in the region of provinces of Granada and Almería in Spain. It contains the highest point of continental Spain, Mulhacén at 3478 m above sea level....
, which is univoltine.
Host-plants
In the UK, the following species are used:- Common Cow-wheatMelampyrum pratenseMelampyrum pratense or Common Cow-wheat is a plant species of the genus Melampyrum. This plant has an interesting relationship with ants. It produces a sugary liquid from small glands under its petals, which wood ants feed on...
(Melampyrum pratense) — the only foodplant in woodland, and principal foodplant on heathland - Common FoxgloveDigitalis purpureaDigitalis purpurea , is a flowering plant in the family Plantaginaceae , native to most of Europe.-Description:...
(Digitalis purpurea) — secondary foodplant on heathland - Ribwort Plantain (Plantago lanceolata) — major foodplant on grassland
- Germander Speedwell (Veronica chamaedryas) — major foodplant on grassland
- Greater Plantain (P. major) — secondary foodplant
- Ivy-leaved Speedwell (V. hederifolia) — secondary foodplant
- Thyme-leaved SpeedwellVeronica serpyllifoliaVeronica serpyllifolia is a perennial flowering plant in the plantain family. It can be found on most continents.-Description:The leaves are oval, borne on creeping stems...
(V. serpyllifolia) — secondary foodplant - YarrowYarrowAchillea millefolium or yarrow is a flowering plant in the family Asteraceae, native to the Northern Hemisphere. In New Mexico and southern Colorado, it is called plumajillo, or "little feather", for the shape of the leaves. In antiquity, yarrow was known as herbal militaris, for its use in...
(Achillea millefolium) — secondary foodplant
Additional host-plants used elsewhere in Europe are:
- Alpine PlantainPlantagoPlantago is a genus of about 200 species of small, inconspicuous plants commonly called plantains. They share this name with the very dissimilar plantain, a kind of banana. Most are herbaceous plants, though a few are subshrubs growing to 60 cm tall. The leaves are sessile, but have a narrow...
(Plantago alpina) - Mountain SpeedwellVeronica (plant)Veronica is the largest genus in the flowering plant family Plantaginaceae, with about 500 species; it was formerly classified in the family Scrophulariaceae...
(Veronica montana) - Heath SpeedwellVeronica officinalisVeronica officinalis is a species of Veronica, native to Europe and western Asia....
(V. officinalis) - Spiked SpeedwellSpiked SpeedwellVeronica spicata is a species of the genus Veronica.It is the county flower of Montgomeryshire in the United Kingdom. Cultivated varieties include blue , red and white ....
(V. spicata) - Small Cow-wheatMelampyrumMelampyrum is a genus of about 10-20 species of herbaceous flowering plants in the family Orobanchaceae, native to temperate regions of the Northern Hemisphere...
(Melampyrum sylvaticum) - Rusty Foxglove (DigitalisDigitalisDigitalis is a genus of about 20 species of herbaceous perennials, shrubs, and biennials that are commonly called foxgloves. This genus was traditionally placed in the figwort family Scrophulariaceae, but recent reviews of phylogenetic research have placed it in the much enlarged family...
ferruginea) - Yellow FoxgloveDigitalis luteaDigitalis lutea, a Yellow Foxglove, is a short-lived perennial plant.The flowers are tubular, yellow, with brown dots on the inside of the corolla. Flowers are borne beginning in late spring, then sporadically throughout the summer and fall....
(D. lutea) - Common ToadflaxLinaria vulgarisLinaria vulgaris is a species of toadflax , native to most of Europe and northern Asia, from the United Kingdom south to Spain in the west, and east to eastern Siberia and western China...
(Linaria vulgaris)
Subspecies and variation
Up to eight subspeciesSubspecies
Subspecies in biological classification, is either a taxonomic rank subordinate to species, ora taxonomic unit in that rank . A subspecies cannot be recognized in isolation: a species will either be recognized as having no subspecies at all or two or more, never just one...
are recognized in Europe:
- M. a. athalia – from the Atlantic coast to the BosporusBosporusThe Bosphorus or Bosporus , also known as the Istanbul Strait , is a strait that forms part of the boundary between Europe and Asia. It is one of the Turkish Straits, along with the Dardanelles...
(TurkeyTurkeyTurkey , known officially as the Republic of Turkey , is a Eurasian country located in Western Asia and in East Thrace in Southeastern Europe...
) (altitude 0–2200m), excluding the ranges of the following subspecies - M. a. norvegica AurivilliusPer Olof Christopher AurivilliusPer Olof Christopher Aurivillius was a Swedish entomologist.Aurivillius was born at Forsa. He was the Director of the Natural History Museum in Stockholm and he specialised in Coleoptera and Lepidoptera. He was, for a long time, the Secretary of the Royal Swedish Academy of Science...
1888 – FennoscandiaFennoscandiaFennoscandia and Fenno-Scandinavia are geographic and geological terms used to describe the Scandinavian Peninsula, the Kola Peninsula, Karelia and Finland...
(altitude 0–800 m) - M. a. celadussa FrühstorferHans FruhstorferHans Fruhstorfer was a German explorer, insect trader and entomologist who specialised in Lepidoptera. He collected and described many new species of exotic butterflies, especially in Seitz's Macrolepidoptera of the World...
1910 – northern IberiaIberian PeninsulaThe Iberian Peninsula , sometimes called Iberia, is located in the extreme southwest of Europe and includes the modern-day sovereign states of Spain, Portugal and Andorra, as well as the British Overseas Territory of Gibraltar...
and the Sierra NevadaSierra Nevada (Spain)The Sierra Nevada is a mountain range in the region of provinces of Granada and Almería in Spain. It contains the highest point of continental Spain, Mulhacén at 3478 m above sea level....
, southern France, southern SwitzerlandSwitzerlandSwitzerland name of one of the Swiss cantons. ; ; ; or ), in its full name the Swiss Confederation , is a federal republic consisting of 26 cantons, with Bern as the seat of the federal authorities. The country is situated in Western Europe,Or Central Europe depending on the definition....
, ItalyItalyItaly , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...
(including SicilySicilySicily is a region of Italy, and is the largest island in the Mediterranean Sea. Along with the surrounding minor islands, it constitutes an autonomous region of Italy, the Regione Autonoma Siciliana Sicily has a rich and unique culture, especially with regard to the arts, music, literature,...
) (altitude 0–2600 m) - The transitional zone between subspecies athalia and celadussa is broad—up to 150 km in places
- M. a. dictynnoides (Hormuzaki 1898) – south-west Europe?
- M. a. licufuga (Fruhstorfer 1917) – south-east Europe
- M. a. reticulata Higgins 1955 – AltaiAltai KraiAltai Krai is a federal subject of Russia . It borders with, clockwise from the south, Kazakhstan, Novosibirsk and Kemerovo Oblasts, and the Altai Republic. The krai's administrative center is the city of Barnaul...
- M. a. baikalensis (Bremer 1961) – southern SiberiaSiberiaSiberia is an extensive region constituting almost all of Northern Asia. Comprising the central and eastern portion of the Russian Federation, it was part of the Soviet Union from its beginning, as its predecessor states, the Tsardom of Russia and the Russian Empire, conquered it during the 16th...
to AmurAmurThe Amur or Heilong Jiang is the world's tenth longest river, forming the border between the Russian Far East and Northeastern China .-Course:...
? - M. a. hyperborea Dubatolov 1997 – MagadanMagadanMagadan is a port town on the Sea of Okhotsk and gateway to the Kolyma region. It is the administrative center of Magadan Oblast , in the Russian Far East. Founded in 1929 on the site of an earlier settlement from the 1920s, it was granted the status of town in 1939...
, Kamchatka
Many forms and subspecies of M. a. athalia have been described, but are best regarded as "ecological variants" and intermediates in clinal variation. Bulgaria
Bulgaria
Bulgaria , officially the Republic of Bulgaria , is a parliamentary democracy within a unitary constitutional republic in Southeast Europe. The country borders Romania to the north, Serbia and Macedonia to the west, Greece and Turkey to the south, as well as the Black Sea to the east...
has f. boris Frühstorfer, with heavier marginal borders, alongside the typical nominate form.
In central Sweden
Sweden
Sweden , officially the Kingdom of Sweden , is a Nordic country on the Scandinavian Peninsula in Northern Europe. Sweden borders with Norway and Finland and is connected to Denmark by a bridge-tunnel across the Öresund....
and Finland
Finland
Finland , officially the Republic of Finland, is a Nordic country situated in the Fennoscandian region of Northern Europe. It is bordered by Sweden in the west, Norway in the north and Russia in the east, while Estonia lies to its south across the Gulf of Finland.Around 5.4 million people reside...
, M. a. norvegica f. lachares Frühstorfer has finer black markings; transitional forms between f. lachares and typical M. a. norvegica occur in southern Sweden.
In Portugal
Portugal
Portugal , officially the Portuguese Republic is a country situated in southwestern Europe on the Iberian Peninsula. Portugal is the westernmost country of Europe, and is bordered by the Atlantic Ocean to the West and South and by Spain to the North and East. The Atlantic archipelagos of the...
and north-west Spain
Spain
Spain , officially the Kingdom of Spain languages]] under the European Charter for Regional or Minority Languages. In each of these, Spain's official name is as follows:;;;;;;), is a country and member state of the European Union located in southwestern Europe on the Iberian Peninsula...
, M. a. celadussa f. biedermanni Querci is larger than typical M. a. celadussa, with finer black lines in the outer half of the wing, but a thicker discal line; transitional forms occur in central west Spain. In Sierra Nevada, f. nevadensis Verity
Ruggero Verity
Ruggero Verity was an Italian entomologist who specialised in butterflies.Verity was born in Florence on 20 May 1883, the elder son of Richard Henry Manners Verity and his wife Matilda daughter of Cav. Sebastiano Fenzi and Emily Verity...
is golden-yellow with fine black markings (it is also univoltine
Voltinism
Voltinism is a term used in biology to indicate the number of broods or generations of an organism in a year. The term is particularly in use in sericulture, where silkworm varieties vary in their voltinism....
, whereas typical M. a. celadussa is bivoltine at low altitude). In "southern" Europe, f. tenuicola Verity occurs in late broods—it is small and its black markings reduced.