The meadow brown (''Maniola jurtina'') is a butterfly found in the

Palearctic realm The Palearctic or Palaearctic is the largest of the eight biogeographic realms of the Earth. It stretches across all of Eurasia north of the foothills of the Himalayas, and North Africa. The realm consists of several bioregions: the Euro-Sibe ...

. Its range includes Europe south of 62°N,

Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...

eastwards to the

Urals The Ural Mountains ( ; rus, Ура́льские го́ры, r=Uralskiye gory, p=ʊˈralʲskʲɪjə ˈɡorɨ; ba, Урал тауҙары) or simply the Urals, are a mountain range that runs approximately from north to south through European ...

Asia Minor Anatolia, tr, Anadolu Yarımadası), and the Anatolian plateau, also known as Asia Minor, is a large peninsula in Western Asia and the westernmost protrusion of the Asian continent. It constitutes the major part of modern-day Turkey. The re ...

Iraq Iraq,; ku, عێراق, translit=Êraq officially the Republic of Iraq, '; ku, کۆماری عێراق, translit=Komarî Êraq is a country in Western Asia. It is bordered by Turkey to Iraq–Turkey border, the north, Iran to Iran–Iraq ...

Iran Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Turkmeni ...

North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...

and the

Canary Islands The Canary Islands (; es, Canarias, ), also known informally as the Canaries, are a Spanish autonomous community and archipelago in the Atlantic Ocean, in Macaronesia. At their closest point to the African mainland, they are west of Morocc ...

. The

larvae 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 ...

feed on grasses.

Description

There is marked

sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...

in this

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 ...

. The upperside of the male is uniformly light brown with a black ocellus centered white at the apex of the forewing, while the female has a tawny patch more or less extended around this ocella. The underside forewing is ochre-colored bordered with dark beige with the same ocelli at the apex in the male, while the hindwing is greyish to brown with a more or less orange band in the female. The males are also much more active and range far about, while females fly less and often may not move away from the area where they grew up.

Description in Seitz

Above dark brown: the apical ocellus minutely centred with white, being in the male bordered with dull dark yellow, and standing in the female in an ochre-yellow half band, which becomes narrower behind and does not reach the hindmargin. The upperside of the live male has often a splendid metallic gloss and bears a broad scent-patch below the cell. Underside of hindwing in the male dark brown, with a hardly perceptible middle band, in the female grey-brown, with a broad, pale, proximally golden-brown-bordered curved band. In several forms throughout Europe Apart from the forms characterized by the disappearance of ocelli or the appearance of accessory there are firstly the albinos which have received in which the blackish ground-colour is replaced by dirty white, while the reddish yellow halfband has remained as such. In others the reddish yellow halfband on the forewing of the female is pale to ivory-white. Specimens have been described in which the band of the male is very prominent on a silky dust-grey upperside. Aberration ''cinerea'' has a bluish gloss on the dark upperside; the hindwing is strongly dentate, tinged with pink on the underside, and bears two eye-dots In hot summers one not seldom meets with specimens in which the reddish yellow colour has increased, this colour being represented by a yellowish red dusting on the apical area in the male and in the female by a yellowish red area in the disc of the hindwing. ubspecies''hispulla'' Hbn. (47b) from Southern Europe, has these characteristics in a still more pronounced degree, and is, besides, generally broader-winged than the nymotypical jurtina — ''fortunata'' Alph. (47 c) is a yet paler form. The live male has in its apical area a magnificent golden gloss on a deep black ground, in the female the ground-colour above is reduced by the extension of the reddish yellow. Moreover, the form is much larger and the basal area of the hindwing is so darkened below that the light discal band contrasts vividly. Seitz. A. in Seitz, A. ed. Band 1: Abt. 1, ''Die Großschmetterlinge des palaearktischen Faunengebietes, Die palaearktischen Tagfalter'', 1909, 379 Seiten, mit 89 kolorierten Tafeln (3470 Figuren) A variable number of smaller eyespots are usually found on the hindwing undersides. These may number up to twelve per butterfly, with up to six on each wing, but sometimes none. The factors that govern polymorphism in this trait are not resolved, although a number of theories have been proposed (Stevens 2005). On the other hand, the

evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

ary significance of the upperwing eyespots is more obvious: The more active males have a markedly more cryptic upperside pattern, whereas the females have more opportunity to present their eyespots in a sudden display of colors and patterns that presumably startle

predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...

s so the butterfly has a better chance of escaping. Some specimens are bi-pupilled. File:ButterflyOnFlower.jpg, Maniola jurtina female on

buttercup ''Ranunculus'' is a large genus of about almost 1700 to more than 1800 species of flowering plants in the family Ranunculaceae. Members of the genus are known as buttercups, spearworts and water crowfoots. The genus is distributed in Europe, ...

flower. File:Coenonympha pamphilus qtl2.jpg,

Small heath Small Heath is an area in south-east Birmingham, West Midlands, England situated on and around the Coventry Road about from the city centre. History Small Heath, which has been settled and used since Roman times, sits on top of a small hill. ...

(for comparison) File:Großes Ochsenauge Maniola jurtina 3.JPG, Female File:Meadow brown butterfly (Maniola jurtina) male underside bipupilled.JPG, Female File:Meadow brown butterfly (Maniola jurtina) male underside no spots.jpg, Female File:Große Ochsenauge, Maniola jurtina, mating.JPG, Mating File:Meadow brown butterflies (Maniola jurtina) mating.jpg, Mating File:Maniola jurtina caterpillar.jpg, First

instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...

caterpillar File:Maniola jurtina pupa.jpg, Pupa South1906Plate84

Similar species

Similar species are the

gatekeeper A gatekeeper is a person who controls access to something, for example via a city gate or bouncer, or more abstractly, controls who is granted access to a category or status. Gatekeepers assess who is "in or out", in the classic words of manage ...

(which prefers to rest with its wings open) and the

small heath Small Heath is an area in south-east Birmingham, West Midlands, England situated on and around the Coventry Road about from the city centre. History Small Heath, which has been settled and used since Roman times, sits on top of a small hill. ...

(which is smaller). More similar are the dusky meadow brown (''

Hyponephele lycaon ''Hyponephele lycaon'', the dusky meadow brown, is a butterfly species belonging to the family Nymphalidae.

'' smaller, male

androconia The external morphology of Lepidoptera is the physiological structure of the bodies of insects belonging to the order Lepidoptera, also known as butterflies and moths. Lepidoptera are distinguished from other orders by the presence ...

l area obliquely directed and divided into three parts by veins Cu1 and Cu2, female with two ochreous ringed eye-spots) is otherwise very similar, '' Pyronia janiroides'', ''

Hyponephele moroccana ''Hyponephele'' is a genus of butterflies of the subfamily Satyrinae in the family Nymphalidae. Most range from Europe into central Asia with a few in northern Africa. They are commonly called meadowbrowns. Species Listed alphabetically:< ...

'', ''

Hyponephele lycaon ''Hyponephele lycaon'', the dusky meadow brown, is a butterfly species belonging to the family Nymphalidae.

'' and ''

Hyponephele lupina ''Hyponephele lupina'', the Oriental meadow brown or branded meadowbrown, is a butterfly of the family Nymphalidae. It is found in North Africa, south western Europe, southern Russia, Asia Minor, southern Siberia, Middle Asia, Iran and from Balo ...

''. Most problematic is the

cryptic species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...

of '' Maniola'' ('' Maniola telmessia'', '' Maniola nurag'', '' Maniola chia'', ''

Maniola halicarnassus ''Maniola'' is a genus of butterflies within the family Nymphalidae.

'' and '' Maniola cypricola'').

Description of egg, larva and pupa

The egg, laid on a blade of grass as shown (Plate 85), is upright and ribbed; the top is flattened, with an impressed ring thereon. Color, whitish green inclining to brownish yellow as it matures, and marked with purplish brown. The caterpillar is bright green, clothed with short whitish hairs; there is a darker line down the back, and a diffused white stripe on each side above the reddish spiracles; the anal points are white. Head rather darker green, hairy. The chrysalis is pale green, marked with brownish on the wing covers, the thorax is spotted with blackish, and the points on the body are brownish. Suspended, and with the old skin attached. (South, 1906).

Food plants

Recorded larval food plants include rough meadow grass (''

Poa trivialis ''Poa trivialis'' (rough bluegrass; ''UK:'' rough-stalked meadow-grass or rough meadow-grass), is a perennial plant regarded in the US as an ornamental plant. It is part of the Poaceae, grass family. Description It is very common in meadows an ...

''), smooth meadow grass (''

Poa pratensis ''Poa pratensis'', commonly known as Kentucky bluegrass (or blue grass), smooth meadow-grass, or common meadow-grass, is a perennial species of grass native to practically all of Europe, North Asia and the mountains of Algeria and Morocco. Altho ...

''), ''

Festuca ''Festuca'' (fescue) is a genus of flowering plants belonging to the grass family Poaceae (subfamily Pooideae). They are evergreen or herbaceous perennial tufted grasses with a height range of and a cosmopolitan distribution, occurring on every ...

'' species, bents (''

Agrostis ''Agrostis'' (bent or bentgrass) is a large and very nearly cosmopolitan genus of plants in the grass family, found in nearly all the countries in the world. It has been bred as a GMO creeping bent grass. Species * '' Agrostis aequivalvi'' ...

'' species), and cock's-foot (''

Dactylis glomerata ''Dactylis Glomerata'' is the sixth studio album by Swedish doom metal band Candlemass, released in 1998. This was their first album released since their split in 1994. The album was originally destined to be the second release by Candlemass ...

''), false brome (''

Brachypodium sylvaticum ''Brachypodium sylvaticum'', commonly known as false-brome, slender false brome or wood false brome, is a perennial grass native to Europe, Asia and Africa. It has a broad native range stretching from North Africa to Eurasia. The bunchgrass is ...

''), downy oat-grass and '' Helictotrichon pubescens''. Less specific records of '' Poa'', ''

Bromus ''Bromus'' is a large genus of grasses, classified in its own tribe Bromeae. They are commonly known as bromes, brome grasses, cheat grasses or chess grasses. Estimates in the scientific literature of the number of species have ranged from 100 ...

'', ''

'', '' Milium'', ''

Brachypodium ''Brachypodium'' is a genus of plants in the grass family, widespread across much of Africa, Eurasia, and Latin America. The genus is classified in its own tribe Brachypodieae. Flimsy upright stems form tussocks. Flowers appear in compact spi ...

'', '' Lolium'', ''

Avena ''Avena'' is a genus of Eurasian and African plants in the grass family. Collectively known as the oats, they include some species which have been cultivated for thousands of years as a food source for humans and livestock. They are widesprea ...

'', ''

Alopecurus ''Alopecurus'', or foxtail grass, is a common and widespread genus of plants in the grass family. It is common across temperate and subtropical parts of Eurasia, northern Africa, and the Americas, as well as naturalized in Australia and on vari ...

'' and ''

Anthoxanthum ''Anthoxanthum'' (Latinised Greek for "yellow blossom"), commonly known as hornworts, vernal grasses, or vernalgrasses, is a genus of plants in the grass family. The generic name means 'Yellow flower' in Botanical Latin, referring to the colo ...

''. Adults feed on

nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...

from a wide spectrum of plants including ''

Centaurea ''Centaurea'' () is a genus of over 700 species of herbaceous thistle-like flowering plants in the family Asteraceae. Members of the genus are found only north of the equator, mostly in the Eastern Hemisphere; the Middle East and surrounding reg ...

'', ''

Cirsium ''Cirsium'' is a genus of perennial and biennial flowering plants in the Asteraceae, one of several genera known commonly as thistles. They are more precisely known as plume thistles. These differ from other thistle genera ('' Carduus'', ''Silyb ...

'', ''

Leontodon ''Leontodon'' is a genus of plants in the tribe Cichorieae within the family Asteraceae, commonly known as hawkbits. Their English name derives from the mediaeval belief that hawks ate the plant to improve their eyesight. Although originally on ...

'', ''

Erica Erica or ERICA may refer to: * Erica (given name) * ''Erica'' (plant), a flowering plant genus * Erica (chatbot), a service of Bank of America * ''Erica'' (video game), a 2019 FMV video game * ''Erica'' (spider), a jumping spider genus * Eric ...

'', ''

Rubus ''Rubus'' is a large and diverse genus of flowering plants in the rose family, Rosaceae, subfamily Rosoideae, with over 1,350 species. Raspberries, blackberries, and dewberries are common, widely distributed members of the genus. Most of these ...

'', '' Heracleum'', ''

Eupatorium ''Eupatorium'' is a genus of flowering plants in the family Asteraceae, containing from 36 to 60 species depending on the classification system. Most are herbaceous perennials growing to tall. A few are shrubs. The genus is native to temperate ...

'', (''

sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...

'') ''

Origanum ''Origanum'' ( ) is a genus of herbaceous perennials and subshrubs in the family Lamiaceae, native to Europe, North Africa, and much of temperate Asia, where they are found in open or mountainous habitats. A few species also naturalized in sc ...

'', ''

Senecio ''Senecio'' is a genus of flowering plants in the daisy family (Asteraceae) that includes ragworts and groundsels. Variously circumscribed taxonomically, the genus ''Senecio'' is one of the largest genera of flowering plants. Description Morp ...

'', ''

Scabiosa ''Scabiosa'' is a genus in the honeysuckle family (Caprifoliaceae) of flowering plants. Many of the species in this genus have common names that include the word scabious, but some plants commonly known as scabious are currently classified in r ...

'', ''

Succisa ''Succisa'' is a genus of flowering plants in the family (biology), family Caprifoliaceae. Species include the devil's-bit scabious, ''Succisa pratensis''. Species , Plants of the World Online accepted three species: *''Succisa pinnatifida'' La ...

'', ''

Ligustrum A privet is a flowering plant in the genus ''Ligustrum''. The genus contains about 50 species of erect, deciduous or evergreen shrubs, sometimes forming small or medium-sized trees, native to Europe, north Africa, Asia, many introduced and nat ...

'' and ''

Filipendula ''Filipendula'' is a genus of 12 species of perennial herbaceous flowering plants in the family Rosaceae, native to the temperate regions of the Northern Hemisphere. Well-known species include meadowsweet (''Filipendula ulmaria'') and dropwort ( ...

''.

Flight time

The meadow brown is

univoltine Voltinism is a term used in biology to indicate the number of broods or generations of an organism in a year. The term is most often applied to insects, and is particularly in use in sericulture, where silkworm varieties vary in their voltinism. ...

(one generation per year) and adults emerge over a long period (spring, summer and autumn). Some individuals have a short larval development time and produce late adults. This is

genetically Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar working i ...

controlled.

Habitat

Because of the vast distribution habitats are hard to define but broadly are forest edge,

forest-steppe A forest steppe is a temperate-climate ecotone and habitat type composed of grassland interspersed with areas of woodland or forest. Locations Forest steppe primarily occurs in a belt of forest steppes across northern Eurasia from the eastern ...

and meadow steppe habitats up to 2,000 m above sea level, cultivated lands (meadow, forest plantations, parks and orchards).

Subspecies

*''Maniola jurtina hispulla'' (Esper, 1805) Portugal, Spain *''Maniola jurtina hyperhispulla'' (Thomson, 1973) Maltese Archipelago (Gozo) *''Maniola jurtina jurtina'' (Linnaeus, 1758) Europe (type locality Sweden), Spain *''Maniola jurtina janira'' Linnaeus, 1758 central Europe, Russia, Latvia, Poland, Slovakia, Hungary, Ukraine, middle and southern Urals, Kazakhstan, western Siberia (Kurgan) *''Maniola jurtina strandiana'' Oberthür, 1936 south Europe, Caucasus Major and Minor Crimea, Armenia, Azerbaijan *''Maniola jurtina persica'' LeCerf, 1912 (=? ''ghilanica'' LeCerf, 1913) Kopet-Dagh, Azerbaijan, Turkmenistan *''Maniola jurtina phormia'' (Fruhstorfer, 1909) Slovenia

References

* (2005): The role of eyespots as anti-predator mechanisms, principally demonstrated in the Lepidoptera. ''

Biological Reviews The Cambridge Philosophical Society (CPS) is a scientific society at the University of Cambridge. It was founded in 1819. The name derives from the medieval use of the word philosophy to denote any research undertaken outside the fields of law ...

'' 80(4): 573–588. (HTML abstract) *Andrea Grill, Rob de Vos, Jan van Arkel, 2004 The shape of endemics: Notes on male and female genitalia in the genus Maniola (Schrank, 1801), (Lepidoptera, Nymphalidae, Satyrinae) ''Contributions to Zoology'', 73 (4) (2004
online here

External links

Satyrinae of the Western Palearctic
{{Taxonbar, from=Q754668 Maniola Butterflies of Africa Butterflies of Asia Butterflies of Europe Butterflies described in 1758 Taxa named by Carl Linnaeus