Brassy Ringlet
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The brassy ringlets are a species group of ringlet butterflies in the genus ''
Erebia ''Erebia'' is a Holarctic genus of brush-footed butterflies, family Nymphalidae. Most of the about 90–100 species (see also below) are dark brown or black in color, with reddish-brown to orange or more rarely yellowish wing blotches or band ...
''.Warren (1936) Though closely related, their
monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...
is not completely resolved. Still, the brassy ringlets are taxa similar to ''E. tyndarus'' – the Swiss brassy ringlet –, and in many cases certainly close relatives. A notable trait of their genus is an ability to adapt well to cold and somewhat
arid A region is arid when it severely lacks available water, to the extent of hindering or preventing the growth and development of plant and animal life. Regions with arid climates tend to lack vegetation and are called xeric or desertic. Most ar ...
habitat, like
taiga Taiga (; rus, тайга́, p=tɐjˈɡa; relates to Mongolic and Turkic languages), generally referred to in North America as a boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruce ...
or regions with
alpine climate Alpine climate is the typical weather (climate) for elevations above the tree line, where trees fail to grow due to cold. This climate is also referred to as a mountain climate or highland climate. Definition There are multiple definitions of ...
. Optimal habitat in Eurasia, where most of the brassy ringlets are found, therefore occurs in two distinct belts – in the very north of the continent and in the Alpide belt – in
interglacial An interglacial period (or alternatively interglacial, interglaciation) is a geological interval of warmer global average temperature lasting thousands of years that separates consecutive glacial periods within an ice age. The current Holocene in ...
s, and in glacials in one
periglacial Periglaciation (adjective: "periglacial", also referring to places at the edges of glacial areas) describes geomorphic processes that result from seasonal thawing of snow in areas of permafrost, the runoff from which refreezes in ice wedges and ot ...
ic belt at lower altitude, in places interrupted by dry wasteland and deserts.Albre ''et al.'' (2008)


Description

Brassy ringlets are mid-sized members of their genus, with a length of 17–22 mm (roughly two-thirds to one inch). Like their relatives, they have black eyespots on their forewings. The largest eyespots are placed, two on each side, near the wingtip, and have a white dot in the center; they are surrounded by a
copper Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkis ...
-colored patch that is lighter than the blackish-brown upperwings. In some, a third black spot, much smaller and without the white dot, is present at the opposite end of the reddish patch. The hindwings have no black spots in many, but in some taxa there are a few (2-4 or so) black dots, usually without white in the center, paralleling the outer margin. If hindwing spots are present, they are sometimes surrounded by a lighter brown field like the forewing spots, sometimes not. The wing undersides are cryptically colored, usually in grayish-brown and often with a noticeable band arching through the central hindwing, the rest of which has a silvery sheen which makes these species quite recognizable in flight. The eyespots and the forewing patch surrounding them are found on the forewing undersides also; if a hindwing pattern is present, it may or may not show up on the under hindwings either whole or in parts. The common name derives from the
brass Brass is an alloy of copper (Cu) and zinc (Zn), in proportions which can be varied to achieve different mechanical, electrical, and chemical properties. It is a substitutional alloy: atoms of the two constituents may replace each other with ...
y luster of the dark wing uppersides. The valvae have a series of external spines pointing
distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
ly, with the first spine pair usually being largest.


Evolution

While the details are still somewhat conjectural, a quite robust evolutionary scenario has been developed for the brassy ringlet group: As it seems, the origin of this group is perhaps south of the
Central Asia Central Asia, also known as Middle Asia, is a subregion, region of Asia that stretches from the Caspian Sea in the west to western China and Mongolia in the east, and from Afghanistan and Iran in the south to Russia in the north. It includes t ...
n or more likely in the
Balkans The Balkans ( ), also known as the Balkan Peninsula, is a geographical area in southeastern Europe with various geographical and historical definitions. The region takes its name from the Balkan Mountains that stretch throughout the who ...
region. Probably around 1
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
during the
Pre-Pastonian Stage The Pre-Pastonian Stage or Baventian Stage (from Easton Bavents in Suffolk), is the name for an early Pleistocene stage used in the British Isles. It precedes the Pastonian Stage and follows the Bramertonian Stage. This stage ended 1.806 Ma (millio ...
, the original population expanded north. During an
interstadial Stadials and interstadials are phases dividing the Quaternary period, or the last 2.6 million years. Stadials are periods of colder climate while interstadials are periods of warmer climate. Each Quaternary climate phase is associated with a Ma ...
, the southern montane
metapopulation A metapopulation consists of a group of spatially separated populations of the same species which interact at some level. The term metapopulation was coined by Richard Levins in 1969 to describe a model of population dynamics of insect pests in ...
and the one to the north which ranged across the
Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...
n
taiga Taiga (; rus, тайга́, p=tɐjˈɡa; relates to Mongolic and Turkic languages), generally referred to in North America as a boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruce ...
split. Two major southern populations were established some 800,000-700,000 years ago during the
Pastonian Stage The Pastonian interglacial, now called the Pastonian Stage (from Paston, Norfolk), is the name for an early or middle Pleistocene stage used in the British Isles. It precedes the Beestonian Stage and follows the Pre-Pastonian Stage. Unfortunately t ...
, when the habitat belt moved uphill, cutting off
gene flow In population genetics, gene flow (also known as gene migration or geneflow and allele flow) is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent a ...
between major mountain ranges. Coincident with the
Günz-Mindel interglacial The Cromerian Stage or Cromerian Complex, also called the Cromerian (german: Cromerium), is a stage in the Pleistocene glacial history of north-western Europe, mostly occurring more than half a million years ago. It is named after the East Anglian t ...
, about 600,000-500,000 years ago some more distinct local populations diverged in the south. Meanwhile, the northern population had been spreading across Europe to the Pyrenees and deep into
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive 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 part of ...
. With the ice – which had then been covering much of northern and central Europe – retreating at the end of the
Elsterian Stage The Elster glaciation (german: Elster-Kaltzeit, ''Elster-Glazial'' or ''Elster-Zeit'') or, less commonly, the Elsterian glaciation, in the older and popular scientific literature also called the Elster Ice Age (''Elster-Eiszeit''), is the oldest k ...
between 400,000 and 300,000 years ago, the taiga population and their relatives in the mountains of central and western Europe became isolated for good. Finally, at the end of the Riss/Saale glaciation about 130,000 years ago, Alpine and Siberian populations fragmented further. The former stayed rather distinct while more subsequent
gene flow In population genetics, gene flow (also known as gene migration or geneflow and allele flow) is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent a ...
occurred between the latter; this difference is probably an effect of habitat
topography Topography is the study of the forms and features of land surfaces. The topography of an area may refer to the land forms and features themselves, or a description or depiction in maps. Topography is a field of geoscience and planetary sci ...
. A peculiar case is the Colorado alpine, the
nominate subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
of ''
Erebia callias ''Erebia callias'', the Colorado alpine, is a member of the Satyridae subfamily of the Nymphalidae butterflies. It is found in alpine areas of Wyoming and Colorado in the U.S. Rocky Mountains as well as various mountain ranges in eastern Asia. ...
''. This isolated
Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in straight-line distance from the northernmost part of western Canada, to New Mexico in ...
population has been lumped with the Siberian brassy ringlets as they are almost alike morphologically. Though one might suspect stronger differentiation and perhaps marked cryptic speciation across the wide range, the Rocky Mountains population is apparently a very recent isolate. Its ancestors apparently crossed over the Bering Strait at the end of the Wisconsinian glaciation, about 15,000 to 10,000 years ago. Thus, brassy ringlets are present on the North American continent quite exactly for the same length of time as a significant human population.


Systematics and taxonomy

Occasionally, as few as three species of brassy ringlets are accepted, or these are treated as superspecies. ''E. tyndarus'' is used for populations from 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 C ...
and
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, Sw ...
, and '' E. ottomana'' for those of the
Balkans The Balkans ( ), also known as the Balkan Peninsula, is a geographical area in southeastern Europe with various geographical and historical definitions. The region takes its name from the Balkan Mountains that stretch throughout the who ...
and the
Caucasus The Caucasus () or Caucasia (), is a region between the Black Sea and the Caspian Sea, mainly comprising Armenia, Azerbaijan, Georgia, and parts of Southern Russia. The Caucasus Mountains, including the Greater Caucasus range, have historically ...
. However, in this case the Caucasus population would also seem to warrant recognition at the same taxonomic rank as the others, as '' E. iranica''. In addition, the related butterflies around
Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72 degrees north latitude in the Chukchi Sea; and on the south by the tip ...
have been united in the '' E. callias'' superspecies. Therefore, though ''E. callias'' might be split up on grounds of
biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
and
ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...
, the molecular and morphological data are still more in favor of retaining a single species. Based on analyses of mtDNA NADH dehydrogenase
subunit Subunit may refer to: *Subunit HIV vaccine, a class of HIV vaccine *Protein subunit, a protein molecule that assembles with other protein molecules *Monomer, a molecule that may bind chemically to other molecules to form a polymer *Sub-subunit, a ...
5 and CoxII
sequence In mathematics, a sequence is an enumerated collection of objects in which repetitions are allowed and order matters. Like a set, it contains members (also called ''elements'', or ''terms''). The number of elements (possibly infinite) is calle ...
, ecological, and morphological data, the following species and well-distinct subspecies ought to be recognized: * ''
Erebia calcaria ''Erebia calcaria'', or Lorkovic's brassy ringlet, is a butterfly of the family Nymphalidae. It is found in the Alps. The species inhabits southern exposed slopes with alpine grassland interspersed with rocks. Screes without vegetation or only a ...
'' Lorkovic, 1949 – Lorkovic's brassy ringlet * ''
Erebia callias ''Erebia callias'', the Colorado alpine, is a member of the Satyridae subfamily of the Nymphalidae butterflies. It is found in alpine areas of Wyoming and Colorado in the U.S. Rocky Mountains as well as various mountain ranges in eastern Asia. ...
'' Edwards, 1871 ** ''Erebia'' (''callias'') ''altajana'' Staudinger, 1901 ** ''Erebia'' (''callias'') ''callias'' – Colorado alpine ** ''Erebia'' (''callias'') ''sibirica'' Staudinger, 1881 ** ''Erebia'' (''callias'') ''simulata'' Warren, 1933 * '' Erebia cassioides'' (Reiner & Hohenwarth, 1792) – Common brassy ringlet ** ''Erebia'' (''cassioides'') ''arvernensis'' Oberthür 1908 – western brassy ringlet ** ''Erebia'' (''cassioides'') ''carmenta'' Fruhstorfer, 1907 – western brassy ringlet ** ''Erebia'' (''cassioides'') ''macedonica'' Buresch, 1918 * ''
Erebia graucasica ''Erebia'' is a Holarctic genus of brush-footed butterflies, family Nymphalidae. Most of the about 90–100 species (see also below) are dark brown or black in color, with reddish-brown to orange or more rarely yellowish wing blotches or ban ...
'' Jachontov, 1909 * ''
Erebia hispania ''Erebia hispania'', the Spanish brassy ringlet, is a species of butterfly of the family Nymphalidae, endemic to the Sierra Nevada of southern Spain. The taxon ''rondoui'' (from the Pyrenees), previously considered as a subspecies of ''Erebia hi ...
'' Butler, 1868 – Spanish brassy ringlet * '' Erebia iranica'' Grum-Grshimailo, 1895 * '' Erebia nivalis'' Lorkovic & Lesse, 1954 – de Lesse's brassy ringlet * ''
Erebia ottomana ''Erebia ottomana'' is a small butterfly found in the East Palearctic (Greece, Bulgaria, Balkans, Alps, Asia Minor) that belongs to the browns family. Description from Seitz ''Erebia ottomana'' H.-Schaff. (37 i) is the largest of all the '' ...
'' Herrich-Schäffer,
851 __NOTOC__ Year 851 ( DCCCLI) was a common year starting on Thursday (link will display the full calendar) of the Julian calendar. Events By place Asia * Bagrat II Bagratuni, Armenian prince and leader of a rebellion against the Abbasi ...
/small> – Ottoman brassy ringlet ** ''Erebia'' (''ottomana'') ''benacensis'' Warren 1933 * ''
Erebia rondoui ''Erebia rondoui'', the Pyrenees brassy ringlet, is a species of butterfly in the family Nymphalidae. It is found in France and Spain,Erebia transcaucasica'' Warren, 1950 (previously in ''E. graucasica'') * '' Erebia tyndarus'' – Swiss brassy ringlet


See also

*
Species concept The species problem is the set of questions that arises when biologists attempt to define what a species is. Such a definition is called a species concept; there are at least 26 recognized species concepts. A species concept that works well for se ...
*
Species problem The species problem is the set of questions that arises when biologists attempt to define what a species is. Such a definition is called a species concept; there are at least 26 recognized species concepts. A species concept that works well for se ...


Footnotes


References

* (2008): Molecular phylogeny of the ''Erebia tyndarus'' (Lepidoptera, Rhopalocera, Nymphalidae, Satyrinae) species group combining CoxII and ND5 mitochondrial genes: A case study of a recent radiation. '' Molecular Phylogenetics and Evolution'' 47(1): 196–210. (HTML abstract) * (1936): ''Monograph of the genus'' Erebia. British Museum of Natural History, London. {{DEFAULTSORT:Brassy Ringlet