Brassy ringlet
Encyclopedia
The brassy ringlets are a "species group" of ringlet butterflies in the genus Erebia
. Though closely related, their monophyly
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
habitat, like taiga
or regions with Alpine climate
. 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
s, and in glacials in one periglacial
ic belt at lower altitude, in places interrupted by dry wasteland and deserts.
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
-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 greyish-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 hindwind 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
y lustre of the dark wing uppersides. The valva
e have a series of external spines pointing distally, with the first spine pair usually being largest.
As it seems, the origin of this group is perhaps south of the Central Asia
n or more likely in the Balkans
region. Probably around 1 million years ago during the Pre-Pastonian Stage, the original population expanded north. During an interstadial, the southern montane
metapopulation
and the one to the north which ranged across the Eurasia
n taiga
split.
Two major southern populations were established some 800.000-700.000 years ago during the Pastonian Stage, when the habitat belt moved uphill, cutting off gene flow
between major mountain ranges. Coincident with the Günz-Mindel interglacial, 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
. With the ice – which had then been covering much of northern and central Europe – retreating at the end of the Elsterian Stage between 400.000-300.000 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
occurred between the latter; this difference is probably an effect of habitat topography
.
A peculiar case is the Colorado Alpine, the nominate subspecies of Erebia callias
. This isolated Rocky Mountains
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 15.000-10.000 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
.
. E. tyndarus is used for populations from the Pyrenees
and Alps
, and E. ottomana for those of the Balkans
and the Caucasus
. 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 have been united in the E. callias
superspecies. Therefore, though E. callias might be split up on grounds of biogeography
and ecology
, the molecualr and morphological data are still more in favor of retaining a single species.
Based on analyses of mtDNA NADH dehydrogenase
subunit
5 and CoxII sequence
, ecological, and morphological data, the following species and well-distinct subspecies ought to be recognized:
Erebia
Erebia is a Holarctic genus of brush-footed butterflies, family Nymphalidae. Most of the about 90–100 species are dark brown or black in color, with reddish brown to orange or more rarely yellowish wing blotches or bands...
. Though closely related, their monophyly
Monophyly
In common cladistic usage, a monophyletic group is a taxon which forms a clade, meaning that it contains all the descendants of the possibly hypothetical closest common ancestor of the members of the group. The term is synonymous with the uncommon term holophyly...
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
Arid
A region is said to be arid when it is characterized by a severe lack of available water, to the extent of hindering or even preventing the growth and development of plant and animal life...
habitat, like taiga
Taiga
Taiga , also known as the boreal forest, is a biome characterized by coniferous forests.Taiga is the world's largest terrestrial biome. In North America it covers most of inland Canada and Alaska as well as parts of the extreme northern continental United States and is known as the Northwoods...
or regions with Alpine climate
Alpine climate
Alpine climate is the average weather for a region above the tree line. This climate is also referred to as mountain climate or highland climate....
. 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
Alpide belt
The Alpide belt is a mountain range which extends along the southern margin of Eurasia. Stretching from Java to Sumatra through the Himalayas, the Mediterranean, and out into the Atlantic, it includes the Alps, the Carpathians, the mountains of Asia Minor and Iran, the Hindu Kush, the Himalayas,...
– in interglacial
Interglacial
An Interglacial period is a geological interval of warmer global average temperature lasting thousands of years that separates consecutive glacial periods within an ice age...
s, and in glacials in one periglacial
Periglacial
Periglacial is an adjective originally referring to places in the edges of glacial areas, but it has later been widely used in geomorphology to describe any place where geomorphic processes related to freezing of water occur...
ic belt at lower altitude, in places interrupted by dry wasteland and deserts.
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 eyespotsEyespot (mimicry)
An eyespot is an eye-like marking. They are found on butterflies, reptiles, birds and fish. In members of the Felidae family , the white circular markings on the backs of the ears are termed ocelli, and they are functionally similar to eyespots in other animals.Eyespots may be a form of...
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 (color)
Copper is a reddish brown color that resembles the metal copper.At right is displayed the color copper.The first recorded use of copper as a color name in English was in 1594.-Pale copper:...
-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
Crypsis
In ecology, crypsis is the ability of an organism to avoid observation or detection by other organisms. It may be either a predation strategy or an antipredator adaptation, and methods include camouflage, nocturnality, subterranean lifestyle, transparency, and mimicry...
colored, usually in greyish-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 hindwind 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
Brass is an alloy of copper and zinc; the proportions of zinc and copper can be varied to create a range of brasses with varying properties.In comparison, bronze is principally an alloy of copper and tin...
y lustre of the dark wing uppersides. The valva
Valva
In biology, a clasper is a male anatomical structure found some groups of animals, used in mating.Male cartilaginous fish have claspers formed from the posterior portion of their pelvic fin which serve as intromittent organs used to channel semen into the female's cloaca during mating...
e have a series of external spines pointing distally, 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
Central Asia is a core region of the Asian continent from the Caspian Sea in the west, China in the east, Afghanistan in the south, and Russia in the north...
n or more likely in the Balkans
Balkans
The Balkans is a geopolitical and cultural region of southeastern Europe...
region. Probably around 1 million years ago during the Pre-Pastonian Stage, the original population expanded north. During an interstadial, the southern montane
Montane
In biogeography, montane is the highland area located below the subalpine zone. Montane regions generally have cooler temperatures and often have higher rainfall than the adjacent lowland regions, and are frequently home to distinct communities of plants and animals.The term "montane" means "of the...
metapopulation
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 1970 to describe a model of population dynamics of insect pests in agricultural fields, but the idea has been most...
and the one to the north which ranged across the Eurasia
Eurasia
Eurasia is a continent or supercontinent comprising the traditional continents of Europe and Asia ; covering about 52,990,000 km2 or about 10.6% of the Earth's surface located primarily in the eastern and northern hemispheres...
n taiga
Taiga
Taiga , also known as the boreal forest, is a biome characterized by coniferous forests.Taiga is the world's largest terrestrial biome. In North America it covers most of inland Canada and Alaska as well as parts of the extreme northern continental United States and is known as the Northwoods...
split.
Two major southern populations were established some 800.000-700.000 years ago during the Pastonian Stage, when the habitat belt moved uphill, cutting off gene flow
Gene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...
between major mountain ranges. Coincident with the Günz-Mindel interglacial, 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
Siberia 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...
. With the ice – which had then been covering much of northern and central Europe – retreating at the end of the Elsterian Stage between 400.000-300.000 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
Gene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...
occurred between the latter; this difference is probably an effect of habitat topography
Topography
Topography is the study of Earth's surface shape and features or those ofplanets, moons, and asteroids...
.
A peculiar case is the Colorado Alpine, the nominate subspecies of Erebia callias
Erebia callias
The Colorado Alpine is a member of the Satyridae subfamily of Nymphalidae. It is found in alpine Wyoming and Colorado in the Rocky Mountains as well as various mountain ranges in eastern Asia....
. This isolated Rocky Mountains
Rocky Mountains
The Rocky Mountains are a major mountain range in western North America. The Rocky Mountains stretch more than from the northernmost part of British Columbia, in western Canada, to New Mexico, in the southwestern United States...
population has been lumped with the Siberian brassy ringlets as they are almost alike morphologically
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
. Though one might suspect stronger differentiation and perhaps marked cryptic speciation
Cryptic species complex
In biology, a cryptic species complex is a group of species which satisfy the biological definition of species—that is, they are reproductively isolated from each other—but whose morphology is very similar ....
across the wide range, the Rocky Mountains population is apparently a very recent isolate. Its ancestors apparently crossed over the Bering Strait
Bering Strait
The Bering Strait , known to natives as Imakpik, is a sea strait between Cape Dezhnev, Chukotka Autonomous Okrug, Russia, the easternmost point of the Asian continent and Cape Prince of Wales, Alaska, USA, the westernmost point of the North American continent, with latitude of about 65°40'N,...
at the end of the Wisconsinian glaciation, about 15.000-10.000 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
Models of migration to the New World
There have been several models for the human settlement of the Americas proposed by various academic communities. The question of how, when and why humans first entered the Americas is of intense interest to archaeologists and anthropologists, and has been a subject of heated debate for centuries...
.
Systematics and taxonomy
Occasionally, as few as three species of brassy ringlets are accepted, or these are treated as superspeciesSuperspecies
A superspecies is a group of at least two more or less distinctive species with approximately parapatric distributions. Not all species complexes, whether cryptices or ring species are superspecies, and vice versa, but many are...
. E. tyndarus is used for populations from the Pyrenees
Pyrenees
The Pyrenees is a range of mountains in southwest Europe that forms a natural border between France and Spain...
and Alps
Alps
The Alps is one of the great mountain range systems of Europe, stretching from Austria and Slovenia in the east through Italy, Switzerland, Liechtenstein and Germany to France in the west....
, and E. ottomana for those of the Balkans
Balkans
The Balkans is a geopolitical and cultural region of southeastern Europe...
and the Caucasus
Caucasus
The Caucasus, also Caucas or Caucasia , is a geopolitical region at the border of Europe and Asia, and situated between the Black and the Caspian sea...
. 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 have been united in the E. callias
Erebia callias
The Colorado Alpine is a member of the Satyridae subfamily of Nymphalidae. It is found in alpine Wyoming and Colorado in the Rocky Mountains as well as various mountain ranges in eastern Asia....
superspecies. Therefore, though E. callias might be split up on grounds of biogeography
Biogeography
Biogeography is the study of the distribution of species , organisms, and ecosystems in space and through geological time. Organisms and biological communities vary in a highly regular fashion along geographic gradients of latitude, elevation, isolation and habitat area...
and ecology
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...
, the molecualr and morphological data are still more in favor of retaining a single species.
Based on analyses of mtDNA NADH dehydrogenase
NADH dehydrogenase
NADH dehydrogenase is an enzyme located in the inner mitochondrial membrane that catalyzes the transfer of electrons from NADH to coenzyme Q...
subunit
Protein subunit
In structural biology, a protein subunit or subunit protein is a single protein molecule that assembles with other protein molecules to form a protein complex: a multimeric or oligomeric protein. Many naturally occurring proteins and enzymes are multimeric...
5 and CoxII sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...
, ecological, and morphological data, the following species and well-distinct subspecies ought to be recognized:
- Erebia calcaria Lorkovic, 1949 – Lorkovic's Brassy Ringlet
- Erebia calliasErebia calliasThe Colorado Alpine is a member of the Satyridae subfamily of Nymphalidae. It is found in alpine Wyoming and Colorado in the 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 cassioidesErebia cassioidesThe Common Brassy Ringlet is a member of the Satyrinae subfamily of Nymphalidae. It is found in the Balkans, the eastern and western Alps, central Italy, the Pyrenees and the Cantabrian mountains...
(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 Jachontov, 1909
- Erebia hispania Butler, 1868 – Spanish Brassy Ringlet
- Erebia iranica Grum-Grshimailo, 1895
- Erebia nivalisErebia nivalisde Lesse's Brassy Ringlet is a member of the Satyrinae subfamily of Nymphalidae. It is found throughout the Alps of southern Austria with a remote population in central Switzerland....
Lorkovic & Lesse, 1954 – de Lesse's Brassy Ringlet - Erebia ottomana Herrich-Schäffer, [1851] – Ottoman Brassy Ringlet
- Erebia (ottomana) benacensis Warren 1933
- Erebia rondoui Oberthür 1908 (previously in E. cassioides)
- Erebia transcaucasica Warren, 1950 (previously in E. graucasica)
- Erebia tyndarusErebia tyndarusThe Swiss Brassy Ringlet, Erebia tyndarus, is a European brush-footed butterfly species of the subfamily Satyrinae. It is found above the treeline, at heights of 1,200-2,700 meters ASL, in the Alps of Switzerland, Liechtenstein and Austrian Vorarlberg, and adjacent regions.-Description and...
– Swiss Brassy Ringlet