Erebia tyndarus
Encyclopedia
The Swiss Brassy Ringlet, Erebia tyndarus, is a Europe
an 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 Austria
n Vorarlberg
, and adjacent regions.
Erebia
has a wingspan around 35 mm. The blackish-brown upperwings typical of this genus; they have a brass
y sheen in this and related species, hence the name. The upper forewings, and sometimes the upper hindwings also, bear a coppery
patch, near the tip in the former and running parallel to the outer margin in the latter.
In this patch there are black eyespots
with a white dot in the middle, two immediately next to each other in the forewings and 3-4 more evenly spaced ones in the hindwings. However, the hindwing spots may be reduced or entirely absent, particularly in the males which also have a darker coloration overall. The underwings are crpytically
colored in lighter greyish brown. A darker band arches between the forward and back margins of the hindwing, which has a prominent silvery sheen also found in closely related species of Erebia and conspicuous at a distance when these butterflies fly around in the sun. The reddish patches and the eyespots of the forewing also occur on the underwings, those of the hindwings usually do not.
The adults are found only for a brief time during summer, mainly in July and August. The Swiss Brassy Ringlet is univoltine and its caterpillar
s feed on Poaceae
grass
es, especially Sheep's Fescue
(Festuca ovina), Mat-grass (Nardus stricta
), and various other fescue
s (Festuca) and meadow-grasses (Poa). They overwinter and pupate on the ground around May/June.
The Swiss Brassy Ringlet belongs to the brassy ringlet
group of its genus
Erebia
. These are found in taiga
and Alpine
habitat across Eurasia
, with one population in comparatively recent times colonizing North America
. The brassy ringlet group, like many Erebia, has a complicated taxonomy
, with scores of "variants" having been named in the late 19th and early 20th century. Most of these in fact were simply aberrations, local or seasonal forms. As this was realized, the brassy ringlets of the Alps
and Pyrenees
were united in E. tyndarus, either as a superspecies
or as a species
.
In recent times, this question has been restudied, including the DNA sequence
data available nowadays. The view that only a single species of this non-migratory butterfly with restricted habitat
preferences occurs in far-flung localities like the Sierra Nevada
of southern Spain
or the Julian Alps
of Slovenia
is generally considered obsolete.
Rather, it seems that quite a few proper species are involved. The Swiss Brassy Ringlet in particular is part of a cryptic species complex
from the Alps
. These diverged at the end of the Riss/Saale glaciation roughly 130,000 years ago, when the available habitat
shifted up the mountains. The ancestors of the Alpine brassy ringlets had lived in the lowlands when the Alps were entirely covered by ice, and as the ice retreated, their typical habitat
moved upwards, and the butterflies with them. Thus, different populations became isolated in different mountain ranges and with gene flow
between them restricted started to evolve
into distinct species.
The brassy ringlets from the Alps, the closest relatives of E. tyndarus, consist of three further species:
The last is presumably closest to the last common ancestors of the Alpine brassy ringlets.
Europe
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...
an brush-footed butterfly species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
of the subfamily Satyrinae
Satyrinae
Satyrinae, the satyrines or satyrids, commonly known as the Browns, is a subfamily of the Nymphalidae . They were formerly considered a distinct family, Satyridae. This group contains nearly half of the known diversity of brush-footed butterflies...
. It is found above the treeline, at heights of 1,200-2,700 meters ASL
Above mean sea level
The term above mean sea level refers to the elevation or altitude of any object, relative to the average sea level datum. AMSL is used extensively in radio by engineers to determine the coverage area a station will be able to reach...
, in the 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....
of Switzerland
Switzerland
Switzerland 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....
, Liechtenstein
Liechtenstein
The Principality of Liechtenstein is a doubly landlocked alpine country in Central Europe, bordered by Switzerland to the west and south and by Austria to the east. Its area is just over , and it has an estimated population of 35,000. Its capital is Vaduz. The biggest town is Schaan...
and Austria
Austria
Austria , officially the Republic of Austria , is a landlocked country of roughly 8.4 million people in Central Europe. It is bordered by the Czech Republic and Germany to the north, Slovakia and Hungary to the east, Slovenia and Italy to the south, and Switzerland and Liechtenstein to the...
n Vorarlberg
Vorarlberg
Vorarlberg is the westernmost federal-state of Austria. Although it is the second smallest in terms of area and population , it borders three countries: Germany , Switzerland and Liechtenstein...
, and adjacent regions.
Description and ecology
This mid-sized member of the genusGenus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
Erebia
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...
has a wingspan around 35 mm. The blackish-brown upperwings typical of this genus; they have a 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 sheen in this and related species, hence the name. The upper forewings, and sometimes the upper hindwings also, bear a coppery
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:...
patch, near the tip in the former and running parallel to the outer margin in the latter.
In this patch there are black eyespots
Eyespot (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...
with a white dot in the middle, two immediately next to each other in the forewings and 3-4 more evenly spaced ones in the hindwings. However, the hindwing spots may be reduced or entirely absent, particularly in the males which also have a darker coloration overall. The underwings are crpytically
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 in lighter greyish brown. A darker band arches between the forward and back margins of the hindwing, which has a prominent silvery sheen also found in closely related species of Erebia and conspicuous at a distance when these butterflies fly around in the sun. The reddish patches and the eyespots of the forewing also occur on the underwings, those of the hindwings usually do not.
The adults are found only for a brief time during summer, mainly in July and August. The Swiss Brassy Ringlet is univoltine and its 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 feed on Poaceae
Poaceae
The Poaceae is a large and nearly ubiquitous family of flowering plants. Members of this family are commonly called grasses, although the term "grass" is also applied to plants that are not in the Poaceae lineage, including the rushes and sedges...
grass
Grass
Grasses, or more technically graminoids, are monocotyledonous, usually herbaceous plants with narrow leaves growing from the base. They include the "true grasses", of the Poaceae family, as well as the sedges and the rushes . The true grasses include cereals, bamboo and the grasses of lawns ...
es, especially Sheep's Fescue
Sheep's Fescue
Sheep's Fescue or Sheep Fescue is a species of grass.-General Description:It is a perennial plant sometimes found in acidic ground, for example in the Portlethen Moss, Scotland and mountain pasture, throughout Europe and eastwards across much of Asia; it has also been introduced to North...
(Festuca ovina), Mat-grass (Nardus stricta
Nardus stricta
Nardus stricta is a densely tufted, tough, wiry perennial plant species belonging to the family Poaceae , and found throughout much of the world. It exists on heath, moorland, hills, mountains; on sandy to peaty soils...
), and various other fescue
Fescue
Festuce is a genus of about 300 species of perennial tufted grasses, belonging to the grass family Poaceae . The genus has a cosmopolitan distribution, although the majority of the species are found in cool temperate areas...
s (Festuca) and meadow-grasses (Poa). They overwinter and pupate on the ground around May/June.
Systematics, taxonomy and evolution
Brassy ringlet
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...
group of its genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
Erebia
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...
. These are found in 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...
and Alpine
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....
habitat across 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...
, with one population in comparatively recent times colonizing North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...
. The brassy ringlet group, like many Erebia, has a complicated taxonomy
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...
, with scores of "variants" having been named in the late 19th and early 20th century. Most of these in fact were simply aberrations, local or seasonal forms. As this was realized, the brassy ringlets of the 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 Pyrenees
Pyrenees
The Pyrenees is a range of mountains in southwest Europe that forms a natural border between France and Spain...
were united in E. tyndarus, either as a superspecies
Superspecies
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...
or as a species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
.
In recent times, this question has been restudied, including the DNA 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...
data available nowadays. The view that only a single species of this non-migratory butterfly with restricted habitat
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...
preferences occurs in far-flung localities like 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....
of southern 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...
or the Julian Alps
Julian Alps
The Julian Alps are a mountain range of the Southern Limestone Alps that stretches from northeastern Italy to Slovenia, where they rise to 2,864 m at Mount Triglav. They are named after Julius Caesar, who founded the municipium of Cividale del Friuli at the foot of the mountains...
of Slovenia
Slovenia
Slovenia , officially the Republic of Slovenia , is a country in Central and Southeastern Europe touching the Alps and bordering the Mediterranean. Slovenia borders Italy to the west, Croatia to the south and east, Hungary to the northeast, and Austria to the north, and also has a small portion of...
is generally considered obsolete.
Rather, it seems that quite a few proper species are involved. The Swiss Brassy Ringlet in particular is part of a cryptic species complex
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 ....
from the 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....
. These diverged at the end of the Riss/Saale glaciation roughly 130,000 years ago, when the available habitat
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...
shifted up the mountains. The ancestors of the Alpine brassy ringlets had lived in the lowlands when the Alps were entirely covered by ice, and as the ice retreated, their typical habitat
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...
moved upwards, and the butterflies with them. Thus, different populations became isolated in different mountain ranges and with 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 them restricted started to evolve
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...
into distinct species.
The brassy ringlets from the Alps, the closest relatives of E. tyndarus, consist of three further species:
- 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....
(de Lesse's Brassy Ringlet), found in the highest regions of the Central Eastern AlpsCentral Eastern AlpsThe Central Eastern Alps comprise the main chain of the Eastern Alps with its highest peaks, located between the Northern Limestone Alps and the Southern Limestone Alps, from which they differ in geological composition....
of AustriaAustriaAustria , officially the Republic of Austria , is a landlocked country of roughly 8.4 million people in Central Europe. It is bordered by the Czech Republic and Germany to the north, Slovakia and Hungary to the east, Slovenia and Italy to the south, and Switzerland and Liechtenstein to the... - Erebia calcaria (Lorkovic's Brassy Ringlet), from the Julian AlpsJulian AlpsThe Julian Alps are a mountain range of the Southern Limestone Alps that stretches from northeastern Italy to Slovenia, where they rise to 2,864 m at Mount Triglav. They are named after Julius Caesar, who founded the municipium of Cividale del Friuli at the foot of the mountains...
which straddle the borders of Austria, 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...
and SloveniaSloveniaSlovenia , officially the Republic of Slovenia , is a country in Central and Southeastern Europe touching the Alps and bordering the Mediterranean. Slovenia borders Italy to the west, Croatia to the south and east, Hungary to the northeast, and Austria to the north, and also has a small portion of... - 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...
(Common Brassy Ringlet), which occurs on meadows somewhat further downhill than the other species and consequently is widely distributed in the rest of the Alps.
The last is presumably closest to the last common ancestors of the Alpine brassy ringlets.