Twinflower
Encyclopedia
Linnaea is a plant 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...

 in the family Caprifoliaceae (the Honeysuckle family)
Caprifoliaceae
The Caprifoliaceae or honeysuckle family is a clade consisting of about 800 dicotyledonous flowering plants, with a nearly cosmopolitan distribution; centres of diversity are found in eastern North America and eastern Asia, while they are absent in tropical and southern Africa.They are mostly...

. The genus includes a single, generally boreal
Boreal
Boreal may refer to*Boreal ecosystem**Boreal climate, the climate found in a region of boreal forests, and designated Dfc, Dwc or Dsc in the Köppen climate classification scheme.*boreal forest*Boreal forest of Canada*Boreal Bluet...

 to subarctic
Subarctic
The Subarctic is a region in the Northern Hemisphere immediately south of the true Arctic and covering much of Alaska, Canada, the north of Scandinavia, Siberia, and northern Mongolia...

 woodland subshrub
Subshrub
A subshrub or dwarf shrub is a short woody plant. Prostrate shrub is a similar term.It is distinguished from a shrub by its ground-hugging stems and lower height, with overwintering perennial woody growth typically less than 10–20 cm tall, or by being only weakly woody and/or persisting...

 species, Linnaea borealis, commonly known as Twinflower (sometimes written Twin Flower).

This plant was a favorite of Carl Linnaeus, founder of the modern system of biological nomenclature, for whom it was named.

Classification

The genus Linnaea is now considered to include only a single, circumboreal species, Linnaea borealis, with three subspecies generally recognized:
  • Linnaea borealis subsp. borealis - Europe
  • Linnaea borealis subsp. americana - North America (formerly classified as the species Linnaea americana)
  • Linnaea borealis subsp. longiflora - Asia, and western North America (from Alaska
    Alaska
    Alaska is the largest state in the United States by area. It is situated in the northwest extremity of the North American continent, with Canada to the east, the Arctic Ocean to the north, and the Pacific Ocean to the west and south, with Russia further west across the Bering Strait...

     to California
    California
    California is a state located on the West Coast of the United States. It is by far the most populous U.S. state, and the third-largest by land area...

    )


The genus Linnaea is usually treated in the Caprifoliaceae
Caprifoliaceae
The Caprifoliaceae or honeysuckle family is a clade consisting of about 800 dicotyledonous flowering plants, with a nearly cosmopolitan distribution; centres of diversity are found in eastern North America and eastern Asia, while they are absent in tropical and southern Africa.They are mostly...

, along with Lonicera, Symphoricarpos
Symphoricarpos
Symphoricarpos, with common names in English of Snowberry, Waxberry or Ghostberry, is a small genus of about 15 species of deciduous shrubs in the honeysuckle family, Caprifoliaceae. All species are natives of North and Central America, except one native to western China...

, and related genera, but sometimes classified in its own family Linnaeaceae, along with such similar genera as Abelia
Abelia
Abelia is a genus of about 15-30 species and many hybrids in the honeysuckle family Caprifoliaceae Some authors, including the Angiosperm Phylogeny Group, considers Abelia and related genera to belong instead in the segregate family Linnaeaceae, also including such genera as Linnaea, Abelia,...

, Dipelta, Kolkwitzia, and Zabelia, but not Lonicera or Symphoricarpos.

Nomenclature

Linnaea is one of the few kinds of plants or animals to be named after Carl Linnaeus, who knew it well from his explorations of Lapland
Lapland
Sápmi is the name of the cultural region traditionally inhabited by the Sami people. Sápmi is located in Northern Europe and includes the northern parts of Fennoscandia. The region stretches over four countries: Norway, Sweden, Finland, and Russia. Sápmi is the name in North Sámi, while the Julev...

 in northern 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....

 in 1732. It was his favorite plant, and as young man, he named it "Linnæa" (now transliterated as "Linnaea"). However, in his (first published in 1735), Linnaeus instead called this plant "Rudbeckia", for Olaus Rudbeck
Olaus Rudbeck
Olaus Rudbeck was a Swedish scientist and writer, professor of medicine at Uppsala University and for several periods rector magnificus of the same university...

 and his son Olof Rudbeck
Olof Rudbeck the Younger
Olaus Rudbeckius, junior or Olof Rudbeck , Swedish explorer and scientist, son of Olaus Rudbeck Sr, Rudbeck Jr. succeeded his father as professor of medicine at Uppsala University. Young Rudbeck was an able botanist and ornithologist who took his doctor's degree in Utrecht in 1690...

, Lapland explorers who knew the species. (Later, in his of 1753, Linnaeus used the genus name Rudbeckia
Rudbeckia
Rudbeckia is a plant genus of 23 species in the family Asteraceae. The species are commonly called coneflowers and black-eyed-susans; all are native to North America and many species are cultivated in gardens for their showy yellow or gold flower heads.The species are herbaceous, mostly perennial...

instead for a quite different plant in the Asteraceae
Asteraceae
The Asteraceae or Compositae , is an exceedingly large and widespread family of vascular plants. The group has more than 22,750 currently accepted species, spread across 1620 genera and 12 subfamilies...

, still known by that name).

The genus name Linnaea was first published by Linnaeus' teacher Jan Frederik Gronovius
Jan Frederik Gronovius
Jan Frederik Gronovius was a Dutch botanist notable as a patron of Linnaeus....

, with the name formalized under the modern system of botanical nomenclature
Botanical nomenclature
Botanical nomenclature is the formal, scientific naming of plants. It is related to, but distinct from taxonomy. Plant taxonomy is concerned with grouping and classifying plants; botanical nomenclature then provides names for the results of this process. The starting point for modern botanical...

 by Linnaeus himself in 1753 in his , where Linnaeus also formally described and named the species Linnaea borealis, with the epithet "" referring to its northern distributon.

The common name "twinflower" for Linnaea borealis refers to the plant's paired flowers.

Description

The perennial
Perennial plant
A perennial plant or simply perennial is a plant that lives for more than two years. The term is often used to differentiate a plant from shorter lived annuals and biennials. The term is sometimes misused by commercial gardeners or horticulturalists to describe only herbaceous perennials...

 stems of Linnaea are slender, pubescent, and prostrate, growing to 20 -0 long, with opposite evergreen
Evergreen
In botany, an evergreen plant is a plant that has leaves in all seasons. This contrasts with deciduous plants, which completely lose their foliage during the winter or dry season.There are many different kinds of evergreen plants, both trees and shrubs...

 rounded oval leaves
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

 3–10 mm (0.118110236220472–0.393700787401575 ) long and 2 millimetre broad. The flowering stems curve erect, to 4 centimetre tall, and are leafless except at the base. The flower
Flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...

s are paired, pendulous, 7 millimetre long, with a five-lobed, pale pink corolla.

Ecology and distribution

Linnaea borealis has a circumpolar
Circumpolar
The term circumpolar may refer to:* circumpolar navigation: to travel the world "vertically" traversing both of the poles* Antarctic region** Antarctic Circle** the Antarctic Circumpolar Current** Subantarctic** List of Antarctic and subantarctic islands...

 distribution in moist subarctic, boreal, or cool temperate forest
Forest
A forest, also referred to as a wood or the woods, is an area with a high density of trees. As with cities, depending where you are in the world, what is considered a forest may vary significantly in size and have various classification according to how and what of the forest is composed...

s, extending further south at higher elevations in various mountain
Mountain
Image:Himalaya_annotated.jpg|thumb|right|The Himalayan mountain range with Mount Everestrect 58 14 160 49 Chomo Lonzorect 200 28 335 52 Makalurect 378 24 566 45 Mount Everestrect 188 581 920 656 Tibetan Plateaurect 250 406 340 427 Rong River...

s, in Europe
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...

 south to 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....

, in Asia
Asia
Asia is the world's largest and most populous continent, located primarily in the eastern and northern hemispheres. It covers 8.7% of the Earth's total surface area and with approximately 3.879 billion people, it hosts 60% of the world's current human population...

 south to northern 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...

, and in North America south to northern California
California
California is a state located on the West Coast of the United States. It is by far the most populous U.S. state, and the third-largest by land area...

 and to Arizona
Arizona
Arizona ; is a state located in the southwestern region of the United States. It is also part of the western United States and the mountain west. The capital and largest city is Phoenix...

 and New Mexico
New Mexico
New Mexico is a state located in the southwest and western regions of the United States. New Mexico is also usually considered one of the Mountain States. With a population density of 16 per square mile, New Mexico is the sixth-most sparsely inhabited U.S...

 in the west, and to West Virginia
West Virginia
West Virginia is a state in the Appalachian and Southeastern regions of the United States, bordered by Virginia to the southeast, Kentucky to the southwest, Ohio to the northwest, Pennsylvania to the northeast and Maryland to the east...

 (and formerly Tennessee
Tennessee
Tennessee is a U.S. state located in the Southeastern United States. It has a population of 6,346,105, making it the nation's 17th-largest state by population, and covers , making it the 36th-largest by total land area...

) in the Appalachian Mountains
Appalachian Mountains
The Appalachian Mountains #Whether the stressed vowel is or ,#Whether the "ch" is pronounced as a fricative or an affricate , and#Whether the final vowel is the monophthong or the diphthong .), often called the Appalachians, are a system of mountains in eastern North America. The Appalachians...

 in the east.

Linnaea borealis is self-incompatible
Self-incompatibility in plants
Self-incompatibility is a general name for several genetic mechanisms in angiosperms, which prevent self-fertilization and thus encourage outcrossing...

, requiring cross-pollination
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...

 to produce viable seed
Seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...

s; since pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

 dispersal is usually not far, genetic individuals can become reproductively isolated. Regardless of seed production, Linnaea plants in a particular area often spread by stolon
Stolon
In biology, stolons are horizontal connections between organisms. They may be part of the organism, or of its skeleton; typically, animal stolons are external skeletons.-In botany:...

s to form clonal patches
Vegetative reproduction
Vegetative reproduction is a form of asexual reproduction in plants. It is a process by which new individuals arise without production of seeds or spores...

 of the same genotype
Genotype
The genotype is the genetic makeup of a cell, an organism, or an individual usually with reference to a specific character under consideration...

. Such clonal stands of Linnaea can be long-persisting, in some places remaining extant even if seed is not produced or if seedling
Seedling
thumb|Monocot and dicot seedlingsA seedling is a young plant sporophyte developing out of a plant embryo from a seed. Seedling development starts with germination of the seed. A typical young seedling consists of three main parts: the radicle , the hypocotyl , and the cotyledons...

 germination or establishment does not occur.

The species was presumably common in areas south of its present range during times of Pleistocene ("Ice Age") glaciations
Pleistocene
The Pleistocene is the epoch from 2,588,000 to 11,700 years BP that spans the world's recent period of repeated glaciations. The name pleistocene is derived from the Greek and ....

, and its clone-forming perennial growth habit has allowed it to survive the subsequent millennia locally within this former range in various high-elevation or otherwise cool and moist habitats, including algific talus slope
Algific talus slope
Algific talus slopes comprise a very rare, fragile ecosystem and habitat stated to exist only in the Driftless Area of Minnesota, Wisconsin, Illinois, and especially, Iowa...

s with persisting underground periglacial ice
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...

.

Conservation

While the three subspecies of Linnaea borealis are all considered widespread, abundant, and secure in their main, northern ranges, all three subspecies are of conservation concern near the subspecies' range edges or at more southerly, disjunct
Disjunct distribution
In biology, a taxon with a disjunct distribution is one that has two or more groups that are related but widely separated from each other geographically...

 sites.

In Great Britain
Great Britain
Great Britain or Britain is an island situated to the northwest of Continental Europe. It is the ninth largest island in the world, and the largest European island, as well as the largest of the British Isles...

, Linnaea borealis ssp. borealis is listed as "nationally scarce", growing mainly in open pine woodlands
Caledonian Forest
The Caledonian Forest is the name of a type of woodland that once covered vast areas of Scotland. Today, however, only 1% of the original forest survives, covering in 84 locations. The forests are home to a wide variety of wildlife, much of which is not found elsewhere in the British...

 in Scotland
Scotland
Scotland is a country that is part of the United Kingdom. Occupying the northern third of the island of Great Britain, it shares a border with England to the south and is bounded by the North Sea to the east, the Atlantic Ocean to the north and west, and the North Channel and Irish Sea to the...

 and northernmost 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...

. Forester
Forester
250px|thumb|right|Foresters of [[Southern University of Chile|UACh]] in the [[Valdivian forest]]s of San Pablo de Tregua, ChileA forester is a person who practices forestry, the science, art, and profession of managing forests. Foresters engage in a broad range of activities including timber...

s consider this plant to be an indicator species
Indicator species
An indicator species is any biological species that defines a trait or characteristic of the environment. For example, a species may delineate an ecoregion or indicate an environmental condition such as a disease outbreak, pollution, species competition or climate change...

 of ancient woodland
Ancient woodland
Ancient woodland is a term used in the United Kingdom to refer specifically to woodland that has existed continuously since 1600 or before in England and Wales . Before those dates, planting of new woodland was uncommon, so a wood present in 1600 was likely to have developed naturally...

s, often found in association with Creeping Lady's Tresses. It is found in about 50 sites around the country, with most situated in the woods around the Cairngorms
Cairngorms
The Cairngorms are a mountain range in the eastern Highlands of Scotland closely associated with the mountain of the same name - Cairn Gorm.-Name:...

; the southernmost locations are four sites in Northumberland
Northumberland
Northumberland is the northernmost ceremonial county and a unitary district in North East England. For Eurostat purposes Northumberland is a NUTS 3 region and is one of three boroughs or unitary districts that comprise the "Northumberland and Tyne and Wear" NUTS 2 region...

 and one in County Durham
County Durham
County Durham is a ceremonial county and unitary district in north east England. The county town is Durham. The largest settlement in the ceremonial county is the town of Darlington...

. The sparseness of the sites is responsible for the continued decline of the flower in the country. In Scotland
Scotland
Scotland is a country that is part of the United Kingdom. Occupying the northern third of the island of Great Britain, it shares a border with England to the south and is bounded by the North Sea to the east, the Atlantic Ocean to the north and west, and the North Channel and Irish Sea to the...

, 37% of L. borealis patches studied consisted of a single genotype
Genotype
The genotype is the genetic makeup of a cell, an organism, or an individual usually with reference to a specific character under consideration...

, reproducing clonally vegetatively
Vegetative reproduction
Vegetative reproduction is a form of asexual reproduction in plants. It is a process by which new individuals arise without production of seeds or spores...

 but not producing viable seed. This is a conservation concern because without viable seed, the species may not be able to re-populate restored habitat, and may not be able to adapt to climate change by establishing new populations.

In the United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...

, Linnaea borealis ssp. americana is of conservation concern in several states along or near the southern edge of the species' range, including Arizona
Arizona
Arizona ; is a state located in the southwestern region of the United States. It is also part of the western United States and the mountain west. The capital and largest city is Phoenix...

, Iowa
Iowa
Iowa is a state located in the Midwestern United States, an area often referred to as the "American Heartland". It derives its name from the Ioway people, one of the many American Indian tribes that occupied the state at the time of European exploration. Iowa was a part of the French colony of New...

, Massachusetts
Massachusetts
The Commonwealth of Massachusetts is a state in the New England region of the northeastern United States of America. It is bordered by Rhode Island and Connecticut to the south, New York to the west, and Vermont and New Hampshire to the north; at its east lies the Atlantic Ocean. As of the 2010...

, North Dakota
North Dakota
North Dakota is a state located in the Midwestern region of the United States of America, along the Canadian border. The state is bordered by Canada to the north, Minnesota to the east, South Dakota to the south and Montana to the west. North Dakota is the 19th-largest state by area in the U.S....

, Pennsylvania
Pennsylvania
The Commonwealth of Pennsylvania is a U.S. state that is located in the Northeastern and Mid-Atlantic regions of the United States. The state borders Delaware and Maryland to the south, West Virginia to the southwest, Ohio to the west, New York and Ontario, Canada, to the north, and New Jersey to...

, and West Virginia
West Virginia
West Virginia is a state in the Appalachian and Southeastern regions of the United States, bordered by Virginia to the southeast, Kentucky to the southwest, Ohio to the northwest, Pennsylvania to the northeast and Maryland to the east...

, and was known historically but now considered extirpated
Local extinction
Local extinction, also known as extirpation, is the condition of a species which ceases to exist in the chosen geographic area of study, though it still exists elsewhere...

 or possibly so in Illinois
Illinois
Illinois is the fifth-most populous state of the United States of America, and is often noted for being a microcosm of the entire country. With Chicago in the northeast, small industrial cities and great agricultural productivity in central and northern Illinois, and natural resources like coal,...

, Indiana
Indiana
Indiana is a US state, admitted to the United States as the 19th on December 11, 1816. It is located in the Midwestern United States and Great Lakes Region. With 6,483,802 residents, the state is ranked 15th in population and 16th in population density. Indiana is ranked 38th in land area and is...

, New Jersey
New Jersey
New Jersey is a state in the Northeastern and Middle Atlantic regions of the United States. , its population was 8,791,894. It is bordered on the north and east by the state of New York, on the southeast and south by the Atlantic Ocean, on the west by Pennsylvania and on the southwest by Delaware...

, Ohio
Ohio
Ohio is a Midwestern state in the United States. The 34th largest state by area in the U.S.,it is the 7th‑most populous with over 11.5 million residents, containing several major American cities and seven metropolitan areas with populations of 500,000 or more.The state's capital is Columbus...

, Rhode Island
Rhode Island
The state of Rhode Island and Providence Plantations, more commonly referred to as Rhode Island , is a state in the New England region of the United States. It is the smallest U.S. state by area...

, and Tennessee
Tennessee
Tennessee is a U.S. state located in the Southeastern United States. It has a population of 6,346,105, making it the nation's 17th-largest state by population, and covers , making it the 36th-largest by total land area...

.

In Canada
Canada
Canada is a North American country consisting of ten provinces and three territories. Located in the northern part of the continent, it extends from the Atlantic Ocean in the east to the Pacific Ocean in the west, and northward into the Arctic Ocean...

, Linnaea borealis ssp. longiflora is considered of conservation significance in the Yukon Territory
Yukon
Yukon is the westernmost and smallest of Canada's three federal territories. It was named after the Yukon River. The word Yukon means "Great River" in Gwich’in....

, along the eastern edge of its range, where ssp. americana is widespread and abundant.

Since many of the outlying southern sites for Linnaea borealis are in habitats that are at high elevations or otherwise in cooler microclimates than the surrounding general landscapes, ongoing and prospective climate change
Global warming
Global warming refers to the rising average temperature of Earth's atmosphere and oceans and its projected continuation. In the last 100 years, Earth's average surface temperature increased by about with about two thirds of the increase occurring over just the last three decades...

 has become a significant concern for the conservation of this species in such places, such as Ice Mountain
Ice Mountain
Ice Mountain is a mountain ridge and algific talus slope that is part of a preserve near the community of North River Mills in Hampshire County, West Virginia, United States....

 in West Virginia, a low-elevation algific talus slope
Algific talus slope
Algific talus slopes comprise a very rare, fragile ecosystem and habitat stated to exist only in the Driftless Area of Minnesota, Wisconsin, Illinois, and especially, Iowa...

 with persisting buried ice.

Cultural references

Linnaeus took Linnaea borealis as his own personal symbol when he was raised to the Swedish nobility in 1757. In his (1737), Linnaeus had used Gronovius' name Linnaea as an example to advocate the use of commemorative personal names as botanical names:
Linnéa or Linnea is a female given name
Given name
A given name, in Western contexts often referred to as a first name, is a personal name that specifies and differentiates between members of a group of individuals, especially in a family, all of whose members usually share the same family name...

 of Swedish
Swedish language
Swedish is a North Germanic language, spoken by approximately 10 million people, predominantly in Sweden and parts of Finland, especially along its coast and on the Åland islands. It is largely mutually intelligible with Norwegian and Danish...

 origin, derived either from the name of the Swedish botanist Carl Linnaeus or from the Linnaea plant, named in his honor.

The flower of Linnaea borealis is the provincial emblem of Småland
Småland
' is a historical province in southern Sweden.Småland borders Blekinge, Scania or Skåne, Halland, Västergötland, Östergötland and the island Öland in the Baltic Sea. The name Småland literally means Small Lands. . The latinized form Smolandia has been used in other languages...

 in 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....

, Linnaeus' home province.
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