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''Linnaea borealis'' is a
species In biology, a species is the basic unit of Taxonomy (biology), 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 ...
of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. They include all forbs (flowering plants without a woody stem), grasses and grass-like plants, a vast majority of ...
in the family Caprifoliaceae (the honeysuckle family). Until 2013, it was the only species in the genus '' Linnaea''. It is a boreal to subarctic woodland subshrub, commonly known as twinflower (sometimes written twin flower). This plant was a favorite of
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...
, founder of the modern system of
binomial nomenclature In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, b ...
, for whom the genus was named.


Description

The perennial stems of ''Linnaea borealis'' are slender, pubescent, and prostrate, growing to long, with opposite
evergreen In botany, an evergreen is a plant which has foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, which ...
rounded oval leaves long and broad. The flowering stems curve erect, to tall, and are leafless except at the base. The
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanism ...
s are paired, pendulous, long, with a five-lobed, pale pink corolla.


Taxonomy

''Linnaea borealis'' was first formally described by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...
in 1753 in ''
Species Plantarum ' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genera. It is the first work to consistently apply binomial names and was the ...
''. It was then the sole species in the genus '' Linnaea''. The genus name had been used earlier by the Dutch botanist Jan Frederik Gronovius, and was given in honour of Linnaeus. Linnaeus adopted the name because ''Linnaea borealis'' was his favourite plant. ''Linnaea borealis'' is considered to be a single
circumboreal The Circumboreal Region in phytogeography is a floristic region within the Holarctic Kingdom in Eurasia and North America, as delineated by such geobotanists as Josias Braun-Blanquet and Armen Takhtajan. It is the largest floristic region i ...
species, with three generally recognized subspecies: *''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 ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S ...
to
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the ...
) The English name "twinflower" for ''Linnaea borealis'' refers to the plant's paired flowers.


Ecology and distribution

''Linnaea borealis'' has a circumpolar distribution in moist subarctic, boreal, or cool temperate
forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological function. The United Nations' ...
s, extending further south at higher elevations in various
mountain A mountain is an elevated portion of the Earth's crust, generally with steep sides that show significant exposed bedrock. Although definitions vary, a mountain may differ from a plateau in having a limited summit area, and is usually higher t ...
s, in
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located enti ...
south to the
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, ...
, in
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an ...
south to northern Japan, and in North America south to northern
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the ...
and to
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a U.S. state, state in the Southwestern United States. It is the list of U.S. states and territories by area, 6th largest and the list of U.S. states and territories by population, 14 ...
and
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe, New Mexico, Santa Fe , LargestCity = Albuquerque, New Mexico, Albuquerque , LargestMetro = Albuquerque metropolitan area, Tiguex , Offi ...
in the west, and to
West Virginia West Virginia is a state in the Appalachian, Mid-Atlantic and Southeastern regions of the United States.The Census Bureau and the Association of American Geographers classify West Virginia as part of the Southern United States while the ...
(and formerly
Tennessee Tennessee ( , ), officially the State of Tennessee, is a landlocked state in the Southeastern region of the United States. Tennessee is the 36th-largest by area and the 15th-most populous of the 50 states. It is bordered by Kentucky to ...
) in the
Appalachian Mountains The Appalachian Mountains, often called the Appalachians, (french: Appalaches), are a system of mountains in eastern to northeastern North America. The Appalachians first formed roughly 480 million years ago during the Ordovician Period. The ...
in the east. ''Linnaea borealis'' is self-incompatible, requiring cross-pollination to produce viable
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
s; since
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...
dispersal is usually not far, individuals and clonal colonies can become reproductively isolated. Regardless of seed production, ''Linnaea'' plants in a particular area often spread by stolons to form clonal patches of the same
genotype The genotype of an organism is its complete set of genetic material. Genotype can also be used to refer to the alleles or variants an individual carries in a particular gene or genetic location. The number of alleles an individual can have in a ...
. Such clonal stands of ''Linnaea'' can be long-persisting, in some places remaining extant even if seed is not produced or if seedling germination or establishment does not occur. The species was presumably common in areas south of its present range during times of
Pleistocene The Pleistocene ( , often referred to as the '' Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed ...
("Ice Age") glaciations, 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 slopes comprise a very rare, fragile ecosystem and habitat initially stated to exist only in the Driftless Area of Minnesota, Wisconsin, Illinois, and especially, Iowa.+Algific Taiga and boreal forests in the United States Ecore ...
s with persisting underground periglacial ice.


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 sites. In
Great Britain Great Britain is an island in the North Atlantic Ocean off the northwest coast of continental Europe. With an area of , it is the largest of the British Isles, the largest European island and the ninth-largest island in the world. It is ...
, ''Linnaea borealis'' ssp. ''borealis'' is listed as "nationally scarce", growing mainly in open pine woodlands in
Scotland Scotland (, ) is a country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a border with England to the southeast and is otherwise surrounded by the Atlantic Ocean to th ...
and northernmost
England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe ...
.
Forester A forester is a person who practises forestry, the science, art, and profession of managing forests. Foresters engage in a broad range of activities including ecological restoration and management of protected areas. Foresters manage forests ...
s consider this plant to be an indicator species of ancient woodlands, often found in association with
creeping lady's tresses ''Goodyera repens'', an orchid in the genus '' Goodyera'', is called by the common name creeping lady's-tresses in Anglophone Europe and dwarf rattlesnake plantain or lesser rattlesnake plantain in North America. It is a green underground creepe ...
. It is found in about 50 sites around the country, with most situated in the woods around the Cairngorms; the southernmost locations are four sites in
Northumberland Northumberland () is a ceremonial counties of England, county in Northern England, one of two counties in England which border with Scotland. Notable landmarks in the county include Alnwick Castle, Bamburgh Castle, Hadrian's Wall and Hexham Ab ...
and one in County Durham. The sparseness of the sites is responsible for the continued decline of the plant in the country. In
Scotland Scotland (, ) is a country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a border with England to the southeast and is otherwise surrounded by the Atlantic Ocean to th ...
, 37% of ''L. borealis'' patches studied consisted of a single
genotype The genotype of an organism is its complete set of genetic material. Genotype can also be used to refer to the alleles or variants an individual carries in a particular gene or genetic location. The number of alleles an individual can have in a ...
, reproducing clonally vegetatively 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 The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 U.S. state, states, a Washington, D.C., federal district, five ma ...
, ''Linnaea borealis'' ssp. ''americana'' is of conservation concern in several states along or near the southern edge of the species' range, including
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a U.S. state, state in the Southwestern United States. It is the list of U.S. states and territories by area, 6th largest and the list of U.S. states and territories by population, 14 ...
,
Iowa Iowa () is a U.S. state, state in the Midwestern United States, Midwestern region of the United States, bordered by the Mississippi River to the east and the Missouri River and Big Sioux River to the west. It is bordered by six states: Wiscon ...
,
Massachusetts Massachusetts (Massachusett language, Massachusett: ''Muhsachuweesut assachusett writing systems, məhswatʃəwiːsət'' English: , ), officially the Commonwealth of Massachusetts, is the most populous U.S. state, state in the New England ...
,
North Dakota North Dakota () is a U.S. state in the Upper Midwest, named after the indigenous Dakota Sioux. North Dakota is bordered by the Canadian provinces of Saskatchewan and Manitoba to the north and by the U.S. states of Minnesota to the east, S ...
,
Pennsylvania Pennsylvania (; (Pennsylvania Dutch: )), officially the Commonwealth of Pennsylvania, is a state spanning the Mid-Atlantic, Northeastern, Appalachian, and Great Lakes regions of the United States. It borders Delaware to its southeast, Ma ...
, and
West Virginia West Virginia is a state in the Appalachian, Mid-Atlantic and Southeastern regions of the United States.The Census Bureau and the Association of American Geographers classify West Virginia as part of the Southern United States while the ...
, and was known historically but now considered extirpated or possibly so in
Illinois Illinois ( ) is a state in the Midwestern United States. Its largest metropolitan areas include the Chicago metropolitan area, and the Metro East section, of Greater St. Louis. Other smaller metropolitan areas include, Peoria and Roc ...
,
Indiana Indiana () is a U.S. state in the Midwestern United States. It is the 38th-largest by area and the 17th-most populous of the 50 States. Its capital and largest city is Indianapolis. Indiana was admitted to the United States as the 19th ...
,
New Jersey New Jersey is a U.S. state, state in the Mid-Atlantic States, Mid-Atlantic and Northeastern United States, Northeastern regions of the United States. It is bordered on the north and east by the state of New York (state), New York; on the ea ...
,
Ohio Ohio () is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. Of the List of states and territories of the United States, fifty U.S. states, it is the List of U.S. states and territories by area, 34th-l ...
,
Rhode Island Rhode Island (, like ''road'') is a state in the New England region of the Northeastern United States. It is the smallest U.S. state by area and the seventh-least populous, with slightly fewer than 1.1 million residents as of 2020, but i ...
, and
Tennessee Tennessee ( , ), officially the State of Tennessee, is a landlocked state in the Southeastern region of the United States. Tennessee is the 36th-largest by area and the 15th-most populous of the 50 states. It is bordered by Kentucky to ...
. In
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tota ...
, ''Linnaea borealis'' ssp. ''longiflora'' is considered of conservation significance in the
Yukon Yukon (; ; formerly called Yukon Territory and also referred to as the Yukon) is the smallest and westernmost of Canada's three territories. It also is the second-least populated province or territory in Canada, with a population of 43,964 as ...
Territory, 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 In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
has become a significant concern for the conservation of this species in such places, such as Ice Mountain in West Virginia, a low-elevation
algific talus slope Algific talus slopes comprise a very rare, fragile ecosystem and habitat initially stated to exist only in the Driftless Area of Minnesota, Wisconsin, Illinois, and especially, Iowa.+Algific Taiga and boreal forests in the United States Ecore ...
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's name ''Linnaea'' as an example to advocate the use of commemorative personal names as botanical names: The flower of ''Linnaea borealis'' is the provincial flower of
Småland Småland () is a historical province () in southern Sweden. Småland borders Blekinge, Scania, Halland, Västergötland, Östergötland and the island Öland in the Baltic Sea. The name Småland literally means ''Small Lands''. The Latinized ...
,Visit Småland
Retrieved 25 February 2012.
the home province of Linnaeus.


References

*''The
Linnaeus Link Project Linnaeus Link is an international collaboration between libraries with significant holdings of material by or relating to Carl Linnaeus (1707–1778), his students, and his legacy. Aims and objectives The project's main aim is to provide a com ...
'' in the spring 2005 edition of ''Nature First'', the magazine for
Natural History Museum A natural history museum or museum of natural history is a scientific institution with natural history collections that include current and historical records of animals, plants, fungi, ecosystems, geology, paleontology, climatology, and more ...
members. *''Species and habitat conservation'' from Plantlife.org.u

an

* Twinflower species profil


External links

* {{Use British English, date=July 2020 Caprifoliaceae Alpine flora Flora of Europe Flora of North America Commemoration of Carl Linnaeus