Linnaea borealis
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''Linnaea borealis'' is a
species In biology, a species is the basic unit of 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 the appropriate s ...
of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
in the family
Caprifoliaceae The Caprifoliaceae or honeysuckle family is a clade of dicotyledonous flowering plants consisting of about 860 species, in 33, to 42 genera, with a nearly cosmopolitan distribution. Centres of diversity are found in eastern North America an ...
(the honeysuckle family). Until 2013, it was the only species in the genus ''
Linnaea ''Linnaea'' is a plant genus in the honeysuckle family Caprifoliaceae. Until 2013, the genus included a single species, ''Linnaea borealis''. In 2013, on the basis of molecular phylogenetic evidence, the genus was expanded to include species f ...
''. It is a boreal to
subarctic The subarctic zone is a region in the Northern Hemisphere immediately south of the true Arctic, north of humid continental regions and covering much of Alaska, Canada, Iceland, the north of Scandinavia, Siberia, and the Cairngorms. Generally, ...
woodland
subshrub A subshrub (Latin ''suffrutex'') or dwarf shrub is a short shrub, and is a woody plant. Prostrate shrub is a related term. "Subshrub" is often used interchangeably with "bush".Jackson, Benjamin, Daydon; A Glossary of Botanic Terms with their Der ...
, 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, the ...
, 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, bot ...
, for whom the genus was named.


Description

The
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
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 A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
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 mechani ...
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, the ...
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 ''Linnaea'' is a plant genus in the honeysuckle family Caprifoliaceae. Until 2013, the genus included a single species, ''Linnaea borealis''. In 2013, on the basis of molecular phylogenetic evidence, the genus was expanded to include species f ...
''. The genus name had been used earlier by the Dutch botanist
Jan Frederik Gronovius Jan Frederik Gronovius (also seen as Johann Frederik and Johannes Fredericus) (10 February 1690 in Leiden – 10 July 1762 in Leiden) was a Dutch botanist notable as a patron of Linnaeus. John Clayton, a plant collector in Virginia sent him many ...
, 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 in t ...
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 U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
) 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 The subarctic zone is a region in the Northern Hemisphere immediately south of the true Arctic, north of humid continental regions and covering much of Alaska, Canada, Iceland, the north of Scandinavia, Siberia, and the Cairngorms. Generally, ...
, 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 (topography), summit area, and ...
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 Continent#Subcontinents, subcontinent of Eurasia ...
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, Sw ...
, 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 area ...
south to northern
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
, and in North America south to northern
California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
and to
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...
and
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ker ...
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 Bur ...
(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 th ...
) 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. They ...
in the east. ''Linnaea borealis'' is
self-incompatible Self-incompatibility (SI) is a general name for several genetic mechanisms that prevent self-fertilization in sexually reproducing organisms, and thus encourage outcrossing and allogamy. It is contrasted with separation of sexes among individuals ...
, requiring
cross-pollination Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, ...
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 angiospe ...
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 gametophyt ...
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
stolon In biology, stolons (from Latin '' stolō'', genitive ''stolōnis'' – "branch"), also known as runners, are horizontal connections between organisms. They may be part of the organism, or of its skeleton; typically, animal stolons are external s ...
s 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 A seedling is a young 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 (embryonic root), the hypocotyl (embry ...
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 (geology), 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 fina ...
("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 slopes with persisting underground
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 o ...
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 the ...
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 b ...
.
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 to ...
s consider this plant to be an
indicator species A bioindicator is any species (an indicator species) or group of species whose function, population, or status can reveal the qualitative status of the environment. The most common indicator species are animals. For example, copepods and other sma ...
of
ancient woodland In the United Kingdom, an ancient woodland is a woodland that has existed continuously since 1600 or before in England, Wales and Northern Ireland (or 1750 in Scotland). Planting of woodland was uncommon before those dates, so a wood present in 16 ...
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 The Cairngorms ( gd, Am Monadh Ruadh) are a mountain range in the eastern Highlands of Scotland closely associated with the mountain Cairn Gorm. The Cairngorms became part of Scotland's second national park (the Cairngorms National Park) on 1 S ...
; the southernmost locations are four sites in
Northumberland Northumberland () is a 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 Abbey. It is bordered by land on ...
and one in
County Durham County Durham ( ), officially simply Durham,UK General Acts 1997 c. 23Lieutenancies Act 1997 Schedule 1(3). From legislation.gov.uk, retrieved 6 April 2022. is a ceremonial county in North East England.North East Assembly About North East E ...
. 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 the ...
, 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 Vegetative reproduction (also known as vegetative propagation, vegetative multiplication or cloning) is any form of asexual reproduction occurring in plants in which a new plant grows from a fragment or cutting of the parent plant or spec ...
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 states, a federal district, five major unincorporated territorie ...
, ''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 state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...
,
Iowa Iowa () is a state in the 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: Wisconsin to the northeast, Illinois to the ...
,
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 Native Americans in the United States, indigenous Dakota people, Dakota Sioux. North Dakota is bordered by the Canadian provinces of Saskatchewan and Manitoba to the north a ...
,
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, ...
, 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 Bur ...
, and was known historically but now considered
extirpated Local extinction, also known as extirpation, refers to a species (or other taxon) of plant or animal that ceases to exist in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinct ...
or possibly so in
Illinois Illinois ( ) is a U.S. state, state in the Midwestern United States, Midwestern United States. Its largest metropolitan areas include the Chicago metropolitan area, and the Metro East section, of Greater St. Louis. Other smaller metropolita ...
,
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 s ...
,
New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north and east by the state of New York; on the east, southeast, and south by the Atlantic Ocean; on the west by the Delaware ...
,
Ohio Ohio () is a state in the Midwestern region of the United States. Of the fifty U.S. states, it is the 34th-largest by area, and with a population of nearly 11.8 million, is the seventh-most populous and tenth-most densely populated. The sta ...
,
Rhode Island Rhode Island (, like ''road'') is a U.S. state, state in the New England region of the Northeastern United States. It is the List of U.S. states by area, smallest U.S. state by area and the List of states and territories of the United States ...
, 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 th ...
. 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 tot ...
, ''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 A microclimate (or micro-climate) is a local set of atmospheric conditions that differ from those in the surrounding areas, often with a slight difference but sometimes with a substantial one. The term may refer to areas as small as a few squa ...
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 E ...
has become a significant concern for the conservation of this species in such places, such as
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. It was designated a National Natural Landmark in 2012. Ice Mo ...
in West Virginia, a low-elevation algific talus slope 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 fo ...
,Visit Småland
Retrieved 25 February 2012.
the home province of Linnaeus.


References

*''The Linnaeus Link Project'' 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