''Linnaea borealis'' is a
species
A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
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 and easte ...
(the honeysuckle family). It is the only species in the genus ''Linnaea''.
It is a
boreal to
subarctic
The subarctic zone is a region in the Northern Hemisphere immediately south of the true Arctic, north of hemiboreal regions and covering much of Alaska, Canada, Iceland, the north of Fennoscandia, Northwestern Russia, Siberia, and the Cair ...
woodland
subshrub
A subshrub (Latin ''suffrutex'') or undershrub is either a small shrub (e.g. prostrate shrubs) or a perennial that is largely herbaceous but slightly woody at the base (e.g. garden pink and florist's chrysanthemum). The term is often interch ...
, commonly known as twinflower (sometimes written twin flower).
This plant was a favourite of
Carl Linnaeus
Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
, founder of the modern system of
binomial nomenclature
In taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, altho ...
, after whom the genus was named.
Description
The
perennial
In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...
stems of ''Linnaea borealis'' are slender, pubescent, and prostrate, growing to long, with opposite
evergreen
In botany, an evergreen is a plant which has Leaf, foliage that remains green and functional throughout the year. This contrasts with deciduous plants, which lose their foliage completely during the winter or dry season. Consisting of many diffe ...
rounded oval
leaves
A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
long and broad. The flowering stems curve erect, to tall, and are leafless except at the base. The
flower
Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s are paired, pendulous, long, with a five-lobed, pale pink corolla.
''L. borealis'' is
self-incompatible, requiring
cross-pollination to produce viable
seed
In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s; since
pollen
Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
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, a stolon ( from Latin ''wikt:stolo, stolō'', genitive ''stolōnis'' – "branch"), also known as a runner, is a horizontal connection between parts of an organism. It may be part of the organism, or of its skeleton. Typically, animal ...
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 ...
.
Taxonomy
''Linnaea borealis'' was first formally described by
Carl Linnaeus
Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
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 genus, genera. It is the first work to consistently apply binomial nomenclature ...
''.
It was 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 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 ( ) is a non-contiguous U.S. state on the northwest extremity of North America. Part of the Western United States region, it is one of the two non-contiguous U.S. states, alongside Hawaii. Alaska is also considered to be the north ...
to
California
California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
)
The English name "twinflower" for ''Linnaea borealis'' refers to the plant's paired flowers.
In 2013 Christenhusz ''et al.'' concluded that the genus ''
Abelia'', as then circumscribed, was polyphyletic, and proposed merging ''Abelia'' and several other genera in tribe Linnaeeae into ''Linnaea''.
Wang, Landrein, ''et al.'' proposed reorganizing the tribe into several new and existing genera, and keeping ''Linnaea'' a monotypic genus.
[Wang H-F, Landrein S, Dong W-P, Nie Z-L, Kondo K, Funamoto T, et al. (2015) Molecular Phylogeny and Biogeographic Diversification of Linnaeoideae (Caprifoliaceae s. l.) Disjunctly Distributed in Eurasia, North America and Mexico. PLoS ONE 10(3): e0116485. doi:10.1371/journal. pone.0116485] The circumscription proposed by Wang, Landrein, ''et al.'' is currently accepted by
Plants of the World Online
Plants of the World Online (POWO) is an online taxonomic database published by the Royal Botanic Gardens, Kew.
History
Following the Convention on Biological Diversity, the Royal Botanic Gardens in Kew launched Plants of the World Online i ...
.
[
]
Distribution and habitat
''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 hemiboreal regions and covering much of Alaska, Canada, Iceland, the north of Fennoscandia, Northwestern Russia, Siberia, and the Cair ...
, boreal, or cool temperate forest
A forest is an ecosystem characterized by a dense ecological community, community of trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, ...
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 continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
south to the Alps
The Alps () are some of the highest and most extensive mountain ranges in Europe, stretching approximately across eight Alpine countries (from west to east): Monaco, France, Switzerland, Italy, Liechtenstein, Germany, Austria and Slovenia.
...
, in Asia
Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
south to northern Japan
Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
, and in North America south to northern California
California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
and to Arizona
Arizona is a U.S. state, state in the Southwestern United States, Southwestern region of the United States, sharing the Four Corners region of the western United States with Colorado, New Mexico, and Utah. It also borders Nevada to the nort ...
and New Mexico
New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
in the west, and to West Virginia
West Virginia is a mountainous U.S. state, state in the Southern United States, Southern and Mid-Atlantic (United States), Mid-Atlantic regions of the United States.The United States Census Bureau, Census Bureau and the Association of American ...
(and formerly Tennessee
Tennessee (, ), officially the State of Tennessee, is a landlocked U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders Kentucky to the north, Virginia to the northeast, North Carolina t ...
) in the Appalachian Mountains
The Appalachian Mountains, often called the Appalachians, are a mountain range in eastern to northeastern North America. The term "Appalachian" refers to several different regions associated with the mountain range, and its surrounding terrain ...
in the east.
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 ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
("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 ice.
Conservation
While the three subspecies of ''L. 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 north-west coast of continental Europe, consisting of the countries England, Scotland, and Wales. With an area of , it is the largest of the British Isles, the List of European ...
, ''L. borealis'' ssp. ''borealis'' is listed as "nationally scarce", growing mainly in open pine woodlands in Scotland
Scotland is a Countries of the United Kingdom, country that is part of the United Kingdom. It contains nearly one-third of the United Kingdom's land area, consisting of the northern part of the island of Great Britain and more than 790 adjac ...
and northernmost England
England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
. Forester
A forester is a person who practises forest management and 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. Fores ...
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 woodlands, 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 () are a mountain range in the eastern Scottish Highlands, Highlands of Scotland closely associated with the mountain Cairn Gorm. The Cairngorms became part of Scotland's second national parks of Scotland, national park (the Cairn ...
; the southernmost locations are four sites in Northumberland
Northumberland ( ) is a ceremonial counties of England, ceremonial county in North East England, on the Anglo-Scottish border, border with Scotland. It is bordered by the North Sea to the east, Tyne and Wear and County Durham to the south, Cumb ...
and one in County Durham
County Durham, officially simply Durham, is a ceremonial county in North East England.UK General Acts 1997 c. 23Lieutenancies Act 1997 Schedule 1(3). From legislation.gov.uk, retrieved 6 April 2022. The county borders Northumberland and Tyne an ...
. The sparseness of the sites is responsible for the continued decline of the plant in the country. In Scotland
Scotland is a Countries of the United Kingdom, country that is part of the United Kingdom. It contains nearly one-third of the United Kingdom's land area, consisting of the northern part of the island of Great Britain and more than 790 adjac ...
, 37% of ''L. borealis'' patches studied consisted of a single genotype, 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 (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
, ''L. borealis'' ssp. ''americana'' is of conservation concern in several states along or near the southern edge of the species' range, including Arizona, Iowa
Iowa ( ) is a U.S. state, state in the upper Midwestern United States, Midwestern region of the United States. It borders the Mississippi River to the east and the Missouri River and Big Sioux River to the west; Wisconsin to the northeast, Ill ...
, Massachusetts
Massachusetts ( ; ), officially the Commonwealth of Massachusetts, is a U.S. state, state in the New England region of the Northeastern United States. It borders the Atlantic Ocean and the Gulf of Maine to its east, Connecticut and Rhode ...
, North Dakota
North Dakota ( ) is a U.S. state in the Upper Midwest, named after the indigenous Dakota people, Dakota and Sioux peoples. It is bordered by the Canadian provinces of Saskatchewan and Manitoba to the north and by the U.S. states of Minneso ...
, Pennsylvania
Pennsylvania, officially the Commonwealth of Pennsylvania, is a U.S. state, state spanning the Mid-Atlantic (United States), Mid-Atlantic, Northeastern United States, Northeastern, Appalachian, and Great Lakes region, Great Lakes regions o ...
, and West Virginia, and was known historically but now considered extirpated
Local extinction, also extirpation, is the termination of a species (or other taxon) in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinctions.
Local extinctions mark a chan ...
or possibly so in Illinois
Illinois ( ) is a U.S. state, state in the Midwestern United States, Midwestern United States. It borders on Lake Michigan to its northeast, the Mississippi River to its west, and the Wabash River, Wabash and Ohio River, Ohio rivers to its ...
, Indiana
Indiana ( ) is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Lake Michigan to the northwest, Michigan to the north and northeast, Ohio to the east, the Ohio River and Kentucky to the s ...
, New Jersey
New Jersey is a U.S. state, state located in both the Mid-Atlantic States, Mid-Atlantic and Northeastern United States, Northeastern regions of the United States. Located at the geographic hub of the urban area, heavily urbanized Northeas ...
, Ohio
Ohio ( ) is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Lake Erie to the north, Pennsylvania to the east, West Virginia to the southeast, Kentucky to the southwest, Indiana to the ...
, Rhode Island
Rhode Island ( ) is a state in the New England region of the Northeastern United States. It borders Connecticut to its west; Massachusetts to its north and east; and the Atlantic Ocean to its south via Rhode Island Sound and Block Is ...
, and Tennessee.
In Canada
Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
, ''Linnaea borealis'' ssp. ''longiflora'' is considered of conservation significance in the Yukon
Yukon () is a Provinces and territories of Canada, territory of Canada, bordering British Columbia to the south, the Northwest Territories to the east, the Beaufort Sea to the north, and the U.S. state of Alaska to the west. It is Canada’s we ...
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
Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
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 with persisting buried ice.
In culture
Linnaeus took ''L. 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:
The flower of ''Linnaea borealis'' is the provincial flower of Småland
Småland () is a historical Provinces of Sweden, 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 la ...
,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 scientific collection, collections that include current and historical records of animals, plants, Fungus, fungi, ecosystems, geology, paleo ...
members.
*''Species and habitat conservation'' from Plantlife.org.u
an
* Twinflower species profil
External links
*
{{Authority control
Caprifoliaceae
Caprifoliaceae genera
Monotypic asterid genera
Alpine flora
Flora of Northern America
Flora of temperate Asia
Flora of Europe
Plants described in 1753
Taxa named by Carl Linnaeus
Commemoration of Carl Linnaeus