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Biology

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Twinflower is a perennial, and the evergreen leaves last some two years before being shed and replaced. It can spread by seed or vegetatively, the seeds being dispersed by animals and birds, whilst new shoots come from plants at least five years old. The plant is shallow-rooted, susceptible to both drought and fire, but it can re-establish itself on fire-damaged or disturbed ground. However, it is believed that the seeds do not survive for long within the seedbank. Twinflower has the ability to produce vegetative clones of itself, although these do not appear to be able to self-fertilise.
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Conservation

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The twinflower is listed as a priority species in the UK Biodiversity Action Plan (UK BAP). The plant is known to have declined considerably in recent years, and is now found on some 50 sites in north-east Scotland. There used to be a small population in northern England, thought to have arrived with pine seedlings. Native pine woodland is categorised as a priority habitat under the UK Biodiversity Action Plan, and many populations of twinflower in Scotland are on designated sites, so the plant enjoys a fair measure of protection. There are also populations in cultivation, notably at the Royal Botanic Gardens, Edinburgh, and the botanic gardens of both Edinburgh and Aberdeen Universities. However, it is felt that more should be done to improve this attractive plant's chances of survival in this country. A number of projects are underway to variously assess the true status of twinflower, and study the problems of sexual regeneration. This may include plans to re-locate specimens to different areas in an attempt to overcome the problem of non-fertilisation between plant clones, and help produce viable seed stock. There are hopes that more can be learned about the plant's biology by exchanging information with botanists from other countries where twinflower grows. The Forestry Commission are one of the partners working on the plans to conserve twinflower, and they are the main UK contact agency for this species. Plantlife Scotland and Scottish Natural Heritage are also engaged in research work for this attractive species.
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Description

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Twinflower is a member of the honeysuckle family and, like some of its relations, it is a creeper. However, twinflower grows much closer to the ground than honeysuckle, creeping beneath other low-growing plants with its thread-like stem. It is an evergreen perennial, with small oval leaves about a centimetre in length, and delicate bell-shaped flowers. As its name suggests, the flowers usually occur in pairs (although, there may only be one), drooping from a long stalk and coloured white or pink, with red stripes inside the bell. The plant's scientific name Linnaea refers to the fact that it was named in honour of the 18th century Swedish naturalist, Carl Linné, often called 'the father of modern taxonomy'. Linné, who called himself 'Linneaus', developed the system that is still used to identify all species of plant, animal, fungus, single and simple multi-cell organisms and bacteria – the five Kingdoms of life on Earth. Every newly discovered organism (and newly discovered fossil) is identified and classified with two names (or three if it is discovered to be a variation of an existing species), derived from Latin or Greek. The names usually describe one of the species' characteristics, (such as borealis – meaning 'of the north'). A full classification of the species will place it in one of the five Kingdoms. From there, its other characteristics determine that species' place within a number of other sub-categories (such as those shown above). Linneaus was typically self-deprecatory in response to the honour of having twinflower named after him. He wrote; "Linnaea was named by the celebrated Gronovius and is a plant of Lapland, lowly, insignificant and disregarded, flowering but for a brief space - from Linnaeus who resembles it."
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Habitat

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The plant occurs in both native and planted Scots pine woodland, preferring light shade and acidic soils.
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Range

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Twinflower is found over Northern Europe, up to an altitude of 2,400 metres. It is also found in the Alps, and the Caucasus and Carpathian mountain ranges. In the UK, it is limited to the north-eastern part of Scotland where it still grows as a native plant, although there was a small introduced population in the north of England.
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Status

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Classified as Nationally Scarce in the UK.
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Threats

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The main threats to twinflower come from damage during forestry operations, although some think it might improve the plant's survival within an area through the dispersal of plant fragments. Overgrazing by sheep and deer can be a threat as well, although a lack of grazing can lead to the plants being crowded out by more dominant vegetation. There is also the risk of loosing small isolated populations of related plants.
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Associations

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Foodplant / feeds on
epiphyllous, subiculately pseudothecial stroma of Metacoleroa dickiei feeds on leaf of Linnaea borealis

Foodplant / sap sucker
nymph of Metatropis rufescens ssp. linnaeae sucks sap of Linnaea borealis

In Great Britain and/or Ireland:
Foodplant / spot causer
epiphyllous, black, shining pycnidium of Septoria coelomycetous anamorph of Septoria linnaeae causes spots on live leaf of Linnaea borealis

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Broad-scale Impacts of Plant Response to Fire

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More info for the terms: fire use, prescribed fire, restoration

The following Research Project Summaries provide information on prescribed
fire use and postfire response of plant community species including twinflower:

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bibliographic citation
Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Common Names

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twinflower
northern twinflower
western twinflower
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Description

provided by Fire Effects Information System Plants
More info for the terms: capsule, duff, forest, fruit, litter, root crown, shrub, stolon

Twinflower is a native, evergreen, dwarf shrub [20].  It is creeping or
trailing in growth form, with numerous short aerial stems rising from
the stolon.  With time, stolons may become shallowly buried beneath
litter and duff layers [17]; maximum depth of twinflower stolons in a
mixed conifer old-growth forest of central Oregon was 0.11 inch (0.27
cm) below the soil surface [2].  Aerial stems become woody with age but
rarely exceed 0.12 inch (0.30 cm) in diameter.  The root crown of these
stems is positioned at or just beneath the duff or soil surface [17].
Twinflower has a shallow, fibrous network of roots with their growing
points within and slightly below the duff layer [56].  In the central
Oregon study mentioned above, average root depth was 0.11 inch (0.28 cm)
[2].  Twinflower fruit is a small, dry, one-seeded capsule [30,62].
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Distribution

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The typical subspecies of twinflower is distributed from eastern Alaska
across Siberia and northern Europe [35].  Pacific twinflower occurs
along the West Coast from southern Alaska to northern California
[35,55].  American twinflower is distributed from interior Alaska across
Canada to Newfoundland and south to northern Arizona and New Mexico,
South Dakota, Indiana, West Virginia, and New Jersey [28,35,41].
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bibliographic citation
Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Fire Ecology

provided by Fire Effects Information System Plants
More info for the terms: duff, fire regime, seed

Twinflower is a fire avoider [56]. During fire, small patches of
twinflower in draws, moist duff, or other protected places usually
escape burning [15,17].  Twinflower most commonly establishes in burn
areas from stolons produced by these unburned plants.  It also
establishes from animal-dispersed seed [3,56].

FIRE REGIMES :
Find fire regime information for the plant communities in which this
species may occur by entering the species name in the FEIS home page under
"Find FIRE REGIMES".
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Fire Management Considerations

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More info for the terms: forbs, forest, fuel

Brown and Marsden [11] have developed an equation for estimating fuel
weight of twinflower and other small woody plants, grasses, and forbs in
coniferous forests of western Montana and northern Idaho.  Brown [10]
developed a method of determining bulk densities of nonuniform surface
fuels in subalpine fir/twinflower and other forest types of that region.
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Growth Form (according to Raunkiær Life-form classification)

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More info on this topic.

More info for the term: chamaephyte

Chamaephyte
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat characteristics

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More info for the terms: peatland, xeric

Twinflower grows in soils derived from a variety of parent materials.
Soil texture and nutrient levels also vary, and soil moisture levels
range from xeric to hydric [13,16,30,59].  The pH range of
twinflower-supporting soils in peatland bogs of Minnesota and
Saskatchewan is 4.0 to 7.0 [31,39].  Twinflower occurs on all aspects
[18].  It is found at the following elevations:

                                  feet            meters
northern ID; western MT      2,200 - 5,900      670 - 1,800 [13,25]
southern CO; northern NM     7,900 - 9,800    2,400 - 2,900 [19]
UT                           6,000 - 9,600    1,830 - 2,900 [67]
CA                           4,000 - 8,000    1,200 - 2,400 [55]
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat: Cover Types

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This species is known to occur in association with the following cover types (as classified by the Society of American Foresters):

     1  Jack pine
     5  Balsam fir
    12  Black spruce
    13  Black spruce - tamarack
    15  Red pine
    16  Aspen
    18  Paper birch
    20  White pine - northern red oak - red maple
    21  Eastern white pine
    22  White pine - hemlock
    23  Eastern hemlock
    24  Hemlock - yellow birch
    25  Sugar maple - beech - yellow birch
    26  Sugar maple - basswood
    27  Sugar maple
    30  Red spruce - yellow birch
    31  Red spruce - sugar maple - beech
    32  Red spruce
    33  Red spruce - balsam fir
    35  Paper birch - red spruce - balsam fir
    37  Northern white-cedar
    38  Tamarack
    51  White pine - chestnut oak
    60  Beech - sugar maple
   107  White spruce
   201  White spruce
   202  White spruce - paper birch
   203  Balsam poplar
   204  Black spruce
   205  Mountain hemlock
   206  Engelmann spruce - subalpine fir
   210  Interior Douglas-fir
   211  White fir
   212  Western larch
   213  Grand fir
   215  Western white pine
   216  Blue spruce
   218  Lodgepole pine
   251  White spruce - aspen
   252  Paper birch
   253  Black spruce - white spruce
   254  Black spruce -  paper birch
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat: Ecosystem

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This species is known to occur in the following ecosystem types (as named by the U.S. Forest Service in their Forest and Range Ecosystem [FRES] Type classification):

   FRES10  White - red - jack pine
   FRES11  Spruce - fir
   FRES18  Maple - beech - birch
   FRES19  Aspen - birch
   FRES20  Douglas-fir
   FRES21  Ponderosa pine
   FRES22  Western white pine
   FRES23  Fir - spruce
   FRES24  Hemlock - Sitka spruce
   FRES25  Larch
   FRES26  Lodgepole pine
   FRES27  Redwood
   FRES28  Western hardwoods
   FRES38  Plains grasslands
   FRES44  Alpine
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat: Plant Associations

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This species is known to occur in association with the following plant community types (as classified by Küchler 1964):

More info for the terms: bog, forest

   K001  Spruce - cedar - hemlock forest
   K002  Cedar - hemlock - Douglas-fir forest
   K003  Silver fir - Douglas-fir forest
   K004  Fir - hemlock forest
   K005  Mixed conifer forest
   K006  Redwood forest
   K008  Lodgepole pine - subalpine forest
   K011  Western ponderosa forest
   K012  Douglas-fir forest
   K013  Cedar - hemlock - pine forest
   K014  Grand fir - Douglas-fir forest
   K015  Western spruce - fir forest
   K016  Eastern ponderosa forest
   K017  Black Hills pine forest
   K018  Pine - Douglas-fir forest
   K020  Spruce - fir - Douglas-fir forest
   K021  Southwestern spruce - fir forest
   K028  Mosaic of K002 and K026
   K029  California mixed evergreen forest
   K052  Alpine meadows and barren
   K066  Wheatgrass - needlegrass
   K067  Wheatgrass - bluestem - needlegrass
   K093  Great Lakes spruce - fir forest
   K094  Conifer bog
   K095  Great Lakes pine forest
   K096  Northeastern spruce - fir forest
   K106  Northern hardwoods
   K107  Northern hardwoods - fir forest
   K108  Northern hardwoods - spruce forest
   K109  Transition between K104 and K106
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Immediate Effect of Fire

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Twinflower is killed even by low-intensity fire [9,17,23,50,65].
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Importance to Livestock and Wildlife

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Bighorn sheep of Alaska and caribou of northern Canada use twinflower as
incidental forage [7,61].  It provides 9.7 percent of winter diets of
Roosevelt elk of British Columbia, Washington, and Oregon, but only 0.5
percent of their summer diets [40].  It is listed as a food item of
ruffed grouse of Idaho [36].
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Key Plant Community Associations

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More info for the terms: forest, grassland, habitat type, hardwood, taiga

Twinflower occurs in several grassland and many hardwood and coniferous
forest types.  It is named as a dominant understory or indicator species
in numerous published classifications.  A partial listing is as follows:

Preliminary plant associations of the southern Oregon Cascade Mountain
   Province [4]
Preliminary plant associations of the Siskiyou Mountain Province [5]
Ecology of wetlands in Big Meadows, Rocky Mountain National Park,
   Colorado [12]
Forest habitat types of northern Idaho:  a second approximation [13]
Classification of montane forest community types in the Cedar River
   drainage of western Washington, U.S.A. [18]
A classification of forest habitat types of northern New Mexico and
   southern Colorado [19]
Classification, description, and dynamics of plant communities after
   fire in the taiga of interior Alaska [24]
Forest vegetation of the Black Hills National Forest of South Dakota and
   Wyoming:  a habitat type classification [34]
Plant communities of Voyageurs National Park, Minnesota, U.S.A. [46]
Forest habitat types of Montana [57]
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Life Form

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More info for the terms: forb, shrub, vine

Forb-Shrub-Vine
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Nutritional Value

provided by Fire Effects Information System Plants
Dry matter nutrient content of twinflower in Manitoba is 7.0 percent
protein and 39.3 percent acid detergent fiber.  Estimated dry matter
digestibility for caribou is 59.5 percent [60].
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Occurrence in North America

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     AK  AZ  CA  CO  CT  ID  IL  IN  ME  MD
     MA  MI  MN  MT  NH  NJ  NM  NY  ND  OH
     OR  PA  RI  SD  UT  VT  WA  WV  WI  WY
     AB  BC  MB  NB  NF  NT  NS  ON  PE  PQ
     SK  YT
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Phenology

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Twinflower blooms from June through September throughout most of its
range [30,36].  Flowers last about 7 days, and fruits mature
approximately 36 days after flowering [32].  The leaves persist for 2
years [21]; season of leaf abscission was not reported in the
literature.
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Plant Response to Fire

provided by Fire Effects Information System Plants
More info for the terms: cover, density, duff, frequency, relative density

Twinflower sometimes colonizes new areas after fire.  In northwestern
Montana, broadcast burning was conducted to remove slash following
logging of subalpine fir (Abies lasiocarpa).  Twinflower had previously
been absent from the site.  Twinflower seedlings first appeared at
postfire year 6, showing 1 percent ground cover.  At postfire year 9,
twinflower cover was still at 1 percent [65].  Broadcast burning was
also conducted at a nearby site where prefire twinflower cover was 8
percent.  The fire removed 11 percent of the duff, and all existing
twinflower was killed.  As with the previously mentioned fire,
twinflower seedlings first established at postfire year 6, showing 1
percent cover.  By postfire year 8, twinflower cover at this site had
increased to 8 percent [65].

In Pacific silver fir (Abies amabilis)-subalpine fir forests of central
British Columbia, twinflower frequency on 4- to 22-year-old burns was 60
percent.  Frequency on 37- to 75-year-old burns was 70 percent [26].

Two consecutive annual, low-intensity prescribed fires were conducted on
the Petawawa Experimental Station in Ontario.  Prefire relative
twinflower density was 9.65 percent.  After the first fire, twinflower
relative density lowered to 0.14 percent.  It dropped to 0.11 percent
after the second fire [52].
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Post-fire Regeneration

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More info for the terms: root crown, secondary colonizer

   Surface rhizome/chamaephytic root crown
   Secondary colonizer - off-site seed
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Regeneration Processes

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More info for the terms: density, forest, litter, seed

Vegetative reproduction by stolons is the primary method of twinflower
regeneration [2,21].  Twinflower first produces stolons at 5 to 10 years
of age [21].

Sexual reproduction is uncommon, but seedlings are occasionally found in
burned or other disturbed areas [21,65].  Pollination is effected by
native bees and syrphid wasps [6,30]; rarely, plants are self-fertile
[30].  Twinflower produces abundant seed [29], which apparently does not
persist in seed banks.  A study in a paper birch-balsam fir (Betula
papyrifera-Abies balsamea) forest on Mont Jacques-Cartier in Quebec
showed a twinflower seed density of six seeds per square meter, with
none of the seeds proving viable [54].  In British Columbia, Kellman
[42] found only one viable twinflower seed in 34 samples collected from
the litter, A, and B soil horizons.  Twinflower seed attaches to the
fur, hides, or feathers of animals, which serve as dispersal agents.
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Regional Distribution in the Western United States

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This species can be found in the following regions of the western United States (according to the Bureau of Land Management classification of Physiographic Regions of the western United States):

    1  Northern Pacific Border
    2  Cascade Mountains
    3  Southern Pacific Border
    4  Sierra Mountains
    5  Columbia Plateau
    6  Upper Basin and Range
    8  Northern Rocky Mountains
    9  Middle Rocky Mountains
   10  Wyoming Basin
   11  Southern Rocky Mountains
   12  Colorado Plateau
   15  Black Hills Uplift
   16  Upper Missouri Basin and Broken Lands
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Successional Status

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More info for the terms: climax, forest, peat, presence

Twinflower tolerates a wide spectrum of light intensity.  It grows in
full daylight in alpine meadows and peat bogs [12,39], but subcanopy
light has been measured at 2 percent of full daylight in a closed-canopy
mountain hemlock-western redcedar (Tsuga heterophylla-Thuja
plicata)/twinflower type near Vancouver, British Columbia [51].

Twinflower is found in recently disturbed, seral, and climax plant
communities.  Irwin [37] reported it as "abundant" after shelterwood
cutting in a western hemlock-western redcedar forest of northern Idaho.
Several authors have noted its presence in various climax forest types
[13,17,18,24,25,38].

In a study of understories of young (30-80 yrs), mature (80-195 yrs),
and old-growth (195-900 yrs) Douglas-fir forests of the Cascade Range of
Washington and Oregon, percent occurrence of twinflower by forest age
was as follows [63]:

     young:       89
     mature:      80
     old-growth:  98
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Taxonomy

provided by Fire Effects Information System Plants
The currently accepted scientific name of twinflower is Linnaea borealis
L. [28,33,55,62,62]. It is a monotypic genus [28]. Recognized
subspecies are as follows:

Linnaea borealis ssp. borealis [35]
Linnaea borealis sps. americana (Forbes) Hult. [35,45] (American twinflower)
Linnaea borealis ssp. longiflora (Torr.) Hult. [35,55,62] (Pacific twinflower)

Some authorities [33,62] recognize L. b. ssp. borealis and L. b. ssp.
longiflora as the only distinct subspecies of twinflower.
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Howard, Janet L. 1993. Linnaea borealis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Linnaea borealis

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Linnaea borealis is a species of flowering plant 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, founder of the modern system of binomial nomenclature, for whom the genus was named.

Description

The leaves are under 1cm long, with a few shallow teeth on the upper half.

The perennial stems of Linnaea borealis are slender, pubescent, and prostrate, growing to 20–40 centimetres (7.9–15.7 in) long, with opposite evergreen rounded oval leaves 3–10 millimetres (0.12–0.39 in) long and 2–7 mm (0.079–0.276 in) broad. The flowering stems curve erect, to 4–8 cm (1.6–3.1 in) tall, and are leafless except at the base. The flowers are paired, pendulous, 7–12 mm (0.28–0.47 in) long, with a five-lobed, pale pink corolla.[1][2]

Taxonomy

Linnaea borealis was first formally described by Carl Linnaeus in 1753 in Species Plantarum.[3] 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.[4]

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 to California)

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

Ecology and distribution

Linnaea borealis ssp. longiflora in the Mount Baker-Snoqualmie National Forest, Washington, U.S.
Linnaea borealis may form long-persisting clonal colonies

Linnaea borealis has a circumpolar distribution in moist subarctic, boreal, or cool temperate forests, extending further south at higher elevations in various mountains, in Europe south to the Alps, in Asia south to northern Japan, and in North America south to northern California and to Arizona and New Mexico in the west, and to West Virginia (and formerly Tennessee) in the Appalachian Mountains in the east.[5] [6]

Linnaea borealis is self-incompatible, requiring cross-pollination to produce viable seeds; since pollen dispersal is usually not far, individuals and clonal colonies can become reproductively isolated.[7] Regardless of seed production, Linnaea plants in a particular area often spread by stolons to form clonal patches of the same genotype. 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 ("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 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, Linnaea borealis ssp. borealis is listed as "nationally scarce", growing mainly in open pine woodlands in Scotland and northernmost England. Foresters consider this plant to be an indicator species 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 southernmost locations are four sites in Northumberland and one in County Durham. The sparseness of the sites is responsible for the continued decline of the plant in the country. In Scotland, 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.[7]

In the United States, Linnaea borealis ssp. americana is of conservation concern in several states along or near the southern edge of the species' range, including Arizona, Iowa, Massachusetts, North Dakota, Pennsylvania, and West Virginia, and was known historically but now considered extirpated or possibly so in Illinois, Indiana, New Jersey, Ohio, Rhode Island, and Tennessee.

In Canada, Linnaea borealis ssp. longiflora is considered of conservation significance in the Yukon 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 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.

Cultural references

Rue Linnea Borealis, Cogne (Aosta Valley)

Linnaeus took Linnaea borealis as his own personal symbol when he was raised to the Swedish nobility in 1757. In his Critica Botanica (1737), Linnaeus had used Gronovius's name Linnaea as an example to advocate the use of commemorative personal names as botanical names:

it is commonly believed that the name of a plant which is derived from that of a botanist shows no connection between the two...[but]...Linnaea was named by the celebrated Gronovius and is a plant of Lapland, lowly, insignificant, disregarded, flowering but for a brief space — after Linnaeus who resembles it.[8][9]

The flower of Linnaea borealis is the provincial flower of Småland,[10] the home province of Linnaeus.

References

  1. ^ Klinkenberg, Brian, ed. (2014). "Linnaea borealis". E-Flora BC: Electronic Atlas of the Plants of British Columbia [eflora.bc.ca]. Lab for Advanced Spatial Analysis, Department of Geography, University of British Columbia, Vancouver. Retrieved 2015-01-24.
  2. ^ Giblin, David, ed. (2015). "Linnaea borealis". WTU Herbarium Image Collection. Burke Museum, University of Washington. Retrieved 2015-01-24.
  3. ^ "Linnaea borealis L." International Plant Names Index (IPNI). Royal Botanic Gardens, Kew; Harvard University Herbaria & Libraries; Australian National Botanic Gardens. Retrieved 2018-01-22.
  4. ^ Christenhusz, Maarten J.M. (2013). "Twins are not alone: a recircumscription of Linnaea (Caprifoliaceae)". Phytotaxa. 125 (1): 25–32. doi:10.11646/phytotaxa.125.1.4.
  5. ^ Sullivan, Steven. K. (2015). "Linnaea borealis". Wildflower Search. Retrieved 2015-01-24.
  6. ^ USDA, NRCS (n.d.). "Linnaea borealis". The PLANTS Database (plants.usda.gov). Greensboro, North Carolina: National Plant Data Team. Retrieved 2015-01-24.
  7. ^ a b Scobie, A. R.; Wilcock, C. C. (2009). "Limited mate availability decreases reproductive success of fragmented populations of Linnaea borealis, a rare, clonal self-incompatible plant". Annals of Botany. 103 (6): 835–846. doi:10.1093/aob/mcp007. PMC 2707897. PMID 19181748.
  8. ^ Stafleu, Frans A. 1971. Linnaeus and the Linnaeans: the Spreading of their Ideas in Systematic Botany, 1735–1789. Utrecht: International Association for Plant Taxonomy. ISBN 90-6046-064-2. p. 83.
  9. ^ Core, Earl L. (1975), The Wondrous Year: West Virginia Through the Seasons, Grantsville, West Virginia: Seneca Books, pg 45.
  10. ^ Visit Småland Archived 2012-09-18 at archive.today Retrieved 25 February 2012.

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Linnaea borealis: Brief Summary

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Linnaea borealis is a species of flowering plant 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, founder of the modern system of binomial nomenclature, for whom the genus was named.

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