Index of Species Information

SPECIES:  Linnaea borealis

Introductory

SPECIES: Linnaea borealis
Photo by Rob Routledge, Sault College, Bugwood.org
AUTHORSHIP AND 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/ [].

Revisions: Photo added on 28 April 2014.

ABBREVIATION : LINBOR SYNONYMS : NO-ENTRY SCS PLANT CODE : LIBO3 LIBOL COMMON NAMES : twinflower northern twinflower western twinflower TAXONOMY : 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. LIFE FORM : Forb-Shrub-Vine FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY

DISTRIBUTION AND OCCURRENCE

SPECIES: Linnaea borealis
GENERAL DISTRIBUTION : 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]. ECOSYSTEMS :    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 STATES :      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 BLM PHYSIOGRAPHIC REGIONS :     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 KUCHLER PLANT ASSOCIATIONS :    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 SAF COVER TYPES :      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 SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : 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]

MANAGEMENT CONSIDERATIONS

SPECIES: Linnaea borealis
IMPORTANCE TO LIVESTOCK AND WILDLIFE : 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]. PALATABILITY : NO-ENTRY NUTRITIONAL VALUE : 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]. COVER VALUE : NO-ENTRY VALUE FOR REHABILITATION OF DISTURBED SITES : NO-ENTRY OTHER USES AND VALUES : NO-ENTRY OTHER MANAGEMENT CONSIDERATIONS : NO-ENTRY

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Linnaea borealis
GENERAL BOTANICAL CHARACTERISTICS : 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]. RAUNKIAER LIFE FORM : Chamaephyte REGENERATION PROCESSES : 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. SITE CHARACTERISTICS : 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] SUCCESSIONAL STATUS : 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 SEASONAL DEVELOPMENT : 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.

FIRE ECOLOGY

SPECIES: Linnaea borealis
FIRE ECOLOGY OR ADAPTATIONS : 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]. POSTFIRE REGENERATION STRATEGY :    Surface rhizome/chamaephytic root crown    Secondary colonizer - off-site seed

FIRE EFFECTS

SPECIES: Linnaea borealis
IMMEDIATE FIRE EFFECT ON PLANT : Twinflower is killed even by low-intensity fire [9,17,23,50,65]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : 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]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : The following Research Project Summaries provide information on prescribed fire use and postfire response of plant community species including twinflower: FIRE MANAGEMENT CONSIDERATIONS : 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.

REFERENCES

SPECIES: Linnaea borealis
REFERENCES :  1.  Achuff, Peter L. 1989. Old-growth forests of the Canadian Rocky Mountain        national parks. Natural Areas Journal. 9(1): 12-26.  [7442]  2.  Antos, Joseph A.; Zobel, Donald B. 1984. Ecological implications of        belowground morphology of nine coniferous forest herbs. Botanical        Gazette. 145(4): 508-517.  [17417]  3.  Archibold, O. W. 1980. Seed input into a postfire forest site in        northern Saskatchewan. Canadian Journal of Forest Research. 10: 129-134.        [4506]  4.  Atzet, Thomas; McCrimmon, Lisa A. 1990. Preliminary plant associations        of the southern Oregon Cascade Mountain Province. Grants Pass, OR: U.S.        Department of Agriculture, Forest Service, Siskiyou National Forest. 330        p.  [12977]  5.  Atzet, Thomas; Wheeler, David L. 1984. Preliminary plant associations of        the Siskiyou Mountain Province. Portland, OR: U.S. Department of        Agriculture, Forest Service, Pacific Northwest Region. 278 p.  [9351]  6.  Barrett, Spencer C.; Helenurm, Kaius. 1987. The reproductive biology of        boreal forest herbs. I. Breeding systems and pollination. Canadian        Journal of Botany. 65: 2036-2046.  [6624]  7.  Bentz, Jerry A.; Woodard, Paul M. 1988. Vegetation characteristics and        bighorn sheep use on burned and unburned areas in Alberta. Wildlife        Society Bulletin. 16(2): 186-193.  [15276]  8.  Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals,        reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's        associations for the eleven western states. Tech. Note 301. Denver, CO:        U.S. Department of the Interior, Bureau of Land Management. 169 p.        [434]  9.  Bradley, Anne Foster. 1984. Rhizome morphology, soil distribution, and        the potential fire survival of eight woody understory species in western        Montana. Missoula, MT: University of Montana. 183 p. Thesis.  [502] 10.  Brown, James K. 1981. Bulk densities of nonuniform surface fuels and        their application to fire modeling. Forest Science. 27(4): 667-683.        [13269] 11.  Brown, James K.; Marsden, Michael A. 1976. Estimating fuel weights of        grasses, forbs, and small woody plants. Res. Note INT-210. Ogden, UT:        U.S. Department of Agriculture, Forest Service, Intermountain Forest &        Range Experiment Station. 11 p.  [5030] 12.  Cooper, David J. 1990. Ecology of wetlands in Big Meadows, Rocky        Mountain National Park, Colorado. Biological Report 90(15). Washington,        DC: U.S. Department of the Interior, Fish and Wildlife Service. 45 p.        [16106] 13.  Cooper, Stephen V.; Neiman, Kenneth E.; Roberts, David W. 1991. 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